Freshwater Fishes of Iran

Introduction

Revised:  06 December 2011

Contents

The main chapters are:-

1. Introduction - includes as one file Contents, Acknowledgements, Purpose, Materials and Methods, History of Research, Fisheries, Geography, Climate, Habitats, Environmental Change, Drainage Basins, Scientific Names, Fish Structure, Collecting Fishes, Preserving Fishes, Checklists, Glossaries, and Quotes.

2. Keys - active but still under construction.

3. Species Accounts - see below for links.

4. Bibliography 1711-2011

Alphabetical Links to Genera

Abramis     Acanthalburnus     Acanthobrama     Acanthopagrus     Acipenser     Alburnoides     Alburnus     Alosa     Anatirostrum     Anguilla    Aphanius     Aspidoparia    Aspiolucius    Aspius    Atherina     Babka     Barbus     Barilius     Benthophiloides     Benthophilus     Blicca     Boleophthalmus     Capoeta     Capoetobrama     Carasobarbus     Carassius     Carcharhinus     Caspiomyzon     Caspiosoma     Channa     Chanos     Chasar     Chondrostoma     Clupeonella     Cobitis     Coregonus     Crossocheilus     Ctenopharyngodon     Cyprinion     Cyprinus     Esox     Gambusia     Garra     Gasterosteus     Glossogobius     Gobio     Glyptothorax     Hemiculter     Hemigrammocapoeta     Heteropneustes     Huso     Hypophthalmichthys     Hyrcanogobius     Ilamnemacheilus     Iranocichla     Iranocypris     Knipowitschia     Kosswigobarbus    Labeo     Lepomis     Leucaspius     Liza     Lota     Luciobarbus     Mastacembelus     Mesogobius     Mesopotamichthys     Metaschistura     Micropterus     Misgurnus     Morone     Mugil     Mullus     Mylopharyngodon     Mystus     Neogobius     Oncorhynchus     Oreochromis     Oryzias     Oxynoemacheilus     Parabramis     Paracobitis     Paraschistura     Pelecus     Perca     Periophthalmus     Petroleuciscus     Pimephales     Platichthys     Ponticola     Proterorhinus     Psetta     Pseudogobio     Pseudorasbora    Pseudoscaphirhynchus     Pungitius     Rhinogobius     Rhodeus     Romanogobio     Rutilus     Sabanejewia     Salmo     Salvelinus     Sander     Scardinius     Schizocypris     Schizopygopsis     Schizothorax     Seminemacheilus     Silurus     Squalius     Stenodus     Syngnathus     Tenualosa     Tilapia     Tinca     Tor     Triplophysa     Turcinoemacheilus     Vimba     Xiphophorus    

Alphabetical Links to Families

Acipenseridae     Adrianichthyidae     Anguillidae     Atherinidae     Bagridae     Carcharhinidae     Centrarchidae     Chanidae     Channidae     Cichlidae     Clupeidae     Cobitidae     Cyprinidae     Cyprinodontidae     Esocidae     Gasterosteidae     Gobiidae     Heteropneustidae     Lotidae     Mastacembelidae     Mugilidae     Mullidae     Nemacheilidae     Percichthyidae    Percidae     Petromyzontidae     Pleuronectidae     Poeciliidae     Salmonidae     Scophthalmidae     Siluridae     Sisoridae     Sparidae     Syngnathidae

Marine species entering fresh water from the Persian Gulf and Sea of Oman can be accessed through a Marine List in the Checklists of the Introduction.
 


Google                         
WWW
www.briancoad.com

Some sections of this work are incomplete but have been posted as is. They will be updated and queries resolved as time permits. Apart from files not yet complete, distribution maps, some figures and habitat photographs are the main items to be added. A question mark may appear in the text appended to items that need to be checked by me. Numerous queries have led me to post incomplete material.

This work has been carried out over a period of 40 years, starting in 1971. I arrived in Iran in January 1976 and, in that year, 7 articles were published strictly on Iranian fishes (3 on parasites, 1 on pesticides, 1 on fisheries, 1 describing the blind white fish and 1 a summary of the latter; 2 were in Farsi). A generation later in 2006, over 160 articles on Iranian fishes appeared, along with hundreds of relevant works from neighbouring countries, works on the aquatic environment in Iran and works on taxonomy and systematics relevant to Iran. The study of fishes is now a very active field within Iran and the Middle East and much of the newer literature is easily available on-line (see Bibliography). Accordingly, 2010 is the last year that this work was updated although some systematic and taxonomic studies may still be incorporated.
 


Acknowledgements

More names to add?

A wide range of people in Iran, Canada and elsewhere have assisted me in this work over more than 40 years. Inevitably, I will have forgotten some names, which I regret. Some people I never met formally, an example being the gentleman nattily dressed in suit by a stream near Kazerun who jumped fully-clothed into the water to help me catch fish. Numerous other Iranians have assisted my studies and this website is dedicated to them.

The staff at the Department of Biology, Shiraz (then Pahlavi) University helped me in numerous ways to collect fishes during a three-year tenure as an Associate Professor. Dr. Bahman Kholdebarin was Chairman of the Department for much of my time in Iran and it is only through his support that I was able to make the collections that enabled this work to be done. The Research Council of Pahlavi University funded field trips and is gratefully acknowledged for this support. Collections were made with the help of drivers and assistants and their efforts over long periods in the field are gratefully acknowledged. They include H. Assadi, M. H. Jaferi, Sh. Mansoorabadi, A. Shirazi, A. Tofangdar and N. Yaghar. Various other people assisted too and are mentioned below under the Pahlavi University name.

Studies on Iranian fishes since my residence in Iran have been supported by grants from the Canadian Museum of Nature, Ottawa (CMN, fish collection acronym CMNFI), by assistance from staff there including Noel Alfonso, Jadwiga Frank, C. G. Gruchy, Sylvie Laframboise, Alison Murray, Claude Renaud and Michèle Steigerwald, and by a wide range of students and volunteers. The staff in the CMN library searched out all the numerous and varied papers on fishes in Iran and neighbouring countries without which this synthesis would not be possible. One paper took six years to locate and arrived in the form of a microfilm from the Soviet Union. I am particularly indebted to Victor Adomaitis who kindly volunteered for the unrewarding task of scanning hundreds of images and converting them to thumbnails and usable files. Mollie MacCormac carried on this task, making a wide variety of images available for the website.

In particular, I should like to acknowledge the support and encouragement of the late Dr. D. E. McAllister, Curator of Fishes, CMN over many years, in terms of training and education, both formal and informal, of financial and moral support, and in practical terms in the ways and means of collecting, cataloguing, identifying, and studying fishes, and of getting things done.

Co-authors are evident in the Bibliography and their added expertise made several studies possible (authors marked by a * are first authors on some papers with me). These are in temporal order F. Krupp, H. F. Alkahem, R. J. Behnke, G. B. Delmastro, M. Kuru, M. Sarieyyüpoglu, F. Papahn, L. A. J..Al-Hassan, A. Abdoli, N. Najafpour, *D. González-Solís, *F. Moravec, Y. Keivany, J. Holcík, J. W. Atz, M. Naderi, *H. Ahnelt, *N. A. Hussein, *T. S. Ali, K. E. Limburg, B. Ya. Vilenkin, N. Keyzer-de Ville, *H. R. Esmaeili, T. T. Nalbant, and *T. Hrbek.

Various people and their organisations are mentioned below separately for their particular assistance; these are in alphabetical order.

Dr. Asghar Abdoli collected numerous specimens including exotics and allowed me to incorporate these discoveries in several papers.

Dr. P. Bănărescu, Institutul de Biologie, Bucureşti has communicated much information in detailed letters on fishes in the Middle East as well as loaning and exchanging specimens, for all of which his assistance is acknowledged.

Dr. R. J. Behnke, Colorado State University, Fort Collins is gratefully acknowledged for his extensive loans of, and access to, collections he and associates made. These are listed more fully in the Materials and Methods.

Prof. Dr. P. G. Bianco, University of Naples, allowed me free access to materials, including types, in his possession at the University of Naples and his hospitality is acknowledged.

Dr. N. Bogutskaya and Dr. A. Naseka, Laboratory of Ichthyology, Zoological Institute, Academy of Sciences, St. Petersburg are thanked especially for their hospitality, access to collections, data analyses and interpretations on Iranian fishes.

Dr. C. E. Bond, Department of Fisheries and Wildlife, Oregon State University, Corvallis allowed extensive loans of fishes from Iran under his care and these materials are listed in the Material and Methods (see Contents).

Staff at the Fish Section, British Museum (Natural History) (now the Natural History Museum) have loaned materials and hosted visits on numerous occasions; their help has been much appreciated for the extensive collections are a required study to understand the Iranian fauna. They include Dr. K. Banister, B. Brewster, P. Campbell, O. Crimmen, S. Davidson, Dr. P. H. Greenwood, A.-M. Hodges, G. Howes, J. Maclaine, Dr. N. Merrett, Dr. D. Siebert, Dr. E. Trewavas, A. Wheeler and Dr. P. J. P. Whitehead.

Dr. T. Hrbek, Washington University School of Medicine, St. Louis is acknowledged for his complementary studies on tooth-carps using molecular techniques.

Dr. M. Kasparek and Prof. Dr. R. Kinzelbach kindly appointed me to the Advisory Board of the journal Zoology in the Middle East which has given me an interesting and valuable overview of studies in that region.

Dr. Yazdan Keivany translated abstracts of his manuscript reports and first posted my bibliography of Iranian freshwater fishes on the internet - a stimulus to this work! He is continuing collaborative efforts aimed at improving knowledge on the ichthyofauna of Iran.

Dr. Bahram Kiabi, Gorgan University of Agricultural Sciences and Natural Resources is thanked for various items of information on fishes, translations and gifts of Farsi articles and many interesting fish specimens. His efforts at facilitating collegiality and his students have formed the core of modern university researchers on the fishes of Iran.

Dr. F. Krupp, Johannes Gutenburg-Universität Mainz and Forschungsintitut Senckenberg (NaturMuseum Senckenberg), Frankfurt am Main contributed a wide variety of information on Middle Eastern fishes, sent me copies of his theses and in his letters provided many stimulating points of discussion which helped me clarify my views on the fishes. His published works are a model for students on fishes in that region. He, with Prof. Dr. Kinzelbach, kindly invited me to the Symposium on the Fauna and Zoogeography of the Middle East in Mainz, 1985.

Nasser Najafpour, Iranian Fisheries Research and Training Organization, Jahad-e Sazandegi,Ahvaz was instrumental in arranging visits to Iran and associated field trips. His enthusiastic cooperation in the field resulted in many interesting new specimens and his studies on distributions of fishes in Khuzestan have been very important for this web site. The team at Ahvaz is acknowledged below individually and in teaching me Farsi names of fishes. J. Gh. Marammazi was head of that team and his hospitality and efforts to bring me to Iran are gratefully acknowledged.

Dr. T. T. Nalbant, National Museum of Natural History "Grigore Antipa", Bucharest, is currently studying loaches I collected in Iran.

Staff at the National Museum of Natural History (Smithsonian Institution), Washington arranged loans of specimens and allowed access to the collections during several visits. They include K. A. Bruwelheide, Dr. B. B. Collette, S. Jewett, S. Karnella and Dr. J. T. Williams.

Staff at the Fischsammlung, Naturhistorisches Museum Vienna have also loaned materials and hosted visits and their assistance has been essential to studies on Iranian fishes based on the collections of J. J. Heckel. They include Dr. H. Ahnelt, Dr. E. Mikschi, Dr. B. Herzig and Dr. R. Hacker.

Dr. J. G. Nielsen and Dr. P. R. Möller, Zoological Museum, University of Copenhagen facilitated access to collections despite the "orkan".

Dr. P. Bartsch and Mrs. C. Lamour, Museum für Naturkunde, Humboldt-Universität zu Berlin for access to collections.

M. Rabaniha and F. Owfi, Persian Gulf Fisheries Research Centre, Bushehr and Iranian Fisheries Research and Training Organization, Jahad-e Sazandegi, Tehran, kindly copied the MMTT catalogue for me and showed me various specimens of fishes from their work in Bushehr Province and southern Iran.

Dr. Jalal Valiallahi provided stimulating discussions on the limits and the content of the genus Barbus sensu lato in Iran while working at the CMN as well as a variety of photographs of these sometimes immense fish. Prof. Dr. H. Wilkens, Zoologisches Institut und Zoologisches Museum der Universität Hamburg kindly loaned materials and facilitated two visits to the museum to examine materials.

Various people collected material for me or made gifts of material, sent specimens for identification, identified material, allowed access to collections under their care, made loans of material, provided other useful data and general information, and exchanged ideas. These are listed below in alphabetical order with their affiliations at the time of their contribution (sometimes only email addresses were known; and apologies if any titles are missing):-

K. Abbasi, Gilan Fisheries Research Centre, Bandar Anzali, H. A. Abdolhay, Tehran, I. M. Abd, Nature Iraq, Baghdad, Iraq, A. Abdoli, Fisheries Research Centre, Sari and Gorgan University of Agricultural Sciences and Natural Resources, S. Abdolmalaki, Gilan Fisheries Research Centre, Bandar Anzali, S. M. A. Abdullah, Iraq, Dr. T. Abe, University Museum, University of Tokyo, Dr. M. Abedi, Savadkooh University, H. Abyot, Iranian Fisheries Research and Training Organization, Jahad-e Sazandegi, Ahvaz, T. K. Aday, Iraq, Dr. A. Adhami, Muze-ye Melli-ye Tarikh-e Tabi'i, Tehran, A. Afzali, Bandar Abbas, Fikret Ahsenböre, Turkey, Dr. A. Akbary Pasand, University of Zabol, Zabol, A. Alamdari, Organization of the Environment, Shiraz, A. A. Al-Attar, Basrah University, A. W. Al-Hakim, University of Nottingham, L. A. J. Al-Hassan, School of Biological Sciences, University of Auckland, S. A. S. Al Hatimy, Oman Natural History Museum, Muscat, W. Al-Baharna, Directorate of Fisheries, Bahrein, Dr. N. M. Ali, Biological Research Centre, University of Baghdad, Dr. T. S. Ali, University of Basrah, S. Alinejad, Offshore Fisheries Research Centre, Chah Bahar, Iranian Fisheries Research and Training Organization, Jahad-e Sazandegi, H. R. Alizadeh, Iranian Fisheries Research and Training Organization, Jahad-e Sazandegi, Tehran, A. R. Al-Jafery, Department of Hydrobiology, Baghdad, Dr. H. Alkahem, King Saud University, Riyadh, M. A. Al-Mukhtar, Fisheries Research Centre, Ahvaz, Dr. A. J. Al-Rudainy, University of Baghdad, Iraq, Dr. A. Al-Shamma'a, Ministry of Science and Technology, Iraq, Nisreen Alwan, Forschungsinstitut Senckenberg, Germany, Prof. O. A. Amin, Arizona State University, Tempe, Dr. F. Andreone, Museo Regionale di Scienze Naturali, Torino, Dr. R. Arai, National Science Museum, Tokyo, G. Arbocco, Museo Civico di Storia Naturale "Giacomo Doria", Genova, Dr. J. D. Archibald, Yale University, Connecticut, Dr. N. B. Armantrout, Portland, Oregon, Dr. G. Arratia, University of Kansas, Lawrence, S. Asadollah, Isfahan University of Technology, A. Ashraf, Encyclopædia Iranica, Columbia University, New York, Dr. J. W. Atz, Department of Herpetology and Ichthyology, American Museum of Natural History, New York, Prof. S. Balik, Ege University, Izmir, Prof. E. Balletto, Istituto di Zoologia, Genova, G. A. C. Balma, Museo Civico di Storia Naturale, Carmagnola, Dr. K. Banister, Fish Section, British Museum (Natural History), London, A. J. Bardhun, Shiraz, D. M. Bartley, Food and Agriculture Organization, Rome, Dr. V. V. Barsukov, Zoological Institute, Academy of Sciences, Leningrad, M. L. Bauchot, Laboratoire d'Ichtyologie générale et appliquée, Muséum National d'Histoire Naturelle, Paris, R. Beck, COFAD GmbH, Tutzing, Dr. W. C. Beckman, Opelousas, Louisiana, Dr. A. Ben-Tuvia, Hebrew University of Jerusalem, Dr. M. Berberian, Uinversity of Cambridge, Dr. P. Berrebi, Université Montpellier, Dr. A. D. Berrie, Freshwater Biological Association, Wareham, Dr. E. Bertelsen, Zoologisk Museum, Copenhagen, Prof. Dr. P. G. Bianco, Universita degli Studi di l'Aquila, K. L. Bist, Government Postgraduate College, Gopeshwar, J. Bohlen, Academy of Sciences, Libechov, Dr. J. E. Böhlke, Academy of Natural Sciences, Philadelphia, Dr. A. H. Bornbusch, Duke University, Durham, Dr. J. Briggs, King Faisal university, Dammam, Dr. K. E. Carpenter, Food and Agriculture Organization, Rome, L. A. Cloutier, Department of the Environment, Tehran, Dr. D. Coffey, Pahlavi University, Shiraz, Dr. M. J. Collares-Pereira, Museu Bocage, Lisbon, Dr. J. T. Collins, Museum of Natural History, University of Kansas, Lawrence, Dr. L. J. V. Compagno, J. L. B. Smith Institute of Ichthyology, Grahamstown, Dr. B. B. Collette, National Museum of Natural History, Washington, G. H. Copp, Centre for Environment, Fisheries and Aquaculture Science, Lowestoft, Dr. L. Cornwallis, Oxford, A. S. Creighton, Division of Fishes, Museum of Zoology, University of Michigan, Ann Arbor, Dr. E. J. Crossman, Department of Ichthyology and Herpetology, Royal Ontario Museum, Toronto, E. L. Daniel, Encyclopædia Iranica, Columbia University, New York, F. Darvishi, Mazandaran, S. Deeb, American University of Lebanon, Beirut, S. Dehqan-Mediseh, Iranian Fisheries Research and Training Organization, Jahad-e Sazandegi, Ahvaz, Dr. G. B. Delmastro, Museo Civico di Storia Naturale, Carmagnola, M. Desoutter, Laboratoire d'Ichtyologie générale et appliquée, Museum National d'Histoire Naturelle, Paris, Dr. M. M. Dick, Museum of Comparative Zoology, Harvard University, Cambridge, P. Dickinson, National Zoological Garden, Al Ain, Abu Dhabi, W. A. Dill, Davis, California, J. Dominique, Freshwater and River Ecology Reserach Unit, Villeurbane, Dr. P. Dugan, Penang, Malayasia, M. Doroudi, Iranian Fisheries Research and Training Organization, Jahad-e Sazandegi, Bandar-e Lengeh, Dr. J. D. Durand, ESA CNRS, Villeurbane, Dr. G. Ekingen, Veteriner Fakultesi, Elazig, O.Elter, Museo ed Istituto di Zoologia Sistematico, Universita di Torino, Dr. B. Elvira, Ministerio de Agricultura y Pesca, Madrid, G. El Zein, Université Libanaise, Ksara, Dr. F. Erk'akan, Hacettepe University, Ankara, Dr. W. N. Eschmeyer, Department of Ichthyology, California Academy of Sciences, San Francisco, Gh. Eskandary, Fisheries Research Centre, Jahad-e Sazandegi, Ahvaz, Dr. H. R. Esmaeili, Shiraz University, D. Evans, IUCN, Cambridge, K. Evans, Pahlavi University, Shiraz, K. Fakhro, Directorate of Fisheries, Bahrein, R. Fatemi, Tehran, Dr. A. M. Fazel, Natural Resources Faculty, Tehran University, Karaj and Natural History Museum, Department of the Environment, Tehran, , H. Fazly, Fereydun Kenar, Mazandaran, R. F. Field, Muscat, Dr. E. Firouz, Tehran, Dr. W. Fischer, Food and Agriculture Organization, Rome, J. Fitzpatrick, Food and Agriculture Organization, Rome, Dr. R. Fricke, Staatliches Museum für Naturkunde in Stuttgart, P. A. M. Gaemers, Rijksmuseum van Geologie en Mineralogie, Leiden, M. D. Gallagher, Oman Natural History Museum, Muscat, M. Geerts, Swalmen, The Netherlands, Prof. Dr. R. Geldiay, Ege University, Izmir, Dr. C. George, Union College, Schenectady, Dr. H. Ghadirnejad, Iranian Fisheries Research and Training Organization, Jahad-e Sazandegi, Tehran, A. Ghamoosi, Shahid Beheshti University, Tehran, S. M. Ghasempouri, Tarbiat Modares University, Noor, Dr. D. I. Gibson, British Museum (Natural History), London, D. Golani, Zoological Museum, Hebrew University of Jerusalem, Dr. M. Goren, Tel Aviv University, S. Gorgin, Shiraz, Dr. B. Groombridge, UNEP World Conservation Monitoring Centre, Cambridge, Dr. S. H. Gruber, University of Miami, J. M. Gunn, University of Ottawa, R. Haas, California State University, Fresno, M. Hafezieh, Research Centre for Natural Resources and Animal Husbandry, Jahad-e Sazandegi, Shiraz, Dr. J. Halpern, Pahlavi University, Shiraz, Dr. K. E. Hartel, Museum of Comparative Zoology, Harvard University, Cambridge, S. S. Hasan, University of Basrah, Dr. M. R. Hassannia, Jahad-e Sazandegi, Tehran, M. R. Hemami, Isfahan University of Technology, D. M. Herdson, The Laboratory, Plymouth, E. Holm, Department of Ichthyology and Herpetology, Royal Ontario Museum, Toronto, Dr. R. A. Hinrichsen, Shad Foundation, Seattle, A.-M. Hodges, Fish Section, British Museum (Natural History), London, M. L. Holloway, Fish Section, British Museum (Natural History), London, L. Honarmond, University of Tehran, Dr. J. Holčík, Institute of Zoology, Slovak Academy of Sciences, Bratislava, Drs. F. and Sh. Hosseinie, Shiraz University, Dr. C. Hubbs, University of Texas, Austin, Dr. J. Huber, Muséum National d'Histoire Naturelle, Paris, J. Hull, University Museum, Oxford University, Dr. N. A. Hussain, Marine Science Centre, University of Basrah, Ch. Izadi, Research Centre for Natural Resources and Animal Husbandry, Jahad-e Sazandegi, Shiraz, Gh. Izadpanahi, Dr. B. Jalali, ABZIGOSTAR, Tehran, Dr. S. Jahromi, Pahlavi University, Shiraz, Dr. S. Jamili, Iranian Fisheries Research and Training Organization, Jahad-e Sazandegi, Tehran, Gh. A. Jasimi, Iranian Fisheries Research and Training Organization, Jahad-e Sazandegi, Ahvaz, Dr. M. N. Javed, Government College, Lahore, Dr. K. C. Jayaram, Zoological Survey of India, Calcutta, K. Jazebizadeh, Iranian Fisheries Research and Training Organization, Ahvaz, Dr. J. B. Jensen, Pahlavi University, Shiraz, Dr. R. K. Johnson, Field Museum of Natural History, Chicago, W. J. Jones, Al Ain, U.A.E., Dr. H. G. Kami, University of Tehran, J. M. Kapetsky, Food and Agriculture Organization, Rome, Dr. M. H. Karim Koshteh, University of Guelph, Dr. M. Kasparek, Kasparek Verlag, Heidelberg, Dr. E. J. Keall, Royal Ontario Museum, Toronto, Dr. A. Keyvanfar, Centre national de Transfusion sanguine-Institut, Paris, Dr. G. Khalaf, Lebanese University, Mansourieh-el-Metn, Dr. N. R. Khamees, University of Basrah, S. Khera, Punjab University, Chandigarh, A. Khodady, Shahid Chamran University, Ahvaz, Prof. Dr. R. Kinzelbach, Zoologisches Institut, Darmstadt, Dr. W. Klausewitz, Forschungsintitut Senckenberg, Frankfurt, Dr. W. L. Klawe, Inter-American Tropical Tuna Commission, Scripps Institution of Oceanography, La Jolla, Dr. M. Kottelat, Zoologsiches Staatsammlung, Munich, Dr. A. Kownacki, Laboratory of Water Biology, Polish Academy of Sciences, Krakow, Dr. S. O. Kullander, Swedish Museum of Natural History, Stockholm, Dr. K. Kuronuma, Tokyo University of Fisheries, Dr. M. Kuru, Hacettepe University, Ankara, P. Lamothe, Hydro Québec, Montréal, Dr. K. J. Lazara, US Merchant Marine Academy, Kings Point, New York, A. Lealmonfared, Shahid Beheshti University, Tehran, Dr. R. E. Lee, Pahlavi University, Shiraz, Dr. K. E. Limburg, State University of New York, Syracuse, Dr. R. Littman, University of Hawaii, Honolulu, Prof. Dr. H. Loffler, Vienna, R. Lolea, Gorgan University, J. Long, Department of Fisheries and Wildlife, Oregon State University, Corvallis, O. Lucanus, Montreal, Dr.Mabee, Department of Zoology, Duke University, Durham, A. A. Mahdi, University of Basrah, A. Mahjoor Azad, Shahid Beheshti University, Tehran, Dr. P. S. Maitland, Institute of Terrestrial Ecology, Edinburgh, Dr. H. Malicky, Biologische Station Lunz, L. Maltz, Tel Aviv University, J. Mansoori, Iranian Fisheries Research and Training Organization, Jahad-e Sazandegi, Ahvaz, J. Gh. Marammazi, Iranian Fisheries Research and Training Organization, Jahad-e Sazandegi, Ahvaz, R. Martino, American Killifish Association, Dr. M. Masoumian, Iranian Fisheries Research and Training Organization, Jahad-e Sazandegi, Tehran, Dr. A. Matinfar, Iranian Fisheries Research and Training Organization, Jahad-e Sazandegi, Tehran, Y. Mayahi, Iranian Fisheries Research and Training Organization, Jahad-e Sazandegi, Ahvaz, Dr. R. L. Mayden, Department of Biological Sciences, University of Alabama, Tuscaloosa, J. J. McAniff, National Underwater Accident Center, University of Rhode Island, Kingston, M. McDavitt, Alexandria, Virginia, S. Mickleburgh, Fauna and Flora Preservation Society, London, H.Meeus, Belgische Killifish Vereniging, Wommelgen, R. Mehrani, Lorestan Research Centre of Natural Resources and Animal Science, Khorramabad, Dr. A. G. K. Menon, Zoological Survey of India, Calcutta, Dr. S. N. Messieh, UNDP, Abu Dhabi, Dr. F. T. Mhaisen, University of Baghdad, Dr. A. Miller, Royal Botanic Garden, Edinburgh, I. D. Miller, United States-Saudi Arabian Joint Commission, New York, Dr. P. Miller, University of Bristol, Dr. R. R. Miller, Division of Fishes, Museum of Zoology, University of Michigan, Ann Arbor, Dr. A. A. Mirhosseyni, National Natural History Museum, Baghdad, Dr. M. R. Mirza, Lahore, A. Mobaraki, Department of the Environment, Tehran, M. R. Mohaghegh, Tehran, M. Mohammadi, Gorgan Agricultural and Natural Resources University, Dr. S. Moini, Department of the Environment, Tehran, Dr. B. Mokhayer, University of Tehran, Dr. K. Molnár, Veterinary Medical Research Institute, Hungarian Academy of Sciences, Budapest, Dr. F. Moravec, Institute of Parasitology, Czechoslovak Academy of Sciences, Prague, E. Morin, SOGREAH, Echirolles, Dr. E. O. Murdy, Bureau of Oceans and International Environmental and Scientific Affairs, Washington, Dr. G. S. Myers, Scotts Valley, California, M. Naderi, Mazandaran Fishery Research Centre, Sari, S. Naem, Faculty of Veterinary Medicine, Urmia University, A. Nasrollahzadeh, Gilan, Prof. Dr. C. M. Naumann, Universität Bielefeld, Dr. S. Nazeeri, Iranian Fisheries Research and Training Organization, Jahad-e Sazandegi, Khorramabad, R. B. Nehring, Department of the Environment, Tehran, N. Niameymandi, Persian Gulf Fisheries Research Centre, Bushehr, Dr. H. Nijssen, Instituut voor Taxonomisch Zoölogie, Zoölogisch Museum, Universiteit van Amsterdam, M. Nikpaey, Iranian Fisheries Research and Training Organization, Jahad-e Sazandegi, Ahvaz, N. Nouri, Iranian Fisheries Research and Training Organization, Jahad-e Sazandegi, Tehran, Dr. O. Oliva, Charles University, Prague, Dr. H.-J. Paepke, Museum für Naturkunde der Humboldt-Universität, Berlin, Dr. A. Paltrinieri, World Health Organization, Muscat, F. Papahn, Shahid Chamran University, Ahvaz, Dr. L. R. Parenti, National Museum of Natural History, Washington, J. Parkinson, Edmonton, A. Parsamanesh, Iranian Fisheries Research and Training Organization, Ahvaz, D. Peck, IUCN, Gland, T. Petr, Food and Agriculture Organization, Rome, H. Piri Zirkohy, Gilan Fisheries Research Centre, Bandar Anzali, Dr. E. P. Pister, Desert Fishes Council, Bishop, California, S. P. Platania, Colorado State University, Fort Collins, T. Plosch, Ganderkesee, L. Podshadley, Department of Ichthyology, California Academy of Sciences, San Francisco, Dr. M. Pourgholam, Iranian Fisheries Research and Training Organization, Jahad-e Sazandegi, Sari, M. Price, Division of Fishes, Museum of Zoology, University of Michigan, Ann Arbor, Dr. G. S. Proudlove, Department of Environmental Biology, University of Manchester, T. A. Qureshi, Technical Institute for Agriculture, Amara, M. Rabbaniha, Persian Gulf Fisheries Research Centre, Bushehr, Dr. H. Rahimian, University of Tehran, Dr. M. Ramin, Iranian Fisheries Research and Training Organization, Jahad-e Sazandegi, Tehran, F. M. Razi, Nature and Wildlife Museum, Tehran, Dr. W. J. Rainboth, University of California, Los Angeles, R. W. Redding, Museum of Zoology, University of Michigan, Ann Arbor, D. Rees, BBC, London, Dr. K. Relyea, Kuwait Institute for Scientific Research, H. Rezai, Tehran, Dr. S. Rezvani Gilkolaei, Iranian Fisheries Research and Training Organization, Jahad-e Sazandegi, Tehran, S. Richards, Murray, Utah, Dr. T. R. Roberts, Kasetsart University, Bangkok, A. Roohi, Sabzevar Teaching and Training University, Sabzevar, Khorasan, Dr. I. Rostami, Shahid Chamran University, Ahvaz, B. Saadallah, Iraq Natural History Museum, Baghdad, M. A. G. Saadati, Department of the Environment, Mashhad, H. Saadoni, Iranian Fisheries Research and Training Organization, Jahad-e Sazandegi, Ahvaz, H. R. A. Sabet, Iranian Fisheries Research and Training Organization, Tehran, A. R. Saeed, University of Kerman, E. Saderigh-Nejad Massouleh, Iranian Fisheries Research and Training Organization, Jahad-e Sazandegi, Khorramabad, H. Safikhani, Iranian Fisheries Research and Training Organization, Jahad-e Sazandegi, Ahvaz, Dr. A. Salnikov, Institute of Zoology, Academy of Sciences, Ashkhabad, Dr. A. Samaie, Muse-ye Melli-ye Tarikh-e Tabi'i, Tehran, B. Sanford, Montrose, Colorado and Port Ludlow, Washington, Dr. A. Sanyal, Zoological Survey of India, Calcutta, Dr. M. Sarieyyüpoglu, Firat Üniversitesi, Elazig, Dr. A. Savari, Faculty of Oceanography, Shahid Chamran University, Ahvaz, M. Sayfali, Shahid Beheshti University, Tehran, D. A. Scott, Dursley, Gloucestershire, Dr. D. E. Sergeant, Arctic Biological Station, Ste-Anne de Bellevue, Quebec, Gh. Shakhiba, Iranian Fisheries Research and Training Organization, Ahvaz, A. J. Shams, Directorate of Fisheries, Bahrein, Dr. I. Sharifpour, Iranian Fisheries Research and Training Organization, Ahvaz, J. W. Sherman, Academy of Natural Sciences, Philadelphia, Dr. A. Shiralipour, Pahlavi University, Shiraz, Dr. I. Q. Siddiqui, King Faisal University, Al Hasa, Dr. P. Skelton, Fish Section, British Museum (Natural History), London, Dr. G. R. Smith, Museum of Zoology, University of Michigan, Ann Arbor, Dr. W. F. Smith-Vaniz, Academy of Sciences, Philadelphia, M. Soleymani, Green Front of Iran, Tehran, N. Statman, Dr. A. N. Svetovidov, Zoological Institute, Academy of Sciences, Leningrad, Dr. C. C. Swift, Natural History Museum of Los Angeles County, Dr. F. Terofal, Zoologische Sammlung des Bayreischen Staates, Munich, M. V. Tofighi, Iranian Fisheries Research and Training Organization, Jahad-e Sazandegi, Tehran, A. Torfi, Iranian Fisheries Research and Training Organization, Jahad-e Sazandegi, Ahvaz, Dr. W. Torke, Institut fur Urgeschichte, Tübingen, Dr. E. Tortonese, Museo Civico di Storia Naturale, Genova, Dr. R. A. Travers, Fish Section, British Museum (Natural History), London, R. G. Tuck, Muze-ye Melli-ye Tarikh-e Tabi'i, Tehran, Dr. H. Türkmen, Istanbul Üniversitesi, Dr. E. Unlu, University of Dicle, Diyarbakir, Dr. I. Unsal, Istanbul Üniversitesi, T. Valinasab, Fisheries Research and Training Organization, Jahad-e Sazandegi, Tehran, Dr. J. Valiallahi, Tarbiat-e Modarres, Noor, >W. van Neer, Royal Museum of Central Africa, Tervuren, Prof. Dr. R. Victor, Sultan Qaboos University, Muscat, Prof. Dr. W. Villwock, Zoologisches Institut und Zoologisches Museum, Hamburg, Dr. V. D. Vladykov, University of Ottawa, A. Vosughi, Iranian Fisheries Research and Training Organization, Jahad-e Sazandegi, Tehran, B. Waaland, Pahlavi University, Shiraz, P. Walczak, Department of the Environment, Tehran, Dr. B. A. Whitton, University of Durham, Dr. R. Winterbottom, Department of Ichthyology and Herpetology, Royal Ontario Museum, Toronto, Dr. G. H. Wossughi, University of Tehran, Dr. T. C. Young, Royal Ontario Museum, Toronto, M. Zapater, Zaragoza, A. R. Zeanaie, Payam-e Noor University, Bandar Abbas.

Individual Iranians, too numerous to mention here, kindly enunciated carefully and repeatedly Farsi fish names for my cloth ear. However it would be remiss not to mention staff at the Iranian Fisheries Research and Training Organization, Ahvaz including N. Najafpour, Gh. Marammazi, Gh. Eskandari, and M. A. Al-Mukhtar, as well as E. Firouz, Tehran, B. Kiabi and A. Abdoli, Gorgan Agricultural and Natural Resources University, and Y. Keivany, University of Alberta, Edmonton.

And finally I must thank my wife Sylvie and son Nicholas for supporting me in my obsession with fishes from Iran and Nick for constructing the index page for this website and linking it to the internet.
 


Purpose

This work is meant to provide a guide to the freshwater fishes of Iran. There are no modern keys to this fauna, some available books are incomplete or cursory treatments or outdated, and the detailed and diverse scientific literature is widely scattered in time, languages and journals. Iran lies at a region of major zoogeographical interchange and has a diverse and interesting ichthyofauna about which comparatively little is known. An accurate identification is a pre-requisite for further scientific studies and this website aims to serve that purpose and to be an introductory guide to the fishes. The guide is aimed at a mixed audience, including scientists familiar with ichthyology to whom some introductory sections of this work will be superfluous, and those whose knowledge of fishes is embryonic or who may have limited access to literature sources.

This work has been carried out over a period of 40 years from my first studies on Iranian fishes in 1971 at the University of Ottawa on collections made by V. D. Vladykov along the Caspian coast, continuing during a three-year residence in Iran from January 1976. In that year, 7 articles were published strictly on Iranian fishes (3 on parasites, 1 on pesticides, 1 on fisheries, 1 describing the blind white fish and 1 a summary of the latter; 2 were in Farsi). In 2006, 160 articles on Iranian fishes appeared, along with many relevant works from neighbouring countries, works on the aquatic environment in Iran and works on taxonomy and systematics relevant to Iran. The study of fishes is now a very active field within Iran and the Middle East. Accordingly, 2010 is the last year that this work is updated although some systematic and taxonomic studies may still be incorporated.

Literature on fishes of Iran can be found in Zoological Record (Pisces) and at the Scientific Information Database (or SID at http://www.sid.ir/En/Index.asp)which has lists of publications in Iranian journals and abstracts, both in English, as well as in Farsi.

The main Introduction contains several explanatory sections. These sections include detailed methods of counting and measuring characters, an explanation of scientific names of fishes, details of fish structure so that keys can be readily understood, ways of capturing and preserving fishes and recording the all-important collection data, and how to identify fishes. This introductory part also includes a brief review of the history of research on Iranian fishes and descriptions of the environment including geography, climate, habitats, environmental change and drainage basins.

The bulk of the text is the Species Accounts which serve to identify, describe and map the distribution of each species. Families of fishes follow Nelson (2006) with genera and species arranged alphabetically within each family. Each Species Account is comprised of the following parts: the scientific name, common names, sections on systematics, key characters, morphology, sexual dimorphism, colour, size, distribution, zoogeography, habitat, age and growth, food, reproduction, parasites and predators, economic importance, conservation, further work, sources, and an illustration and a distribution map.

The biological information may be cursory. Many species are poorly known and their biology has not been studied, especially within Iran. Some information is available for species shared with Turkey and Iraq and I have tried to incorporate this literature as being less well known or accessible. Many Caspian Sea basin species are shared with Europe and the former U.S.S.R., are comparatively well-known and have an extensive literature, often summarised in books, bibliographies and synopses. It is not known in many cases if their biology in Iran is similar. Iranian populations are often referred to distinct subspecies and occur at the southern limit of the species range. Only a brief, summary account of their biology is therefore given from synoptic literature sources. Biological information generally is a brief summary of literature and readers should consult the original papers for more details.

Some anecdotal biological information is added from my field collections where spawning individuals were noted or gut contents examined superficially. Most fish spawn in the spring. Feeding habits can often be deduced from morphology. Fish with an arched and ventral mouth, horny jaw edge, elongate gut and black peritoneum are feeders on detritus and aufwuchs scraped from rocks. Most fish with a simple, s-shaped gut feed on invertebrates such as crustaceans and aquatic insect larvae. A few fish with molar pharyngeal teeth have a diet of molluscs whose shells are crushed by the heavy teeth. Some fish are piscivorous and have an appropriate jaw shape and streamlined appearance suitable for catching and holding their fish prey. Fish with elongate and numerous fine gill rakers filter phytoplankton or zooplankton from the water column. Very few fish feed on macrophytes (large plants).

Checklists summarise the diversity of the ichthyofauna. Glossaries explain both ichthyological terms for those new to the science and Farsi and geographical terms for those unfamiliar with that language. A Bibliography comprises books and papers referred to in the text and other relevant works, which form a good general basis for the serious student of Iranian freshwater fishes.
 


Materials and Methods

1. Materials

The descriptions in this work are founded on original observations of material and a consideration of the literature. The sources of this material are various museums which house a scattering of Iranian species including in particular the Natural History Museum, London (formerly the British Museum (Natural History)), the Naturhistorisches Museum Wien, and the Zoological Institute, St. Petersburg which are depositories for older type material, but the bulk of the research has been based on four collections. The first of these was made by V. D. Vladykov during 1961 and 1962 when he was an Inland Fisheries Biologist under the Expanded Programme of Technical Assistance of the Food and Agriculture Organization, UN. This material was deposited in the National Museum of Natural Sciences, Ottawa (now the Canadian Museum of Nature) and consists mainly of specimens from the Caspian Sea basin. The second collection was made by employees of the Department of the Environment, Tehran, and N. B. Armantrout and R. J. Behnke. Half this collection was placed in the National Museum of Natural History, Tehran (Muze-ye Melli-ye Tarikh-e Tabi'i) and half was retained by R. J. Behnke and formed the basis of Saadati's (1977) thesis at Colorado State University, Fort Collins. This collection covered the whole of Iran except the Caspian and Sistan basins. Through the courtesy of Dr. Behnke I have been able to examine this material in Fort Collins and make extended loans for study in Ottawa. The Muze-ye Melli-ye Tarikh-e Tabi'i collection is small (examined in 1995; catalogue 2000) and not as diverse as the Fort Collins material. Oregon State University contains a collection of fishes made by W. Kinunen, S. Bullock, R. RaLonde and P. Walczak, who were members of the Peace Corps in Iran (some of this collection was deposited at the Smithsonian Institution, Washington, which helped to fund the collection and transport of specimens). Dr. Carl Bond kindly loaned me much of this material for long periods. This collection was from all parts of Iran. The last collection, comprising the bulk of the material, was made by me from 1976 to 1979 while I was teaching at Pahlavi (now Shiraz) University in Shiraz. This collection is housed in the Canadian Museum of Nature, Ottawa (formerly NMC, now CMNFI), and covers all of Iran except the extreme northeast and northwest. Field trips were funded by the Research Council of Pahlavi University. Subsequently various Iranian colleagues have sent me specimens and these too are incorporated in the present work. Principal among these were materials collected by Asghar Abdoli (then based in Golestan) and Nasser Najafpour and associates of the Iranian Fisheries Research Organisation (IFRO), Ahvaz. These collections together effectively cover all the major drainages of Iran and provide the best foundation yet assembled for a study on this ichthyofauna.

All material stored at the Canadian Museum of Nature, Ottawa was examined in 45% isopropyl alcohol. Preservative was later changed to 70% ethanol. The Canadian Museum of Nature also stores extensive field records including slides, numerous data sheets on most species (counts and measurements including x-ray plates), an extensive literature base including translations from foreign languages, and comparative specimens and literature from other countries in Southwest Asia.

2. Methods

Specimens collected by me were caught by any means that presented themselves. Gear used included seines of various lengths and mesh sizes (much repaired and patched!), gill-nets of various stretch meshes (sometimes used as seines), cast-nets of several diameters (thrown skilfully by others and poorly by me), by hand, and by purchase from small boys and anglers using a variety of techniques (of angling on their part and of persuasion on mine to extract catches from their possession). The object was to sample any water body for all the kinds of habitat found there within the limitations of a hasty schedule and the available equipment. Most habitats were visited for less than one hour, but in the small springs and streams, which comprise the bulk of Iranian fresh waters outside the large rivers and lakes of Khuzestan and Sistan and the deep waters of the Caspian Sea, this was more than adequate to catch a good and varied sample of most species. This was borne out by repeated visits of longer duration to certain localities near Shiraz. Pools and flowing sections were seined, gill-netted or cast-netted. Riffle areas were also attacked in this fashion or seines were used to block off sections of riffle and upstream rocks disturbed by kicking to scare secretive species like loaches into the fixed net. In small streams a dip-net was placed downstream of individual rocks which were kicked over and the net scooped along the stream bed. Cast-nets proved particularly useful in rocky streams which had little open water. Draped over the rocks and only partly in the water, they nevertheless caught large and fast specimens which were unobtainable by seining. The available fishing gear was less effective on large rivers and on the Caspian Sea. Here boats, long gill-nets and trawl gear would have been most useful. The collections are poor in inhabitants of the main current of large rivers and in the deep water species of the Caspian Sea. Larger specimens in major water bodies undoubtedly evaded my nets with ease; some samples of larger individuals were available from other collections and by purchase from commercial fisheries.

Several criteria were used to select specimens for counts and measurements. Where few specimens were available, all were counted and measured. Where several hundred specimens were available selection was by size (usually larger fish; sometimes much smaller fish as well for comparison with adult values), by sex to ensure an adequate representation of males and females, and by locality where geographical variation was examined. Badly damaged or grossly deformed specimens were excluded but there was no (conscious) selection for "ideal" specimens.

Wherever a putative species was collected from more than one drainage basin and material diversity permitted, a comparison was made between the drainage basins. This work is continuing and details of methods and materials are to be seen in published results. Students of Iranian fishes should note that the application of sufficient statistical "weight" will reveal differences between drainage basin samples and this is especially true of a desert and semi-desert country like Iran. Springs and streams may have been colonised by only a few founders. A small population sampled in the lower reaches of a stream may not have had any contact with conspecifics higher up in the stream for many generations. Conversely, several seasons of heavy rain may have afforded recent opportunities for contact and gene exchange. A one-time sample from a stream may therefore give a quite inaccurate picture of the character suite of that population. Whether any of the differences detected have systematic significance requires careful consideration. For example, Balletto and Spano (1977) described 9 subspecies of Garra tibanica in the southwest of the Arabian Peninsula using Principal Components Analysis. This has been termed "statistical overkill" by Alkahem and Behnke (1983). Also Krupp (1983) has observed that samples of Garra rufa from the same locality collected in different years or seasons varied in several characters. Description of subspecies based on limited material requires a great deal of care therefore.

There are various methods of measuring and counting anatomical features of fishes. The ones I have used are outlined below. They are based on Hubbs and Lagler (1958) and Trautman (1981). Some particular characters are outlined in papers by me in the Bibliography.

The method of counting fin rays differs from that in use in North America since unbranched and branched rays are counted separately. A "III,8" count in the European literature would be "9" in the system advocated by Hubbs and Lagler (1958), i.e. the soft ray count is increased by one to convert from the "European" to the "American" system. The bulk of the work on fishes of southwest Asia follows the European system and I have adopted this methodology to facilitate comparisons, although eschewing Roman numerals.

A) Meristic characters

In this book, scale counts, number of gill rakers and of vertebrae are usually expressed as ranges based on literature sources since frequency counts are rarely given. A separate section gives counts on Iranian fish examined by me followed by a frequency in parentheses (..). Fin ray counts often show strong modes, but citing the mode alone would be misleading. Pharyngeal tooth formula is often a modal value from the literature; loss of or incomplete development of major or minor row teeth is not uncommon, so counts may vary quite markedly.

Scale counts and paired fin ray counts were made on the left side of each fish. In some instances, such as a badly deformed fin or where scales on the left were mostly missing, counts were made on the right. These instances were rare and restricted to species with low sample sizes.

Not all meristic characters had equal sample sizes; some material from other museums was not available for x-rays, large series of pharyngeal tooth counts was not often available because removal of arches damages specimens, some specimens were damaged in certain characters, time did not always permit all characters to be counted, some species are well-known and additional data from Iran is clearly a subset of widely gathered data, some species were examined in detail to address systematic problems, and so on.

1) Vertebrae

All vertebrae were counted including the hypural plate as one vertebra. In Cypriniformes and Siluriformes, the four Weberian vertebrae were included in the count. Almost all counts were made from radiographs.

2) Gill rakers

All rakers on the first gill arch were counted. A lower limb count in the literature includes any raker at the angle of the upper and lower limbs. Gill raker counts presented something of a problem when comparing specimens of disparate sizes. The smaller fish often had very small rakers at each end of the arch. These were easily missed or torn off when cleaning a debris-encrusted arch. Removal of arches for a more careful examination may also damage or destroy the finer rakers which are intimately associated with the tissues adjacent to the arches. Alizarin preparations can be of assistance, but the finer rakers may have no bony content and thereby be omitted. Counts of juvenile fish may therefore give lower values than counts for larger fish, whether this be due to an increase in gill raker number with age or because rakers are more easy to count in larger fish. This kind of variation is only critical where this character is being used in species identification or in analyses meant to define and relate species.

3) Pharyngeal teeth

The teeth of the modified fifth gill arch in Cyprinidae were counted in each row and given as a formula from left to right. A count of 2,5-4,2 consists of two teeth in both the outer left and outer right rows, five teeth in the inner left row and four teeth in the inner right row. Pharyngeal teeth rows in Iranian cyprinids varied from one to three on each side. In certain cases, it was evident from the presence of a socket that a tooth had been lost. The count then included that tooth.

4) Fin rays

a) Dorsal and anal fins

Fin ray counts were divided into two types. One count is of spines or hardened soft rays or any unbranched, unpaired unsegmented rays and this is usually given in Roman numerals in the literature. In deference to some Iranian unfamiliarity with Roman numerals, the spine count is given in Arabic numerals in this text. Spine count included rudimentary rays which, at the anterior dorsal and anal fins, may be obscured by flesh or scales requiring some probing or dissection. Radiographs were often useful to confirm counts made under a microscope. The second count is of soft rays and is also indicated by Arabic numerals. These rays are usually branched, flexible, segmented and laterally paired. The last two unbranched rays often arise from a single internal base and were then counted as one. This is generally the case in Cyprinidae. The branched ray count is the most diagnostic and variable in such fishes. Some families contain species with more than one dorsal fin. The first dorsal fin may be composed of spines and the second dorsal fin of spines and soft rays. In such species the count is given separately for each fin.

b) Caudal fin

The branched caudal fin rays only were counted. Dorsal and ventral to these central rays are a series of unbranched rays which become progressively smaller and may be obscured by flesh and scales where the caudal fin attaches to the caudal peduncle. Counts in other works often comprise the branched rays plus one dorsal and one ventral unbranched ray. Caudal fin ray counts are remarkably uniform within families. In Cyprinidae the count is almost always 17, except for occasional variants. Garra persica was unique in having a strong modal count of 16 branched caudal fin rays.

c) Paired fins

Paired fin ray counts can be separated into unbranched and branched rays. A small splint in some species at the origin of the paired fins was excluded from the count. There is usually one unbranched ray which is not included in counts cited here. The branched ray counts were the most important and are the ones given here. However, in the pectoral fin the innermost rays were often difficult to discern and may increase with age.

5) Scales

a) Lateral line count

The first scale counted was that scale contacting the pectoral girdle. The count continued along the flank following the pored scales and including small, additional scales lying between the large, regular scales as well as any unpored scales. The small, additional scales were relatively rare occurrences and any obviously abnormal fish - those with healed injuries for example - were not counted. The count terminated with the scale lying over the end of the hypural plate as determined by flexing the caudal fin. Some works recommend inclusion of a scale overlying the flexure only if most of its exposed field is closer to the body than to the caudal fin. Since the flexure of the caudal fin produces a relatively broad groove, this is difficult to judge in smaller fish. Therefore, the most posterior scale whose exposed surface touched the groove was the last scale counted. I have also continued the count onto the caudal fin in some species for a total count as this sometimes proved useful in comparison with counts in older literature.

b) Scales above the lateral line

This count commenced with the scale at the origin of the first dorsal fin and continued down and back to, but not including, the lateral line scale. Any scale partially or wholly straddling the dorsal fin origin was counted as one scale. The count followed the natural scale row and included any small or irregular scales in the row.

c) Scales below the lateral line

This count commenced with the scale at the origin of the anal fin, followed the natural scale row up and forward to, but not including, the lateral line scale and included any small or irregular scales. In this, and the previous count, it sometimes proved necessary to shift the counting row because of the scale arrangement. This was always a backward shift. In some instances there were several scales at the anal fin origin which overlapped each other very closely. All these were counted and account for the large degree of variation in counts between individuals of some species.

d) Scales between the lateral line and the pelvic fin origin

This count was made as in the above count.

e) Predorsal scale rows

All rows of scales between the origin of the dorsal fin and the head were counted just below the mid-line of the back on the upper flank. The final "row" at the occiput may consist of a single scale. This method was used because scales on the mid-line may be small and irregular, obscured by heavy pigment, or absent.

f) Caudal peduncle scales

This was the lowest count of the scale rows around the caudal peduncle, usually at its narrowest point. Both lateral line scales were included. Scale rows were counted even when the scale arrangement was such that occasional alternate rows touched. This count may be quite consistent between individuals of a species, but it may also vary markedly. The variation depended on the presence of large scales dorsally and ventrally on the caudal peduncle connecting the flank scale rows. When such large scales were present bridging over the top and bottom of the caudal peduncle, the total count could be, e.g. 12, but in some individuals two or more smaller scales occupied their positions so that the scale count jumped to 16.

B) Morphometric characters

All measurements were to the nearest 0.1 mm using dial calipers. Measurements were taken on the left side unless a left fin, for example, was badly deformed or broken. Badly deformed specimens were not measured. Distortions due to preservation, such as a gaping mouth or expanded gill covers, were gently adjusted to as natural a position as possible. The following list explains how the various measurements were taken. All measurements were taken in a straight line and not over the curve of the head or body.

1) Total length

From the anteriormost part of the head to the tip of either lobe of the caudal fin when that fin is normally splayed.

2) Standard length

From the anteriormost part of the snout (even when the lower jaw projects) to the end of the hypural plate (the end of the plate is found by flexing the caudal fin; in small fish it may be seen by shining a strong light through the caudal region). Standard length can be an inaccurate measurement. The end of the hypural plate is obscured by scales, flesh and caudal rays. Its position is determined by flexing the caudal fin; this flexure is taken to be the end of the hypural plate. Small fish have thin, delicate bones and the flexure may be at the anterior base of the hypural plate, at the origin of the caudal fin rays which articulate with and overlap the end of the hypural plate, or even between the last whole vertebra and the hypural plate. Large fish have a broad flexure which can give a variety of measurements by independent observers. Fortunately, in this study most fish were comparatively small and strong illumination helped to discern the end of the hypural plate. For larger fish I can only plead an attempt at consistency.

3) Head length

From the anteriormost part of the snout to the bony margin of the opercle (excluding the opercular membrane).

4) Body depth

Maximum straight line depth excluding fins or fleshy and scaly structures at fin bases

5) Body width

Maximum distance from one side of the body to the other.

6) Head depth

From the occiput vertically to the breast or lower head surface.

7) Head width

The distance between the opercles when in their normal, closed position. The opercles are gently pressed into a closed position if greatly dilated.

8) Snout length

From the anteriormost part of the snout or upper lip at the mid-line to the bony front margin of the orbit.

9) Orbit diameter

Greatest diameter between the bony rims of the orbit. This distance is not always horizontal.

10) Postorbital length

Greatest distance between the posterior bony orbit margin and the bony opercular margin.

11) Interorbital width

Least bony width between the orbits over the top of the head in a straight line.

12) Predorsal length

From the base of the anteriormost dorsal fin ray to the tip of the snout or upper lip.

13) Prepelvic length

From the base of the anteriormost pelvic fin ray to the anteriormost point on the head (snout or upper lip).

14) Preanal length

From the base of the anteriormost anal fin ray to the anteriormost point on the head (snout or upper lip).

15) Length of caudal peduncle

The oblique distance from the insertion of the anal fin to the mid-point of the end of the hypural plate.

16) Depth of caudal peduncle

The least depth of this structure from the mid-line of the ventral surface.

17) Length of the longest dorsal and anal fin rays

From the structural base of the ray to its tip.

18) Length of the dorsal and anal fin bases

From the anteriormost ray base (the origin of the fin) to the point where the fin membrane contacts the body behind the last ray (the insertion of the fin).

19) Length of the pectoral and pelvic fins

From the extreme base of the uppermost, outermost or anteriormost ray to the tip of the fin.

20) Distance between pectoral and pelvic fin bases

Used principally in Cyprinidae and Cobitidae, this and the following measurement are from the extreme base of the anteriormost, uppermost or outermost ray of the appropriate fin to the anterior base of the next fin.

21) Distance between the pelvic and anal fin bases

As above.

22) Length of fin spine

From the base of the spine to its tip. In pungent spines, as in catfishes, this excludes soft rays or membranes distal to the sharp tip, but in more flexible spines, which may taper gradually as in Cyprinidae, this measurement includes the soft tip.
 


History of Research

Written records extend back to the third millennium B.C. in Mesopotamia, the plain shared between Iran and Iraq. The Uruk IV symbol for fish dates to 3100 B.C. or 5050 B.P. Later cuneiform writing on clay tablets refer to fishes and attempts have been made to identify the species, with variable results (Scheil, 1918; Diemel, 1926; Civil, 1961: Landsberger, 1962; Salonen, 1970; Sahrhage and Lundbeck, 1992). About 324 Sumerian and Babylonian fish names have been identified referring to about 90 species (some of which are marine). Fish played a prominent part in every day life, both as food and as religious symbols (van Buren, 1948; Salonen, 1970; de Moor, 1998).

Fishing regulations had set penalties and fishing rights were leased. Guilds of fishermen existed and transport to cities with marketing was organised. Fish were sun-dried, salted, pickled, fermented and possibly smoked. Fishermen had to deliver part of their catch to the temples or as duties. Surplus fish were sold to the public. Consumption of fish was prohibited on certain days (Sahrhage and Lundbeck, 1992). See also Freshwater Fishes of Iraq website here.

The Babylonian Epic of Creation mentions nets and splitting fish for drying. Amulets and cylinder seals depicting fish are common. A hymn which praises Ishtar of Uruk gives the result of her favour as "whole channels are filled with fish, the channels swarm with fish and with dates". Fish were offered as sacrifices to gods and as part of funeral rites, as symbols of life and its renewal, and of fertility (Wright, 1990). The amount of fish required was clearly stipulated and whether it should be fresh, roasted or dried. The commoner species were requested by the basketful but rarer species were requested by numbers so a practical knowledge of diversity existed in the distant past. So numerous were sacrificial offerings that at Uruk I the floor of a room or court was covered with a thick layer of fish scales and fatty waste that gave it a deep golden-yellow tinge. Some areas had layers of compacted fish, 4-5 cm thick, comprising skeletons, skin and scales, indicative that these were not kitchen wastes but were sacrifices (van Buren, 1948). An Assyrian king would have 10,000 fish served at a banquet, although these were cheaper food items and the Sumerians favoured large, plant-eating carps from muddy pond bottoms (de Moor, 1998).

Archaeological remains containing fish bones at Abu Salabikh, Iraq, dated to 3000 B.C. (and summarised for south Mesopotamia), have been identified to include Barbus (= Luciobarbus) esocinus, Barbus (= Tor) grypus, B. (= Luciobarbus) kersin, B. (= Carasobarbus) luteus, Barbus (= Mesopotamichthys) sharpeyi, B. (= Luciobarbus) xanthopterus, Aspius vorax, Acanthobrama (presumably A. marmid), Cyprinion sp., Alburnus sp., Silurus triostegus, Mystus pelusius, Mastacembelus mastacembelus, Liza abu, Acanthopagrus sp., and Tenualosa ilisha.

Radcliffe (1926), Salonen (1970) and Sahrhage and Lundbeck (1992) review fishing in Assyrian and Sumerian-Akkadian times using nets, spears, traps, weirs and copper hooks and line. Contracts concerned with fish ponds date from the reign of Darius II, in 422 B.C., and with fishing in 419 B.C. He also discusses Ea, the god of water dating back to Sumerian times, for which a fish-god or man-fish was a symbol, still to be seen on ancient monuments in Iran (see also Green (1986)). The Middle Elamite rock relief at Tall-i Bakun near Persepolis in Fars depicts a river filled with fish but these are highly stylised and not identifiable to species.

Fish do appear on bowls and other objects or in the round from archaeological collections and some are illustrated below courtesy of F. Biglari and the National Museum of Iran:-

 

Vessel 5th millennium B.C., Susa, Khuzestan     Vessel 5th millennium B.C., Tal-e Shoqa, Fars     Rython, 3rd millennium B.C., Tal-e Shoqa, Fars 

                   

 

Jar, 4th millennium B.C., Choqa Mish, Khuzestan         Chlorite vessel, 3rd millennium B.C., Jiroft, Kerman     Ivory or bone 5th millennium B.C., Susa, Khuzestan

Sassanian bowl with fish, San Antonio Museum of Art (Wikimedia Commons).  Gold and silver Sassanian plate showing a fishing party, Iran Bastan Museum, Tehran (Wikimedia Commons). 

 

A'lam (1999b) briefly reviews fish in pre-Islamic Persian lore but most, if not all, the fishes referred to are unidentifiable today. Illustrations of fishes often occur in art work but are generally unidentifiable to species. One example is a 14.5 cm, 12th century bowl from Iran in the Victoria and Albert Museum, London. The bowl has shoals of fish in a rotating design painted in black slip on a frit ware bowl under a turquoise clear glaze (www.iranian.com/Arts/July97/Design/Page6.html, downloaded 10 June 1997). Governmental revenue from the Caspian fisheries have been recorded as early as 820-873 under the Taherids. Alam (no date) summarises the history of fisheries in Iran.

The Arabic work Aja'ibu-l-Makhluqat or "Wonders of Creation" by Zakariya b. Muhammad b. Mahmud al-Kammuni al-Qazwini published in 1263 A.D. and later translated into Persian and enlarged in 1275, records sharks entering rivers at the head of the Persian Gulf to Basrah on the Tigris and comments on their ferocity and their teeth like points of spears, swords or saws. Other Arabic and Persian works contain few recognisable species of freshwater fishes although the tenth century Kitab al-Tabikh from Baghdad contains some fish names such as bunni (= probably Mesopotamichthys sharpeyi) and shabbût (= probably Tor grypus)(Perry, 1998). Probably the best example of an early "scientific" Islamic work on zoology is the fourteenth century "Nuzhatu-l-Qulub" or "Hearts Delight" by Hamdullah Al-Mustaufi Al-Qazwini (translated into English by Stephenson (1928)). Only the "tarikh" is identifiable as a freshwater fish - Alburnus tarichi from Lake Van in modern Turkey.

Generally paintings of fish on historic items are insufficiently detailed to allow identification to species (see Stchoukine (1936) for some examples). However an interesting painting of a fish is found on a Persian miniature of the fourteenth century stored in the Metropolitan Museum of Art, New York (Dimand, 1934). The painting shows Jonah leaving the mouth of a fish. A colour figure of this painting is found in Gould and Atz (1996), although the image is reversed and a corrected colour version is in Coad et al. (2000). The painting is from Rashid ad-Din's Jami` al-Tawarikh or "Universal or World History" which contains accounts of various historical and mythical events, including the history of China and Mongolia, the Bible and incidents in the lives of Mohammad and Buddha. As Dimand (1934) points out, this book was highly favoured by Persian painters of the fourteenth century and several copies exist, the earliest being 707 A.H. (= 1307 A.D.). The painting, dating to about 1400 A.D., shows Jonah being cast up by a fish. The text on Jonah's arms however reads "The disk of the sun entered into darkness" on the left arm and "Jonah entered the mouth of the fish" on the right arm. The former, which was taken from the Gulistan (= Flower Garden) of Sa`di written in 1258, being a more poetic rendering of the latter. The angel, however, appears to be offering the naked Prophet a garment, and this, as well as the proximity of terrestrial vegetation, suggests he is leaving the mouth of the fish.

The fish undoubtedly was copied by the Persian artist from Chinese paintings (Rice, 1976; Blair, 1995). It most closely approximates some kind of carp but its mouth has been enlarged to accommodate the squatting figure, and the opercular opening approaches the eye too closely to make it a recognisable rendition of any particular species. There also are two dorsal fins (not found in any member of the carp family), and the pectoral fins are located too far from the head. Nevertheless, the fish does exhibit a number of well-observed features such as symmetrical, overlapping scales on the body with smaller ones on the caudal peduncle, paired and median fins with fin rays, and the absence of head scales and teeth.

In modern Iran, the fish is still a symbol of prosperity, blessings, abundance and happiness at Now Ruz, the Persian New Year on 21 March, when a live fish from a store (usually a goldfish) or local stream is kept in a bowl. In Persian mythology the earth is balanced on the horn a gigantic cow and as the new year starts the cow throws the earth from one horn to the other. The movement of the fish in the bowl when this happens shows that the new year has begun (Noorbaksh, 1995). Anahita, the ancient god of water, watched over people in their dealings with water and fish (Sajaadyeh, 1995).

A general survey of natural history studies in the Muslim world is given by Mirza (1983), an Islamic approach to the environmental crisis by Zaidi (1981), and Islamic principles for conservation by Ba Kader et al. (1983).

Travelers from Europe often wrote up accounts of their visits to Persia and some commented on the fishes although such comments were mostly of a general nature and species were rarely identified. An exception is the trout near Tehran and some of the older comments on these populations are given in the species description. A summary and translation into English of the earlier accounts may be found in Pinkerton (1758-1826). Adam Olearius noted that the king leased fishing in the rivers entering the Caspian. The lessees blocked the river from September to April near the mouth to catch migrating fishes. Outside this area anyone was free to fish. Sir John Chardin, in a series of English and French editions from 1686 to the early nineteenth century of his Description of Persia and Other Eastern Nations, briefly mentioned fishes (see quote at the beginning of this work, taken from Sykes (1927)) as did Fraser (1825; 1834), both authors observing the lack of diversity in a water-poor country but commenting on the presence of fishes in qanats. Continuing in full the abbreviated quote from Fraser (1825) at the beginning of this work:-

" I may remark as a curious fact in zoology, that many of the cannauts, both here (i.e. at Neyshabur) and at Shahrood, swarmed with fish, some of which were of considerable size. When it is remembered that these are not natural, but artificial sources of water, brought from under ground for distances of many hundreds and even thousands of yards; and that the water, after issuing into the open air, has but a short course, being either entirely absorbed in irrigating the cultivation, or lost in the barren plain beyond it, and thus having no communication whatever with any large or permanent body of water, it seems difficult to account for the presence of these fish. The natives say that they are to be found in most considerable cannauts, but are never put into them by the hand of man. It may be added, that we saw no fish in any of the mountain streams on the southern face of the Elburz, although some that we crossed were clear, and of considerable depth. The Russian soldiers, who catch these fish, observe the same fact, all are taken in cannauts, not in the natural streams. They are a leather-mouthed fish, of no great delicacy, but perfectly sweet and wholesome".

Cornelius Bruyn (1652-1719) (or Corneille LeBrun, de Bruin) depicts several fishes from his journey through Russia and Persia, mostly from the Persian Gulf, but including one called "sjir-majie" (= shir mahi or milk fish) which Heckel (1843b) identifies as Capoeta trutta and states that it is from Esfahan. Capoeta trutta is not found near the city of Esfahan. This illustration appears in volume 1, page 185, plate 69 of the Amsterdam edition in French published in 1718. However a reading of the text and examination of the illustration (slides kindly provided by Martine Desoutter of the Muséum national d'Histoire naturelle, Paris) show that the fish cannot be identified so clearly. No scales are shown and the colour pattern is unusual and unlike any Iranian freshwater fish. The colour pattern is vaguely reminiscent of Barbus lacerta, although much exaggerated. The illustration is possibly based on a Barbus or a Capoeta species. The author was in Esfahan on 23 November 1703 when describing the fish but the specimen is mentioned in the same paragraph as a "Lezard de mer....prend dans le Golfe Persique" and I take this to mean that the fish too may come from a locality on or near the Persian Gulf rather than the neighbourhood of Esfahan as Heckel (1843b) has it.

Floor (2003) devotes some considerable space to fisheries in Qajar Iran, not repeated here. The most important were the Caspian caviar fishery but also dried mullets were exported. Mullet were caught on mats stretched across a stream, the shadow of the mat causing the mullet to jump to avoid it and thus becoming stranded on the mat surface. The Russians controlled much of the Caspian fishery although there were also Persian concessionaires.

Scientific works relevant to Iran begin with the Systema Naturae, 10th edition, by Carolus Linnaeus (1701-1778) published in 1758 and in which scientific naming in zoology has its beginning. Linnaeus adopted many of the names from the system developed by Petrus Artedi (1705-1735) who, on a visit to Amsterdam to examine a collection of fishes from the East and West Indies, drowned in one of the canals. Genera subsequently found in Iran include Acipenser, Perca, Cobitis, Silurus, Salmo, Esox, Atherina, Mugil, Cyprinus, and Syngnathus and various species were described in these and other genera. After this date a variety of papers were published by authors in many countries describing fishes scientifically and some of these fishes were eventually found to occur in Iran, as with the Linnaean genera and species. Examples include Marc Elieser Bloch (1723-1799), a physician who began to devote himself to ichthyology at the age of 56, and Johann Gottlob Schneider (1750-1822) who collaborated with Bloch and published their "Systema Ichthyologiae" in 1801 after Bloch's death. This work contains all known species at that time (Bloch also wrote "Naturgeschichte der ausländischen Fische, 1785-1795) and in these works appear such Iranian species as diverse as the Indian stinging catfish, Heteropneustes fossilis, and the snakehead, Channa gachua (see Karrer et al., 1994); Johannes Müller (1801-1858) and Friedrich Gustav Jacob Henle (1807-1885) who published their "Systematische Beschreibung der Plagiostomen" in 1838-1841, the classical work on sharks and their relatives; Antoine Risso (1777-1845), an apothecary, who published in 1810 his "Ichthyologie de Nice" in which are described two mullet species (Liza aurata and L. saliens) and an atherinid (Atherina boyeri - see A. caspia) and in a later work (1826) the pipefish (Syngnathus abaster - see S. caspius) which are now recorded from Iran; and lastly Franz Steindachner (1834-1919), director of the "Kaiserlich-Königliches Naturhistorisches Hof-Museum (or Imperial-Royal Natural History Court-Museum - now the Naturhistorisches Museum at Vienna), who wrote so copiously on fishes from all over the world that any systematist eventually must consult his works, e.g. for the description of Schizopygopsis stoliczkae (1866) and Nemacheilus (= Oxynoemacheilus) angorae (1897)(see Kähsbauer, 1959; Adler, 1989; Herzig-Straschil, 1997). A number of fish species are named by others for Ferdinand Stoliczka (1838-1874), who collected extensively in the Himalayas and was appointed naturalist to the Second Mission to Yarkand, but who died on the way to Leh through hardships encountered on this journey (see Day, 1876; 1878).

Fish descriptions from the Middle East begin with the work of Fredrik Hasselquist (1722-1752) in his "Iter Palaestinum eller Resa til Heliga Landet Förrättad ifrån År 1749 till 1752" or "Voyage to the Holy Land Undertaken from the Year 1749 to 1752" which was published by Linnaeus in 1757 after Hasselquist "Succumbed to the fatigues and cares of the Journey" (Günther, 1869). Although this work appeared before Linnaeus' 10th Edition and is thus rejected as far as scientific nomenclature goes, it still contains recognisable and scientific descriptions of fishes.

Alexander Russell, physician to the British Factory at Aleppo from 1742?-1753, gave an account of four undescribed fishes from modern Syria in 1756 (see Russell (1794) for greater detail and illustrations) of which Mystus pelusius and Mastacembelus mastacembelus were later found in Iran. The descriptions in this work are attributed to Daniel Carl Solander (1736-1782) and to Sir Joseph Banks (1743-1820) and Solander respectively (Wheeler, 1958). Since then a number of works have appeared on Middle East fishes and although many were restricted to Syria, the Jordan River basin or drainages of Anatolian Turkey they often contain descriptions of species also found in Iran (see Bibliography).

Peter Simon Pallas (1741-1811) and Johann Anton von Güldenstädt (1745-1781) described species from the Caspian Sea basin but outside Iranian waters (Pallas, 1771, 1776, 1787, 1814; Güldenstaedt, 1772, 1773, 1778). von Güldenstädt was a naturalist on the expedition led by Pallas charged with exploring the Russian Empire of Catherine II. Pallas travelled to the Urals and eastwards while Güldenstädt went south to the Caucasus, only returning to St. Petersburg seven years later (Mearns and Mearns, 1988). Güldenstädt died in St. Petersburg at only 36 years of age from fever, his resistance weakened by diseases caught in the Caucasus. Pallas based some of his descriptions on the work of Samuel Gottlieb Gmelin (1743, 1744 or 1745-1774), an explorer and Professor of Botany at St. Petersburg employed by the Russian government who visited Gilan and Mazandaran in 1770-1772, living at Anzali for some months. Gmelin died a captive of a Caucasian chieftain, the Khan of Khaïtakes. A translated account in English of his travels in northern Iran is given by Floor (2007). It includes descriptions of fishes and fishing methods such as cast nets and gill nets.

Other important eighteenth and early nineteenth century authors describing and collecting fishes eventually found in northern Iran include A. Lovetzky and Johann Friedrich Brandt (1802-1879), Director of the Zoological Museum at St. Petersburg, who worked on sturgeons and described respectively Acipenser nudiventris and Acipenser gueldenstaedtii, and Karl Eduard von Eichwald (Eduard Ivanovich Eikhval'd) (1795-1876) who travelled to the Caucasus and Caspian Sea including Iran (1825-1826) and collected fishes although he was prevented from landing at Anzali by the Persian Governor. Eichwald's "Fauna Caspio-Caucasica" (1841) was of particular importance as it carried descriptions of new species and records of a variety of other fishes. Édouard Ménétries (= Menestrier) (1802-1861) was Curator of the Zoological Collection at St. Petersburg and collected fishes in the Caucasus during 1829-1830 and reached the Talish Mountains (Kuhha-ye Tavalesh). He listed a number of species found in the Caspian Sea and its tributaries in his Catalogue (1832). Alexander von Nordmann (1803-1866) described the fishes of the Black Sea in 1840 including gobies (Gobiidae) since found in the Caspian Sea and the herring Clupeonella cultriventris (= caspia) and the minnow Rutilus frisii.

Further to the east, there were Francis Buchanan (1762-1829) (see also under Scientific Names below) whose work on the fishes of the Ganges River in India with 269 species published in 1822 contains species later found at the westernmost extremity of their range in south-eastern Iran such as Aspidoparia morar (Gudger, 1924), and John McClelland (1805-1875) who described fishes collected by William Griffith (1810-1845) with the Army of the Indus in Afghanistan including the Helmand River basin which shares waters with Iran (McClelland, 1842). Some material was lost or badly damaged and the descriptions are "inadequate and highly confusing" (Hora, 1933).

Several authors worked on marine fishes in the Indian Ocean and Red Sea, describing species eventually found to penetrate or live in fresh waters of southern Iran. First among these was Petrus Forsskål (1732-1763), a Swedish member of a Danish expedition to the Red Sea in 1762 (Nielsen, 1993). Forsskål and four of his companions died and it was left to the sole survivor, Carsten Niebuhr (1783-1815), to publish Forsskål's fish descriptions posthumously in 1775. Some of Forsskål's specimens survive as dried skins in the Zoological Museum of Copenhagen. Forsskål was the describer of the milkfish, Chanos chanos. Wilhelm Peter Eduard Simon Rüppell (1794-1884) of the Senckenberg Museum, Frankfurt collected fishes in the Red Sea in 1822 and published "Fische des rothen Meeres" in his "Atlas zu der Reise im nördlichen Afrika" (1828-1830) followed by further field work in 1831 resulting in a second "Fische des rothen Meeres" in Neue Wirbelthiere zu der Fauna von Abyssinien gehörig (1835-1838). Rüppell described the tooth-carp Lebias dispar (= Aphanius dispar) now found throughout southern Iran. Later works are summarised by Dor (1984) and Dor and Goren (1994) for the Red Sea. The Persian Gulf fishes have received attention although there has been no comprehensive review of the fauna and its literature. Some principal works on this marine fauna include Blegvad and Loppenthin (1944), White and Barwani (1971), Randall et al. (1978), Relyea (1981), Sivasubramanian and Ibrahim (1982), Fischer and Bianchi (1984), Al-Baharna (1986), Kuronuma and Abe (1986) Asadi and Dehqani Posterudi (1996), and A'lam (1999a).

However, the most important early work on the Middle East and specifically on Iran is that of Johann Jakob Heckel (1790-1857), Inspector at the Imperial Royal Court Collection of Natural History in Vienna. He described the collections sent by Theodor Kotschy (1813-1866) to Vienna from "Syria" which includes such places as the Quwayq (= Coic, Kueik or Kuweiq) and Orontes rivers near Aleppo and Antioch, Damascus, the Jordan River, Mosul on the Tigris River and Kurdistan (Herzig-Straschil, 1997). In addition, collections were made in Iran from around Shiraz including the streams of the Maharlu basin in the Shiraz valley, the Kor River basin north of Shiraz, the Mand River (= Qarah Aqaj) which drains to the Persian Gulf and Lake Perishan (= Famur) near Kazerun. (Note that measurements used by Heckel are the "Wiener Zoll" = 26.34 mm comprising 12 "Linien" (= 2.195 mm) as opposed to the English inch (= 25.40 mm) from information courtesy of Dr. Barbara Herzig, Naturhistorisches Museum Wien). Heckel's descriptions appeared in Joseph Russegger's "Reisen in Europa, Asien und Afrika" in 1843 (volume 1, part 2) for the "Süsswasser-Fische Syriens" continued in 1846-1849 as a "Naturhistorischer Anhang" followed by "Die Fische Persiens gesammelt von Theodor Kotschy" (both in volume 2, part 3). The Syrian collections contained a number of species later found in Iran. In total 70 species were described or mentioned from "Syria" and many of the specimens are still to be found in excellent condition in the Naturhistorisches Museum, Wien. Note that these collections contained numerous specimens (and still do) while the catalogue in Vienna lists relatively few, presumably those which Heckel intended to be the type series. Heckel's publications often do not give accurate counts of the specimens on which the species is founded. It is not always evident which specimens are types and the whole series from a type locality is regarded as syntypes.

The dating of Heckel's works is not clear for the "Naturhistorischer Anhang" and the "Die Fische Persiens..." parts which have 1846-1849 on the cover. According to the International Code of Zoological Nomenclature the final date is the correct one if it cannot be demonstrated that parts of the work have their own dates. The copies of Heckel's works I have seen (mostly xeroxes) do not seem to have individually dated parts or sections and so I have used 1849 for the date whereas many earlier authors have used 1846. This does not have any significant taxonomic complications as there are no other works with potential synonyms in this date range.

The nominal Iranian species numbered 22 and these too may be found in Vienna. Of 89 species described from Syria and Iran (two were deemed to be found in both countries and a third is listed merely as the trout), 72 were described as new species by Heckel, although all are not now recognised as valid. Heckel's new species from Iran may be summarised as follows:-

1. Barbus barbulus (= Luciobarbus barbulus)

2. Systomus albus var. alpina (= Carasobarbus luteus)

3. Scaphiodon amir (= Capoeta damascina)

4. Scaphiodon niger (= Capoeta damascina)

5. Scaphiodon macrolepis (= Capoeta aculeata)

6. Scaphiodon saadii (= Capoeta damascina)

7. Cyprinion tenuiradius

8. Discognathus crenulatus (= Garra rufa)

9. Alburnus iblis (= Alburnus mossulensis)

10. Alburnus schejtan (= Alburnus mossulensis)

11. Alburnus caudimacula (= Alburnus mossulensis)

12. Alburnus megacephalus (= Alburnus mossulensis)

13. Cobitis persa (= Oxynoemacheilus persus)

14. Acanthopsis linea (= Cobitis linea)

15. Lebias sophiae (= Aphanius sophiae)

16. Lebias punctata (= Aphanius sophiae)

17. Lebias crystallodon (= Aphanius sophiae)

In all, only 4 new species were discovered according to the modern interpretation of these taxa. In addition the following 21 species (under their modern names) described from Syria and Iraq by Heckel have since been found in Iran: Acanthobrama marmid, Aspius vorax, Barbus lacerta, Carasobarbus luteus, Luciobarbus esocinus, Luciobarbus kersin, Luciobarbus pectoralis, Luciobarbus xanthopterus, Capoeta trutta, Alburnus mossulensis, Chondrostoma regium, Cyprinion kais, C. macrostomum, Garra rufa, G. variabilis, Squalius lepidus, Tor grypus, Ovynoemacheilus frenatus, Silurus triostegus, Aphanius mento and Liza abu. Heckel therefore described 25 of the species now known from Iran, the highest proportion of the fauna by a single scientist.

Some of this material was sent on exchange or as gifts to other museums although it is not always clear in their records whether the material comprises types, e.g. the Muséum national d'Histoire naturelle, Paris contains specimens marked from Vienna or Heckel of Alburnus sellal from Persepolis (sic, possibly a Heckel species re-identified as sellal)(1638), Chondrostoma regium from Mosul (1635), Cyprinion kais from Mosul (1641), Cyprinion tenuiradius from Perse (1640), Garra rufa obtusa from the Tigris (1633), Garra rufa rufa from the Orontes (1634), and Squalius lepidus from Mosul (1636). The Museum für Naturkunde, Universität Humboldt, Berlin (ZMB) has some Heckel types listed as such, plus additional material marked as from the Wiener Museum with type localities such as Aleppo and Mosul but without dates. Some of these may also be part of Heckel's material but are not indicated as types in the catalogue. The Senckenberg Museum, Frankfurt also holds some Heckel material. All this additional material has not been investigated in detail by me as to type status, although some has been examined in these museums as indicated in the species descriptions.

At the time Heckel's descriptions came out a series of 22 volumes was being published in Paris covering all the fishes then known. This work by Baron Georges Léopold Chrétien Frédéric Dagobert Cuvier (1769-1832) and Achille Valenciennes (1794-1865) appeared from 1828 to 1849 and was a seminal work in ichthyology, the "Histoire naturelle des poissons" (see Bauchot et al. (1990) for more details). It contained new species and summaries of descriptions by other authors for a total of over 4500 fishes. New species from Iran were collected by Pierre Martin Rémi Aucher-Éloy (1793-1838), a French botanist and printer, who traveled extensively in Iran from 1835-1838, eventually dying at Julfa in Esfahan from "an excess of zeal for natural sciences" (Jaubert, 1843; Cuvier and Valenciennes, 1828-1849 (1844:298); Bauchot et al., 1990). In 1835 he traveled from Baghdad to Hamadan, Esfahan, Tehran and Tabriz and in 1837-1838 he visited Shiraz, Bushehr, Bandar Abbas and the Bakhtiari mountains. The fishes he collected were Leuciscus maxillaris (= Alburnus mossulensis), Leuciscus albuloides (= ? Alburnus chalcoides) and Chondrostoma aculeatum (= Capoeta aculeata) but collection data were poor, stating only "rivers of Persia".

A similar work was undertaken by Albert Carl Ludwig Gotthilf Günther (1830-1914) whose "Catalogue of the Fishes of the British Museum" in 8 volumes appeared from 1859 to 1870 and contained new descriptions and reviews of earlier works with over 6840 species described and over 1680 doubtful species mentioned. New species from Iran or later found there were Barbus (= Luciobarbus) subquincunciatus and Hemigarra elegans. Günther also founded the Zoological Record, an annual index of the zoological literature.

Several other works appeared between these major, synoptic works of Heckel, Cuvier and Valenciennes and Günther and the next major work on Iranian fishes by Berg (1949) and these are outlined below.

Graf Eugen Keyserling joined a scientific expedition in 1858-1859 sent by the Russian Imperial Government to explore Khorasan under the direction of the acting privy councilor N. Chanikoff. The difficulty of baggage transport limited the quantity of alcohol Keyserling could carry and early fish collections spoiled. However he did draw cyprinid fishes from nature and gave good descriptions of 9 new species and reported 2 others from what is now northwest and western Afghanistan south of Esfahan, Yazd and Khabis near Kerman. Only one of his new species is now regarded as a distinct species, namely Squalius latus.

Filippo de Filippi (1814-1867) an Italian zoologist, Professor at Turin and Director of the Museum (1848-1865), accompanied an Italian embassy to Persia in 1862 visiting Tabriz, Qazvin, Tehran, Rasht and the Caspian Sea. His companion the Marquis Giacomo Doria collected fishes as far south as Shiraz. Seventeen species were described from the Caspian basin and inland waters of Iran although locality data were poor in some instances (Coad, 1985). Seven species were described as new of which 2 are still regarded as full species (Acanthalburnus microlepis and Cobitis aurata).

Albert Günther, referred to above, also described collections and new species from the borders of Iran presented to the Natural History Museum (formerly the British Museum (Natural History)), London. The earliest of these was the collection made by William Henry Colvill at Baghdad which Günther referred to 9 extant species in 1874, including a freshwater shark, and 2 new species, Barbus (= Mesopotamichthys) sharpeyi and Macrones colvillii (= Mystus pelusius). Barbus faoensis (= Mesopotamichthys sharpeyi) was described from Fao (= Faw) in another paper in 1896. The Afghan Delimitation Commission was dispatched by the British government to mark the western borders of Afghanistan. J. E. T. Aitchison was appointed Naturalist and made collections, mostly on the Afghan side of the border, from Sistan to the Hari Rud which were described in 1889 by Günther. Seven species were discovered, 3 new, of which only Paraschistura kessleri is still recognised as valid. Robert Theodore Günther (1869-1940) was the first curator of the Lewis Evans Collection (1924) which later became the Oxford Museum for the History of Science in 1935. In the summer of 1898 he made collections of a variety of animals and fossils in the Lake Orumiyeh (= Urmia) basin, including fishes, through the assistance of the Persian authorities and the Archbishop of Canterbury's Mission to the Assyrian Christians. These were described by Albert Günther in 1899 and comprised 6 species already described elsewhere and 4 new species which are still regarded as valid names, with the exception of Leuciscus gaderanus (= Petroleuciscus ulanus also described in this work). The papers of R. T. Günther, containing some notes on fishes, were examined in the New Bodleian Library, University of Oxford in 2007.

Karl Fedorovich Kessler (1815-1881) was a Russian zoologist who helped organise the St. Petersburg Society of Naturalists in 1868 and later became its President for 11 years. Kessler worked on fishes of the Volga River and in 1877 published his important monograph on the "Fishes of the Aral-Caspian-Pontic Ichthyological Region". Kessler described in this and earlier works a number of species now found in Iran including the still valid species Caspiomyzon wagneri, Clupeonella grimmi, Alburnus filippii, Luciobarbus brachycephalus, Capoeta buhsei (from "Persia", apparently near Tehran (Berg, 1949)), Chondrostoma oxyrhynchum, Oxynoemacheilus brandti, Paracobitis longicauda, and Pungitius platygaster, plus a number of other species since synonymised and other valid species reported from the Caspian Sea basin but not yet recorded from Iran.

Francis Day (1829-1889), Inspector-General of Fisheries in India and Burma, was the leading nineteenth century ichthyologist of the Indian subcontinent, attaining this position from his initial career as a medical officer with the Madras establishment of the East India Company when fishes were but a hobby. His numerous studies have some items of relevance to Iran and his 1875-1878 monograph "The Fishes of India" with its 1888 Supplement and the two-volume "Fishes" in the Fauna of British India series contain useful data and descriptions of over 1400 species.

Henri Emile Sauvage (1844-?) described in 1882 and 1884 the fishes collected by Ernest Chantre of the Lyon Museum on a scientific expedition to Syria, upper Mesopotamia, Kurdistan and the Caucasus including several new species from the borders of Iran, namely Silurus chantrei (= S. triostegus ?) from the Kura River of the Caspian Sea basin (but Berg (1948-1949) suggests that this species was collected in Syria or the Tigris basin but without any explanation), Barbus microphthalmus from the Kura River (= Luciobarbus mursa) and Labeobarbus euphrati from the Euphrates River (= Luciobarbus esocinus).

Oscar von Grimm described two species of herrings (Clupeidae) from the Volga River at Astrakhan (Alosa kessleri and A. saposchnikowii), now known also from Iran.

Aleksandr Mikhailovich Nikol'skii (1858-1942) described in three papers the fishes collected by N. A. Zarudnyi (see below) in Iran. Nikol'skii was primarily a herpetologist, head of the herpetological department of the Zoological Museum of the Academy of Sciences in St. Petersburg, and later professor at Kharkov University in the Ukraine (Adler, 1989). These included the first record of Channa orientalis from Iran and the new species Capoeta fusca, Capoeta nudiventris (= C. fusca), Capoeta gibbosa (= C. capoeta), Aspiostoma zarudnyi (= Schizothorax zarudnyi), Barbus bampurensis (= C. watsoni), Cyprinion kirmanense (= C. watsoni), Nemacheilus (= Paraschistura) bampurensis, Nemacheilus (= Paraschistura) sargadensis, Discognathus rossicus (= Garra rossica) ?

Serghyei Nikolaevich Kamenskii of Kharkov University described in 1899-1901 "Die Cypriniden der Kaukasusländer" in two volumes which described a number of new species notably in the genus Barbus since synonymised.

Nikolai Andreevich Borodin (1866-1937) was Chief Specialist in Fish Culture in the Department of Agriculture and Professor in the Petrograd Agricultural College and later an exile in the U.S.A., becoming Curator of Fishes in the Harvard Museum of Comparative Zoology. He wrote a number of articles on the sturgeons and herrings of the Caspian Sea and discovered such new species as Acipenser persicus, Alosa braschnikowii, Clupeonella engrauliformis check others?. In 1908 he co-authored with E. K. Suvorov "Caspian herrings and their commercial exploitation", the results of the Caspian Expedition of 1904. Suvorov described Alosa curensis.

Erich Zugmayer (1879-?) collected fishes along the Mekran coast of what is now Pakistani Baluchistan describing, in 1912, 6 freshwater species including 5 new ones from internal and Sea of Oman basins close to or shared with those of Iran, namely at Panjgur in the Mashkel (= Mashkid) River drainage and the Dasht River drainage. A later work (1913) added additional records for Baluchistan. The specimens were deposited in the Zoological Museum, Munich (Zoologische Staatssammlung, München) but all fishes were destroyed in World War II on 25 April 1944 (Fritz Terofal, pers. comm., 1981; Neumann, 2006). Single type specimens were deposited in the Naturhistorisches Museum Wien (NMW) and the Zoological Survey of India, Calcutta (ZSI) of Labeo macmahoni (NMW 81256), Scaphiodon daukesi (NMW 19784, ZSI F8028, ZSI F8032), and Nemacheilus (= Paraschistura) baluchiorum (NMW 19851). None of the species has been collected in Iran.

William Thomas Blanford (1832-1905)(Anonymous, 1905) accompanied the Persian Boundary Commission in 1872, publishing a two-volume account in 1876. The Commission mapped the boundary between Persia and Baluchistan. Major (later Sir) Oliver St. John, with a collector from the Indian Museum, Calcutta, also made collections from 1869-1871. Fish collections were minor and not included in Blanford's books. Part of the collections was described by J. T. Jenkins in 1910 from material deposited in Calcutta. Blanford and St. John marched from Gwadar through Jalk, Bampur and Kerman to Shiraz, with Blanford carrying on alone through Esfahan to Tehran. One new species is from what is now Pakistani Baluchistan, close to the Iranian border in the Nihing-Dasht drainage (Scaphiodon baluchiorum = Cyprinion watsoni) while the remaining material, comprising 3 new species of tooth-carps, is from the neighbourhood of Shiraz. Further discussion about the tangled nomenclatural history of these little fishes can be found in the relevant Species Accounts.

(Thomas) Nelson Annandale (1876-1924) was founder and then Director of the Zoological Survey of India (Anonymous, 1925; Kemp et al., 1925; Adler, 1989). He and a co-author reviewed the fishes of Sistan (1920) collected by Colonel Sir A. Henry McMahon and other officers of the Seistan Arbitration Commission of 1901-1904 and by officers of the Zoological Survey of India in the winter of 1918. Nine species were described, one of which, (Nemacheilus macmahoni), formed the basis for a new genus, Adiposia, since synonymised with Nemacheilus and now Paracobitis. The McMahon collection had been examined by Charles Tate Regan (1878-?), later to be Director of the British Museum (Natural History), London (now the Natural History Museum) who found 2 new species out of 5 collected in his 1906 work (Scaphiodon macmahoni (= Cyprinion watsoni) and Nemacheilus rhadinaeus (= Paracobitis rhadinaea)), by Banawari Lal Chaudhuri of the Indian Museum, Calcutta in 1909 who reported a new loach (Nemacheilus macmahoni (= Paracobitis rhadinaea)) and by Annandale in 1919 who described 2 new species of Discognathus, D. adiscus (= Crossocheilus latius) and D. phryne (= Garra rossica).

Annandale's co-author on the "Fish of Seistan" was Sunder Lal Hora (1896-1955) who was to become the leading ichthyologist of India on a par with Hamilton and Day, and Director of the Zoological Survey of India.

A. Ya. Nedoshivin and B. S. Iljin produced two lengthy papers in Russian in 1927 and 1929 on fishery capture data for Iranian waters, forming an important historical record.

Alfons Gabriel and his wife collected fishes in the neighbourhood of Bandar-e Abbas including the Genu hot spring and the Baschakird Mountains. This material was described in 1929 by Maximilian Holly of the Naturhistorisches Staatsmuseum in Vienna and contained Cyprinodon (= Aphanius) ginaonis and Barbus baschakirdi (= Cyprinion watsoni) from fresh waters.

Viktor Pietschmann (1881-1956), originally Steindachner's assistant and later (1919-1946) in charge of the fish collection at the Naturhistorisches Museum Wien, described Mugil pseudotelestes (= Liza abu) and Glyptothorax steindachneri (identification uncertain) from the Tigris River basin in Iraq based on materials collected on the Mesopotamian Expedition in 1910 (Kähsbauer, 1957).

Lev Semenovich Berg (1876-1950) was a leading Soviet physical geographer and biologist. From 1930 until his death, he was head of the "Special Laboratory of Ichthyology" of the Zoological Institute of the Academy of Sciences of the U.S.S.R. in Leningrad and an Academician (Oliva, 1977). His contributions to the ichthyology of the former U.S.S.R. and to that of Iran appeared in a number of shorter articles and in lengthy monographs from the late nineteenth century onwards. The shorter works are listed in the Bibliography and include descriptions of such new species as Alosa sphaerocephala, Barilius mesopotamicus, Alburnus atropatenae, Garra persica, Nemacheilus cristatus (= Metaschistura cristata), Glyptothorax kurdistanicus, Anatirostrum profundorum, Knipowitschia caucasica and Knipowitschia iljini. His summary work "Freshwater Fishes of the U.S.S.R. and adjacent countries" was published in 1948-1949 and in English translation in 1962-1965 and has much of relevance to northern Iran, although the taxonomy is now dated. His 1940 work on the "Zoogeography of freshwater fish of the Near East" placed that fauna in context and included Iran but it was his 1949 work "Freshwater Fishes of Iran and adjacent countries" which has been the major modern work on Iranian fishes south of the Caspian Sea basin and the Lake Orumiyeh basin. This was based on collections deposited in the U.S.S.R. Academy of Sciences Zoological Institute in Leningrad (acronym ZIL, now St. Petersburg, Russia with the acronym ZISP). The collections had been made by two Russian biologists. The first of these was Nikolai Alekseevich Zarudnyi (1859-1919), a zoologist and ornithologist who made four journeys to Iran for which he was awarded medals and the Przheval'skii Prize by the Russian Geographical Society. His first journey in 1896 was to Kuchan, Sistan and Mashhad, his second in 1898 was to eastern Khorasan and Beluchistan, the third (1900-1901) was to Khorasan, Sistan and Beluchistan including the Bampur region and the Makran, and the last journey (1903-1904) was to Gorgan, western Khorasan, western Kuhistan, southern Irak-Ajemi and Khuzestan. Zarudnyi's material had previously been examined and described by Nikol'skii (see above). The second biologist was P. V. Nestorov who worked with the Turko-Persian Demarcation Commission in 1914 and collected fishes in the Tigris basin along the present Iran-Iraq frontier.

The Zoological Museum of the Lomonosov Moscow State University (MSU) contains collections from the Caucasus and Transcaucasia including the Kura River basin and Azerbaijan but none apparently from Iran (Verigina, 1991).

Anton Bruun (1901-1961 - see Spärck (1962)) was the lead author on the description of Iranocypris typhlops, the Iranian cave fish, later the reason and subject of popular books and articles by Anthony Smith (see Bibliography).

Relevant works since 1950 can be found in the Bibliography and encompass a wide range of papers and books of varying quality and utility. There has been a rapid increase in studies on fishes of Iran, starting in the 1990s. Prior to 1900, this Bibliography lists less than 100 publications relevant to this work, many not strictly on Iranian fishes. On a decadal basis, it is only in the 1960s that publications exceed 100 and by the 1990s are an order of magnitude larger.

Several books have appeared in recent years in Farsi on Iranian freshwater fishes and include "Freshwater Fishes" by Vossughi and Mostajeer (1994), "Identification of some freshwater fishes of Khuzestan Province" by Najafpour (1997), "Atlas of Iranian Fishes. Gilan Inland Waters" by Abbasi, Valipour, Talebi Haghighi, Sarpanah and Nezami (1999), "Freshwater Fishes of Iran" by Mohammadian (1999), "The Inland Water Fishes of Iran" by Adoli (2000), "A Guide to the Fauna of Iran" by Firouz (2000; in English as "The Complete Fauna of Iran", 2005), "Iranian sturgeons in the Caspian Sea (Systematic, biology, artificial propagation, biomass evaluation and conservation, fishing and production of caviar" by Keyvan (2003),  "Freshwater fishes of Khuzestan Province (Part II)" by Najafpour (2003), "Fish Species Atlas of South Caspian Sea Basin (Iranian Waters)' by Naderi and Abdoli (2004), "A Biological Review of Caspian Sturgeons" by Sarafraz and Akbarian (2005), "Applied Ichthyology" by Hedayatifard and Ramezani (2007), "Biodiversity of Fishes of the Southern Basin of the Caspian Sea" by Abdoli and Naderi (2009), and, in English, "Fishes of Tehran Province and adjacent areas" by Coad (2008).

A report on water laws and institutions in Iran was authored by Dezfouli (1996) and gives some background on legislation affecting fish habitats through regulation of water abstraction and pollution prevention.

Several general works on zoogeography of fishes have encompassed Iran as part of their study. These include Berg (1933b; 1940), Banarescu (1960; 1977; 1992b) and Por and Dimentman (1989). Most of Iran is part of the West Asian area, which includes southern Anatolia, the Levant, and the Arabian Peninsula, or an Iranian Province which excludes the Caspian Sea, Lake Orumiyeh and Persian Gulf and Sea of Oman drainages. Berg (1940) lists the following districts within the Iranian Province: the Tehran District (= Namak Lake basin here), the Turkmen District (= includes the Tedzhen or Hari River basin here), the Sistan District (= Sistan basin here), and a Fars District (= the rest, or the basins Dasht-e Kavir, Esfahan, Kerman-Na'in, Sirjan, Lake Maharlu, Kor River, Hamun-e Jaz Murian, Hamun-e Mashkid, Dasht-e Lut, and Bejestan here). The Caspian Sea drainage is regarded as a separate area. The fauna is a mixture of elements from the European (western Palaearctic), the Mediterranean, southern Asia, High Asia and Africa and should be regarded as a transitional region (various views briefly summarised in Mirza (1994b; 1995)). Zoogeography is dealt with here in the individual Species Accounts with some mention in the drainage basin accounts.

A brief history of Afghanistan ichthyology is given in Coad (1981d) and Petr (1999), of Pakistan in Mirza (1978) and Bilqees et al. (1995). Literature, and therefore history, on Turkey is summarised in Coad and Kuru (1986) and Fricke et al. (2007), and on Iraq and the Tigris-Euphrates basin in Coad and Al-Hassan (1989). Much of the earlier Russian literature on the Caspian Sea and adjacent waters is given in Romanov (1955).
 


Fisheries

Freshwater fisheries are increasing in Iran and with this exploitation there is a commensurate need for an understanding of the whole ichthyofauna. Coad and Abdoli (1996) and Coad (1998; 1999) review the biodiversity of Iranian freshwater fishes. Reviews of fisheries, including aquaculture, can be found in the magazine Abzeeyan, e.g. Anonymous (1992c) and Madbaygi (1992), at the Food and Agriculture Organization of the United Nations website (www.fao.org), at www.agri-jahad.org, the Iranian ministry concerned with fisheries, at the Caspian Environment Programme (CEP), Baku, Azerbaijan at www.caspianenvironment.org and in various articles such as Matinfar and Nikouyan (1995), Nash (1997a, 1997b), Mehrabi (2002), Sadeghi and Agheli (2002), Saeedi (2002) and Alam (no date). Additional information is found under each of the Species Accounts, in particular for sturgeons (Acipenseridae), the most valuable fishery.

Fisheries data from various sources (and sometimes the same source) are not always compatible or comparable. The data should be treated as indicative of trends and relative fishing pressure between species. Some years may have been inadequately reported, data is incomplete, sources for figures are disparate, poaching levels have varied, and low numbers may not reflect actual catches.

Early accounts of fisheries along the Caspian shore of Iran are given by Nedoshivin and Iljin (1927; 1929), Vladykov (1964) and Keddie (1971). The freshwater fish catch increased from 6954 tonnes/year in 1974-1976 to 24,613 tonnes/year in 1984-1986, a 254% increase and five times the world average (Gleick, 1993). Inland fisheries finfish production was 30,924 tonnes in 1986 and in 1992 Iran had an inland capture fishery of 40,000 t, as did Turkmenistan; Kazakhstan had 80,000 t, Uzbekistan 27,439 t, Azerbaijan 36,371 t, Iraq 4400 t, and Armenia 4500 t (Food and Agriculture Organization, Rome, Inland Water Resources and Aquaculture Service, Fishery Resources Division, 1995a). The Caspian Sea fisheries grew from 25,987 t to 98,000 t in the decade 1990-2000 (www.agri-jahad.org, downloaded 3 November 2003). Saheli (1999) gives figures that show total aquatic production was dominated by Persian Gulf and Sea of Oman fisheries in 1995 at 63%, the Caspian Sea occupied 15% and inland waters 15%, the remainder being from international waters. Petr and Marmulla (2002) give an average catch of 30,000 t for 1995-1999 in inland waters. Kilka was the most important factor for increased catches in the Caspian and aquaculture in inland fisheries. The catch in 1998 was 75,000 t for inland waters (IRNA, 15 June 1999) - catch records vary between sources but give a general idea of the importance of freshwater fisheries. The value of all fish production in Iran rose to 1046 billion rials in 1996 from 171 billion rials in 1989 (Tehran Times, 27 July 1998). Freshwater landings increased from 22,177 t in 1985 to 115,000 t in 1994 (Food and Agriculture Organization, Fisheries Department, 1996). Cold and warm water fish production was 67,000 t in 2001 with per capita annual consumption at 5.2 kg. Production was expected to rise to 220,000 t in 2000-2005 (IRNA, 11 November 2001). Per capita yields for inland capture fisheries in kilogrammes after Food and Agriculture Organization, Rome, Inland Water Resources and Aquaculture Service, Fishery Resources Division (1995a) was as follows and shows marked increases over these years:-

1987 1988 1989 1990 1991 1992
0.321 0.329 0.342 0.444 1.038 0.667

These values compare with neighbouring countries as follows for the same period:- Iraq (range 0.182-0.672), Turkey (0.666-0.903), Afghanistan (0.079-0.102) and Pakistan (0.773-0.874). Per capita supply of cultured fish was 1.3 kg in 2003 while capture fisheries yielded 5.1 kg (Food and Agriculture Organization, Fisheries Department, 2006). This same publication gives fish consumption in kilogrammes per capita as follows:-
 
1969-1971 1979-1981 1990-1992 1995-1997 2000-2002
0.7 1.5 4.4 4.7 4.7

Catches in the Caspian Sea for 1991 and 1992 were 3036 t and 2692 t of sturgeons respectively, 13,817 and 21,527 t of kilka (herrings of the genus Clupeonella, family Clupeidae), and 18,571 and 16,873 t of bony fishes. The herring catch reached 51,000 t in 1994 from none 10 years previously (Food and Agriculture Organization, Fisheries Department, 1996). The FAO also records that the silver carp catch went from none in 1989 to 24,720 t in 1994. In inland waters the catches of warm water fish were 19,947 t and 21,462 t, of cold water fish 579 t and 775 t (both presumably from fish farming) and from "natural resources" 24,905 t and 20,183 t. These catches (totals 80,855 t and 83,512 t) are less than the totals for the marine catches in the Persian Gulf and Sea of Oman at 277,000 t and 271,000 t but are still significant (Abzeeyan, Tehran, 5(9):III, 1995).

In 1996, the total Caspian Sea catch was 58,000 t while the southern, marine fisheries reached 265,000 t. The gross value of all catches (1995) including marine fish and shrimps was U.S.$45 million while fish imports were at $65 million. Caviar made up nearly 60% of exports in 1994 and nearly half of imports are fish meal. The industry had 111,800 primary employees in 1995, including about 8000 fish farmers. Most fish (70%) is eaten fresh, 15% is frozen and canned, with some smoked or salted and the remainder is made into fish meal (Food and Agriculture Organization, Fishery Country Profile, 1997, at www.fao.org/waicent/faoinfo/fishery/fcp/irane.htm). In 1998, the annual fish catch was listed as 65,000 t with the aim of raising the catch to 110,000 t by the end of the 1995-1999 economic development plan. It was estimated that 150,000 t could be obtained from 500,000 ha of ponds and dam reservoirs (IRNA, 23 October 1998).

TACIS (2002) demonstrates the growth in catches in the Caspian Sea basin of Iran as follows. The kilka catch was 2000 tonnes per year in 1932-1959, 63,300 t/y in 1996-1998, mullets 390 t/y growing to 4560 t/y, and total catch 7440 t/y to 81,360 t/y. Nezami et al. (2000) gives the following figures for fish harvested from Caspian coastal provinces in Iran:-

Golestan:-
 
Species/Year 1997-98 1998-99
Rutilus frisii 174,869 kg 191,680 kg
*Rutilus rutilus 20,124 kg 18.025 kg
Mugilidae 43,016 kg 229,487 kg
Cyprinus carpio 229,734 kg 260,890 kg
Other 2712 kg 10,529 kg
Total 470,455 kg 710,611 kg

*May include R. caspicus as these taxa were not distinguished.

This province demonstrates a great variation in mullet catch between years.

Mazandaran (1998):-
 
Species tonnes
Cultured fishes 12,363
Rutilus frisii 2174
Mugilidae 1533
Clupeonella (kilka) 31,583
Other bony fishes 374
Total 48,027

Gilan (1997):-
 
Species tonnes
Clupeonella (kilka) 36,077
All bony fishes 2813
Acipenseridae (sturgeons) 264
Total 39,154

Unauthorised fishing in Gorgan Bay in the southeastern Caspian was estimated at 167,681 kg in 2000-2001 (Kamran, 2006). Mullets (Liza aurata and L. saliens) comprised 35.7% of the catch.

The biomass of fishes in the Iranian Caspian is estimated at 556,530 t, 12.7% of the total for the sea, with a fish density of 50.6 tonnes/nautical mile (the lowest values of any Caspian state)(Ivanov and Katunin, 2001). The Caspian Environment Programme (1998) gives the following tables for bony fish production in the Iranian Caspian Sea (tonnes) in recent years:-
 
Year/Species Kilka
(Clupeonella spp.)
Rutilus
frisii
Mugilidae Salmo
trutta
(= caspius)
Cyprinus
carpio
Sander
lucioperca
Abramis
brama
*Rutilus
rutilus
Alosa pontica
(= kessleri)
Silurus
glanis
Others Total
1973 1013 2.63 927.3 2.9 93.5 2.2 0.3 22.5 2 6 19.2 2091.53
1974 1170 338.6 403.5 1.3 101.6 2.8 - 34.6 2 10 20.6 2085
1975 1286 695.7 963.4 1.4 84.4 9 0.3 29.5 4.5 6.5 27.8 3108.5
1976 900 1231.8 2004.6 1.1 47.4 6.8 2.4 94.8 5.5 5.5 33 4332.9
1977 1261 530.6 1297.9 1.5 40.1 11.2 1 18.6 2 5 36.5 3205.4
1978 771 191.1 373.8 0.7 13 2.8 0.06 3.6 - 2.5 9.8 1368.36
1979 836 84.1 352.4 0.6 69.6 0.4 - 11.9 - 0.1 2.6 1357.7
1980 619 158.2 1411.7 0.3 69.6 - - 71.2 0.1 - 3.5 2333.6
1981 1341 252.1 408.3 0.4 129 1.6 - 217.4 0.4 2.5 9.7 2362.4
1982 798 342.3 2674.7 1.1 128.4 13.5 - 915.5 10.4 3.5 15.7 4903.1
1983 621 277.9 1637.7 0.7 160.2 4.1 - 108.6 1.6 3.5 16.7 2832
1984 1517 252.3 1219.5 1.2 173.4 3.5 - 384.4 20.3 3.5 17.2 3592.3
1985 1828 174.5 1402.9 1.1 16.4 0.7 - 200.5 34.8 3.5 10 3672.4
1986 2450 110.4 177.2 0.7 3.4 0.16 - 27.4 71.9 3.5 1.7 2846.36
1987 4389 162.7 109 0.5 19.5 0.2 - 6 13 3.8 10.5 4714.2
1988 4700 5000 1750 0.5 20 5 0 100 16 3.5 105 11,700
1989 7902 6500 2380 - - 5 - 130 30 - 2068 015
1990 8814 8500 1503 110 - 10 - 100 30 1000 3671 23,738
1991 13,817 12,000 2500 130 - 100 - 120 35 1000 2686 32,388
1992 21,527 12,000 2200 130 - 100 20 120 35 1000 1445 38,577
1993 28,730 12,727 5135 1 - 16 17 714 893 670 2155 51,058
1994 51,000 9277 2809 1 - 95 29 1366 720 28 2475 67,800
1995 41,000 8435 5014 13 - 10 5 1178 490 5 650 56,800
1996 57,000 9222 2554 8 - 6 3 878 330 22 2477 72,500

*May include R. caspicus as these taxa were not distinguished.

Abdolmalaki and Psuty (2007) give figures over a wide range of years for Iranian coastal catches in the southern Caspian Sea as follows:-

Catch and frequency 1927-1936 1937-1946 1947-1956 1957-1966 1967-1976 1977-1986 1987-1996 1997-2003
Total recorded catch (t) 8959 7224 4986 3262 5547 5384 16,903 16,201
Sander lucioperca (%) 29.7 1.7 1.0 0.2 0.4 0.1 0.1 0.2
Sturgeon meat + caviar (%) 13.4 8.8 16.3 50.9 40.9 34.2 9.4 5.0
Cyrpinus carpio (%) 9.8 8.5 1.8 2.5 2.6 1.1 6.3 6.1
Rutilus frisii kutum (%) 12.2 43.0 24.9 25.8 17.8 19.8 53.2 45.4
*Rutilus rutilus (%) 20.7 25.5 18.8 0.7 0.8 2.3 5.8 6.1
Alosa spp. (%) 1.9 6.2 14.7 2.9 0.3 0.2 3.2 3.9
Liza aurata and L. saliens (%) 0 1.8 20.9 15.8 36.1 42.2 19.7 28.9
Other species (%) 12.3 4.5 1.6 1.2 1.1 0.2 2.5 4.4

*May include R. caspicus as these taxa were not distinguished.

The Statistical Center of Iran (www.iranworld.com/Indicators/isc-t023.asp, downloaded 4 April 2005) gives kilka catches for 1997 as 60,400 t, for 1998 as 85,000 t and for 19919.79 as 95,000 t.

The bony fish catches in the Iranian Caspian Sea waters for 1999-2000 were given by D. Ghaninejad (5th International Symposium on Sturgeon, Iranian Fisheries Research Organizatio, 9-13 May 2005, Ramsar). Beach seine cooperatives took 11,170 t and the total catch, allowing for poaching, was estimated at 16,860 t. The total kutum (Rutilus frisii) catch was 1400 t and this species had an estimated biomass in Iranian waters of about 22,000 t. The catch of Liza aurata was estimated at 3559 t with about 22% undersized and the biomass estimated at 11,100 t. Cyprinus carpio biomass was very low and was estimated at 4200 t. The Rutilus rutilus (presumably includes R. caspicus) catch was estimated at 1340 t for 2000-2001, mostly poached with gill nets, and Sander lucioperca at 18 t for the same period, mostly undersized and immature. The total catch of Abramis brama was estimated to be 17 t, again undersized and immature.

Catches in the Caspian Sea showed no differences between 7 regions based on catch-per-unit-effort (cpue) (Mirzajani et al., 2005). Catches varied from 88 to 459 kg/cpue for 1991-92 and 31-418 kg/cpue for 1994-95. In 2000-01, the Anzali region had the highest values, significantly different from the Astara-Hashtpar and east of Gilan province regions.

Beach seines are known as pareh in Farsi. Beach seine cooperatives increased from 68 in 1989 to 151 in 2004 while the numbers of fishers doubled from 6000 to 12,000. About 85-100 people are members of each beach seine cooperative. The beach seines are 1000 m long, with a cod-end 10-15 m wide and 100 m long and with a mesh size legally fixed at 30 mm (smaller meshes are used too). They are hauled in by tractors. Although there are minimum sizes for fish retention, e.g. 34 cm fork length for Sander lucioperca, fisheries do retain smaller ones for home consumption or even marketing  (Abdolmalaki and Psuty, 2007). Some further details on Sander lucioperca catches are given in the appropriate Species Accounts.

Salehi (2008c) summarises the Iranian Caspian fisheries for bony fishes. In 2006 the industry employed more than 10,000 fishermen with 142 co-operatives managing the industry. Average yearly production was over 18,000 t for 1980-2006. Landings of Rutilus kutum were estimated to average 46.6% of the total bony fish catch from 1983 to 2006 due to the stock enhancement project for this species. Average fingerling production of this species from 1981 to 2006 was 191,776,000 fish (17,536,000 for sturgeon, 18,024,000 for Abramis brama and 11,012,000 for Rutilus rutilus). Beach seines are back in use as the gill nets of the 1980s were found to adversely affect sturgeon stocks. Each net may require up to 100 people and a tractor to operate. Re-introduction of beach seines partly accounts for catches rising from 17,629 t in 1993 to 21,845 t in 2005 and 23,802 t in 2006. Ghaninezhad and Abd Almalaki (2009) give further details on bony fish exploitation in the Caspian Sea and Alyan (2010) comments on declines in the fishery.

Caviar and sturgeon catches from the Statistical Center of Iran (www.iranworld.com/Indicators/isc-t023.asp, downloaded 4 April 2005) were as follows (note that the Iranian years run from March to March, so the western years are an approximation here and in the above table) :-

Year Beluga caviar Beluga meat Asetra caviar Asetra meat Sevryuga caviar Sevryuga meat
1995 (1374) 6 135 68 516 108 512
1996 (1375) 7 165 96 669 92 461
1997 (1376) 5 126 81 550 65 324
1998 (1377) 6 168 92 684 59 348
1999 (1378) 4 141 57 569 36 290

The whole fisheries industry, including the Persian Gulf marine fin fisheries and shellfish, received an investment of 500 billion rials by government and 800 billion rials by the private sector, apparently for the period 1989-1993. Nine billion rials were allocated to aquaculture by the government in 1993, planned to rise to 23 billion rials in 1994, and to 210 billion rials in the next five-year economic development plan. In 1995, 200 billion rials were allocated to preparation and provision of infrastructure activities for fish farming (http://netiran.com/news/IranNews/html/9503131INEC.html). A national project to expand fish farming within a six-year period would raise annual production by 50,000 t, create 30,000 jobs, earn $50 million a year and increase consumption of fish to 10 kg per person (IRNA, 22 January 2000). Consumption of fish in Iran is estimated at 5 kg per capita, having risen from 1 kg in the decade prior to 1999 and is expected to rise to 6.5 kg in the next five-year economic plan (by the year 2000) and to 10 kg by 2004 (later revised to 8.5 kg by 2005 (IRNA, 25 September 2000)). Per capita consumption of fish increased due to increased production but also a government policy of lower prices than for meat and poultry (IRNA, 6 March 1999; 31 May 1999). In 1993, 350,000 t of seafood products were produced comprising 30% of the country's protein requirements and a sevenfold increase over catches before the Islamic Revolution in 1979 (Abzeeyan, Tehran, 4(9):VI, 1993). The annual fisheries output was expected to reach 1 million tons by the year 2004 from a 1999 level of 400,000 tons (IRNA, 6 March 1999). Fish exports were expected to earn Iran $400 million and create 150,000 jobs by 2004. The 1999-2000 government budget allocated 300 billion rials to fisheries (IRNA, 6 March 1999). In 1998, Rana and Bartley (1998) report the average per capita fish consumption in Iran to be 4.5 kg, low compared to the world average of 13.5 kg. The Government's plan is to increase consumption to 6.5 kg by the year 2020 which would require an increase in fishery production from 382,000 t in 1995 to 670,000 t; these amounts conflicting with news reports.

Adeli and Shaabanpour (2007) looked at consumption of aquatic products in Tehran in 2001 and 2005. Per capita consumption rose from 2.8 to 3.46 kg, 16.6% of people preferred more packaged food, and farmed aquatics were consumed more than other products, live rainbow trout being preferred the most. Salehi and Mokhtari (2008) investigated attitudes in fish consumption among Iranian nutrition experts. The experts listed various factors such as fish market expansion, advertisements and promotions, health factors, and quality and trust in the seller as having effects on the increase of fish consumption in Iran.

The Caspian Sea at this time produced 60,000 t and other inland waters 59,000 t. These waters would have production increased to 420,000 t by 2020. Aquaculture has a high priority in this plan and expanded at 8.2% per year during 1990-1996, the value in 1996 being U.S.$306.6 million for a production of 30,000 t. However aquaculture production for 1988 was only exceeded in 1995 (www.fao.org/fi/publ/circular/c886.1/wasia3.asp).

Over 975 million fingerlings were released into the Caspian Sea and inland waters from hatcheries or given to fish farmers to be cultured in ponds during the first five-year plan, 1989-1993. During the next five-year economic plan, the catch was expected to increase to 2.6 million t from 1.309 million t and 1.9 billion fingerlings would be released (Abzeeyan, Tehran, 4(9):V, 1993). The "Iranian Fisheries Research and Training Organization" was expected to have a budget of 35 billion rials by the end of 1993, indicative of the importance attached to developing fisheries in Iran (Abzeeyan, Tehran, 4(5):IV, VII, 1993).

Prior to the Islamic Revolution in 1979, the Iranian fisheries were divided into two companies, known as Shilat in Farsi, a northern one centred on the Caspian Sea and a southern one centred on the Persian Gulf. The combined companies, known as the Iranian Fisheries Organization or Shilat, were under the Jihad-e Sazandegi Ministry, starting in 1987. Jihad-e Sazandegi translates as "Construction Crusade" and is indicative of the attempt to develop the fisheries to serve the growing population of Iran. The Organisation is now known as Jihad-e Agriculture as of the year 2000. The Iranian Fisheries Research and Training Organization officially commenced its activities in 1990 and is now known as the Iranian Fisheries Research Organization . It has departments of Research, Training, Scientific Information and Administration and Research Centres at Bandar Anzali and Sari in the north of Iran and at Bushehr, Bandar Abbas, Ahvaz, Bandar Lengeh and Chahbahar in the south. A general account of the fisheries and their organization in Iran is given at http://netiran.com/press/economy-domestic/html/000000XXDE0090.html which was available on the net on 14 April 1997 and a more recent version was at www.netiran.com/php/artp.php?id=1609, downloaded 19 July 2004.

Aquaculture is now of major significance. Demand for fishery products is expected to outstrip that available from fisheries (Salehi, 2003). Iran is a major producer of Chinese carps (Billard and Berni, 2004). For the year 1986-1987 aquaculture production was the largest in Southwest Asia and in 1992 at 42,420 t, it represented 50% of the production for West Asia and by value it was 62% (Food and Agriculture Organization, Rome, Inland Water Resources and Aquaculture Service, Fishery Resources Division, 1995b). Yearly cultured fish production climbed from 4753 t in 1985, to 15,000 t in 1986, 18,000 t in 1987, 33,684 t in 1988, 39,913 t in 1989, and to 45,134 t in 1990. In 1995, Iran had 32% of the main aquaculture production in West Asia (among Turkey, Israel, Iraq and Syria) although it had been 47% in 1984. The decline was due to a slower growth rate. The 1995 production was 29,000 t (Shehadeh, 1997). However other sources differ with a freshwater aquaculture production of 13,615 t for 1995 according to the Food and Agriculture Organization, Rome, Fisheries Department and Network of Aquaculture Centres in Asia-Pacific Bangkok (1997). This source summarises action plans and national objectives for aquaculture. The year 2005-2006 had 96,000 tons of warm and 32,000 tons of cold water production (Iran Daily, 10 May 2006).

The Food and Agriculture Organization, Rome, Inland Water Resources and Aquaculture Service, Fishery Resources Division (1995b) also gives different figures for a range of years:-

Year 1984 1985 1986 1987 1988 1989 1990 1991 1992
tonnes (t) 18,369 17,776 20,930 24,820 28,900 31,000 45,134 20,226 42,420
$U.S. x 1000 36,988 62,217 94,650 164,201 251,500 299,000 446,876 208,298 424,534
% West Asia t 47.72 44.58 48.54 50.80 50.15 50.87 57.76 35.84 50.30
% West Asia $ 33.23 44.54 51.26 63.47 63.40 66.32 71.23 49.33 62.04

The Caspian Environment Programme (1998) gives annual production (in thousands) of the main cultured fish species in government and private hatcheries as follows:-

Year/Species Rutilus
frisii
Acipenseridae Cyprinus
carpio
Salmo
trutta
(= caspius)
Oncorhynchus
mykiss
Abramis
brama
Sander
lucioperca
Total
1978 11,857.4 3244.8 - - - - - 15,102.2
1979 2637.8 2911.4 - - - - - 5549.2
1980 - - 3003.5 - - - - 3003.5
1981 405 2044 5 - - - - 2454
1982 280 1016.2 811.7 - - - - 4637.1(sic)1
1983 - 25,335.3 1028.9 2185.8 - - - 28,550.2(sic)*
1984 28,342.2 1104.7 5036.5 - 570 - - 35,053.5(sic)*
1985 38,000 1132.1 12,836.1 - 1804.5 - - 53,772.8(sic)*
1986 51,704.9 2283.6 20,831 - 1565.2 - - 76,384.8(sic)*
1987 72,000 3040 19,044 - 3012 - - 97,096
1988 84,306.7 3157.5 50,021.9 50 50 - - 138,036.3(sic)2
1989 140,158 3149 61,176 - 7280 - - 211,763
1990 156,268 4343 93,377 155 5389 66 118 259,716
1991 109,843 6608 84,208 155 4979 2275 1630 209,693(sic)*
1992 144,680 3457 42,709 360 1834 5929 2443 200,782(sic)3
1993 100,047 4176 73,321 335 7401 5524 1160 191,964
1994 142,734 6295 104,089 640 8423 10,350 2888 275,418(sic)*
1995 117,919 9125 112,824 800 11,937 11,217 2270 266,092
1996 142,092 12,456 130,371 424 28,940 8478 2414 325,175

*Total from CEP (1998), not quite accurate; 1 = 2107.9; 2 = 137,586; 3 = 201,412.

Aquaculture production was expected to reach 110,000 t by 1999 (Abzeeyan, Tehran, 6(8):V, 1995) although reports in 2001 list a figure of 90,000 t. The production target for 2006 was 550,000 t, an increase of 1800% over 1995 (Shehadeh, 1997). These figures conflict with the ones in the table above*. The following table from www.agri-jahad.org, downloaded 15 November 2002 gives somewhat different figures for production of aquatic farms but it is not always clear whether the same values and methods of organising data are being used:-

Description/Year 1996 1997 1998 1999 2000
Number of Farms 3330 3647 3801 4524 -
Area (ha) 558,151 516,268 741,592 819,052 -
Production (tonnes) 65,000 65,000 72,000 67,800 66,000

Hosseinzadeh (2003) gives the following figures in tonnes for total fisheries production in Iran (note that southern waters are marine captures):-
 
Year/Area Caspian Sea Southern Waters Inland Waters Total
1978 3724 25,500 3219 32,443
1987 14,401 130,000 15,000 159,401
1989 21,193 239,000 40,490 300,683
1990 25,978 247,000 42.040 315,018
1991 34,596 248,000 45,131 327,727
1992 40,769 271,000 42,420 354,189
1993 52,768 272,000 44,123 368,891
1994 69,700 235,000 45,300 350,000
1995 58,300 265,000 59,000 382,300
1996 74,100 260,920 65,000 400,020
1997 76,200 259,000 65,000 400,200
1998 101,500 226,500 72,000 400,000
1999 110,000 234,200 67,800 412,000

Hosseinzadeh (2003) also gives warmwater fish (major carps, see below) production by province. Average production (tonnes/ha) increased as follows: 1989 (1 t/ha), 1990 (1.5), 1991 (1.5), 1992 (2.8), 1993 (3.0), 1994 (3.1), 1995 (3.3), 1996 (3.5), 1997 (3.4), 1998 (3.5) and 1999 (3.6). Coldwater fish production (primarily rainbow trout, Oncorhynchus mykiss) was as follows in tonnes:-
 
Province/Year 1989 1990 1991 1992 1993 1994 1995 1996 1997 1998 1999
Fars 219 118 104 148 203 410 350 491 717 1066 1174
Tehran 297 302 250 308 283 365 368 495 339 638 691.8
Bakhtiari 25 39.5 70 70 105 220 271 381 468 707 1104.6
Mazandaran 20 57 97 150 140 141 170 196 346 740 844.5
Azarbayjan (west) 4 4 30 22 25 104 64 84 108 234 257.7
Lorestan - - - 3 2 11.6 11 68 131 319 670
Bovir Ahmadi va Kohkiluyeh 6 9 9 53 45 39 24 52 43 124 239.2
Khorasan 18 17 18 21 32 38 35 38 55 88 174.5
Others 0 0 0 0 0 0 39 93 303 1078 1876
Total 589 546.5 578 775 835 1328.6 1332 1898 2510 4994 7032
Average (kg/cu m) - 9.5 9.5 9.5 9.3 10.3 10.7 11.2 12 12 -

The website www.iranseafoodexpo.ir/portion.asp, downloaded 9 February 2006, gives the following production of freshwater fishes, presumably in tonnes, with some obvious rounding of figures and conflicts with figures above:-

Year 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003
Cold water 835 1200 1500 1900 2510 4994 7000 9000 12,170 16,026 23,137
Warm water 43,288 44,728 51,554 63,229 61,964 66,137 55,862 52,987 53,843 79,084 67,811

Carp culture is the most important fisheries subsector according to Salehi (1999, 2004a). Chinese major carps are reared in hatcheries and, at about 8 days of age, they are transferred to nursery ponds. At about 10 g in weight they are transplanted into water bodies or grown out to market size (1 kg) in farm ponds (Saheli, 1999). Salehi's 1999 thesis gives an economic, marketing and consumer study of carp culture in Iran in the 1990s, concentrating on Cyprinus carpio. He maps fish culture facilities and hatcheries, gives production of carps by species and by provinces, and also gives an overview of Caspian fisheries apart from carps. However carp culture is more generally used in the sense of the Chinese major carps (Cyprinus carpio, Hypophthalmichthys molitrix, Ctenopharyngodon idella and Hypophthalmichthys nobilis, often reared in polyculture. C. idella commands the highest price followed by H. molitrix with C. carpio the cheapest. Polyculture stocking in natural and artificial water bodies is usually 28-32% Cyprinus carpio, 40-50% Hypophthalmichthys molitrix, 5-10% H. nobilis and the rest Ctenopharyngodon idella. Average yields varied from 43 kg/ha in 1993, to 40 kg/ha in 1994 to 49 kg/ha in 1995. Higher yields are cited by Salehi (2004a) at 1540 kg/ha in 2001 but this may be for growth in summer months and special condition. Total carp production was 54,000 t in 2001 (but see below after FAO, also from Salehi). Salehi's data differ from those of Hosseinzadeh (2003) above. The following figures are in tonnes:-
 
Species/Year 1991 1993 1995 1997 1999 2000 % growth 1990-2000
Cyprinus carpio 5502 4206 6561 5435 4600 7000 27
Hypophthalmichthys nobilis 983 1052 1269 1360 1150 1500 53
Hypophthalmichthys molitrix 10,019 12,619 15,228 16,310 13,800 17,000 70
Ctenopharyngodon idella 3143 3155 3942 4078 3450 2000 -36
Total 19,647 21,032 27,000 27,138 23,000 27,500 40

Production by major fish-culturing provinces from Salehi (2004a) for carps is as follows:-
 
Province/Year 1991 1993 1995 1997 1999 2001 % share in 1995 % share in 2001
Khuzestan 9119 6019 2830 12,000 4309 200 11 0.8
Gilan 6689 2164 1445 1360 1029 1270 6 4.8
Mazandaran and Golestan 1958 3813 8975 10,060 9518 15,700 36 60.9
Sistan and Baluchestan 4353 3000 4600 4200 11,307 0 19 0
Fars 216 2657 1320 1450 743 400 5 1.5
West Azarbayjan 875 1065 1633 1800 1905 2350 7 9
Others 1693 3539 4036 3915 5007 5865 16 23
Total 24,903 22,257 24,836 34,785 33,818 25,785 100 100

New aquaculture developments are reported regularly, e.g. see Abzeeyan, Tehran, 7(4):IV-VI, 1996; Aavakh-Kismi, 1996). The share of aquaculture compared with total fisheries production more than doubled between 1980 and 1987, from 5.5% to 12% due to high private sector investment while the monetary value climbed from 10.9% to 22.2%. Aquaculture is concentrated in Gilan, Mazandaran, Khuzestan and Markazi or Tehran provinces where 96% of the total number of existing establishments are found and 87% of total production (Ahmadi, 1993). Various other areas of the country are taking on fish culture plans, e.g. Anonymous (1991b; www.irna.com/newshtm/eng/08151227.htm, IRNA, 29 July 2000) - Lorestan Province; Anonymous (1992b) - Chahar Mahall va Bakhtiari Province; Anonymous (1996) - Kermanshahan Province; Islamic Republic News Agency (19 October 1997) - Ilam Province). In 1992 there were over 8047 ha of ponds and 503,500 ha of natural and semi-natural reservoirs. Consumption of aquaculture products was 800 g and over 10,400 people were employed in private sector aquaculture (Emadi, 1993a). The number of warm-water fish farms in 1996 was 3736 with an area of 7989 ha and the number of cold-water fish farms was 79 with an area of 164,984 ha (Iranian Fisheries Research and Training Organization Newsletter, 17:4-5, 1997). Lorestan Province produced 772 t of farmed fish in 1997 with 1000 t predicted for 1998 and a long-term goal of 21,000 t worth 156 billion rials and 10,000 jobs. In 1997, 50 fish farms were under construction along with 125 pools for fish culture purposes and 10 billion rials were invested (Tehran Times, 22 September 1998). Yazd Province produced 36 t of trout from ponds, 16 t of this from saline water, in 1997. In Dehshir and Marvast, 250 t were to be cultured with 200 t in salt water. For 1999, 500 t were forecast for this province (Tehran Times, 17 September 1998). The Azadegan Fish Farm south of Ahvaz was scheduled to produce 70,000 t of cold and warm water fishes annually from 342 pools of 15 or 40 ha, employing 4250 people directly and 13,000 indirectly, and with a gross revenue of 305 billion rials annually (IRNA, 11 November 1998). In the Iranian year ending 20 March 2002, warmwater fish culture produced 3843 t and coldwater culture 12,169 t (www.irna.com, downloaded 6 November 2002). Confusingly, the warmwater fish production in the year ending 20 March 2003 was expected to be 30,000 t according to IRNA (17 December 2002), and compare tables above.

The following table from www.agri-jahad.org, downloaded 15 November 2002 shows production of fry of various species in thousands:-


Description/Year 1996 1997 1998 1999 2000
Kutum 142,092 154,367 143,361 147,879 147,437
Sturgeons 12,456 21,626 24,557 18,857 18,279
Carps 130,371 113,172 33,785 99,493 116,398
Salmon 424 349 510 412 400
Trout 28,940 28,651 75,378 71,930 115,166
Bream 8478 12,995 13,792 14,231 14,325
Perch (probably zander) 2414 3800 3615 4257 3931
Other - 15,800 13,896 10,977 16,900
Total 325,175 350,760 308,894 368,036 432,836

Kutum or whitefish (Rutilus frisii) is very popular in Iran and has local cultural significance, hence the effort expended. Carps presumably includes the common carp (Cyprinus carpio) and other major carps (Hypophthalmichthys spp., Ctenopharyngodon) farmed in numerous localities as is rainbow trout (Oncorhynchus mykiss) which probably accounts for most, if not all, of trouts above. The salmon is Salmo caspius, difficult to re-establish its Caspian Sea migratory stocks because of habitat changes.

Integrated rice-carp farming and trout farming during the post-harvest period is also being developed. In 1999 rice-field farming yielded 126 t of fish, as well as fertilising the fields and controlling the rice stem borer (Petr and Marmulla, 2002). Salehi and Momen Nia (2006) analysed the benefits of fish and rice integrated culture in Iran and found it would increase farmer's profits and reduce the need for fertilisers and pesticides.

Drought conditions have severely affected fish farming in parts of Iran, e.g. the warm-water farming in Golestan and Mazandaran provinces which lost $6.5 million in 2006 because of low rainfall and the subsequent drought. Output shrank by 5000 tons in Mazandaran and 1000 tons in Golestan and projected growth of 15-20% was not attained. This report, from www.agriculturenews.net, downloaded 2 February 2007, noted that Mazandaran alone accounts for 30% of Iran's farm fish production.

Various studies have been carried out on aquaculture facilities or fish farms in Iran, aimed at improving the yield and combating problems. For example, Ebrahimzadeh Mousavi and Khosravi (2001) found the toxigenic fungi Aspergillus flavus, Alternaria spp., Penicillium spp. and Fusarium spp. at a fish farm for common, grass and silver carp in northern Iran. Shahsavani et al. (2001) found carp pox in common, grass, silver and bighead carp in a fish farm in Mashhad; Fathiazad et al. (2002) found clove oil to be a suitable substitute anaesthetic for MS-222 (which has side effects and 21-day withdrawal period) in juvenile Cyprinus carpio, Hypophthalmichthys molitrix and Ctenopharygodon idella; Abtahi et al. (2002) found the LC50 of clove essence was no different from MS-222 for cultivated Acipenser persicus, Oncorhynchus mykiss and Cyprinus carpio; Rabani and Nourouzi (2002) studied the quality of the water output from the Neka Power Station in the eastern Caspian basin for its possible use in warmwater carp culture, finding it suitable except for dissolved oxygen levels; Yakhchali and Mahmudihesar (2002) surveyed abundance of Ichthyophthirius multifilis (a protozoan causing white spot disease) in coldwater fish farms in West Azarbaijan and Seyed Moratzaei et al. (2002) studied this parasite's in vitro culture; Ebrahimzadeh et al. (2003) examined polyculture of female grass carp x male bighead carp with silver, bighead and common carp (final weight gain was not different between hybrids and grass carp, for example); Ghomi Marzdashti and Azari Takami (2004) studied effects of polyculture of silver, common, grass and bighead carp (only bighead showed increased growth, for example); Safari (2006) sampled bacteria on 51 farms and examined their use in improving chemical conditions; Esteki (2006) determined the best conditions for manuring fish farms; Rahmani and Ehsani (2006) studied ion exchange and air stripping methods for removing ammonium, which can kill fish in culture systems; Ghorbani Vaghei and Ahmadi (2007) studied the diversity and abundance of of macrozoobenthos at three fish farms for Chinese carps in Gilan; etc..

Parasites of fishes are common in aquaculture and wild-caught fishes; the species are detailed in each of the Species Accounts. Clostridium botulinum is present in coastal areas of northern Iran and is a potential food hazard if preservation is inadequate. Contamination rate was 10% in Sander lucioperca and 6.66% in Salmo trutta (= caspius if native) (Tavakoli and Razavilar, 2003; Tavakoli and Tabatabei, 2005), 2.2% of smoked carp, 1.1% of fresh carp, 1.1% of smoked kutum and 1.1% of osetr caviar (R. S. E. Khandaghi in 5th International Symposium on Sturgeon, Iranian Fisheries Research Organization, 9-13 May 2005, Ramsar).

Shariati and Nikfetrat (2005) survey the attitudes of fishermen to stock enhancement and conservation efforts in Gilan Province and found a significant positive attitude. Overfishing and illegal fishing were commonly cited as major problems. Emami and Hosseini (2004) also assessed the participation of fishery cooperatives from Sari in preserving fish resources.

Marketing fish in Iran was discussed at www.shilat.com (downloaded 28 February 2007) and in Salehi (2006) including such items as product quality, availability, variety, safety, price control, shelf-life, size control, consumption behaviour, prices, among others. Adeli et al. (2010) found households in Tehran bought farmed fish 11 times per year, with trout having the highest demand, and reviewed factors preferred by consumers such as live fish and price decrease in competition with wild fish.

Quliyev (2006) details fish farming in the neighbouring country of Azerbaijan with relevance to Iranian Caspian Sea basin species.
 


Geography

Iran is the second largest country in Southwest Asia (after Saudi Arabia with less than 20 freshwater fish species), has an area of 1,648,000 sq km and ranks fourteenth in the world in size, nearly as large as the British Isles, France, Italy and Spain combined (Firouz et al., 1970). It lies between latitudes 25°N and 40°N and longitudes 44°E and 63°E. Its northern border is shared with the former U.S.S.R. (Armenia (35 km long) and Azerbaijan (611 km) in the west opposite Iranian Azarbayjan, and Turkmenistan (992 km) in the east opposite Mazandaran, Golestan and Khorasan) and includes the southern part of the Caspian "Sea", by far the world's largest lake (436,284 sq km) and one of the deepest (1025 m). The Iranian coastline extends for 740 km. The eastern border is shared with Afghanistan (936 km) and Pakistan (909 km). The southern border fronts on the Sea of Oman and the Persian Gulf, a coastline of 2440 km. The western border is with Iraq (1458 km) in the south and Turkey (499 km) in the north. Much of Iran lies at an average altitude of about 1000 m, a feature found only in a few countries world-wide. Only Khuzestan, the Caspian Sea coast and the Persian Gulf coast form lowlands. These lowlands are quite narrow, often less than 20 km wide. Mountains are the most prominent feature of the Iranian landscape. The two major chains are the Alborz or Elburz, which rim the Caspian Sea basin in the north, and the Zagros which form a chain down the western side of the country. Inland of these chains lies the Iranian plateau, which is flanked on the east and south by lesser chains of mountains. The country has been likened to a bowl or saucer. This central plateau has extremely high summer temperatures and often very cold winters. The deserts of this plateau are barren and among the driest in the world. Rain falls only in winter. The terminal basins for streams and springs may be dry for years. There are extensive salt crusts, known as kavirs, over black, slimy mud and large areas are composed of hard, gravel plains known as dashts, prominently the Dasht-e Kavir and the Dasht-e Lut. Water is scarce in these regions, often restricted to small streams and springs. Larger rivers have their source in distant mountains. Between the Tigris and the Indus, only the Hirmand River on the Afghanistan border is large enough to be a river on a world scale - various "rivers" in the intervening area are really small streams easily fordable on foot for much of the year.

Iran topographic map from Wikimedia Commons.

 

Iran, satellite view (NASA and Wikimedia Commons).

Iran, satellite view (NASA and Wikimedia Commons).

 

The total renewable water resource of Iran is estimated as 137.5 km3/year. 9 km3/year are through transboundary rivers such as the Hirmand, Tedzhen and Aras and about 10 km3/year is surface runoff to other countries notably Iraq. More than 1900 km or 22% of Iran's borders are rivers (Chavoshian et al., 2005).

Fisher (1968) gives a general, physical geography and Breckle (1983) gives a general account of the features and life (excepting fishes) of deserts and semi-deserts in Iran. Barthold (1984) gives an historical geography of Iran and Yarshater (continuing) has many articles on geographical features. Geological literature is summarised in Dürkoop et al. (1979) and Davoudzadeh (1997).

It is pertinent here to interject a note on geographical names. Transliteration of Farsi place names into English is possible by more than one system. This results in variant spellings for geographical features in articles and on maps of Iran. For convenience, I have followed the official standard names approved by the U.S. Board on Geographic Names. The Board publishes a gazetteer for Iran with a designation of the geographical place (e.g. lake, populated place, stream, spring, etc.) and its latitude and longitude. The latest gazetteer is available from the Defense Mapping Agency, Combat Support Center, Washington, D.C. 20315-0010. Some literature localities could not be identified from maps or gazetteers. They are placed in quotes (".....").

I have not included the diacritical marks used in the Board's system. They would be of little help to those unfamiliar with Farsi and perhaps unnecessary to those who are. Needless to say, there are variant diacritical marking systems and in any case pronunciation varies throughout Iran.

The situation is further complicated by transliterations into other European languages and readers should be aware of this when reading non-English papers on Iran or Iranian fishes, e.g. the English Shiraz is Chiraz in French, and Genu, the type locality of Aphanius ginaonis, has such variants as Ginau, Genow, Gueno, Geno, and finally Ginao from the German transliteration, hence the trivial name. As if this were not enough, the vagaries of political fortune are writ large upon the face of Iran (which used to be Persia). Bandar-e Pahlavi has reverted to its older name of Anzali (often spelt Enzeli on older maps), Reza'iyeh to Orumiyeh (= Urmia in older English literature), and Shahreza to Qumisheh after the fall of the Pahlavi Dynasty in 1979. Other variants are Bandar-e Khomeyni (formerly Bandar-e Shahpur), Bakhtaran (formerly Kermanshah), and Khuninshahr or City of Blood (formerly Khorramshahr or City of Joy, and again Khorramshahr). I have retained names current for the years 1976-1979 recorded in the Board's gazetteer (1984). One exception is the province of Hormozgan (or Hormozdgan) which I have preferred for its brevity over the older name on some maps of Saheli-ye Jazayer va Banader-e Khalij-e Fars va Darya-ye Oman! The province of Mazandaran is now split into two with the eastern part termed Golestan, and Khorasan and Markazi have also been split up. Iranian governments have a distressing tendency to change the names and borders of provinces. The provinces used here are as follows, being what existed when the data was compiled:-

Ardabil

Azarbayjan-e Bakhtari (= Azarbayjan-e Gharbi or West Azarbayjan)

Azarbayjan-e Khavari (= Azarbayjan-e Sharqi or East Azarbayjan)

Gilan

Mazandaran (now split to include Golestan in the east)

Kordestan

Zanjan

Semnan

Khorasan

Kermanshahan (or Bakhtaran)

Hamadan

Markazi (= Central or Tehran; sometimes split into Tehran and a southeast part called Markazi)

Qazvin

Qom

Esfahan

Ilam (or Ilam va Postkuh)

Lorestan

Khuzestan

Chahar Mahall va Bakhtiari

Bovir Ahmadi va Kohkiluyeh (or Boyer Ahmadi-ye Sardsir va Kohkiluyeh)

Fars

Yazd

Kerman

Bushehr (or Khalij-e Fars)

Hormozgan (or Hormozdgan or Saheli-ye Jazayer va Banader-e Khalij-e Fars va Darya-ye Oman)

Sistan va Baluchestan

Another complication is the tendency for long rivers to have several names along their course, sometimes taken from the nearest population centre, and for locally used names to be different from map or gazetteer names. Names also vary with language and through time. One of the major rivers of Fars Province appears on maps as the Mand River, but near Shiraz it is called by its Turkic name Qarah Aqaj (also transliterated Qara Aghach, Qareh Aghaj, Qara Agach, Qareh Aqaj, Qareh Aqach, Kara Agach, and Kara Agaj). The Kor River, also in Fars, is known in older papers as the Araxes River which is not the same as that forming the border between Iran and the former U.S.S.R. (which anyway is often spelt Aras or Araks!).

The early geological history of Iran and neighbouring areas has necessarily affected the distribution of fishes, facilitating dispersal or hindering it, isolating or joining species. Some historical features are discussed under the appropriate drainage basin descriptions below or under the relevant genus or species but others are more widespread and are briefly outlined here. Sources include in particular Wolfart (1987) but also Harrison (1968), Takin (1972), Falcon (1974), Stöcklin (1968, 1974a, 1974b), Krinsley (1970), Stoneley (1974), Kashfi (1976), Shearman (1976), Booth (1977), Jackson and Wood (1980), Berberian and King (1981a, 1981b), Haynes (1981), Rögl and Steininger (1984), Šengör (1984), Oosterbroek and Arntzen (1992), Rögl (1998; Rögl, 1999), and Adams et al. (1999). There have been no cladistic analyses of taxa on which history can be determined. Zoogeographical analyses are based on present day distribution and suppositions on relationships. During the Cretaceous and through the Early Oligocene the Tethys Sea, several thousand kilometres wide, extended from the Mediterranean Sea to the Indian Ocean, separating the Afro-Arabian and Eurasian continents. Afro-Arabia was part of Gondwanaland. The usual assumption is that Iran belongs to Eurasia, perhaps with Central Iran a microcontinent or island or as a northern continuation of Arabia, and with East Iran a microcontinent or peninsula of Eurasia. Förster (1976), however, maintains that Central Iran, and probably North Iran, were part of Gondwana. The Tethys covered much of what is now Iran and was a barrier to the movement of freshwater fishes. The ocean regressed during the Late Oligocene except for a Euphrates-Persian Gulf furrow and the Zagros and Makran troughs. Continental sediments were deposited in endorheic basins of Iran. The Tethys closed in the Middle to Late Miocene as evidenced by mammal migrations between Asia and Africa. The establishment of continental conditions over Iran has been continuous since the Late Miocene except for an inundation in the Late Pliocene in the Zagros trough and the Makran coastal region. There may also have been an early Miocene connection between Arabia and Iran/Iraq allowing movements of freshwater fishes (Adams et al., 1999). Iran is therefore composed of parts of Gondwana, which was the continent south of the Tethys, welded to the northern continent and parts of the Eurasian plate (such as the central and eastern Iranian microcontinent). The northeastward movement of the Arabian Plate caused the closure of the Tethys and led to the folding which in the Miocene/Pliocene orogenies formed the Zagros Mountains, a prominent feature of western Iran important in zoogeographic studies of fishes (see Kashfi (1976) for an opposing view). The Zagros orogeny is related to the opening of the Red Sea which formed a barrier to fish dispersal. The Alborz Mountains are a northern part of the Alpine-Himalayan orogen of which the Zagros are a southern part and started to rise in the upper-lower Pliocene (Krinsley, 1970; Stöcklin, 1974). A continuous land-bridge between Eurasia and Africa has been in existence since the upper Miocene, facilitating freshwater fish dispersal. Hora (1937) and Menon (1957) refer to wet, marshy, tropical conditions and headwater captures along the whole southern face of the Himalayas and westwards during the Pliocene and early Pleistocene facilitating the spread of fishes from the east to Iran. Hora (1937) and Briggs (1987) consider that cyprinids entered Africa from southeast Asia 18-16 MYA, in the early Miocene, while other groups moved through Iran and the Arabian Peninsula beginning in the early Eocene. Kosswig (1951; 1952; 1955a; 1955b) notes the similarity at the generic level between Indian and African fishes, e.g. the cyprinids Barilius, Garra and Labeo, indicating that these fishes arrived in Africa from India after the desiccation of the Syrian-Iranian Sea in the Pliocene. The primary route, according to Kosswig and to Por (1987), was a northern one around the barrier of the Persian Gulf and Sea of Oman via northern Arabia, Syria and the Levant. Cooling conditions in these areas during the Pliocene and especially the Pleistocene glaciations, and arid climates at times, were unsuitable for tropical forms. These movements left a selection of fishes in what is now Iran including the cyprinid Garra, the sisorid catfish Glyptothorax and the spiny eel Mastacembelus.

The Pleistocene fore-deep of the Himalayas may have had connections with the Tigris-Euphrates basin which extending down the Persian Gulf as a river valley. The Tigris-Euphrates basin formed during the Pliocene and was colonised by primary freshwater fishes no earlier than the late Pliocene (Krupp, 1983). Movements of fishes into Iran from the west and north were also affected by the presence of the Tethys Sea and a brief account is given under the genus Barbus sensu lato which has been studied in this regard.

The present picture of the Arabian peninsula is of an arid desert unsupportive of fish life. The presence of fishes in Arabia and the Levant, and even Africa, with apparent relationships to fishes from Iran and the east indicate that fishes must once have traversed this area. Movements of fishes are thought to have been in a northern arc around the Fertile Crescent or its earlier version. However this modern picture is perhaps illusory as there is evidence of a more hospitable environment in the Arabian Peninsula at various times in the past. Wadis were active during "pluvial" periods of the Pleistocene as evidenced by deposition of fluvial material (Al-Asfour, 1978). One of these wadis drained much of central Arabia to the Kuwait area. The "Kuwait River" once ran from the Hijaz Mountains in western Saudi Arabia northeastwards for about 850 km to drain into the Persian Gulf via a vast delta occupying much of modern Kuwait. The river was 8 km wide and over 15 m deep along most of its length (Hamblin, 1987; Anonymous, 1993b). This river last ran between 11,000 and 6,000 years ago and could have provided a highway for fish dispersal. Earlier rivers of this nature dating to the Late Miocene (Forey and Young, 1999; Hill and Whybrow, 1999; Friend, 1999), the Pliocene (Gerson, 1982), and others like it in other parts of the peninsula, as well as shallow lakes (e.g. Lake Mundafan in the Rub' al Khali at 36,000-17,000 B.P. and again at 9000-6000 B.P.) would have facilitated transfer of species across the Arabian Peninsula, today an impassable desert for fishes, e.g. at the height of the Würm glaciation 40,000 years ago (Chapman, 1971; McClure, 1976; Al-Sayari and Zötl, 1978; Brice, 1978; Jado and Zötl, 1984; Wagstaff, 1985). A freshwater connection between Iran and Arabia was almost continuous from 70,000 to 20,000 years B.P. (Krupp, 1983). However no fish remains have been found in the late Pleistocene lakes although freshwater molluscs are frequent, Hippopotamus remains are reported and Neolithic fish hooks have been found in Al Hasa in eastern Saudi Arabia. Incomplete Miocene freshwater fish fossils are reported from the Jizan basin in the Tihama north of the Saudi Arabian-Yemen border (Brown, 1970). One was identified as a Barbus and the other as a Tilapia. Both these identifications are of such a general nature (see account on the genus Barbus and related genera for example) as to throw little light on past history or relationships with modern taxa. The Lower Miocene fauna of Al-Sarrar at 15-17 MYA, northwest of Dhahran in eastern Saudi Arabia, contains pharyngeal teeth thought to be Barbus sensu lato, and more interestingly several thought to be Labeo (Thomas et al., 1982). This latter genus is not now found in the Middle East but occurs in the Indian subcontinent and Africa. The Late Miocene Baynunah Fauna of Abu Dhabi in the United Arab Emirates contains Clarias, Bagrus shuwaiensis and Barbus sensu lato in a river connected with an ancestral Tigris-Euphrates system (Forey and Young, 1999). These fossils tend to confirm the hypothesis that fishes of Asian origin reached Africa through the Middle East and could have taken what may be termed a southern route across the Arabian Peninsula. However Forey and Young (1999) point out that the modern Arabian fauna may not have a history stretching back to the Miocene but is due more to a re-invasion after a loss of an earlier fauna. The modern Iranian fauna, in part, may be a remnant of movements at various times yet to be resolved in the absence of species-level phylogenies.
 


Climate

The general climate of Iran is based on Bobek (1952), Ganji (1960, 1968), Taha et al. (1981), "Aquastat" from the Food and Agriculture Organization, Rome (www.fao.org/ag/agl/aglw/aquastat/iran.htm) and www.bibliothecapersica.com/articlenavigation/index.html, under ab (= water) and climate, downloaded 24 December 2004. Kouchoukos et al. (1998) give an overview of climatology for Southwest Asia based on satellite datasets. Precipitation, its amount, nature and seasonality, is important in determining the water regime and thus the habitats for fishes. Iran is sparsely vegetated, both naturally and through the agency of man, and the air temperature and amount of insolation has a direct effect on water temperatures. Insolation is continuous through summer days when clouds are a rarity over much of Iran and the weather remains settled for weeks at a time.

In general, the climate of Iran can be classified as arid to semi-arid, with more than 80% of the country characterised by less than 250 mm annual rainfall. Mountain ranges block off the interior of Iran and give extremely continental conditions except for the narrow littoral zones on the Caspian shore and the Persian Gulf. Summers are hot and dry with little change from day to day. Three main climatic types are found: warm, temperate and rainy with a dry summer in the Caspian coastal area, dry, hot desert in the central plateau, and dry, hot steppe in the rest of the country. Humidity is generally low because of the altitude, much of Iran being over 1000 m average height. Coastal regions along the Persian Gulf have a high humidity, especially in summer. Wind patterns are deflected by the Zagros and Alborz ranges in the west and north. Summer winds are mainly north and northwest over much of northern and central Iran and are hot, dry, and strong for long periods. The Sistan "Wind of 120 Days" from the northwest blows from the end of May to September continuously and is very hot, dry and sand-laden. The "shamal" blows from the northwest over Khuzestan and coastal regions of the Persian Gulf from February to October, most intensely in summer. These summer winds undoubtedly contribute to the desiccation and, in some cases, filling-in of water courses. In the south the winds are west and southwest.

Temperature varies greatly over Iran with latitude and altitude, as well as with the seasons. Winter lows are found in January and summer highs in July in general, with the Zagros and Alborz mountains and the Caspian shore having maximum temperatures in August as a result of the influence of altitude and the sea. The mean monthly temperatures for January at 15 selected stations across Iran (Ganji, 1968) had a range of -1°C to 20°C, average about 8°C. For July these figures are 25 to 37°C, average 30°C. The annual range is 14C° at Jask on the Sea of Oman and 30.5C° at Mianeh in East Azarbayjan. Outside the coastal areas of the Caspian and Gulf, the annual range is considerable, and daily ranges also are large. Nights can be very cold in the northeast, less so on the plateau. Some areas, like the Khuzestan plains, have maximum temperatures over 50°C (53°C at Gatvand near Dezful; possibly over 55°C in the interior, hotter than anywhere else on earth) in summer while in the northwest in winter the temperature can fall below -30°C (to a low of -36°C at Bijar in Kordestan). Five temperature provinces have been delineated for Iran: the Caspian zone along the littoral which has a low annual temperature range; the Persian Gulf zone which has a low annual range but high values; the Zagros zone with a much higher range than the first two zones and a very low January mean; the Alborz zone which is similar to the Zagros but has higher temperatures and a greater range; and the interior zone with the greatest annual range coupled with relatively high values.

Precipitation falls in winter as snow on the mountains of the north and west. The highest mountains remain snow-covered year round. The plateau also receives snow but it does not last long and there is no snow along the Persian Gulf coast. Rain falls mainly in November to May with a mean annual of 416 mm, although the Caspian littoral is much higher and the interior plateau much less. Rain is uncommon from May to October over most of Iran. Maximum rain is found on the outward slopes of the Alborz and Zagros ranges where the mean annual rainfall is more than 1200 mm, 1950 mm at Anzali. The plateau has less than 120 mm annually, Sistan less than 70 mm, and Mirjaveh on the Pakistani border only 48 mm annually. The Caspian littoral has rain in every month at some localities. The plateau receives most of its rain in spring, the Caspian in autumn, and the Gulf coast in winter. The result of this pattern of rainfall is heavy runoff in spring with silt-laden floods and erosion a feature. Many streams marked on maps are actually dry for much of the year. Even a major, interior basin river like the Zayandeh which flows through Esfahan does not reach its terminal basin for much of the year.

A review of modern and historical floods in Iran can be found in Mazra'eh, News, Analytical and Educational Monthly, No. 10, January 1998 at www.netiran.com/Htdocs/Clippings/DEconomy/980100XXDE05.html. Devastating floods occurred in 2001, after several years of drought, in Gilan, Golestan and Khorasan provinces (IRNA, 11 August, 14 August, 4 September 2001).

Droughts occur and can be devastating for fish habitats. The drought years 1999-2001 were the worst in 30-40 years and resulted in a United Nations Technical Mission (see ReliefWeb, 22 August 2000, UN Office for the Coordination of Humanitarian Affairs (OCHA) at www.reliefweb.int; Foghi, 2004). Various effects were noted including the drying of 2500 qanats in Yazd, in southern Fars groundwater became saline, the Latian, Lar and Karaj dams near Tehran had water reserves of 51 million cu m, down from 173 million cu m for the same period in the previous year and were within about 2 months of drying up, several lakes and wetlands of international importance dried out (Bakhtegan-Neyriz and surrounding wetlands, Hamun-e Saberi, south end of Hamun-e Puzak and Gav Khuni), rivers dried completely (Hirmand River and its terminal lake), the Dez and Karkheh rivers in Khuzestan were depleted by 70% in 2001, water rationing was implemented in Tehran and 30 other cities, and lower water levels in rivers that retained flow had reduced oxygen affecting fish (IRNA, various news reports, 2001). In East Azarbayjan, 190 ha of 220 ha used for fish breeding were useless through drought (IRNA, 29 August 2001). Marshes south of Lake Orumiyeh near Mahabad encompassing 30,000 ha dried up (IRNA, 25 August 2001). Water reserves behind dams in Khorasan were depleted by 65% in 2001, the precipitation rate having declined by 40% in the period November 2000-August 2001 (IRNA, 3 September 2001).

Abbaspour and Sabetraftar (2005) reviewed Iranian drought cycles and found arid conditions were experienced for 13 of the previous 23 years. Drought affected fishes in the drying of wetlands where hundreds of thousands of fish died, in Sistan 8-12,000 tons of fish were lost as the lakes dried up, in Fars fish losses were reported from the Kor River, in East Azarbayjan 174 ha of fish culture farms were damaged, and rivers draining to the Persian Gulf lost fishes including migratory species.

The nature of the drainages of Iran is directly related to climate. The Alborz Mountains in the north block movement of moisture to the south while the Zagros Mountains in the west block moisture from that direction. The southeast monsoon is almost completely dry before it reaches eastern Iran. In consequence the best watered parts of Iran lie on its northern and western fringes and the interior becomes drier from west to east and north to south. Interior rivers exist in large part because of mountain ranges which store water as snow, in the case of the Hirmand River and the Sistan lakes, far removed from Iran.

There has been many studies on past climates in Iran and neighbouring countries, attempting to link climate with past environmental conditions in the Late Pleistocene-Holocene. The Early to Middle Pleistocene, however, is practically unknown for the Middle East and is not dealt with here (Butzer, 1978). Past environments have significance for fish habitats, distributions and zoogeography. The brief summary below is based on Butzer (1957, 1958a, 1958b, 1961, 1975, 1978), Bobek (1959), Whyte (1961), Hutchinson and Cowgill (1963), van Zeist and Wright (1963), van Zeist (1967), Wright et al. (1967), Krinsley (1970), Diester-Haass (1973), Turnbull and Reed (1974), Nützel (1976), van Zeist and Bottema (1977, 1982), Wright (1977; 1983), Ganji (1978), Neumann and Sigrist (1978), van Zeist and Woldring (1978), Woosley and Hole (1978), Farrand (1979), Storch (1980), Coad (1980c), Kay and Johnson (1981), Lamb (1982), Neumann (1993), Qin and Yu (1998); Griffiths et al. (2001); Stevens et al. (2001); Snyder et al. (2001); this being by no means an exhaustive listing of the studies in this field nor is the below a critical assessment of conflicting views. Evidence for these past environments is taken from a number of studies in different fields. The Pleistocene ice has been gradually withdrawing from its last maximum at 20,000 B.P. and the remains of ice fields and glacial moraines can be used to determine former conditions such as the snowline. The advance and retreat of deserts and the use and abandonment of settlements are indicative of changes. Such erosional physical features as dry riverbeds and other riverine structures, alluvial fans, sand dunes, and aeolian deposits all give clues to environmental change. The extent and level of lakes and playas have been widely studied as indicators of climatic fluctuations. Pollen and other organisms associated with lake sediments can be used to trace changing conditions and finally historical records can be analyzed.

Glacial deposits in the outward slopes of the Zagros and Alborz mountains indicate that the snowline was 600-800 m lower than today, perhaps as much as 1800 m in some areas, and as much as 1500 m at Shir Kuh near Yazd and Kuh-e Jupar near Kerman in south-central Iran. Lowered snowlines cannot be explained by temperature alone but were probably due to much greater precipitation. Winter would have been longer and colder in the Pleistocene, more snow would accumulate and summers may have been cloudier. The runoff period would have been longer and river habitats could have been less prone to desiccation in late summer.

The climate in the Zagros Mountains of the late Quaternary in Iran has been examined by means of sediment analyses from lakes Zaribar and Mirabad and for nearby Turkey at Lake Van. Pollen, chemistry, sediments, diatoms, cladocerans, ostracods and palaeobotany all confirm geological studies. The last glacial maximum (the Würm) at about 20,000 B.P. led to local glaciation, a depression in the snow line and absence of trees. The climate was cool and relatively dry, with less precipitation than today. The cooler temperatures meant less evaporation, more runoff and filling of intermontane lakes. The Caspian Sea and Lake Orumiyeh were much larger than today, being 78 m and 55 m higher. As the glaciers receded, the land environment or life zones moved up the mountains. The significance of this for fishes is unknown; there were few trees and the environment may have resembled modern denuded conditions. There may have been a higher flow than later when trees developed to hold runoff and before man chopped them down. However bushes could have retained water and reduced silt load in rivers. By 12-14,000 B.P the evidence from Zaribar and Mirabad indicates a warming climate but without increased precipitation. Indeed rainfall may have been less than today, reducing river flows and perhaps habitats for fishes. This arid period was succeeded by a more humid period. An increase in precipitation at Lake Van did not take place until 6500 B.P., about 4000 years later than in western Iran. Climate changed not only through time but also geographically, just as today. Regional variations mask general statements about earlier climate for Iran and the outline given here is perhaps best seen as indicative that change occurred. The humid period was followed by a period of less rainfall, and then in the late Holocene by an increase in rainfall. The last 3000 years have been humid with perhaps two, short, arid episodes. Southern Iran may have been cool and comparatively moist when the highlands were moderately cold and relatively dry. Climate probably changed markedly over short periods. Short cold phases are recorded for Europe in the last several thousand years, e.g. from about 1400 to 1230 B.C., associated with rises in lake levels. Similar events may occurred in Iran. Barley harvest dates in Babylonia derived from clay tablets indicate they were 10-20 days earlier in the period 1800-1650 B.C. and 10-20 days later in 600-400 B.C. It is concluded that the former period was warmer and the latter cooler than today.

Pluvial conditions as recognised for more northerly areas of Europe probably did not occur in Iran during the Pleistocene although summers may have been less dry because of greater cloudiness and lower temperatures and evaporation. Lake levels were probably higher 18,000-20,000 years ago (Roberts and Wright, 1993). Krinsley (1970), in his study of playas in Iran, concluded that the climate was semi-arid rather than pluvial in the period of maximum cold during the Pleistocene. Lakes, which occupied endorheic basins and could have facilitated local fish movements, dried up as the climate warmed with the retreat of ice sheets and glaciers and evaporation exceeded precipitation. These shallow lakes were found along the inner mountain front or within basins which received greater discharges. As distance from the mountains increased, there were only intermittent lakes and finally playas. An immense lake filling much of central Iran, as proposed by earlier authors, seems unlikely. Generally conditions over Iran appear to have varied as much, if not more, in the Pleistocene as they did in recent centuries through the agency of man. Conditions 9000 years ago were probably drier than today (Roberts and Wright, 1993). The fishes may have been selected for an ability to survive highly variable conditions in terms of stream flow, temperature, silt load, local fluctuations in lake levels and salt content, etc.

The greenhouse effect is apparent in Iran, a rise in temperature caused by various man-made and released gases. Nasrallah and Balling (1993) show a temperature increase of 0.09-0.23C°/decade, mean 0.18C°/decade, from 1950-1990.
 


Habitats

The major rivers of Iran drain the two mountain chains which retain enough snow or collect enough rainfall to ensure a constant and appreciable flow. Afshin (1994) summarises the rivers of Iran. All rivers in Iran are fordable on foot when not in spring flood with the exception of the Aras and Safid rivers of the Caspian basin, the Hirmand river of Sistan and the large rivers of Khuzestan. Most rivers marked on maps are in reality small streams, with very shallow and clear water. There is little vegetation on the banks, and fishes, if present, can be seen with ease. A significant proportion of fish habitat is occupied by small streams, springs and qanats. Large freshwater lakes or marshes are absent except in Sistan, the Caspian basin and the plains of Khuzestan. Most large lakes on maps are salty and do not support a fish fauna. A number of dams have been built and more are planned (see Bagley (1976), Coad (1980c) and "Aquastat" from the Food and Agriculture Organization, Rome (http://www.fao.org/ag/agl/aglw/aquastat/iran.htm)) and these form important lacustrine habitats. In 1994, 27 storage dams were in operation with a capacity of 39.2 km3 and a further 24 were under construction with a capacity of 11.5 km3 (see also below for more on dams). In 2002 Iran was building 68 dams and the construction of a further 120 dams were being considered as 33% of the country's water resources were wasted (IRNA, 2 January 2002). Manouchehri and Mahmoodian (2002) briefly review environmental impacts of dams in Iran.

The streams may have their origin in a mountain, a spring or a qanat, but they hold in common a clarity of water, a bare pebble bed, small dimensions (one to a few metres wide and a few centimetres deep) and often a short course. They may join another stream but are often lost in marshes, tapped for irrigation and lost in fields or become absorbed by the friable and porous ground. Many streams are intermittent, with flow near their mountain source, dry sections and perhaps a flow near their mid-course, with subsequent absorption into the ground. Heavy aquatic vegetation is not common and most plant material is a thin encrusting layer on the bottom. Banks are often bare of riparian vegetation and streams are fully exposed to insolation. Summer temperatures are often high as a result (30°C and more) yet at higher altitudes streams can be icy cold even in summer and the typical blue-grey of snow-fed water. Spring floods can be disastrous, scouring out the stream beds and dumping heavy silt loads (Melville, 1984). Spring fed streams of shorter course are not affected because they have a small catchment area and may well provide a refuge for fishes. The clean water of springs attracts human settlement and these waters are often blocked off to form ponds or cisterns with water led off through artificial channels subject to drying as requirements change. Streams and rivers may also be impounded, forming small ponds or lakes. Bridges often have small pools beneath them and this may be the deepest (at ca. 1 m) and most shaded section of a stream.

Marsh areas may be associated with springs. Reeds and other vegetation develop downstream of the source and may be quite extensive, occupying several square kilometres. Some areas of marsh are ponded and provide habitat for larger species as well as shelter for young. Extensive marshes, lakes and lagoons are developed in Sistan, the Caspian basin and Khuzestan, all fed by major rivers (50+ m wide and 3+ m deep) draining vast areas of land. These areas vary widely with season and flood dramatically in spring, inundating vast tracts of land. The rivers and associated marsh-lake complexes provide the major freshwater food fishing areas in Iran. The Sistan marshes have been described in Annandale (1921) and Annandale and Hora (1920), the Caspian shore by Schüz (1959) and the lowlands of southern Iraq by Rzóska (1980) and by Thesiger (1985) and Young (1989).

Conservation of aquatic habitats in Iran has been part of a general programme for biotic conservation summarised in Firouz (1974; 1976), Firouz and Harrington (1976), Ashtiani-Zarandi (1990) and Kahrom (2000). The Ramsar Convention on Wetlands of International Importance was named after the city of Ramsar in northern Iran where the first conference was held in January 1971. Iran has more Ramsar listed sites than any other country in Southwest Asia (Scott, 1993). In 1977 there were 11 Park-e Melli (National Parks), 4 Asar-e Tabii Melli (National Nature Monuments), 24 Manatgheh-Hefazat Shodeh (Protected Regions or Areas) and 31 Panahgah-e hayat-e Vahsh (Wildlife Refuges) offering varying degrees of protection to the fish fauna (Firouz et al., 1970; Yachkaschi, 1976; Köpp and Yachkaschi, 1978; Majnunian, 1985). The 1993 United Nations List of National Parks and Protected Areas at "www.wcmc.org.uk/data/database/un_combo.html lists" 7 National Parks, 2 National Nature Monuments, 41 Protected Areas and 18 Wildlife Refuges and the National Report of the Islamic Republic of Iran for the Convention on Biological Diversity (Department of the Environment, Tehran) lists 11 National Parks, 47 Protected Areas, 25 Wildlife Refuges, 5 National Nature Monuments, 9 MAB (Man and Biosphere) Sites and 20 Ramsar Sites.

Seven Ramsar sites are priorities for urgent action with the causes, namely:- Alagol, Ulmagol and Ajigol lakes (impact of agricultural development), the Anzali Mordab (Talab) complex (falling water levels and increased eutrophication leading to the rapid spread of the reed Phragmites australis, south end of Hamun-e Puzak (water inflow could be reduced because of dam construction in Afghanistan), Hamun-e Saberi and Hamun-e Hirmand (dam construction in Afghanistan), Neyriz lakes and Kamjan Marshes (drought and agricultural activities), Shadegan Marshes and mudflats of Khor al Amaya and Khor Musa (chemical pollution from the Iran-Iraq war), and Shurgol, Yadegarlu and Dorgeh Sangi lakes (war and drought effects) (www.ramsar.org/ram_rpt_37e.htm, downloaded 28 July 2000).

The status of the fish fauna in Iran was assessed by Coad (1980c) and Kiabi et al. (1999) and compared with other areas by Moyle and Leidy in Fiedler and Jain (1992). The percentage of the total fauna under some form of threat was assessed at 22%, a figure which was lower than most other areas examined.

Iran has several unusual habitats for fishes and these are described below.

i) Hot springs

A number of hot springs are reported from Iran (Waring, 1965; Joneidi et al., 1971?; www.bibliothecapersica.com/articlenavigation/index.html, under ab-e garm, downloaded 24 December 2004). Some of the hot springs marked on maps are not hot, e.g. the spring at Tafresh (ca. 34°44'N, 50°02'E) was only 19°C (and fishless). Some springs produce water at relatively high temperatures, but since these temperatures are also seen in nearby streams they are not regarded as "hot", e.g. a spring near Farrashband (28°53'N, 52°06'E) at 30°C.

Only the true hot spring at Genu (27º26'N, 56º20'E) is known to contain fish including Aphanius ginaonis, Cyprinion watsoni and Garra persica (Coad, 1980b). A hot spring on the slopes of Kuh-e Bazman (the mountain is at 28°04'N, 60°01'E) is rumoured to contain tooth-carps (Cyprinodontidae).

The Ab-e Garm (literally hot water) at Genu emerges at 41°C and was partially enclosed by brickwork associated with a hammam or bath-house. The altitude of the spring is about 400 m. Its stream is 10-15 m wide near the source and the bed is composed of stones and pebbles covered by lime-green algal mats and strings. Only Aphanius ginaonis was found at the hot spring, not in the main flow but along the stream margins and in many minor subsidiary springs which emerge a few metres from the main spring. These minor springs had a mud bottom, were as shallow as 1 cm and had soap and food debris pollution in 1977. Side springs and stream margin near the source were 37-40ºC. The other species (along with A. ginaonis) were found below a cascade and have no access to the hotter parts of the spring and stream. A. dispar is recorded from the spring by Werner (1929) but this has not been confirmed by my collections. The water is clear and colourless, but there is a strong smell of sulphur. Flow is 30 l/sec. The chemistry of this spring as given by Joneidi et al. (1971?) was : pH = 6.2, conductivity 14,000 us, dry residue at 180ºC = 9933 mg/l, H2S = 34 (? p.p.b.), r (reacting value) Ca = 22.4, r Mg = 9.9, r Na + K = 6.1, total cations 162.1 (sic), r Cl = 147, r SO4 = 15.4, r HCO3 = 4.6, total anions = 166 (sic), SiO2 = 10 mg/l, NH4 = 0.7 (no units given), NO3 = 22 (no units given). There were traces of CO2 and no measurable Fe, NO2, or CO3. The hot spring lies in the Genu Protected Area (Biosphere Reserve) which is described by Zehzad et al. (1997).

ii) Caves

Iran is replete with caves but thus far only one has been found to contain a fish fauna. This cave lies about 12 km north of the railway station Tang-e Haft in Lorestan at 33°05'N, 48°36'E. Two species are found here, Iranocypris typhlops (Cyprinidae) and Paracobitis smithi (Nemacheilidae) (Bruun and Kaiser, 1944; Movaghar, 1973; Greenwood, 1976; Smith, 1978; 1979; Coad, 1996c; Proudlove, 2001; Romero and Paulson, 2001). The cave lies in the Dez River drainage of the Tigris River basin and its connection to nearby surface water is intermittent. The cave is the surface outlet of a subterranean limestone system and the captures may represent strays from underground. B. Sandford (pers. comm., 1979) stated that there is some evidence of recent collapse in the cave system and thus the habitat may be endangered but it is difficult to assess the extent and nature of underground fissures in the rock.

iii) Qanats

Qanats are an unusual yet important habitat for fishes in Iran. An account of their fishes with an extensive bibliography is given in Coad (1996h); additional literature on this unique environment not referenced there includes Kuros (1943), Aisenstein (1947), Feylessoufi (1959), Nesbitt and Bawa (1960; 1961), de Menasce (1966), Jentsch (1970), Nadji (1970; 1972a; 1972b), Braun (1974), Goblot (1979), Hartl (1979), Sajjadi (1982), Goldsmith and Hildyard (1984), Behnia (1988), McLachlan (1988), Beaumont et al. (1989), Harwit (1990), Razavi (1991), Coad (1994b), Koocheki (1996), Liaqati (1997), Salim Manshadi et al. (1997), Afkhami (1998), English (1998), Aminpouri (2002), www.netiran.com/Htdocs/Clippings/DEconomy/200629XXDE05.html, downloaded 8 August 2002), Foltz (2002), Floor (2003), Wessels and Hoogeveen (2003), and qanats at www.waterhistory.org, and at www.bibliothecapersica.com/articlenavigation/index.html, under abyari (irrigation), downloaded 24 December 2004.

The word qanat has various suggested origins including a derivation from the Akkadian for "reed" according to Goblot (1979) in contrast to others listed in Coad (1996h).

Over 20% of the irrigated area of Iran is fed by qanats (Redding and Midlen, 1991) and numbers as high as 60,000 have been estimated. They are essentially horizontal wells which tap groundwater and provide a continual, low gradient flow of fresh water. Qanats are an advantageous habitat for fishes in several ways. The water temperature is not subject to the extremes found in natural waters, shade within the qanat provides protection against predators on adults, young and eggs and against insolation, the gradient and water flow are gentle, and a certain amount of food is provided by kitchen scraps since dishes, cooking containers and implements are washed in the jube or channel and food is cleaned and trimmed there. A school of fish will quickly gather at a washing site and maintain station in clouds of detergent in order to pick up scraps of food. Attempts to imitate washing movements will attract fish momentarily but they soon dart off when no food is forthcoming. The garden environment with trees and other vegetation provides shade, energy input from leaf fall and garbage items, and facilitates development of an invertebrate fauna as a food source. Aufwuchs on rock surfaces provide a food source along with the associated invertebrate fauna. The Zoroastrian community, once widespread in Iran, has a ceremony known as com-e mahi or "meal for the fishes" in which bread and dried fruit are thrown into running water as a libation (Boyce, 1977). Feeding of scraps to fish is also seen in Moslem communities and boys regularly attempt to attract and catch fish using any available food material and primitive fishing gear (personal observations; Edwards, 1971).

Qanats are now rapidly being replaced by pump-wells which are faster and easier to excavate but do not provide fish habitat. Pump-wells often dry up qanats and natural springs by lowering the water table (Razavi, 1991; Anonymous, 2001b; Aminpouri, 2002). Also schemes to restrict water flow from qanats for conservation reasons will presumably affect the available habitat for fishes (Salim Manshadi et al., 1997).

The qanat fishes comprised 25 species in Coad's study (1996h), 40% of the fauna on the plateau of Iran. The number of species per qanat ranges from 1 to 6 although 88% of qanats have only 1-2 species. Areas with little surface water and low in diversity have 94% of the species occurring in qanats while better-watered areas with more diversity have only 29% of species in qanats. The qanat fauna is dominated by the Cyprinidae, which comprises 76% of the ichthyofauna. The qanat fauna is a subset of the basin in which the qanat occurs, comprising small species, broadcast spawners, lacking in specialised food requirements (usually scrapers of aufwuchs or feeding on invertebrates), non-migratory, and widely tolerant of environmental conditions.

The fishes in qanats are caught by local people for food but given the restricted size of this habitat and of most fishes found in them, this is not a significant dietary item. In the seventeenth century qanat fishes "were not esteemed as they never saw the light and were used only for medicinal purposes to cause vomiting" (Ferrier, 1996, quoting Jean Chardin). In the 1950s villagers in Iran believed that qanat fish lived forever and needed no food, only their own eggs (www.iras.ucalgary.ca/~volk/sylvia/qanat.htm, downloaded 24 June 2002).

Colonisation is both natural, since loaches are unlikely to be seen and caught by local people, and deliberate, since larger cyprinids are found in qanats remote from any surface water. These fish are hardy, already living in high temperature environments, and are easily transported for Now Ruz celebrations. At the Zard-Abieh qanat in Shahrud, a local man remembered putting fish into the qanat 60 years ago from one now dry (H. Rahimian, pers. comm., 2000).

iv) Salt streams and lakes

Salt lakes are common in Iran and are mostly too saline to support a fish fauna. They are discussed in a world context by Williams (1996). Fishes do exist in tributary streams (which may be saline in varying degrees). Rivers and springs around salt lakes are therefore isolated from one another and might be expected to give rise to unique populations of fishes. However all these salt lakes are shallow and liable to desiccate such that tributary streams and springs can connect and allow faunal interchanges once the lake level falls.

Many streams in Iran are highly mineralized or even salt to taste yet these support fishes which are usually regarded, at least at the family level, as salt intolerant. Salinity tolerance studies have not been carried out on Iranian fishes. The Caspian Sea is at one-third sea water (12-13‰) yet typical "fresh" water species can be found there, e.g. Cyprinus carpio.

v) Sacred waters

A number of springs in Iran are said to be "sacred" and their fish then attain a degree of importance on account of their inaccessibility to ichthyologists. Howz or tanks at Qumisheh (32°01'N, 51°52'E) were supposed to hold sacred fish, decorated with gold rings, according to John Fryer in 1698 and John Chardin in 1711, but G. N. Curzon in 1892 mentioned that the gold rings were gone and by 1978 so were the fish. A sacred tank or artificial reservoir at Soh contained fish deemed to be holy. Visitors were expected to purchase bread to feed these fishes (Anderson, 1880).

The most important "sacred" fish are those of Sa'di's Tomb in Shiraz (29°37'N, 52°35'E) which were described by Heckel (1849b) as new species Scaphiodon saadii (= Capoeta damascina) and Discognathus crenulatus (= Garra rufa). The water is a stream (?qanat) under the tomb and part is expanded into a hawz-e mahi or fish pond. Fish have been present here since at least the early nineteenth century as they are mentioned briefly by Waring (1807). Official permission was gained to collect fishes in Sa'di's tomb for study but sampling was actively discouraged by local people. Sa'di was supposed to punish any killing of these fishes with death but the traveller Chardin was able to catch some to eat by monetary means. Some of these fish too were reputedly decorated with gold rings (Ouseley, 1819-1823); regrettably my captures were not.

vi) Mordab

A mordab is a fresh or brackish water lagoon area found along the Caspian Sea coast (literally "dead water", the Russian equivalent is liman). The Anzali Mordab at 37°26'N, 49°25'E is the best known (Firouz, 1968b) and was formerly called the Pahlavi Mordab. The more modern term is "talab" (= pool or marsh, which lacks the association with death) but the older literature refers to mordab and the term is still in common use. The Anzali Mordab is about 30 km long and 4-8 km wide with clear water of only 1.5 m average depth. Much of the area is covered by Phragmites reeds and other plants and only about 15% is open water. Variations in Caspian Sea level and water abstraction from feeder streams will affect the mordab level and size. In the 1930s the mordab was 4 to 8 m deep (Vladykov, 1964) and the fall in level has severely affected the spawning migrations of fishes and the habitat for developing young. The mordab is the principal breeding ground for Rutilus frisii kutum and is also important for several other species. Further details are given below under the description of the Caspian Sea basin.

vii) Wetlands

Wetlands were originally studied and protected as feeding and overwintering grounds for important waterfowl but they do protect fish populations which might otherwise be threatened. Access and hunting is forbidden or restricted and often fishing too. Anonymous (1971), Carp (1972) and Dugan (1993) list and describe various wetlands in Iran of international importance principally:-

See also Scott (1995) where latitude-longitudes are often slightly different.

Lower Atrak River and Alagol Lake (37°21'N, 54°35'E)

Farahabad and Larim Sahra (36°45'N, 53°05'E)

Zarrin Kola (36°43'N, 53°00'E)

Bisheh Sar (36°36'N, 52°43'E)

Fereydun Kenar (36°40'N, 52°31'E)

Bandar-e Farahnaz Lagoon (37°25'N, 49°57'E)

Khalij-e Gorgan (36°50'N, 53°40'E)

Anzali Mordab (37°25'N, 49°30'E)

Nur Gol (38°00'N, 48°33'E)

Neyriz Lakes (29°30'N, 53°40'E)

Lake Parishan (Famur)(29°26'N, 51°50'E)

Khuzestan Marshes (30°30'N, 49°30'E)

Dasht-e Arjan (29°35'N, 52°00'E)

Lake Kopibalbalch, Hassanlu Marsh, Yadergarlu Marsh and surrounding marshes (37°00'N, 45°30'E)

Lake Bishovan (37°09'N, 54°52'E)

Amirkelayeh (37°17'N, 50°12'E)

Coastal lagoons north of Gomishan (37°15'N, 54°00'E)

Seyed Mahalleh (36°45'N, 53°00'E)

Sistan lowlands (31°00'N, 61°10'E)

Additional wetlands not of international importance were listed as follows:-

Safid Rud Reservoir (36°45'N, 49°24'E)

Astara (38°25'N, 48°50'E)

Bahr-e Zaribar (35°32'N, 46°07'E)

Soltanabad Marshes (29°30'N, 52°35'E)

Lake Maharlu (29°30'N, 52°50'E)

Dasht-e Mogan (39°30'N, 47°30'E)

Araxes River (39°10'N, 45°20'E)

Agh Gol (39°55'N, 44°47'E)

Rud-e Shur (35º50'N, 50°25'E)

Zarghan and Lapu'i Marshes (29°50'N, 52°50'E)

Hasanzadeh Kiabi et al. (2004) list the top 13 wetlands as Choghakhor, Mand River, Hamun-e Saberi-Hirmand, Khorekhoran, Gandoman, Orumiyeh, Hawr al Azim, Gorgan Bay + Miankaleh + Lapoo, Shadegan, Helleh estuary, Anzali, Arghan of Parishan, and Hamun-e Puzak.

Other wetlands are mentioned in the appropriate drainage basin account.

The biotopes of Iran are summarised in the figure below:-

Biotopes of Iran from Wikimedia Commons.

Biotopes of Iran (and adjacent countries)
 
   Central Persian desert basin
   Forest steppe
   Eastern Anatolian forest steppe
   Elburz forest steppe
   Hyrcanian Caspian forests
   Kopet Dag forest steppe
   Kopet Dag semi desert
   Desert Caspian lowlands
   Azerbaijan steppe
   Middle East steppe
   Mesopotamian desert
   Arabian desert
   Tigris-Euphrates salted alluvial marshes
   Persian Gulf desert
   South Persian desert and semi-desert
   North Pakistan sandy desert
   Central Afghan woodlands
   Oman Gulf desert and semi-desert
   Baluchistan Woodlands


 


Environmental Change

There is evidence for changes in the environment and therefore fish habitats during historical times. Many of these changes are man-made and are on-going. References to change, e.g. habitat loss, pollution, effects of exotic species, etc., can be found in each of the Drainage Basin files. This topic has been reviewed in general by Coad (1980c) and the references therein, with papers cited under Climate above, are relevant.

Peritore (1999) gives a general overview of ecological conditions and attitudes to the environment, Foltz (2001) reviews environmental initiatives, Afasiabi (2003) reviews the environmental movement in Iran and Valeolahy (2000) reviews the factors affecting the abundance of fishes and suggests measures for conservation. Jawad (2003) gives an account of the impact of environmental change on Iraqi fishes which has implications for fishes in neighbouring waters of Iran.
 


Drainage Basins

The drainage basins of Iran are shown in the Figure. The delimitation of these basins is somewhat arbitrary. Iran is a mountainous country and much of it is desert. There are thousands of small springs and streams with no present or recent connection to other water bodies. Practical considerations require a large scale and I have divided the country into 19 major basins based on field work, maps, fish distributions, history of research, works on hydrography and areas deemed important for an understanding of zoogeography.

There are two main types of basin, exorheic where the rivers and lakes drain to the sea and endorheic, where rivers drain to an internal basin such as a lake, or are lost in the desert, and have no connection with the sea. The exorheic basins all fringe the southern part of Iran. The bulk of the basins, in number (15) and area (about 78.1% of Iran), are endorheic. These plateau basins lie at an average altitude of 800 m, alternating with mountains ridges at an average of 2000 m. The salt lakes and flats of these basins are fed primarily by groundwater rather than rain (Issar, 1967) and water is lost by evaporation. Wolfart (1987) makes the valuable point that Quaternary environments in the closed or endorheic basins of arid Southwest Asia often have marine and brackish fossils. These are not evidence of marine invasions but of the increasing salinity derived from the mineral content of rainwater. As the water evaporates it leaves behind the minerals and over ten thousand years or less a saline environment develops.

www.bibliothecapersica.com/articlenavigation/index.html, under drainage, downloaded 24 December 2004 gives four main drainages for Iran as follows:-
 
Drainage Area (sq km) %
Caspian Sea 193,161 11.9
Lake Orumiyeh 54,747 3.4
Persian Gulf 335,864 21.9
Interior 1,626,520 61.8
Total 2,210,292 100

with the interior drainages as follows:-

Drainage Area (sq km) %
Qom (Namak Lake) 92,332 9.0
Damghan 19,863 1.9
Dasht-e Kavir 200,747 19.6
Mashhad (Tedzhen River or Hari Rud) 43,496 4.3
Bejestan Highlands 91,349 8.9
Dasht-e Lut 166,160 16.2
Sistan 90,813 8.9
Jaz Murian 75,193 7.4
Yazd 105,291 10.3
Esfahan 97,802 9.6
Zagros Mountains (Tigris River) 39,702 3.9
Total 1,022,748 100

Van der Leeden (1975) summarises water resources of Iran with discharges of principal rivers at various recording stations, lists of major dams and reservoirs, and resources and demand. www.bibliothecapersica.com/articlenavigation/index.html, under ab (= water), downloaded 24 December 2004 also lists major dams and gives a general overview of hydrology and has descriptions of various rivers under their names. McLachlan (1988) also considers water resources in Iran. Some of the earlier dam projects are described by Justin and Taleghani (1955). Later dam projects can be located by a search at "Netiran.com". Prior to the Islamic Revolution 13 dams had been built in Iran but the five-year development plan (1990-1995) designed 110 dams of which 22 were under construction in 1993. 60 dams have been constructed after the 1979 revolution (IRNA, 31 August 1998).

"Aquastat" from the Food and Agriculture Organization, Rome (www.fao.org/ag/agl/aglw/aquastat/iran.htm) gives an overview of Iranian water resources and water abstraction and is updated at intervals. The total domestic, industrial and agricultural water abstraction was estimated at 70 km3 in 1993, 51% of the renewable water resources. Annual abstraction from aquifers (57 km3) is more than the estimated safe yield of 46 km3. An additional 39 km3 is used annually, 20 km3 for electricity production, 11 km3 for flood control and 2 km3 for control and thence environmental protection of downstream parts of rivers, the remainder being surplus. The increasing demands will have serious effects on the water supply and hence the fish fauna. Nikravesh (1997) estimates, based on water consumption and population growth, that Iran will be added to the U.N. list of countries facing water shortages in the year 2025.

Kuros (1943) gives accounts of historical water resources and the problems of water supplies in Iran. Lambton (1953) gives an account of the allocation of water resources in Iran for irrigation. This latter work is important for an understanding of restrictions on fish habitats, e.g. in qanats, reservoirs, rivers and springs. Beaumont (1981) reviews management of water resources in the Middle East and places the Iranian resources in a wider context. Anonymous (1961c) and Beaumont (1974) outline water resource development in Iran, the construction of dams, abstraction for irrigation by traditional and modern means, and the demands of industry and domestic consumers of water. All these affect the habitat of fishes, often in deleterious ways. Noori (1966) describes the hydrology of surface water in Iran. Pirnia (1951), Anonymous (1961c) and Beaumont (1973b) give accounts of the river regimes in Iran with discharges and runoffs at various recording stations. Peak discharges occur in March to May because of snowmelt. Very low flows occur in summer because of the lack of precipitation, and because of abstraction for irrigation, and flow is mostly from groundwater sources. Most rivers are really streams for much of the year as minimum flows for principal rivers are 0.16-451 cu m/sec, average about 36 cu m/sec. The Caspian rivers are the only ones which lack a distinctive annual rhythm and show flows closely related to precipitation throughout the year. The areas with the largest runoff values are in the northern and central Zagros Mountains and in the Alborz Mountains while lowest runoff values per unit area are found around the deserts in central Iran. In the Zagros and Alborz, annual runoff values can attain more than 300,000 cu m per sq km. Löffler (1956; 1961) studied the limnology of several of the major basins within Iran. The Ramsar Convention on Wetlands has a report on the Islamic Republic of Iran (No. 37, at www.ramsar.org/ram_rpt_37e.htm, downloaded 4 May 2001).

Peritore (1999) gives a general overview of ecological conditions and attitudes to the environment in Iran. Zohary (1963) gives a general account of the vegetation of Iran. A general description of Iran, its structure and drainage can be found in Harrison et al. (1945), Neumann (1953), Fisher (1968) and Krinsley (1970). Water policy development is summarised in Aminipouri (2002). A description of natural areas in Iran, including a list of National Parks and Protected Rivers, can be found in Zehzad et al. (2002). The Protected rivers are the Jajrud and Karaj in the Namak Lake basin, and the Chalus, Sardab, Lar and Haraz rivers of the Caspian Sea basin.

The basins are as follows:-

Exorheic Basins:- Gulf, Hormuz, Makran, Tigris River

Endorheic Basins: Bejestan, Caspian Sea, Dasht-e Kavir, Dasht-e Lut, Esfahan, Hamun-e Mashkid, Hamun-e Jaz Murian, Kor River, Lake Maharlu, Lake Orumiyeh, Namak Lake, Sirjan, Sistan, Tedzhen River, Kerman-Na'in

Exorheic Basins

Gulf

This basin comprises rivers which drain the southern Zagros Mountains to the head of the Persian Gulf, but which are not now tributaries of the Tigris River nor are they the salt streams of Hormuz. None of these rivers has a significant fishery. At its northern edge, the Zohreh River flows across the Khuzestan plains and is close to Tigris River tributaries. Other major rivers are the Helleh, which debouches into the Gulf north of Bushehr (28°59'N, 50°50'E) and the Mand or Qarah Aqaj (= the classical Sitakos), which, with its tributaries, drains much of Fars Province to the Gulf south of Bushehr. Near Shiraz it is known as the Qarah Aqaj or Kavar River. The Band-e Bahman, a weir or small dam on this river near Kavar, is probably pre-Islamic.

The Mand River is 480 km long and occupies a basin of about 60,000 sq km. Its flow is reduced by a low snow cover (although there can be torrential spring flow), water seepage, evaporation and abstraction for irrigation purposes. Discharge has been estimated to range from 10-2025 cu m (Merchant and Ronaghy, 1976). It is also polluted near Kavar (29°11'N, 52°44'E) by sewage and agriculture residues and does dry up to a series of isolated pools there. A fish kill, numbering in the many thousands, occurred in the Mand near Shiraz in 1977 and was attributed to chemicals used in spraying against malarial mosquitos. The people hired to spray village houses either dumped quantities of the chemical into the river to reduce their work load or washed out containers in the river (Coad, 1980c). Temperature range is at least 20C° between winter and summer. The delta of the Mand is a Protected Area of 46,700 ha. There are thin oxbow lakes and associated marshes

The Mand has a number of tributaries, at least two of which are called Shur (= salt) River. Conductivity near Firuzabad on the Shur River is 695-715 µM/cm but rises to 20,000 µM/cm below salt domes further downriver. The more southerly headwaters are close to those of the Shur River of the Hormuz basin between Darab (28°45'N, 54°34'E) and Fasa (28°56'N, 53°42'E). The headwaters of the Mand lie north-west of Shiraz near Kuh-e Tabask at 2318 m (29°52'N, 51°49'E) and there are a series of springs in this area called Chehel Chashmeh (= Forty Springs) which feed the Mand. Nearby is the Dasht-e Arzhan (29°39'N, 51°58'E), a small enclosed basin with a flooded plain encompassing about 24 sq km at maximum. It is fed by small springs and streams. The water is fresh since swallow holes in the southeast corner of the plain drain water away with a salt flushing effect. Shiraz was once "chiefly supplied with fish from this lake" (Ouseley, 1819-1823) but it does not now support such a copious ichthyofauna. A report from Reuters (8 June 2000) cites a fish kill numbering in the hundreds of thousands from the "Arjang lagoon, in a suburb of the southern city of Shiraz", presumably this lake, after it dried up (www.iran-sabz.org/news/fish2.htm). The Haft Barm-e Kudian lie about 20 km north of Dasht-e Arzhan at 29°49'N, 52°02'E at 2200 m. The seven lakes lie in rolling country and the largest is about a 1 sq km. Some may dry up in certain years but fish were found suggesting that there is a perennial water supply (Cornwallis, 1968a). Scott (1995) says the southern 5 lakes generally dry out completely in summer. In winter the lakes freeze over. They are about 2-3 m deep and some are slightly saline. These lakes have been stocked with Esox lucius, Hypophthalmichthys molitrix, Ctenopharyngodon idella and Gambusia holbrooki.

Surber (1969) gives some spot data on pH, total alkalinity, calcium-magnesium hardness, chlorides and free CO2 in the Mand basin. Near Firuzabad, the concentration of total dissolved solids is 333 mg/l while near Jahrom it reaches 6937 mg/l, indicating how there can be great variations in habitat within the same river basin over short distances, depending on local geology.

The Zohreh River and its tributary the Shul, are over 400 km long and have their headwaters near Kuh-e Barm Firuz at 3673 m (30°25'N, 51°58'E) whose northern flank spawns the Khersan River, a Karun tributary in the Tigris basin. Its basin is estimated to be 15,500 sq km. The Kowsar Dam at Gachsaran is 337.5 m high, its crest is 126 m and the reservoir capacity is 450 million cu m (http://netiran.com/news/IRNA/html/941126IRGG10.html). Gorjipoor et al. (2007) carried out a limnological investigation of the Zohreh River.

The Helleh River receives the Dalaki (205 km) and Shapur (231 km) rivers which drain the lower Zagros ranges west of Shiraz. Its basin is estimated to be 20,300 sq km (Shiati (1989) gives 10,000 sq km) and includes Lake Famur. Shiati (1989) gave an account of salinity in the rivers of this basin. Saline springs and salt domes increase the salinity about 10 times as the rivers flow down from the mountains. Total dissolved solids in the upper reaches of this basin are 366 mg/l, rising to 4219 mg/l in the lower reaches. Geological sources of sulphur also add to the chemical make up of these waters. There are no important sources of industrial pollution along these rivers but humans, domestic animals and agriculture are the main pollution sources. The levels of pollution are in the acceptable range (Gh. Izadpanahi, pers. comm., 1995). Aquaculture in the area (Helleh and Mand river basins) has not had obvious effects on coastal water quality (Omidi, 2006). The delta of the Helleh River is a complex of brackish and fresh marshes and lagoons with a maximum depth of 3.5 m. It is the largest freshwater marsh system on the Persian Gulf coast in southern Iran. It is designated as a Protected Area (42,600 ha). This area developed in the early 1970s when the main river channel was diverted onto the coastal plain.

A cave at Bishapur above the Shapur River is reputed to house a deep lake full of fish but this has not been investigated and may only be a local legend (Mounsey, 1872).

Endorheic Lake Famur, Perishan or Parishan (29°31'N, 51°48'E) is a particular feature of the Gulf basin which encompasses 42 sq km at about 820 m near Kazerun, is fed by about 80 fresh and brackish springs with a discharge of about 800 litres/second and supports a fish fauna near the springs. In years of heavy rainfall the fresh areas expand only to contract in dry years.

The ringing marshes are eutrophic and have halophytic plants of the genera Salsola, Kochia, Camphorosma and Halocnemum along with extensive reedbeds of Phragmites communis and Typha. This marshy shore attained 31°C in early June when air temperature was 43°C. Maximum depth is about 6 m, falling in summer to 3.87 m. pH is 7-8. The drainage basin encompasses about 290 sq km. Conductivity is 5 to 6,000 micromhos.

Södergren et al. (1978) recorded pollution in fish from this lake and the Shapur and Kupor rivers. Only small amounts of the organochlorine chemical p,p'-DDE were found in the lake but the rivers had very high levels of DDT and its metabolites DDE and TDE. At this time DDT was used for indoor spraying against malaria-infected mosquitos and insecticide containers were cleaned in the rivers after spraying.

An account of the lake is given in Farsi by Maafi (1996a; 1996b; 1996c). The lake is eutrophic and low concentrations of oxygen periodically cause fish mortalities. The reed beds are set on fire to increase the available agricultural land and this results in a sediment input with the consequent decrease in water depth, fingerling habitat destruction, and fish mortality through sediments clogging gills. Overfishing is also a problem. Wastewater and sewage enter the lake untreated and this enhances algal growth and eutrophication. Fishery ponds are established west of Lake Parishan resulting in exotic escapes. During periods of low rainfall, Parishan becomes a shallow saline lake and presumably fish habitat is limited to the immediate vicinity of freshwater springs.

Lake Parishan and the nearby Dasht-e Arjan (29°37'N, 51°59'E) are a Ramsar Site (World Conservation Monitoring Centre, 1990). They lie within the Arjan National Park and International Reserve which encompasses 65,750 ha as established in 1973. However the Park has been downgraded to a Protected Area of 52,800 ha with the Ramsar Site being the wetlands of Lake Parishan at 4200 ha and Dasht-e Arjan at 2400 ha (Khan et al., 1992). Dasht-e Arjan at 1950 m is a shallow, eutrophic freshwater lake fed by runoff, precipitation and the Salmon springs. The lake area in winter may be 1950 ha but shrinks in summer to a few hundred hectares. It dried completely in 2001. There is an outflow through swallow-holes in the south-east, traditionally linked to Lake Parishan. The lake margin and the spring-fed marshes have Phragmites communis, Typha and Juncus along with aquatic vegetation. Dasht-e Arjan is cooler than the environs of Lake Parishan because of its higher altitude - 15-35°C in summer and -10-15°C in winter as opposed to 22-40°C and 5-15°C.

As well as the rivers described above, springs and qanats are important in the Gulf basin. The Dalaki mineral springs have a temperature range of 30-38°C and a discharge of 200l/s. They are at 130 m above sea level and their hydrology, geology and chemistry is reviewed in Kompani-Zare and Moore (2001). The fishes in this area have not been investigated.

Rabbaniha et al. (2003) surveyed the larval ichthyofauna in the Farakeh Creek estuary area in the northern Gulf and found 15 families to be represented, Clupeidae, Gobiidae and Sillaginidae making up almost 94% of the catch.

The Shabankareh Dam is a diversion dam in the lower Helleh River basin and several other dams have been planned for this basin. Small canals or diversions are also present in this basin (Borowicka, 1958).

Berg (1940) places this basin, the Hormuz basin and the Makran basin as part of the Sind Province of the Indian Subregion of the Sino-Indian Region. Its eastwards extent is the lower and middle Indus River. The Iranian portion is called the Southern Iranian District. Small southern Iranian rivers belonged to a single river basin in the Pliocene, facilitating dispersal according to Berg.

Hormuz

The Hormuz (or Hormozgan) basin comprises a number of intermittent streams and rivers which drain to the Straits of Hormuz. None of the rivers has a significant fishery. The basin has a catchment of 55,800 sq km. Rainfall is low and sporadic at this southern end of the Zagros Mountains and streams are not always perennial. Qanats are an important feature and there is a hot spring (41°C) at Genu (27°26'N, 56°20'E) just north of Bandar-e Abbas. This area of Iran is rich in salt domes rising to over 1200 m above the surrounding land surface and consequently surface water is often contaminated and stream banks are rimed with salt (Lehner, 1944; Shearman, 1976; Kent, 1979). Some of the islands off this coast are salt plugs, e.g. Hormuz Island. Temperatures in winter are high in the lower streams, 15-33°C, and must be much higher in summer. These warm and saline streams are home to the endemic cichlid, Iranocichla hormuzensis, and so are distinguished from the fresh waters to the north, east and west. This species has been collected in the Minab River where my collections in the 1970s did not find it. The Minab River was therefore included in the Makran basin but may well form the easternmost part of this basin. However the possibility of an introduction of this species to the Minab cannot be ruled out.

Salt dome, Zagro Mounatins, from NASA and Wikimedia Commons.

Salt domes and salt glaciers, southern Iran, NASA and Wikimedia Commons.

Salt domes and salt glaciers, southern Iran, NASA and Wikimedia Commons.

 

The principal river is the Kul with its tributary the Shur (= salt) River. The upper reaches of the Shur lie west of Darab (28°45'N, 54°34'E) and mountains here exceed 3000 m. The headwaters of the Shur approach those of the eastern tributaries of the Mand River basin. The lower valleys parallel the coast and drain eastwards. The Rasul River is a tributary of the Kul, while the Mehran River drains directly into the sea. The Mehran delta lies in the Hara Protected Area (Biosphere Reserve) described by Zehzad et al. (1998). The offshore islands such as Qeshm, are poor in fresh water, but have not been explored. A number of streams cross the plain east of Bandar-e Abbas (27°11'N, 56°17'E) draining the Kuh-e Furgun at 3279 m and associated ranges. Although many streams are salty, a freshwater oasis is found at Sar Khun (27°23'N, 56°26'E).

Several islands in the Persian Gulf are included as part of this basin. The largest island is Qeshm but it lacks rivers although there are some small dams to collect rainwater runoff (A. R. Zeanaie, pers. comm., 1999). Species observed are Aphanius dispar, a mudskipper and the introduced Gambusia holbrooki. Water temperatures reach 32°C.

Qeshm Island and adjacent coast including the Mehran and Kul rivers (left and centre) (NASA and Wikimedia Commons).

Qeshm Island and adjacent coast, including the Mehran and Kul rivers (left and centre) (NASA and Wikimedia Commons).

Makran

The Makran is the coastal region of southeastern Iran between the Straits of Hormuz and the Pakistan border. In the west of this region the relief runs in a north-south direction parallel to the coast but from Jask eastwards the relief runs west-east, again paralleling the coast, to the Pakistan border. The rivers and streams of the Makran all drain to the sea at the Straits of Hormuz and the Sea of Oman. The inland Hamun-e Jaz Murian basin is isolated by mountain ranges reaching peaks in excess of 2000 m. The coastal drainages are often incised and the larger watercourses pass through tangs over 1000 m deep (Harrison, 1968).

I have not seen the watercourses between Jask and the upper Geh (= Nikshahr, Kaeyr or Kalar) River drainage (mouth is at 25°37'N, 60°08'E) but descriptions by Harrison (1941) indicate they are similar to other areas of Makran. It seems probable that only the Minab and Sarbaz Rivers have, or nearly have, a perennial and continuous flow along most of their course. Even these rivers are quite shallow and the Sarbaz in particular is easily fordable on foot along its entire length (ca. 280 km). The Minab River flows over a shorter course (ca. 220 km) than the Sarbaz, but has a greater flow regime. At Minab (27°09'N, 57°05'E) and at Rudan (27°26'N, 57°12'E) the Minab River was up to 100 m wide with an estimated maximum depth in pools of 2-3 m. The lower Sarbaz River was a series of shallow, muddy pools in the bottom of a canyon with some water flowing over sills connecting the pools (in early December 1977). The lower Sarbaz has been designated a Wetland of International Importance. In its middle and upper course the Sarbaz varied from a very shallow and narrow stream connecting pools (some of which were fishless) to what must be termed a river in the semi-desert environment of Baluchestan, with a width of 10 m, a depth of about 1 m and fast current. The rockfill embankment Pishin Dam built over the rivers Pishin and Sarbaz is 63 m high, has a crest length of 400 m and can store 175 million cu m of flood waters (http://netiran.com/news/IRNA/html/930418IRGG10.html).

The other streams of the Makran have little running water, often become isolated pools a kilometre or more apart, and regularly dry up along much of their length. Several rivers between the Mazavi (= Geru) River (mouth is at 26°56'N, 56°56'E) and the port of Jask are named and marked prominently on maps, but these were all dry in their lower reaches in late November 1976. Some flow in their upper reaches is to be expected, but its extent will depend on topography and recent climatic conditions. A dam and irrigation network is to be constructed on the Jaghin River east of Jask (IRNA, 26 June 2000).

Coad (1997a) combined the basins of the Makran, Dasht-e Lut, Hamun-e Jaz Murian, Mashkel and the Pakistani Pishin Lora as a single entity, expanding on earlier work by Mirza (1980). Mirza proposed the name Gedrosia for the Baluchistan Plateau west of the Central Brahui and Hala Ranges in Pakistan. The easternmost river along the Makran coast is the Hingol in Pakistan. East of this river the fauna becomes much more diverse at all taxonomic levels and the fauna is an Indus River one. In the north, the Pishin Lora River basin lies partly in Pakistan and partly in Afghanistan. Beyond this basin to the north and northwest lies the Registan Desert and then the Sistan basin, with its distinctive faunal mix including schizothoracines (Schizothorax, Schizocypris and Schizopygopsis) and a crested loach (Paracobitis rhadinaea). To the northeast lies an area designated as Yaghistan by Mirza (1980), with its unique faunal association. The westernmost river is the Dasht, whose upper reaches cross the Iranian border. The western limit of Gedrosia is the Mashkel River basin which has several tributaries from Iran. Coad (1997a) proposed that the limits of Gedrosia be extended westwards to encompass the Iranian part of the Mashkel basin, along coastal Makran as far west as the Minab River, and internally to include the Hamun-e Jaz Murian and southern Dasht-e Lut basins. West of the Minab River, the fauna was deemed to be unique in having an endemic cichlid, Iranocichla hormuzensis and in having members of such Euro-Mediterranean and Southwest Asian (= Middle East) cyprinid genera as Barbus sensu lato, Chalcalburnus (= Alburnus), Leuciscus (= Squalius) and the cobitid genus Cobitis not found further east. However specimens of Iranocichla hormuzensis have been collected from the Minab River by H. R. Esmaeili (examined by me in 1997) and this river may properly belong to the Hormuz basin. I did not collect this species in the 1970s and it is possible that the record is an introduction since that time from adjacent rivers as there have been many accidental movements of fishes in Iran associated with fish farming.

Generally basins within Gedrosia appear most closely related to their geographical neighbours and support the argument for containing these endorheic basins in one division. No basins are strongly and uniquely linked although Makran and Hamun-e Jaz Murian uniquely share Garra persica and Channa gachua, and Mashkel and Makran uniquely share Aspidoparia morar and Paraschsitura baluchiorum.

At the species level Gedrosia is most closely related to the adjacent Yaghistan and Indus basins to the east, then to the adjacent Sistan and Hormuz basins, and least of all to the remoter Tigris-Euphrates basin. Its principal relationships are eastern, to some extent northern and very little to the west.

The generic pattern is different from the species one. The Sistan basin has the highest share of genera, followed by Yaghistan and Hormuz. The Indus and Tigris-Euphrates share far fewer genera but they have a greater diversity (5.8 and 2.3 times that of Gedrosia). It is therefore not surprising that Gedrosia shares proportionately more genera with immediately neighbouring basins whose fauna at the generic level is also limited. However, omitting genera found in all basins or unique to a single basin, reveals that Yaghistan and Indus share 5 of 7 such genera exclusively with Gedrosia. Only Capoeta shows a different pattern being found in the western basins but not Yaghistan and Indus. The last genus is Crossocheilus which is found in the Indus, Yaghistan and Sistan basins. Therefore, generic level comparisons also show that Gedrosia is most closely related to the east.

The transitional nature of Gedrosia is evidenced by its having the distributional limits of certain wide-ranging species. This is most notable for species reaching their westernmost limits, namely Aspidoparia morar, Crossocheilus latius, Channa gachua, Labeo dero, Puntius sophore, and Tor putitora (the last three not recorded from Iran). Species are probably limited by environmental conditions such as temperature in comparison with the warm waters of South Asia. However a significant factor, as recognised by local people, must be the poor physical condition of Baluchistan. Freshwater marshes, lakes and large rivers are all absent. Desiccation of water bodies is common and many streams are intermittent. Habitat diversity for fishes is severely limited. All the common fish species are non-predatory - most fishes feed on small insects or scrape aufwuchs from the rocky stream beds.

In contrast to western limits, only one species has a distribution which is principally Southwest Asian and reaches its eastern limit in Gedrosia, namely Capoeta damascina. The remaining species have distributions which are centred on Gedrosia and immediately adjacent basins. There is also a link northwards in that some species have an extensive north-south distribution, namely Garra rossica, Paraschistura kessleri and P. sargadensis.

One of the most interesting features of Gedrosia is its paucity of fishes. Diversity is low, presumably a result of the physical conditions noted above, compounded by desiccation and during climatic variations both past and present. Gedrosia is presumably an important former route of dispersal for taxa from South and Southeast Asia to Southwest Asia and beyond. The significant absences are of taxa found in the Tigris-Euphrates basin to the west and in the Indus basin to the east.

At the family level, five families are found both west and east, but not in, Gedrosia. These are Cobitidae, Bagridae, Siluridae, Sisoridae and Mastacembelidae. No cobitid or silurid genera are shared. They may be quite ancient and their absence from Gedrosia is by a vicariant event or their dispersal was via a northern route to the Tigris-Euphrates and separately to the Indus. The most significant absences are of such genera as Mystus in the Bagridae, Glyptothorax in the Sisoridae, Mastacembelus in the Mastacembelidae (Mastacembelus is not found in eastern Iran and hence does not have a continuous range throughout the Orient (pace Travers (1984)), and also Barilius in the Cyprinidae. The last three genera are found in drainages entering the upper Persian Gulf separate from the Tigris-Euphrates basin but probably had a recent connection with that basin during the Pleistocene lowering of sea levels when the Gulf was drained.

Berg (1940) suggested that fish dispersal across this region was facilitated by the coastal rivers of Iranian and Pakistani Baluchestan being part of a single river system in the Pliocene, since submerged by subsidence. This distribution of these genera is not, therefore, a remnant of the dispersal across Iran from Asia. It is possible that the Pleistocene fore-deep of the Himalayas had connections with the Tigris-Euphrates basin which extending down the Persian Gulf as a river valley. Hora (1937) and Menon (1957) refer to wet, marshy, tropical conditions and headwater captures along the whole southern face of the Himalayas and westwards during the Pliocene and early Pleistocene facilitating the spread of fishes from the east to what is now Southwest Asia (= Middle East) and Africa. However, it is here considered unlikely that the Tigris-Euphrates and Gedrosian rivers were once tributary to the Indus when sea levels were lower during glaciations as the Gulf of Oman descends to an abyssal plain at 3340 m as noted above. These taxa probably reached the Tigris-Euphrates basin across the Iranian land mass and subsequently became extinct as desiccation increased. Their absence from Gedrosia is probably by loss.

Hora (1937) and Briggs (1987) consider that cyprinids entered Africa from southeast Asia 18-16 MYA, in the early Miocene, while other groups moved through Iran and the Arabian Peninsula beginning in the early Eocene. Kosswig (1951; 1952; 1955a; 1955b) notes the similarity at the generic level between Indian and African fishes, e.g. the cyprinids Barilius, Garra and Labeo, indicating that these fishes arrived in Africa from India after the desiccation of the Syrian-Iranian Sea in the Pliocene. The primary route, according to Kosswig and to Por (1987), was a northern one around the barrier of the Persian Gulf and Sea of Oman via northern Arabia, Syria and the Levant. Cooling conditions in these areas, and presumably too in Gedrosia, during the Pliocene and especially the Pleistocene glaciations, and arid climates at times, were unsuitable for tropical forms.

Potential endemic taxa are Cyprinion milesi, Paraschistura bampurensis (in Iran), Labeo gedrosicus, Labeo macmahoni, Paraschistura baluchiorum, and Triplophysa brahui (in Pakistan). The systematic position, as species, of Cyprinion milesi and Labeo gedrosicus need further study, and the distributions of the three nemacheilid species are in contention. Endemism may be relatively high or low dependent on the resolution of these problems.

Fishes in the easternmost part of the basin have a unique predator to contend with among Iranian species. The gandoo (marsh crocodile or mugger, Crocodylus palustris) is found in the Sarbaz, Khaju and Bahu Kalat rivers including the Pishin Dam, makeshift lagoons and fish culture ponds. It is feeds on Cyprinus carpio and Periophthalmus (Crocodile Specialist Group Newsletter, IUCN, 18(1), WWW Edition, downloaded 16 December 1999 from www.flmnh.ufl.edu/natsci/herpetology/newsletter/news181b.htm; report by A. Mobaraki; A. Mobaraki, pers. comm., 2000). The Cyprinus carpio are escapees from fish farms.

Tigris River

The Tigris-Euphrates basin is the largest and most important river system between the Nile and the Indus. Details of its biology can be found in Rzóska (1980) but comparatively little was based historically on the Iranian part of this basin although Nümann (1966) gave some limited data on chemical and physical parameters. There is now an increasing number of studies on environmental conditions in Iran. Studies on limnology and pollution were restricted mostly to waters of Iraq, but probably apply equally well to Iran, certainly as far as those marshes which cross the border are concerned and for the Shatt al Arab, part of which forms the southern border of Iran and Iraq. Such studies include Cressey (1958a), Jacobsen and Adams (1958), Al-Hamed (1966c), Mohammed (1965; 1966), Salonen (1970), Al-Saadi and Arndt (1973), Al-Sahaf (1975), Al-Saadi et al. (1975), Arndt and Al-Saadi (1975), Antoine and Al-Saadi (1982), Maulood et al. (1979; 1981; 1993), Sarker et al. (1980), Saad (1978a; 1978b), Saad and Antoine (1978a; 1978b; 1978c; 1982; 1983), Saad and Kell (1975), Kell and Saad (1975), Al-Hamed (1976), Al-Daham et al. (1981), Huq et al. (1981), Schiewer et al. (1982), Antoine (1983), DouAbul et al. (1987; 1987; 1988), Abaychi and Al-Saad (1988), Abaychi et al. (1988), Mohamed and Barak (1988), Al-Saadi et al. (1989), Hussain et al. (1991), Kassim and Al-Saadi (1995), Partow (2001), among others. Ionides (1937) describes the river regimes of the Tigris-Euphrates basin, MacFadyen (1938) the water supplies, El Kholy (1952) the hydrology of the Tigris River, Buringh (1957) the physiographic regions, shores and irrigation systems on the lower Mesopotamian plain, and Al-Khashab (1958) the water budget of the Tigris-Euphrates basin, mainly referring to waters in Iraq. Scott (1995) gives details of wetlands in Iraq, some of which border and/or are contiguous with Iranian wetlands, and whose general ecological features are very similar. Shapland (1997) reviews water disputes in the Middle East although western Iranian rivers flow out of the country and are not likely to be affected apart from any losses in shared habitats or refuges in border areas.

Por and Dimentman (1989) regard the Tigris-Euphrates or Mesopotamian basin as a cradle for inland aquatic faunas. A proto-Euphrates collected water from the Levant and had contacts with the Black and Caspian sea drainages before the Pliocene orogeny. Berg (1940) points out that the upper reaches of the Tigris-Euphrates basin today lie on a plateau close to the upper reaches of the Caspian Sea basin. The basin acted as an area where African and Asian species could meet or transit such as the cichlid Iranocichla. These connections were interrupted in the early Pliocene by orogeny, rifting and desert formation. Banarescu (1977) and Por and Dimentman (1989) regard the area to be a zoogeographic crossroads with elements from the Palaearctic such as the cyprinid genera Leuciscus (= Squalius) and Chondrostoma, Mediterranean genera such as the cyprinid Acanthobrama (although Krupp (1987) refers to this genus as Palaearctic, of Mesopotamian origin), and Oriental genera such as the cyprinid Garra and the spiny eel Mastacembelus. Al-Rudainy (2008) and Coad  (2010) are recent accounts of the fishes in Iraq.

Khalaji-Pirbalouty and Sari (2004) studied the biogeography of amphipods crustaceans in the central Zagros Mountains. They consider habitat diversification and climatic fluctuations to be the principal factors influencing species diversity and endemism in this area, with the mountains acting as a barrier to species distribution. Endemism is evident in lizards, plants and amphipods as well as fish.

An analysis by Coad (1996f) shows that this basin is mainly Black-Caspian Sea basin in its connections, with minor links to Asia and possibly Africa. Numbers of families, genera and species shared between the Tigris-Euphrates and neighbouring basins are summarised in this analysis. Relatively few taxa appear to have made the transition between Asia and Africa or survived subsequent climatic and habitat changes.

Certain families are absent from the Tigris-Euphrates but are found in the Indus and the Nile (Notopteridae, Schilbeidae, Clariidae, Anabantidae, Channidae). These are assumed to be of Gondwanic origin and are separated today by plate tectonic movements. A representative of the Channidae is found in eastern Iran but this species is at the western limit of its range there. Only two families are shared between the three basins but are not found to the north, Bagridae and Mastacembelidae, and the relationships of the two species in these families are with the Indus (Travers, 1984).

At the generic level, some have dispersed into eastern Iran from the Indus and other eastern basins but have not reached the Tigris-Euphrates basin, presumably for reasons of time or lack of suitable environmental conditions, e.g. Aspidoparia, Crossocheilus, schizothoracines. However two genera have reached the Tigris-Euphrates (Glyptothorax, Barilius) and Howes (1982) considers Cyprinion to be related to the eastern genus Semiplotus. Barilius resembles Indus and other eastern species superficially although its relationships have not been fully worked out. Assuming that these taxa dispersed westward from the Indus and the east, the route must be determined. All but Cyprinion are absent from much of Iran, including the bagrid Mystus and the mastacembelid Mastacembelus referred to at the family level above (Mastacembelus is not found in eastern Iran and hence does not have a continuous range throughout the Orient (pace Travers (1984)). It is unlikely that rivers of the Tigris-Euphrates basin were once tributary to the Indus when sea levels were lower during glaciations as the Gulf of Oman descends to an abyssal plain at 3340 m. I suspect, but cannot prove, that these taxa reached the Tigris-Euphrates basin across the Iranian land mass and subsequently became extinct as desiccation increased. Many of the rivers in southern and eastern Iran today are very small, regularly dry up and some are highly saline. They may be unsuitable for these taxa. Barilius, it should be noted, appears to prefer, in Asia and the Tigris-Euphrates basin, large lowland rivers and its dispersal across Iran is difficult to envisage by headwater capture (the other genera can be found in small streams at higher altitudes as well as lowland rivers). However Berg (1940) suggested that fish dispersal across this region was facilitated by the coastal rivers of Iranian and Pakistani Baluchestan being part of a single river system in the Pliocene, since submerged by subsidence. The presence of Mastacembelus and Barilius in western Iranian basins is attributed to headwater capture and/or colonisation from the Tigris-Euphrates basin when Gulf rivers were tributary to an expanded Tigris-Euphrates basin during lowered sea levels in glacial times. This distribution of these genera is not, therefore, a remnant of the dispersal across Iran from Asia.

At the generic level, only Garra is found from the Indus to the Nile and in the Tigris-Euphrates basin. Menon (1964) suggests that Garra reached the Tigris-Euphrates basin and Africa in two "waves" from Asia, the first wave being in the Miocene to the Tigris-Euphrates basin, the second through southern Arabia to Africa during the Pliocene. Karaman (1971) disputes Menon's Garra waves based on anatomy and zoogeography. Garra presumably dispersed from Asia to Africa via the Tigris-Euphrates basin and the Levant. The apparent continuous distribution of Garra across southern Arabia is not borne out in systematic analyses by Krupp (1983). Garra (and Cyprinion) species of southeastern Arabia are clearly related to southern Iranian species, having crossed the Persian Gulf when it was drained during the Pleistocene and part of an extended Tigris-Euphrates basin. Southwestern Arabian species (and a Barbus species) are a mixture of African and Levantine elements. Krupp (1983) found no evidence in his studies for the Arabian Peninsula serving as a transition area in an exchange of freshwater fishes between Asia and Africa.

Nemacheilus sensu lato also has a similar wide distribution but is probably polyphyletic and detailed revisionary works are enabling adequate zoogeographical analyses to be made. The systematics of loaches in the Middle East is a contentious subject (Por and Dimentman, 1989). The absence of nemacheilid species from southern Arabia also argues for a dispersal route through the Tigris-Euphrates basin as these cryptic fishes are found today in many small streams throughout Southwest Asia and are unlikely to have been eliminated from southern Arabia through desiccation.

The only Nile (or east African) genus present in the Tigris-Euphrates basin is Barbus sensu lato. Certain members of this polyphyletic genus in Southwest Asia are characterised by sharing 6 branched anal fin rays, last unbranched dorsal fin ray a smooth spine, large scales, few gill rakers, high dorsal fin ray counts, reduced barbel numbers, compressed body, and other characters which set them apart from European Barbus as a monophyletic group, probably related to east African species (suggested by Banister (1980)). These former Barbus species are found from southwestern Arabia (but not southeastern Arabia), through the Levant and the Tigris-Euphrates basin to rivers at the Strait of Hormuz in Iran. They may represent an African element in the fauna of the Tigris-Euphrates and may reflect the route of the cichlid Iranocichla or its ancestor from Africa to the Strait of Hormuz. Bănărescu (1992b) considers African elements in Southwest Asia to be the oldest, of at least Miocene age.

A significant proportion of the families and genera in the Tigris-Euphrates basin is also found in the Black-Caspian Sea basin. Such widespread, northern cyprinid genera as Alburnoides, Alburnus, Aspius, Alburnus, Chondrostoma, and Leuciscus (= Squalius) reach their southern limit in the Tigris-Euphrates basin (and neighbouring Iranian basins) suggesting that they reached the Tigris-Euphrates basin from the north.

The presence of Glyptothorax in the Black Sea basin of Anatolia (Coad and Delmastro, 1985) is a recent event through headwater capture from the Tigris-Euphrates basin and thus far is the only example of a clearly-defined Indus genus reaching the Black-Caspian seas basin. It is probably an example, in reverse, of the colonisation of the Tigris-Euphrates basin in recent times from the Black-Caspian seas basin. Headwaters of a number of Tigris-Euphrates basin rivers interdigitate with the upper reaches of Black-Caspian seas basin rivers, e.g. the Aras River of the Caspian Sea and the Kizilirmak of the Black Sea with the Euphrates near Erzurum and Sivas respectively; the Qezel Owzan of the Caspian Sea with Tigris River tributaries. Headwater capture is common in the Zagros Mountains (Oberlander, 1965) and in Anatolia and pluvial conditions in the past would have facilitated fish dispersal. Por and Dimentman (1989) mention direct connections of a proto-Euphrates with Black Sea and Caspian sea fluviatile drainages before the Pliocene orogeny which would serve to allow entry of taxa to the Tigris-Euphrates basin. Direct connections were interrupted by the early Pliocene as orogeny, rifting and desertification took hold. Almaça (1990) has reviewed possible routes for Barbus sensu lato species into Iran and the Tigris-Euphrates basin from the north via what is now Anatolia and east of the Caspian Sea dating from the early Oligocene. A continuous route for exchange of taxa has been possible since the upper Miocene, almost 12 million years ago. These routes have been variously available down to modern times for Barbus sensu lato and other taxa as exemplified by some species being in common between the Black-Caspian seas basin while others are distinct but related at the generic level. Bănărescu (1992b) considers that northern or European elements penetrated to the Tigris-Euphrates basin earlier than Asian ones, and that this partially explains their prevalence.

Iranian internal and Gulf basins and the Levant show evident affinities with the Tigris-Euphrates basin. The ichthyogeography of the Levant has been dealt with by Krupp (1987) and will not be reviewed here. Krupp considers that parts of the Levant were colonised separately via branches of the Tigris-Euphrates river system. Iranian basins to the west of the Tigris-Euphrates basin have a very similar fauna to that of the Tigris-Euphrates at the species level. The diversity falls off rapidly with distance (Coad, 1987). Headwater capture in the Zagros Mountains is an evident route for species found in common with the Tigris-Euphrates basin but not in Iranian rivers draining separately to the Gulf. The draining of the Gulf during Pleistocene lowering of sea levels enabled Tigris-Euphrates basin fishes to colonise tributary Iranian rivers now separated by a rise in sea level. The melting of the Laurentide ice sheet and drainage of Lake Agassiz in Canada caused this rise in sea level world-wide, including the shallow Persian Gulf (Perkins, 2002). By about 11,500 years B.P., the Gulf was filled with present shorelines attained shortly before 6000 B.P. and exceeded by 1-2 m (Lambeck, 1996).

Por and Dimentman (1989) regard the Mesopotamian subregion or Tigris-Euphrates basin as one of the most isolated major freshwater areas in the world. However, as Coad (1997f) points out, endemism is only at the species level and diversity is low with only about 52 primary division species in 7 families, 34 species of which are Cyprinidae.

The Zagros Mountains form the western flank of Iran and store water as snow. The higher peaks are snow-capped even in summer. Zard Kuh, for example, reaches 4548 m (32°22'N, 50°04'E). Rivers drain south and west to become tributaries of the Tigris River in Iraq or its confluence with the Euphrates River, the Shatt al Arab (known as the Arvand (= swift) Rud in Iran). The Shatt al Arab has a course of 190 km to the head of the Persian Gulf and is navigable by ocean-going ships. It forms part of the Iran-Iraq border. The origin of the Tigris River is the Hazar Gölü of Elazig (38°41'N, 39°14'E) between the Murat Nehri and the Euphrates. It flows south-east, forming a short section of the border of Syria with Turkey, before entering Iraq to parallel, roughly, the course of the Euphrates River. It is a larger and swifter river than the Euphrates because of its left bank tributaries from Iran. The Tigris is over 1900 km long (1851 km and 2032 km are extremes cited in the literature). It is the 81st river in size in the world. The Tigris-Euphrates basin encompasses 784,500 sq km of which 19% or 146,000 sq km lies in Iran (Gleick (1993) gives 238,500 sq km and 27% for Iran and 884,000 sq km for the whole basin; the Iraqi Government in the same publication gives 378,834 sq km for the Tigris basin alone with Iran's share 28.8%). Iran contributes 7% of the water supply of this immense basin. The Tigris catchment is 166,155 sq km.

The Tigris is an alkaline river (pH 7.8-8.2) with a total hardness of 200-350 mg/l. Water temperatures range from 8.5°C in January to 31.4°C in August. The flow pattern of the Tigris and its tributaries has a sharp peak in April at about 9 billion cu m, falling rapidly to about 1 billion cu m from August to October before beginning to rise again. The water level may fall by as much as 2 m over the summer. Interannual variation in spring flood levels are marked. Approximate streamflows over the past 6000 years are given by Kay and Johnson (1981) based on proxy data from paleoenvironmental sources. They found an increase in streamflow over this period. The southern province of Khuzestan in the Tigris river basin with 9% of Iran's surface area has an estimated 37% of its surface water flow.

The Shatt al Arab is under tidal influence up to 110 km from the mouth. Its waters are therefore strongly mineralised. Salinity varies with distance from the sea. Crops are irrigated by means of the tidal rise which is used to push fresh water into the fields (Harrison, 1942; Gholizadeh, 1963; Gholizadeh and Fatemi, 1969). This has obvious effects for the fish fauna and its composition as well as for increased salinisation of habitats. There are appreciable diurnal and seasonal fluctuations in physico-chemical conditions. Tidal waters probably penetrated far inland through the Holocene as evidenced by faunal remains in boreholes of the Hammar Formation (MacFadyen and Vita-Finzi, 1978). Al-Hassan and Hussain (1985) describe the hydrological parameters affecting the penetration of marine fishes into the Shatt al Arab. Recently an increase in the Tigris River discharge has decreased salinity in the Shatt al Arab: previously marine species were common at Basrah in Iraq but they became rare, Carassius auratus appeared in Basrah fish market and Cyprinus carpio was caught in large numbers down to the estuary (N. A. Hussain, in litt., 1994). Pollution is widespread in the Shatt al Arab from industrial, agricultural and untreated human wastes. Hussain et al. (2001) evaluate environmental degradation in the Iraqi portion of the Shatt Al-Arab and its effects on the fish fauna.

The principal Iranian tributaries of the Tigris River are the Little Zab River (= Zab-e Kuchek) which drains a small stretch of mountains south of Lake Orumiyeh, and the Diyala River (= Sirvan River) which drains the western mountains of Kordestan. The Diyala River is 442 km long. A principal tributary of the Diyala in Iran is the Qeshlaq River which flows through Sanandaj (35°19'N, 47°00'E). The river is polluted from Sanandaj and from agriculture wastes (Jafari Salim et al., 2009). The Qeshlaq or Vahdat Dam near Sanandaj has a fauna including Alburnus mossulensis, Barbus lacerta, Capoeta damascina, C. trutta, Ctenopharyngodon idella, Cyprinus carpio, Gambusia sp., Pseudorasbora parva and Squalius cephalus (Barzegar and Jalali, 2006). Nine species of parasites were found on this fauna, notably Ligula intestinalis, which is detrimental to native and food fishes.

Lake Zaribar, Zarivar or Zeribar is a permanent freshwater body with fringing reed beds and extensive marshes lying at 1435 m in the Diyala River drainage just west of Marivan at 35°32'N, 46°08'E. It has an area of 8 sq km and a maximum depth of 6 m and an average depth of 2.5-3.5 m. Reputedly the lake is fed by 600-700 springs. At high water it overflows into a small river at its southern end. In winter it often freezes over. It was damaged in the Iran-Iraq War suffering rocket and missile hits and chemical warfare (Scott, 1995; 1997). There is a small resort at the southeast corner of the lake, the surrounding land has livestock grazing and agriculture with drainage channels for the peripheral marshes, forests are cut for fuel, and there is waterfowl hunting and fishing. Exotic fish species have been introduced by a government organization, including Alburnus alburnus (= hohenackeri), Ctenopharyngodon idella, Cyprinus carpio (in two varieties), Hemiculter leucisculus, Hypophthalmichthys molitrix, H. nobilis, Pseudorasbora parva and Gambusia holbrooki. Native fish include Barbus lacerta, Capoeta buhsei (sic), Leuciscus (= Squalius) cephalus and Mastacembelus mastacembelus (Scott, 1997). Jalali et al. (2002) add the species Capoeta damascina (possibly the correct identification of the C. buhsei listed above but Esmaeili et al. (2010) include C. barroisi), Carassius auratus (given as Carassius gibelio by Esmaeili et al. (2010), and Chalcalburnus (= Alburnus) sp.

A number of minor streams also cross the Iran-Iraq border, but the principal rivers drain through anticlines in spectacular gorges or tangs, funnelling the waters of the Zagros onto the Khuzestan plains through a narrow gap near Dezful (32°23'N, 48°24'E). Stream flows in late winter are at least ten times that of summer and 116,500 sq km of mountains and three big rivers debouch onto 38,800 sq km of plain. Lowlands may be inundated for more than 100 days. Early accounts of floods in Mesopotamia, dating back to Sumerian times almost 5000 years ago, are discussed by Mallowan (1964). Floods can encompass close to 100,000 sq km in Iran and Iraq at the head of the Persian Gulf (Naff and Matson, 1984). Progressive clearing of woodland over the last 7000 years increased runoff, causing higher and more severe floods, soil erosion, increased turbidity in streams and higher sedimentation (Wagstaff, 1985). Erosion is three times the world standard rate at 30 tonnes/hectare and will rise twofold over the next ten years (IRNA, 20 December 1998). All these must have, and continue, to affect the fishes in this and other basins, favouring those species able to cope with these conditions. Even artificial habitats such as small dam reservoirs in Chahar Mahall and Bakhtiari are affected by high sedimentation rates and their utility as fish habitat must be affected (Mousavi and Samadi-Boroujeni, 1998).

The main river is the Karun, with a catchment of 67,340 sq km (Naff and Matson, 1984) and a length of 820 km. It now drains to the Shatt al Arab but once drained directly into the Persian Gulf. The Karun is also connected to the Gulf via the Bahmanshir River, paralleling the Shatt al Arab, and enclosing Abadan Island. The Bahmanshir is the only river along the Persian Gulf coast to have a significant fishery. A physicochemical study of the Bahmanshir was carried out by Faal (2009). The Karun headwaters are extensive and lie near both the Esfahan and Kor River basins. The environmental conditions in a headwater dam, the Hanna Reservoir, in the Karun basin are described by Esteky (2001), two-thirds of the reservoir being covered by macrophytes. The Dez River is a Karun tributary and is 400 km long. The Karkheh River (with the Cherdavel, Kashkan, Qareh Su, Gav Masiab and Simareh in its upper reaches) is 320 km long, but is lost in the Hawr-al-Azim marshes of the Tigris after draining 43,000 sq km. Sutcliffe and Carpenter (1967) described runoff from the Karkheh basin. The Karkheh and Dez flows were depleted by 70% in 2001 during a drought and it was thought that these rivers might dry completely (Foltz, 2002). The marshes along the Karkheh and Dez rivers, with oxbow lakes and riverine forest, are a habitat now rare in southern Iran and Iraq outside protected areas. The severe drought of the year 2000 dried up the natural Dez reservoirs south of Dezful (www.irna.com/newshtm/eng/08130315.htm, IRNA, 29 July 2000). The Karkheh Dam, 20 km northwest of Andimeshk, has a crest 3030 m long, a height of 127 m and is the sixth largest dam in the world with a capacity of 7.8 billion cu m, nearly a third of the total dam capacity for the country. The dam is meant to produce electricity, for fish farming and to control floods and drought (IRNA, 17 April 2001; 19 April 2001; Aftab Yazd, Tehran, 346(18 April 2001, 7 pp.; Sadegi, 2003). The Qareh Su near Kermanshah is about 30 m wide and less than a metre deep at its deepest. The Qareh Su or "black water" derives its name from its transparency over a dark, pebbly bed, distinguishing it from the muddy rivers of the lowlands. The Qareh Su is the Classical Choaspes, the water of which the ancient monarchs of Persia carried with them on their military expeditions for its taste, a superiority confirmed by Buckingham (1829). The Gawshan Dam is located at Kamyaran near Kermanshah on the "Gaweh" River and is scheduled for completion in 2002. The dam will be 136 m high and the complex includes a 19 km long tunnel for water transfer (http://netiran.com/news/IranNews/html/94111305INEC.html). Other dams include the 40 million cu m Zarivar Dam in Marivan and the 563 million cu m Kavoshan Dam 35 km south of Sanandaj (http://netiran.com/news/TehranTimes/html/95111803TTPL.html).

Partow (2001) lists 18 dams in the Tigris basin of Iran, either constructed or planned, and these will affect the environment markedly in changing flow regimes, impounding water and eliminating fluvial habitat, removing silt, affecting temperature downstream, causing salinisation as return water from irrigation projects flows back into rivers, and so on. The Karkheh Dam is planned to carry water via pipeline over land (330 km in length) and under the sea (210 km) to Kuwait. The supply rate would be 200 million gallons per day (Partow, 2001) or 300 million cu m (www.irna.com, downloaded 29 January 2003).

The Dez (formerly Mohammed Reza Shah Pahlavi) Dam on the Dez River at 32°38'N, 48°28'E contains 3350 million cu m of water (another source states 60 billion cu m) and has a maximum surface area of 4000 ha. Surface water temperatures can exceed 30°C while at 50 m plus depths it is 15-16°C in summer. Its original life span was estimated at 100 years but this had to be reduced to less than 50 years because of the rapid accumulation of sediment from erosion. Sediment prevents development of a bottom fauna and steep banks with water fluctuations limit vegetation. Nümann (1966, 1969) gives some limnological information on this reservoir. Nümann (1966) recommended introduction of Acanthobrama terraesanctae and Tilapia galilaea from Israel to the reservoir, and later Sander lucioperca and even trout. Sabzalizadeh (2006) gives a description of the ecology of this reservoir and Eskandari et al. (2007) a description of fish populations. Capoeta trutta, Barbus (= Tor) grypus and Barbus (= Luciobarbus) esocinus were the most numerous species and the fauna includes the exotics, Carassius auratus, Hypophthalmichthys molitrix and Oncorhynchus mykiss. There is also a diversion dam, the Sadd-e Gotvand. The Gotvand Dam is under construction and will be 180 m high with a reservoir capacity of 4,500 million cubic metres making it the second largest dam in Iran (sic) (IRNA, 25 January 2000). The 205 m high Karun-3 Dam near Izeh, to be completed in the year 2001 (filling actually started in 2003 - www.netiran.com, downloaded 15 November 2004), is a major hydroelectrical plant as is the Karun-4 Dam (Shahid Abbaspour) near 25 km northeast of Masjed-e Soleyman (http://netiran.com/news/IranNews/html/95040822INPL.html). A major dam is also planned at Shushtar (IRNA, 26 September 1998). A tunnel is planned from the Dez River to Golpayegan to supply water to Markazi Province in central Iran (www.iranmania.com, downloaded 19 January 2004).

Some literature refers to the Seymarreh-Karasu-Gamasiab (and variant spellings) as an important complex of rivers. These are the Simareh, Qareh Su and Gav Masiab in gazetteers. A giant dam is planned for the Simareh (IRNA, 26 September 1998). Nümann (1966) notes pollution in these rivers from an oil refinery and sugar factory which decreased fish populations, a condition exacerbated through the use of explosives, insecticides and herbicides by local people to catch fish. He also lists explosive usage on the rivers Khairabad and Zohreh.

Lake Mirabad lies in the basin of the Karkheh at 33°05'N, 47°43'E. While it measures only 100 by 200 m it is important for establishing past vegetation and environments based on sediment cores (Griffiths et al., 2001). The Hashelan or Hashilan Marsh at 34°33'N, 46°55'E occupies 260 to 400 ha (accounts differ) northwest of Kermanshah at about 1310 m. It is a complex of permanent spring-fed pools and associated marshes with much submerged, floating and emergent vegetation. The surrounding plains are heavily grazed and cultivated and ducks are hunted in the marshes. The Sabz Ali spring feeding the marsh has an average annual discharge of 323.4 l/sec, range 208.3-442.5 l/sec, highest in March and lowest in September. The total average volume of water in the marsh is estimated at 1.02 x 107 (Karami et al., 2001). Local people and those from Kermanshah fish in the marsh. A drought in 2008 severely affected the Hashilan Marsh (www.payvand.com/news/08/aug/1152.html, downloaded 5 July 2009).

A truck carrying diethyl hexanoyl plunged into the Kashkan River, a Karkheh tributary in Lorestan, 15 km from Pol-e Dokhtar resulting in the poisoning of thousands of fish on 13 April 1998 (IRNA, 14 April 1998; Brief on Iran, 880, 16 April 1998). The river suffered an oil slick in October 2001 when the Khuzestan-Tehran pipeline fractured 4 km from Pol-e Dokhtar. Oil pollution caused a fish kill numbering about 70,000 fish in the Kambel River near Gachsaran, a centre of oil production (Tehran Times, 24 November 2002). Varkouhi and Sobhani (2005) and Varkouhi (2007) studied the presence of various pollutants in the livers of fishes in the Khorramabad River. The Meymeh River in Ilam has some pollution from urban and rural sewage, and this may potentially increase (Cheraghi et al., 2007).

The Jarrahi River is a southern Karun tributary from the east. The Marun River is a major Jarrahi tributary. The Marun and Jarrahi feed the Shadegan Marshes, the largest Iranian wetland according to Kurdistani and Bajestan (2004). The Marun Reservoir Dam northeast of Behbahan was scheduled for completion in 1996 with a crest of 345 m (IRNA, 11 November 1998) but was to be completed in 2004 with a crest of 175 m and containing 1.2-1.3 billion cu m of water (IRNA, 12 January 1999; IRNA, 5 February 2002). There are also four diversion dams on the Marun and one of these, the Jazaeen, has a fishway but fish are trapped downstream of it during their migration (sic)(Kurdistani and Bajestan, 2004). Other dams in this system lack a fishway. Later Kurdistani and Bajestan state that there are no migratory fishes in the Marun, only resident species (which presumably undergo local movements blocked by dams). They mention Barbus (= Tor) grypus and Barbus (= Luciobarbus) pectoralis as the affected species. The Jareh Dam on the Zard River northeast of Ramhormoz dates back to the Sassanid era and is still in use (IRNA, 26 June 2000).

The Karun has the greatest mean discharge, followed by the Dez and Karkheh. The Karun mean discharge is the largest in Iran. The Karun carries a heavy silt load with a hundredfold increase during flood. The Karun discharge ranges from 207 cu m per second to 2225 cu m/sec, average 1100 cu m/sec, while the Dez is 63-1227 cu m/sec, average 288 cu m/sec. The Jarrahi range is 8-770 cu m/sec, average 78 cu m/sec. These figures vary among different sources indicating fluctuations between years and gauging stations; however the relative importance of these rivers is shown. The peak discharge of the Karun is in April, with high values also in March and May; the lowest discharge is in October when flow is only about a ninth of the peak. The combined Tigris-Euphrates-Karun in flood carries five times the load of the Nile (Fisher, 1968). Most of this is deposited north of Basrah (30°30'N, 47°47'E) and much is lost to evaporation in the marshes, e.g. of 27 cu km of discharge into the Persian Gulf through the Shatt al Arab, 22 cu km is from the Karun River. 22 million metric tons of dissolved chemicals are deposited each year and hence there are siltation and salinity problems in the lower parts of this basin.

The Karun River on the Khuzestan plains was examined in 1992 for various parameters and at various localities (courtesy of the Iranian Fisheries Research and Training Organization, Ahvaz). It has a pH of 7.07-8.85, mean 8.17, dissolved oxygen 5.6-12.38 p.p.m., mean 9.29 p.p.m., bicarbonate 79.3-214.72 p.p.m., mean 154.4 p.p.m., carbonate 0.6-21, mean 5.53 p.p.m., total alkalinity 1.9-3.8 meq/l, mean 2.84 meq/l, carbonate hardness 5.32-10.64 p.p.m., mean 7.95 p.p.m., total hardness 168-474 p.p.m., mean 287 p.p.m., ash residue 40-1142 p.p.m., mean 425 p.p.m., chloride 45.4-518.3 p.p.m., mean 207.28 p.p.m., total dissolved solids 226-1374 p.p.m., mean 696 p.p.m., sulphate 43.75-325 p.p.m., mean 101.73 p.p.m., calcium 33.63-101.7 p.p.m., mean 61.8 p.p.m., magnesium 16.8-78.24 p.p.m., mean 33.8 p.p.m., phosphate 0.05-4 p.p.m., mean 0.24 p.p.m., iron trace to 0.32 p.p.m., mean 0.069 p.p.m., manganese trace to 0.657 p.p.m., mean 0.483 p.p.m., and nitrate trace to 0.657 p.p.m., mean 0.039 p.p.m. Esmaili et al. (1999) report heavy metals in water, sediments and fish from the Karun River and Jafarzadeh-Haghiehi et al. (2005) report on the poor water quality of the river. Haghighi and Arabi (2010) modeled water exploitation of this river for fish farms, tracing heavy metal pollution and concluding where water could be safely withdrawn.

As lowlands at the head of the Persian Gulf receive waters from this vast drainage basin, floods occur, increasing the depth and extent of marshes. Flood waters may increase depths by 1-1.5 m, with 2-3.5 m in more permanent basins. Most of the large marshes lie in Iraq, but the Hoveyzeh or Hawr-al-Azim marshes are on the border, and occupy 3000 sq km at high water. They are fed by the Karkheh and other rivers from Iran. Construction of the Karkheh Dam in Iran (and pipeline water transfer to Kuwait) will reduce input of water to this marsh, compounded by canal construction and draining of marshes in Iraq. Additionally, irrigation return waters will be salinised (Partow, 2001). A dam has been built by Iran across the Hoveyzeh marsh to retain water on the border with Iraq. Marsh temperatures range from 15°C in January to 31°C in August and fish may retreat to deeper areas or move upriver at the higher temperatures. Flooded marshes tend to be warmer than rivers in winter. The Shatt al Arab has temperatures of 32°C in July and 16°C in December.

Floods are often a feature of these southern rivers and some loss of fish stocks must occur as they recede. For example, the Jarrahi and Zohreh rivers overran their banks in November 1994 after torrential rains causing widespread flooding (http://netiran.com/news/IranNews/html/94112109INEV.html).

The Zagros Mountains consists of tightly packed ranges in the Tigris basin trending north-west to south-east. A trellis drainage pattern is imposed on this. The tangs, their formation and the drainage pattern are described by Harrison (1937) and Oberlander (1965; 1968a). These deep defiles may exceed 2400 m in depth with vertical walls of 300 m splitting anticlinal mountain ranges instead of taking apparently easier routes around their ends. They may well be barriers to the movement of less vagile fish species or a highway into the interior for those with some dispersal ability. Tangs formed because an antecedent drainage over lower relief was gradually uplifted at a rate slow enough to permit streams to cut through ridges and retain the original pattern of drainage once the softer material was washed out of the valleys between the anticlines.

The uppermost parts of the basin show evidence of headwater captures and this orogenic zone is very unstable. The divide between endo- and exo-rheic basins is not the snowline of the Zagros but is east of it, so streams must first cross the Zagros peaks to start on their journey to the Persian Gulf.

Springs are important in the mountains, tapping aquifers and helping to maintain river flow. The Karun River traditionally has its source in springs. Keivany et al. (1992) surveyed 72 springs in Chahar Mahall va Bakhtiari Province, in the upper Karun River basin, and found them suitable for trout culture with a potential production of about 6000 tonnes per year. Flows varied from 50 to 4000 l/second, temperature from 6 to 15°C, pH from 6.2 to 7.8, conductivity from 128 to 570 mMoh/cm, total alkalinity from 20-220 meq/l, total hardness from 140-250 mg/l, oxygen from 7 to 11 mg/l, carbon dioxide from 5 to 20 mg/l (falling rapidly to less than 2 mg/l within a few tens of metres of the spring source), H2S 0 mg/l, Cl- 1-28 mg/l, SO4-- 14-135 mg/l, PO4-- 0.1-0.3 mg/l, Ca++ 16-82 mg/l, Mg++ 3-34 mg/l, K+ 0.2-1.0 mg/l, Na+ 0.5-1.5 mg/l, Fe+++ 0-0.06 mg/l, Fe++ 0 mg/l, NO2- 0-0.2 mg/l, NO3- 0-13 mg/l, NH4+ 0-0.5 mg/l, and HCO3- 48-220 mg/l. Springs (or sarabs) in Kermanshah Province have been described by Khatami and Shayegan (2003) and are regarded as a significant water supply for rivers. Sarabs are used for drinking water and irrigation, and are threatened by pollution and fish farms. Qanats are also found, in drier parts of the basin, but they are not as significant for fish habitat as in other parts of Iran.

Marshes and ponds as well as seasonally flooded arable land along the Karun River in the lowlands of Khuzestan provide temporary and permanent habitats for fishes. Some are reviewed below.

The "Hawr-e Bmdej"? or "Sadi Shavour" Marshes lie between the Karkheh and Dez rivers northwest of Ahvaz at 31°45'N, 48°36'E and encompass 12,000 ha. This is the most extensive permanent freshwater marsh with tall reeds of Phragmites and Typha in Khuzestan. There is relatively little open water. Some parts are being drained for agriculture, a continuing trend for marshes with concomitant loss of fish habitat. The "Hamidieh" plains at 31°20'N, 48°20'E comprise 20,000 ha of seasonally flooded (winter) plain and arable land along the Karkheh River. Hamidieh Lake, an old oxbow of the Karkheh, is included in this area and is permanent fresh water. The lake is 3 ha and has extensive reed beds.

The "Susangerd" Marshes or Hawr-e Susangerd at 31°45'N, 47°55'E are northwest of Ahvaz near the Iraqi border and form the extreme eastern edge of the Hawr-al-Azim, most of which lies in Iraq. The marshes occupy about 30,000 ha and are made up of permanent and seasonal fresh and brackish marshes and seasonally flooded arable land. The marshes are on the floodplain of the Karkheh River. Irrigation projects, grazing by livestock, reed cutting and fishing all occur here. Parts of the marsh were damaged by the Iran-Iraq War. The Iran-Iraq marshes declined in area from 1089 sq km to 758 sq km from 2000 to 2002 and was predicted to dry up in 5 years from 2002 because of the Karkheh Dam. Reports conflict since once the dam was full, a relatively normal flow regime would help maintain the marshes. Restocking with 490,000 Barbus (= Mesopotamichthys) sharpeyi and and Barbus (= Luciobarbus) xanthopterus took place in this marsh (www.shilat.com, downloaded 12May 2006).

The Shadegan Marshes at 30°20'N, 48°20'E occupy 282,500 ha (Jones, www.ramsar.org/lib_dir_2_3.htm, downloaded 4 April 2000) gives 296,000 ha) and form the southern part of the seasonal floodplain of the Dez, Karun and other rivers at the head of the Persian Gulf. There are adjacent tidal mudflats. The central and southern part of the marshes are part of a Ramsar Site, along with the mudflats (World Conservation Monitoring Centre, 1990). Sabzalizadeh and Amirineia (2003) give some physical and chemical characteristics of 5 sample stations in this marsh. Range of pH was 7.2-9.4, maximum water temperatures occurred in July and August. Maximum levels of dissolved oxygen were found in November and February but were more than 5 p.p.m in most cases, optimum for fish growth and reproduction. The water quality was hard and brackish. The whole area may dry out in late summer, a natural condition exacerbated by dams and irrigation schemes on the major inflowing rivers. In a November 2000 visit, much of this area was dry although it had been flooded in 1999. When the marsh dries, fish concentrate in the deeper pools where they are easily caught, even the smaller ones. The marsh is re-colonised from the rivers. Fishing is important. The fishes of this marsh in order of abundance are kopur, shirbot, touyeni, esbele, binni, berzem, biah and very few himri and gattan (Y. Mayahi, pers. comm., 2000). Rice paddies occupy part of the Ramsar Site and reed cutting, fishing and grazing goes on. There is extensive reed cutting, some livestock grazing, some rice paddies and potential pollution from main roads, shipping and oil terminals. Over 100,000 ha were contaminated with oil from a leaking pipeline in 2000 and 35,000 cu m of refinery wastes were dumped in the marsh in 2004 (www.payvand.com, downloaded, 5 September 2006). Esmaeili Sari et al. (2001) detail the damages resulting from the war's oil pollution when 20% of the emergent vegetation was destroyed. Chemical weapon use occurred here in the Iran-Iraq War and acid rain fell from the burning of the Kuwaiti oilfields in the Gulf War. About 10% of the marshes were destroyed (Anonymous, 1988b; Scott, 1993; Jones, www.ramsar.org/lib_dir_2_3.htm, downloaded 4 April 2000). The Shadegan Wildlife Refuge, encompassing 296,000 ha, is on the threatened list for National Parks since it was substantially damaged in the Iran-Iraq War, both physically and by chemical agents.

The principal fishes appearing on fish stalls in Ahvaz from marshes such as Shadegan are Luciobarbus xanthopterus, Liza abu, Mesopotamichthys sharpeyi and Cyprinus carpio as well as cultured Hypophthalmichthys molitrix as escapes or plantings. Farm ponds in Khuzestan have Luciobarbus barbulus, Ctenopharyngodon idella, Hypophthalmichthys molitrix and Cyprinus carpio. Hawr-al-Azim, Hawr-al-Hoveyzeh and the Shadegan marshes are important refuges for fishes in Khuzestan (Korki, 1992; N. Najafpour, pers. comm., 1995). 490,000 fingerlings of Barbus (= Luciobarbus) sharpeyi and Barbus (= Luciobarbus) xanthopterus were stocked in this marsh in 2005, a 40% increase over the previous year (www.iranfisheries.net, downloaded 30 November 2005). Various studies on fish parasites have been carried out in southwest Iran (Khuzestan Province) and these are mostly dealt with under the Species Accounts. Mortezaei et al. (2008), for example, collected fishes from the Haw-al-Azim, Shadegan Marsh and Karun River and recorded such nematodes as Rhabdocona denudata, R. fortunatowi, Rhabdocona sp., Proleptinae, Cucullanus sp., Pseudocapillaria tomentosa, Philometra karunensis, Philometra sp., Anisakis sp. and Contracaecum sp. from 10 fish species.

Izeh and "Shiekho" lakes at 31°52'N, 49°54'E occupy 1400 ha in the Zagros foothills. These small freshwater lakes are shallow with extensive sedge marshes. Izeh is the deeper of the two with much more open water. They are fed by runoff and springs. Shiekho, the larger lake, is almost overgrown with emergent vegetation except where cattle have grazed and trampled areas leaving some open water. Some fishing occurs in the lakes and water is abstracted for irrigation.

"Choghakor" or "Chaghakhour" Marsh or Wetland at 31°55'N, 50°54'E lies in upper Karun River drainage in the Zagros Mountains in Chahar Mahal va Bakhtiari at ca. 2100-2270 m and occupies 1600 ha. Maximum depth in spring and winter is 2 m but in summer it is almost entirely dry and overgrown with emergent vegetation. Construction of a dam may enable a more permanent marsh to exist (Taqvaie and Ramezani, 2002) although others consider dam construction to be a threat to the habitat and its diversity as the habitat changes from a wetland to a lake (Ebrahimi and Moshari, 2006). After the dam was built, water depth increased from 1.5 m to 6 m or more. Mousavi Nadoushan et al. (2008) record introduction of cyprinids, which along with water level fluctuations and agricultural discharge, caused serious changes in trophic states. Fish yield potential was estimated at 34.4 kg.ha-1. Rahimi and Raeisi (2009) found lead and cadmium levels in fish tissues from this marsh exceeded tolerance limits established by the European Commission. These high concentrations probably came from misuse of phosphate fertilisers in local agriculture. Fadaei Fard et al. (2001) recorded Alburnus mossulensis, Capoeta aculeata, C. damascina, Carassius auratus, Chondrostoma orientale (sic), Cyprinus carpio, Hypopthlalmichthys molitrix and Aphanius vladykovi from this marsh area. The parasites Dactylogyrus lenkorani, D. extensus, Gyrodactylus sp., Diplostomum spathaceum, Allocreadium isoporum, Ichthyophthirius multifilis, Trichodina sp., Myxobolus sp., Lernaea sp., Rhabdocona sp. and Acanthocephalorhynchoides cholodkowski were recorded from these fishes with Cyprinus carpio with about 88% infestation and Aphanius vladykovi parasite -free.

Gandoman Marsh or Lagoon at 31°50'N, 51°07'E at 2250 m and occupying 1500 ha (or 1200 ha, Khan et al. (1992) or 3510 ha Taqvaie and Ramezani (2002)) is a similar habitat but it has a stream running through it. "Sulegan" wetland or marsh in the same area encompasses 164 ha and is spring fed. These marshes have been proposed as a Ramsar Site although not yet formally designated (Scott and Smart, 1992). Raissy et al. (2010) record Alburnus alburnus, Capoeta aculeata, C. damascina, Carassius auratus, Cyprinus carpio and Chondrostoma regium from this lagoon, parasitised by Ichthyophthirius mulitifilis, Trichodina sp. (Ciliophora), Myxobolus musayevi, Myxobolus sp. (Myxozoa), Dactylogyrus extensus, D. lenkorani (Monogenea), Diplostomum spathaceum, Tylodelphys clavata (Digenea), and Argulus foliaceus and Lernaea cyprinacea (Crustacea), with 77.7% of fish infected with at least one of these.

The southern areas of this basin are areas with high temperatures and large cities (Abadan in Iran and Basrah in Iraq). Adjacent waters are highly polluted with sewage, agricultural waste and other chemicals (e.g. see DouAbul et al., 1988; Diagomanolin et al., 2004; Karamouz et al, 2005; Afkhami et al., 2007). The increased use of motor boats has led to oil pollution. DDT is still sprayed against malarial mosquitos on stagnant pools adjacent to the main river course leaving a brown stain on the rocks (observations in 1995; a letter of complaint to the appropriate agency carrying out this spraying by the Iranian Fisheries Research and Training Organization elicited no response). Scott (1995) records sale of Chloridrin, a persistent insecticide, to residents of the Hawr-al-Hoveyzeh in Iran as a means of poisoning large numbers of fish for sale. Phytoplankton blooms are common and in canals the chlorosity increases, transparency decreases and pH is reduced because of the dying plant material. The Shatt al Arab is more affected by physical factors as it is an estuary. Historical problems with salinisation of soils (and presumably water) extend back 5000 years in southern Mesopotamia including Khuzestan, a consequence of over-irrigation and inadequate drainage (Goldsmith and Hildyard, 1984). The irrigation systems rose and fell with the vicissitudes of history. There was a large-scale irrigation network in Khuzestan during the Sassanian period (A.D. 226-639), lost through conflict and natural disasters after this date and reconstructed in modern times (Adams, 1962).

A theory has been advanced that the silt-laden discharge of the Tigris-Euphrates-Karun rivers has built out a delta into the Persian Gulf. The head of the Gulf would have reached Baghdad and Samarra about 7000-6000 B.P. and since then the land area is supposed to have extended some 200 km southward. The present plains would not then have been as extensive and rivers from Iran would have entered directly into the Gulf. The Admiralty Naval Staff (1918), Mason et al. (1944), Adams (1962), Hansman (1978), Maltby (1994) and Lambeck (1996) provided illustrations of this recession of the head of the Persian Gulf in historic times along with details of historical and archaeological evidence. The sea coast was then supposedly as far inland as Ahvaz in Iran for example. Lees and Falcon (1952) proposed that in fact downwarping occurs under the weight of sediment. Certainly the silt load has not built up a land surface. The coastline, under this theory as interpreted by Fisher (1968), has been constant since the end of the Pliocene and presumably as a marsh habitat for fishes too. However Lees and Falcon did state that there were advances and retreats through historic and prehistoric time. Ionides (1954), Larsen (1975) and Nützel (1975) refuted Lees and Falcon and maintained that marine clays and silts indicate a marine embayment as far inland as Amara in Iraq (31°50'N, 47°09'E) and that the third millennium cities of Ur and Eridu have left cuneiform sources placing them on the sea although now they are 100 km from the head of the Persian Gulf. Lees and Falcon did not take into account sea level changes such as the postglacial rise of 100 m and interglacial rises of 30-100 m. Active growth of a delta at the head of the Gulf over the last 20,000 years may only have occurred from 10,000 to 2000 B.P. and again in the last 300 years. Subsidence levels are probably not as great as postulated (Vita-Finzi, 1978). Nevertheless, there were probably marshes to the north and they may have just become more available and extensive in recent centuries (Aqrawi, 2001). As Larsen and Evans (1978) and Wagstaff (1985) point out, the Persian Gulf shoreline at the head of the Gulf has been affected by, and rendered difficult to interpret by, a complex of factors including confusion of marine and freshwater fossils in an estuarine environment, subsidence, eustatic sea level fluctuations, local seismic activity, climate and therefore hydrologic changes, and cultural changes such as irrigation. Jacobsen (1960) detailed some of the changes in the courses of rivers and canals, based on evidence of ancient settlements which were presumed to be linearly arranged along water courses. Mallowan (1964) also maps some ancient river courses. The fish fauna has evidently had to cope with a changing availability of habitat through the post-glacial period. Floods and changes in river courses over this time have no doubt facilitated movement of fishes between Iran and the Tigris-Euphrates basin. It seems unlikely that the separate entry of rivers from Iran into the Gulf would have led to isolation of the faunas to any significant degree.

Canals and other irrigation structures have long been a feature of the Mesopotamian plains, forming habitats for fishes dating back thousands of years (Bagley, 1976). Their loss through natural and man-made disasters must have affected fish populations but sufficient natural habitat no doubt remained to ensure survival. The construction of dams upstream in Turkey and the large scale, modern drainage programmes in Iraq bordering Iran such as the "Three River Project" are drying up the extensive marsh systems and these are regarded as an eco-disaster leading to desertification in Iraq and adjacent regions of Iran (North, 1993; Pearce, 1993, 2001; Ryan, 1994; National Geographic, 185(4):unnumbered page, 1994; Scott, 1995; Munro and Touron, 1997; Maltby, 1999; Partow, 2001; www.amarappeal.com/documents/Draft_Report.pdf, downloaded 15 November 2001). The 32 km long "Fish Lake" was constructed as a barrier to Iranian attacks on Basrah. The Iranians dug several drainage ditches from "Fish Lake" northeast of Basrah to the Karun River, to dry up land for infantry attacks on Basrah. This whole marsh area of about 17,000 sq km, is the most important wetland in the Middle East and one of the top ten in the world. The Central and Al-Hammar marshes in Iraq by 2001 have had 97% and 94% of their land converted into bare ground and salt crusts. Less than one-third of the Hawr al Hawizeh (= Hawr al Azim in Iran) survives. It was estimated in the 1990s that the marsh area would be a desert within a decade and this seems to be an accurate assessment. The effects on the fishes in Iran are unknown but much habitat is being lost which could have served as a reserve against loss in Iran through natural and man-made changes.

The Iran-Iraq War of 1980-1988 severely damaged the Hawr al Hawizeh in Iraq, and presumably to some extent in Iran. Bombs and shells, chemical weapons, pollution, burning of reed beds, reed cutting and armoured boats used to smash through obstructing reeds all had deleterious effects (Scott, 1995). The Iraqi shores of this hawr have been drained by dyke construction and river control presumably for military reasons in this border area. Some marsh will survive in Iran because it is fed from wholly Iranian rivers but Iran News (19 February 1995) reports that draining of Iraqi marshes will lead to desertification inside Iran. Details on the restored Hawizeh marsh and its fishe sin Iraq can b found in Mohamed et al. (2008) and Abd et al. (2009).

The Southeast Anatolia Project (known as GAP after its Turkish acronym) incorporates 21 dams and 19 hydroelectric facilities including the massive Ataturk Dam on the Euphrates completed in 1993. It plans to draw off one-third of the waters originating in Turkey and will also use water from the Tigris River (Ottawa Citizen, 10 November 1994; Morris, 1992; Biswas, 1994; Beaumont, 1998). The reduction in flow for Iraq may reach 60%, especially when water is taken from the Euphrates or ath-Thawrah Dam (its reservoir is Lake Assad) at Tabqa in Syria (Vesiland, 1993). This will have major downstream effects, less so in Iran than in Syria and Iraq, but flow into the Shatt al Arab shared between Iran and Iraq will be greatly decreased perhaps allowing greater penetration of saline water and restricting migrations of fishes.

Between 20 and 15 thousand years ago, the Persian Gulf was dry as water was locked up in ice-caps and sea level was 110-120 m lower than today (Sarnthein, 1972; Kassler, 1973; Nützel, 1975; Al-Sayari and Zötl, 1978; Vita-Finzi, 1978). The floor of the Gulf was then thought to be a generally waterless, flat depression with a few swampy tracts rather than a "Garden of Eden" as has been proposed. A marine transgression occurred between 12 to 8 thousand years ago and by 6 thousand years ago the present sea-level was attained. Streams now isolated from the Tigris River basin by the sea in the Gulf and Hormuz basins would have been tributary to an extended Shatt al Arab, extending 800 km down the gulf to form an estuary at the shelf margin in the Sea of Oman, now under 110 m of sea. Earlier regressions no doubt occurred and facilitated the movement of fishes.

Construction of fish farms is widespread throughout this basin in Iran. For example in Lorestan Province, 772 tonnes were produced by the Lorestan Province Fishery Company in 1997, 50 fish farms were under construction and 125 pools built for aquaculture uses. The long-term aim was to increase fish production to 20,000 tonnes worth 156 billion rials and employing 10,000 people (Tehran Times, 22 September 1998). In Chahar Mahall va Bakhtiari Province in the highlands of this basin, 4360 tons of trout fingerlings were produced with plans to produce 8000 tons in future years (Tehran Times, 14 March 2005). The Indian carps Cirrhinus mrigala, Labeo rohita and Catla catla are being reared in aquaculture stations and are potential escapees into the natural environment (Gilkolaei, 2007).

Berg (1940) places this basin in the Mesopotamian Transitional Region, since the boundaries of three zoogeographical regions meet here, namely the Holarctic (i.e. its Palaearctic part), Sino-Indian (= Oriental) and the African (= Ethiopian). The Mesopotamian Transitional Region includes the Tigris and Euphrates basins and the Quwayq River, Syria, forming a single Mesopotamian Province. The province is transitional between the Mediterranean Subregion and the Indian Subregion. Genera such as Leuciscus (= Squalius) Aspius, Chondrostoma and Alburnus point to a Mediterranean or European association while such genera as Glyptothorax, Barilius, Mystus and Mastacembelus point to an Indian association.

Endorheic Basins

Bejestan

This basin comprises the drainages of the eastern highlands north of Birjand (32°53'N, 59°13'E) flanked by the Dasht-e Kavir basin to the west, the Dasht-e Lut and Sistan basins to the south, the Tedzhen to the north and the Afghan border to the east. The Tedzhen basin is separated by three ranges, from west to east, the Kuh-e Sorkh (35°30'N, 58°36'E) at 3017 m, the Kuh-e Bizak (35°11'N, 60°20'E) and the Kuh-e Khvaf at 2517 m east of Khvaf (34°33'N, 60°08'E). These receive snow in winter from moist Caspian Sea air. The highlands are relatively low compared with other parts of Iran and nowhere exceed 3000 m except for the Kuh-e Sorkh. The lowest points are in the sumps on the Afghan border at about 610 m. There are a number of minor sumps and the drainage patterns have been described as indeterminate. The total area is about 82,000 sq km. Tectonism commonly causes drainage disruptions (Krinsley, 1970).

The distinction of the western parts of the basin from the Dasht-e Kavir basin is somewhat arbitrary since the Kavir-e Namak near Bejestan (34°31'N, 58°10'E) lies at a similar level to the Kavir-e Bozorg and is separated by only a low rise in the land. This kavir receives intermittent streams from the east and north. The Bejestan basin does receive tributaries from Afghanistan but these are minor and do not begin to approach the input received by the Sistan and Tedzhen basins from the east. Streams drain mostly to the east, to three small terminal basins straddling the border; from north to south these are the Namakzar-e Khvaf, the Daqq-e Patargan and the Daqq-e Tondi.

The Dasht-e Lut basin to the south is separated by the drainage divide of the Birjand-Qa'in highlands, which trend north-west to south-east. Kuh-e Kalat is at 2605 m (34°18'N, 58°22'E) in the north-west and altitudes of 2779 m are reached in the south-east.

This whole basin has seasonal streams and a few springs with qanats a prominent feature. Water temperatures in qanats is 22-25°C year round and their is little fluctuation in water flow and chemical composition. Springs in contrast are influenced by the local geology and have a variable chemical composition, as well as being influenced by climate and pollution (Ruttner-Kolisko, 1964; 1966).

Caspian Sea

The Caspian Sea (Darya-ye Khazar, Darya-ye Mazandaran) basin is here taken to include both the rivers draining to that sea and the sea itself within Iranian territorial waters. This basin, in its land part, is elongate, extending from the Turkish border almost to the Afghan border and only acquires some width where the Safid River and its tributaries penetrate the Alborz Mountains in the west. According to Pirnia (1951) the Caspian basin in Iran (excluding the sea) encompasses 182,100 sq km while according to Zakeri (1997) this figure is 256,000 sq km, 15.5% of the whole country. Zakeri (1997) records 864 small and large rivers, including the Safid River with a catchment of 67,000 sq km. Much of the information on the Caspian Sea itself is restricted to waters of the former U.S.S.R. and there is relatively little on Iranian territorial waters. Rozengurt and Hedgpeth (1989), Kosarev and Yablonskaya (1994), Mandych (1995), Golubev (1996) and Ivanov (2000) summarise much of the recent Soviet literature, a general review is given by Mamaev (2002) and Bogutskaya et al. (2008) review early investigations of the sea and its fish biodiversity with special emphasis on the 1904 expedition led by N. M. Knipovich.

An ongoing and developing source of information on this sea, the surrounding land, its history, its management, biodiversity strategy and action plan, and a wide sweep of environmental problems is the Caspian Environment Programme (CEP), Baku, Azerbaijan at www.caspianenvironment.org. This site has numerous documents and reports on-line, some with authors, e.g. Katunin (2000), Ivanov and Katunin (2001), ERM-Lahmeyer International GmbH, DHI Water & Environment and GOPA Consultants (2001a), others appearing under CEP or TACIS (Technical Assistance to the Commonwealth of Independent States, European Union), e.g. TACIS and UNDP (2000), TACIS (2002), CEP (1998, 2000b, 2002). These reports include information on the fishes and fisheries but are best referred to for the interactions between people and the environment. Kiabi et al. (1999) describe the wetlands and rivers of Golestan Province at the southeast corner of the Caspian Sea. Razavi (1999) gives an introduction to the ecology of the sea in Farsi. Nezami et al. (2000) and CEP (2001) give recent general descriptions of the Iranian Caspian coastal zone, the important rivers, wetlands, water quality, climate, pollutants, and fisheries. www.bibliothecapersica.com/articlenavigation/index.html, under Caspian Sea, downloaded 24 December 2004 also gives an overview of this basin. Nadim et al. (2006) review the management of coastal areas in the Caspian Sea. Nasrollahzadeh (2010) reviews the ecological challenges facing thus enclosed sea and Allahyari (2010) the social sustainability of fishery cooperatives in Gilan..

The Caspian Sea is the largest "lake" or inland water body in the world at 436,284 sq km, a surface area encompassing 18% of the total area of all lakes in the world, about the same area as Great Britain (other surface area figures are 378,400 sq km, 384,400 sq km and 390,000 sq km - data of this nature varies quite markedly between apparently authoritative sources). The volume is 78,100 cu km, 44% of the total volume of inland lakes of the world. Its north-south extent is 1204 km and width is 204 to 566 km. The shoreline, including islands, extends for 7000 km, 1000 km of which is Iranian. The catchment area is 3.6 million sq km. Dumont (1998) presents arguments for this water body being a true lake and not a sea.

Caspian Sea with eastern edge of Black Sea on right and Kara Bogaz Gol on right. Lake Urmia (= Orumiyeh ) is at the lower left (turquoise) and Lake Van in Turkey lies to its west. Lake Sevan in Armenia is to the north of Orumiyeh. From NASA and Wikimedia Commons.

Caspian Sea with eastern edge of Black Sea on right and Kara Bogaz Gol on right. Lake Urmia (= Orumiyeh ) is at the lower left (turquoise)
and Lake Van in Turkey lies to its west. Lake Sevan in Armenia is to the north of Orumiyeh. From NASA and Wikimedia Commons.

 

North, Middle and South Caspian basins are recognised, divided by shoals. Iranian waters fall within the South Caspian Basin which occupies 148,700 sq km and is separated from the Middle Caspian by the Apsheron Bank. The South Caspian holds over 65% of the sea's water and is the deepest basin, to -1000 m in depressions, average - 325 m. The northern basin holds only 1% of the water.

The sea receives 291 cu km from river run-off and 87 cu km from precipitation but loses 374 cu km from evaporation and 11 cu km to overflow into the Kara Bogaz Gol (Gerasimov, 1978b). The Volga River accounts for 76.3% (82% according to Dumont (1995)) of the inflow of rivers, the Kura River 4.9%, the Ural River 3.7%, the Terek River 3.2% and the remaining rivers including all those of the Iranian shore 11.9%. Iranian rivers account for only 5% of the Caspian inflow, Iran has 7% of the catchment area, 14% of the coast, contributes 3% of the settling solids, and 2% of the fishery (Badakhshan and Shayegan in Glantz and Zonn, 1997). The Volga has its headwaters near Moscow and is 3688 km long with a catchment area of 1,360,000 sq km and a mean annual flow at Volgograd of 8380 cu m/sec. The Volga is of prime importance in the Caspian Sea basin to migratory fishes as a spawning site and the biology of these species has been studied extensively. Often these studies provide the basis for much of the knowledge of Iranian fishes to the south.

Zenkevi(t)ch (1957; 1963) and Barimani (1977) have reviewed the geography, hydrology and biology of the Caspian Sea, Moiseev (1971) summarises the living resources of the whole sea, Karpinsky (1992) aspects of the benthic ecosystem, and Knipovich (1921), Iljin (1927a), and Nevraev (1929) give accounts of Iranian coastal waters and regional fisheries in the early twentieth century. Zahmatkesh (1993) describes the gammarids and bottom sediments, Fallahi (1993) the plankton and Soleimani (1994) the benthic fauna in Iranian waters. Mamaev (2002) is a recent general overview.

Water balance for this sea depends on a delicate balance of inflow, evaporation, precipitation, climate, and abstraction for human needs. Water 10 m deep or shallower has a bottom of sand and gravel while at greater depths of 50-100 m clay and softer sediments increase. There is more sand in these greater depths off Gilan compared with off Mazandaran.

Maximum depth is 1025 m, mean depth is 184 m, and depth below sea level is -28 m (-27.66 m averaged over the past 2,500 years according to Dumont (1998)). There are natural water level fluctuations - the figure cited is from 1983; in 1978 it was -29.02 m, the lowest recorded since observations began (Voropaev and Velikanov, 1985). Petr (1987) has pointed out that a decline below -28.5 m would result in a change in salinity distribution and in water currents mixing riverine and sea water. A decline in productivity would follow. A fall of only 1 m would cause a 60% reduction in fish food supply and, since this fall poses barriers to migration to better feeding grounds, a further 20% loss in food supply. Recently however, since 1978, the sea has begun to rise, by 2.1 m from 1978 to 1993 to -26.95 m, with a possible rise of 3 m in the next 25 years. Vaziri and Borghei (1995) give an average rise of 1.2 cm a month for the period 1986-1993. The sea rose 26 cm in 1994. However, over the past 2500 years the sea level has not exceeded -25 m and is not anticipated to do so in the near future; the level is cyclical (Rychagov, 1997; Gorji-Bandpy and Hooman, 2004). The reason for the rise is probably a climatic shift (Mandych, 1995; Shayegan and Badakshan, 1996; Kobori and Glantz, 1998) but a sheen of oil from pollution may be helping in the reduced evaporation of 7-10% observed over two decades. Tectonic shifts of the sea floor may also be a contributing factor. Predictions of water level changes have proved unreliable so schemes to ameliorate rises or falls are unwarranted and could be catastrophic (Abuzyarov, 1999). Georgievskiy (2001) however, predicts a lowering of the sea level to -27.6-28.9 m by the year 2030 from -27.0 m in 2000. Klige and Myagkov (1992) examined the water balance of the Caspian Sea and predicted a rise in sea level to 1995-1997 and then future declines of the order of several metres in the next century.

The rise in water level is engulfing buildings including industrial sites which will pollute the waters of the Caspian further. Iranian towns and cities damaged include Babolsar, Tonekabon, Ramsar, Ashuradeh, Bandar-e Torkoman, Anzali, Astara and Kolachai (Zonn in Glantz and Zonn (1997)). Fish caught near Nowshahr in 1999 were contaminated with oil pollutants (Tehran Times, 1 November 1999). The complex of chemical, petrochemical and metallurgical plants at Sumgait near Baku in Azerbaijan produces 335,000 tonnes of mostly toxic waste including dioxins. Hundreds of waste lakes of oil near Baku are being slowly engulfed by the rising Caspian. Nasrolazadeh Saravi (2001) and Khatoonabadai and Dehcheshmeh (2006) describe oil pollution in Iranian coastal waters although it is much less than near Baku, particularly in Mazandaran and Golestan. Heavy metals enter down the major rivers from mining and industry and the effects from the Kura River may have rendered the coast of Azerbaijan almost untenable for life (Bickham, 1996; Pohlman and Naismith, 1996; Rowe, 1996). Radioactive waste, both liquid and solid, is found in low lying depressions around nuclear power plants and is liable to enter the Caspian (Rodionov, 1994; Dumont, 1995).

On the plus side, sturgeons may benefit from easier access to spawning grounds (Ottawa Citizen, 9 July 1994; 3 July 1995) but this is probably offset by the pollution load of the major spawning rivers.

In contrast to the recent rise in sea level, a series of reports have appeared in past scientific and popular literature on the falling level of the Caspian Sea and diversionary schemes to combat this (e.g. Kovda, 1961; Lamb, 1977; Hollis, 1978; Gribbin, 1979; Micklin, 1979; 1986; Golden, 1982; Rich, 1982; 1983; Voropaev and Kosarev, 1982; Voropaev and Velikanov, 1985; Pearce, 1984; Ryan, 1986; Perera, 1989; Rozengurt and Hedgpeth, 1989; among others). The Caspian dropped 2.3 m between 1930 and 1962 and area has decreased by 10% or 40,000 sq km. Recent historical levels appear to be between -25 and -26 m, average -25.8 m. Changes in level of the Caspian due to natural or other causes in historical and pre-historical times have been reviewed above. Fall in the sea level increases salinity, destroys habitat and blocks spawning migrations, although some effects are less in the southern, Iranian Caspian because of the larger water mass. The Volga accounts for 76% (some reports say more than 80%) of the river input to the Caspian Sea. The Volga is now extensively dammed, as are other rivers in this basin, and its waters used for industry and agriculture. There are 8 large dams on the Volga, the largest being the Kuibyshevskaya with a reservoir area of 6450 sq km and a total volume of 58 cu km. Dams in the Caspian basin provide almost one third of the hydropower of the former U.S.S.R. (Rozengurt and Hedgpeth, 1989). Flow into the Caspian has been cut by at least 25% and in spring, the time of spawning migrations, by as much as 37% for the Volga-Kama systems. Berka (1990) reviewed the effects of water level changes on the northern Caspian fisheries. The North Caspian was designated as an "ecological disaster area" in 1992 because of water pollution input from the Volga. The delta is eutrophic with cyanobacterial blooms being common, affecting fish survival (Saiko in Glantz and Zonn, 1997).

The decline in sea level has been reversed in recent years and a rise of nearly 2 m was reported and, in Turkmenistan, a shoreline advance of 2-3 km in places (Rich, 1991; Anonymous, 1992a; Golub, 1992; Ottawa Citizen, 9 July 1994; Priroda, 5:3-25, 1994). This will have positive effects for some fisheries and wetland conservation but negative effects on recent, low-lying construction including oil refineries and wells in Azerbaijan and a nuclear waste dump in Turkmenistan which would cause massive pollution from oil and radioactive compounds (Pearce, 1995). Environmental hazards to the fisheries caused by sea level rise include eutrophication from farmland covered by the sea, pesticides and herbicides from inundated farmland, salt water penetration into wetlands, input of solid municipal and industrial wastes and vegetation, destruction of fish habitat, and input of soil altering the ecosystem (Shayegan and Badakhshan in Glantz and Zonn, 1997).

It has been suggested that the rise in sea level is due, in part, to seepage from the Aral Sea basin and that this could be halted by setting off underground explosions. This smacks of the large-scale alteration to the environment favoured by Soviet planners to combat the fall in sea level - both are grandiose and have unknown consequences for the environment. Climate change is probably a major factor abetted by the closing off of the Kara Bogaz Gol (responsible for an estimated 40-45 cm rise alone) and diversion of Siberian rivers into the Ural River in the northeastern Caspian (Khan et al., 1992).

Much of the former southern U.S.S.R. is water poor and a solution to this and the falling Caspian level has been advocated. This would involve diversion of north flowing Siberian rivers at a cost $40 billion. The potential for environmental damage on a local and even global scale caused this scheme to be shelved in 1986. The project involved excavations using nuclear explosives, drowning of forests and construction of canals thousands of kilometres long. Reduced flow into the Arctic Ocean could affect ice cover which influences atmospheric pressure and circulation patterns over the whole northern hemisphere. This Soviet plan has recently been revived (Pearce, 2004).

There is an abundance of historical and other evidence for variations in Caspian Sea level and its connections with other water bodies in both recent times and over several million years (Huntington, 1907; Ehlers, 1971; Lamb, 1977; Gerasimov, 1978b; Hsü, 1978; Coad, 1980c; Rögl and Steininger, 1984; Wossugh-Zamani (1991c); Oosterbroek and Arntzen, 1992; Sal'nikov, 1995; Mamedov, 1997; Rychagov, 1997; Caspian Environmental Programme, 2000; Grigorovich et al., 2003; Kotlík et al., 2008). Brooks (1949) maintains that the Oxus (= Amu Darya) flowed into the Caspian in the 14th century instead of the Aral Sea. Shnitnikov (1969) and Gerasimov (1978a) report flow along the Uzboi channel north of the Iranian border into the Caspian from the Aral Sea basin at several periods from the third millennium B.C. to the 16th century. Sal'nikov (1998) illustrates connections between the Amu Darya and the Caspian Sea from the Pleistocene to the 20th century. The connection between the Caspian and Amu Darya and Aral Sea was interrupted about 20,000 years ago when the Amu Darya turned north, was reconnected about 10,000 years ago, and essentially interrupted about 4000 years ago. These regular contacts have resulted in an Aral Sea ichthyofauna with "weakly pronounced endemics", although the Amu Darya ichthyofauna has a number of clearly defined endemics which are not yet found in the Caspian Sea basin (but see below under Tedzhen River basin). Dunin-Barkovsky (1977) records level fluctuations of up to 50 m during the Holocene due to variations in the general moistening of Eurasia and intermittent warming and cooling variously associated with changes in precipitation and evaporation. Ice melt from the Fennoscandian ice cap, as late as 4000 B.C., added large volumes of water to the Caspian and an overflow to the Black Sea was then possible. Berg (1948-1949) maintains that Atherina presbyter (=caspia) and Syngnathus caspius entered the Caspian at about this time. Some fishes, such as Salmo trutta (as then recognised), are probably immigrants from Arctic regions and certain cyprinoids and percids are freshwater immigrants. Bianco (1990; 1995b) points out that, at every glacial- interglacial ice melting phase, a network of connected rivers and lakes allowed primary freshwater fishes to disperse in the northern Palaearctic. Other fishes are relicts of earlier transgressions. Such species as herrings (Clupeidae), gobies (Gobiidae) and possibly sturgeons are believed to have evolved from the marine fauna of the Tethys Sea which ran from the modern Atlantic to the Indian Ocean before the Sarmatian basin formed. The uplift of eastern Anatolia and the Alborz in the Early Miocene between 20 and 17 million years ago (MYBP) closed a seaway from the Indo-Pacific which had extended into the Eastern Paratethys (= Black-Caspian-Aral sea in modern terms). The connection reopened in the Middle Miocene 16.8-16 MYBP) but by the Late Miocene a Sarmatian basin was cut off from the open seas and developed a unique marine fauna (Ekman, 1953). This was mostly lost as salinity decreased from freshwater input and a new fauna developed. A series of connections and breaks with the Black Sea, Mediterranean Sea and the Atlantic Ocean in various combinations with brackish and freshwater episodes gave varying opportunities for faunal interchanges and evolution. The Caspian fauna differs from the Mediterranean one because its only communication was via the Black Sea which acted as a "filter". When the Black and Caspian seas were well connected, the link to the Mediterranean was broken, and when the Black and Mediterranean seas were connected, the Caspian connection was not well developed. Mamedov (1997) and Rychagov (1997) review late Pleistocene and Holocene changes in Caspian Sea level, Chepalyga (1984) and Gerasimov (1978b) review water level changes and connections with the Black Sea over the last 80,000 years, Kosarev and Yablonskaya (1994) and Mandych (1995) for the last 500,000 years and Grigorovich et al. (2003) for the last 12.5 million years. Bianco (1990) gives an overview of the palaeohistory of the Paratethys Basin. Fluctuations in water level are correlated with climate changes Kotlík et al. (2008) using multiple gene phylogeography found the Black and Caspian seas supported separate populations of Rutilus frisii during the last glaciation, although this separation was not complete and gene exchange occurred, with the majority of migrations in the Pleistocene.

The total Caspian Sea drainage area is said to be 3,700,000 sq km, about 25% of the continental land mass of the U.S.A. (Rozengurt and Hedgpeth, 1989). The basin includes about one fifth of the crops and one third of total industrial output of the former U.S.S.R. (Rozengurt and Hedgpeth, 1989). Its northernmost waters are north of St. Petersburg (= Leningrad) in Russia while its southernmost waters rise on the flanks of the Zagros Mountains in Iran. This ranges from the subarctic to the subtropical region and is very diverse in climate and geology. Natural runoff in the South Caspian Basin ranges from 8 to 18 cu km while in the North Caspian it is 207-375 cu km. However the North Caspian is very shallow (mean 4-5 m, maximum 20-25 m) compared to the south Caspian (mean 325-334 m, maximum 980-1025 m). This is also reflected in the volume, 400-700 cu km compared to 49,000-77,500 cu km. Salinity is about 12-13‰, increasing in isolated bays and decreasing near river mouths. Summer temperatures in the south reach 27°C and in winter 9°C but the northern parts ice over. The Gorgan River area reached 30.9°C (Laloei, 2006). Surface water temperatures for the South Caspian are reported as 7.0-10.3°C in winter, 7.9-14.0°C in spring, 25.0-29.0°C in summer and 12.0-19.0°C in autumn (Rozengurt and Hedgpeth, 1989). These authors also report salinity ranges of 12.5-13.0, 12.3-13.2, 12.6-13.6 and 12.3-13.5‰ for the same seasons, oxygen levels of 7.0-7.8, 7.0-8.2, 5.0-6.0 and 6.0-8.0 ml/l, and pH values of 8.48, 8.44, 8.44 and 8.50. Vertical mixing occurs down to 50-150 m in the South Caspian (Mellat-Parast, 1992). There is little oxygen below 200-300 m and no fish life although changes to the hydrological regime of the Volga have increased aeration and oxygen content of deeper layers in the south Caspian, down to 600-800 m. The Caspian has no tides but sustained winds can cause seiches, local and temporary rises in sea level. There is a current along the Iranian shore from west to east. The shelf along the Iranian coast is narrow (6-10 km) and steep (Kosarev and Yablonskaya, 1994). Beaches are usually sand with shell gravel on the bottom further out. The extreme western coast has some shingle beaches and west of Alamdeh in the central part is some rocky shore but there are no major cliffs or headlands. The shore has coastal dunes, spits and bars with lagoons inland, either brackish or fresh, grading into the higher and dryer foothills.

Much of the coast was once forested, but it has been actively cleared and marshes reclaimed as rice paddy. Rice paddies are now being investigated for fish cultivation. About 300-500 kg of carp "seed" and a 10% increase in paddy production per hectare was recorded during the rice cultivation season. Extending this into the fall gave a production of 750-1000 kg of fish and duck and in winter 5.5-8.0 t of rainbow trout (Tehran Times, 1 October 2000). Gilan is attempting a production of 2 kg of trout per sq m of paddy field, with the aim of harvesting 46,000 t of fish (IRNA, 14 November 2001). Mazandaran has the highest farm fish production in Iran at 28,000 tonnes (2006-2007) and is expected to reach 50,000 t by 2010 (www.mehrnews.ir, downloaded 8 February 2007). The area of forests in northern Iran has been reduced from 3.4 million hectares in 1962 to 1.8 million hectares in 1977 and about 1 million hectares or less in 1995. In Gilan, 975,000 cu m of wood from the forests are burnt annually by cattle breeders for heating or cooking purposes or for production of dairy products. Additionally 450,000 cu m of wood are used for industrial purposes. Reforestation cannot keep up with the losses and forests have been reduced by half over the past 50 years (Barzegar, The Agricultural and Cattle Breeding Publication, No. 761, 22 December 1997, from www.netiran.com/Htdocs/Clippings/Deconomy/971222XXDE01.html). As a result floods now occur with destruction of fish habitat after 30-40 hours of rain where previously no flooding occurred after even 4 days of rain (Hamshahri, Tehran, 628, 20 February 1995). Abstraction of water for irrigation (60% of water use) has severely reduced water levels and runoff rates necessary for reproduction of fishes. Estuarine habitats have been degraded inhibiting the survival of eggs, larvae and juveniles of anadromous and semi-anadromous fishes (the latter are species which spawn in the lower stretches and deltas of rivers where salinity is optimal at 8 g/l for many commercial species, e.g. Sander lucioperca, Cyprinus carpio, Rutilus caspicus). Over 90% of coastal streams along the Caspian shore are dry in July in Iran because of irrigation demands. As a result larvae of spring spawners are flushed into fields where they die, migration and late summer spawning of Aspius aspius and Luciobarbus brachycephalus are obstructed, and Salmo caspius and Rutilus frisii kutum populations are depleted because they cannot spawn in the shallow, warm, weed-choked water. Nursery and reproductive areas for Abramis brama, A. sapa, Blicca bjoerkna, Aspius aspius, and Sander lucioperca among others are confined because of their low tolerance to salinities above 7-8‰. Without an adequate runoff, the sea encroaches on the estuary. Nasri-Chaari (1994) cites physical obstacles, sand removal from river banks, overfishing and water pollution for declines in fish migration in recent years.

An earlier, general work including fishes of the Iranian Caspian Sea and coast is Berg (1948-1049). More recent works are the atlas of the fish species in the Iranian Caspian Sea in English and Farsi by Jolodar and Abdoli (2004) and that on the biodiversity of the southern basin by Abdoli and Naderi (2009).

The commercially important species of fish were summarised in Abzeeyan, Tehran, 5(7):VII-IX (1995) and are divided into sturgeons (Acipenseridae, 4 species) and bony fishes (3 species of kilkas in the genus Clupeonella of the family Clupeidae; herrings or Alosa spp. also in Clupeidae; 5 species of the family Cyprinidae namely Rutilus frisii, Cyprinus carpio, Abramis brama, Rutilus rutilus (and presumably R. caspicus) and Aspius aspius; 2 species of mullets, family Mugilidae, Liza auratus and L. saliens; a member of the perch family, Percidae, namely Sander lucioperca; and a member of the salmon family, Salmonidae, namely Salmo trutta (= caspius)). About 70% of Rutilus frisii is caught in Gilan Province, while 60% of mullets and 75% of sturgeons are caught in Mazandaran Province. More than 50% of the sturgeon catch is Acipenser stellatus and 10% is Huso huso, the remainder being A. gueldenstaedtii and A. persicus, with a yearly catch for all sturgeons of about 2500 tonnes. Sturgeon fishing is carried out by the government and no private sector fishing is allowed because of the value of this fishery and the need for careful management. Accidentally caught sturgeon must be released or turned over to the government operation. Ivanov (2000) summarises the biological resources of the Caspian Sea from a Russian perspective with some comparative figures from Iran. Generally, catches in Iranian waters are always less than those in former Soviet Union countries combined. A particular exception is Rutilus frisii (safid mahi), an esteemed fish in Iran.

About 25% of the Iranian total fish catch is from the Caspian coastal area (CEP) and figures for the Iranian Caspian Sea in tonnes are:-

Year All fish species Kilka Sturgeon flesh Caviar
1976/77 8,428 1131 2368 221
1981/82 10,466 1341 1914 234
1986/87 11,084 2384 2500 303
1991/92 34,596 13,817 2208 283
1992/93 40,598 21,527 2198 262
1993/94 52,768 28,730 1170 217
1994/95 69,700 51,000 1700 218
1995/96 58,300 41,000 1500 182
1996/97 74,100 57,000 1600 195
1997/98 76,200 60,400 1300 151
1998/99 101,500 85,000 1200 157

The fish harvest from the southern Caspian coast of Iran for the 7 month period October 1999-April 2000 dropped by 11% over the same period from the year before, from 8630 t to 7710 t (IRNA, 10 May 2000). The decline was attributed to a rise in fish prices which encouraged illegal fishing and to habitat loss. The value for the whole Caspian fisheries is given as $6 billion by Nezami et al. (2000). A proposal for a Caspian Fisheries Commission is given by TACIS (1999; 2000b) and ERM-Lahmeyer International GmbH et al. (2001b). It would aim to conserve and utilise the living aquatic resources, including the management of fish stocks such as kilka, herrings and mullets, as well as the famous sturgeons. These species all have transboundary stocks requiring cooperative management between countries. Articles aim to protect traditional fishing for sturgeon along the Iranian coast, establish state monopolies for the export of caviar, set up cooperative research programmes to conserve sturgeon species, establish annual total allowable catches and fishing regulations, and so on.

About 50,000 tonnes of kilkas are caught each year by the Industrial Fishing Company and fishing cooperatives using deep conical nets and air lifting with artificial lights as attractants. About 20,000 t of other species are caught by licensed cooperatives using beach seines and gill nets although a report in IRNA (27 March 2000) cites more than 16,000 t including whitefish (Rutilus), Mugilidae, Cyprinidae, "anchovy" (sic), bream (Abramis) and zander (Sander). An account of site selection for beach seining is given by Zanoosi (1993). Beach seining has been restricted to the period from sunrise to 8 p.m., and to 10 p.m. in Miankaleh (www.iranfisheries.net, downloaded 14 November 2006). The 1994-1995 finfish catch (excluding sturgeon and kilka) using gill nets, coastal purse seines and beach seines, was 17,000 t, perhaps over 22,000 t with the illegal catch included. About 87% of this catch is Rutilus frisii kutum, Liza auratus and Liza saliens (Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 37, 1996). Gill nets showed a 39% decline compared to the previous year and beach seines were 16% less. Rutilus frisii kutum comprised 53%, mullet 39% and others 8% of the total catch (Abzeeyan, Tehran, 6(5, 6):IV, 1995). The harvest from the southern Caspian Sea coast dropped 11% in the year 2000 from the same seven month period in the preceding year, to 7710 t, as a consequence of poaching, neglect of river maintenance, and substandard capture methods (IRNA, 10 May 2000). The catch in Golestan Province rose from 470 t in 2000 to 3278 t in 2005, attributed to artificial propagation, restrictions on beach seining, training about closed seasons and beach seine standards, increased fishing effort, and a favourable climate (www.iranfisheries.net, downloaded 14 November 2006).

There are 5 regional fishing centres namely Bandar Anzali with 14 fishing stations, Keyashahr with 12 stations, Babolsar with 13, Ashuradeh with 9 and Nowshahr with 9 (Iranian Fisheries Research and Training Organization Newsletter, 7:7, 1995). The Caspian Environment Programme (2001c) gives 15 stations for Bandar Anzali, 9 for Keyashahr, Babolsar, Ashuradeh and Nowshahr for sturgeon fisheries. Fixed gill nets are used with a standardised mesh. The Ashuradeh Peninsula, where more than half of Iran's caviar is processed, was threatened by the rising Caspian Sea in a 1991 radio report. A 1995 agreement between Iran, Azerbaijan, Turkmenistan, Kazakhstan and Russia gives each nation an exclusive fishing zone of 20 nautical miles from shore (Iranian Fisheries Research and Training Organization Newsletter, 7:7, 1995).

Inland freshwaters of Gilan are divided into three categories by Bakhshizod-Mahmoodi (1996): natural and impounded ponds, the Safid River reservoir, and wetlands. The ponds are used primarily for cyprinid and acipenserid culture, the reservoir is fished by seining, by spreading wheat grains in littoral areas to attract fish and by using the shemshad or shaghoul net (a giant dip-net), and the wetlands are fished by seining, by the salik or mashak (cast-nets), by the la'kesh (drifting gill net using one and two boats), by fixed gill nets, by the shemshad and by angling (for ordak mahi).

Pollution is an important factor in the ecology of the sea, from offshore oil drilling, ship discharges of oil wastes and contaminated water as well as garbage and even discharges from ship collisions, radiation from underground, non-military explosions and nuclear waste dumped in inflowing rivers (radiation levels are 100 times above normal (Time, 1 November 1993)), manure and pesticides from farming on the surrounding land mass, city waste water, sewage and garbage, industrial wastes including mercury and other heavy metals, discharges from water desalinating plants, extraction of minerals such as sodium sulphate, mirabelite and espomite, and untreated sewage (see Sardar (1979), Nuhi and Khorasani (1981), Coad (1980c), Khalili (1994), Raiss-Tousi (1999), Namazi (2000), Abaee (2001), Charamlambous (2001), Laloei (2006), Zeynali (2009) and Saeidi et al. (2010) for Iranian problems and acceptable levels of some elements; Anonymous (1988c), Edwards (1994), Specter (1994) and Kasymov and Rogers (1996) for former Soviet waters; Stone (2000b) is a recent, short general overview).

Data collected in 1991 showed the Caspian Sea received effluents comprised of 3000 tonnes of oil products, 28,000 t of sulphites, 315,000 t of chlorides, 200,000 t of tar and 25,000 t of phenols (Namazi, 2000). In Daghestani rivers, the same author records heavy metals, pesticides, phenol, arsenic, boron and selenium, among others, at 60-100 times the maximum permissible for fisheries. The oil industry is considered to be the main source of ecological problems in the Caspian Sea (Karpyuk, M. and Shavandin, V. 1996. Astrakhaners on the Caspian Sea. International Affairs, 42(1) from http://home.eastview.com/ia/42_01_15.htm). Prospecting uses blasting operations which have caused sturgeon deaths on more than one occasion. A single offshore well during its life releases into the water 30-120 tonnes of oil, 200-1000 t of sand, clay and other waste and 150-400 t of drilling mud paraffin fractions, baryta, lime, detergents, emulsifiers and lubricants. The ecology is affected 5-12 km from each well. The oil industry in the Caspian has reserves estimated at $4 trillion and a new oil rush will further contaminate the sea.

Charamlambous (2001) concludes that municipal wastewater from 11 million people is the primary pollutant in Iranian coastal waters. Industrial discharge accounts for 31%. of organic loading, the rest being municipal discharge. The most industrialised area is around Rasht with waste going into the Anzali Mordab. The Zarjub River in Rasht is the most polluted river in Gilan, and possibly in Iran (Ghodrati et al., 2007). TACIS (2000c) reports that in Gilan, 32 of 36 major cities discharge wastewater untreated into a river and 89 of 90 industries discharge treated wastewater to a river. Ayati (2003) also reviews pollution in the mordab. Mirkou (2001) details agro-chemical usage along the Caspian shore comprising various fertilisers and pesticides. Naderi Jeloudar et al. (2007), Varedi et al. (2007) and Amirkolaie (2008) describes the environmental impact on the Haraz River of aquaculture waste water discharge from rainbow trout farms - pollution levels in this instance were generally too low to have a significant impact of the river system although phosphorus loading was increased and levels varied with activity rates of the farms.

Chlorinated pesticides have been used in anti-malarial campaigns throughout Iran and to eliminate pests on cotton, rice and other products in Mazandaran. Herbicides and pesticides are widely used in rice paddies. DDE, DDT, DDD, Lindane, Dieldrin, Eindrin and Kelthane have been identified in such rivers as the Babol and Chalus (Annual Report, 1995-1996, Iranian Fisheries Research and Training Organization, Tehran, p. 11-13, 1997). Ebadi and Shokrzadeh (2006) examined Rutilus frisii, Vimba vimba (= V. persa), Clupeonella delicatula and Liza aurata for lindane at Chalus, Babolsar, Khazarabad and Miankaleh but levels detected were less than the FAO/WHO recommended permissible intake and were no cause for public concern. Similar studies on DDT and DDE and on chlorobenzilate from the same sites and fish and levels were also less than the permissible intake (Shokrzadeh and Ebadi, 2005; 2006). Shokrazadeh et al. (2009) also found that levels of Lindane in dorsal muscle of safid mahi, kefal, kuli and kilka species were less than FAO/WHO recommended intake. The Chalus River also contains various heavy metals, such as lead, zinc, copper, iron, cadmium and chromium from mining activities (Annual Report, 1995-1996, Iranian Fisheries Research and Training Organization, Tehran, p. 18, 1997). Zeynali et al. (2009) demonstrated the presence of copper and zinc in muscle tissues of Liza aurata, Rutilus frisii kutum and Cyprinus carpio from Chalus, Anzali, Rudsar and Fereydoon Kenar in the Caspian Sea basin although levels were acceptable for human consumption. Hashemy-Tonkabony and Asadi Langaroodi (1976) have shown the presence of DDE, DDT, TDE, Dieldrin, Lindane, Aldrin and Heptachlor in a wide variety of Caspian fishes in Iran. However, Ebadi and Shokrzadeh (2006) examined Rutilus frisii, Alburnus, Clupeonella and Liza species in Mazandaran for the organochlorine pesticide lindane and found levels in muscle tissues to be less than FAO and WHO recommended permissible intake and so were not a public concern. Rutilus frisii, Cyprinus carpio, Liza species and Acipenser stellatus were tested for DDT, aldrin and heptachlor with only the latter slightly elevated above standard levels at Hashtpar (Iran Daily, 11 January 2006). Phytoplankton diversity in the western Caspian Sea fell from 74 to 40 species, biomass from 8.7 to 2.1 g/ sq m and biomass of benthic organisms in coastal areas fell from 1724 g/ sq m in 1961 to 21 g/sq m in 1969 (Clark, 1986). These declines were noted particularly in the nursery grounds for sturgeon, Abramis brama, Esox lucius and Cyprinus carpio among other fish species. In the 1980s, catches of Abramis brama, Cyprinus carpio, Rutilus rutilus (presumably R. caspicus) and Sander lucioperca fell by as much as 80% and Salmo trutta (= caspius) and "shad" had almost disappeared. It was estimated that for 1985, 10,200 tonnes of oil products and 104,200 t of sewage were dumped in the sea. One-fourth (or 40 billion cubic metres) of all the wastewater in Russia enters the Caspian Sea and petrochemical factories alone release 67,000 t of waste annually (Anonymous, 1988c; Platt, 1995; Hamshahri, Tehran, 3 (639), 7 March 1995). Salinity increased as more water was taken for irrigation - two-thirds of the Terek and Kura flows did not reach the sea (Markham, 1989). In Iran, sewage is discharged into the Caspian Sea from coastal towns, and via rivers, from towns inland. Industrial solid wastes enter the sea through the larger rivers such as the Safid, Gohar and Siah as well as the Anzali Mordab complex. The use of agricultural chemicals such as fertilisers and pesticides has led to pollution, e.g. in Gilan Province 88,851 t of fertilisers were used in the year 1992-1993, an 18.7% increase over the previous year. A survey of 30 towns in Gilan shows that 80% of rubbish dumps are located by rivers, marshes or the coast (Hamshahri, Tehran, 3 (639), 7 March 1995). An estimated 200,000 fish were killed in the Kacha River, a branch of the Siyarud in Rasht, poisoned from a dump in the Saravan region which receives 390 t of rubbish daily. Heavy rains had washed poison into the river (Tehran Times, 7 October 1998). As many as 1000 trout (presumably mahi azad, Salmo caspius) died in the Kileh River in Mazandaran from release of wastes from a dairy manufacturer; sand extraction was also blamed for affecting fish populations (Iran Daily, 21 July 2005).

The biology of the Volga River and its effects on the Caspian ecology has been reviewed by Rozengurt and Hedgpeth (1989) and Pavlov and Vilenkin (1989). This river is of critical importance for marine fisheries. Fish production is less in the central and southern parts of the sea as nutrient supply comes from upwelling and circulation rather than a riverine input. However the Volga has effects even here, changing the Caspian Sea from its regime in the 1950s. Abstraction of water for irrigation, industry and household use caused salinity increases of about 0.2-0.3‰, increased aeration of deep layers and in their oxygen content down to 600-800 m by as much as 2-3 ml/l due to convection and thermal winter mixing, an increase in the euphotic zone to 50 m and depths open to total photosynthesis to 100 m, a decrease in organic matter and its vertical gradient, and an increase in wind-driven circulation and its effects on temperature and salinity layers. In the period 1956-1972, the Caspian Sea was transformed from a fishery based on valuable species (listed above) to one dependent on kilka which now occupies 80% of the catch (or 107 times the catch in 1930). Even including the kilka, catches in the 1970s were 245 x 103 tonnes or only 37% of the 1913 catch. The catch of Caspian herrings (a complex of species in the family Clupeidae) ceased to exist commercially by the 1970s and in fact was banned. In 1967-1972 it was 0.6-2.1 x 103 compared to 56-62 x 103 in 1945-1953 or 82-307 x 103 in 1900-1917 (Rozengurt and Hedgpeth, 1989). Moghim et al. (1994) report that, in the southern areas of the Caspian Sea, nearly 90% of the catch is composed of Rutilus frisii, Liza saliens and Liza aurata (with biomasses of 24,000, 7000 and 2400 t respectively and maximum sustainable yields of 7000, 2900 and 960 t  respectively). The Volga is a major pollutant of the Caspian Sea, carrying sewage, agricultural waste, PCBs, petrochemical wastes, tannery waste, etc. from a population base of 60 million people (Golub, 1992). In 1989, 40 million t of polluted wastewater entered the Caspian via the Volga River, more than a quarter of all the wastewater of Russia (http://www.oneworld.org/patp/pap_overview.html). A report in 1995 gives the volume of pollutants and industrial wastes entering the Caspian Sea each year as 11 billion cu m. Russia accounts for 50%, Azerbaijan 16% and Iran 11% (http://netiran.com/news/IRNA/html/950731IRGG17.html).

The Volga-Don canal in the former U.S.S.R. connected the Caspian Sea with the Black Sea in 1952 and formed an invasion route for various benthic organisms while others came in attached to boats transported by rail or were deliberately introduced (Kasymov, 1982). The molluscs Abra ovata and Mytilaster lineatus, two invaders, accounted for over 90% of the total benthic biomass. Invaders provided 95.1-99.3% of the total benthic biomass in the western part of the south Caspian Sea in 1976. East of the mouth of the Safid River, the Azov-Black Sea molluscs Abra ovata and Cerastoderma lamarcki accounted for 80% of total benthic biomass. In Gorgan Bay, 99.9% of the benthos fauna is comprised of invaders. The Volga is also connected to the Baltic and White seas via the White Sea-Baltic Canal opened in 1933 (Pavlov and Vilenkin, 1989).

The fisheries may well collapse if the 10 cm long ctenophore or comb jelly Mnemiopsis leidyi from the northwestern Atlantic Ocean enters the Caspian Sea via the Volga-Don canal in ballast water. It reached the Black Sea in the early 1980s and destroyed the local pelagic food chain (Travis, 1993; Dumont, 1995; Pearce, 1995; GESAMP, Joint Group of Experts on the Scientific Aspects of Marine Environmental Protection, 1997; Negarestan et al., 2002; Kideys, 2002a; 2002b; 2003). The ctenophore eats fish eggs and larvae directly as well as zooplankton and crustaceans which are foods for fish (Bagheri et al., 2005). The Black Sea fish catches fell 90% in 6 years and the biomass of the ctenophore reached an estimated 900 million tonnes, ten times the world annual fish catch (or 1 billion t, about equal to the world fish catch - sources differ). The wet weight biomass of the whole Black Sea at times was 95% ctenophore. This suggestion of the mid-1990s was borne out, as detailed below. A continuing series of reports, magazine articles and studies on this invader are not all cited here.

The earliest report for the Caspian appears to be in 1995 by the Iranian Fisheries Research Organization (Bilio and Niermann, 2004; www.caspianenvironment.org/mnemiopsis/mnem_attach13.htm). The Islamic Republic News Agency (IRNA) on 12 May 1998 reported that a number of jellyfish had been observed in the Caspian Sea recently, presumably brought in the ballast of oil tankers, and its occurrence is documented by Esmaili Sari et al. (1999) and in numerous other studies by this author and co-authors. Various studies on the biology of the comb jelly and its impacts have been carried out in the Iranian Caspian Sea including, e.g. Movahedinia et al. (2002), Esmaeili et al. (2003), Yussefian (2002) and Moghim and Rouhi (2009).

The kilka fisheries are now threatened by the comb jelly which spread through the entire sea by the year 2000. J. Muir (http://news.bbc.co.uk/hi/english/world/middle_east/newsid_1453000/1453117.stm, downloaded 30 August 2001), Kideys (2002b) and Kideys and Moghim (2003) report a 50% drop in kilka numbers with catches down from 3-6 t per night to half a tonne for one boat. A 50% decrease in kilka catches meant a minimum U.S.$15 million loss to the fishermen (Kideys and Moghim, 2003). Iran's kilka fishery fell from 85,000 t in 1999 to 15,000 t in 2004 and losses exceed $125 million (Stone, 2005a). Ghafar Zadeh and Honar Bakhsh (2008) summarise the economic consequences for Iran. This comb jelly can double in size in one day, reaches maturity in 2 weeks and then produces 8,000 young every day. Maximum abundance reached 5122 individuals per square metre in October 2001 and biomass 1024.5 g/sq metre in august-October 2002 (Roohi et al., 2003; Bagheri, 2004; 2006). The fisheries may recover somewhat after the comb jelly population collapses (Tidwell, 2001b). The website www.caspianenvironment.org/mnemiopsis/index. htm, downloaded 9 April 2003 and Dumont (2002) have extensive information on this problem and Stone (2002b) and IFRO Newsletter (29:4, 2001) confirm a severe depression in kilka and herring stocks. Beroe ovata, a comb jelly that preys on Mnemiopsis, is being cultured in Iran (Kideys, 2002b; Kideys et al., 2004; Rezvani Gilkolaei et al., 2005; Mirzajani, 2006; Mirzajani et al., 2007) and does not appear to feed on other organisms in the Iranian Caspian (Iranian Fisheries Research Organization Newsletter, 38:3, 2004). Reproduction and growth are slower, and mortality higher, than in the Black Sea, due either to the lower salinity in the Caspian Sea water or damage to individuals during transportation for the experiments. If this comb jelly fails to control Mnemiopsis, the introduction of the exotic American species, the butterfish (Peprilus triacanthus), known to feed on ctenophores has been advocated but this fish could also feed on other fishes (Harbison, 2002; Bilio and Niermann, 2004). The complex politics of the nations surrounding the Caspian have prevented the introduction of Beroe (Stone, 2005a).

The Kara Bogaz Gol ("Black Throat Bay"), an eastern arm of the Caspian Sea in Turkmenistan, is 160 km long by 140 km broad (18,389 sq km) but only 2-3 m deep. It acts as a salt precipitator. This water body was blocked off by a dam to conserve the water lost in it by evaporation in 1980. The Caspian Sea has a net annual water deficit of 15 cu km with 5 cu km being lost through the Kara Bogaz Gol alone (Rich, 1982; 1983). However this resulted in salts being spread by the winds, ruining fish spawning grounds and fish farms in the Caspian basin, and ultimately would lead to the salinisation of the Caspian Sea. A dike has now been constructed to allow some flow into the Kara Bogaz Gol and allow the flushing effect to operate. The refilling process over 3 years prevented a 35 cm rise in the Caspian Sea level (Dumont, 1995). Use of this water body to reduce level rises in the Caspian Sea and prevent flooding has been proposed (Wardlaw, 2001). Fish which enter the salty Kara Bogaz Gol lose their swimming capacity, become blind and thrashing about often come to lie on the shore. Birds eat them but those that are missed become salted and dried and may be preserved for a year or so. The Turkmenistan government re-established natural flow into the Kara Bogaz Gol in 1992 because of the Caspian Sea level rise (Zonn in Glantz and Zonn (1997)).

The Caspian coastal plain in Iran runs for almost 650 km from Astara (38°26'N, 48°52'E) in the west to Bandar-e Torkeman (= Bandar-e Shah) (36°56'N, 54°06'E) in the east. This plain has a width of about 25-32 km, but is as narrow as 2 km in places, although it opens out in the east. The Alborz Mountains are almost 1000 km long, on average less than 100 km wide but very high. Damavand reaches 5766 m - an estimate - at 35°56'N, 52° 08'E and is the highest of any mountain to the west of it in Europe and Asia. It has a continuous snow cover. There are persistent snow fields and Alam Kuh at 4849 m has small icefields. The north or Caspian slope is very steep and streams tend to be short and torrential, fed by snow melt and year-round rain. However there are some longer rivers and the principal ones are detailed below. There are about 128 small to large rivers along the Caspian shore. Nümann (1966) gives some limited biological, chemical and physical data on these streams based on spot recordings. Surber (1969) gives values of total alkalinity and calcium-magnesium hardness for a number of streams and reservoirs along the Caspian shore. Most were moderately to relatively hard and therefore productive for aquatic organisms such as insect larvae on which fish feed. The Caspian Environmental Programme (2001b) gives an overview of habitats and biodiversity along this Iranian shore. Environmentally managed areas are listed along with factors affecting their status under the headings of development, drainage, land use alteration, pollution, destruction of vegetation, over-grazing, mining, hunting and fishing, exotics, dams, and roads. Of 123 fish species only 10 or just over 8% are protected with one protected species on the verge of extinction.

Most rivers along the Caspian shore have less than 30% of their discharge in the two wettest months and 40% in the six driest months so discharge is well distributed through the year. In contrast, the Gorgan River at the eastern end of the Caspian basin has 70% of its discharge in the two wettest months, figures comparable with drier areas such as Azarbayjan at 50-60% and the Zayandeh and Kor rivers at 40-60%. Annual discharges can vary markedly, e.g. the Lar River had 545 mm on its basin area in 1949-1950 and 1560 mm in 1950-1951 (Ghahraman, 1958).

The Aras (= Araxes or Araks) is a tributary of the Kura River of Azerbaijan. The Kura rises in Turkey and is 1510 km long. The Aras forms the border between Iran and the former U.S.S.R. (now Azerbaijan and Armenia) for 430 km and has its source near Erzurum (39°55'N, 41°17'E) in Anatolia and the headwaters of the Euphrates River. Its total length is 1072 km. The Aras can be wide and meandering with braided channels and backwaters. Depth range of the Aras is 0.5-4.0 m, average 2.5 m (Zakeri, 1997). The Araxes or Aras Dam was a joint Iranian-Soviet project on this river. Iranian authorities stocked the dam with 1.8 million fingerlings (species not specified) weighing over 10 g each in 1997 to enhance fish farming (Islamic Republic News Agency, 29 December 1997). Akh Gol occupies 600 ha at 820 m in the Aras River valley in northeastern Iran (Scott, 1995). It comprises a small brackish lake with associated marshes and springs and rains to the Aras 5 km away. The area is being converted to agriculture and the lake is being drained. Principal tributaries of the Aras in Iran are the Qareh Su (= black water, draining easily eroded, volcanic soil) draining from the Kuhha-ye Sabalan at 4810 m (38°15'N, 47°49'E) near Ardabil (38°15'N, 48°18'E) and the Qotur River draining past Kuh-e Zaki at 3079 m on the Turkish border through Khvoy (38°33'N, 44°58'E) to the Azerbaijan border near Jolfa (38°57'N, 45°38'E). The Aras and the Safid are two of the three largest rivers in Iran (with the Karun River of Khuzestan). The Kura-Araks basin encompasses 225,000 sq km of which 28,000 sq km or 12.4% is found in Iran (Gleick, 1993). Azerbaijan discharges 303 million cu m of waste into the Caspian Sea annually according to Golub (1992), presumably through the Kura and other major rivers.

Derzhavin (1929a) gave an interesting account of the formation of a new channel of the Aras north of the Iranian border in 1896 which led to the freshening of the Kyzylagach Bay. This favoured migrations of fishes into the Kura River. However, irrigation schemes on the Mugan steppe severely reduced catches as well as causing salinisation of soil. Water abstraction prevented entry of adequate numbers of sturgeons onto the Kura spawning grounds. This type of water usage is paralleled along the Caspian shore in Iran with deleterious effects on a variety of sedentary and migratory fish species.

The Safid (= Sefid or White from its sediment load, up to 60 g/l) River is the only one to completely pierce the Alborz Mountains and has a considerable basin (54,100 sq km) on the plateau. Various sources give differing accounts of its length, up to 800 km. The Safid has the greatest mean discharge of Iranian Caspian rivers, over three times that of the Heraz, the next most important. In flood the Safid discharge is twice that of the Karun, but its minimum is less than a tenth, because the Karun drains a greater area with higher elevations and a more extensive snow pack. The Safid discharge is 4000 cu m per second at maximum, falling to only 15 cu m per second. An average discharge is 182.17 cu m per second. There used to be two freshets before the dam was constructed at Manjil, one fed by spring snow melt in March-May and one by rainfall in the autumn. The rise in water levels and increased sediment load attracted sturgeons, in particular Acipenser persicus. The catch of this species and A. gueldenstaedtii in the Safid River area reached 733,127 kg in 1927/1928 representing 46,500 fish and a caviar yield of 120,958 kg (Vladykov, 1964).

The width of the Safid River varies from 100 to 250 m and depth from 2 to 8 m. The average instant yield is 128.79 m/sec, range 76.5-288.5 m/sec. The average annual yield is 3,998.4 million cu m (Zakeri, 1997).

The Safid is formed from the Qezel Owzan from the west and the Shah River from the east which meet on the plateau and flow through a narrow gorge. This gorge is dammed by what was named the Shahbanou Farah Dam at Manjil (now the Safid or Manjil Dam) (dam height 106 m, length 425 m; reservoir 1860 million cu m, surface area 56 sq km maximum, 14 sq km minimum, maximum depth 80 m, minimum 30 m, summer temperature 24°C, winter 7°C, pH 7.8, 31 g/l turbid materials, Cl- 229 mg/l, SO4 178 mg/l). Strong water level fluctuations prevent the development of a belt of vegetation and the heavy sedimentation inhibit a bottom fauna. Khodjeini and Mohamed (1975) detailed the rate of sediment accumulation in this dam, 757 cu m/sq km/year, evidence of severe erosion of a devegetated drainage basin. The reservoir was half filled with sediment after only 20 years despite an expected life span of 100 years. The reservoir is apparently drained at intervals to remove some of the accumulated sediment. This would severely affect littoral spawning and feeding habitats for fishes. Nümann (1966, 1969) gives details on the limnology of this reservoir. The dam decreased turbidity in the river, raised water temperatures at the river bed in summer and caused marked diurnal temperature changes. This prevented ascent of Salmo caspius to the upper reaches and the dam itself prevented ascent of Rutilus caspicus. Nümann (1966) recommended introducing Sander lucioperca, Acanthobrama terraesanctae (a Levantine species) and cichlids to the reservoir.

Sarpanah et al. (2004) found 45 species and subspecies in the Safid River basin with 29 of these economically important. Thirteen species were migratory, 11 species estuarine and the rest resident. Thirty-six species were recorded as endemics (presumably native) with rest exotics and migrants. The Boojagh National Park near the estuary of the Safid River in Gilan has 25 species and subspecies of fish (Khara et al., 2004).

Lower dams on the Safid, such as the Tarik (10 m high) and the Sangar (3 m high), divert water for irrigation purposes on the Gilan plain, the former through a 16.7 km long tunnel. Construction of the Alamut Dam in the upper reaches of the Safid River basin would affect such species as Luciobarbus mursa, prized for sport fishing, which would need full habitat protection to survive (Aghili et al., 2008). Salmo trutta would not need protection as its habitat is confined to a stretch of river above the dam.

The Safid breaks up into distributaries near its mouth and its flow is carried off into a complex of canals and irrigation ditches. The Safid has changed its delta several times, (Vladykov, 1964). In 1911 it shifted 2-3 km east from the fishing post of 12 Bahman to Hasan Kiadeh. An account in Farsi on the Safid River is given by Wossugh-Zamani (1991b).

The headwaters of the Qezel Owzan lie in Kordestan, near the Iraqi border, and so drain part of the northern Zagros Mountains as well as areas near Lake Orumiyeh such as the Kuh-e Sahand (37°44'N, 46°27'E), mountains near Hamadan (34°48'N, 48°30'E) and the southern slopes of the Alborz Mountains. The Qezel Owzan is about 550 km long. The Taham Dam project 12 km northwest of Zanjan lies in the Qezel Owzan basin on the Taham Chay. This dam is to be 120 m high with a crest length of 450 m and a capacity of 82.7 million cu m. The fish fauna behind the earth dam at "Maljiq", 50 km southwest of Hashtrud in the upper Qezel Owzan basin, suffered severely in the drought of the year 2000. Twenty-five tonnes of fish died after the reservoir dried up (www.irna.com/newshtm/eng/09151847.htm, IRNA, 30 July 2000).

The Shah River is much shorter (ca. 175 km) than the Qezel Owzan and drains the southern Alborz as far east as Takht-e Soleyman at 4819 m (36°22'N, 50°58'E).

The 500 ha Bandar Kiashahr Lagoon (= Bandar-e Farahnaz) Ramsar Site (World Conservation Monitoring Centre, 1990) at 37°25'N, 49°19'E east of the mouth of the Safid Rud was a freshwater coastal lagoon and swamp fed by two streams from the Safid Rud to the west and draining to the Caspian Sea via a channel to the north. The recent rise in Caspian Sea level has converted this area into a bay of the sea as it was in the 1950s before the fall in sea level (Khan et al., 1992). The lagoon bed is sand and mud and the water was oligotrophic except near the marshes to the west. There were reedbeds of Phragmites communis, Typha and Juncus, now restricted to the extreme west end. There were several factors affecting this habitat including a fishery with a fish-processing warehouse, grazing, reed cutting, irrigation abstraction and recreational activities. It was an important spawning and nursery ground for fishes (effects of recent changes unknown) and is still an important centre for commercial fishing.

The Heraz (or Haraz) River drains the Alborz east of Tehran and has a number of longitudinal tributaries in the mountains. These depend on snow melt and are cold even in summer. Fishes are reported to be present in these high streams, but were not easily caught. The Heraz debouches onto a plain and splits up into distributaries. It is polluted from rainbow trout farms (Kazemzadeh Khajuie et al., 2002) and heavy metals (lead and cadmium) are present in fish (Riahi Bakhtiyari, 2001; 2002). Banagar et al. (2008, 2009) record the fish biodiversity as 20 species in 9 families, dominated by cyprinids at 67.2% and with 70% of species resident, the rest anadromous. Exotics are Oncorhynchus mykiss, Carassius auratus, Liza saliens, Gasterosteus aculeatus and Gambusia holbrooki.

The Tajan or Tadjan River was studied by Ro(o)shan Tabari (1995; 1996) who reported on its hydrology and biology. Its mouth lies at 36°49'N, 53°05'E. The maximum flow is in April, decreasing from May onward. In April 1989 flow was 45 cu m/sec falling rapidly to 0.11 cu m/sec in June. Over 70% of the fishes are anadromous with sturgeons being the most important species (Acipenser persicus, A. gueldenstaedtii and Huso huso). Salmo caspius is the most important species in the upper reaches. Other species found in this river are Cyprinus carpio, Alburnus sp. (presumably Alburnus hohenackeri), Capoeta capoeta, Luciobarbus capito, Vimba vimba (= V. persa), Alburnus chalcoides, Rutilus frisii, Rutilus rutilus, Liza sp., Gobiidae, and Esox lucius. Rural, agricultural and industrial pollutants are found in the Tajan and affect the fishes along with dams and other physical obstacles, sand removal and overfishing. The Shahid Rajaee Reservoir Dam, inaugurated in 1997, is found on this river 41 km south of Sari (http://netiran.com/news/IRNA/html/951016IRGG15.html and http://netiran.com/news/IRNA/current.html#HLNO4). The Independent (London) reported on 13 July 1994 that tens of thousands of fish died in this river after poachers poured poison into it about 9 miles (14.4 km) above the estuary. Dead fish covered the river bed for 6 miles (9.6 km).

The south-eastern corner of the Caspian Sea receives two major rivers, the Gorgan and the Atrak or Atrek (ancient Sarnois). Their courses are roughly east-west and parallel each other with the Atrak forming part of the border with Turkmenistan. The Atrak is 495 km long (with 145 km of this in Turkmenistan; Nezami et al. (2000) state 715 km for the Atrak) and the Gorgan 240 km. The Gorgan drains 10,200 sq km and has an average discharge of 9.39 cu m per second (cf. Safid River with 182.17 cu m per second; the Chalus River, directly north of Tehran, has a discharge of 12.65 cu m per second). The Voshmgir or Sangarsavar Dam at 37°12'N, 54°45'E on the Gorgan stores 60 million cu m of water. The water level fluctuates markedly, banks are steep and there is little emergent vegetation. The Golestan Dam (same as preceding?) is 20 km north of Gonbad-e Qabus on the Gorgan River and has a capacity of 86 million cu m. Keivany et al. (1990; http://gause.biology.ualberta.ca/Keivany/bsc/html - 1996) report an irregular pH range for the Gorgan River from 6.3 to 7.9 with an average of 7.1. Temperature range was 8 to 33°C. Conductivity varied greatly from 667 to 10,000 µM/cm, with an average of 875 µM/cm. Chlorides, especially sodium chloride, were the most abundant soluble salts. Total dissolved solids varied from 21 mg/l to 4300 mg/l in an inverse relationship with water volume. Water volume at the dam inlet varied from 2 to 75 m3/second and almost 52% of the sediments entered the dam during a high flood. Water quality was assessed as polluted. The major fish species were Cyprinus carpio, Barbus barbus (sic - possibly Luciobarbus capito), Alburnus spp., Cobitis taenia, Gambusia affinis, and Carassius carassius (sic - presumably C. auratus). A fish kill noted by Coad (1980c) in 1978 was attributed by local informants to careless insecticide spraying on fields neighbouring the Gorgan. Newspaper and radio reports variously stated that 200 barrels of a highly toxic chemical spilled into the river when a truck overturned and that the chemical, identified as Turbidan from the Trintext chemical plant, was dumped by a technician commissioned to get rid of the waste product (Kayhan International, 7 May 1978).

The Atrak headwaters are close to those of the Tedzhen basin. The Atrak basin comprises about 40,000 sq km. The Atrak is only about 10-15 m wide and about 0.5 m deep over much of its lower course. It only reaches the Caspian Sea during floods. A tributary of the Atrak from Turkmenistan is the saline Sambar River, about 203 km long. Petr (1987) reports that efforts were being made to divert this river so as to increase the water quality in the Atrak. The fresh section of the Atrak has a conductivity of 2362 µS and the saline section 23,500 µS. The Caspian Sea off the Atrak River is an important fishery economic zone. Gasan-kuli or Hasan Kuli is a town in Turkmenistan near the Iranian border referred to in fishery reports from this area. The catch of Rutilus caspicus, Cyprinus carpio and Sander marinum was nearly 1.44 x 104 tonnes with only 1.9% being accounted for by Clupeonella cultriventris (= caspia). However by 1972 the catch of the commercially important species had declined to 1.5% and the less desirable Clupeonella had increased to 5.73 x 104 t or 98.3% of the catch. The causes were reduction in the Atrak runoff through irrigation withdrawals, pollution from agriculture, overfishing in the sea and the drop in sea level. Flows of the Atrak did not reach the sea in 1984, 1986, 1990 and 1991and spawning of species using the lower reaches did not occur (Caspian Environmental Programme, 2000).

There are 5 lakes along the Atrak, fed by the river, which have been recently dyked to improve water retention. Their fauna is dominated by native cyprinids. The lowest lake is saline and they range in size from 400 to 2500 ha.

The lakes Alagol or Ala-Gol at 37°21-22'N, 54°35'E, Ulmogol, Alma-Gol or Ulmagol 37°24-25'N, 54°38-39'E and Ajigol or Adji-Gol at 37°24-25'N, 54°40'E comprise a Ramsar Site (World Conservation Monitoring Centre, 1990; Scott, 1995) near the frontier with Turkmenistan just east of the Caspian Sea. Alagol occupies 1400 ha (Scott (1995) states 900 ha) and both Ulmogol and Ajigol 200 ha (Scott (1995) states 280 ha and 360 ha respectively). The Alagol Lake is slightly saline with a mud and sand bottom. It is fed by springs, seepage and precipitation and may dry out completely in summer. It overflows westwards when full. Vegetation is sparse with Juncus, Carex and grasses mainly in the northeast and small patches of Phragmites communis. It is oligotrophic and vegetation poor. The other two lakes have seasonal fresh water fed by precipitation and have a mud and clay bottom. They are eutrophic and water levels vary greatly so that they may dry up completely. Ulmogol has little vegetation such as Juncus, the duckweed Lemna, Phragmites communis, Alhagi and algae while Ajigol has extensive Phragmites reedbeds at its eastern end and abundant submerged vegetation. Fishing occurs in the lakes and the habitats are affected by cattle grazing and reed cutting. Water is abstracted for irrigation and for a fish hatchery. In Alma-Gol and Ala-Gol, 90.91% and 82.18% of the total frequency of fishes was comprised of exotic species. Hemiculter leucisculus was the most frequent in Alma-Gol (58%) and Adji-Gol (16.82%) and Carassius auratus in Ala-Gol (77.6%). Other exotics were Gambusia holbrooki, Pseudorasbora parva and Cyprinus carpio (Patimar and Kiabi, 2005; Patimar, 2007). Patimar (2008) details the environment of these lakes and lists six native species (Alburnus alburnus (= hohenackeri), Barbus (= Luciobarbus) capito, Capoeta capoeta, Cyprinus carpio, Rutilus rutilus and Atherina boyeri (= caspia)) and 4 introduced species (Carassius auratus, Hemiculter leucisculus, Pseudorasbora parva and Gambusia holbrooki), variously distributed among the lakes.

The Qareh Su (= Gharesoo) is another river entering the Gorgan Mordab. In its upper reaches it has a rocky bed and a fauna of Paracobitis malapterura, Capoeta capoeta and Alburnoides cf. bipunctatus, resembling the grayling zone of Europe. The central part of the river dries up (the barbel zone) while the lower river (bream zone) is brackish from gulf input, has high temperatures and pollution. This lower zone has Carassius auratus, Alburnus alburnus (= hohenackeri), Cyprinus carpio, Pseudorasbora parva, Gambusia holbrooki and Gasterosteus aculeatus with Atherina boyeri (= caspia), Neogobius kessleri (= Ponticola gorlap), Neogobius melanostomus, Neogobius pallasi, Knipowitschia caucasica, and Liza saliens feeding in the estuary, and Acipenser stellatus, Alburnus chalcoides, Cyprinus carpio, Rutilius rutilus (= R. caspicus) and Vimba vimba (= V. persa) migrating into the river for reproduction.

Incheh Borun Lake at 37°13'N, 54°30'E is a small and isolated freshwater body of 50 ha about 40 km north of Gorgan. Lake Bibishervan at 37°09'N, 54°52'E and Lake Eymar at 37°08'N, 54°52'E are two more small isolated freshwater lakes occupying 300 ha and 250 ha respectively. All three lakes lie on a cultivated plain. The fish faunas of these lakes are unknown.

The Golestan National Park lies between Bojnurd and Gonbad-e Qabus and is divided by the Tehran-Mashhad highway. The Iran Nature and Wildlife Magazine (volume 3, 1999; downloaded from its English website) states that fish in the Doogh River include rainbow trout and Umbra krameri (sic), both exotics. The latter species is an error of translation from Farsi to English of common names (B. Kiabi, pers. comm., 23 February 2000). A description of the park is given by Kiabi et al. (1994) and of the Madar-Su Stream in the park, which has been studied ichthyologically, by Mikaeili et al. (2005).

The Anzali (= Enzeli or Pahlavi) Mordab (37°26'N, 49°25'E) is a freshwater to brackish lagoon (Firouz, 1968b) separated from the Caspian Sea by a sandy barrier about 1 km wide. The more modern term is "talab" (= pool or marsh, which lacks the association with death) but the older literature refers to mordab and the term is still in common use. It is surrounded by ab-bandans such as the Selke Ab-bandan of 360 ha at 37°24'N, 49°29'E which is protected as a Wildlife Refuge. Ab-bandans are a feature of the Caspian coastal plain, being a shallow and artificial freshwater impoundment managed in winter for duck hunting and in summer as an irrigation reservoir. Safaian and Shokri (2003) describe ab-bandans in Mazandaran based on 423 of these features and Khorasani and Rokni (2001) examined two Mazandaran ab-bandans in particular. The Anzali Mordab complex of 15,000 ha is a Ramsar Site and this includes the whole mordab, the Siah-Kesheem marshes, Selke Ab-bandan and several other ab-bandans. The main mordab comprising open water is 26 km long and 2.0-3.5 km wide encompassing about 11,000 ha. Reed beds extend the eastern limit by a further 7 km. The Siah-Kesheem (or Siah-Keshim) Protected Region has a lagoonal surface area of 4500 ha (Khara, 1994; 6700 ha in Scott, 1995) and is about 12 km long by 4.5 km wide. It lies to the southwest of the main mordab, of which is was probably once part, and is fed by the Esfand River. Note that Khan et al. (1992) state that the Anzali Mordab is unprotected except for the Siah-Kesheem Protected Region and the Selke Ab-bandan of 360 ha. A description of the Siah-Keshim Protected Area is given by Riazi (1996) and of the wetland generally by Monawari (1990). Pollution in the Sia-Keshim Wetland is reviewed by Ganjidoust et al. (2009). Important fishes are listed as Sander lucioperca, Cyprinus carpio, Silurus glanis and Esox lucius (Iran Nature and Wildlife Magazine, 5, www.neda.net/inwm/no.5/english/pre_sites/pre_sites01.html, downloaded 8 March 2000).

The main mordab is drained by the Sowsar Roga, Pir Bazar Roga, "Raste-Khaleh" (? Rasteh Kenar) Roga, Nahang Roga and Pahlavi or "Koulivar" (? Kolver) Roga over a distance of about 4 km to the Caspian Sea. Warm, dense and saline sea water is able to penetrate up these effluent rivers for as much as 10 km, which generally have low flow because of water abstraction and seasonally low precipitation, because of the rise in sea level since 1977. Fresh water flows across the surface of the saline water mixing at depths of 0.5-2.0 m. Salt water contamination is always a danger as more water is abstracted in this heavily populated and farmed area (Kimball, 1973; Kimball and Shayegan, 1973; Sharifi, 2006). Abdolmaleki (1994) gives some data on the benthic macrofauna of this lagoon. Hosseinpour (1995) surveys the zoobenthic resources of the Siahdarvishan and Pasikhan, two principal rivers which enter the lagoon. Other entering rivers are the "Bohambar, Chakoor and Esfand".

Forest clearance around the mordab, rice production and other agriculture, dams and weirs on inflowing rivers, river bed erosion through decline in Caspian Sea level, influx of pesticides such as Diazinon (Talebi, 1998), Paraquat, Glyphosite, and chemical fertilisers, domestic and agricultural sewage, excessive aquatic plant growth and natural decay of vegetation (Nezami and Khodaparast, 1996; Filizadeh and Khodaparast, 2005), phytoplankton blooms, some toxic (Nejatkhah et al., 2003) anionic surfactants (Dadaye Ghandi et al., 2005), siltation from deforestation of feeder streams, introduction of exotic species of fish and plants such as Azolla (Iran Daily, 2 November 2006), grazing for livestock, reed cutting for mats, fences and building materials, and a high urban population growth of 4.6% per year, all affect the habitat and the marsh is highly eutrophic (Mirzajani et al., 2010). These factors also contribute to the fall in commercial fishing success. In the 1930s the catch was dominated by the valuable Rutilus frisii kutum but in the 1990s the catch was 50-75 times lower and the mordab now has a low value to fisheries. The situation is compounded by the absence of effective fishery management. The introduced Carassius auratus dominates catches. The mordab was a principal breeding ground for Rutilus frisii kutum, Abramis brama and Cyprinus carpio, and to a lesser extent Sander lucioperca, and was an important habitat for Esox lucius. Fish kills occur, more than 100,000 dying in August 1997 due to a lack of oxygen after "torrential rain and the growth of aquatic herbs had created an unsuitable environment" (a Reuters report) and more fish died in 2005 (Iran Daily, 21 August 2005). Ghahraman and Atar (2003) concluded that the wetland is dying.

The bottom of the shallow west basin was completely covered by perennial submerged vegetation in the early 1970s (Chara, Nitella, Ceratophyllum, Myriophyllum, Hydrilla, and Vallisneria). Water chestnut (Trapa natans) was the predominant floating plant and covered the central basin in 1966. The Caspian lotus, Nelumbium caspium is found all across the lagoon and is a significant part of the standing stock. Phragmites, Sparganium and Typha are emergent plants which engulfed open water. Reeds were formerly cut extensively for building purposes but are now replaced by sheet metal and cement blocks. Falling Caspian Sea water level and eutrophication from domestic sewage and fertilizers aided plant growth. The fern, Azolla filiculoides, was introduced as an additive to cattle feed and rice cultivation from the Philippines in 1986. It soon entered the mordab from the rice fields and mats up to 20 cm thick covered much of the open water in 1991 (Holčík and Oláh, 1992; Filizadeh, 2002). Dense growths of macrophytes have contributed to declines in commercial fish catches as spawning grounds have decreased, eutrophication is enhanced, and light penetration is decreased and so oxygen declines. There are about 200 sq km of marshes and 30 sq km of shallow open water fed by rivers from the Alborz Mountains. The area of open water in 1989 was only 22.5% of that in the late 1930s (Holčík and Oláh, 1992). However the rise in Caspian Sea level since 1978 has led to a salt water intrusion during the summer months when the Caspian level is at its highest and freshwater input from rivers is at its lowest. Deeper and more saline water may well inhibit plant growth in the future (Khan et al., 1992).

The marsh is only a few metres higher than the Caspian Sea and had a maximum depth of 2.5 m in the early 1970s. Caspian Sea level fluctuations have serious effects on the level of the mordab and hence its utility as a habitat for fishes. The optimum level for the fish industry in general in the Caspian basin is given as -27±1 m (Mandych, 1995). The rise in Caspian Sea level since 1977 is gradually returning the mordab to its supposed, natural brackish state and may improve the fisheries situation which had declined over the last 50 years. Emergent and submergent aquatic macrophytes were decreasing and such fish as Atherina boyeri (= caspia), Alosa caspia, Liza aurata, Syngnathus caspius and Clupeonella cultriventris (= caspia) were increasing in numbers since 1989. However the fishery will require extensive engineering and management innovations to recover.

Hydrorybproject (1965), Kimball (1973), Kimball and Shayegan (1973), Kimball and Kimball (1974), Hagh-Panah (1992), Holčík and Oláh (1992) and Caspian Environmental Programme (2001c) give details of the limnology of the marsh. Water temperatures vary seasonally from 0° to 28.8°C (average about 16ºC) and dissolved oxygen from 0 to 17.5 mg/l for example. Phytoplankton blooms have killed fish in the mordab, e.g. on 5 June 1997 when dissolved oxygen in the western part was at 0-0.2 mg/l and hydrogen sulphide was at 2.0-2.5 mg/l (Iranian Fisheries Research and Training Organization Newsletter, 17:7, 1997).. Conversely, low phytoplankton populations have probably resulted in lowered fish catches. High water temperatures and chlorophyll inactivation through high light levels reduce the numbers of phytoplankton and hence zooplankton, on which fish feed, also decline. Higgins (1973) found that DDT levels in sturgeon, sturgeon caviar, Cyprinus carpio and Rutilus frisii taken near Anzali were not hazardous to humans in flesh (0.2-1.8 p.p.m.) or in caviar (0.05 to 2.5 p.p.m.), both less than the limit for edible fishes set by the U.S. Food and Drug Administration at 5 p.p.m., but that the level in the caviar was a serious threat to sturgeon reproduction. DDT was more concentrated in the eggs because of their fats and oils in which DDT is more soluble. Certain heavy metals, lead and silver, were potentially harmful to the fishes also. Pourang (1995, 1996), Amini Ranjbar (1998), CEP (2001a) and Sartaj et al. (2005) describe heavy metal concentrations (lead, chromium, copper, cadmium, zinc, manganese and nickel) in fish, surficial sediments and various macroinvertebrates of the Anzali wetland. Levels in Carassius auratus and Esox lucius were below recommended levels for human consumption. Carassius auratus, Cyprinus carpio, Esox lucius and Hypophthalmichthys molitrix in the Anzali Mordab have zinc (5.39-27.98, mean 17.28 p.p.m.), cadmium (0-0.08, mean 0.0251 p.p.m.), cobalt (0-1.67, mean 0.6935 p.p.m.), lead (0.11-2.95, mean 1.04 p.p.m.) and mercury (0.113-0.63, mean 0.3 p.p.m.) in their muscle tissues (Annual Report, 1995-1996, Iranian Fisheries Research and Training Organization, Tehran, p. 46-47, 1997). Nadim (1977) found the highest mercury levels in Caspian Sea fish were 0.51 and 0.36 mg/kg in Rutilus frisii and Esox lucius respectively with the lowest in Liza aurata at 0.07 mg/kg. As the acceptable limit was 0.5 mg/kg, mercury contamination in fish was not considered a problem. The lowest zinc concentration was in H. molitrix, the highest lead concentration was in C. carpio and the highest cobalt concentration in C. auratus but concentrations were less than those set by WHO as significant. Södergren et al. (1978) reported on pollution with organochlorines in Esox lucius from the mordab and found this predatory fish to have accumulated the DDT metabolite p,p'-DDE, suggesting that this occurred over considerable time and was not a recent event. DDT did not appear to be incorporated in the pelagic food chain, although it has been used for agriculture and vector control problems. Most DDT probably attaches to clay and soil particles and settles out on the mordab bottom. These authors also recorded DDT from sturgeon species and their eggs in Iranian waters. Pollution continues to be a problem in this heavily populated, industrial and farming region. Heavy rains in October 1995 swept industrial wastes including heavy metals such as lead and zinc, agricultural waste and domestic sewage into the mordab. A fish kill resulted as evidenced by the mordab being covered with floating dead fish. The kill was attributed to the heavy metals and to oxygen depletion (http://netiran.com:80/news/IRNA/html/941029IRGG01.html).

Mercury concentrations in fish and fishermen's hair were studied from the Caspian shore by Zolfaghari et al. (2008). The mean hair mercury concentration was below the WHO threshold level and there was a weak correlation between number of fish meals per month and mercury levels. Levels in Vimba vimba (= V. persa), Rutilus rutilus (possibly R. caspicus), R. frisii, Liza spp., Carassius auratus and Esox lucius exceeded US EPA guidelines.

Amini Rad (2001) assesses the socio-economic importance of fisheries in Bandar Anzali. Fishes are very popular food items there with an average consumption of 11.3 kg, 70% more than in the rest of Iran. White fish (safid mahi, Rutilus frisii) was 1.5 times more expensive than mullets (Mugilidae), 2.6 more than other species and almost 28 times kilka.

Gorgan (= Asterabad or Astrabad) Bay (36°40'N, 53°50'E) is 56 km long by 16 km long and is brackish (8.7-10.0‰) because of input from rivers although Bayrami et al. (20030 give 16 p.p.t. The bay encompasses about 400 sq km. A general description is given by Zanusi (1995) who considers it to be the second richest resource for caviar in the Caspian Sea after the Volga River. The Caspian Environmental Programme (2001c) gives an average surface water temperature of 19.1ºC, oxygen from 2.4 to 11.1 mg/l, pH 8.0-8.5 and total dissolved solids 11.23 mg/l in February to 15,052 mg/l in March. The bay's ecology has been changed by the recent rise in sea level which resulted in storm surges over the sand bar between it and the Caspian Sea. The construction of the Voshmgir Dam on the Gorgan River in 1970 also had an effect, reducing the amount of fresh water to the river mouth which provided spawning areas for Cyprinus carpio and Rutilus rutilus (presumably R. caspicus). Over 40% of the total sturgeon fishing in the Caspian Sea is centred on Bandar-e Torkeman. There is also a black market in sturgeon products. Authorised fishing resources shrunk by 33% from 1993-1994 to 1994-1995 through unauthorised fishing, lack of controls and decrease in controlled sturgeon reproduction. The authorised catch in 1994 for the region from the Neka River to the Turkmenistan border was 1500 tonnes and the unauthorised catch was probably of similar size. The caviar production was 57,000 kg.

The bay once had a valuable Rutilus rutilus (sic = R. caspicus) fishery with an annual catch of 4000 t per year about 20-30 years ago but this has disappeared (Petr, 1987). The bay is now dominated by Mugilidae (CEP, 1998). The catch in the Voshmgir reservoir was 60 t in 1986 although it may improve with stocking programmes. Lalouie (1993) surveyed the hydrobiology of the bay and found an average pH of 8.3, similar to the sea proper as were alkalinity and total hardness. Water temperatures ranged from 5°C to 30°C annually. Pollution from urban and industrial sewage and pesticides is present.

Gorgan Bay is believed to be an important nursery ground for Liza aurata, a major food fish, although an exotic. Cage and pen culture operations in the bay may result in escapes of exotics that could affect native species. On three separate occasions, cages capsized in storms releasing millions of Oncorhynchus mykiss fingerlings (www.ramsar.org/ram_rpt_37e/htm, downloaded 4 May 2001).

The area of the Miankaleh Peninsula, Gorgan Bay and the nearby freshwater Lapoo-Zaghmarz Ab-bandans is designated as a Ramsar Site (World Conservation Monitoring Centre, 1990). The Miankaleh Wildlife Refuge encompasses 81,180 ha and is part of the Miankaleh Protected Region (97,200 ha). Jones (www.ramsar.org/lib_dir_2_3.htm downloaded 14 April 2000) gives 68,800 ha for the Wildlife Refuge. The Miankaleh wetland may encompass 40,000 ha, not the larger figures as originally designated (Khan et al., 1992). The bay has a sand and mud bottom and is oligotrophic. There are extensive marshes along the southern and eastern shores which flood in fall and winter. These marshes are eutrophic from agricultural runoff and stream and irrigation channel inputs. The bay vegetation comprises principally glasswort (Salicornia), sedges (Carex) and rushes (Juncus) with some small reedbeds of Phragmites communis. The ab-bandans have extensive reedbeds of Phragmites communis with stands of reedmace (Typha) and abundant submerged vegetation. Several factors will affect the ichthyofauna including irrigation requirements limiting freshwater flow into the bay and ab-bandans, a fish processing plant at Ashuradeh with associated wastes, a new road along the peninsula which facilitates access and potentially increased pollution and poaching, reed cutting, heavy livestock grazing, agricultural wastes, aquaculture ponds using exotics, fishing by local people and a proposed nuclear power plant. The whole area is an important nursery and breeding ground for fishes. The ab-bandans are not protected although they are within the Ramsar Site. The two shallow ab-bandans occupy 950 ha at 36°50'N, 53°17'E northwest of Behshahr. They are fed by irrigation ditches and drain east into Gorgan Bay.

The Gomishan Marshes at 37°15'N, 53°55'E extends along the eastern shore of the Caspian Sea from Gomishan north and northwest to the Turkmenistan border. There are about 4850 ha of brackish lagoons and marshes, their brackish nature occasioned by the rise in Caspian Sea level. There is agriculture, livestock grazing and waterfowl hunting. The fish fauna is mostly unknown but the area is probably and important breeding ground for the commercially important mullet Liza aurata (www.ramsar.org/ram_rpt_37e.htm, downloaded 4 May 2001), for Rutilus rutilus (presumably includes or is R. caspicus) and for Sander lucioperca, and the latter two are open to hydrocarbon pollution (Ghasempouri and Esmaili Sari, 2002).

The Astara lagoon at the western end of the Caspian coast of Iran is separated from the Caspian Sea by a sand bar, and is flooded across this bar during winter storms. The lagoon encompasses about 950 ha and is fed by a river during August to March, reducing its salinity to about 7 p.p.m. There is a rich growth of aquatic plants and the area has potential for fishing and aquaculture (Petr, 1987). Lavandavil Marsh at 38°20'N, 48°50'E is found about 10 km south of Astara and lies within a Protected Area of 949 ha. It is a small swampy woodland and freshwater marsh with extensive stands of Juncus. Abbasabad Dam at 38°23'N, 48°50'E south of Astara is a 45 ha water storage reservoir. Nur or Neur Gol at 38°00'N, 48°33'E in the northwest Alborz Mountains is a 200 ha freshwater lake at 2300 m about 50 km south of Astara. It lies within the Lisar Protected Area which includes the whole watershed of the Lisar River. The lake drains north to an Aras River tributary but freezes over for about 6 months each year. The submergent vegetation is rich. Rainbow trout (Oncorhynchus mykiss - see account of this species) were introduced to the lake in the early 1970s in an attempt to start a sport fishery. There is also a number of permanent and seasonal lakes along the Sabalan Mountain range which lies partly in this basin and partly in the Lake Orumiyeh basin and these are known to have fishes (www.netiran.com, downloaded 17 June 2004).

The "Lapu" Lake, about 20 km northeast of Sari in Mazandaran, is an example of a smaller water body along the Caspian shore, covering about 100 ha with a maximum depth of about 2.5 m, perhaps 3.5 m in winter (Petr, 1987). There is a rich assortment of aquatic plants. In 1985, 90,000 fingerlings of common carp or kopur (Cyprinus carpio), grass carp (Ctenopharyngodon idella) and silver carp (Hypophthalmichthys molitrix) were stocked and 120,000 fingerlings were added in 1986. A good harvest was reported in 1986. There is a wide variety of reservoirs on the Caspian shore, varying in size from about 10 to 400 ha. Some completely dry out in summer when water demands are high but others are stocked with common carp, silver carp and, to a lesser degree, grass carp. There are also populations of native fishes such as kopur Cyprinus carpio and ordak mahi (Esox lucius) but not in commercial quantities.

The "Amirkelayeh" Lake or Lagoon is located between the cities of Lahijan, Langarud and Kiashahr at 37°17'N, 50°12'E. It is an example of a larger, freshwater lagoon as it encompasses 1230 ha, being 4.5 km long and up to 1.7 km wide. The lake is in the Amirkelayeh Wildlife Refuge and is a Ramsar Site (World Conservation Monitoring Centre, 1990). Average depth is only 1.6 m although some areas reach 4 m (Scott (1995) states 3-4 m on average but up to 6 m). The lake is fed by springs and precipitation and is eutrophic. It lies above the 1980s rise in water level of the Caspian Sea (Khan et al., 1992). It may flood into marshes or the Caspian Sea via a small stream into a channel of the Safid River but is above the recent (1990s) rise in Caspian Sea level. Vegetation is Phragmites communis and Typha with abundant submerged and floating plants such as Nelumbium, Lemna, Potamogeton, Hydrilla, Myriophyllum and Ceratophyllum. The fishes comprise Esox lucius, Sander lucioperca, Carassius sp. (listed as Crucian carp, probably C. auratus), Blicca bjoerkna, Syngnathus caspius, Pungitius platygaster, Silurus glanis, Rutilus rutilus, Cyprinus carpio, and Tinca tinca. Ctenopharyngodon idella has been introduced (Nejatsanatee, 1994).

The Fereidookenar or Fereydun Kenar Marshes at 36°35'N, 52°31'E lie 13 km southwest of Babolsar and occupy 1000 ha. These marshes are artificial, being a damgah or shallow impoundment for duck hunting and water storage. They are one of the best protected wetlands along the Caspian shore as the local duck hunters aggressively restrict access (Khan et al., 1992). There are fringing reed beds of Phragmites australis and Typha with abundant floating and submerged vegetation.

"Seyed Mohalli, Zarin Kola (both at 36°44'N, 53°00'E) and Larim Sara (36°45'N, 53°03'E)" are ab-bandans and associated marshy areas found north of Sari and east of the Tajan River mouth. The first two occupy 600 ha and the last one 1000 ha. Aquatic vegetation is rich, both submerged and floating, and there are extensive stands of Typha and Phragmites. Construction of a large dam on the Tajan will result in an associated network of irrigation canals which may cause ab-bandans to be neglected. The ab-bandans, although artificial, have more of the character of a natural marsh than irrigation channels. Much of this area of the coastal plain has been converted to agriculture which destroys natural wetlands so ab-bandans take on a disproportionate importance as a refuge for wildlife including fishes.

Various dams have been built or are under construction in this basin including the Gourchye Embankment Dam 15 km southeast of Ardebil with a capacity of 20 million cu m, the Yamchi Dam 20 km southwest of Ardebil and the Gaybeglou Dam 40 km south of Meshgin Shahr in East Azarbayjan Province, the Maku Dam with a 150 million cu m capacity in West Azarbayjan and the Agh Chay or Ziaeddin Dam near Khvoy (http://netiran.com/news/IRNA/html/950914IRGG06.html; http://netiran.com/news/IRNA/html/950914IRGG10.html; http://netiran.com/news/IranNews/html/96102201INEC.html). The Neka Power Plant in the eastern Caspian basin entrains a large amount of debris and algae that prevent effective physical systems of fish protection from entrainment. An electrical fish protection system is used instead. Inflatable rubber dams are now being constructed in the lower reaches of rivers, e.g. the Babol, to block the rise in Caspian Sea level such that agricultural water intakes will not be contaminated with saline water. The effects of these dams on fish migrations and biology is unknown (www.satujo.com/english/barrage/dams4.htm, downloaded 20 December 2002).

Qanats and springs are not a feature of this basin as in so many other parts of Iran, except for the drier areas drained by the Qezel Owzan and other streams of the plateau and in the drier valleys of the east away from the rainfall of the Alborz-backed Caspian lowlands. One particular artificial habitat for fishes in the lowlands are the ab-bandans, shallow freshwater marshes maintained as habitat and overwintering areas for waterfowl and for conserving water for rice fields (Beaumont and Neville, 1968). Some ab-bandans around the Anzali Mordab were set aside as refuges for waterfowl and incidentally would protect some fish species threatened by the draining of marshes. Construction of irrigation dams will also lead to abandonment of ab-bandans. Ab-bandans and damgah (ponds made specifically for duck trapping) have declined in number but still encompass 10,000 ha (Scott, 1995).

Extensive stocking of commercially important species in the sturgeon (Acipenseridae) and carp (Cyprindiae) families takes place annually in the Caspian waters of Iran. These are detailed under the Species Accounts. Varedi and Fazli (2005) examined the rivers Shirud, Tonekabon, Larim, Tajan and Goharbara of Mazandaran for the physico-chemical properties of estuarine water in 2000-2001. Only the Shirud and Tonekabon met U.S. Environmental Protection Agency standards for release of fingerlings, the other rivers failing because of water abstraction and improper land use development.

Introduced species based on a summary by Mamaev (2002) include Liza aurata and L. saliens (Mugilidae), Platichthys flesus (Pleuronectidae, apparently not surviving), Psetta maxima maeotica (Scophthalmidae, as Rhombus maeoticus in TACIS (2002) and probably not surviving), Scomber scombrus (Scombridae, not often recorded elsewhere in the literature (an example is TACIS (2002), probably not surviving), Engraulis encrasicholus (Engraulidae, probably not surviving), Anguilla anguilla (Anguillidae), Gambusia affinis (Poeciliidae), Oncorhynchus keta , O. kisutch, O. gorbuscha and Salmo salar (Salmonidae), and Ctenopharyngodon idella, Hypophthalmichthys molitrix, H. nobilis (Cyprinidae). The Indian carps Cirrhinus mrigala, Labeo rohita and Catla catla are being reared in aquaculture stations and are potential escapees into the natural environment (Gilkolaei, 2007). Sal'nikov (2009) reports the capture of an Atractosteus sp. (Lepisosteidae), a North American gar, on the Turkmenistan coast of the Caspian Sea.

A wide variety of parasites have been recorded from fishes in this basin and these are mostly dealt with in the Species Accounts. Pazooki et al. (2008), for example, recorded 7 monogenean species from 11 fish species in the Aras, Zangbar and Ghotor rivers of northwest Iran, namely Dactylogyrus extensus, D. chramuli, D. lenkorani, D. kendalanicus, Silurodiscoides siluri, Diplozoon megan and Gyrodactylus varicorhini.

Zoogeographically, Berg (1940) considers this part of Iran to belong to the Kura-Iranian sector of the Caspian District of the Ponto-Caspian-Aral Province. This fauna is very similar to that of the Kura River although certain genera are absent, even in the Safid - a major river, such as Chondrostoma, Gobio and Leucalburnus.

Dasht-e Kavir

This basin occupies an immense area of north-central Iran, over 200,000 sq km in the rain shadow of the Alborz Mountains. Mahdavi and Anderson (1983) detailed the qanat water supply of the margins of this basin. Intermittent streams drain to several kavirs which are grouped together under this basin for convenience. The principal kavirs are the Damghan Kavir in the north, the Sabzevar Kavir in the north-east and the Kavir-e Bozorg (or Great Kavir) occupying much of the basin, being about 450 km in east-west extent and 250 km in north-south extent. The Kavir-e Bozorg receives waters exiting from other kavirs. The principal streams entering this basin drain the Alborz Mountains and their eastern extensions in Khorasan. The Alborz peaks exceed 4000 m and even to the east the Kuh-e Binalud (36°30'N, 58°55'E) attains 3416 m near Neyshabur (36°12'N, 58°50'E) while the lowest points are at an altitude of 650 m. The Damghan Kavir receives two major streams from the Alborz, the Damghan River and the Hasanabad River, and other streams dry up in early summer. The Sabzevar Kavir has numerous small and temporary streams which feed it as well as two major streams, the Mureh River, 320 km long, and its tributary, the Kalshur River, 240 km long. The Kalshur drains the Kuh-e Binalud and flows west to meet the south flowing Mureh. These rivers drain areas rich in salt domes and samples taken show water to be saline and some streams are fishless. Qanats support fishes in this area although the fish only emerge at night in some cases. Ruttner-Kolisko (1964; 1966) and Ruttner and Ruttner-Kolisko (1972; 1973) studied the chemistry and limnology of natural springs and qanats in a mountain area separating this basin from the Bejestan basin. Several factors were found to affect the limnology. Climatic factors were temperature, precipitation and evaporation, edaphic factors were geology, salt content of soil and intensity of waterflow, and pollution by man and animals was a factor. There was a range in salinity from low (<15 mval/l) to high (>120 mval/l). Qanat discharges in this area were 20-50 l/sec. Springs were small and many were dammed to form small pools for livestock.

These large central basins of Iran were once thought to be desiccating lake basins. However more recent studies have shown that although there may have been shallow lakes, e.g. saline Lake Damghan, and rivers carried more flow and were perhaps more closely linked than today, there was no extensive and continuous freshwater lake over the whole of central Iran that could have facilitated fish dispersal. While the hills received increased rainfall, the central deserts remained arid during Pleistocene "pluvials" and cold phases (Bobek, 1959; Scharlau, 1968; Krinsley, 1970).

Dasht-e Kavir (NASA and Wikimedia Commons).

Dasht-e Kavir (NASA and Wikimedia Commons).

Dasht-e Lut

The Dasht-e Lut basin of south-central Iran is ringed by mountains yet has the lowest point on the plateau at 205 m in the Namakzar-e Shahdad. The central portions of this basin are some of the most barren and inhospitable in Iran or indeed the world. Conrad and Conrad (1970) and Gabriel (1938) give descriptions of this desert basin. Intermittent streams drain the mountain ranges around Kerman east to the namakzar or namaksar (= salt waste), north from mountains near Bam (29°06'N, 58°21'E) such as the Kuh-e Jebal Barez (28°30'N, 58°20'E) and Kuh-e Bazman (28°04'N, 60°01'E) which delimit the northern edge of the Hamun-e Jaz Murian basin, west from the slopes of the active volcano Kuh-e Taftan (28°36'N, 61°06'E) and south from the mountain ranges near Birjand (32°53'N, 58°13'E). High points include the Kuh-e Hazaran west of Bam and south of Kerman at 4402 m. Such heights retain snow and have more abundant precipitation which feed streams at least in the mountains. However many minor and some apparently major streams marked on maps are completely dry. Much of the water is absorbed into the ground and tapped by qanats. The Shah River at Birjand is dry through most of the year (Fisher, 1968). Tabas (33°36'N, 56°54'E) at the northern end of this basin has numerous qanats (Krinsley, 1970) but I have not seen samples from this area.

The Shahdad River is presumably in this basin based on maps and supplies water to Kerman and some nearby villages. One sample station was polluted by wastes from a rainbow trout farm (Rezaei Tavabi et al., 2009). The Tahrud is an important stream which drains the Hazaran to a small sump in the south of the Dasht-e Lut basin and has a continuous flow which becomes subsurface well east of Bam (compare maps). Its maximum map extent approaches 250 km. In the mountains, the Tahrud is 1-8 m wide and up to 50 cm deep. Water temperature was a warm, 15°C on a cool December day.

The Dasht-e Lut includes the largest sand dune field in Iran (ca. 10,000 sq km) which has developed through aeolian erosion. Sand dunes block roads and may well fill in or divert streams.

Qanats in this basin can have water temperatures much higher than the few surface streams. One qanat near Bam had a temperature of 25°C in a snowstorm, yet stream temperatures below 10°C are not uncommon.

Esfahan

The principal feature of this basin is the Zayandeh River which rises in the Zagros Mountains east of Zard Kuh at 4548 m (32°22'N, 50°04'E) and flows east for about 300 km to its terminal basin, the Batlaq-e Gavkhuni at 32°20'N, 52°47'E, a salt marsh with a salinity of 315‰ (Löffler, 1961) and an average depth of about 1 m (www.netiran.com/php/artp.php?id=1615, downloaded 19 July 2004). The salt marsh can dry up in summer. Wetlands associated with the terminal basin are a Ramsar Site of 43,000 ha (or 37,000 ha; sources vary as does the size of the marsh seasonally and annually). Associated marshes at the river delta and along its banks are fresh to brackish. These marshes are fed by flooding and by irrigation canals but dry up in late spring or early summer. Flooded areas often freeze over in winter. There is little natural marsh vegetation and flooding occurs over degraded steppe and cultivated land. Water is diverted for irrigation and for domestic and industrial uses. It  receives pollution from Esfahan and other urban sources. Esfahan is a major oasis city on the Zayandeh at 32°40'N, 51°38'E with a population over 1 million, famous for its bridges (pol in Farsi) among other sites.

Zayandeh River at Si-o-Se Pol (Photo by Farokh Behmardi from Wikimedia Commons).

Zayandeh River at Si-o-Se Pol (Photo by Farokh Behmardi from Wikimedia Commons).

Zayandeh River at Pol-e Khaju in winter (Wikimedia Commons).

Zayandeh River at Pol-e Khaju in winter (Wikimedia Commons).

 

The Zayandeh basin encompasses about 30,480 sq km and is connected to the upper Karun River basin (which drains to the Persian Gulf) by the Kuhrang Tunnel constructed in 1953 although first proposed in the early sixteenth century (Fitt, 1953; Afifi, 1966; IRNA, 5 February 2002). Two additional tunnels are under construction (Stoltz, 2002). A hydroelectric dam at Godar-e Langar (also known as Karun-4) would also supply piped water to Esfahan 300 km away if it is completed (Whitley and Gallagher, 1995). Dams have deleterious effects on a riverine fish fauna and are often stocked with exotic species. The upper Karun has not been well explored for endemic taxa. Mean annual flow of the Zayandeh is estimated at 1.2-1.45 billion cu m, used mostly for agriculture but an increase in population and industry has necessitated dam construction (Shah Abbas Kabir or Sadd-e Zayandeh Rud, capacity 1450 million cu m) and diversion schemes. The dam is an oligo- mesotrophic water body based on phytoplankton studies (Shams and Afsharzadeh, 2009). There is also the Hana Dam on the Hana River at Semirom with a height of 35 m and a capacity of 45 million cu m (http://netiran.com/news/IRNA/html931003IRGG04.html) and the Izadkhast dam to the southwest of the Batlaq-e Gavkhuni (www.irna.com/newshtm/eng/12003142.htm, IRNA, 2 July 2000). As well as man-made diversions, the upper Zayandeh basin has captured headwaters from systems tributary to the Persian Gulf. The Shah Abbas dam has reduced the natural flood flows downstream and little water now enters the salt desert.

Plans have been made to transfer Zayandeh River water from the Band-e Cham-e Asseman to Yazd's Shahneh Reservoir by pipeline over a distance of 375 km (Hamshahri, Tehran, 629:5, 22 February 1995). 78 million cu m of water will be transferred annually and this will decrease the habitat for fishes in the Zayandeh River basin.

Spring flow is at least 1700 cu m per second, but this drops to 28 cu m per second in autumn (Oberlander, 1968b). Discharge peaks in April with low values in September-October and decreases dramatically downstream after abstraction, evaporation and infiltration (Beaumont, 1981). The Zayandeh can be forded on foot at Esfahan in summer and Buckingham (1829) reported it to be dry. It dried again in 2000, 2001 and 2003 under drought conditions, partly through water abstraction upstream for irrigation and partly through aqueducts to other desert cities (Rafsanjan and Yazd) not in the Esfahan basin (Anonymous, 2001b; Foltz, 2002; newspaper reports). The river is polluted by city sewage, local wastes dumped directly into the river, and industrial wastes (Moghadam, 1976; Al-Hashimi, 1987; Tehran Times, 15 September 1997). 172,000 cu m of industrial pollutants enter the river daily. Pollutants include phosphorus, nitrogen, lead, nickel, zinc, organic substances, iron, manganese, oil products, mineral and organic dyes and the sewage from villages with a population of 900,000 people. Nadim (1977) found the highest mercury levels in fish were 0.19 mg/kg. As the acceptable limit was 0.5 mg/kg, mercury contamination in fish was not considered a problem. The flow is 1.45 billion cu m annually of which 1.1 billion cu m is used for agriculture, 150 million cu m for industry and the remainder is used as drinking water. The basin has a high demand for water supplies and has been under stress in this regard for the last 50 years. It will be unable to meet water demands in less than 15 years (Salemi and Heydari, 2006).

Ouseley (1819-1823) noted numerous small "bleak" and caught several carp-like fish up to 12-14 inches long (ca. 30-36 cm) in the deeper waters around the bridges over the Zayandeh at Esfahan.

The Batlaq-e Gavkhuni and marshes on the lower Zayandeh are a Ramsar Site, the lake occupying 12,000 ha, permanent marsh 1000 ha and temporary marsh 30,000 ha (World Conservation Monitoring Centre, 1990) or 47,000 ha (Mehrabi, 2004). It lies at 1470 m and has an average depth of 1 m. The Batlaq (= salt lake or marsh, gavkhuni = cowshed because cattle are put out to pasture in the marshes) is fishless but the marshes have a freshwater character depending on the input from the Zayandeh River. The substrate is silt and mud. Much of the marsh has been converted for agriculture. Flooded areas may freeze over in winter. The salt lake is said not to dry out completely (Mehrabi, 2004) although flows were down to 10-100l/s in the dry years 2000-2002 and the lake was dried out (Esteky, 2006).

As with all plateau basins, this one also has springs and qanats which contain fishes. Surber (1969) gives some data on total alkalinity and calcium-magnesium hardness in this basin and characterises it as moderately hard.

Fish farms have been developed in Esfahan Province (Tehran Times, 31 October 1999). Thirteen cold water and 10 warm water fish farms are expected to yield 490 t of fish, rising to 18 cold water and 15 warm water farms by the year 2000 with a yield of 760 t.

Hamun-e Jaz Murian

The Hamun (= marshy lake, in this instance) is dry for most of the year, but fills with fresh water in winter (Harrison, 1941). Its extent is presumably variable, depending on rainfall. It lies at an altitude of about 300 m, with a still-subsiding depression within the Jaz Murian plain, and is ringed by mountains.

The two major rivers flowing into the Hamun are the Halil (or Haliri) River, known as the Kharan or Zar Dasht River in its upper reaches, which flows from the neighbourhood of Kuh-e Laleh Zar at 4374 m lying to the northwest, and the Bampur River which flows towards the Hamun from the east but follows a southerly course in its upper reaches (Tipper, 1921). The source of the Bampur River lies between 1000 and 1500 m. The Halil is a longer river (ca. 390 km) than the Bampur (ca. 315 km) with a stronger and more continuous flow. However, this river was nearly dry downstream of the Jiroft Dam and there was only minimum flow upstream in 2008 during a drought (Atabak Mahjoor Azad, pers. comm., 6 October 2008). There is a 130 m high dam on the Halil, the Jiroft Dam, 40 km upriver of Jiroft. A flood water storage dam at Bazman is 37 m high with a capacity of 3.3 million cu m (www.irna.com, downloaded 26 January 2003). Discharge is only 1-3 m3/second in summer. Floods occur (including an historical one which destroyed Jiroft in 1000 A.D., and one in 1993) and river discharge can reach 800 m3/second in 15 hours with an 18 m rise in reservoir level in 40 hours and massive sediment transport with turbidity reaching 280 gr/liter (sic) (www.stucky.ch/publication/JIRFLOOD.htm downloaded 19 July 1999). The Bampur River in late November and early December was flowing in its upper reaches near Karevandar and around Iranshahr and Bampur but was dry between these two areas. Judging from its width and depth below Bampur it probably did not reach the Hamun by surface flow. Most rain at Iranshahr falls in January and February (15 and 52 mm respectively) with none in the remaining months except for rare summer monsoonal rains (Ganji, 1960). Irrigation and canal schemes in the Bampur basin suffer from erosion and siltation problems as elsewhere in Iran (Borowicka, 1958).

The Hamun-e Jaz Murian basin is ringed by much smaller streams draining the surrounding mountains. These are all very small, e.g. the Ughin River was as narrow as 30 cm and maximum depth in pools was about 50 cm when sampled on 4 December 1977.

Hamun-e Mashkid

The Hamun-e Mashkid (= Mashkel) lies within Pakistan with its western edge on the border with Iran. In this instance hamun means a salt waste. The mountain ranges in this area of Iran are parallel with the Iran-Pakistan border and run in a northwest-southeast direction.

The Mashkid River rises to the east of the mountains ringing the Hamun-e Jaz Murian basin and flows east into Pakistan where it receives a right bank tributary, the Rakhshan River, before turning north to flow into the Hamun-e Mashkid. Its total length is ca. 430 km. Two tributaries of the Mashkid within Iran are the Rutak River and the Simish (= Sunish River) which drain the lowlands between Kuh-e Birag (27°35'N, 61°20'E) and the Badamo Range (27°38'N, 62°08'E) from the northwest to enter the Mashkid River southeast of Saravan (27°22'N, 62°20'E). The upper Mashkid River is a small mountain stream, probably with a perennial flow. The lower reaches of this river, and of the Simish, comprise a series of muddy pools of varying size. Some of these pools were isolated and fishless in early December 1977, while larger ones, perhaps 1 km long, contained some emaciated specimens. In this area fish are found more abundantly in perennially flowing qanat streams.

The Tahlab River and its tributaries drain the eastern slopes of the mountains south of Zahedan. The Tahlab flows in a southeasterly direction into the Hamun over a ca. 160 km course. It was dry between Zahedan and Mirjaveh (29°01'N, 61°28'E) in early December 1977. The Ladiz River is a short (ca. 80 km) right bank tributary of the Tahlab flowing from Kuh-e Taftan. In its lower reach it was a small stream flowing in the bottom of a deep and wide canyon. The stream banks were white with salt deposits.

Kor River

This basin occupies 26,440 sq km north and east of Shiraz at a lowest altitude of ca. 1525 m. Its lowest part is occupied by a chloride lake, the third largest lake in Iran, composed of two parts, a northern basin known as Narges or Tashk and a southern basin known as Neyriz = (Niriz) or Bakhtegan. The two basins are not always connected and the southern basin is saltier because major freshwater input is from the north. Löffler (1956; 1957; 1959; 1968; 1981) gives details of this lake. The lake area varies between 1210 and 2400 sq km, with a maximum depth of 1.1-1.7 m and a mean depth of 0.5 m. Salinity is 13.7-101.6 gl-1 and temperatures range from 15°C to 45°C in the shallows. The lake is reported to have dried out completely in 1871, 1933 and 1966 (Cornwallis, 1968a) and in 2000 (www.irna.com/newshtm/eng/05142727.htm, IRNA, 26 July 2000). Löffler (1993) considers that this lake may dry out permanently in the near future if abstraction of water from the Kor River for irrigation continues to grow. The drought in 2003 reduced Lake Bakhtegan to a series of puddles. Fluctuations in lake levels affect the freshwater faunas of springs, including fishes, which drain into the lake: high levels swamp the springs with water too saline for fishes to survive. Low levels, however, allow streams to connect and exchange faunas on the lake bed so they are not as isolated as they might appear.

Lakes Tashk and Bakhtegan (centre) with Lake Maharlu on left (NASA and Wikimedia Commons).

Lakes Tashk and Bakhtegan (centre) with Lake Maharlu on left
(NASA and Wikimedia Commons).

 

Bobek (1963) suggests that there may have been an outflow from this basin to the Gulf at the south-east corner of the lake which was cut off at the end of the Pleistocene by alluvial fans. However Krinsley (1970) maintains that any outlet was closed by the late Pliocene.

Major rivers are the Kor (= the classical Araxes) and its tributary the Pulvar (or Sivan) (= the classical Medus) which rise in the Zagros Mountains to the north and north-west and drain to the north-west corner of Lake Tashk. These mountains are high enough (Kuh-e Dinar at 4432 m and 30°50'N, 51°35'E) to have a snow cover and thus there is a continuous flow throughout the year. However in summer water does not reach the lake because of the demands of irrigation. Drainage and irrigation canals run through the basin on the plains at the north end of the lake. Several springs feed marshes, notably the Lapu'i marshes, a wetland of 150 sq km to the north-west of the Kor-Pulvar junction, the Zarqan marshes of 4 sq km, an extension of the Lapu'i marsh (both now severely damaged by construction of a drainage canal as part of the Dorudzan or Sadd-e Daryush-e Kabir (dam) at 30°15'N, 52°20'E, a project on the Kor River), the "Gomun", "Gumoon", "Gumoo" or "Sangare" marshes of 2 sq km at the north-west corner of Lake Tashk and the Sahlabad marshes of 5 sq km on the south-east coast of Lake Bakhtegan (Cornwallis, 1968a; 1968b). The Band-e Amir or Kamjan Marshes at 29°40'N, 53°05'E are formed at the delta of the Kor River and encompassed about 100 sq km but the Daryush-e Kabir Dam severely restricts the water flow to these marshes. A dam on the Bolaghi Gorge is proposed which would affect the flow of the Pulvar but is being opposed on archaeological grounds (www.netiran.com, downloaded 4 October 2004).

The fish, Aphanius sophiae, is found in these marshes and springs, but suffers predatory attacks in an unusual way. The greater flamingo stirs up mud in its feeding and this releases H2S, CO2, and CH4, suffocating the fish and making them easy prey for herons.

The Neyriz Lakes and Kamjan Marshes are a Ramsar Site (World Conservation Monitoring Centre, 1990; Khan et al., 1992) although the Kamjan Marsh area may be deleted because of drought and other factors such as rice, wheat and cotton growing and livestock grazing. The "Cheghakhur" and "Gandoman" marshes in Chahar Mahall and Bakhtiari Province will be substituted for the Kamjan Marshes as a listed Ramsar Site (Khan et al., 1992). The "Gumoon" marshes have been partially drained for irrigation and for conversion into aquaculture ponds (Khan et al., 1992).

The Ghadamghah spring-stream system at 30°15'N, 52°25'E and 1660 m altitude has been described by Esmaeili et al. (2007) and is a regional hotspot for biodiversity. The fishes present are Petroleucsicus persidis (Cyrpinidae), Cobitis linea (Cobitidae), Seminemacheilus tongiorgii, Oxynoemacheilus farsicus (Nemacheilidae), Aphanius sophiae (Cyprinodontidae) - all Iranian endemics, and Alburnus mossulensis, Capoeta aculeata and Capoeta damascina.

The Daryush-e Kabir Dam on the Kor River contains 990 million cu m of water, is 24 km long and about 9.5 km wide. Its conductivity is 363 µS compared to Lake Bakhtegan at 105,900 µS and consequently it can support a fish fauna. Band-e Amir on the Kor River is a diversion dam over 1000 years old and also provides a small reservoir habitat for fishes (Houtum-Schindler, 1891). At least three other dam sites have been proposed in this basin (Tang "Boraghi" (= Tang-e Boraq), "Tang Bulak" and "Ghaderabad" (= Qaderabad)). Surber (1969) gives some spot data on pH, total alkalinity, calcium-magnesium hardness, chlorides and free CO2 in this area. Water is relatively hard. Concentrations of total dissolved solids vary between 202 mg/l and 436 mg/l in the rivers compared to a range of 333-6937 mg/l in the Gulf basin.

Kaftar Lake at 30°34'N, 52°47'E is at ca. 2300 m in the Zagros Mountains northeast of Shiraz. It occupies 4700 ha (500 ha in Khan et al. (1992)) and is a shallow, semi-permanent freshwater lake which can dry out completely in summer and is frozen over in winter. The annual mean water temperature is 14.4°C, the mean maximum 23.5°C and the mean minimum about 2°C (B. Jalali, pers. comm., 1999); and Nowrouzi and Valavi (2011) give various physicochemical parameters. Lake water has been proposed for irrigation usage in the past and a recently proposed earthen dam would reduce the lake area by half (Scott, 1995). It has a mixed ichthyofauna of native species and exotics. The fishes recolonise from springs and the main river entering the lake and are also stocked.

The Kor River basin also contains qanats. Some of these flank the Pulvar River, for example, and serve to bring water to fields above the incised river bed.

Pollution in this basin has been recorded by Merchant and Ronaghy (1976) where industry discharges waste untreated into surface and ground waters. Waste from a sugar mill killed 1 million fish in 1994 and a further 500,000 fish died in 1996 from industrial waste (http://www.iran-e-azad.org/english/noi/noi-83.html or News on Iran, 83, 15 November 1996). A fish kill was reported from the Pulvar River in 1978, polluted by wastes from a food factory (Coad, 1980c). Peritore (1999) and Moussavi and Saber (1999) record the Kor River receiving organic wastes from animal processing plants, ammonium and mercury from petrochemical complexes and such heavy metals as cadmium, chromium and arsenic from electronics manufacturers. Ebrahimi et al. (2008) and Taherianfard et al. (2008)  report lead and mercury levels in Cyprinus carpio and Capoeta spp. to be less than the maximum allowable by the European Union but still of concern. Ebrahimi and Taherianfard (2010a, b), however, found that levels of arsenic, cadmium, lead and mercury for these species were higher than permissible for human consumption.

Channels started in 1981 to provide more agricultural land drain through the Kamjan Marshes to Lake Tashk and the Kharameh Marshes to Lake Bakhtegan. Much of the marsh habitat has been destroyed. The "Gumoon" Marsh has been drained for agriculture and fish ponds.

Miller (1985) reports on deforestation in this part of Iran during the fourth to second millennium B.C. Even marsh areas were probably treed before demands for charcoal and construction materials increased. The fish faunas must have adapted to increased insolation and any species sensitive to higher marsh and stream temperatures would become less common.

Lake Maharlu

The Maharlu basin is the valley of Shiraz (29°36'N, 52°32'E) and encompasses about 4100 sq km. Lake Maharlu is at an altitude of about 1460 m, has an estimated average area of 220 sq km, a maximum depth variously cited as 0.5 and 3 m, a salinity of 124‰ or 304.95 gl-1 and is fishless. The lake dried out completely in 1967 (Cornwallis, 1968a). The lake is fed by minor streams and springs around its margin. The Khoshk River flowing through Shiraz is dry for much of the year or composed mostly of polluted wastes from businesses, domestic sources, industry and agriculture (Kafilzadeh et al., 2007). The basin also has a number of qanats. Stream temperatures vary between 8°C in January to 32°C in June while qanats can be warm, e.g. at Sarvestan (29°16'N, 53°13'E) in December a qanat was 25°C. Surber (1969) gives some spot data on pH, total alkalinity, calcium-magnesium hardness, chlorides and free CO2 in this area.

The basin is separated by only a small rise from the Mand River of the Gulf basin, but is treated separately here because fish collections have been focused on this valley as Shiraz is the major city of southern Iran.

Major fresh to brackish springs and their associated marshes (Ab-e Paravan (2.5 sq km), Barm-e Shur (1.5 sq km) and Soltanabad (7 sq km)) are concentrated at the northern end of the lake (Cornwallis, 1968a). Larger springs have pools which are about 2 m deep and reed beds of Phragmites and Typha, some of which are cut. Livestock grazing occurs. Amphibious tanks were tested in Barm-e Shur, stirring up anoxic bottom mud and leading to a fish kill.

Numerous small springs around the lake are isolated from one another by the intervening hypersaline water. Lake levels fluctuate markedly and allow streams to meet on the exposed salt flat when the water level is low. At high levels, salty water invades the lower springs and eliminates their fishes, which only recolonise when the lake level falls again and connection is made with a stream from a spring which was above the last rise in lake level. One spring had a salinity of 34‰ at the source when the lake had risen to "invade" the spring. Aphanius persicus were concentrated close to the source but would attempt to evade capture by swimming into the salt lake where salinity was 180‰. Their excursions into water of this salinity was brief and fish paled visibly while darting in and out. Another spring was replete with tooth-carps at 144‰. Temperature on 8 June 1976 at one spring was 27°C at the surface and 32°C on the bottom, at about 1 m depth.

Lake Orumiyeh

Lake Orumiyeh (= Reza'iyeh, Urmia, Urmi, Urumiyeh or Darya-e Shahi) lies in north-west Iran and is the only Iranian lake large enough to appear on general maps of the world. This lake is a Ramsar Site and includes Orumiyeh National Park. Brackish marshes in the northeast, northwest and southern shores probably support some fishes but the lake itself is too salty.

Lake Orumiyeh (NASA and Wikimedia Commons).

Orumiyeh lies at about 1275-1300 m (accounts vary), is about 128-149 km long and 40-60 km wide. This thalassohaline lake has a surface area of 4750-6100 sq km, a volume of 29.4 cu km, a mean depth of 4.9-6.0 m, a maximum depth of 16 m, and a temperature range of -1.3-27.5°C. Lake level can rise as much as 2 m in one season, as it did in the winter of 1968-1969. It is a sodium chloride-sulphate system with a salinity up to 340.0 gl-1 (but mostly 217-235gl-1) and consequently is fishless (Abich, 1856; von Seidlitz, 1858; Rodler, 1887; De Mecquenem, 1908; Plattner, 1955; Vladykov, 1964; Kelts and Shahrabi, 1986; Ghaheri et al., 1999; www.neda.net/inwm/no.6/english/geology/geology01.html, downloaded 10 July 2000; Van Stappen et al., 2001; Eimanifar and Mohebbi, 2007; Karbassi et al., 2010). Initially the lake was probably fresh (Admiralty Naval Staff, 1918). A causeway has divided the lake into two parts since 1989; a gap allows a limited exchange between the two parts. Its drainage basin approaches 57,000 sq km (or 51,786 sq km, authors differ) and the lake is the terminal basin for a number of streams and rivers. Annual inflow is 6900 x 106 m3 (Ghaheri et al., 1999). During spring runoff a freshwater plume covers large areas over the saline lake near river mouths. Prominent perennial streams include the Zarrineh River (230 km long) entering from the south and draining part of the northern Zagros with a range in discharge of 10-510 cu m per second with the Tata'u or Simineh River (145 km) as a major tributary, the saline Aji Chay or Talkheh (= bitter) River from the east draining the flanks of Kuhha-ye Sabalan at 4810 m (38°15'N, 47°49'E) and Kuh-e Sahand at 3710 m (37°44'N, 46°27'E), and the smaller streams from the west such as the Zowla (= "Zola") Chay (84 km), Nazlu Chay (85 km), Shahr (= "Shaher") Chay (70 km), Baranduz Chay (70 km) and Gadar (= "Qader") Chay (100 km) (Günther, 1899). Both the Zarrineh and the Talkheh exceed 200 km in length. The Talkheh River has a hardness of 820 mg/l according to Surber (1969), who also gives values of total alkalinity and calcium-magnesium hardness for a number of streams and lakes around Tabriz. The Talkheh floods extensively in the spring and forms large marshes. Most streams were relatively hard like the Talkheh although some were soft such as the Basmenj Chay draining Kuh-e Sahand at 70 mg/l.

Lake Kobi (= Ghopi) is a Ramsar Site lying at 36°57'N, 45°52'E and 1240 m altitude in this basin. It is south of Lake Orumiyeh and northeast of Mahabad. It comprises the fresh to brackish lake and associated but discontinuous marshes of about 1200 ha. The endorheic lake is shallow with a maximum depth of 1.5 m and a mud bottom. It is fed by precipitation and springs, and when full floods marshes to the north. It freezes over in winter. The lake is eutrophic and has reedbeds of Phragmites communis and abundant submerged vegetation. Livestock grazing and wildfowl hunting occur.

The Shur Gol and the "Yadegarlu" (= Yadergarlu) and "Dorgeh Sangi" endorheic lakes are at 37°00', 45°26-35'E south of Lake Orumiyeh and northwest of Mahabad at 1290 m are also a Ramsar Site comprising 2500 ha of lakes and associated marshes. They are fed by precipitation, springs and small streams. Shur Gol at 2000 ha is surrounded by the Hassanlu Marshes. Its water is brackish to saline. The eutrophic marshes flood in fall and winter and have abundant submerged vegetation. "Yadegarlu" is a shallow freshwater lake of 350 ha with abundant submerged vegetation and a surrounding of eutrophic sedge marshes. It may dry out in summer. It apparently suffered in the Iran-Iraq war (Jones, www.ramsar.org/lib_dir_2_3.htm, downloaded 4 April 2000) and may be deleted as a Ramsar Site. "Dorgeh Sangi" is 150 ha in extent and is a shallow freshwater and eutrophic lake. All three lakes may freeze over in winter. Reed cutting, grazing and waterfowl hunting occurs in this complex and some drainage of wetlands for agriculture may occur (Khan et al., 1992).

"Gerde Gheet" (Gordeh Git) and "Mamiyand" (= Meimand?) at 37°02'N, 45°40'E are freshwater marshes south of Lake Orumiyeh and north of Mahabad occupy 500 ha at 1300 m. The marshes are covered by Phragmites. Waterfowl hunting occurs here and some livestock grazing.

The "Ghara Gheshlaq" freshwater marshes at 37°10'N, 45°50'E occupy 400 ha at 1290 m south of Lake Orumiyeh and north of Mahabad. The water is about 1 m deep, eutrophic and freezes over in winter. Large parts of these marshes were drained by the "Mahabad Multipurpose Drainage and Irrigation Project" in the 1970s despite environmental concerns. Cornwallis (1976) notes both the draining of these marshes and the cessation of freshwater discharge from the Mahabad River. He also points out the likelihood of chemical contamination from agriculture, choking by vegetation and the probable use of herbicides. He recommends introduction of Ctenopharyngodon idella and Hypophthalmichthys molitrix. The marshes have been proposed as a Ramsar Site.

Lagoons in the Mahabad area dried in the year 2000 (www.irna.com/newshtm/eng/05142727.htm, IRNA, 26 July 2000).

Gori Gol or Lake Gory at 37°5'N, 46°42'E is a fresh to brackish lake near Tabriz occupying 120 ha at 1950 m. Depth is 2-3 m on average. It is a Ramsar Site (World Conservation Monitoring Centre, 1990; Scott, 1995). It is fed by precipitation, springs and small streams, with overflow through a small stream. The lake freezes over in winter. The submerged vegetation is abundant and there are extensive reedbeds of Phragmites communis, Juncus, Carex and Scirpus. It is under pressure from the population of the major city of Tabriz through sport fishing and wildfowl hunting as well as reed cutting and cattle grazing.

Qanats are found in this basin where surface water is saline. About 225 million cu m of water are produced annually by qanats and wells on the northern and eastern coast of the lake (Alamouti, 1966). Dams are found on the Zarrineh River and on the river which flows through Mahabad paralleling the Zarrineh. The Mahabad Dam has a fish catch of 130 tons (sic) annually and 300,000 fingerlings (species unspecified) were stocked to save the fish reserves from possible extinction (IRNA, 7 January 1999). The Mahabad reservoir has a leech fauna (Codonobdella trunata, Parcanthobdella livanowi, Baicalobdella torquata, Piscicola geometra) which may affect local fish farms and fish populations elsewhere if fish are transplanted (Abdi, 1999: www.mondialvet99.com, downloaded 31 May 2000). The Nowruzlu Dam on the Zarrineh is at 36°55'N, 46°10'E, occupying 1000 ha at 1260 m. It is water storage reservoir with heavy input from surrounding farming activities. The Alavian Dam near Maragheh is 80 m high, 935 m long and has a reservoir of 145 million cu m (http://netiran.com/news/IRNA/html/951214IRGG11.html). The Nahand Reservoir Dam northeast of Tabriz was inaugurated in 1995 with a capacity of 30 million cu m and a second dam, the Shahid Madani also near Tabriz, was under construction. Other dams include those at Ahar, Tabriz, Hashtrud, Hasanlu, Mianeh (= Onligh) and Heris which were scheduled to be completed in the period 1995-2000 (http://netiran.com/news/TehranTimes/95121601TTEC.html and www.irna.com/newshtm/eng/12003142.htm, IRNA, 2 July 2000). ? check basins for dams

The Hassanlou Reservoir Dam at Naqadeh was to open in 1998 with a height of 10 m, a crest of 5160 m (sic) and a capacity of 107 million cu m (http://netiran.com/news/IRNA/html/950915IRGG06.html). A total of 6 reservoir dams and 10 dams for re-directing water flow will decrease water input to the lake by 1.04 billion cu m by 2014. The volume of surface water has fallen from 42 to 22 billion cu m since 1995. The lake salt has increased to more than 260g/l, up from about 185 g/l. The lake may well dry up by 2014 (IRNA, 10 September 2001).

Khorasani et al. (2004) determined the environmental consequences of the construction of a dam on the Shahr Chai, a river 12 km southwest of Orumiyeh. Recommendations were made as to discharge and it was noted that fisheries potential would increase because of the reservoir.

Löffler (1993) details the eutrophication threat to this lake since a traffic embankment was built across the lake 35 km north of Orumiyeh in 1990. Untreated sewage from Orumiyeh will pollute the southern part of the lake.

Pollution occurs in various localities on a sporadic basis such as the Godar River in Naqadeh where a fish kill numbering in the thousands was reported (Tehran Times, 18 July 1999). Haji Hassani et al. (2004) found that levels of Ni, Pb and Cu in the Talkheh Rud were higher than acceptable limits for fish culture while Cr and Fe were lower. The river receives waste water from agricultural and industrial activities.

Water reservoirs behind the Mahabad, Miandoab and Shahid Kazemi dams were stocked with 3.6 million fish fry (species not specified) from the Pol-e Dasht Complex in 2000. This aquaculture site has the capacity to produce 4 million fry. West Azarbayjan produces over 600 tons of fish annually (Tehran Times, 2 January 2001).

Lake Orumiyeh is the largest natural habitat for brine shrimp in the world and, since 2000, is has been harvested, processed and used to feed sturgeon in hatcheries (www.worldfishingcompanies.com/html/us/world.report.html?id=1, downloaded 23 October 2001).

The lake was formed during the late Pliocene-Pleistocene and lies at 1275-1300 m and may well have had a Pleistocene connection to the Caspian Sea basin although this is in dispute (Scharlu, 1968; Schweizer, 1975). Pleistocene shorelines from 30 to 115 m above the present level have been confirmed, and the lake covered twice its present area, but this would not permit an external discharge. Berg (1940) reports benches at levels of about 1800 m, 1650-1550 m and 1500-1360 m, which may represent shorelines, and a level of about 1570 m would have had an outlet to the Aras River basin through the Kara-tepe Pass in the northwest and across the plain near Khvoy. Saadati (1977) suggests two connections with the Caspian Sea, an early one in the Pliocene to early Pleistocene resulting in endemic species and a later one in the late Pleistocene resulting in species which are the same as the Caspian or only subspecifically distinct. Stream capture may have allowed the entry of some species in recent times as evidenced by a Salmo cf. trutta/caspius population.

Fish farming is extensive in West Azarbayjan. In the Iranian year ending 20 March 2002, 840 tonnes of coldwater fish were produced and 3000 t of warmwater fish (Tehran Times, 24 November 2002).

Günther (1899) details a method of catching fish used in the rivers of this basin. Flour and the pounded berries of Cocculus indicus are mixed with butter to form a stiff paste. Small pellets of the paste are thrown into slow flowing water and after 10-15 minutes, if the fish are feeding, they will begin to swim at the surface in small circles or lie helpless in the shallows and are then easily scooped up. Some fish can recover from the poison. There is no effect on humans if poisoned fish are eaten.

Berg (1940) considers that this basin falls within his assignment of the Iranian shore of the Caspian Sea. Species in common include Leuciscus (= Squalius) cephalus, Barbus lacerta, Gobio (= Romanogobio) persus, Capoeta capoeta, Alburnoides bipunctatus (sic), and Silurus glanis, and Acanthalburnus urmianus is related to A. microlepis. Groombridge (1992) notes that the ichthyofauna of this region is badly in need of re-examination. Naseka (2010) recognises Urmia (Orumiyeh) Lake as a District within a West Asian Transitional Region related zoogeographically to the East Transcaucasian District (southern Caspian Sea area from the Kura River to the Atrek River). Both these Districts are linked to Iranian endorheic basins, including those listed as ecoregions in Abell et al. (2008), namely Namak, Kavir, Lut, Esfahan and Sistan, plus Kavir, Kor, Sirjan, Maharlu Kerman-Na'in and Jaz Murian basins in this work.

Namak Lake

This basin is flanked by the Alborz Mountains to the north and the Zagros Mountains to the west. On the east is the vast expanse of the Dasht-e Kavir basin and on the south such ranges as the Kuh-e Karkas at 3899 m (33°27'N, 51°48'E). The basin encloses about 87,600 sq km.

NASA and Wikimedia Commons.

A small sump near Arak (34°05'N, 49°41'E) is included as part of this basin as it is not separated by any major landform. A second salt lake is the Howz-e Soltan by the Tehran-Qom road and this lies in the same depression as the much larger Namak Lake south of Tehran. The lowest part of this basin is at 765 m and is covered by water in spring but this generally evaporates by the middle of summer.

The proximity of the capital, Tehran, to the rivers of this basin and its rapid growth in population and industry has led to many water diversionary schemes (Anonymous, 2003). A proposed dam northwest of Tehran would be the largest man-made lake in the country and the Middle East (sic) (Nouri et al., 2005). Much of this basin lies in Markazi or Central Province which has 42 dams of varying sizes. The Abbasabad Embankment Dam in Khomein, for example, is 36 m high, has a crest of 260 m and has a reservoir of 25,000 ha (IRNA, 3 February 1999).

The principal river in the west draining the Alborz south towards the Namak Lake is the Karaj River. Average temperatures of the Karaj River at the dam site before construction ranged from 2.5°C in January to 16.4°C in August (Nümann, 1966). Rieben (1954) and Hariri (1966) give details of surface and ground water in this river basin. The Amir Kabir Dam on the Karaj contains 205 million cu m of water and feeds through pipelines to Tehran. The reservoir has an area of 4 sq km at high water, 1.1 sq km at low water. Vladykov (1964) and Nümann (1966, 1969) give some details on the limnology of this reservoir, particularly temperature regimes. The Karaj has a discharge of 124 cu m per second in spring but this falls to 4.2 cu m per second in autumn. 55.6% of the annual discharge occurs during spring. There is no vegetation because of the steep rock sides and water fluctuations. Nümann (1966) recommended stocking the Karaj reservoir with Coregonus sp., Sander lucioperca, Acanthobrama terraesanctae (a Levantine species) and cichlids from Israel as environmental conditions and plankton levels were suitable. Nadim (1977) found the highest mercury levels in fish from the Karaj were 0.05 mg/kg. As the acceptable limit was 0.5 mg/kg, mercury contamination in fish was not considered a problem.

The Abhar River and its tributaries drain the land west of Tehran and south of Qazvin (36°16'N, 50°00'E). Its headwaters approach those of the Zanjan River, a Caspian Sea tributary. The course of the Abhar is about 350 km from its headwaters to the terminal sump. The lower part of this river is known as the Shur and is salty. Sewage and untreated factory wastes, as much as 40,000 cu m, flowed into the streams around the city of Qazvin although waste-water and sewage treatment plants are offsetting this problem (http://netiran.com/news/IRNA/html/941220IRGG05.html).

Other rivers draining the Alborz are much shorter. The Jajrud (Jaj, Jaji or Jaje River) to the east of Tehran is dammed at Latian (95 million cu m) for the Tehran water supply also. The Jajrud discharge is 60.5 cu m per second in spring and 1.5 cu m per second in autumn. Nümann (1966) reports fish kills in the thousands for Capoeta buhsei on turbid spring floods of this river. Khorasani (2001) give an environmental survey of this river. Mirzaei et al. (2010) give details of Eurasian Otters feeding on Alburnoides bipunctatus (sic, = namaki), Squalius cephalus and Capoeta spp. in the Jajrud. The Band Ali Khan River flows from the Khasrang Mountain (as does the the Jajrud which it receives) and its branches on the Varamin Plain are used for irrigation. Much of this river is polluted from wastes in the Jajrud and Tehran's sewage floodway (Rohani, 2004; Kashefi Alasl and Zaeimdar, 2009). The Lar River, a Caspian Sea tributary, was scheduled for diversion via a massive tunnel into the Jajrud (Marwick and Germond, 1975a; 1975b). This would affect flow in the Heraz River of the Caspian Sea basin and plans to offset this involved weirs and canal construction no doubt with the usual deleterious effects on fishes. These major projects are a far cry from the days in the twentieth century when Tehran depended solely on qanats for its water supply (Rieben, 1954).

The Namak Lake receives the Qareh Su (Gharechay), which flows north of Qom, and the Qom River from the Zagros Mountains. Discharge of both these rivers is about 312 cu m per second in flood falling to about 4 cu m per second in October (Oberlander, 1968b). The Qareh Su exceeds 400 km in length. The Qom River has captured headwater streams of Persian Gulf drainage. The Golpayegan River near Golpayegan has a storage reservoir, the Sadd-e Shah Esma`il. Borowicka (1958) gives some early figures on siltation and irrigation requirements. The Haroon Canal had diverted water for irrigation from the Golpayegan River for over 1000 years, and during the summer and fall all river water entered this canal. The Ghadir or Qadir Dam near Saveh has a volume of 290 million cu m of water. The 15th Khordad Dam is located 80 km south of Qom on the Qom-Delijan road (http://netiran.com/news/IranNews/html/95030718INPL.html). The Khandab Diversionary Dam is near Arak (http://netiran.com/news/IRNA/html/951217IRGG09.html).

Egglishaw (1980) gives some details on the water quality and environment of rivers and streams of this basin. Imandel et al. (1978) recorded ground water pollution by detergents in Tehran, where there was no method of sewage disposal other than discharge to wells and seepage pits. Södergren et al. (1978) reported on pollution with organochlorines in the Karaj and Latian reservoirs. Capoeta buhsei, Oncorhynchus mykiss, Alburnoides bipunctatus (= namaki) Coregonus sp. had accumulated the DDT metabolite p,p'-DDE, particularly in the Latian Reservoir. Direct removal of plants for fuel and laying bare the roots of such thorny plants as "giavan" for extracting gum tragacanth leading to plant loss has caused soil loss by erosion, gullying and affected recharge of groundwater. Poor farming practices on steep slopes has also led to the loss of topsoil such that runoff is too fast for infiltration of rain and snow (Rieben, 1954). These factors causing silting of reservoirs, added silt input to rivers and reduced groundwater recharge with consequent reduction in spring and qanat flows, all detrimental to fish habitats. Some areas of southern Tehran receive 300 kg/ha/yr of sulphate ions as acid rain which lowers river pH and has effects on the fish fauna (Salahi Kojoure, 1997). An effluent leak from a power station in the Vian area of Hamedan sent 40-50,000 litres of furnace oil into 1 km of river in the Qareh Su basin (Iran Network 1, Persian TV, 1730 GMT, 2 January 2000). Monavari and Mardani (2007) record the effects of sewage from fish culture ponds in this basin on water quality in the Jajrud, most factors being within acceptable limits except coliform bacteria.

Qanats are still a major feature of this basin. Alamouti (1966) records 260 qanats producing 99 million cu m per year on the Varamin Plain (35°20'N, 51°39'E), 220 qanats producing 161 million cu m per year on the Karaj Plain and 600 qanats producing 200 million cu m per year on the Qazvin Plain. However numerous pump wells have led to the drying of qanats and a complex irrigation system has reduced groundwater recharge (Beaumont, 1974). Alibekov (1994) gives a Russian account of qanats in Central Asia and also refers to those around Tehran in the Namak basin.

The Qazvin area has more than 20 aquarium fish farms producing over 2 million fish (www.tehrantimes.com, downloaded 28 July 2004). Waters in this area drain also to the Caspian Sea and there may be potential for escapes of exotics.

Berg (1940) refers this basin to his Tehran District of the Iranian Province. He notes that some drainages are close to those of the Caspian Sea basin and that the fauna may be of quite recent origin, rather than the Pliocene advocated by Derzhavin (1934) for Salmo trutta or presumably now caspius). Saadati (1977) considered that the fish fauna of this basin was not derived from movements through a large freshwater lake connecting all the tributaries. Some species came from the Caspian Sea basin and others from the Esfahan and Tigris River basins. The basin may also have served as a "filter-bridge" allowing such species as Capoeta aculeata, Capoeta capoeta and the progenitor of Capoeta fusca to reach the Dasht-e Kavir basin.

Sirjan

The Sirjan basin extends south-east of the Esfahan basin and parallels the Kerman-Na'in basin. It is named for the town of Sirjan at 29°28'N, 55°42'E which lies at the edge of the largest salt flat in the basin. It is somewhat higher than the Esfahan basin which is at 1300 m, being 1448-1710 m. It is distinguished from the Esfahan basin by its lack of a significant river. There are four major sumps in this basin, strung out along its length at regular intervals, and the northern two are connected as are the southern two. The sumps are fed by intermittent streams. Qanats and minor springs are found in this basin which has not been extensively explored. The sump in the north near Abarqu (31°08'N, 53°17'E) receives streams from the west (Kuh-e Bul at 3661 m and 30°48'N, 52°45'E) and from the east (Khar Kuh at 3512 m and 31°39'N, 53°46'E, and Shir Kuh at 4074 m and 31°37'N, 54°04'E). The southern basins near Sirjan receive their streams from lower elevations.

Sistan

The Sistan (= Seistan) basin straddles the Iran-Afghanistan border and is a north-west to south-east oval in shape. It comprises a number of minor streams and qanats flowing from the west and the Birjand highlands, but these are rapidly absorbed or run for only a few days each year. Its most obvious feature is the vast hamun or swamp comprising open freshwater lakes, reed beds or neizar, and the rivers that feed the lakes. This is a major oasis of fresh water surrounded by hundreds of kilometres of arid plains. Huntington (1905a; 1905b), Annandale (1919a), Ahmadi and Wossughi (1988), Noorbakhsh (1993), Mansoori (1994), Ibrahimzadeh (1995), Scott (1995), Weier (2002),  CIRSPE (2006a) and van Beek et al. (2008) give descriptions of this basin. Note that Weier's (2002) statement (repeated in various newspaper reports and in UNEP (2003)) that there is nearly 140 species of fish in Sistan is an error by an order of magnitude! The native ichthyofauna comprises a mixture of endemic species, species related to or conspecific with high-altitude species from Central Asia and species from Baluchestan in the wider sense. There is little relationship to species from Iran to the west. Variations in water level and crowded conditions lead to disease and parasite outbreaks in the fishes (Mansoori, 1994).

The principal river is the Helmand (or Hirmand) which flows from the Paghman Mountains just west of Kabul to end in Sistan after a journey of 1400 km. Along with the Hari or Tedzhen, this is the only major river entering Iran. Snow and rain in the Hindu Kush mountains ultimately reaches Sistan at 427 m from heights of 5300 m. The Helmand is the most important river between the Tigris and the Indus and drains an area of 386,000 sq km of which 78,000 sq km or 20.2% lies in Iran (Gleick, 1993).

The Helmand produces 1700-2000 cu m per second in flood and 56 cu m per second in the dry season. The average annual flow is 78 cu m per second. The river varies between 200 and 900 m in width and between 2 and 5 m in depth. The annual water income to Iran is about 6 billion cu m but this varies markedly and was 14,740 million cu m in 1970-1971 and 1976-1977 and 600 cu m in 1985-1986 (Mansoori, 1994). UNEP (2003) gives the following flows in million cu m:-

1991-2 1992-3 1993-4 1994-5 1995-6 1996-7 1997-8 1998-9 1999-2000 2000-1
2211.7 1783.8 529.5 829.7 1023.8 908.7 2193 258.8 114.1 48

As it enters the Sistan depression, the Helmand splits into several branches which feed the swamps, the two main ones being the Sistan feeding the Hamun-e Helmand (also Hirmand or Hamun Lake) in Iran and the Parian feeding the Hamun-e Puzak (or Parian) lying mostly in Afghanistan. The northern part of the Hamun-e Helmand is called Hamun-e Sabari, or Lake Sistan, which lies half in Afghanistan and half in Iran, and the southern part is called Hamun-e Hirmand. Hamun-e Sabari receives water from the Farah River and overflow from Hamun-e Puzak. The Hamun-e Hirmand receives water from the southern or Sistan branch of the Helmand River and overflow from Hamun-e Sabari. Other rivers flowing from Afghanistan are the Harut, Khospas and Khash but their flow is minor and intermittent compared to the Helmand (Gabriel, 1938). The whole lake area of Sistan is often called the Hamun Lake.

Sistan lakes from NASA and Wikimedia Commons.

The plentiful natural flow of the Helmand is reduced by irrigation dams in Afghanistan; the Arqhandab and Kajaki dams extract about half of the 12 billion cu m which enter the Afghan plain (Michel, 1973; Mansoori, 1994; Mojtahedzadeh, 2001). A third dam is under construction in Afghanistan without environmental considerations being taken into account (World Conservation Monitoring Centre, 1990). The proposed Kamal Khan Dam on the Helmand in Afghanistan and the "Sistan Drainage and Irrigation Completion and Rehabilitation Project" in Iran would lower water level in the lake complex. There are also plans to divert water from the Sistan area to the city of Zahedan in the south. The Char-Neimeh (or Chahnimeh) Lake is a depression used as a water reservoir and is filled from the Parian branch of the Helmand. It has a surface area of 4,700 ha and is used for irrigation and fish culture but does reduce flow into the hamuns. However floods in spring 1991 destroyed the Kajaki Dam and associated irrigation controls and the lakes were more extensive than they had been in over a decade. Rainfall in Afghanistan increased flow of the Helmand in 2003 and some flooding was expected in Sistan (www.irna.com, downloaded 23 April 2003). The Helmand was dry at the Iran-Afghanistan border in 2004 (Gall, 2004). Sadeq (1999) lists several factors which are threatening the Hamun Lake namely, fluctuation in incoming water, sedimentation, exotic species, urbanization and increased population pressure on the hamun resources.

The south end of Hamun-e Puzak and the contiguous Hamun-e Sabari (or Lake Hamun) are Ramsar Sites (World Conservation Monitoring Centre, 1990). The Lake Hamun Ramsar Site is on the threatened list of National Parks (Anonymous, 1988b).

Puzak is very shallow, with maximum depth of less than 4 m, and is the first of the Sistan lakes to flood and may never dry out completely unlike the other lakes (Khan et al., 1992; Scott, 1995). This lake has extensive reed beds of Phragmites australis with associated submerged Ceratophyllum demersum and relatively little open water. Reeds are cut as forage for cattle, burnt to improve grazing for livestock, used for boats, for wind-breaks and for cooking and heating. Local people engage in fishing.

The Helmand is very turbid and deposits 8 g of silt for each litre of water (Fisher, 1968). The sediment load in 1975-1976 was 15,149,000 t and in 1985-1986 280,000 t (Mansoori, 1994). Drinking water looks like milk! (personal observations, 1977). Rain accounts for little input to the lake, the annual mean precipitation over 12 years being only 51 mm, most rain falling within 10-15 days (Mansoori, 1994). UNEP (2003) reports evidence of pesticide pollution in the Helmand and the swamps, e.g. dieldrin.

The lake bottom in Iran is clay and silt and the waters are markedly alkaline. Water at the edges of the reed swamp were 31°C in early May, warmer than the inflowing rivers and the irrigation ditches which were only 22°C at this time. Annandale (1919a) and Mansoori (1994) give a brief chemistry of Sistan water. There are marked variations in conductivity, temperature, pH, oxygen, alkalinity and hardness between sites. Conductivity ranges from 1280 to 64,000 mmhos (sic), pH from 7.5 to 9.15, oxygen from 0.64 to 11 mg/l, alkalinity from 3.6 to 165 mval and hardness (CaCO3) 180 to 3500 mg/l in Mansoori's water samples from the Hamun Lake.

Evaporation lowers the water level each year and is caused by extreme heat and the famous Bad-e Sad-o Bist Ruz (Wind of 120 Days) which approaches 200 km per hour. This wind causes serious erosion and marching sand dunes often block streams causing them to change channel. Evaporation has been measured at 4 m per year because of temperatures over 40°C in July (Mansoori, 1994). Refilling occurs in February-June and in flood years various hamuns are joined together into one vast lake. 75% of flooding occurs in March-May. There are about 3900 sq km of seasonal lake and marsh at a maximum, dropping to 1930 sq km in July-January. The maximum flood zone is about 200 km long and 20 km wide, but the lakes have dried up completely, or almost so, at least 5 times in the past 100 years, e.g. in 1907, 1962 for 5 years, 1970-1971, 1984 for 4 years, 1988-1989, and 1998-2002, with major fish kills resulting (Tate, 1910; Harrington, 1976; Costantini and Tosi, 1978; Anonymous, 1992a; Khan et al., 1992; MacFarquahar, 2001; Foltz, 2002; Weier, 2002; www.netiran.com, downloaded 18 June 2002). There was a big flood in March 1989, spring 1990 and an exceptional flood in February/March 1991 (Khan et al., 1992). The lakes filled in 2005 (E. Penning, pers. comm., 28 July 2005). Mansoori (1994) mentions historical floods, e.g. in 1247 A.D., and droughts, e.g. in 835 A.D. UNEP (2003) gives satellite photographs showing variations in water extent. The fish fauna can recolonise newly-flooded marsh areas from the Helmand but population numbers in the hamun vary greatly between years.

Sistan showing variations in water extent, U.S. Geological Survey and Wikimedia Commons.

The centre of the hamun is only about 2-3 m deep on average with a maximum depth of 5 m at highest water level (www.bibliothecapersica.com/articlenavigation/index.html, under hamun, downloaded 24 December 2004, gives 11 m). Overflow spills into the salt flat Gowd-e Zereh of Afghanistan through the Shelah River. This flushing effect probably prevents this endorheic basin from becoming saline. The Shelah was reduced to isolated and fishless pools in May 1977. The Gowd-e Zereh is at 467 m at its lowest point.

Extensive canals and ditches form a network over Iranian Sistan and serve to irrigate and drain fields. These waters contain fish, but may dry up. The Hirmand is dammed to feed the major canals. The open lake areas are fringed by reed beds comprised of Typha, Phragmites and Scirpus which are concentrated at the ends of the detrital cones of the river deltas (Costantini and Tosi, 1978). Mansoori (1994) and Ibrahimzadeh (1995) report an absence of Phragmites in area which was two-thirds covered in previous studies, drought being advanced as the causative agent along with cattle grazing (Khan et al., 1992). Usually the reeds recover after drought but in 1991 this did not happen (probably the effects of introduced Ctenopharyngodon idella on the young shoots since fenced areas excluding fish show successful reed growth). Two million fish were introduced in early January 1992 near Kuh-e Khvajeh. Scott (1995) also suggests that local people may have dug up tubers to use as fuel. A major fish and bird kill occurred in November 1994 but the cause was never ascertained (Scott, 1995).

Agricultural land around the Sistan lakes is being abandoned because of increasing soil salinity. Wind-blown salt is becoming a problem in summer and the area might suffer the same fate as the Aral Sea (Scott, 1995). A new road running between the Sabari and Helmand lakes in the Ramsar Site may impede water flow despite bridges having been constructed. A canal between Puzak and Sabari will also have major hydrological impacts.

Curiously, both the open lake and the reed beds are poor in fish but channels among the reeds and areas at the edge of reed beds are productive. The effluents of the Helmand are particularly productive and provide a refuge for fish if the lakes dry out. Annandale in Annandale and Hora (1921) gives an interesting account of the fisheries of the Sistan lakes in the early years of the 20th century. Only one species, Schizothorax zarudnyi, was pursued (q.v.) using reed boats or skiffs called tutin which were still in evidence in the 1970s. The introduction of exotic species resulted in an increased fish catch in the 1980s and 1990s and the number of active fishermen was 1090 (Abzeeyan, Tehran 5(5):III, 1994, M. H. Karim Koshteh, in litt., 2003). However, Ibrahimzadeh (1995) reports that there is no fish catch in the lake. Local people took more fish as the population increased (4% per annum, with added impact from Afghani refugees), as transport facilities improved and as animal husbandry decreased through degradation of reed beds (M. H. Karim Koshteh, in litt., 2003). The Islamic Republic News Agency (IRNA, 22 March 2000) reports a catch of 7000 tonnes from the Hamun Lake; the following figures are from M. H. Karim Koshteh (in litt., 2003):-
 
Year 1990 1991 1992 1993 1994 1995 1996 1997 1998 1999 2000
tonnes 2790 3520 4380 4106 3543 5998 4251 3900 6044 12,000 2426

Meijer (2006) gives an estimated catch in a semi-wet year as high as 21,840 tons although official figures give 9000 t. Variations reflect drought conditions, the year 2000 being particularly severe. Fluctuations in catches make the fishery a difficult occupation. Gilkolaei (2007) estimates a commercial catch of stocked fish at 22.5-45,000 tons/year in the whole Sistan basin.

Sistan has fish farming in various water bodies. In 2005, 1.3 million juveniles of grass carp, common carp, bighead and silver carp produced by the Zahak hatchery were stocked in farms (www.iranfisheries.net, downloaded 17 January 2005; CIRSPE, 2006b). Goldfish and silver carp are exotics found in the hamuns (E. Penning, pers. comm., 28 July 2005). CIRSPE (2006a) also lists Rutilus frisii, Abramis brama and Sander lucioperca, all Caspian Sea basin species, as being present in Sistan but this may be an error. Gilkolaei (2007) discusses breeding of Schizothorax zarudnyi, culture of Ctenopharyngodon idella, Hypophthalmichthys molitrix and Oncorhynchus mykiss and ornamental fish breeding in this basin.

Berg (1940) places this basin in his Sistan District of the Iranian Province. It excludes the upper reaches of the Hirmand River. The schizothoracine fauna is particularly characteristic and had its origins either by descent from higher altitudes during the Pleistocene glaciations (favoured by Berg) or are autochthonous as the forms at high altitudes in the Pamirs and Himalayas rose with mountain building.

Tedzhen River

The Tedzhen River is the more familiar, international name and is used here. In Iran this major river is known as the Harirud or Hari River. The Tedzhen rises in the Selseleh-ye Kuh-e Baba of Afghanistan and flows west for about 490 km before turning north as the Iran-Afghanistan border for 160 km. Along with the Hirmand and Aras, this is the only major river entering Iran. At Sarakhs (36°32'N, 61°11'E) it enters Turkmenistan and is known there as the Tedzhen, and is eventually lost in the Karakum desert. The river is usually dry even at Sarakhs (Barthold, 1984). Most of the water in the Tedzhen remains in Afghanistan where it is used for irrigation of the Herat valley. Spring floods (March-April) can increase flow ten-fold for short periods of time. The Jam River is a southern tributary from Iran, draining the mountains around Torbat-e Jam (35°14'N, 60°36'E) and the Kashaf River is a northern Iranian tributary draining past Mashhad from the northern slopes of the Kuh-e Binalud (3416 m at 36°30'N, 58°55'E) and the southern slopes of the Kuh-e Hazar Masjed (3146 m at 36°52'N, 59°26'E). The Kashaf is about 310 km long. Its discharge is comparable to, if not as great as, central Zagros streams and is larger than the plethora of minor streams draining the Alborz (Oberlander, 1968b). The upper reaches of the Kashaf approach those of the Atrak River, a Caspian Sea tributary, and are separated by only a small upfold. This area is very unstable with frequent earthquakes. The catchment area for the Tedzhen basin approaches 45,000 sq km (Pirnia, 1951).

Bazangan Lake between Mashhad and Serakhs (36°17'N, 60°29'E) is the largest natural lake in northeast Iran with an area of 690,000 sq m and a maximum depth of 6-11 m. It is hyposaline oligotrophic with low phyto- and zooplankton communities, and with a corresponding low diversity of fishes (Ghassemzadeh, 2004).

Najafpoor et al. (2007) give a water quality assessment for the Kashaf River and note its use for water supply, agriculture fishing and recreation. Pollution from agriculture, and from industrial and municipal wastes at Mashhad, is recorded. Supersaturation from excessive plant life and low night-time levels of dissolved oxygen through respiration could lead to fish kills in the Kashaf.

A number of minor streams drain northward from the Koppeh Dagh (= Kopet Dagh or Kopetdag) in the west, a range which straddles the border of Iran and Turkmenistan in this north-eastern part of Iran, and from the Hazar Masjed and intervening ranges in the east. These have not been collected. The Iranian tributaries of the Tedzhen have not been well collected either, but there is data on the fish fauna from both Afghanistan and the former U.S.S.R. (now Turkmenistan). Coad (1981d) lists fishes from Afghanistan, and Aliev et al. (1987; 1988), Starostin (1992) and Salnikov (1994) fishes from Turkmenistan. Aliev et al. (1987) list rare and endangered species in Turkmenistan.

There is evidently a strong possibility of exotic species from Turkmenistan entering Iranian waters via the Tedzhen drainage. Fishes, including exotics, are farmed along the basin of the Karakum Canal, a 1372 km long diversion from the Amu Darya. Some of these exotics can be expected to enter the Tedzhen River basin via its delta and eventually the Caspian Sea basin via the Atrek River through runoff and collector canals (Sal'nikov, 1995; 1998). Potential exotics for Iran from the Karakum Canal include Pseudoscaphirhyncus kaufmanni (Acipenseridae), Alburnoides taeniatus, Aristichthys nobilis (= Hypophthalmichthys nobilis), Aspiolucius esocinus, Aspius aspius iblioides, Barbus (= Luciobarbus) capito conocephalus, Capoetobrama kuschakewitschi, Carassius auratus gibelio, Chalcalburnus (= Alburnus) chalcoides aralensis, Ctenopharyngodon idella, Hemiculter eigenmanni (= leucisculus), Hypophthalmichthys molitrix, Mylopharyngodon piceus, Parabramis pekinensis, Pseudogobio rivularis, Pseudorasbora parva, Rhodeus ocellatus, Rutilus rutilus aralensis (all Cyprinidae), Cobitis aurata aralensis, Misgurnus anguillicaudatus (Cobitidae), Barbatula oxiana (Nemacheilidae), Gambusia holbrooki (Poeciliidae), Oryzias latipes (Oryziatidae), Channa argus warpachowskii (Channidae), Micropercops cinctus (Odontobutidae), and Rhinogobius brunneus or Rhinogobius similis (Gobiidae). A Rhinogobius species is now found in Iran (Coad and Abdoli, 2000a; Abdoli et al., 2000). Other species not native to the Tedzhen basin but found elsewhere in Iran are also reported such as Acipenser nudiventris (Acipenseridae), Pelecus cultratus (Cyprinidae), and Sander lucioperca (Percidae). Cyprinus carpio stocks are a mix of native and Chinese imports. Silurus glanis (Siluridae) has also been introduced along with carp from the Amu Darya although it is also native. Sal'nikov (1995; 1998) also lists other species which may penetrate the canal eventually. These exotics have a great potential to cause devastation in the native fauna through competition and through genetic swamping of related taxa.

The fauna of the Tedzhen basin is found in rivers and streams as well as springs and qanats. Dams include the Barzou, 40 km north of Shirvan, which is 85 m high with a crest of 325 m and the Shirnin Darreh north of Bojnurd which produces 60 million cu m of water for irrigation (Iran News, 17 September 1997). A dam is scheduled for completion in 2005 at the Iran-Turkmenistan border. It will have a capacity of 1,250 million cu m of water (IRNA, 3 September 1999).

Berg (1940) places this basin as a part of his Turkmen District of the Iranian Province (other parts include the Murgab River of Afghanistan and Turkmenistan and northslope streams of the Kopet Dagh in Iran and Turkmenistan). He considers that the Hari River once belonged to the Amu Darya basin of Central Asia.

Kerman-Na'in

The Kerman-Na'in basin extends from Ardestan (33°22'N, 52°23'E) in the north-west to Kerman (30°17'N, 57°05'E) in the south-east. It is an elongate series of small basins combined here for convenience and named for two major towns at the ends of the basin. Its length exceeds 600 km and its maximum width is 175 km. An almost continuous range of mountains, paralleling the Zagros, flanks this basin on the west, while the eastern edge is lower and abuts the Dasht-e Kavir and Dasht-e Lut basins, particularly in the north-east. The Kerman-Na'in basin lies at a similar altitude to the other interior basins, ca. 1000 m.

In the south-east, streams drain the mountains ringing Kerman, such as the Kuh-e Hazaran at 4420 m (29°30'N, 57°18'E), the Kuhpayeh at 3142 m (30°35'N, 57°15'E), and the Kuh-e Masahim at 3600 m (30°21'N, 55°20'E), to a sump just west of Bafq (31°35'N, 55°24'E). These streams bear names such as Namak and Shur and may well be inhospitable to fishes. Several streams between Kerman and Yazd marked prominently on maps were dry in January. Irrigation requirements may have reduced their flow and most of the fishes from this area are to be found in qanats. Qanats have temperatures in this region of 17-21°C in January and have been studied in one village by Smith (1953; 1979).

Around Yazd streams drain the Shir Kuh at 4074 m (31°37'N, 54°04'E) and the Khar Kuh at 3512 m (31°39'N, 53°46'E) but there is no major terminal sump. Some of the streams enter the Bafq sump while others drain north to a sump near Na'in (32°52'N, 53°05'E) which also receives intermittent streams from around Na'in.

Intermittent streams from the Kuh-e Karkas at 3899 m (33°27'N, 51°48'E) drain to a sump near Ardestan but, as in the southern parts of this basin, are not a prominent feature of the landscape and fishes are mostly to be caught in qanats.

The underground water resources of Yazd Province have been examined in a newspaper article (Hamshahri, Tehran, 629:5, 22 February 1995) and, although the province is not the same area as the drainage basin outlined here, it is indicative of the underground water resources of this part of Iran. These resources comprise 1751 subterranean water canals (probably this means qanats), 2084 semi-deep wells and 897 deep wells with an annual discharge of 1100 million cu m of underground water. The authorised capacity is 893 million cu m and the excess removal has resulted in an annual drop in the water table of 70 cm. In addition, chemical and biological pollution of groundwater is a continuing problem and these factors too will affect fish survival.

Much of the fish fauna of the Kerman-Na'in basin appears to be restricted to qanats, although there may be a fauna in high mountain streams not readily accessible by road.
 


Scientific Names

The common names of fishes vary with language between countries and within a country with local usage. This problem is overcome to the scientists' satisfaction by the scientific name, consisting of two words, the genus name and the specific or trivial name. A genus, e.g. Luciobarbus, may contain many species but each species is a unique combination of Luciobarbus and a specific or trivial name. This scientific name is used the world over whatever the local common name may be. It is always written in Latin script and the genus and trivial names are derived from and spelt according to rules of grammar in Latin and Classical Greek. Both these languages are "dead" so the rules and spelling are fixed and not subject to change with time as modern languages are. It is generally felt that the advantages of this system outweigh the unfamiliarity of Latin and Greek words and grammar for most people.

As an example of the scientific name, we can consider the first species dealt with in this work, the Caspian lamprey or Volga lamprey Caspiomyzon wagneri (Kessler, 1870). The scientific name is underlined or set in italics or bold face to denote its scientific status. Caspiomyzon is the genus name and wagneri the trivial or specific name. This species was first described by Kessler in a publication dated 1870. Kessler placed this new species in the genus Petromyzon but L. S. Berg later published reasons for placing it in a distinct genus, Caspiomyzon. The parentheses around the author (or first describer of the new species) and the date of description indicate that its generic allocation has been changed.

Sometimes the author of a work (paper or book) is not the describer of the new species, e.g. the multi-volume work by G. Cuvier and A. Valenciennes (1828-1849, Histoire naturelle des poissons, 22 volumes - see above) continued to appear after Cuvier died. For many years, the species author appeared in taxonomic works as "Valenciennes in Cuvier and Valenciennes" to indicate that Valenciennes described the species but the description appeared in a volume of the book on whose title page both Cuvier and Valenciennes appeared as authors. Bailey (1951) determined who authored which species in this case. The trend now is to cite simply a single name, Valenciennes in this example, and this is seen in "FishBase" and "Catalog of Fishes".

Another example of a confusing author name involves Francis Buchanan (see above for details of fishes described by him). His name also appears as Hamilton or Hamilton-Buchanan or Buchanan-Hamilton - the name Hamilton was assumed on succeeding to property in Scotland from his mother, formerly a Miss Hamilton.

The scientific name is also used to show relationships between species and it can therefore be changed if views on the relationships of the species are changed according to the "International Code of Zoological Nomenclature". The Fourth Edition of the Code came into effect on 1 January 2000. Errors also arise in giving species scientific names and these must be corrected by name changes according to the Code. The Code is complicated and detailed explanations based on fishes may be found in Eschmeyer (1998; this Catalog of Fishes is now online). Some of the more common reasons for name changes are given below.

A single species may be described twice, either by the same person or by two people. At the time of these descriptions it was genuinely believed that there were two species but subsequent studies showed that they were the same. This error often arises with confusion between juveniles and adults and between males and females which may be quite different in appearance. Older collections from remote areas often comprised only a few specimens and could be in rather poor condition by the time they came into the hands of an ichthyologist and were described scientifically. An example of confusion of males and females of the same species is found in the genus Aphanius. In 1910, J. T. Jenkins described three species of Aphanius (under the genus Cyprinodon as it was recognised then) from near Shiraz. This material had been collected in 1872 by W. T. Blanford and was comprised of 10 specimens, mostly in good condition. The three species were Cyprinodon blanfordi, C. persicus and C. pluristriatus. The first of these was thought by Berg (1949) to be a female Aphanius sophiae and the latter two to be males differing in characters not now considered to be specifically important. I have a differing opinion! It is also possible, where two people are concerned, that the author who published his description later was ignorant of the first author's work. The first name published has priority and the second name is called a synonym and is no longer used. There may be several synonyms for a species. These are listed in the species descriptions. There is also the problem of misidentification of specimens. When these specimens are available for study identification can be confirmed (or amended) but often specimens are discarded or lost. These errors too may be listed in a synonymy. Krupp (1984a) gives a synonymy for Aphanius cypris which amply illustrates how a scientific name may be mis-applied (there are 89 uses of names which all refer to one species in Krupp's opinion). A. cypris is now thought to be correctly named A. mento.

Occasionally the same name is given to two distinct species because the later author was not aware that the name had already been used. The name of the species described first is called a senior homonym and is retained while the later species name, the junior homonym, must by replaced.

The genus name of a species can be changed because an ichthyologist, who has studied the species and its relatives in detail, considers that it is more closely related to another species or group of species with a distinct genus name. A similar case was discussed above with Caspiomyzon wagneri where a new genus was erected for this species. In both cases, parentheses are placed around the author's name and the date of description to indicate that the genus name used has been changed. The species placed in a different genus will retain its trivial or species name unless this trivial name is already in use in the different genus. Homonymy has then occurred and the species which has priority retains its trivial name and a replacement name must be given to the more recently described species. It is not unusual for scientists to disagree about the interpretation of the same data and a species may have a long and complex career being switched from genus to genus as publications advocate one view or another of its relationships.

There is a higher classification which groups together related genera into Families, Families into Orders and Orders into Classes. The vast majority of Iranian freshwater fishes belong to the Class Actinopterygii, the ray-finned bony fishes, with only the Caspian lamprey in the Class Cephalaspidomorphi. Some sharks penetrate freshwater and these belong to a third Class, Chondrichthyes or cartilaginous fishes.
 


Fish Structure

A knowledge of fish anatomy is essential in identifying specimens. The head of a fish carries a number of structures. The eyes are without eyelids although sharks have a protective membrane, the nictitating membrane, which acts as an eyelid. Eye size varies with age within a species but can also be a distinguishing characteristic between species. There are nostrils, for detecting odours, on the snout, that part of the head before the eyes. Nostrils are blind sacs and do not connect with the mouth cavity. Their position and shape may be useful characters. Barbels are slender, fleshy structures on the snout or chin used for touch and taste. Their presence, number, position and length are important characters. Sharks and sturgeons have a small opening near the eye called the spiracle, not found in the bony fishes. Teeth may be found variously on the tongue, roof and floor of the mouth and even in the throat. The pharyngeal teeth of Cyprinidae are often useful characters in identification and may be dissected out from the posterior part of the gill cavity under the operculum using dissecting equipment. This requires some practice to avoid damaging the specimen too extensively. Some teeth are sharp and pointed for piercing and holding prey, while others are rounded and heavy for crushing food items covered by a protective shell. The side of the head behind the eye is the gill cover in bony fishes, composed mainly of one bone, the opercle, which protects the gills. The gill cover opens posteriorly; bony fishes have one opening on each side of the head, but lampreys have seven rounded openings and sharks five to seven vertical slits. The cheek is the area between the gill cover and the eye. Spines and scales may be found at various places on the gill cover and cheek. A membrane is found below the gill cover, supported by thin slivers of bone, the branchiostegals, and connected with the gill cover on the other side of the head. Under the gill cover lie the gills which serve in gaseous exchange. Gill rakers on the front of each gill arch serve to prevent food from damaging the gills and direct food into the gut. Rakers may be short and widely spaced where food items are large and easily deflected, or long and close together where food items are minute like plankton.

The head leads directly to the body; there is no neck. The body is made up mostly of a trunk. The caudal peduncle or tail stem starts behind the anal fin and ends at the tail fin. The number and presence of different types of fins on the body varies with the species of fish and is often a useful character for identification. The back may carry 1-3 dorsal fins and an adipose fin between the last dorsal fin and the tail fin. The tail (or caudal) fin is at the end of the body and may be forked, square cut, rounded, pointed, lanceolate or lunate. Its skeletal structure may be almost symmetrical or upturned at the end. This upturn is obvious in sharks and sturgeons which also have a large upper lobe to the tail fin and a smaller lower lobe. The anal fin, or fins, lies on the underside of the body surface behind the vent which is the exit for the intestine, kidney ducts and gonads. The pectoral fins are found behind the gill cover on each side of the body and a pair of pelvic fins are behind (abdominal), below (thoracic), or in front (jugular) of the pectorals on the lower body surface. An axillary pelvic scale above the pelvic fin streamlines the fin when it is pressed against the body. The pectoral fin may also have an axillary scale. All the fins except the fleshy adipose fin are supported by rays. Soft rays are flexible and jointed while spines are rigid, pointed and unjointed. The number of soft rays and spines in the various fins is very useful for identification.

Most fishes have a body covering of scales which may extend onto the head and certain fins. Notable exceptions are the catfish families Bagridae, Siluridae, Sisoridae and Heteropneustidae, which are completely naked. Rounded, smooth scales are called cycloid and are found in less advanced bony fishes. Large cycloid scales may easily detach, as in herrings (Clupeidae) but small cycloid scales can be embedded and hard to see as in the eel (Anguillidae). Ctenoid scales bear small teeth on the posterior margin and feel rough to the touch. Such scales are found in the more advanced bony fishes such as Percidae. Sturgeons have heavy bony plates called scutes. Sharks have placoid scales which can be so rough as to scrape the skin off a human. The teeth of sharks are modified placoid scales. Scales grow with the fish, laying down rings of material as do trees. In areas with a change of seasons, the growth rings are widely spaced during the summer growing season and cramped together in winter when growth is slow. Fish age can be determined from these rings. The energy expended in spawning is reflected in the scales which may resorb the edge producing a spawning check. A fish which lives and grows slowly in fresh water and then migrates to the rich feeding grounds of the sea will have this history reflected in the spacing of the growth rings. Scales can be "read" to reveal much about the life history of an individual fish. The scales also bear radii, or radiating lines, and their distribution can be useful in identification along with other scale characters such as shape and focus (growth origin) position. The scales are covered by an almost undetectable layer of skin. The skin contains mucus cells which give the fish a slippery feel and colour cells which give the fish its colour. Some fish are characteristically more slimy than others, e.g. the eel. Most fish have a distinctive colour pattern but this can change with age, maturity, behaviour, background, between sexes, and after death.

Fishes have a sensory lateral line system which runs along the flank and a similar system on the head. The extent and development of these systems varies with the species of fish. The lateral line is a tube in the skin with openings to the outside through pores in the scales. A lateral line pore count is often used in separating fish species.

The internal structure of a fish may be summarised as follows. The gills and teeth have already been mentioned. After these structures, the mouth cavity narrows to an oesophagus which passes to a straight, U- or J-shaped stomach. Pyloric caeca, which produce enzymes, may be attached at the junction of the stomach and intestine in some fishes and counts of these caeca are used in identification of some species. The intestine ends at the vent. The length of the intestine varies with the diet. Fishes which feed on plant material have long guts while those that feed on animals have a short, often s-shaped intestine. Fish have a liver, a reddish organ at the front of the body cavity. The liver may be very large in sharks and form a significant part of the body weight. There may be a small, green gall-bladder associated with the liver. The swimbladder (gasbladder) is a gas-filled sac with thin walls lying near the top of the body cavity where it functions as a buoyancy organ and can be used to transmit sounds to the brain or even produce sounds by means of special drumming muscles. The swimbladder shape has been used to characterize species. Some fishes have a poorly developed swimbladder or none at all, since they live on the bottom of stream beds and must avoid being swept away. Just below the backbone above the swimbladder are two long, dark-coloured kidneys and below these are the ovaries, which may be filled with eggs, or the testes which produce the sperm. A small urinary bladder lies at the end of the body cavity. The body cavity is lined with a membrane which may vary in colour from silvery-white to jet black. The main body muscles are in the form of W-shaped, interlocking blocks and this arrangement helps produce the sinuous body movements by which fish swim.

Lampreys (Petromyzontidae) differ in structure from bony fishes. They lack true jaws and have a round, suctorial mouth armed with teeth. There is a single nostril on top of the head rather than a pair on each side. There are no scales or paired fins (pectorals and pelvics). There are seven rounded gill openings in a row behind each eye. The larval lamprey is called an ammocoete and lives buried in fine sediments, filter feeding minute particles from the water. In this stage it lacks teeth and the eye is poorly developed.

Sharks also have a somewhat different structure from bony fishes. Some species produce living young rather than eggs, while in others the embryo is laid in a horny egg-case known as a mermaid's purse when it washes up on a beach. Male sharks have claspers derived from the pelvic fins, which serve to ensure that sperm are delivered to the female. The length of time food stays in the gut of sharks, and also sturgeons, is increased by a spiral valve. The food follows the spiral around rather than going straight through the gut and so there is more time for digestion and absorption. There is no swimbladder in sharks, which have to swim constantly to stay above the bottom. Sharks produce teeth in multiple rows, and as older teeth at the front of the jaw fall out, new ones move forward to replace them.

The skeleton includes the skull comprising the cranium, which contains the brain, the jaws, gill arches, operculum and other associated bones. The cranium also contains small objects known as otoliths in the inner ear. These aid in sensing change of direction and in balance. Otoliths can be characteristic of species. There is a vertebral column with ribs anteriorly enclosing and protecting the body cavity and its contents. The number of vertebrae is a useful character and can be counted easily, without damage to the fish, by taking x-rays. A tail skeleton supports the tail fin and the pectoral and pelvic girdles support their respective fins. There are fin supports too for the dorsal and anal fins. Lampreys, sharks and sturgeons have a skeleton composed of cartilage, a substance not as strong as bone, but when impregnated with salts (like shark teeth) are remarkably effective.

Most characters used for fish identification are external for convenience. The most used internal characters are gill raker counts, pharyngeal teeth counts, gut shape and body cavity lining colour, pyloric caeca counts and vertebral counts.

The general structure and biology of fishes is covered in various general works. Coad (1993; 1995b) gives a list of general ichthyology texts and the Dictionary of Ichthyology describes various anatomical terms.
 


Collecting Fishes

Collecting methods and literature are summarised by Coad (1993; 1995b). Luck plays a part even in scientific collecting as discovery of new species in areas previously sampled demonstrates (e.g. P. G. Bianco did not collect Petroleuciscus persidis in Fars, while I found several populations of that previously unknown species; conversely he found several populations of Cobitis linea, previously known only from badly damaged type specimens, while I found none!). Repeated visits to areas already sampled may prove rewarding.

It is essential that a collector obtain the necessary licences for scientific purposes. The Department of the Environment, Tehran, issues licences for set periods and areas. There are about 70 National Parks, National Nature Monuments, Wildlife Refuges and Protected Areas or Regions where special licences to collect in these biotic reserves are only issued if there is no threat to endangered species. The Chalus River, Sardab River, Karaj River, Jajrud, Lar-Haraz River and all marshes, wetlands, waterways, deltas and bays along the Caspian Sea coast, and all rivers of Gilan and Mazandaran provinces that enter those waters are protected rivers and wetlands. Penalties for unlawful fishing range up to a year in prison and fines of 50,000 rials. Caspian salmon, cave fishes and trout are protected specifically and by additional fines (Anonymous, 1977-1978).

 


Preserving Fishes

Captured fishes which cannot be identified or seem unusual enough to warrant further attention should be preserved. Labeled, preserved specimens deposited in a museum are a permanent record of species identity and distribution. Some taxa present problems of identification even for experts so that misidentifications are often a nuisance if there is no material to examine. There is a developing aquarium industry in Iran that imports fishes from Singapore and Malaysia. There is a potential for exotics to become established as there are no controls or statistics are not kept (Tehran Times, 28 July 2001). Various exotics are now established through the aquaculture industry (Coad and Abdoli, 1993b). Samples from ecological or experimental studies as well as systematic and distributional works may be preserved and sent to a museum where their identity can be confirmed and where they are available to workers in the future. The National Museum of Natural History, Tehran (Muze-ye Melli-ye Tarikh-e Tabi'i or MMTT) has a small collection of Iranian fishes but it is not extensive enough for systematic studies. Major museums in a number of countries welcome exotic material to enhance the variety of their collections. Acronyms for museum collections can be found in Sabaj Pérez (2010).

Specimens should be preserved whole, without removal of the guts or gills so that no key characters are lost. Specimens may be frozen, or even salted, but the best method and the one used by scientists is to drop fish into 1 part full-strength formalin to kill the fish quickly and then immediately add 9 parts of water to form a 10% preserving solution. Large specimens (larger than about 15 cm) should have a small slit made in the right side of the belly to allow formalin to penetrate the tissues. Ichthyologists cut the right side of the fish and leave the left side undamaged for illustration and scale counting. Hypodermic syringes are used to inject the abdominal cavity and muscle blocks of very large fish with formalin, otherwise the preservative will not penetrate all the tissues before decay sets in. This is especially important in a hot climate like that of Iran. Syringes should have a capacity of up to 100 ml and be capable of taking needles of various sizes. Particular care should be taken when injecting formalin into tissues; the needle should be withdrawn gradually while injecting the formalin solution to avoid a sudden spurt of liquid under pressure from the injection site.

Wherever possible some specimens should be preserved in 95+% ethanol or other appropriate solutions for potential molecular studies. Modern DNA techniques may be the only way to resolve some systematic problems as morphology has proved inadequate.

Formalin should be handled with care as it is a noxious chemical which irritates the eyes and nose and is painful in skin cuts. It may be carcinogenic and repeated exposure can trigger allergic reactions in the skin. Gloves and safety glasses are useful when diluting full-strength formalin. It should only be handled in well-ventilated rooms or in the open air. In the field, care should be exercised in packing specimens for transport so that leakages do not occur. Long-term preservation in formalin is not advisable as the solution becomes acidic and rots the fish. It also wrinkles and hardens the specimens.

Most museums store their specimens in alcohol for the long term. The formalin-fixed specimens are washed briefly in water and then transferred to 45% iso-propyl alcohol or 70% ethanol. These chemicals are pleasanter to work with. Some care should be taken such that specimens are not twisted and bent inside the preserving container. It is difficult to make counts and measurements necessary for identification on badly deformed specimens. Each specimen or group of specimens should have at least an equal volume of preservative as water in the fish tissues tends to dilute the preserving fluid. Specimens may be stepped through 30%, 50% and 70% alcohol solutions to reduce wrinkling and ensure a fuller penetration of alcohol into tissues and a final storage solution of at least 70% ethanol. Ethanol may be difficult to obtain in Islamic countries and undrinkable iso-propyl alcohol can be substituted.

The best containers for long-term storage are made of glass with tightly-sealing polypropylene lids. Plastic containers deteriorate with time and tend to crack. Metal containers and metal lids eventually rust. In the field, large plastic buckets with tightly-sealed lids are less likely to break than glass containers and are not as heavy. Very large fish may require sone sort of drum, such as a clean oil drum but it should be noted that formalin corrodes metal and the drums should be lined with plastic or lacquered. Fluid levels in the collection should be checked regularly and alcohol concentrations maintained at the recommended values or the specimens will deteriorate. Collections should be kept in the dark to reduce fading of pigments and at a constant, cool temperature.

Fish which have been preserved for a week in formalin, more for larger fishes, or transferred to alcohol can be sent to a museum for identification. Glass containers full of formalin or alcohol should not be mailed because of the danger of breakage. The fish should be wrapped in cheesecloth or some other absorbent packaging, with its label, the cheesecloth dampened with preservative, and tightly sealed in several, leak-proof plastic bags before being placed in a padded box for mailing. Spiny fish should be especially well wrapped to avoid puncturing the plastic bags. A tightly-sealed package retains the preservative which keeps the fish in good condition. The box may be labelled "Scientific specimens, no commercial value".

The label is as important as the fish itself. An interesting specimen is of little or no scientific value if there is no locality data. Labels should be written at the time of capture. Faulty memory and good intentions to label specimens later make a poor combination and often result in collections with no data, or worse with incorrect data. The label should bear the place of capture, such as a stream, lake, spring, qanat, etc., including a reference to the nearest town (local names may not be on maps or in gazetteers and some village names are very common, e.g. Hoseynabad, of which there are over 170 in Iran!), latitude and longitude, province, date, name of collector, notes on the habitat and live colour of the specimens, and any other items likely to be useful. Colour photographs of fresh fish are most useful, especially if the fins are pinned erect. Pencil or India ink should be used on stout, waterproof paper which will not disintegrate in liquid. The label must be dropped in the jar with the fish. Labels on the outside of jars always fall off and lids with labels always get put on the wrong jar!

In fact the amount of information which should be usefully recorded cannot be put on a small label. Instead extensive field sheets are used and related to the specimen or sample by a field number. The Canadian Museum of Nature, Ottawa (formerly National Museum of Natural Sciences) has field sheets with over 70 categories which can, potentially, be filled in and some categories have as many as 30 alternatives, e.g. Category 17, Environment includes fresh spring, cave, canal, stream/river, river-lake junction, flooded area, fresh pool, pond, lake, marsh (treeless), swamp (with trees), reservoir, ditch, etc. (see below). As an insurance against loss of field sheets or confusion of numbers, the jar label should carry minimal locality data as well as the field number.

   
 

 


Glossaries

1. Geographical Glossary

The following glossary of geographical and other terms is mostly in Farsi (which includes words taken from Turkic, Kurdish and Arabic) with a few Russian words, all of which appear on maps, in the literature and in the text of the "Freshwater Fishes of Iran" web site. They may be of help to those unfamiliar with these languages and avoid tautologies such as Safidrud River.

There are various diacritical marking systems for Farsi but these do not always transfer accurately across platforms, appearing as strange symbols or gaps in text. I have eschewed the use of these as being irrelevant for native speakers of Farsi and too confusing for those unfamiliar with this language.

ab = water, intermittent stream, stream, spring, lake, well

ab-bandan = shallow, freshwater reservoir on the Caspian plain used for duck hunting in winter and water storage for irrigation in the dry summer

abad = a suffix indicating an inhabited place

ab-e garm = hot spring

ab ambar = cistern

ab anbar = cistern

abshur rud = salt river, common name of salty rivers

anbar = tank

ateshkade = fire-temple (archaeological feature)

av = stream

`ayn = spring

bagh = garden

bahr = sea

Bahr-e Khazar = Caspian Sea

baksh= municipality

band = dam, reservoir, lake, mountain range (old dams for water storage - see sadd for modern dams)

bandar = port, harbour, anchorage, bay

bankari = constructing temporary weirs for water diversion

bar andaz = halting place

barm = marsh, lake, pond

batlaq = marsh, swamp

berkeh = tank, pool, cistern

biaban = desert (also the name of the coastal plain south of the Minab River to the cape of Ra's al-Kuh in Hormozgan Province)

birkat = pool, well, marsh, lake

borj = fort, tower

botlaq = marsh, swamp

caviar = sturgeon eggs as food

çay = stream

centner = 100 kg (used in Russian texts as a measure of commercial catches; sometimes given as 50 kg elsewhere but internal evidence in Russian papers indicates 100 kg is correct)

chah = well, spring, cistern

chai = stream

cham = stream, gorge

chashmeh = spring, well

chay = stream

cheng = hill, mountain, promontory

cheshmeh = spring, well

dag = mountain

dagh = mountain

dahaneh = section of a stream, gorge, pass, defile, water gap

damagheh = cape, promontory

damgah = an artificial, freshwater wetland, maintained primarily as a duck-hunting area but also used for irrigation during the dry summer months

daqq = salt flat, salt depression, salt waste, marsh, intermittent salt lake

darband = gorge or pass

darreh = stream, valley, gorge, ravine

darya = sea, stream, intermittent stream, channel

daryacheh = lake, marshy lake, stream

Darya-ye Mazandaran = Caspian Sea

dasht = plain, desert, steppe, depression, upland, open country, field; usually dry desert with a firm base of pebbles or silts

deh = village

dehkadeh = village

dez = fortress

echbel = eggs of fishes other than sturgeons as food

emamzadeh = tomb, shrine

eskeleh = jetty

estakhr = pool

gadik, gaduk = pass

galal = stream

gardan, gardaneh = pass

godar = pass

garmsir = hot country, winter quarters in the lowlands

ghadamgah = a religious site; usually no fishing allowed.

gharb(i) = west(ern), as in the province Azarbayjan-e Gharbi)

godar = pass

göl = lake, marsh, swamp

gölü = lake, marsh, swamp

gowd = depression

hammam = bath

hamun = marshy lake, salt waste

hawr = marsh, lake

hesar = fort, castle

hor = marsh

howr = marsh

howz = tank, cistern, pond, pool, lake, reservoir, spring

il'men = a shallow, flood-plain lake heavily overgrown with reeds and rushes (Russian)

ishan = hill

istgah = railway station

jabal = hill, mountain

jangal = forest

jar = stream

jazirat = island

jazireh = island

jebal = hill, mountain

jehil = lake

jolgeh = plain

jonub(i) = south(ern)

ju = stream, irrigation channel

jub, jube = irrigation channel, watercourse, gutter, ditch

juy = stream, watercourse

kal = stream, intermittent stream

kalleh = peak

kamar = hill, mountain, ridge

kani = well

karavansara = caravansary

karez = underground irrigation channel

kaur = stream

kavir = salt waste, salt desert, marsh; usually a salt crust over silt deposits which can be fatal mires of slimy mud (= playa)

khalij = bay, gulf

Khalij-e Fars = Persian Gulf

kharabeh = ruins

khirr = stream

khowr = inlet, stream, channel, bay, bight, tidal creek, estuary

khwar = stream

kizil ala = brown trout (see also qezelala)

kotal = mountain pass

kowr = stream

kuh = mountain, range, hill, peak, ridge, spur

kuhha = mountains, range, hills

liman = a brackish bay of the sea, usually at a river mouth, sometimes cut off from the sea but still brackish; also an estuary (Russian, mordab in Farsi)

lut = desert

mahur = hill

mandah = stream

markazi = central, as in Markazi Province

masjed = mosque

mordab = lagoon, backwater, creek, swamp, stagnant water (literally dead water, now replaced by talab)

nahr = river, stream, canal, docking basin

naizar = reed swamp (Sistan)

namak = salt; usually a salt lake with open water or a salt crust but without much mud

namaksar = salt waste

naveh = stream

nawah = stream

nehri = stream

neizar = reed swamp in Sistan

ostan = province, governorate-general

ozero = lake

pal = hill, mountain

paskuh = mountain range

pereval = pass

poshteh = hill, mountain

qabr = tomb

qabrestan = cemetery

qal'at = fort

qal'eh = fortress

qanat = underground irrigation channel; an adit shaft

qasr = fort

qolleh = hill, mountain, peak

ramlat = sandy area

ra's = cape, point, promontory

reka = river

reshteh = mountain, mountain range, hill, spur

reshteh kuh = mountain range

rig = sand area, dunes

riz ab = stream

roga = outflow (in the Enzeli or Anzali Mordab)

rud = river, stream, intermittent stream

rudbar = valley drained by a river with flowing water; place watered by many streams

rudkhaneh = river, river bed, watercourse, intermittent stream

rusta = village, inhabited place

sabkhat = salt marsh, lake

sadd = dam, reservoir (used for modern dams)

saddi qanat = a qanat drawing water from a dam

sahel = coast, beach, shore

sar = cape

sarab = spring (in western Iran), literally "beginning water"

saray = caravansaray

sardsir = cold country, summer quarters in the highlands

sarhadd = frontier

sazhen = a marine sazhen equals 1.83 m (Russian)

selseleh = mountain range, mountains

shahr = town, city

shahrdari = municipality

shahrestan = district

shahzadeh = shrine

shamal(i) = north(ern)

sharq(i) = east(ern), as in the province Azarbayjan-e Sharqi

shatt = large river, bank of a river, stream

saydgah = fishing station, as along the Caspian coast

shebh-e jazireh = peninsula

shekasteh = hill, mountain

shil = a wooden barrier erected across a river for catching fish; hence shilat (in Gilaki, the Persian dialect of Gilan)

shilat = fisheries company; Sherkat Shilat = Northern Fisheries Company concerned with the Caspian Sea; Shilat Jonub = Southern Fisheries Company concerned with the Persian Gulf

shur = salt, brackish, stream

shurab = salt water

shurehzar = salt stream, salt marsh

su = water, stream

suyu = stream

talab = more modern version of mordab.

tall = hill, mountain, spur

tang = pass

tangeh = valley

tappeh = hill, mountain, mound

tell = hill

tepe = hill, mound

vareh = a small dam

vilayet = province

ziarat = shrine

2. Ichthyological Glossary

Technical terms used in ichthyology and in this work are in the Dictionary of Ichthyology and can also be accessed through the Main Page of www.briancoad.com.
 


Quotes

A weak fisherman caught a strong fish in his net and not being able to retain it the fish overcame him and pulled the net from his hand. A boy went to bring water from the torrent. The torrent came and took the boy away. The net brought every time a fish. This time the fish went and carried off the net. The other fishermen were sorry and blamed him for not being able to retain such a fish which had fallen into his net. He replied: 'O brothers, what can be done? My day was not lucky but the fish had yet one remaining'. Moral: A fisherman cannot catch a fish in the Tigris without a day of luck and a fish cannot die on dry ground without the decree of fate. ---- Story 24 from the Gulistan of Sheikh Muslih-uddin Sa'di Shirazi, 1258.

The Caspian sea is marueilous full of fish, but no kind of monstrous fish, as farre as I could vnderstand, yet hath it sundry sortes of fishes which are not in these parts of the world. ---- Principal Navigations, Voyages, Traffiques and Discoveries of the English Nation, Richard Hakluyt, 1599.

Fresh-water Fish is not so plentiful, because there are but few Rivers in Persia, and they take abundance of Water out of them, so that very little Fish can breed there..... There are three sorts of Fresh-water Fish in that large Empire; that of Lakes, that of Rivers, and that of Kerises, or Subterraneous Canals. ---- Sir John Chardin, 1724.

No sea, perhaps, in the world, produces so great a quantity of fish. ---- Said of the Caspian Sea by J. M. Kinneir, 1813.

Thus a man told me that the Caspian Sea, (on the shore of which we conversed) was a Maaden-i-mahi or mine of fish. ---- Sir William Ouseley, 1819.

I may remark as a curious fact in zoology, that many of the cannauts, both here and at Shahrood, swarmed with fish, some of which were of considerable size. ---- James B. Fraser, 1825.

Of fish, in a country which possesses so few rivers, we are not to look for either abundance or variety; nor do the inhabitants make any great use of what they have." ---- James B. Fraser, 1834.

"If you trip over a pebble on the ground, you can be sure that an Englishman put it there" (Persian saying) - as an Englishman I hope there are not too many pebbles in this work, and those that are were inadvertent.

Keys

Introduction

The freshwater fishes of Iran can be identified using these keys, aided by the Species Accounts. The keys should not be used for countries bordering Iran which share many species but also have others not found in Iran. All keys benefit from use and feed-back - please let me know if you encounter problems.

There are two sets of keys in this work. There is a general key to families (although all families are recognisable at a glance in Iran, with a little experience) and a series of keys to genera and species. Genera keys and species keys may be separate if there are many genera and species, or combined in a single key if there are few.

Identification keys are based on couplets, a choice between two alternatives, e.g.

1a. Mouth a large crescent; gill membranes joined to form a free fold over the isthmus = Huso huso
1b. Mouth small and transverse; gill membranes attached to isthmus---> 2

2a. Lower lip continuous, not split in middle; more than 48 lateral scutes, usually 55 or more; barbels fimbriate = Acipenser nudiventris
2b. Lower lip interrupted in the middle; less than 51 lateral scutes, usually less than 45 in Iran; barbels not fimbriate---> 3

If the fish has a large and crescentic mouth, then it is the fil mahi, Huso huso; if not, then the user is directed to the next couplet (2), and so on.

Ideally each couplet has a series of characters which reinforce each other and allow for any loss or damage to characters. Additionally, some characters are "key" but difficult to interpret without experience or are internal and require dissection which is not always possible. In some cases, only one character is available since it must encompass all included species below that point in the key. Since some species are difficult to identify, additional characters are given in brackets [.....]. These additional characters are not unique to the species but, in combination, help to identify the species. Definitions of characters are given in the Dictionary of Ichthyology.

If used properly, a key is more accurate and less time consuming than flicking through pages of text. The disadvantage of keys is that the alternative state in each couplet is not at hand if you only have one fish to examine, and a simple error can lead you widely astray. Some recognised species have overlapping counts for obvious meristic characters, although means and modes are significantly different, and differ in other, subtler ways not readily summarised in a key. Ideally a student of fishes should collect a series of individuals of different sizes and sexes from each locality, wherever conservation demands and practicality permits. A series of about 30-40 specimens allows for character variations dependent on sex and size, and on abnormalities, and also allows for comparative measurements and counts to be made. And more careful examination may reveal more than one species in the sample.

Distribution is often an important aid in assigning samples to a species. Readers should be aware however that fish farming in Iran has led to the translocation, either deliberately or by accident, of species into basins where they are not native. Distribution can appear as a key character when the species is found in basins exclusive of related species. In brackets [.....], distribution is not exclusive but can be an additional character as outlined above.

The most important characters for identification are the general body shape, the number, position and size of the fins, the position and size of the mouth, whether teeth are obvious or not, the number of scales along the flank and the number of rays in various fins, among others. Although colour is often a useful guide, it can also be misleading. Fish vary their colour to match their background or for spawning rituals. In general, it is best to use several characters to identify a fish rather than relying on a single one which can easily be misleading.

Large fishes can be examined for these characters using the naked eye, but various pieces of equipment are necessary for identification of smaller species or juveniles. Hand lenses are of some use in magnifying small characters but by far the best instrument is a binocular microscope which can magnify up to 50 times. Pharyngeal teeth, fin rays and scales can be counted with ease using a microscope. Attachments can be used to take photographs or project images of structures for drawing. Measurements can be taken under a microscope on small specimens to ensure accuracy, and a microscope leaves both hands free to handle the specimen and dissecting tools or calipers. Ichthyologists develop their own techniques for manipulating light sources and specimens for making structures readily visible. I prefer to have two light sources. One of these illuminates the surface of the fish for scale counts and observation of structures. The other bounces light off a white enamel tray into the microscope and is particularly useful for counting fin rays as the light travels through the fin enabling clear distinction of rays.

Two types of forceps are very useful. A large pair (25-35 cm long) enables specimens to be taken out of a jar and sorted without immersing one's fingers. Preservative solutions will irritate the skin and contact should be minimised; some ichthyologists wash the specimen in water before handling, but this may compromise subsequent effectiveness of preservatives. Fine plastic gloves can be worn, but some people develop allergies to latex. A very fine pair of forceps with needle-like points is used to spread folded fins to see the rays and to probe and examine other structures.

Scissors are necessary for slitting the belly and these will vary in size depending on the size of the fish. Fine scissors can be useful in dissection. Very large fish may require a sharp knife or scalpel for dissection or slitting the abdomen. The slit is usually made on the right side of the fish as the left side (head to left) is used for drawings and photographs.

A needle mounted on a wooden or metal handle can be used for cleaning gill arches of debris, clearing flesh from pharyngeal arches or lifting the edges of scales to help in counting them. Most commercial dissecting needles are too blunt and a fine needle can be taped on the end.

Measurements are best made with calipers for accuracy. Dial or electronic calipers are available which measure to an accuracy of 0.1 mm, and are available in several lengths. Very large calipers are usually vernier calipers, but an accuracy of 0.1 mm for large specimens is not required, or even attainable.

Examination of minute scales, debris encrusted gill arches or the lateral and cephalic line canals is facilitated by using compressed air delivered through a glass tube of 1 mm diameter. The air can come from a compressor or aquarium air pump, or even from a hand-squeezed bulb.
 


Key to Families

A little experience will soon make this key to families redundant as all Iranian freshwater fish families can be recognised at a glance. Separate keys are given for families with two or more species (Genera and Species below). Only species which I have seen in Iran, examined museum material of, or have reliable literature records for, are included. The survival of breeding populations of some exotic fishes is uncertain; nonetheless these species are included here.

Drainage basins are given for families with a limited distribution; others are widespread, occurring in all or most basins. "Marine" is used here for drainages entering the Persian Gulf and Sea of Oman including the basins of the Tigris River, Gulf, Hormuz and Makran. The families under this heading are marine but have species that regularly enter fresh water in Iran. The terms Tigris River, Gulf, Hormuz and Makran are restricted here for freshwater residents.

Key characters, e.g. fin ray counts, are restricted to the Iranian species and family members from elsewhere may not key out here.

The families Adrianichthyidae, Mullidae, Percichthyidae and Scophthalmidae have no confirmed records from Iran and are not included in this key (see figures in Species Accounts for distinctive appearance).

* = exotic species; includes species used in aquaculture which may be widely distributed. Note that some species have both native and exotic populations, e.g. Cyprinus carpio.

** = native and translocated. This latter category is liable to change over time as native species are inadvertently or deliberately moved around Iran.

Only families which key out to a single species, or whose included species are all exotics, are marked * or ** here - more speciose families may have both native and exotic components.

Images are of the species mentioned at that point in the key. Where the image is labelled "e.g." then this is a representative species or character for that key couplet.

 

---------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------delete these because of wraparoud or just have short one

--------------------------------------------------50

----------------------------------------------------------------------------------------------------100

 

check books in library for layouts with figures - note wraparound problem

  

1a. Seven lateral gill openings on each side; mouth a sucking disc; no paired (pectoral or pelvic) fins = Petromyzontidae (Caspiomyzon wagneri - Caspian Sea basin)
1b. Less than seven gill openings (1 or 5) on each side; mouth normal; at least pectoral fins present, usually pelvic fins also ---> 2



2a. Five lateral gill slits on each side; scales placoid (small and prickle-like) = Carcharhinidae (Carcharhinus leucas - Marine (Tigris River basin))
2b. One gill opening on each side; scales, when present, cycloid, ctenoid or bony scutes ---> 3

 

  

3a. Body covered with five rows of bony scutes; mouth inferior, behind long snout, with four barbels in front of mouth = Acipenseridae (Caspian Sea basin)
3b. Body without scutes; barbels, if present, not immediately in front of mouth on a long snout ---> 4

4a. Chin with a single barbel at mid-point [no fin spines, 58 or more anal and second dorsal fin rays] = Lotidae (Lota lota - Caspian Sea basin)



4b. Chin without a barbel;  ---> 5

----------------------------------------------------------------------------------

5a. Pelvic fins united to form a disc or funnel = Gobiidae (Caspian Sea, Tedzhen River and Marine basins) ?pelvic disc pic
5b. Pelvic fins present or absent but not formed into a disc ---> 6 

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6a. Pelvic fins absent; body very elongate ("eel-like") ---> 7
6b. Pelvic fins present; body not very elongate ---> 9

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7a. First dorsal fin comprising 30-35 short, sharp spines; flexible snout tip = Mastacembelidae (Mastacembelus mastacembelus - Gulf, Kor River and Tigris River basins)

  

7b. Spines lacking in dorsal fin; snout not flexible ---> 8   

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8a. Body extremely thin, bounded by bony rings; dorsal fin short; snout tube-like with minute mouth = Syngnathidae (Syngnathus abaster - Caspian Sea basin)


8b. Body robust, covered with minute scales; dorsal fin long; snout not tube-like and mouth large = *Anguillidae (Anguilla anguilla) - Caspian Sea basin)

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9a. First dorsal fin with either 3 or 8-12 isolated spines = Gasterosteidae (Caspian Sea basin; introduced)

e.g. Pungitius platygaster


9b. First dorsal fin not composed of isolated spines, spines when present connected by a membrane ---> 10

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10a. Nostrils each with a single pore; lateral line in two parts, the  posterior one lower = Cichlidae (Iranocichla hormuzensis - Hormuz basin)


10b. Nostrils each with two pores; lateral line a continuous line or absent ---> 11

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11a. Eyes on same side of body; body compressed with left side lying on bottom = *Pleuronectidae (Platichthys flesus - Caspian Sea basin)


11b. Eyes on opposite sides of body ---> 12

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12a. Blunt grinding teeth in jaws = Sparidae (Acanthopgarus latus - Marine basins)


12b.Teeth absent or, if present, sharp ---> 13

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13a. Jaws duck-like with strong teeth; dorsal and anal fins far back on body near tail = **Esocidae Esox lucius - Caspian Sea basin; translocated)


13b. Jaws and fins not as above ---> 14

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14a. Barbels absent ---> 15
14b. Barbels present ---> 18

    
Barbels in Paracobitis smithi (dorsal view) and in Barbus lacerta

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15a. First and second dorsal fins widely separate; scales cycloid ---> 16 ?scales
15b. First and second dorsal fins continuous or close together; scales ctenoid ---> 17

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16a. First dorsal fin spines 5 or more (usually 8 or more) and flexible; anal fin spines weak, 1-2 = Atherinidae (Atherina boyeri - Caspian Sea basin)


16b. First dorsal fin spines 4 and very strong; anal fin spines strong, 2-4 (usually 3) = Mugilidae (Caspian Sea and Marine basins)

e.g. Liza abu

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17a. Anal fin spines three or more; first dorsal fin spines rarely 11, usually 10 = *Centrarchidae (Namak Lake basin)

e.g. Lepomis macrochirus

17b. Anal fin spines one or two; first dorsal fin spines 13 or more = Percidae (Caspian Sea basin)

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18a. Three or more pairs of barbels present; no scales or scales minute ---> 19
18b. Barbels two pairs, one pair, or absent; scales present and well developed ---> 24

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19a. Four pairs of barbels present; nasal barbels present = *Heteropneustidae (Heteropneustes fossilis - Tigris River basin)


19b. Three pairs of barbels present; no nasal barbels ? check this--->  20

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20a. A thoracic adhesive apparatus ("sucker") present on the belly between the pectoral fins formed from longitudinal skin folds = Sisoridae (Gulf and Tigris River basins)

Sucker in Glyptothorax silviae

20b. No sucker ---> 21

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21a. Barbels longer than head; no scales; strong pectoral fin spine ---> 22
21b. Barbels shorter than head; scales minute or absent; no pectoral fin spine ---> 23

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22a. Dorsal fin spineless, small and short (3-4 rays) and spineless; anal fin elongate (> 69 rays) = Siluridae (Caspian Sea, Lake Orumiyeh and Tigris River basins)


22b. Dorsal fin with a strong spine, well-developed (7-8 rays); anal fin shorter (6-10 rays) = Bagridae (Mystus pelusius - Gulf, Hormuz and Tigris River basins)

 

---------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------

23a. Spine below eye folding into a groove; head compressed not rounded = Cobitidae (widespread)

e.g. Cobitis taenia   

Cobitis taenia suborbital spine (enlarged) ?just top pic


23b. No spine below eye; head rounded = Nemacheilidae (widespread)

e.g. Oxynoemacheilus kermanshahensis

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24a. Discrete, short adipose fin present = Salmonidae (Caspian Sea, Lake Orumiyeh and Namak Lake basins; widely introduced)

e.g. Salmo caspius
24b. No adipose fin ---> 25

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25a. Dorsal and anal fins long, dorsal with more than 30 rays; head snake-like ---> Channidae (Channa gachua - Hamun-e Jaz Murian basin)


25b. Dorsal and anal fins short, less than 20 rays; head normal ---> 26

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26a. No teeth in jaws; lateral line usually obvious ---> 27
26b. Teeth in jaws; no lateral line pores ---> 28

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27a. Adipose eyelid present; branchiostegal rays 4; alar scales on caudal fin (enlarged scales) = Chanidae (Chanos chanos - Hormuz basin)


27b. Adipose eyelid absent; branchiostegal rays 3; alar scales absent = Cyprinidae (widespread)

e.g. Alburnus filippii

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28a. Head naked dorsally; pelvic fins under dorsal fin = Clupeidae (Caspian Sea and Marine basins)

e.g. Alosa braschnikowii

28b. Head covered with scales dorsally; pelvic fin bases not under dorsal fin ---> 29

---------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------

29a. Teeth conical; anal fin in males enlarged as a copulatory organ; females without sheath around anterior anal fin rays; body slender = *Poeciliidae (widespread)

e.g. Gambusia holbrooki female

                                              Gambusia holbrooki male anal fin

29b. Teeth trifid; anal fin in males normal; females with a sheath around anterior anal fin rays; body robust = Cyprinodontidae (widespread)

e.g. Aphanius vladykovi
 


Keys to Genera and Species

The following keys identify species in the more speciose families and genera. Some keys identify both genera and species, others have separate keys for each genus. Some species are similar and have overlapping characters; distribution is then an additional guide to identity. It should be noted that some species are known to have been introduced to basins outside their natural range, and the possibility exists that other species may be translocated.

Keys are arranged alphabetically by family and by genera within families. * = exotic species and includes species used in aquaculture which may be widely distributed. Note that some species have both native and exotic populations, e.g. Cyprinus carpio, or are native and translocated and marked as **. This latter category is liable to change over time as native species are inadvertently translocated.

--|----|----|--

Key to the Genera and Species of Acipenseridae

Sturgeons are restricted to the Caspian Sea basin and, although aquaculture in internal desert areas has been tried, are unlikely to found as established translocations.

Ventral view of heads of Huso huso, Acipenser nudiventris, A. gueldenstaedtii and A. stellatus. Note that A. persicus is very similar to A. gueldenstaedtii.

1a. Mouth a large crescent; gill membranes joined to form a free fold over the isthmus = Huso huso
1b. Mouth small and transverse; gill membranes attached to isthmus ---> 2

2a. Lower lip continuous, not split in middle; more than 48 lateral scutes, usually 55 or more; barbels fimbriate = Acipenser nudiventris
2b. Lower lip interrupted in the middle; less than 51 lateral scutes (usually less than 45 in Iranian waters); barbels not fimbriate ---> 3

3a. Snout long and narrow (more than 61% of head length); barbels closer to mouth than tip of snout = Acipenser stellatus
3b. Snout shorter and broader (less than 60% of head length); barbels nearer to tip of snout than mouth ---> 4

4a. Back golden-brown, belly yellowish-white; average body depth 12-14% of total length = Acipenser gueldenstaedtii
4b. Back greyish-blue, belly white; average body depth 16.8% of total length = Acipenser persicus

--|----|----|--

Key to the Genera and Species of *Centrarchidae

These two exotic species may not have reproducing populations in Iran. 

1a. Lateral line scales large, 35-50; body compressed in cross section and deep, maximum body depth 1.7-3.0 (usually 2.5 or less) times in body length from snout tip to end of scales [Namak Lake basin] = *Lepomis macrochirus


1b. Lateral line scales small, 58-81; body rounded in cross section and elongate, maximum body depth 2.5-5.0 (usually 3.0 or more) times in body length from snout tip to end of scales [Namak Lake and Tigris River basins] =  *Micropterus salmoides

--|----|----|--

Key to the Genera and Species of Clupeidae

Caspian Sea species have numerous nominal subspecies and keys to these may be found in Berg (1948-1949) and Svetovidov (1952).

1a. Upper jaw without a median notch, rounded when viewed from in front; last two anal fin rays enlarged; lower jaw articulation with skull below or anterior to posterior eye margin; Caspian Sea species ---> 2
1b. Upper jaw with a median notch; last two anal fin rays not enlarged; lower jaw articulation with skull behind posterior eye margin ---> 4

2a. Pectoral fins pointed at tips; head short and wide (interorbital width 16% or more of head length) ---> 3
2b. Pectoral fins rounded at tips; head large and narrow (interorbital width 15.5% or less of head length) = Clupeonella grimmi

3a. Body and belly compressed (body depth about 21-27% of standard length); keeled belly scales evident = Clupeonella caspia


3b. Body cylindrical and belly rounded (body depth 16-19% of standard length); keeled belly scales weakly developed = Clupeonella engrauliformis

4a. Branched pelvic fin rays 8; upper gill rakers overlap lower gill rakers at angle of first arch; Caspian Sea species ---> 5
4b. Branched pelvic fin rays 7; upper gill rakers not overlapping lower gill rakers at angle of first arch; Gulf and Tigris River basins = Tenualosa ilisha

5a. Body deep and compressed; head large and deep, wedge-shaped in anterior view; caudal peduncle short; pectoral fins long ---> 6
5b. Body not deep and not compressed; head not large and deep, not wedge-shaped in anterior view; caudal peduncle not short; pectoral fins short ---> 8

6a. Gill rakers on first arch 60 or more, thin and long, much longer than gill filaments; teeth weakly developed = Alosa caspia


6b. Gill rakers on first arch 45 or less, shorter, equal to or somewhat longer than gill filaments; teeth well developed ---> 7

7a. Upper and lower profiles of head straight; lower jaw protruding and its upper edge straight = Alosa saposchnikowii


7b. Upper and lower profiles of head rounded; jaws equal in length and lower jaw has a crescentic upper edge = Alosa sphaerocephala

8a. Gill rakers 47 or less, thick and coarse = Alosa braschnikowii


8b. Gill rakers 59 or more, may be thin and long but can be coarse and short = Alosa kessleri

--|----|----|--

Key to the Genera of Cobitidae

1a. Caudal fin with 14 branched rays; a row of large and distinct dark spots laterally, speckles above this row tending to form a row too; sides of body not distended in front of dorsal fin in males = Cobitis
1b. Caudal fin with 12, rarely 13, branched rays; speckles above the lateral line not forming horizontal rows; sides of body distended in front of dorsal fin in males = Sabanejewia

Key to the Species of Cobitis

? check spines

1a. Dark brown lateral spots reduced or absent; Hormuz and Kor River basins = Cobitis linea


1b. Large dark and obvious spots along the mid-flank numbering 10-20, usually 16-18; Caspian Sea and Tigris River basins = Cobitis taenia

Key to the Species of Sabanejewia

1a. Row of dark brown spots laterally [branches of suborbital spine differing in length; Caspian Sea and Tedzhen River basins] = Sabanejewia aurata


1b. No row of large dark spots laterally [Caspian Sea basin] ---> 2

2a. A continuous dark streak mid-laterally; branches of suborbital spine differing in length; two dark spots at base of caudal fin = Sabanejewia caspia


2b. Numerous small speckles along flank; suborbital spine strong with branches of similar length; no dark spot at caudal fin base = Sabanejewia caucasica

?pic

--|----|----|--

Key to the Genera and Species of Cyprinidae

?check all genera are in this part of key

?tabulate main charctesr like spine, barbels, sucker?

The cyprinid family is the most speciose in Iranian fresh waters. Members of the family are more easily identified first to genus and then to species. Keys are then shorter and less liable to error in use. Additional characters can be listed under each genus or species which are not unique nor readily incorporated into keys but which in combination help to identify the genus or species. These additional characters are given in brackets. Monotypic genera key out to species in the generic key.

?Add petroleuciscus add Romanogobio, Luciobarbus, Carasobarbus, Kosswigobarbus, Mesoptamichthys, Tor, *Mylopharyngodon

1a. Branchiostegal membranes not attached to isthmus; gill rakers fused together; eyes low on side of head, below midline; suprabranchial organ present = *Hypophthalmichthys spp.


1b. Branchiostegal membranes attached to isthmus; gill rakers not fused; eyes at or above midline of head; suprabranchial organ absent ---> 2

2a. Serrated stiffened ray (spine-like) in the dorsal and anal fins; dorsal fin elongate (? rays or more, usually ?); anal fin origin below dorsal fin ---> 3 see iraq book
2b. No serrated stiffened ray (spine-like) in the anal fin; dorsal fin short to moderately elongate (?-? rays, usually ?); anal fin origin behind dorsal fin end ---> 4

3a. Barbels absent; pharyngeal teeth in one row = *Carassius

pic
3b. Barbels present (two pairs); pharyngeal teeth in three rows; ? and exotic = **Cyprinus carpio

4a. Eyes absent; body pink through lack of pigment; no scales = Iranocypris typhlops
4b. Eyes present; body pigmented; scales present, sometimes restricted to anal area ---> 5

mouth structure *Pseudorasbora parva peculiar short dorsal fin ray *Pimephales promelas other species to pull out here

Aspidoparia morar

Barilius mesopotamicus

*Ctenopharyngodon idella

*Hemiculter leucisculus

Leucaspius delineatus

*Pseudorasbora parva

**Rhodeus sericeus

Scardinius erythrophthalmus

5a. Anus and anal fin base sheathed by markedly enlarged scales ---> 6
5b. Anus and anal fin base not sheathed by markedly enlarged scales ---> 8

6a. Branched anal fin rays 5; scales mostly absent; pharyngeal teeth in two rows =  Schizopygopsis stoliczkai


6b. Branched anal fin rays 6; scales present; pharyngeal teeth in three rows =  

7a. Barbels absent or vestigial; anal fin branched rays 6; pharyngeal tooth formula 2,3,4-4,3,2 = Schizocypris altidorsalis


7b. Barbels present and well-developed; anal fin branched rays 5É see above; pharyngeal tooth formula 2,3,5-5,3,2 = Schizothorax spp.

8a. An adhesive disc prominent on the underside of the head = Garra

Underside of head of Garra persica

8b. No adhesive disc --->

9a. Scaleless keel extending from the throat to the anal fin; lateral line decurved and wavy = Pelecus cultratus
9b. Not as above ---> 10

10a. Barbels present ---> x
10b. Barbels absent ---> 11

 

11.  naked ventral keel Abramis  Alburnoides Alburnus  Blicca  Alburnus  Vimba  (also put alburnoides, alburnus in another couplet)

 

1. Barbels absent + spine in D D ray count [scales large, 29-35 in lateral line; body compressed; Tigris River basin; dorsal fin spine smooth, without denticulations] = Barbus sharpeyi

+ Mesoptamichthys? no barbels

x. Spine in dorsal fin ---> h
    No spine in dorsal fin ---> c

h. Spine smooth; mouth not sector-shaped  ---> j
    Spine with teeth; mouth sector-shaped (u-shaped in young) ---> i

j. Mouth with central tubercles Kosswigobarbus
Mouth without k

k Branched anal fin rays 6, dorsal fin branched rays 10 or more; lateral line scales ? = Cyprinion
   Branched anal fin rays 5; dorsal fin branched rays 9 or less; lateral line scales ? = Capoeta

 

j  = Carasobarbus luteus
= Tor grypus

c. Scales small, more than ?100 in lateral line = Tinca tinca
    Scales larger, less than ? in lateral line ---> q

q. Anal fin branched rays 13 or more = Barilius mesopotamicus
   
Anal fin branched rays less than 13 ---> t

t. Dorsal fin branched rays 7; pharyngeal teeth in two rows = Gobio gobio
    Dorsal fin branched rays 8; pharyngeal teeth in one row ---> s

s. Lateral line complete = Crossocheilus latius
    Lateral line incomplete = Hemigrammocapoeta elegans

  Spine no barbel Acanthobrama marmid

  chck for more see above and Esmaeili list

 

Body and caudal peduncle compressed (caudal peduncle depth at anal fin insertion greater than caudal peduncle width); well-defined spots on the dorsal and caudal fins; Tedzhen River = Gobio gobio
Body only slightly compressed and the caudal peduncle cylindrical (caudal peduncle depth at anal fin insertion less than or about equal to caudal peduncle width); faint spots on the dorsal and caudal fins; Lake Orumiyeh and Caspian Sea = Gobio persus 

 

Key to the Species of Abramis

Both species are found only in the Caspian Sea basin.

1a. Branched anal fin rays 22-30 = Abramis brama


1b. Branched anal fin rays 31-44, mostly 34 or more = Abramis sapa

Key to the Species of Acanthalburnus

1a. Anal fin branched rays 13-19; lateral line scales 60-85; Caspian Sea basin = Acanthalburnus microlepis


1b. Anal fin branched rays 10-13; lateral line scales 50-68; Lake Orumiyeh basin = Acanthalburnus urmianus

 

Key to the Species of Alburnoides

Populations in the Esfahan and Tedzhen River basins are not yet identified to species.

1a. Snout pointed or slightly rounded; mouth terminal or upturned, tip of mouth cleft on level from slightly above middle of eye to upper margin of pupil; lower jaw slightly to moderately projecting relative to upper jaw; junction of lower jaw and quadrate on about vertical through anterior eye margin; Kor River basin = Alburnoides qanati
2b. Snout slightly to markedly rounded; mouth terminal
to subterminal, tip of mouth cleft on level from middle of eye to below lower margin of eye; upper jaw slightly to moderately projecting relative to lower jaw; junction of lower jaw and quadrate on about vertical through about middle of eye ---> 2

2a. Branched anal fin rays 8-11, commonly 9-10; branched dorsal fin rays 7, rarely 8 ---> 3
2b. Branched anal fin rays 10-15, commonly 11-13; branched dorsal fin rays 8, rarely 7 ---> 4

3a. Ventral keel completely scaled; total vertebrae 40-41; abdominal vertebrae 20-22, commonly 21; Lake Orumiyeh basin = Alburnoides petrubanarescui
3b. Ventral keel scaleless from one-third to whole keel length; total vertebrae 38-40, commonly 39; abdominal vertebrae 19-20; Tigris River basin = Alburnoides nicolausi

4a. Ventral keel smoothed, scaled along one-third to whole length; Tigris River basin = Alburnoides idignensis
4b. Ventral keel well-pronounced, almost or completely scaleless ---> 5

6a. Lateral line in alive and preserved fish delineated by dark pigment dots above and below; 13-15 predorsal vertebrae; mouth terminal, tip of mouth cleft on or slightly below middle of eye; Caspian Sea basin = A. eichwaldii
6b. Lateral line in alive and preserved fish somewhat darker than surrounding flank but no strong dark dots outline to canal; 11-13 predorsal vertebrae; mouth almost subterminal, tip of mouth cleft on or below lower margin of eye; Namak Lake basin = Alburnoides namaki

Key to the Species of Alburnus

1a. Dorsal fin branched rays modally 7; strong mid-flank stripe [anal fin branched rays 9-13, usually 10-12; total gill rakers 12-17; lateral line scales 46-64, usually 50-60; Caspian Sea basin] = Alburnus filippii


1b. Dorsal fin branched rays modally 8; no strong stripe in Caspian Sea species ---> 2

2a. Total gill rakers 15-31, usually 19 or more [Caspian Sea basin] ---> 3
2a. Total gill rakers 10-18, usually 16 or less ---> 4

3a. Lateral line scales 54-74, usually 55 or more [anal fin branched rays 12-19; peritoneum light brown; Caspian Sea basin] = Alburnus chalcoides


3b. Lateral line scales 36-53, usually 48 or less [anal fin branched rays 10-21; peritoneum light silvery; Caspian Sea basin and translocated] = **Alburnus hohenackeri

4a. Lake Orumiyeh basin [anal fin branched rays 9-12, usually 10-11; total gill rakers 11-16; lateral line scales 46-63, usually 46-58] = Alburnus atropatenae


4b Tigris River and basins of southern Iran ---> 5

5a. Anal fin branched rays 9-10; upper Tigris River basin near ?; peritoneum silvery [total gill rakers 12-14; lateral line scales 67-83] = Alburnus zagrosensis

pic?
5b. Anal fin branched rays 10-18, usually 11 or more; elsewhere in southern Iran; peritoneum brown to black ---> 6

6a. Lateral line scales 43-58; anal fin branched rays 13-18, usually 14-16 [total gill rakers 10-13; Tigris River basin] = Alburnus caeruleus


6b. Lateral line scales 58-89, usually 60 or more; anal fin branched rays 10-14, usually 11-12 [total gill rakers 11-18; Esfahan, Gulf, Hormuz, Kor River, Lake Maharlu and Tigris River basins] = Alburnus mossulensis

Key to the Species of Aspius

The distinction of these two species has not been examined recently and characters overlap, sample sizes for gill rakers and scales in particular being very small. However, they are found in separate basins.

1a. Lateral line scales 62-105; anal fin branched rays 11-15, usually 12; total gill rakers 8-11; total vertebrae 50-51; Caspian Sea basin = Aspius aspius


1b. Lateral line scales 91-110; anal fin branched rays 10-13, usually 11?; total gill rakers 11-14; total vertebrae 51-53; Tigris River basin = Aspius vorax

  

Key to the Species of Capoeta

1a. Dorsal fin branched rays modally 7 [lateral line scales 42-62; total gill rakers 11-20; Bejestan, Dasht-e Kavir, Dasht-e Lut, Sistan and Tedzhen River basins] = Capoeta fusca


1b. Dorsal fin branched drays modally 8 or 9 ---> 2

2a. Dorsal fin spine strongly developed, longer than head [lateral line scales 68-90; total gill rakers 23-33; Gulf and Tigris River basins] = Capoeta trutta


2b. Dorsal fin spine well-developed to weak, not longer than head ---> 3

3a. Total gill rakers 9-17 [lateral line scales 72-99; dorsal fin spine weak and poorly serrated; Namak Lake basin] = Capoeta buhsei


3b. Total gill rakers 16 or more, usually 18 or more ---> 4

4a. Lateral line scales 36-52, mostly 39-48; [Dasht-e Kavir, Esfahan, Kerman-Na'in, Kor River, Namak Lake and Tigris River basins [total gill rakers 16-25] = Capoeta aculeata


4b. Lateral line scales 46-99; mostly 50 or more ---> 5

5a. Dorsal fin branched rays modally 9; often large black blotches on flank [lateral line scales 60-99; widespread] = Capoeta damascina


5b. Dorsal fin branched rays modally 8, sometimes 9; without black blotches ---> 6

6a. Irregular brown to black speckles on head and flank [lateral line scales 58-82; Gulf and Tigris River] = Capoeta barroisi


6b. Speckles absent; ?  [lateral line scales 46-70; widespread; Tedzhen River fish often with 4 barbels] = Capoeta capoeta


 

Key to the Species of *Carassius

Goldfish have been widely introduced in Iran; presence and distribution of other species is uncertain. ?gibelio

1a. Lateral line scales 25-34, mostly 31 or less; gill rakers 35-54, size dependent and mostly 39 or more in adults; anal fin branched rays modally 5; young never with dark spot on caudal peduncle = *Carassius auratus


1b. Lateral line scales 32-36; gill rakers 23-35, mostly 31 or less; anal fin branched rays modally 6; young usually with dark spot on caudal peduncle = *Carassius carassius

Key to the Species of Chondrostoma

1a. Caspian Sea basin [lateral line scales 50-68] = Chondrostoma cyri


1b. Outside Caspian Sea basin ---> 2

2a. Kor River basin [lateral line scales 49-57; dorsal fin branched rays usually 8] = Chondrostoma orientale

?pic
2b. Tigris River basin [lateral line scales 50-69; dorsal fin branched rays usually 8 or 9] = Chondrostoma regium

 

Key to the Species of Cyprinion ?

 

tabulate characters for comparison see berg?
 

1a. Mouth small with large lateral lobes; cartilage may form a tooth-like structure [dorsal fin branched rays 12-16; total gill rakers 10-15; Gulf and Tigris River basins] = Cyprinion kais

 
 

1b. Mouth without large lateral lobes; cartilage arched and not tooth-like ---> 2

Mouth in Cyprinion macrostomum

2a. Mouth oblique and long in lateral view [dorsal fin branched rays 10-13; total gill rakers 11-12; Hamun-e Jaz Murian, Hormuz and Makran basins] = Cyprinion milesi


2b. Mouth arched in young, transverse in adults ---> 3

3a. Dorsal fin branched rays 9-12, usually 10-11, means 10.0-10.5; southeastern and eastern Iran - Dasht-e Lut, Hamun-e Jaz Murian, Hamun-e Mashkid, Hormuz, Makran and Sistan basins = Cyprinion watsoni

 
3b. Dorsal fin branched rays 11-17, usually 12-15, means 13.1-13.9; southwestern Iran - Gulf, Lake Maharlu and Tigris River basins ---> 4

4a. Dorsal fin spine teeth well-developed, even near spine tip [Gulf and Tigris River basins] = Cyprinion macrostomum


4b. Dorsal fin spine teeth graded in size as near tip and finer [Gulf and Lake Maharlu basins]= Cyprinion tenuiradius

Key to the Species of Garra

?

1a. Caudal fin branched rays modally 16; ? (85.6% for 132 fish, range 15-17) [Hamun-e Jaz Murian, Hormuz and Makran basins] = Garra persica



1b. Caudal fin branched rays modally 17, 16 only rarely ---> 2

2a. Dorsal fin branched rays modally 8 (87.1% for 534 fish, range 6-8);  ? sucker structure [Gulf, Hormuz, Kor River, Lake Maharlu and Tigris River basins] = Garra rufa

rufa sucker pic? from Berg?
2b. Dorsal fin branched rays modally 7 (91.5% for 59 fish, range 6-8) ---> 3

3a. ?; eastern Iran (Bejestan, Hamun-e Jaz Murian, Hamun-e Mashkid, Dasht-e Lut, Makran, Sistan and Tedzhen River basins) = Garra rossica


3b. ?; western Iran (Tigris River basin) = Garra variabilis


Key to the Species of *Hypophthalmichthys

These two species are widely farmed.

1a. Abdomen with a compressed keel extending from the breast (pelvic fins) to the vent; pectoral fins short, not extending past the origin of the pelvic fins; gill rakers a continuous band uniting both sides, roots fused into a spongy mass = *Hypophthalmichthys molitrix


1b. Abdomen with a compressed keel extending from the throat to vent; pectoral fins long, extending past the origin of the pelvic fins; gill rakers free, no spongy mass = *Hypophthalmichthys nobilis

Key to the Species of Kosswigobarbus

1a. Lateral line scales 29-41; total vertebrae 39-40; Tigris River basin = Kosswigobarbus kosswigi


1b. Lateral line scales 24-27; total vertebrae 37-38; A'la River in Khuzestan and the Fahlian River in Fars = Kosswigobarbus sublimus

Key to the Species of Luciobarbus ?

1a. Body covered with large dark spots arranged almost in a quincunx (pattern of five) [Tigris River basin] = Luciobarbus subquincunciatus

ph teeth? her and elsewhere
1b. Body without large spots ---> 2

2a. Head elongate, tapering and depressed anteriorly, pike-like, with postorbital distance in standard length 7.2 or less; adults very large, reputedly over 2 m long [Tigris River and Gulf basins] = Luciobarbus esocinus


2b. Head not as above; not very large, to ? m ---> 3

3a.Northern and northwestern distribution in the Caspian Sea, Lake Orumiyeh and Namak Lake basins ---> 4
3b.Southern and western distribution in the Gulf, Kor River and Tigris River basins ---> 6

4a. Dorsal fin branched rays modally 7 [predorsal length shorter than postdorsal length; lateral line scales 62-90, usually 65-77; total gill rakers 16-25; Caspian Sea basin] = Luciobarbus brachycephalus


4b. Dorsal fin branched rays modally 8 ---> 5

5a. Lateral line scales 51-72; without three lobes to lower lip; upper dark flank clearly delineated from lighter lower flank [predorsal length equal to longer than postdorsal length; total gill rakers 12-19; Caspian Sea basin] = Luciobarbus capito


5b. Lateral line scales 74-103, often 85 or more; lower lip usually with three lobes; body shades from dark to light gradually down flank [?predorsal length; total gill rakers 9-18; Caspian Sea, Lake Orumiyeh and Namak Lake basins] = Luciobarbus mursa

6a. Total gill rakers 7-13 [lateral line scales 57-68; Tigris River basin] = Luciobarbus xanthopterus


6b. Total gill rakers 14 or more ---> 7

7a. Lips markedly fleshy; fourth major row pharyngeal tooth large and molariform [scales?; Gulf, Kor River and Tigris River basins] = Luciobarbus barbulus


7b. Lips not markedly fleshy; ?Fourth major row pharyngeal tooth similar in size to third, not molariform?check [scales?]  ---> 8

8a. Dorsal fin spine strong, arising from an elevated base; dorsal fin origin at or ahead of pelvic fins origin [Gulf, Kor River and Tigris River basins] = Luciobarbus pectoralis

?better pic with stronger spine
8b. Dorsal fin spine present but not markedly strong; dorsal fin origin behind pelvic fins origin [Gulf and Tigris River basins] = Luciobarbus kersin

? pic needed with weaker spine and D further back - see Iraq book for key

Key to the species of Petroleuciscus 

1a. Dorsal fin branched rays 6-7, modally 7; anal fin branched rays 7-9, modally 8; total vertebrae 34-37; pharyngeal teeth 1.5-4.1; Gulf, Hormuz and Kor River  basins = Petroleuciscus persidis


1b. Dorsal fin branched rays 7-9, modally 8 or 9; anal fin branched rays 7-12, modally 9-11; total vertebrae 37-42; pharyngeal teeth usually 2.5-4.2 ---> 2

2a. Anal fin branched rays 9-12, modally 10 or 11; lateral line scales 45-56; total vertebrae 41-42; Esfahan basin = Petroleuciscus esfahani



2b. Anal fin branched rays 7-10, modally 9; lateral line scales 36-45; total vertebrae 37-38; Lake Orumiyeh basin = Petroleuciscus ulanus


 

Key to the Species of Romanogobio

Key by A. Naseka, Zoological Institute, St. Petersburg:- 

1a. Number of lateral line scales 41 to 45 with modes of 42 and 43; total vertebrae 38 to 42 with modes of 40 and 41; connection between the supraorbital and infraorbital head canals usually present; Caspian Sea basin = Romanogobio macropterus
1b. Number of lateral line scales 40 to 42 with modes of 40 or 41; total vertebrae 37 to 40 with modes of 38 and 39; connection between the supraorbital and infraorbital head canals usually absent; Lake Orumiyeh basin = Romanogobio persus


 

Key to the Species of Rutilus

These species occur only in the Caspian Sea basin.

1a. Lateral line scales 47-64, mostly 55-58; swimbladder elongate and conical or pointed posteriorly ---> 2
1b. Lateral line scales 39-48, mostly 42-47; swimbladder rounded posteriorly ---> 3

2a. = Rutilus frisii
2
b. = Rutilus kutum

3a. = Rutilus caspicus
3b. = Rutilus rutilus

Key to the Species of Schizothorax ?

1a. Total gill rakers 24-41[lips thin; Sistan basin] = Schizothorax zarudnyi


1b. Total gill rakers 18 or less ---> 2

2a. lips?; Sistan basin = Schizothorax intermedius


2b. lips thick; Dasht-e Kavir and Tedzhen River basins = Schizothorax pelzami

Key to the Species of Squalius

1a. Flank scales outlined by pigment; anal fin rounded distally [Caspian Sea, Lake Orumiyeh, Namak Lake and Tigris River basins] = Squalius cephalus


1b. Flank scales not outlined by pigment; anal fin truncate or emarginate distally ---> 2

2a. Lower jaw not projecting; Tedzhen River basin = Squalius latus check on fish about lower jaw?


2b. Lower jaw projecting; Tigris River basin = Squalius lepidus

--|----|----|--

Key to the Species of Cyprinodontidae

Almost any sample will contain both males and females, clearly distinguished by colour and pigment patterns.

?isfahanensis and check over key again

1a. Lateral line scales 36-47 [females finely speckled, no lozenge-shaped spot at caudal fin base; Tigris River basin] = Aphanius vladykovi

male  female

1b. Lateral line scales 24-35, mostly 31 or less ---> 2

2a. Total dorsal fin rays 4-7; total anal fin rays 6-10; gut variably coiled [Hormuz basin] = Aphanius ginaonis

male    ?gut pics

2b. Total dorsal fin rays 7-13, usually 9 or more; total anal fin rays 8-13, usually 10 or more; gut regularly coiled ---> 3

3a. Males lemon-yellow with two broad bars on caudal fin; females ? [Gulf, Hamun-e Jaz Murian, Hormuz, Makran and Tigris River basins] = Aphanius dispar


3b. Males not lemon-yellow; ?? ---> 4

4a. Males with blue spots on flank [Tigris River basin] = Aphanius mento

 

male

 

4b. Males without blue spots on flank ---> 5

5a. Females with obvious flank bars [males barred]; Lake Maharlu basin = Aphanius persicus

malefemale
5
b. Females with flank spots ---> 6

6a. ?Females without lozenge-shaped spot at caudal fin base; flank spots large; Tigris River basin = Aphanius mesopotamicus

malefemale

6b. Females with lozenge-shaped spot at caudal fin base; flank spots small; Kor River basin = Aphanius sophiae

malefemale
 

--|----|----|--

Key to the Genera and Species of Gasterosteidae

1a.  Long dorsal fin spines, numbering 3; long pelvic fin spines, dorsal spines exceeding eye diameter in length; scutes (vertical bony plates on flank) large [Caspian Sea, Dasht-e Kavir and Tedzhen River basins] = *Gasterosteus aculeatus


1b.  Short dorsal fin spines, numbering 7-11, alternatively sloping left and right; dorsal spines shorter than eye diameter; scutes small [Caspian Sea basin] = Pungitius platygaster

--|----|----|--

Key to Persian Gulf and Sea of Oman Drainage Species of the Gobiidae

1a. Lateral series scales large, 28-36; eyes not protruding = Glossogobius giuris


1b. Lateral series scales minute, over 90; eyes protruding above dorsal head profile ---> 2

2a. 4-5 first dorsal fin spines; anal fin base and second dorsal fin base 34% or more of standard length; 2 canine teeth internal to the lower jaw symphysis = Boleophthalmus dussumieri


2b.10-14 first dorsal fin spines; anal fin base and second dorsal fin base 27% or less of standard length; no canine teeth internal to the lower jaw symphysis  = Periophthalmus waltoni

--|----|----|--

Key to Caspian Sea Genera and Species of the Gobiidae

? check against checklist for all species included eliminate species not definitely records

Some genera are monotypic or have only a single species in the Caspian Sea basin and so the keys terminates there. Speciose genera have separate keys below. The Iranian shore of the Caspian Sea remains poorly explored in its deeper waters and keys in Miller (2003), Mitrofanov (2003) and Boldyrev and Bogutskaya (2007) should be consulted for specimens which do not key out here (see also Species Accounts for further listings and discussion).

Note that a Rhinogobius species is recorded from the Tedzhen (= Hari) River basin in Iran as an exotic (see Species Accounts for description). This is the only goby outside the Caspian Sea basin and coastal waters of the Persian Gulf and Sea of Oman in Iran.

The following key is modified after Miller in Miller (2003):- ÉRECHECK!!

1a. Suborbital papillae with longitudinal row a immediately below eye and having at least one short side row; cheek with several short transverse rows, none reaching lower eye margin; snout with longitudinal rows s1 and s2 or, if transverse interorbital and snout rows, a perianal organ is present = Knipowitschia

e.g. Knipowitschia iljini

1b. Suborbital papillae in transverse rows; no row a; snout with transverse rows s1 and s2; no perianal organ ---> 2

2a. Row 5i not below level of row 6i; 6i at or opposite end of row d; scales normal; canals present or absent ---> 3
2b. Row 5i below level of row 6i; 6i separated from posterior end of row d by row 5i; scales non-imbricate or bony tubercles and granules or naked; no canals ---> 7

3a. Anterior nostril an elongate tube hanging over lip = Proterorhinus nasalis


3b. Anterior nostril elongate but not overhanging lip ---> 4

4a. Three rows below row b = Mesogobius nonultimus

?pic
4b. Two rows below row b ---> 5

5a. Five rows before row b = Chasar bathybius

?pic

5b. Four rows before row b = 6

6a. = Neogobius
6b. = Ponticola

7a. No chin barbel or cheek flap; snout a duck-bill shape = Anatirostrum profundorum

 
7b. Chin barbel and cheek flap present; snout not a duck-bill shape = Benthophilus

The following key does not use papillae and head canal characters; see figures above. It is modified from Mitrofanov (2003) as translated courtesy of Dmitri Ponomarenko:- RECHECK

1a. Body naked ---> 2
1b. Body covered with regular ctenoid scales ---> 3

2a. Snout narrow and long = Anatirostrum profundorum
2b. Snout regular, not elongated ---> 3

3a. Anterior nostril elongated into a tube that hangs over lip ---> Proterorhinus nasalis
3b. Anterior nostril not as above---> 4

4a. Second dorsal fin short, with less than 12 branched rays; small fishes less than 50 mm ---> 5
4b. Second dorsal long, with more than 12 branched rays ---> 6 

5a. Scales on sides of body, with head, throat, belly and back to second dorsal fin scaleless; eyes lateral; body darkly pigmented without stripes; tail symmetrical without a dark spot at tail base ---> Knipowitschia caucasica
5b. Body fairly fully covered with scales; eyes pointed upwards; body with dark stripes; body glassy and translucent; tail symmetrical without a basal spot; deepwater species ---> Knipowitschia iljini

6a. Sinciput not covered with scales = Mesogobius nonultimus
6b. Sinciput and occiput covered with scales ---> 7

7a. = Neogobius
7b. = Ponticola

Key to the Species Benthophilus

The following key is modified after Pinchuk and Miller in Miller (2004). RECHECK See Boldyrev and Bogut too

1a. One or two dermal barbels behind jaw angle; first dorsal fin with 1-2 spines; tubercles large and high, not all spinous; tubercles in dorsal row 13-15, in ventral row 10-13 = Benthophilus baeri


1b. A dermal fold or lobe behind jaw angle (if absent, tubercles vertically elongated); first dorsal fin spines 3-4 (rarely 2); tubercles in dorsal row 18 or more, in ventral row14 or more ---> 2

2a. Tubercles vertically elongated, curved and crest-like, rear edges spinulose; temporal and occipital region without large tubercles; head narrow, interorbit with median groove between elevated ridges; dermal filaments present or absent; dermal fold behind jaw angle when present narrow, with an acute protuberance; back without brown bands ---> 3
2b. Tubercles conical and tipped by spines ---> 4

3a. Temporal and occipital region of head with granules; tubercles in dorsal row usually 30-33 = Benthophilus ctenolepidus


3b. Temporal region of head naked = Benthophilus pinchuki

?pic

4a. Tubercles distinct, relatively large; granules on temporal and occipital area small and sparse, or if slightly larger then not forming real tubercles; bands present = B. leobergius


4b. Tubercle rows distinct but tubercles relatively small; upper head and body densely covered with very small granules; no dark brown bands = B. macrocephalus

Key to Species of Knipowitschia

The following key is modified after Miller in Miller (2004).

1a. Males with 0-4 flank bars; anterior oculoscapular canals united at posterior interorbit, with a single median pore κ, and canals extending anteriorly to pores λ; preopercular canal present = K. caucasica


1b. Males with 6-10 flank bars; anterior oculoscapular canals more or less separate in midline of posterior interorbit, with pore κ double, and canals extending anteriorly through interorbit of variable extent, typically absent; preopercular canal present or absent = K. iljini

Key to the Species of Neogobius

The following key is modified after Miller and Vasil'eva in Miller (2003).

1a. Posterior nostril markedly distant from edge of orbit; pelvic fin anterior membrane with angular lateral lobes; lobes about one-sixth to almost one-half width of anterior edge of membrane  = N. caspius


1b. Posterior nostril near edge of orbit; pelvic fin anterior membrane with rounded and shallow lateral lobes; lobes not more than one-sixth width of anterior edge of membrane, or lacking entirely ---> 2

2a. At least anterior nape scales cycloid; first dorsal fin with large dark spot at rear; lateral series scales usually 49-55 = Neogobius melanostomus


2
b. Nape scales ctenoid; first dorsal fin without large dark spot; lateral series scales usually 55-70 = Neogobius pallasi

Key to the Species of Ponticola

The following key is modified after Miller and Vasil'eva in Miller (2003). recheck?

1a. Pelvic fin anterior membrane with rounded and shallow lateral lobes; lobes not more than one-sixth width of anterior edge of membrane, or lacking entirely = Ponticola syrman

pic?
1b. Pelvic fin anterior membrane with angular lateral lobes; lobes about one-sixth to almost one-half width of anterior edge of membrane ---> 2

2a. Lateral series scales usually 49-54; lateral lobes of pelvic fin anterior membrane small, not more than one-fifth width of rear edge; upper lip width 0.4-0.67
    lateral preorbital width (lip to orbit); nape scales cycloid; pelvic fin almost reaches the anal fin (0.9 distance) or extends beyond the anal fin
    origin = Ponticola goebelii


2b. Lateral series scales usually 54-76; lateral lobes of pelvic fin anterior membrane large, at least one-fifth width of rear edge; upper lip width at least 0.6
    lateral preorbital width (lip to orbit), if less than 0.75, then nape scales ctenoid; pelvic fin less than nine-tenths distance to anal fin ---> 3

3a. Upper lip not markedly swollen, width about 0.6-0.67 lateral preorbit; interorbital distance 0.8-0.9 eye diameter; caudal peduncle depth 0.67-0.75 length =
    Ponticola gorlap


3b. Upper lip moderately swollen, width about 0.75 to more than length lateral preorbit; interorbital distance 0.4-0.8 eye diameter; caudal peduncle depth  0.75
    to more than length = Ponticola cyrius

--|----|----|--

Key to the Genera and Species of Mugilidae

?other species see Iraq book subviridis and vaigiensis; premaxilla pics

1a. Posterior end of maxilla not curved below tip of premaxilla, but straight; jaw end on line of gape; adipose eyelid well-developed, enclosing eye over much of anterior and posterior fields of iris, so pupil is covered by an oval slit; pyloric caeca 2; Marine, introduced to Caspian Sea = **Mugil cephalus


1b. Posterior end of maxilla curved below tip of premaxilla, visible behind corner of closed mouth; jaw end on below line of gape; adipose eyelid weakly-developed, not reaching pupil of eye; pyloric caeca 6-9 ---> 2

2a. Branched second dorsal fin rays 7; anal branched rays 8; pectoral fin long, reaching dorsal fin level; Gulf, Hormuz and Tigris River basins, translocated to Lake Maharlu basin = **Liza abu  check caeca?


2b. Branched second dorsal fin rays usually 9; anal fin branched rays 9; pectoral fin short, not reaching dorsal fin level; Caspian Sea basin ---> 3

3a. Pyloric caeca equal in length; scales of head and back with one groove; oral edge of preorbital moderately concave ---> *Liza aurata


3b. Pyloric caeca in two groups, 3-5 short and 3-4 long; scales of head and back with 2-7 or more grooves; oral edge of preorbital bone deeply notched = *Liza saliens

--|----|----|--

Key to the Genera and Species of Nemacheilidae

The following key is modified after Prokofiev (2009).

1a. Anus closer to base of pelvic fins than to anal fin origin; pelvic fin origin obviously in advance of dorsal fin origin level [Tigris River and northern Gulf basins] = Turcinoemacheilus kosswigi
1b. Anus closer to anal fin origin than base of pelvic fins, usually at anal fin origin; pelvic fin origin at or slightly behind dorsal fin origin ---> 2

2a. No c---> 3
2b. Fleshy pelvic fin axillary lobe present ---> 5

3a. Males with fine, brush-like patches of breeding tubercles on pectoral fin rays (and on sides of head in some species); 3 radial bones in pectoral fin (if 4 then two external ones are flattened, more or less dilated and at least partly lie over each other) (only visible by dissection or x-ray) = Triplophysa
3b. No such tubercles in males; 4 elongate, cylindrical pectoral fin radial bones ---> 4

4a. Lateral line short and not extending end of pectoral fin level; scales absent; manubrium absent or weakly-developed (in the swimbladder, visible by dissection) [Hormuz and Kor River basins, northwestern Iran] = Seminemacheilus tongiorgii
4b. L
ateral line usually complete (if not, then extends back beyond dorsal fin origin); scales present; manubrium well-developed = Oxynoemacheilus

5a. Cheeks in adult males swollen; no black spot at base of anterior dorsal fin rays; no bars on body = Paracobitis
5b. Cheeks in adult male snot swollen; strong dark-black spot at base of anterior dorsal fin rays; usually bars on body ---> 6

6a. Bars restricted to posterior half of body; well-developed adipose crest supported by 22-25 procurrent caudal fin rays (visible by dissection or x-ray) [pelvic fin axillary lobe short; no sexual dimorphism; Tedzhen River basin] = Metaschistura cristata


6b. Colour pattern on body not as above; adipose crest weak to absent (if present supported by less than 15 procurrent caudal fin rays) ---> Paraschistura

 

 

Ilamnemacheilus longipinnis ADD to key

 

Key to the Species of Oxynoemacheilus

Oxynoemacheilus angorae

   

Oxynoemacheilus bergianus

      

Oxynoemacheilus brandtii

Oxynoemacheilus farsicus

?pic   

Oxynoemacheilus frenatus

   

Oxynoemacheilus kermanshahensis

  

Oxynoemacheilus persus

1a.

Key to the Species of Paracobitis

Paracobitis iranica

?pic

Paracobitis longicauda

Paracobitis malapterura

Paracobitis rhadinaea

?pic

Paracobitis smithi

  

Dorsal and ventral head views

Paracobitis vignai

?pic

1a.

Key to the Species of Paraschistura

Paraschistura bampurensis

Paraschistura kessleri

?pic

Paraschistura nielseni

?pic

Paraschistura sargadensis

.

--|----|----|--

Key to the Genera and Species of Percidae

All naturally distributed in the Caspian Sea basin.

1a. Canine teeth absent; prominent bars on flank; anal fin branched rays usually 8-9, rarely 10; lateral line scales 77 or less = Perca fluviatilis


1b. Canine teeth present; no bars on flank; anal fin branched rays rarely 10, usually 11 or more; lateral line scales 78 or more ---> 2  

2a. More than 18 branched rays in the dorsal fin; interorbital width equal to or less than eye diameter [translocated] = **Sander lucioperca


2b. Less than 19 branched rays in the dorsal fin; interorbital width greater than eye diameter = Sander marinus

--|----|----|--

Key to the Genera and Species of *Poeciliidae

1a. Dorsal fin rays 5-9, usually 7; caudal fin not modified in males; widespread = *Gambusia holbrooki


1b. Dorsal fin rays 11 or more; lower lobe of caudal fin greatly elongated in a sword-shape in adult males; Gulf and Namak Lake basins = *Xiphophorus hellerii

?pic

--|----|----|--

Key to the Genera and Species of Salmonidae

NEEDS to be checked ?

1a. Teeth in lower jaw absent or weak and brush-like; scales large, 13 or less from dorsal fin origin to lateral line, 100 or less in lateral line CHECK; caudal fin clearly forked ---> 2
1b. Teeth in lower jaw strong and conical; lower jaw long, extending back to or past mid-eye; scales small, 19 or more from dorsal fin origin to lateral line, 115 or more in the lateral line CHECK; caudal fin truncate ---> 3

2a. Mouth small? define; snout projects beyond lower jaw; ?, lower jaw not projecting markedly beyond upper jaw CHECK; teeth in roof of mouth few or absent; head length usually 4 times or more in standard length; body not pike-like; Namak Lake basin = *Coregonus lavaretus


2b. Mouth large?; snout not projecting; lower jaw obviously projecting beyond upper jaw; teeth in roof of mouth in broad bands; head length usually less than 4 times in standard length; body pike-like ?fusiform?'^;[Caspian Sea basin = Stenodus leucichthys

3a. Major anal fin rays 12 or more; anterior edge of preoperculum meeting orbital bones; pyloric caeca 140-249 ---> 4
3b. Major anal fin rays 13 or less; gap between anterior edge of preoperculum and orbital bones; pyloric caeca 23-66 ---> 5

4a.  [Caspian Sea basin] = *Oncorhynchus keta


4b.  [widespread]= *Oncorhynchus mykiss

5a. Body with dark spots on light background; vomer with teeth on head and shaft; lower fins without white leading edge ---> 6
5b. Body with light spots on dark background; vomer with teeth on head only; lower fins with white leading edge [Namak Lake basin] = *Salvelinus fontinalis

6a. ? [Caspian Sea, Lake Orumiyeh and Namak Lake basins and translocated]= Salmo caspius


6b. ? [widespread] = *Salmo trutta

--|----|----|--

Key to the Species of Siluridae


1a. Teeth short and weak (not snaggly); upper and lower jaws meeting at an antero-dorsal position; finely serrate or smooth pectoral spine posteriorly; colour dark; Caspian Sea, Lake Orumiyeh and Tedzhen River basins = Silurus glanis

1b.  Teeth robust and long (snaggly, catching on flesh); the upper and lower jaws meet at a dorsal and superior position; distinctly and coarsely serrate pectoral fin spine posteriorly; colour light; Tigris River basin = Silurus triostegus

--|----|----|--

Key to the Species of Sisoridae

1a. Head and body dorso-laterally with striated or elongate tubercles; thoracic adhesive apparatus is wider than long; caudal peduncle short (5.9-6.0 in standard length) [Tigris River basin] = Glyptothorax kurdistanicus

  
1b.Head and body dorso-laterally without striated or elongate tubercles; thoracic adhesive apparatus longer than wide; caudal peduncle  long (4.7-5.2 in standard length).  [Gulf and Tigris River basins] = Glyptothorax silviae

 


Species Accounts

The species dealt with here in detail have all been recorded from Iran and confirmed by specimens. Mention is also made of other species which occur on the borders of Iran or in drainage basins shared with Iran. These have no valid Iranian record but may eventually be found in that country. The listing here is selective from other papers by me on neighbouring countries as a number of species are unlikely to enter Iranian waters because their distributions are too remote, e.g. Cobitis elazigensis from the Tigris-Euphrates basin at Elazig in Turkey or too restricted, e.g. Typhlogarra widdowsoni from a cave in Iraq (see Coad, 1991b). Coad (1995a) gives a more complete listing of species found in waters neighbouring Iran. The most recent checklist on this fauna is by Esmaeili et al. (2010).

The definition of freshwater here includes the southern Caspian Sea which is at one-third seawater and has both nominally marine and freshwater species in its fauna.

The choice of introduced species to include in the Species Accounts is somewhat arbitrary. Soviet authorities introduced a number of species into the Caspian Sea and its tributaries and some of these became well established, spreading to Iranian waters, e.g. Liza aurata and Liza saliens, now commercially important. Other species did not become established but the potential for spread was there and so they are mentioned briefly in the Species Accounts. In northeastern Iran, the Tedzhen River flows into Turkmenistan and a number of exotic species are known from this former Soviet republic (see Aliev et al., 1988; Shakirova and Sukhanova, 1994; Sal'nikov, 1995). I have listed here only ones reported from the Tedzhen River basin and its reservoirs. The Tedzhen (Hari Rud in Iran) connects with the Karakum Canal which harbours a number of exotics as well as species from the Amu Darya. These may be able to colonise Iranian waters should they reach the Tedzhen River but are not included here in the absence of definite records.

A paper in Farsi by Farid-Pak (1957) records the grayling, Thymallus thymallus (Linnaeus, 1758), the lacustrine smelt Osmerus eperlamus eperlamus (sic) m. sprinchus (sic) (= Osmerus eperlanus eperlanus morpha spirinchus Pallas, 1814) and the sculpin Cottus gobio koshewnikowi Grazianov, 1907 from the Caspian coast of Iran but the first two species are distributed in waters remote from Iran and the last has not been recorded south of the Caucasus (Abdurakhmanov, 1962; Abbasov, 1980). They are assumed here to be misreadings of the literature and are not included in the species list.

Some marine species penetrate the fresh waters of southern Iran from the Persian Gulf and Sea of Oman. These species are included in a Marine List under Checklist in the Introduction. They are not included in keys but more detailed descriptions of these fishes can be found in the literature listed in the Bibliography such as Blegvad and Løppenthin (1944), Randall et al. (1978), Kuronoma and Abe (1986) and Assadi and Dehqani Posterudi (1997). Certain marine species do, however, spend a significant part of their life cycle in brackish to fresh water and are given full accounts as freshwater fishes, e.g. Carcharhinus leucas and Tenualosa ilisha. Choice of other marine species to be given a full treatment is dependent on frequency of capture, residence time and distance from the sea.

Coad (1991b; 2010) and the website Freshwater Fishes of Iraq give a list of marine species known from the Tigris-Euphrates basin but these are mostly records from the Shatt al Arab and Hawr al Hammar in Iraq which are under tidal influence. Hussain et al. (1989) give an account of seasonal fluctuations in species composition in the Shatt al Arab, Iraq. Little or nothing is known of the biology of these species in fresh and brackish waters. They are listed here to give an idea of the diversity of species which could be found in Khuzestan and in rivers along the Persian Gulf coast but are not covered in detail unless verified for Iran. Al-Daham and Yousif (1990) list additional species in an Iraqi estuary but do not distinguish the marine species which entered purely fresh water.

Taxonomy and systematics are active disciplines and scientific names of families, genera and species recognised from Iran can change. Older literature will be under the former name and searches for information in such fields as ecology should take this into account. These are described under the appropriate taxon but some significant changes, relevant to the Iranian species only, can be simply summarised as:-

Family changes:-

Cobitidae becomes Cobitidae and Nemacheilidae (the latter formerly Balitoridae).

Gadidae becomes Lotidae.

Generic changes:-

Caspialosa becomes Alosa.

Barbus becomes Barbus, Carasobarbus, Kosswigobarbus, Luciobarbus, Mesopotamichthys, and Tor.

Chalcalburnus becomes Alburnus.

Gobio becomes Gobio and Romanogobio.

Leuciscus becomes Petroleuciscus and Squalius.

Cobitis becomes Cobitis and Sabanejewia.

Nemacheilus becomes Ilamnemacheilus, Metaschistura, Oxynoemacheilus, Paracobitis, Paraschistura, Seminemacheilus, and Triplophysa.

Lebias becomes Aphanius.

Neogobius becomes Babka, Chasar, Neogobius, and Ponticola.

The Species Accounts are arranged by family after Nelson (2006). A higher classification can be found in the Checklist in the Introduction. Each Species Account is comprised of the following parts:

a) Illustration

The species is illustrated by a line drawing which is accurate in respect of body shape, number, position and shape of fins, scales and other structures. This drawing is usually a composite one, based on both a variety of published illustrations and on specimens.

Further illustrations are from various sources as indicated, are of varying quality and format, and may include colour and black and white photographs.

Diagrams may also be found in the Keys to illustrate characters not apparent in the main drawings, such as mouth structure.

b) Map

Distributions are summarized in the form of a map. Often two maps are given, one for the whole of Iran and one zooming in on distribution if restricted to a particular part of the country. The maps are from a world map layer provided by Demis bv (www.demis.nl), accessed through http://linuxgurrl.agr.ca/mapdata/itis/itisrosa.php.

Maps must be examined in conjunction with the text Distribution (see below). Map points are are a reflection of adequately documented museum collections and literature. As such they reflect catchability, ease of identification, rarity, size (large species not as easily preserved in museums as small ones but perhaps better documented, even if only in general), field work, available nets and other equipment, contiguity to research stations and universities, road accessibility, commercial interest, research interests, and so on. However, while bearing all these variables in mind and reading the Distribution summary critically, it is possible to gain a picture of fish distributions and objective rarity of species. Other sources of distributional data are field notes (principally mine and those of V. D. Vladykov) and sight and field records transmitted to me verbally by sources judged to be authoritative.

Note that many of these localities were ascertained in pre-GPS days from maps of varying quality and literature requiring some careful interpretation. Maps available in the field did not always match maps examined later and once I was lost for a whole day. Zooming in reduces accuracy proportionately.

Each symbol may represent more than one record because of the scale of the map or because of repeated visits to the same locality. Localities have not been sampled on a regular basis so population trends cannot be given. The general distribution in Iran and elsewhere is also given textually as outlined below.

The best records are those based on collections in a museum as these can be re-examined should any questions arise about identity and field data notes can be re-assessed for accuracy. However, the data associated with many museum collections are too vague or too contradictory to be included on maps with a locality symbol.

Criteria for inclusion of literature mapping records are as follows:-

1. Accurate identification (e.g. on geographical grounds; uniqueness of species so it could not possibly be anything else; lack of systematic/taxonomic confusion; distinctive characters cited in the text, drawn or photographed; assessed competence of author in identification),

2. Accurate latitude-longitude data. Latitude-longitude may be given by the author or derived by me from the literature based on maps and gazetteers, unique locality names, and my field experience close in time to when the material was recorded (road/river crossings have changed in some areas with new construction after the Islamic Revolution). One exception in accurate latitude-longitude data is that of migratory fish - if reported from a named river then the river mouth can be recorded since the fish pass this point on their migration (but few works mention the extent of upriver migration so no upper limit can be deduced; when an upper limit is given this is spot mapped; then the species is theoretically present in a continuous distribution from mouth to upper limit along the river but this distribution is not filled in and this presence along the river must be assumed from the known migratory habit).

Criteria for exclusion of literature mapping records are as follows:-

1. Generalised localities are not accepted, e.g. Safid River is not accepted since the actual locality along this river is unknown (except migratory fish - see above); landing ports, fish markets and fish farms are not included as localities unless the fish capture site or release site is known,

2. Localities with non-unique names, e.g. Hosseynabad, a common name for many villages; Shur River, a common name for any brackish stream, unless these have accurate qualifying data,

3. Descriptions with internal inconsistencies which cannot be resolved to one locality,

4. Named sites which cannot be found in a gazetteer; this is often a problem with Farsi names transliterated into various European languages with widely differing orthography,

5. Literature records which conflict with original field notes, jar labels or catalogues unless the literature explains why it differs.

Under Sources is a partial list of material examined, most with latitude-longitude. Some material was identified and is used in mapping distributions but lengths were not taken and that material is not listed. Sometimes fish were spirited away to be eaten, fell back in the river, leaped over nets, were kept by another researcher, were seen on market stalls and the source was given verbally, and so on. Collections in Sources may be annotated as "no other locality data" indicating that the collection data could not be interpreted to a latitude/longitude or was internally contradictory.

c) Scientific and Common Names

The use of scientific names is described in the Introduction. Scientific names are dynamic and can change as knowledge of the fishes increases. The ones used here are the latest available.

Common names in Farsi are given with the English translation in parentheses. Obviously some Farsi names are merely a translation from the English common name. Note however that some Iranian names are originally Arabic or Turkic in origin and I have not always been able to track their meaning. Some species have no common name and none has been advocated. Others have a common name which is applied to all members of the same genus (e.g. nemacheilid species are called mar mahi (= snake fish)) but this has not been repeated under each Species Account. The common name in Russian, Arabic, Azarbaijanian, English and from Pakistan is also given to facilitate communication and understanding; these names are in brackets.

There are often many common "book" names for Caspian Sea fishes. This is a result of the Russian designation of subspecies and other categories such as natio. The names are often based on geographical locations. These names are included here, although many of the taxa are not now recognised, as an aid to study of the literature. The names are probably not used locally. Azerbaijani names appear to follow mostly the Russian designations for these subspecies and again may not be truly local names.

The names cited as by J. J. Heckel in Arabic are also of dubious value. They are quite old, often from areas remote from Iran, and may not be in use today. A number of common names whose origin is Arabic are in use in Khuzestan however, although transmogrified into Farsi.

d) Systematics

An extensive synonymy or historical treatment of the mis-application of scientific names is not given. Some earlier names can be found in synoptic works such as Berg (1948-1949; 1949), Coad (1981d; 1985), Krupp (1985) and others. In certain cases, systematic or nomenclatorial problems remain unresolved and these are briefly discussed.

Type locality is given for species originally described from Iran or immediately adjacent waters. This type locality is given as cited in the original text description in quotes ("....") wherever possible. Some type localities are not given in quotes, e.g. middle Caspian Sea, to denote they are a general indication of where the fish was first described - this is usually applied for older literature not at hand or for fishes not described from Iran but nearby waters. The original text, jar labels or catalogues may be compared and interpreted where these are unclear, contradictory or spellings of place names have changed markedly. Most agree well between these three sources and are easily located with due allowance for variant spellings, handwriting skills and transcription errors. Disposition, number and condition of types may vary with time however. Eschmeyer's on-line "Catalog of Fishes" has disposition of types but these records are only as good as the most recent revision of the taxon concerned. Latitude and longitude are calculated for type localities in Iran wherever possible.

Note that transliteration from Russian names often gives variant spellings for authors of species names. Actual dates of publication may vary one or more years subsequent to the date on the journal or article, i.e. publication may be delayed. This may not be evident from an examination of the article but may be known to the author or others familiar with the situation. This has not always been clearly set down in print and accounts for varying publication dates in different sources.

The disposition and condition of type material is given where known along with catalogue numbers. Museum acronyms are from Leviton et al. (1985) but these may change, notably ZIL (Zoological Institute, Leningrad, U.S.S.R.) became ZISP (Zoological Institute, St. Petersburg, Russia) and the British Museum (Natural History), London became the Natural History Museum but retained BM(NH) as its acronym. Note that knowledge of type material in museums changes as the specimens are examined over time. Not all new information is published as it is the result of in-house curatorial work and may only be available in catalogues and jar labels. The information cited here is the most recent available to me.

Subspecies and lower, non-taxonomic categories have received names. Such taxa (and non-taxa) have a narrower range of meristic characters and certain distinguishing other characters compared with the species. Ranges and descriptions apply to the species as a whole, since many subspecies appear to be ill-founded where they have been studied in more detail, and indeed some species are not distinct but members of a wide-ranging and variable species. Certain subspecies may be valid, or their status is undetermined by recent study, and characters for these are given separately, either here or in Key characters or Morphology.

e) Key characters

The characters detailed here will separate the species from any Iranian freshwater fish. These characters (and the keys) should not be used to identify species from countries bordering Iran as they are specific to Iran.

f) Morphology

Under this heading are described a number of features which add to the key characters in describing the fish. Morphometric characters are not often used since the shape of body parts can be seen in the drawing and such characters vary greatly with sex and size in contrast to meristic characters. The accurate explication of morphometric characters depends on comparative statistics and is beyond the scope of this work. The assessment of variation between adults and juveniles or between geographical localities is limited by material and its presentation here by space.

The chief characters summarised here are meristic or countable characters. These include counts of scale, fin rays, vertebrae, gill rakers, and teeth. They are summarised as ranges based on literature sources (including my own data where this expands ranges). In certain cases literature data is extensive and swamps the few specimens available from Iranian waters. The literature ranges give an indication of how variable a species may be in a given character; data on a few Iranian specimens would give a misleading picture of potential variation which future students of Iranian fishes may find. Counts from Iranian specimens made by me are given with frequency in parentheses, e.g. dorsal fin branched rays 7(3), 8(34), 9(5) indicates that 3 fish had 7 branched dorsal fin rays, 34 fish had 8 branched rays and 5 fish had 9 branched rays.

g) Sexual dimorphism

Males and females often differ markedly in appearance, whether in colour, body proportions or in structural features and these are detailed here to obviate misidentifications.

h) Colour

The colour patterns of fresh and preserved specimens including males and females, young and adult, and spawning and non-spawning individuals are given where known. Colour can be a key character in determining the species but is also variable and should be treated with care in identifications. Some fish change colour to match their background or pale in response to a threat. Fish from muddy waters in Iran are often washed out and greyish in colour. Immersion in ice water enhances the colour patterns and some of this is retained in preservative.

i) Size

The maximum reported size is recorded as total length or standard length (if not specified then the source did not indicate which length was measured) and weight where known. These measures are not restricted to Iranian specimens since sample sizes are small for some species and would give a false picture of maximum size.

j) Distribution

This section summarises distribution for the whole range of the species both within Iran and the rest of the world. Within Iran the general distribution is given. The detailed mapped distributions are based on collections or literature with adequate data (see above under Map). Some literature and museum records are given simply as, e.g. "Safid River", which cannot be mapped accurately but can be cited in this section. Some literature records are included here but not every locality based on my field collections as these are summarised on maps. Not every river mentioned in the literature is listed here, as common species are assumed to be widely distributed within a basin; generally only those major rivers or general localities that are in basins without a mapped distribution are cited.

k) Zoogeography

The relationships of the species, its origins and movements in the past are given here, where this has been determined.

l) Habitat

The type of habitat favoured by the species is outlined and includes such factors as altitude, substrate, temperature, salinity, oxygen, flow regime, pH, vegetation, turbidity, pollution resistance, etc. There are few detailed studies of habitat requirements for many species: some can be deduced from morphology. Field data can give a partial picture but are often limited to one time measurements of seasonal and daily variables such as temperature which are necessarily of restricted value. Colour illustrations of habitats are included where available.

m) Age and growth

This section, and the following two sections, either have no information or masses of information. The Caspian Sea basin species are often widely known and have books and numerous papers written about them. There is also a vast "Soviet" literature on some of these species but I did not have the time nor the resources to digest it all. Here only brief summaries can be given and it is not always clear whether the Iranian populations, often at the southern edge of the species range, or recognised as a distinct subspecies, have the same general ecology as European or more northerly "Soviet" populations.

Most species outside the Caspian basin are poorly known ecologically. I have attempted to summarize what is known based on literature in particular from Iraq and Turkey where ecological studies of varying quality have been published on some of the species. Morphology can be used to gain a general picture and knowledge of related species helps.

Generally growth in fishes is fastest in the youngest age groups, slowing with age and with investment in reproduction. Maximum age varies considerably, some small species living only a few years while others are much larger and are reputed to live longer than people. Conventionally, age may be represented by a number then the + sign, e.g. 0+ = a fish in its first year of life, less than one year old; 6+ = a fish between 6 and 7 years old.

n) Food

Diet is reported from literature studies and from brief examination of gut contents by me. Diet varies seasonally, daily, with age, between sexes, and with changes in environmental conditions but most fish concentrate on one or a few major groups. These are scrapers, invertebrates and fishes, and rarely aquatic macrophytes.

o) Reproduction

The spawning season, migrations, egg numbers and diameters, and reproductive behaviours are recorded here. Some migratory behaviour and ages at spawning may be recorded in the the Habitats and Age and growth sections.

p) Parasites and predators

This section contains information on the parasites and predators of the species described. I have recorded only parasites known from Iranian populations. There is a more extensive literature on Iraqi populations (see Mhaisen, 1980; Coad and Al-Hassan, 1989) and on European or Caspian Sea populations (see Romanov, 1955) for species found in Iran. For eastern waters consult Moravec and Amin (1978) on Afghanistan and Mirza (1978) on Pakistan. In the absence of definite records for Iran and in the interests of saving space, I have not cited this extensive literature.

There are a number of piscivorous birds in Iran (see Scott et al. (1975), Behrouzirad (2007) and general field guides) and these take fishes but there seems to be little direct observation on the fish species preferred.

q) Economic importance

Note that fishery information may be given on an annual basis but the year reads 1965-1966 or 1965/66; Iranian years start in March and run across 2 western calendar years.

r) Conservation

This section details conservation measures undertaken or needed for the species. A general survey of conservation status of native Iranian freshwater fishes is given by Coad (2000a).

s) Further work

This section gives some suggestions for knowledge gaps that should be filled.

t) Sources

This section refers to papers or synoptic works on the species in addition to those cited in the text. It should be noted that a number of synoptic works refer to several species in Iran, e.g. Berg's "Freshwater Fishes of the U.S.S.R. and adjacent countries", and these are not listed repetitively under each Species Account although they are to be found in the Bibliography. Web sites or URLs are cited as documentation of statements but it should be noted that these may become broken links and they are not continually verified as active.

Descriptions are based on Iranian specimens wherever possible but additional material from neighbouring countries has also been examined. Meristic counts, for example, are given as frequency distributions for Iranian material while general ranges for these characters are based on Iranian material, on literature and on counts of other specimens listed here briefly. Descriptions are also based on material seen in bazaars or captured in the field but not retained, and on photographs, drawings, field notes of other collectors, and verbal descriptions of other scientists.

Details on collections are on file at the Canadian Museum of Nature, Ottawa and in other institutions as recognised by their acronyms. Locality data is given in short form and the reader is referred to the website of the relevant museum for further information. Locality names are taken from U.S. Board on Geographic Names publications and these may vary from names on labels in museums. The Board names contain both conventional and local Farsi, Arabic and Turkish names of localities. I have interpreted names as best I can and have, for example, retained English names for major water bodies and towns where a strict usage would be bewildering, e.g. Harirud = Tedzhen River, Sefidrud = Safid River, Al Mawsil = Mosul, Darya-ye Mazandaran = Caspian Sea, and so on. Sometimes a collection is annotated as "no other locality data", indicating that no further details are known or localities cited could not be found on maps or in a gazetteer (and thus there is no latitude-longitude). Collections listed as uncatalogued are mostly held in the Canadian Museum of Nature and may eventually receive a catalogue number. The collections listed are those examined for morphology. Map records include these collections, other collections checked for identity and locality only, and literature sources, all kept in a database held at the Canadian Museum of Nature: these would be too lengthy to list here.
 


Petromyzontidae

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Lampreys in the family Petromyzontidae are found in cooler waters of the northern hemisphere, with a few related species in other families in the southern hemisphere. Their origins lie at least 300 million years in the past. There are about 37 lamprey species with only 1 recorded from Iran.

Lampreys are jawless fishes, lacking bone in the skeleton and having 7 pairs of pore-like gill openings. The eel-like body has no pectoral or pelvic fins. There are 1 or 2 dorsal fins and a caudal fin. An anal fin-like fold develops in spawning females. The mouth is a suctorial disc armed with rows of horny teeth. There are also teeth on the tongue. The median nostril, or nasohypophyseal opening, is not connected to the mouth. There is a light-sensitive pineal organ or "third eye" behind the nostril. The skin is covered in mucus which is poisonous to fishes and humans. Lampreys are edible if the mucus is cleaned off.

Their tooth arrangement is used in classification and identification along with the number of myomeres (muscle blocks along the body). Both tooth counts and the number of cusps are used, in particular those on the supraoral lamina (bar above the "mouth", the oesophageal opening), the infraoral lamina (bar below the "mouth") and the row of teeth on both sides of the "mouth". There are various series of smaller teeth and of course teeth on the tongue. Larval lampreys lack teeth and are particularly difficult to identify and their determination often requires specialist knowledge. Characters for the larvae include counts of myomeres and pigmentation patterns.

Lampreys have an unusual life cycle. Adults die after spawning and the eggs develop into a larva, known as an ammocoete, which lacks teeth, has an oral hood, eyes covered by skin, a light-sensitive area near the tail, and is a filter-feeder while buried in mud and silt. Fleshy tentacles in the oral hood are used to extract minute organisms from the water, such as algae (desmids and diatoms) and protozoans. After several years (up to 19 but usually 7 or less), the ammocoete transforms into an adult with enlarged eyes, teeth, a different colour and pronounced dorsal fins. The body shrinks during this metamorphosis and adults are only larger than ammocoetes if they feed. The adult may be a parasite on other fishes and marine mammals, or non-feeding. Individuals of a species may or may not be parasitic and different species may be parasitic or non-parasitic. The non-parasitic species are believed to have evolved from a parasitic species so there tends to be closely related parasitic/non-parasitic species pairs.

Parasitic adults feed mostly on other fishes, attaching to their bodies by suction and using their toothed tongue to rasp through the skin and scales to take blood and tissue fragments. Prey is detected by sight but some lampreys attach to hosts during the night. Perhaps this reduces their own predation risks and enables them to approach their quiescent hosts more easily. Lampreys tend to select larger fish as these survive longer and ensure a good food supply. The flow of blood is aided by an anti-coagulant in lamprey saliva called lamphedrin which also serves to break down muscle tissue. The attack may weaken or even kill the host. Weakened fishes are more prone to diseases and the wound provides an easy path of entry for them. The fish (and marine mammal) species parasitised are varied and reflect availability in the habitat.

Marine lampreys enter fresh water to spawn and freshwater species may move into or up streams. The scientific name of the family means "stone sucker" and the adult mouth is used to hold or suck onto stones as well as on prey. This suction enables the lamprey to maintain position in fast-flowing streams when spawning and even to climb over rapids and small waterfalls. Usually spawning occurs in shallow water with a moderate current, a bottom of gravel and nearby sand and silt for the ammocoetes to live in. Either or both sexes build a nest by moving gravel around with their sucking mouths and by thrashing their bodies. A shallow depression is formed, about 0.5-1.0 metre long. Spawning often occurs in groups and several males may attach to a female with the sucking disc. The process takes several days as only a few white to yellow eggs are laid at a time. The eggs are adhesive.

Adult lampreys are usually caught when attached to a host or when spawning. Electro-shocking will force ammocoetes out of their u-shaped burrows to the surface and immobilize adults. They sometimes attach to boats and occasionally to human swimmers when their skin is cool but are easily removed, perhaps because nobody has left a lamprey on their skin long enough to see if the tongue starts rasping flesh!

Genus Caspiomyzon
Berg, 1906

This genus is characterised by having 2 dorsal fins, an oral disc narrower than the body, teeth are generally low and blunt, the supraoral lamina is small, oval and sometimes has 2 tubercles and rarely 2 teeth, the infraoral lamina has 4-6, usually 5, teeth which may be bicuspid at their tips, there are about 8 small teeth of equal size in the transverse lingual lamina, the exolaterals, anterials and posterials are strong and close together, anterior and endolateral circumorals 9-11, usually 11, and 3 long, papillose velar tentacles are present. The first illustration below shows a notch at the end of the second dorsal fin which is an error.

There is a single species in the genus found only in the Caspian Sea basin. Agnathomyzon Gratzianow, 1906 and its subgenus Haploglossa Gratzianow, 1906 are synonyms of Caspiomyzon (Eschmeyer et al., 1996).

Caspiomyzon wagneri
(Kessler, 1870)

Adult

Caspiomyzon wagneri adult, courtesy of Afshin Afzali, University of Tehran.

Adult

Caspian lamprey, Shirud River, photo courtesy H. Nazari.

Adult

Caspian lampreys, Shirud River, photo courtesy H. Nazari.

Adult

Ammocoete

Disc

Caspian lamprey disc, photo courtesy H. Nazari.

Common names

مارماهي (= mar mahi, meaning snake fish), مارماهي دهان گرد (= marmahi-ye dehangerd, meaning round mouth snake fish), mahi dehangerd, mahi dehangerd daryacheh-ye khazar or dahangerd-e-Daryaye Khazar (= Caspian Sea round mouth fish).

[ilanbaligi or xazar ilanbaligi, djilan-balux or morma in Azerbaijan; kaspiiskaya minoga or Caspian lamprey in Russian; Volga lamprey].

Systematics

The type locality of Petromyzon Wagneri is from the mouth of the Tvertsa to Astrakhan; Oka and Kama rivers and the 3 syntypes (29.0-33.0 cm) are in the Zoological Institute, St. Petersburg (ZISP 31) (Holčík, 1986). The Zoological Museum of Moscow University (ZMMU) has one syntype from the Kura River near Evlakh (P-1393) and one from the Moskva River (P-555) with P-569 from the Volga River near Kazan being lost (Pavlinov and Borissenko, 2001). The Naturhistorisches Museum Wien in 1997 had one specimen listed as "? syntype, ? paratype" (sic) under NMW 61053. Agnathomyzon (Haploglossa) caspicus Gratzianow, 1907 is a synonym.

Key characters

This is the only lamprey species in Iran, easily recognised by the absence of pectoral and pelvic fins, a round, suctorial mouth containing blunt teeth, and 7 branchial openings.

Morphology

Characters of the species are the same as the genus. Trunk myomeres number 53-68 in ammocoetes; and 68(2) or 69(1) in adults from Iran. Ginzburg (1936a) describes ammocoetes from Iran. Renaud et al. (2009) give details of the feedding apparatus. Nazari et al. (2009) found significant differences for morphometric, but not meristic, characters, between fish from the Shirud and Talar River, although a principal components analysis showed relatively high overlap.

Sexual dimorphism

Females reach larger sizes than males and have a smaller urogenital papilla. During the spawning migration, the lamprey undergoes certain morphological changes some of which have been linked to sex of the fish. The teeth become blunt, fin size increases, the dorsal fins become almost united at the base in males, and there is a change in colour. The urogenital papilla length in males increases from a mean of 1.1 mm to 4.9 mm and in females from a mean of 0.6 to 1.7 mm.

Colour

Adults are dark grey with a silvery-white belly. Spawning adults become black on the back and flanks with a grey belly covered with dark oval spots, or are an overall golden colour (Hassan Nazari, pers. comm., 28 July2011, see photo above). Ammocoetes are a pale grey to yellowish with a white belly.

Size

Attains 57.5 cm total length and 205.5 g as the adult and 13.0 cm total length as the ammocoete. After metamorphosis of the ammocoete there is a shrinkage in length, the difference between prespawning and spawning adults being on average 22.3% in Iranian samples (Renaud, 1982). There is also a small variety which measures 19-31 cm and can attain sexual maturity at 19.1 cm (forma praecox).

Distribution

Found only in the Caspian Sea and rivers draining to it, in particular the Volga where it had its largest distribution but is now known only as far as the Volgograd Reservoir; also in the Ural, Terek, and Kura rivers. It is recorded in Iran from the upper reaches of the Aras River, and from the Astara to the Gorgan River along the whole Caspian coast. Specific localities include the Aras River, Anzali Mordab and the Nahang Roga, Pir Bazar Roga, Pasikhan River and Siah Darvishan River in the Anzali region, to Kisom on the Safid River, Cheshmkelya east of the Safid River, Tajan River, Sardab River, Haraz River, Babol River, Tonekabon River, Pol-e Rud, Gorgan River, and in most large streams (Derzhavin, 1934; Holčík and Oláh, 1992; Hosseinpour, 1995; Abbasi et al., 1999; Kiabi et al., 1999; Abdoli, 2000; Abdoli and Naderi, 2009). Migrations into the Babol, Gorgan and Sardab rivers are reported by Ghasempouri (1993), the Sardab and Chalus rivers by the Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran (1996), and the Shirud (Nazari and Abdoli, 2010), for example.

Zoogeography

Known only from the Caspian Sea, its relationships remain uncertain and research is ongoing (Claude B. Renaud, pers. comm., 18 May 2007).

Habitat

The habitat of this species in the southern Caspian Sea proper is unknown although some specimens have been caught in the Caspian at 600-700 m (Jolodar and Abdoli, 2004). Larvae burrow 1-2 cm into the river bottom and favour areas where current is moderate at river bends. They can also be found in the centre of rivers or in backwaters. Fine-grained sand with some ooze and detritus is preferred at all stages of larval growth but larger larvae can also be found in a silt-sand bottom with much plant debris and macrophytes. The ammocoetes select and change habitat according to sediment size as they grow. They prefer depths greater than 3 m as protection against drying out, are mostly shallower than 11 m but as deep as 22 m (Ginzburg, 1970), yet in different rivers or at different times will be concentrated in water of markedly different depths, e.g. 30-85 cm versus 6-8 m.

Spawning migrations up the Volga River used to exceed 1500 km but construction of dams now prevents this. The lamprey migrates in schools with the smaller fish arriving in estuaries first. Larger lampreys migrate more quickly and travel further. The speed varies from 1.9 to 15.9 km/day. The migration is triggered by decreasing water temperature and increasing water level. The strongest migration is reported at 6-11°C. Movement upriver only occurs at night, near the surface when dark and on the bottom when the moon is out. During the day, the lampreys hide among stones. Body fat in the Volga delta was 34% but by the time the fish reached the spawning grounds upriver it had declined to 1-2%. In the Kura River of Azerbaijan, the lamprey migrates at the same time as the Caspian salmon (mahi azad, Salmo caspius) and often attaches to the opercular region of this species. The peak of this run is in December and January. The migration in the Volga takes place from the middle of September to the end of December. Migrating lampreys prefer a current velocity of 0.4-0.6 m/sec and stay close to banks and the bottom. Prespawning adults overwinter among stones or in the substrate of rivers. During winter-spring several individuals may be found coiled in a ball under stones (Askerov et al., 2001). They hardly respond to external stimuli such as noise or being handled. Transformed lampreys migrate to the Caspian Sea. Before breeding, males change colour, increase slightly in size, develop their fins, and become much more active (Askerov et al., 2001).

Nazari and Abdoli (2010) note a short fall migration in late September to October with the main migration being in spring (see below). Movement was mostly at night and involved swimming and resting attached to the concrete of a bridge used as the observation post.

Age and growth

The growth rates of metamorphosing lampreys and adults are almost unknown. Length and weight decrease but coefficient of condition increases in spawning as opposed to pre-spawning adults. The shrinkage in mean total length is 18-26%. Females are heavier than males up to about 43 cm but past this point males weigh more. There are 3 age groups of larvae in the Volga (Ginzburg, 1970), with average lengths of 3.1 cm, 6.2 cm and 10.1 cm and 2-4 age groups in the Kura. In their fourth year of life they metamorphose to adults after a downstream migration into the Caspian Sea. Adult life span is at least 1 year and 5 months. Maturity is attained in May and the beginning of June in the Volga, and from May to the end of July in the Kura River. Mature lampreys are mostly 35-41 cm in the Volga and 41-46 in the Kura River. The female lamprey dies after spawning but the male may live longer until sperm production ceases.

Nazari et al. (2010) investigated growth parameters in fish from the Shirud and Talar River. Most fish were were in the 367-369 mm length group, length-weight relationship was positive, high and significant, growth was negatively allometric, the coefficient of condition was higher in females, sex ratio was nearly equal, and growth parameters were similar in the two rivers.

Food

Abakumov (1959) maintains that this lamprey attacks Caspian salmon (Salmo caspius) based on nineteenth century observations by Kessler (1870a) and Kavraiskii (1896-1897). Lelek (1987) also considers it to be parasitic. The lampreys may only have been using Caspian salmon for transport. Certainly the teeth in this lamprey are blunt, unlike those in lamprey species known to parasitise fishes. In contrast, Holčík (1986) states that it is non-parasitic and Ghasempouri (1993) agrees. Renaud (1982) supposes that adults feed on amphipods since juvenile acanthocephalans (Corynosoma sp.) are found in prespawners. This worm has amphipods as the intermediate host. However, Holčík (1986) thinks that the acanthocephalans are swallowed while the adult lampreys are feeding on the internal organs of dead fish they scavenge. Certainly larvae of Corynosoma strumosum (perhaps correctly C. caspicum: B. Kiabi, in litt., 1994) are found only in the body cavity of fishes. Renaud et al. (2009) list it as a carrion feeder but note the well-developed buccal glands which may compensate for the blunt teeth and it may well feed on fishes. The feeding habits of the adult of this species remain to be confirmed by direct observation. Gut contents include aquatic vegetation in Iran and in the Volga delta. Migratory, transforming and spawning lampreys do not feed. The gut diameter decreases from 2.7 mm in prespawners to 1.4 mm in spawners in Iran (Renaud, 1982). Ammocoetes feed on detritus and diatoms.

Reproduction

Ginzburg (1969; 1970) examined the reproduction of this species below the Volgograd Dam on the Volga River and similar conditions may obtain in Iran. The dam has probably increased fecundity by reducing the length of the spawning migration so that the fish have more energy reserves for egg production. A spawning migration exists from December to May with a peak concentration in the second 10 days of February although the catches declined in April at least in part because of the opening of the spillway of the dam. Before the dam was built the migration from the Caspian Sea passed through the delta from mid-October to mid-December, with a peak in December. The fish migrated when water temperatures reached 10-11°C and moved through channels where the current was strongest. Spawning begins at 15-16°C, usually in early June but sometimes at the end of March through to the beginning of July, and temperatures during spawning are usually 15-23°C. Each female produces up to 60,000 turquoise or blue-green eggs and spawns once in her lifetime. Eggs are ovate and diameter reaches 1.5 mm. The eggs are laid on coarse to fine-grained, turquoise sand at a water depth of 3.5-19.0 m, sometimes shallower. The egg colour is cryptic against the sand substrate. Many eggs are carried downstream by the current. A redd is excavated in sand or gravel by the male or by the female (authors differ on this point) and the lamprey attaches to stones by their suctorial disc. The male attaches to the female's head with his disc and wraps his body around hers. The tails of both fish quiver and eggs and sperm are released at the same time. Females release all their eggs but males may spawn again with other females. Ammocoetes hatch after 8-10 days at 17-23°C. Metamorphosis of ammocoetes occurs at 8.0-11.0 cm in October in Iran.

Nazari and Abdoli (2010) examined migration and reproduction in lampreys from the Shirud in the southern Caspian Sea from 16 March to 2 May at 11.0-21.25°C. The most intensive migration was at night (peaking at 2100 and declining to 0300 hours) at 16°C (34.4% of the run). About 75% of the run had  passed by the time water temperature reached 16-17°C. Migration stopped when temperature reached 21°C. Numbers observed each night varied from 1 to 60, average 17, with peak migrations at 26 March to 10 April and 15 April to 25 April. Sex ratio was 1.07:1 in favour of males but not significantly different. Absolute fecundity was31,758-51,198 eggs (mean 41,924 eggs) relative fecundity was 80.3-148.1 eggs/mm length (mean 107.2 eggs/mm length) and 260.8-677.4 eggs/g (mean 397.6 eggs/g). Egg diameter was 0.78-1.15 mm (mean 0.92 mm). The gonadosomatic index of females was 5.83-31.44 (mean 11.22), the peak being in mid-April. Downstream migrating lampreys were spent but no dead ones were noted so some may survive to spawn a second time. Two ammocoetes, 20 and 22 mm long, were found near the mouth of the Shirud River on 18 April 2006 (river bank in a substrate of the sand-mud, water depth <30 cm). They probably belong to the autumn migratory group (Hassan Nazari, pers. comm., 28 July2011).

Parasites and predators

See above under Food. Nazari et al. (2010) also record Corynosoma in their fish. Caspian lampreys are eaten by Silurus glanis, Lota lota, Sander lucioperca, and Huso huso.

Economic importance

This species was consumed and used for oil extraction in the former U.S.S.R. (Thomas, 1961; Ginzburg, 1969). Their fat content is so high that they were once dried and used as candles (Kottelat and Freyhof, 2007) and the high fat level makes them tasty (Askerov et al., 2001). The catch in the Volga-Caspian region was 3,420,000 kg or 33.4 million fish in 1913 but fishing by state organizations ceased after the Volgograd reservoir was constructed. The mean annual catch in Azerbaijan for 1930-1963 ranged from 10 to 269 tonnes. Local fisheries continue but are of little significance. It is not commercially important in Iran for religious reasons but catches of several hundred kilograms can be made in an hour in such rivers as the Gorgan, Babol and Sardab (Ghasempouri, 1993).

This lamprey is ingested medicinally for treatment of haemorrhoids and besmi (sic, ?) by Turkmen of the southeastern Caspian (Hassan Nazari, pers. comm, 29 July 2011).

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use as food, in textbooks and because it is reputedly ichthyosarcotoxic. Intoxication results from eating the flesh, skin or surface mucus of raw or cooked Caspian lamprey, the location of the toxin being uncertain. A biogenic amine is believed to be responsible. Mucus may cause skin irritations. Poisoning can be avoided by soaking the lamprey in brine as cooking alone is insufficient. Symptoms develop in a few hours and include nausea, vomiting, dysenteric diarrhoea, urge to urinate or defecate without ability to do so, abdominal pain and weakness. Recovery takes several days and treatment is symptomatic (Coad, 1979b). However lampreys lack scales and are not eaten in Iran.

Conservation

The Caspian lamprey has been proposed for inclusion in the "Red Book of the U.S.S.R." which forms the basis for measures to protect species (Pavlov et al., 1985) and is listed as "vulnerable" in Europe by Lelek (1987) and Maitland (1991). It is vulnerable because it migrates into rivers which are polluted and dammed and because of its restricted and declining distribution. These conditions apply particularly in Iran, although there is some evidence for spawning based on captures in the 1990s (Holčík and Oláh, 1992).

Kiabi et al. (1999) consider this species to be near threatened in the south Caspian Sea basin according to IUCN criteria. Criteria include medium numbers, habitat destruction, widespread range (75% of water bodies), absent in other water bodies in Iran, and absent outside the Caspian Sea basin. Mostafavi (2007) lists it as near threatened in the Talar River, Mazandaran.

Further work

The question of adult diet remains unresolved and the general biology of this species in Iran needs to be elucidated.

Sources

The main source of information on this species is the summary by Holčík (1986) which should be consulted for further details on morphology and biology.

Iranian material: CMNFI 1970-0511, 7 ammocoetes, ? 30-82 mm total length, Gilan, Shafa River estuary (37º35'N, 49º09'E); CMNFI 1970-0514, 33 ammocoetes, ? mm total length, Gilan, Shafa River estuary (37º35'N, 49º09'E); CMNFI 1970-0515, 18 ammocoetes, ? 25-98 mm total length, Gilan, Shafa River estuary (37º35'N, 49º09'E); CMNFI 1970-0534, 30 ammocoetes, ? mm total length, Gilan, Shafa River estuary (37º35'N, 49º09'E); CMNFI 1970-0535, 14 ammocoetes, ? mm total length, Gilan, Shafa River estuary (37º35'N, 49º09'E); CMNFI 1970-0545, 1 adult? see photo?, ?mm total length, Gilan, Safid River (37º01'N, 49º38'E); CMNFI 1970-0546, 2 adults, 352.0-355.0 mm total length, Gilan, Safid River (no other locality data); CMNFI 1970-0547,6 adults and 2 ammocoetes, ? photos? mm total length, Gilan, Shafa River estuary (37º35'N, 49º09'E); CMNFI 1970-0585, 3 adults, 406.0-455.0 mm total length, Gilan, Nahang Roga River (37º28'N, 49º28'E); CMNFI 1971-0327A, 1 adult (part of trunk), Gilan, Shafa River estuary (37º35'N, 49º09'E); CMNFI 1979-0787, 11 adults, ?mm total length, Gilan, Nahang Roga River (37º28'N, 49º28'E); CMNFI 1980-0118,8 adults, ? mm total length, Gilan, Gelroudkhan River, tributary of the Anzali Mordab (no other locality data); CMNFI 1980-0119, 10 adults, ? mm total length, Gilan, Gelroudkhan River, tributary of the Anzali Mordab (no other locality data); CMNFI 1980-0139, 44 ammocoetes, ? mm total length, Gilan, Golshan River estuary (37º26'N, 49º40'E).

Carcharhinidae
 

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This family, the requiem or ground sharks, contains more than 50 species of large sharks found world-wide in tropical to warm-temperate waters. There is only 1 species in Iranian fresh waters. A second species is reported from an Iranian river under special circumstances and is not regarded as a resident (see Marine List). They are distinguished from other sharks by a complex of characters including having an anal fin; 5 gill slits; 2 dorsal fins; no fin spines; nictitating eyelids; and a scroll intestinal valve. The first dorsal fin base is in front of the pelvic bases; there is a wavy dorsal tail fin margin; well-developed, knife-like teeth with cutting edges; usually no spiracles; and precaudal pits.

This is one of the largest and most economically important shark families. Most members are voracious predators as their common names suggest and they are frequently dangerous to man. Some of these species enter rivers and remain there for long periods causing human fatalities. These sharks are usually viviparous. Food includes a variety of fishes, sharks, rays, squids, crustaceans, marine reptiles, birds and mammals, and carrion and garbage.

Shark flesh can be eaten and is religiously permissible in Iran.

Genus Carcharhinus
Blainville, 1816

There are about 31 species of gray sharks found world-wide but only one regularly enters fresh water in Iran. A detailed definition of the genus is given by Compagno (1988).

Carcharhinus leucas
(Müller and Henle, 1839)

  

Upper and lower tooth

BM(NH)1924.10.1:1, Tigris River at Al Karradah near Baghdad

BM(NH) 1874.4.28:9, Tigris River near Baghdad

Common names

kooseh, kuseh, kooseh-kuli, sag mahi (= dog fish).

[kosetch or kossage, jarjur in Arabic; bull shark].

Systematics

Carcharhinus leucas was originally described from the Antilles.

A number of shark species have been reported as entering rivers of the Tigris-Euphrates basin including Iranian tributaries (Günther, 1874; Day, 1875-1878; Sykes, 1902; Kennedy, 1937; Hunt, 1951; Khalaf, 1961; Mahdi, 1962; Zorzi, 1995; and others). The species appeared under such names as Carcharhinus gangeticus (Müller and Henle, 1839), Eulamia (= Carcharhinus) lamia (Blainville, 1820), and Carcharhinus menisorrah (Valenciennes in Müller and Henle, 1839). A recent revision of carcharhinid sharks by Garrick (1982) cites only Carcharhinus leucas from fresh waters of the Tigris-Euphrates basin and Compagno (1984) concurs. Coad and Papahn (1988) also list specimens which confirm this species to be present.

Key characters

This is the only shark species commonly encountered in Iranian fresh waters and can be recognised by the 5 gill slits, upper caudal fin lobe larger than lower, and the arched mouth armed with teeth on the underside of the head. Distinction from other sharks is given in Compagno (1984).

Morphology

The snout is very short, rounded and ends bluntly. Its length is less than the distance between the nostrils and much less than the mouth width. There are 12-14, usually 13, teeth on each side of a median tooth in the upper jaw and 12-13, usually 12, teeth on each side of a median tooth in the lower jaw. Teeth are heavy, broad, almost triangular, erect near the jaw symphysis but becoming slightly oblique and more concave or notched nearer the mouth corners. The teeth are strongly serrated, more so near the base, and upper teeth more so than lower teeth.

The first dorsal fin lies over or just behind the level of the axil of the pectoral fin. The apex of the first dorsal fin is pointed to somewhat rounded. The second dorsal fin is high, has a short posterior lobe and lies just over the level of the anal fin origin. Pectoral fins are broad and their tips are narrow and pointed. There is no interdorsal ridge (the back is smooth between the dorsal fins). The upper precaudal pit is well-developed while the lower pit is weak.

Sexual dimorphism

Males bear claspers. The pelvic fins are partially modified into grooved, rod-like structures which are held together to form a tube and are used in mating. They are not used to clasp the female but as an intromittent organ. Females are larger than males.

Colour

Fin tips are dusky to black, particularly in young. There is no other obvious colour pattern although the back is darker than the belly, being bluish, grey or brown. Fins are similar in colour to the neighbouring body.

Size

Attains 3.24 m (Garrick, 1982), 3.40 m (Carpenter et al. 1997), rarely to 4.0 m and an estimated weight of over 600 kg (McCord and Lamberth, 2009). Fish in Iranian fresh waters have been estimated as up to 2 m in length but naturally circumstances were not always favourable for an objective and detached size judgment.

Distribution

Sharks have long been known to enter fresh waters in the Tigris-Euphrates basin. Zorzi (1995) records a book by Pausanias, "Guide to Greece", written in the late second century which refers to sharks in "the Euphrates...., which fatten monsters as man-eating as any in existence".

One of the earliest distributional records is found in the Arabic work "Wonders of Creation" by Zakariya al-Qazwini published in 1263 A.D. and later translated into Persian. The sharks were found at Basrah on the Tigris River in what is now Iraq and were cited as formidable because of their voracity and teeth like the points of spears. Shark attacks still occur at Basrah (Coad and Al-Hassan, 1989).

Subsequently reported in the Tigris River above Baghdad about 850 km from the sea (Günther, 1874; Kennedy, 1937) before dams were built. Sykes (1902) saw sharks in the Ab-e Gargar (Karun River in Iran) at Shushtar 420 km from the sea, Wilson (1942) reporting on events in 1908 records sharks from between Shushtar and Ahvaz and near Shushtar, Blegvad and Løppenthin (1944) report then from Khorramshahr, and Hunt (1951) reported them from the Karun River, Khowr-e Bahmanshir and Shatt al Arab (Arvand River). Coad and Papahn (1988) report sharks at Ahvaz on the Karun River about 275 km from the sea as well as further up river at Shushtar and down river in the Khowr-e Bahmanshir.

Zoogeography

This shark is found world-wide in warm temperate to tropical seas and is reported from fresh waters in Africa, Asia, Australia and the Americas.

Habitat

This is a shark of coastal waters such as harbours, bays and estuaries but unusually it will penetrate far up rivers, as far as 4000 km up the Amazon River. It is said to be a sluggish bottom dweller except when attacking prey and in the sea may be found down to at least 150 m although usually at less than 30 m. They are said to invade the Khowr-e Bahmanshir and Karun River of Iran from July to September when freshwater flow is at a minimum and tidal penetration of salt water is at its highest. However they do travel well beyond tidal influence in Iran. Local people along the Bahmanshir River near Tangeh Se in Khuzestan maintain that it is dangerous to swim there because of these sharks. They are occasionally trapped in nets set for Tenualosa ilisha and may be caught on hooks. They are not as common as in the past (N. Najafpour, pers. comm., November 2000).

Age and growth

Maturity in males is attained at 1.60-2.25 m and in females at 1.80-2.30 m. Mature fish are about 6 years old and life span is up to about 14 years.

Food

Food is a wide variety of fishes including tunas, small sharks, and rays, as well as crabs, shrimps, molluscs, cephalopods, sea urchins, turtles, sea birds and mammals. Diet in fresh water has not been investigated in Southwest Asia although Blegvad and Løppenthin (1944) reported that sharks station themselves under the date palms at Khorramshahr to eat the falling dates!

Reproduction

Birth size is about 56-81 cm and takes place in estuaries and river mouths. Females may contain up to 13 embryos and the gestation period is 10-11 months. This species is known to breed in fresh waters, such as Lake Nicaragua in Central America, but there have been no reports of reproduction in the Tigris-Euphrates basin.

Parasites and predators

None are reported for Iran.

Economic importance

This shark has a considerable impact on people using water directly in Khuzestan. A number of severe injuries and fatalities have been reported in fresh waters through shark attacks. The first comprehensive report in modern times was by Hunt (1951) although accounts date back to the thirteenth century (Coad and Papahn, 1988). The latter summarize recorded attacks and add new ones for a total of 34 in the period 1941-1985, of which about half were fatal. Additionally Wilson (1942) reports a woman taken by a shark while drawing water between Shushtar and Ahvaz and a 9 foot (= 2.8 m) near Shushtar which killed two boys and a girl. These Iranian records are a significant proportion of freshwater attacks worldwide, about 28%. A number of soldiers were apparently victims during the Iran-Iraq war but no records have come to light. Men, women and children are attacked as well as horses and sheep, only the massive water buffalo is said to be safe. Many minor attacks and narrow misses are probably not reported. Attacks are said to have declined in recent years since shark oil is no longer used to caulk boats but this is probably a local legend. People were attacked while swimming, paddling, bathing, washing vehicles or fishing. There was no apparent triggering factors for the attacks as victims were dressed in various colours and types of clothing, engaged in various activities and environmental conditions where known varied between attack sites.

Freshwater shark attacks have even appeared in a novel "Harem" by Mossanen (2002).

In other parts of the world, this species has been used for its flesh and fins, as leather, for its liver oil and for fishmeal. Sharks can be eaten by Muslims if "reliable experts confirmed that shark fell into the category of cartilaginous and scaly fish" (http://netiran.com:80/news/IRNA/html/950216IRGG13.html) which appears to be so (netiran.com/news/IranNews/html/95021814INPL.html).

Conservation

This shark appears to still be common in Iranian fresh waters judging from the attacks reported over the past 50 years or more and no conservation measures are needed (or likely to be acceptable to the local population).

Further work

The biology of this species in fresh water is unknown for Iran and Iraq and should be thoroughly investigated as a real hazard to those using rivers of Mesopotamia. Are the sharks permanent residents or seasonal visitors? Detailed records of attacks should be kept and analyzed in an attempt to determine any triggering actions. It may prove possible to make recommendations for use of water resources so as to avoid shark attacks in future.

Sources

Garrick (1982) and Compagno (1984, 1988) for general anatomy and biology.

Comparative material: BM(NH) 1874.4.28:9, 1, ca. 76.8 cm total length, Iraq, Tigris River near Baghdad (ca. 33º21'N, ca. 44º25'E); BM(NH) 1924.10.1:1 1, (head only, recorded length 4 ft 1 inch = 1.25 m), Tigris River at Al Karradah near Baghdad (33º17'N, 44º23'E). 

Acipenseridae
 

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The family is found in Europe, northern Asia and North America with 4 genera and 25 species. The Caspian Sea basin contains 2 genera and 6 native species, with both genera and 5 species recorded from Iran. The Caspian population of sturgeons is the largest in the world (Levin, 1997) and Iran is the world's second largest producer of this resource after the former U.S.S.R. (Josupeit, 1994).

These very large fishes are characterised by 5 longitudinal rows of well-developed, bony plates along the body. There is a dorsal row, a lateral row on each side and a ventro-lateral row on each side. In young fish these plates are sharp and obvious but they become smoother with age and may disappear completely. The unpaired fins have fulcra, or flat bony plates, distinct from the scutes, in front of them. Small plates, grains and denticles cover the remainder of the body and the head is covered by large bony plates. Sturgeons have an elongate snout, an inferior protrusible mouth without teeth in adults, fleshy lips and 4 barbels in a row in front of the mouth (see Keys). The vertebral column turns upward at the end into the upper lobe of the tail (known as a heterocercal tail). The first pectoral ray is a strong spine. There are few gill rakers under a single large gill cover. The skeleton is cartilaginous, there is a spiral intestinal valve, 1 branchiostegal ray, fin rays number more than the underlying basal bones which support them, no gular bones on the lower head surface and a large swimbladder. The karyotype may be complex with a very large number of chromosomes, including the very small microchromosomes, and tetraploidy, e.g. Huso huso, Acipenser nudiventris and A. stellatus have 2n about 120 while A. gueldenstaedtii has 2n about 240 and is a tetraploid. Karyotypes of 120 chromosome species are very similar indicating a slow evolution, correlated with a slow rate of DNA and protein evolution. Hybridization is common, even between genera, and hybrids are fertile and used in aquaculture in Russia (Birstein, 1993). Artyukhin (1995) gives a phylogenetic tree of Acipenser and Huso. Krieger et al. (2008) reviewed the molecular phylogeny of the order Acipenseriformes and found Huso not to be monophyletic, among other unusual placements. They conclude that some revision of classification may be needed. Rastorguev et al. (2008) examined mtDNA for Ponto-Caspian sturgeons, although sample sizes were small, and determined various relationships; Huso was basal with Atlantic species and all species in the gueldenstaedtii complex were closely related.

A general overview of sturgeon systematics and biology is given by Williot et al. (1991) and Billard (2002). Artyukhin (2006) and Peng et al. (2007) summarise the relationships aned biogeography of major clades for the order (Acipenseriformes) which dates back 200 MYA to at least the early Jurassic. A bibliography of sturgeons can be found at www.geocities.com/CapeCanaveral/Hall/1345/sturgbibl.html.

Sturgeons are subject to overexploitation, a problem addressed by Lukyanenko (1992), Vadrot (1990), Bemis and Findeis (1994), Faber (1994), Moghim (1994), Anonymous (1995), Asadollahi (1995), Ivanov et al. (1995; 1995, 1999), Vlasenko (1995), Waldman (1995), Birstein (1996), Emadi (1996a; 1996b), DeSalle and Birstein (1996), Hosseinie (1996), Khodorevskaya et al. (1997), Matthews (1998), Khodorevskaya and Krasikov (1999), G. Strieker (in CNN.com, downloaded 9 March 2002), Speer et al. (2000), Raymakers (2002), Oliver (2003), Harrison (2005), Pourkazemi (2006b), Karayev (2006), Raymakers (2006), and numerous newspaper and magazine articles. The problems for sturgeon survival in the Caspian Sea and other waters have been the subject of numerous popular and scientific articles which cannot all be cited here. A summary of the problems and management recommendations are found in De Meulenaer and Raymakers (1996) and The Sturgeon Quarterly published in New York gives recent information. Caspian populations are Endangered (high risk of extinction in the near future - Acipenser gueldenstaedtii, A. nudiventris, Huso huso) or Vulnerable (high risk of extinction in the medium term future - A. stellatus, A. persicus) (De Meulenaer and Raymakers, 1996). In 1997, the Secretariat of the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES) recommended a proposal to list all sturgeons as a species requiring protection because of overfishing and pollution. This would result in the close regulation of the caviar trade and perhaps a trade ban on beluga caviar. Sales of caviar in airport duty-free shops could end as passengers in a hurry would not be able to obtain the necessary CITES export permits or certificates from national authorities. After 1 April 1998 shipped caviar requires export permits or re-export certificates (Traffic North America, 1(3):14, 1998). In the year 2000, western countries through CITES (Convention on International Trade in Endangered Species) gave Iran, Russia, Kazakhstan, Azerbaijan and Turkmenistan until 31 December to impose quotas on their exports in an effort to save the sturgeon stocks. Failure to comply would result in a ban on caviar sales in the west in the year 2002 (IRNA, 25 June 2001). Australia had already banned caviar while the U.K. banned the import of caviar over 250 g without a permit (IRNA, 26 July 2000; The Times, 1 August 2000). Fishing for sturgeon was halted after the spring 2001 season in all Caspian states except Iran which has a well-managed fishery. Fishing quotas will be established after a survey in the summer of 2001 so as to avoid a complete ban on exports (Ottawa Citizen, 19 June 2001, 22 June 2001). By 2004, as Profitt (2004) points out, the agreement had not been fully implemented. Pourkazemi (2006) considers most sturgeon species in the Caspian Sea will be extinct in the near future.

Stone (2002), Stone and Mervis (2002) and Pearce (2003) give details of a dispute between scientists and CITES which arose when fishing for beluga was allowed in 2002. CITES endorsed Russian figures that showed beluga numbers increased from 7.6 million in 1998, to 9.3 million in 2001 and to 11.6 million in 2002. Scientific critics felt that there may well be less than half a million beluga, the differences being based on estimates on how many fish escape experimental trawling in relation to fish actually caught. The United States banned beluga caviar imports on 30 September 2005 and Russia advocated a moratorium on fishing of the major species (Pala, 2005). In April 2006 a global suspension of trade in caviar and sturgeon products by CITES from the Caspian Sea was extended indefinitely, with only one species allowed, the Persian sturgeon from Iran, Iran being the only country that submitted harvest data for assessment of a sustainable fishery (New York Times (www.nytimes.com), 12 April 2006, downloaded 13 April 2006). The export quota for Iran was set at 100,000 pounds of caviar. Bemis and Findeis (1994) recommend gourmets restrict their purchases of caviar to that from fish farms in order to preserve wild stocks of sturgeons.

There was a two-thirds to three-quarters decline in sturgeon numbers in the Caspian from 1990 to 1995, a result of overfishing and poaching. References cited above, The Sturgeon Quarterly (5(1/2):15, 1997) and various newspaper and popular articles reports (e.g. Boston Globe, 8 June 1997 at www.nd.edu/~astrouni/zhiwriter/97/97060808.htm and New York Times, 23 December 1995 at www.nd.edu/~astrouni/zhiwriter/spool/95122301.htm; Tidwell (2001a)) give details about poaching in former U.S.S.R. waters of the Caspian Sea. In 1996, caviar should have sold for £470/kg in Germany but was available for £100/kg illegally (Nuttall, 1996). Caviar imports to the U.S.A. increased by 100% from 1991 to 1996 (DeSalle and Birstein, 1996). The international market demand for caviar was 450 t in 1995 but the legal production from the Caspian Sea was only 228 t; the deficit being made up in part by poaching (Birstein, 1996). Russia officially exported $25 million worth of caviar in 1999 but smuggling of poached caviar was valued at $250 million (Speer et al., 2000). As a result, natural reproduction in the Volga River, the principal spawning ground in the Caspian Sea has been completely destroyed (Birstein, 1996). Bickham (1996) states that it is highly likely that the native sturgeon stocks of the Kura River are extinct or nearly so and Khodorevskaya et al. (1997) simply record that sturgeons no longer use the Kura and Terek rivers. Water pollution was given as the cause for a fall in catch in Iran from 34 tons in 2000 to 9.2 tons in 2004 (Iran Daily, 27 August 2005). Legally traded caviar fell by almost 70% between 1998 and 2003 but illegal sales probably offset this decline (www.canada.com, downloaded 16 December 2005). The export of Iranian sturgeon was expected to drop 20-25% in the year ending in March 2006 (Iran Daily, 25 December 2005). However caviar exports in the 2005-2006 year were given as 18 tons in a later report, still a drastic fall (Iran Daily, 1 May 2006). The caviar export quota for Iran in 2006 stood at 44.3 tons (Iran Daily, 11 September 2006).

Azerbaijan increased the allowable catch from 4 tonnes to 30 tonnes after independence and generally illegal catches made up 90% of all sturgeon caught (Anonymous, 1996a). The yearly allowable catch for Iranian sturgeon in 1996 was 1500 tonnes but the total catch for the Caspian Sea probably exceeds 40,000 tonnes when all countries are taken into account (Emadi, 1996b). Reduction in stocks was noted in assessments carried out in Iranian waters from 1988 onward and the it was decided to reduce the annual catch in 1996 (Iranian Fisheries Research and Training Organization Newsletter, 14:3, 1996). Iran was auhorised to take 90 tonnes of caviar for export in 2000 but the government reduced this to 70 t as a conservation measure (Speer et al., 2000). A restocking programme in Iranian waters cost about U.S.$33 million and a buyout of 4000 fixed gillnetters cost U.S.$10 million (Bartley and Rana, 1998b). Gill nets were trapping young sturgeon, Salmo caspius, Barbus sensu lato spp., Rutilus spp., and Abramis brama.

Sturgeon fingerling production was 9,124,000 in 1995 and 22 million in 1996-1997 according to the above authors, 25 million according to IRNA (2 February 1999), and 12 million according to Abdolhay and Tahori (1999). However pollution causes losses of 40-50 million fingerlings from a production of 108 million, figures at variance with the preceding (Tehran Times, 5 September 1999). The Iranian Fisheries Company produced 88.1% A. persicus in 1996, 5.4% A. gueldenstaedtii, 2.7% Huso huso, 2.5% A.stellatus and 1.3% A. nudiventris (Abdolhay and Tahori, 1999). Keyvanfar and Khanipour (1999) advocate use of trammel nets to catch broodstock for aquaculture as fish are less stressed. TACIS (2002) and Raymakers (2002) give the following table for sturgeon fingerling releases in Iran (in millions):-
 
Species/Year 1990 1991 1992 1993 1994 1995 1996 1997 1998 1999
A. persicus 4.06 5.92 2.93 3.57 4.66 8.05 11.02 18.75 22.59 17.30
A. gueldenstaedtii - 0.04 - - 0.30 0.52 0.67 0.92 0.42 0.96
A. stellatus 0.36 0.47 0.07 0.30 0.46 0.27 0.22 0.29 0.18 0.13
H. huso 0.14 0.17 0.45 0.30 0.49 0.29 0.34 1.44 0.69 0.41
Total 4.56 6.60 3.45 4.17 5.91 9.13 12.35 21.63 24.56 19.10

Abdolhay and Tahori (2006) give descriptions of hatcheries in Iran and the process of fingerling production, including transportation and incubation techniques, pond and tank culture, release strategies, and strategic development plans. Trial production of larvae first occurred in 1922, reaching about 2 million in 1928 but hatchery production first began in 1971. Sturgeon fingerling production was low between 1981 and 1986 as the focus shifted to Chinese carps and Rutilus frisii. Brood stock are captured in rivers by beach seines or selected from fishery stations in February-March. The fish are checked by sampling eggs and  examining germinal vesicle development. Only suitable fish are injected with ovulation-inducing hormones in March-May over 3-5 days. The fish are killed and the collected eggs are fertilised with diluted sperm (1:200 with hatchery water) to avoid polyspermy as eggs have many micropyles. Eggs are incubated in jars or troughs for 5-10 days and newly emerged larvae are held in circular tanks. Fry are raised in fertilised ponds for 40-60 days until they reach 3-5 g. Fingerlings are released in river deltas in June-July. Release strategies are spot planting of all fish at once, scatter planting at several sites in the same region and trickle planting over a period of time. Fish are captured as adults 10-20 years later at a return rate of 1-3%.

Fingerling production in 1000s was:-

Year/Species

H. huso A. nudiventris A. gueldenstaedtii A. persicus A. stellatus Total
1993 301 no data no data 3570 300 4171
1994 491 no data 300 4662 456 5910
1995 286 no data 522 8049 268 9125
1996 344 102 673 11,018 316 12,455
1997 1437 230 919 18,751 288 21,627
1998 687 678 418 22,586 181 24,552
1999 406 304 722 17,300 132 18,864
2000 1901 114 1327 13,711 226 17,279
2001 641 1782 447 16,278 820 19,970
2002 2404 1819 1816 12,301 1300 19,642
2003 42 1414 0 18,388 196 20,041
2004 1464 1311 617 17,412 314 21,121
Total 11,175 7757 7805 191,682 9774 258,567

Iranian sturgeons and their caviar increased in importance in the 1990s as the Russian caviar trade was taken over by a black market system with poor attention to quality. However caviar production in Iran fell in the 1990s through poaching and oil pollution in other parts of the Caspian Sea. Production was 130 tonnes per year, down from 160 tonnes up to 1989 (IRNA, 31 August 1998; Tehran Times, 13 December 1998). Caviar comprises 50% of the seafood exports from Iran (IRNA, 21 October 1998) and formed 1.2% of Iran's total exports for the first four months of the Iranian year in 1998 (in 1994 it was 62% (Salehi, 1999)). On the 23 October 1998, the Islamic Republic News Agency (IRNA) reported that Iran had stopped exporting caviar to protect the resource, this despite the number of sturgeons in the sea having risen from 6 to 22 million over the past couple of years. The same article reports 22 million sturgeon fingerlings stocked in the Caspian Sea by Iran. The export of 111 tons of caviar in 1998-1999 was worth $29.5 million; catches had been reduced to save the species from extinction (Tehran Times, 1999). The export amount was over 80 tonnes since the beginning of the Iranian year (21 March 1999), a 30% drop in production over the previous year (IRNA, 26 January 2000). The 1999 total export was 90 tonnes of caviar worth 70 million deutschmarks, a monetary increase of 42% (IRNA, 4 May 2000). The 2000 export of caviar was 70 tonnes (or 71.5 t, or 80-90 t, reports vary) worth 100 million deutschmarks (or $34.4 million) with 80% going to Europe, 10% to Japan and the rest to various other countries; in addition 100-200 tonnes of sturgeon meat worth $2-3 million is exported annually (IRNA, 14 July 2001, 7 August 2001, 30 September 2001). The sturgeon catch was 75 t in 2002 with 50 t being exported for U.S$30 million (IRNA, 11 June 2003). Golestan Province produced 43% of Iranian caviar, a 17.5% increase presumably in 2000 over the 1999 catch. There are 295 fishermen using 91 fishing boats (IFRO Newsletter, 29:4, 2001).

Sturgeon stocks were evaluated in Iranian waters in 2000 (M. Moghime and F. Parafkandeh Haghighi, 5th International Symposium on Sturgeon, Iranian Fisheries Research Organization, 9-13 May 2005, Ramsar; Haghighi, 2006; Moghime, 2006). The catch was 855 t yielding 92.5 t of caviar, with Acipenser persicus comprising 472 t, A. stellatus 201 t, H. huso 105 t, A. gueldenstaedtii 48 t and A. nudiventris 31.8 t. The catch-per-unit-effort was A. gueldenstaedtii (0.285 kg), A. persicus (2.296 kg), A. nudiventris (0.089 kg), and A. stellatus (2.941 kg). Mature females comprised A. gueldenstaedtii (80.0%), A. persicus (71.8%), A. nudiventris (51.3%), A. stellatus (74.7%) and H. huso (67.4%). The gonadosomatic value in terms of body weight was A. gueldenstaedtii (9%), A. persicus (11%), A. nudiventris (8%), A. stellatus (14%) and H. huso (2.8%). The catch in Gilan and Golestan provinces was A. stellatus (11%) and H. huso (35%) of the total catch. In Gilan, the catch was made up of A. gueldenstaedtii (44.3%), A. persicus (16.4%) and A. nudiventris (3.8%) and in Golestan these values were 16%, 72% and 0.9% respectively. The average age in Gilan and Golestan respectively was A. gueldenstaedtii (15.5 and 18.5 years), A. persicus (20 and 19.2 years), A. nudiventris (15 and 19.3 years), A. stellatus (13.8 and 13.2 years and H. huso (15.8 and 18 years).

Tavakoli et al. (2007)and Kor et al. (2008) surveyed stocks in the southern (2004-2005) and northern (2006)  Caspian Sea respectively. In the southern Caspian Sea catch of 288 fish total, catch per unit effort was 2 fish in summer and 1.38 fish in winter. The most abundant species was Acipenser persicus with 1.67 fish per trawl in summer (142 fish) and 0.88 in winter (75 fish). A. stellatus was 0.22 and 0.48 fish. No Huso huso were caught in winter and only 4 fish in summer. A. nudiventris comprised only 4 fish too and and A. gueldenstaedtii 3 fish, both in total. Kor et al. (2007) examined the population structure of sturgeons in the coastal waters of Mazandaran, less than 10 m deep, for 2003-2005. The number of fish captured in 2003-2004 was 301 with catch per unit effort (CPUE) being A. persicus 4.07, A. stellatus 0.58, A. nudiventris 0.22, and A. gueldenstaedtii 0.15, and in 2004-2005 the catch was 412 fish with CPUE A. persicus 6.15, A. stellatus 0.23, A. nudiventris 0.12, A. gueldenstaedtii 0.35, and Huso huso 0.02.

The world's leading importer of caviar, Caviar House, with an annual turnover of $100 million took 85% of its caviar from Iran (Lindberg, 1994; Pala, 1994). The value of the caviar fishery in Iran was estimated at U.S.$45 million (Bartley and Rana, 1998a; 1998b) and is the main fish product exported with an international cultural and culinary significance. The caviar industry in Iran is a state monopoly under strict control and has not suffered from poaching to the same extent as happened in the former U.S.S.R. after the collapse of central authorities. There has been some smuggling reported via Bandar Abbas to Ras al Khaimah across the Gulf and re-labelling of Azerbaijani caviar as Iranian to fill Iranian contracts with the U.A.E. An illegal trade in "bazaar" caviar reached a peak of 70 tonnes in 1983, about 50% of the legal exports (Taylor, 1997). This caviar was processed poorly in primitive tins with sealant rings made from old tyres; consequently the price for this product was low. The Iranian government actively sought to suppress this trade and after 10 years of effort reduced smuggling to 2-4 t annually, a level similar to that prior to 1979. In 2003 however, 3.8 t of smuggled sturgeon fish and caviar were reported as confiscated for the previous year (ending 20 March) in Mazandaran (IRNA, 21 April 2003). Evidence of Iranian control of the industry is seen in the 1994 setting of a minimum catch size limit of 1 m on all sturgeon species and limiting fishing sites along the Caspian coast to 90 (Josupeit, 1994; De Meulenaer and Raymakers, 1996). Additionally Iran now stocks more sturgeons from farms than it catches (The Times, London, 8 July 1998). However the BBC News (6 May 1998) reports declines in catches of sturgeon over the past 5-10 years.

The illegal market in caviar has been estimated at £500 million with some caviar fetching up to £20,000 a kilogramme (The Times, 28 December 2006). In Britain, caviar tins must indicate their exact source and without this label will be seized by Customs. The label will carry a species code, source of the caviar, country code, year of harvest, processing plant registration number and lot identification number, all in an attempt to regulate and eliminate sales of smuggled caviar. Much of the smuggled caviar is sold under the counter or to those who have pre-ordered it, or by shops that then state they were unaware of its illegal status.

Although caviar is the main market item for sturgeons, Iran is investigating the use of fillets and smoked and salted A. stellatus in vacuum packs for export (Annual Report, 1995-1996, Iranian Fisheries Research and Training Organization, Tehran, p. 45-46, 1997). Smoked, marinated and canned sturgeon, smoked sturgeon in vegetable oil and frozen fillets are now available (2001) from several Iranian companies. Javanmard and Taghavi (2002) investigated the microbiological and chemical characteristics of these products and only one had a total coliform count more than the European Community standard. Gelatin has also been produced from sturgeon fish skin on an experimental basis (Iranian Fisheries Research Organiztion Newsletter, 30-31:6, 2002; Koochakian Sabour et al., 2001).

All sturgeon species in the Caspian Sea basin are listed as "endangered" or "vulnerable" and are maintained in part by hatchery stocks (http://www.sturgeons.com/htdocs/status.html). Survival and growth of sturgeon fry in the Caspian Sea is reviewed in Farsi by Aslaanparviz (1992).

The countries of the Caspian littoral are attempting to conserve their sturgeon stocks. Even the Swiss company Caviar House has established a hatchery in Iran to increase stocks (Anonymous, 2001a). An agreement for the "Preservation and Exploitation of Live Resources in the Caspian Sea" was made between Iran, Russia, Azerbaijan, Turkmenistan and Kazakhstan in 1996. Luk'yanernko et al. (1999) point out the need for the agreement to recognise that sturgeons are sustained by an ecosytem involving the whole Caspian Sea and the inflowing rivers, that there must be an absolute ban on uncontrolled fishing for sturgeon in the sea and that national quotas must reflect the real contribution of a a particular state to overall sturgeon stocks. Without adequate measures, these authors predict extermination within 5-7 years. Export of caviar is made a monopoly of the governments concerned in an effort to minimize smuggling of low quality caviar. Jenkins (2001) gives reasons why an international trade ban would not necessarily help conserve the sturgeons - most poached caviar is sold within Russia, for example. Sturgeon catches are restricted to rivers and their estuaries and open-sea trawling is banned. The five countries are investing $150 million in a fish farm programme to save the sturgeon from extinction: Russia will have 10 new farms and renovate 8 farms on the Volga River, and both Kazakhstan and Iran have a new farm (Abzeeyan, Tehran, 7(4):II-III, 1996; The Sturgeon Quarterly, New York, 4(4):1, 1996; newspaper reports). Russian strategies for conservation of sturgeon are reviewed in Artyukhin et al. (1999) and the status of the Russian sturgeons is given in Vaisman and Raymakers (2001). Some sturgeon species are now on Appendix 2 of the U.N. Convention on International Trade in Endangered Species of Flora and Fauna (CITES) in an effort to control the import and export of meat and caviar (Pearce, 1997). The U.S. Fish and Wildlife Service, in an attempt to combat overfishing of sturgeons, now requires valid CITES permits for imported caviar (Anonymous, 1998a). DNA tests will be used to confirm the species of sturgeon listed on the shipment and to eliminate illegal mixtures with inferior quality roe. Even cats are now used to detect smuggled sturgeon in Russia. A cat named Rusik is able to detect sturgeon hidden in trucks better than sniffer dogs (National Post, 9 July 2003, p. A12).

The number of adult fish in the Caspian Sea had declined from 142 million in 1978 to 43.5 million fish in 1994. Ivanov et al. (1999) and Khodorevskaya and Krasikov (1999) review the status of stocks in the Caspian. Marked declines are evident and only the Iranian catches are reasonably stable from 1977 to 1994. All species studied in Iranian waters had a very low percentage of fish older than 20 years, are evidently in need of protection (Iranian Fisheries Research and Training Organization Newsletter, 16:4-5, 1997). An initiative to make the sale of caviar from threatened sturgeon species illegal is being proposed by the Species Survival Commission and the IUCN (The Sturgeon Quarterly, New York, 4(4):1, 1996; Morris, 1997). Part of this initiative would involve genetic testing of the caviar as a means of identifying the species of sturgeon. Paddlefish eggs from Montana, U.S.A. costing less than $5 an ounce have been repackaged as beluga caviar in Russia and eastern Europe and sold in the U.S.A. for $50 an ounce. The U.S. Fish and Wildlife Service was to begin monitoring the caviar trade on 1 April 1998 using DNA tests (U.S.A. Today, 18 November 1997, internet edition). Birstein et al. (1998) describe a molecular technique for identifying the species source of commercial caviar (see also Brainard (1998)). They found 23% of species designations by caviar suppliers to be incorrect, indicating possible illegal harvesting and poaching. The Iranian Fisheries Research and Training Organization Newsletter (20:4, 1998) also reports on nuclear DNA amplification and a marker which distinguishes species. Additional research is being carried out on egg identification using ultrastructural characteristics (L. Debus and M. Winkler, 1998, www.uni-rostock.de).

Sturgeons have been fished since the Neolithic, perhaps 6000 or more years ago (Tsepkin, 1986) but only in recent years have the stocks declined significantly. Historical records show it was possible to catch 500 Huso huso weighing 600-1000 kg in about 2 hours in the Volga delta at the end of the eighteenth century (Birstein, 1993). All the sturgeon species were bigger on average and lived longer than now based on archaeological excavations (Tsepkin and Sokolov, 1971). The sturgeon catch in the Caspian Sea declined from 27,400 tonnes in 1977 to 8,900 tonnes in 1990 (Vlasenko, 1995). The catch in 1993 was only 4,200 tonnes because of poaching and pollution of the Volga River (The Sturgeon Quarterly, 3(1):12, 1995). An estimated 90% of the Caspian sturgeons are killed before they mature (Platt, 1995). Catches in Russian waters of the Caspian Sea declined from 7106 tonnes in 1992 to 3426 tonnes in 1993 to 2960 tonnes in 1994 but 90% of the real catch is unreported (16,700 tonnes were reported in 1983 for comparison). The number of adult sturgeons in the Caspian Sea is estimated to have declined from 142 million fish in 1978 to 43.5 million fish in 1994 (De Meulenaer and Raymakers, 1996). The Caspian Sea Sturgeon Ranching Programme of the former Soviet Union helped to sustain fisheries but declines still occurred (Secor et al., 2000).

Catches in Iran, however, increased over a five year period, perhaps because of heavier fishing pressure. Sternin and Doré (1993) cite figures for 1986-1990 of 1690 tonnes, 1759 t, 1851 t, 2051 t and 2021 t, while U.S.S.R. catches over the same period were 21,817 t, 20,991 t, 19,027 t, 16,880 t and 15,056 t. A conflicting study noted a decline from 122,000 sturgeons caught in 1986 to 68,000 in 1993 (Abzeeyan, Tehran, 6(5, 6):IV-V, 1995). De Meulenaer and Raymakers (1996) summarise Iranian catches as 700 to 2500 t in the twentieth century, peaking towards the end of the 1960s, falling to 1000-1500 t in the 1970s and increasing from 1450 t in 1982 to a 1991 high of 3036 t but falling off rapidly to 1700 t in 1994. Josupeit (1994) gives catches in Iran in tonnes from 1982 to 1992 as 1450, 1288, 1557, 1650, 1690, 1759, 1851, 2051, 2645, 3036 and 2692 t. The commercial sturgeon catch in the Safid River delta fell from 6700 tons in 1961 to less than half a ton in 1993 (http://www.oneworld.org/patp/pap_overview.html). Spawning may no longer take place in the Safid River (De Meulenaer and Raymakers, 1996). Zanusi (1995) maintains that over 40% of the total sturgeon fishing in the Caspian Sea is centred on Bandar-e Torkeman in Mazandaran, presumably including the acknowledged black market in sturgeon products. Lewis (1980) gives some information about the Iranian black market in caviar shortly after the Islamic Revolution before controls were re-established. A 400 g tin was selling in Paris black market for $40 compared to $310-315 for the best Russian beluga. Caviar production in the three Caspian coast provinces of Iran for the 1990s were as follows in kg after Nezami et al. (2000):-
 
Year/Province Gilan Mazandaran Golestan
1991 75,974 78,713 128,446
1992 81,520 80,758 99,336
1993 51,480 58,543 83,026
1994 40,368 52,162 87,576
1995 37,241 43,831 70,154
1996 41,743 41,432 79,063
1997 28,641 42,329 58,304

The problem of overexploitation of sturgeons is compounded by their long life span and their use of rivers as spawning grounds such that they are easily caught on this migration from the sea. The migration and spawning is timed differently between species and populations within species. Some sturgeons migrate long distances up rivers while others have a shorter migration. Eggs are deposited on stony or gravel bottoms and hatch after a short incubation. As an example, a study of sturgeon migrations in the Gorgan and Tajan rivers of Iran showed a movement of 2 out of 28 fish caught at one station reached the second station in the Gorgan and no tagged fish reached higher stations in the Tajan - the rest were caught by fishermen (Annual Report, 1995-1996, Iranian Fisheries Research and Training Organization, Tehran, p. 53, 1997). Ramin (1998) studied migration in the Safid River over 35 days in April-May, from the mouth to 30 km upriver for A. gueldenstaedtii, A. persicus and A. stellatus. Shallow water caused by sand-clay deposits and illegal fishing did not prevent successful migration. It was recommended that the Manjil Dam be used to regulate water flow and a total ban on fishing, especially at the mouth, during the March-May spawning season be implemented.

In 1998 the comb jelly, Mnemiopsis leidyi, reached the Caspian Sea via ship ballast and newspapers speculated that the sturgeon populations would be affected, although how was not specified.

The young migrate downstream to feed and grow in the sea. Old reports have sturgeons overwintering in deeper parts of rivers, in a kind of torpor and with a viscous substance coating the body (Baird, 1873). The barbels are highly sensitive and, as soon as they detect food, the tubular mouth protrudes to suck in the prey. Food is benthic organisms although some are predators on larger fishes. Young sturgeon in Iran feed predominately on polychaetes while crustaceans are a minor food item, probably caused by lower oxygen conditions favouring the former (Haddadi Moghaddam and Negaresten, 2003). Pourgholam (1994) reports the coelenterate Polypodium hydriforme from sturgeons caught on the Babol Sar and Bandar-e Torkeman fishing grounds in Mazandaran where up to 25.6% of fishes are infected, particularly Huso huso and Acipenser gueldenstaedtii. This parasite destroys the eggs of sturgeons, affecting reproductive success and the caviar industry (see also Raikova (2002)). Incubated eggs of sturgeons are susceptible to various species of fungi, with up to 70-90% of eggs being lost (Czeczuga et al., 1995). Czeczuga et al. (1995) report 43 species of fungi on eggs of sturgeons from Russian and Iranian Caspian Sea samples immersed in water from a Polish river, lake and pond. Huso huso and Acipenser gueldenstaedtii persicus (sic) eggs carried the fewest species of fungi, about half the load of other sturgeon species. Ghoroghi (1996) reports metacercariae of Diplostomum spathaceum in the lens of 22% of fingerlings on the Shahid Beheshti Fish Farm causing weight loss and mortality. External parasites on sturgeons include Pseudotracheliastes stellatus, Nitzschia sturionis, Diclybothrium armatum, Cystoopsis acipenseris and Diplostomum spathecum with the highest prevalence in Huso huso at 60% and the lowest in Acipenser persicus at 13.9% (A. Hajumoradloo in 5th International Symposium on Sturgeon, Iranian Fisheries Research Organization, 9-13 May 2005, Ramsar). Ghaemi et al. (2006) found strains of mycobacteria in Iranian sturgeons and Mycobacterium marinum can cause fish tank granuloma, a disease in humans although none was found in fishermen.

Many sturgeons in former Soviet waters of the Caspian Sea have developed fatal diseases associated with chemicals such as phenols, waste fluids and air from gas production facilities associated with the petrochemical industry. Both the sturgeon and their caviar are now inedible. Iranian sturgeons are believed to be less affected but since sturgeons migrate they are susceptible to extra-territorial pollution (Golub, 1992).

Sturgeons are some of the most important commercial species in the world, with 90% of the total catch coming from the former U.S.S.R. and only 6% from Iran (but see later under Acipenser gueldenstaedtii where Iranian production of caviar increased in the 1990s). Over 90% of all sturgeons are caught in the Caspian Sea. The proportion of catch is heavily weighted towards the former U.S.S.R. (compared with Iran in parentheses) with figures from 1971 to 1988 as given by Sternin and Doré (1993) being 19,100 tonnes (2400 t) for 1971, 20,400 t (2200 t) for 1972, 24,958 t (1801 t) for 1978, 26,322 t (1578 t) for 1979, 26,697 t (1429 t) for 1980, 26,452 t (1496 t) for 1981, 25,704 t (1450 t) for 1982, 25,570 t (1500 t) for 1983 and 18,470 t (1700 t) for 1988. The Volga River and its delta provided 75% of the commercial sturgeon harvest in the Caspian Sea with Acipenser gueldenstaedtii making up 60-70% of this amount, A. stellatus about 30% and Huso huso 5-6% (Khodorevskaya et al., 1997). Williot and Bourguignon (1991) summarise sturgeon catches in Iran from FAO data for 1965 to 1987 as ranging from a low of 1429 t to a high of 3000 t. Abdolhay and Tahori (2006) summarise catches as follows:-

 
Year Total catch (tonnes) A. stellatus (%)  Osetra* (%) H. huso (%)
1972 1500 34.0 36.3 29.7
1991 3036 49.5 41.0 9.5
1994 1700 49.5 41.0 9.5
1997 1300 35.8 54.3 9.9
2000 1000 35.8 61.0 3.5
2001 870 28.2 69.3 2.5
2004 600 10.7 74.6 14.7

* presumably includes A. persicus and A. gueldenstaedtii

Only 5% of Iranian caviar is consumed in that country, the rest being exported. Domestic prices are very high at about U.S.$340 per kilogramme and the caviar is rationed to 100 g per person. Contraband caviar can be bought at about half this price around Bandar Anzali and at least 30 t are smuggled out of the country each year (The Daily Star, 7 December 2004, www.dailystar.com, downloaded 17 December 2004). In 1996, 95 t out of an estimated 120 t catch was exported although formerly as little as 38.7% was exported as in 1978. Iran is the chief exporter to the European Union, the weight varying from 95 to 125 t from 1988 to 1994 (De Meulenaer and Raymakers, 1996). Caviar exports by year for Iran are given by these authors as:-
 

Year

1988

1989

1990

1991

1992

tonnes

225

249

226

225

169

U.S.$1000

42,155

47,865

46,005

53,800

42,004

U.S.$/kg

187

192

204

239

249

The export volume of caviar for 1997-1998 was 105 tonnes worth 62 million German marks (= U.S.$34 million) (IRNA, 16 March 1998).

Prices outside Iran have fluctuated widely because of large amounts of illegal and often poor quality caviar flooding the world market. Caviar exports are declining in the 1990s reflecting, it is believed, the loss of sturgeon stocks in the Caspian Sea (De Meulenaer and Raymakers, 1996).

Caviar is the main product but the flesh is also eaten (a religious ruling was made in the 1980s to the effect that Iranian ichthyologists had determined sturgeons to be fish with scales - see below). Early reports of poisoning from sturgeon eggs have been attributed to poor preservation and consequent bacterial contamination (Halstead, 1967-1970). The milt of Acipenser sturio contains a toxic substance known as "sturin" and although this species does not occur in Iran a similar toxin may occur in Iranian Acipenser (Coad, 1979b).

The swimbladders of sturgeons have been converted to isinglass, a transparent gelatin used in a variety of products including as a wine and beer clarifier, in jams and jellies and in glass and pottery. Gmelin (2007) mentions that in 1770-1774 people along the Langerud were catching large numbers of sturgeons for their isinglass only, the caviar and flesh not been used. Sabour (2006) found the swimbladder in Iranian sturgeons to weigh 250-285 g in H. huso, 35-92 g in A. stellatus and 85-160 g in Acipenser spp and could be was processed to isinglass at 15-20%. Koochekian et al. (2006) found a higher percentage production of isinglass from A. persicus/A. gueldenstaedtii than in Huso huso or A. stellatus. Recently Iranian scientists have investigated production of leather from sturgeon skin (Iranian Fisheries Research and Training Organization Newsletter, 4:2, 1994; Davarzani, 1995; Iranian Fisheries Research Organization Newsletter, 22:2, 2000; Iran Daily, 17 January 2006). An estimated 1 million square feet of leather could be produced and used in handicrafts, book binding, waterproof products and ornaments.

Various methods to enhance the sturgeon fisheries have been investigated in Iran. Some are given under the Species Accounts and others are summarised here. Experiments with pen culture in Gorgan Bay have been carried out to increase production and with cross-breeding Huso huso and Acipenser stellatus to create new commercial and resistant stocks (Iranian Fisheries Research and Training Organization Annual Report, 1992-93). A Farsi review of sturgeon culture is given by Rasoli (1992). Even surgical procedures under anaesthesia have been tried to remove eggs through a 15-20 cm incision as part of attempts to increase caviar production (Mokhayer, 1993; The Times, London, 8 July 1998). Ultrasonagraphy has been used successfully to distinguish males, females and immature fish without damaging them (Vajhi, Moghim, Veshkini and Masoudifard (1999) www.mondialvert99.com, downloaded 31 May 2000; Moghime, 2006). The accuracy was 97.2% for A. stellatus and 100% for A. gueldenstaedtii, A. nudiventris and H. huso. Vacuum pumps have also been used to breed female Acipenser nudiventris and male A. stellatus. The fish are anaesthetized with xylazine hydrochloride and then eggs and sperm are pumped out, the advantage being that females can be returned alive to the sea (Iranian Fisheries Research and Training Organization Newsletter, 13:5, 1996). Bahmani et al. (2001) compared haematological parameters in Acipenser persicus and Huso huso and how these changed with age. Haemtaological indices give insight into the physiological condition and aid in the selection of broodfish. Cryopreservation of sperm has been carried out as stripping fish late in the season is difficult. Sperm in liquid nitrogen with an extender is viable for 1.5 to 2 years (M. Moghim and H. N. Moghadam in 5th International Symposium on Sturgeon, Iranian Fisheries Research Organization, 9-13 May 2005, Ramsar; Moghadam, 2006). The colour of gill nets used in the capture of sturgeons has been investigated with blue nets having a yield of 42.6%, white 29.8% and green 27.5% (5th International Symposium on Sturgeon, Iranian Fisheries Research Organization, 9-13 May 2005, Ramsar). Studies on the fingerling production of hatcheries include the nature of the phytoplankton community and the benthic biomass, parasitic infections (e.g. Diplostomum sp. on the eyes and Trichodina sp. on the gills were noted at an incidence of 25% and 35.85, productivity (6,509,185 fingerlings produced from 31 March and 28 July 2000 in two hatcheries with some transfer of Huso huso fingerlings from another hatchery), survival rates (56.7% and 25.2% for A. persicus and H. huso respectively), and growth rate and condition factor (generally low). Kami et al. (2005) studied the biology of pond turtles (Emys orbicularis) which live in culture ponds along with sturgeon. One dietary item was Acipenser persicus. The use of probiotics, microbial cells in the diet, used to improve health and thus enhance quality of farmed fish is of potential use in sturgeons as reviewed by Askarian et al. (2006). Bahmani (2006) used both histology and haematology on Acipenser persicus, A. gueldenstaedtii and Huso huso to determine physiological condition  of fish in ponds and rearing tanks, comparing the results with natural conditions (similar) and finding that fibreglass tanks were more suitable than rearing ponds. Banadani (2006) examined the environmental conditions in the Gorgan River, a major site for release of sturgeon fingerlings. Mohseni (2006) studied the effect of stocking density of eggs and larvae in incubators on their survival, growth and appearance of deformities. Increased density reduced survival and growth and increased deformities. Parandavar (2006) compared production of sturgeon from broodfish maintained on farms to those produced from fish taken from the wild. Salehi (2006) analysed the economics of sturgeon fingerling production and found labour costs were 55%, food and fertiliser 14%, maintenance 7% and fertilised eggs 5%. A single fingerling cost 992 rials to produce in Iran, varying between 447 and 1224 rials among hatcheries. Yousefian (2006) gives details of the production of fingerlings at the Shaid Rajii Fish Farm in 2002. This farm produced 2,898,086 or 93.27% of the fingerlings released into the Tajan, Larim, Goharbaran and Sardab rivers. These fingerlings had an average weight of 3.58 g and condition factors were 0.4 for Acipenser persicus, 0.37 for A. gueldenstaedtii and 0.31 for A. stellatus, in total and average grade for the condition factor. Fazlei (no date) summarised the number and quality of fingerlings released into Mazandaran and Golestan provinces. The most important rivers for release were the Gorgan (8,659,377 fingerlings, average weight 2.55 g), Tajan (1,453,410, 4.12 g), Larim (1,211,875, 3.4 g) and Goharbaran (743,561, 3.09 g). A. persicus comprised 87.7% of the fingerlings, A. gueldenstaedtii 6.6%, H. huso 3.3% and A. nudiventris 2.4%. The International Sturgeon Research Institute has developed a food formula based on Iranian sturgeon species. Previously food for aquaculture came from Europe and the domestic version was demonstrated to be superior (Iranian Fisheries Research Organization Newsletter, 58 & 59:2, 2009). The hybrid sturgeon known as bester (female beluga x male sterlet) has been investigated  for expanding sturgeon culture in Iran. Growth was significantly better than in beluga (Iranian Fisheries Research Organization Newsletter, 58 & 59:4, 2009). Jafarian et al. (2009) studied the use of probiotic bacilli to encapsulate Artemia urmiana nauplii and the yeast Saccharomyces cerevisiae used to encapsulate Daphnia magna, Artemia and Daphnia being used as live food for sturgeon larvae. Both treatments increased growth parameters and feeding efficiency in A. persicus, A. nudiventris and H. huso.

Lake Orumiyeh has been used as a source for Artemia urmiana or brine shrimp to be used as a live food in sturgeon aquaculture (Azari Takami, 1987; 1993). Brine shrimp were found to be a better food than white worms or Daphnia, being cheaper and easier to prepare, easier to store as cysts, sturgeon fry showed better growth, pathogens were less, mortality was lower and yield higher. Since 1972 almost 50% of fry diet has been brine shrimp. The large mouths of sturgeon fry enable them to take brine shrimp nauplii and even adults a few days after yolk-sac absorption. Fry are grown to 100-120 mg within 7-10 days and then released into the sea.

Anonymous (1961b) reported on the caviar industry in Iran which at that time was about 5-6% of the world supply. The Food and Agriculture Organization of the United Nations in their Yearbook of Fishery Statistics reported catches of sturgeons from 1980 to 1985 as 1429, 1496, 1450, 1288, 1557 and 1650 tonnes respectively. Soviet catches of sturgeons in all waters, not just the Caspian Sea, ranged from 22,772 to 26,697 tonnes for the same period. Petr (1987) summarised FAO statistics for Iran and gave mean landings of sturgeons as 2300 tonnes (1964-1970), 1800 t (1971-1975), 1500 t (1976-1980), and 1774 t (1980-1985) but some of this data is very approximate being repeats of a 1500 t value as an estimate (see also above for more figures). A pamphlet from the Ministry of Jahad-e Sazandegi (= Construction Crusade or Rural Development), which is charged with fisheries in Iran, gave catches for "caviar fish" of 3036 tons (presumably tonnes) in 1991 and 2692 tons in 1992. The catch in 1995 was 995 tonnes yielding 134 tonnes of caviar with 74% of the catch from Mazandaran province (http://netiran.com:80/news/TehranTimes/html/95122503TTEC.html). Other news reports give the 1995 catch as 142 tonnes of caviar, in 1996 112 t and an estimated 140 t in 1997. The 200 t of caviar produced in 1992 was worth $100 million through export while the 1997 catch was worth only $60 million despite a 50% increase in price. The Tehran Times (30 May 1998) reported that caviar production was reduced from 220 tonnes to 40 tonnes during the previous 6 years to preserve stocks. The allowable catch in 2003 was set at 676.4 t for Iran, a decrease from 685 t in 2002, with caviar exports set at 78.8 t. Figures for other Caspian states were Azerbaijan 130 t (9.1 t of caviar), Russia 429 t (30.3 t), Kazakhstan 216 t (23.18 t), and Turkmenistan 56.25 t (5.85 t)(IRNA, 28 December 2002).

Caviar from Iran commanded a higher price than that from the former U.S.S.R. in the 1990s (Christie, 1995). Catches in the 1952-1957 period yielded an annual average yield of 120 tons (sic, possibly tonnes here and below) of caviar (Kayhan International, 1 December 1962) which agrees closely with the figure given by Job (1961a) of 90-115 tons (sic) annually. Catches from 1965/66 to 1968/69 in Iran rendered 208 to 219 tonnes of caviar annually from 1996 to 2290 tonnes of the three main species fished (A. gueldenstaedtii (presumably including A. persicus), A. stellatus and Huso huso)(Andersskog, 1970). The catch in 1961-1962 was 170 tons (or 178 tons, V. D. Vladykov, in litt., 1966; differing data is not unusual as effectiveness of information gathering varies) and this was the first season when exports to the U.S.A. exceeded that to the former U.S.S.R., 56 to 46 tons, with 58 tons going to Europe and about 10 tons consumed locally. The caviar yield in 1956-1957 was low, at 134 tons (or 137 tons, V. D. Vladykov, in litt., 1966) and averaged 120 tons from 1952-1957 (Kayhan International, 1 December 1962), a decline over levels prior to dissolution of the Iran-Soviet company. White (1988) reported a caviar export of 150 tonnes from Iran with a value of U.S.$20 million out of a 250 tonnes annual production. Caviar yield in airtight containers was 233 tonnes (1981), 204 t (1982), 222 t (1983), 247 t (1984), 304 t (1985), 283 t (1986), 296 t (1987), 281 t (1988), 286 t (1989), and 290 t (1990) (Sternin and Doré, 1993). Production of caviar in the 1990's dropped steadily from 160 tonnes to 120 tonnes as the Caspian became more polluted (Tehran Times, 5 August 1999) and the catch for the year ending in March 2000 was expected to be less than 100 tonnes (Reuters News Service, downloaded 1 September 1999). Pollutants from Russia, Azerbaijan and Kazakhstan include oil spillage from old equipment at offshore sites and 12 million cubic metres of sewage from the Volga. The sewage includes toxic PCBs, phenol, heavy metals, dioxins and DDT as well as household, agricultural and industrial wastes. A 10-year ban on sturgeon fishing would have to be placed into effect to allow stocked sturgeon to mature and breed. Research on qara burun (A. persicus) and uzun burun (A. stellatus) in Iran has shown heavy metal (cadmium, copper, zinc, lead and mercury) density in caviar and flesh to be 10 times less than the global safety standard (IRNA, 15 January 2002; IFRO Newsletter, 28:2, 2001). Pourang et al. (2005) examined all five sturgeon species in Iranian waters and found all toxic trace elements (Cd, Cu, Pb and Zn) to be markedly below international guidelines for human consumption. Kajiwara et al. (2003) demonstrated contamination by organochlorines in Iranian sturgeons. DDT and its metabolites predominated at 180-18,000 ng/g on lipid weight followed by PCBs at 110-1900 ng/g. Generally Huso huso was the most contaminated species and contaminant concentrations were higher in Azerbaijan and Kazakhstan than Iran, the latter having fewer oil wells. Gelodar (2006) evaluated four caviar processing plants for their hygienic standards using the Hazard Analysis and Critical Control Point (HACCP), an internationally recognized food safety system. Those plants following the European Community code had decreased their contamination levels.

70% of Iranian caviar is produced in Mazandaran, 130 tonnes in 1994 (Abzeeyan, Tehran, 6(5, 6):III, 1995) although this conflicts with a report from IRNA for 2 May 1998 where Mazandaran has 35% of the total Iranian output at 44 tonnes for 1997-1998. 95% of the Mazandaran caviar is exported along with 60 of 260 tonnes of sturgeon meat (IRNA, 2 May 1998). The Bandar-e Torkman fisheries organization in Golestan Province (eastern Caspian Sea) planned to process 360 tonnes of sturgeon and 48 tonnes of caviar in 1999-2000 (IRNA, 14 December 1999). Newspaper reports in 1995 gave a value of U.S.$40 million for caviar exports from Iran; another report gave U.S.$50 million for 250 t (Food and Agriculture Organization, Fisheries Department, 1996). This is less than the value of half a day's oil sales but the caviar fishery is a national symbol (Christie, 1995). Mazandaran produced 17 tons of caviar in 10 months in 2003-2004 as well as 140 tons of meat (www.iranmania.com, downloaded 4 October 2004). In the Iranian fiscal year ending 20 March 1998 Iran exported 105 tonnes of caviar worth about U.S.$11 million (Anonymous, 1998b). The 2003 allowed share for Iran was 78.8 t from a catch for the whole Caspian Sea of 148 t (IRNA, 22 September 2003). The quota for all Caspian caviar in 2004 was 125 tons (www.nytimes.com, downloaded 12 October 2004).

The sturgeons were little used after eggs were extracted for caviar although they were sometimes served in small restaurants along the Caspian coast (personal observations; remarkably tough and tasteless too!). Sturgeons were "haram" in Iran, forbidden for religious reasons as scaleless fish although this has been reversed (Caddy, 1984; Anonymous, 1989; saffron, 2002). Most flesh was exported to Russia (RaLonde and Walczak, 1970b) although some is dried and pickled for local consumption (De Meulenaer and Raymakers, 1996) or eaten freshly grilled (personal observations). Fraser (1834) noted thousands of sturgeon carcasses lying on the Safid River banks, discarded after removal of eggs for caviar and swimbladders for isinglass. Export prices in 1995 ranged from $5.00 per kg of fresh sevryuga fillet to $14.50 per kg for smoked beluga (fil mahi) fillet (Abzeeyan, Tehran, 5(9):V, 1995). Shilat now markets sturgeon head-on or headless, gutted, frozen or in any form required by customers. The average processed weight is about 20 kg for Huso huso, 8 kg for Acipenser gueldenstaedtii (probably includes persicus) and 6 kg for Acipenser stellatus. The meat is served roasted or smoked (Shilat advertisement in Seafood International, December 1995). Research has been carried out in Iran on products derived from left-over parts of sturgeons, the intestines for fish sauce, and skin for gelatin (Sabour et al., 2006).

Capture methods, in the early twentieth century, involved large iron-barbed hooks attached to ropes stretched across the river mouth to foul-hook the sturgeon or, further upstream, poles 6-8 feet long armed with an iron hook used to gaff the sturgeon (Fortescue, 1920). Sturgeons may be caught more recently by large shore seines but mostly they are taken by gill nets set 1-3 km out to sea although De Meulenaer and Raymakers (1996) refer to 300 m fixed nets in rivers with passage space at the sides and bottom as the only authorised method in Iran. Trawling in the sea is not allowed in Iranian waters. Sturgeons are taken from March to June and from September to November in each year (Christie, 1995). The autumn season is best for A. gueldenstaedtii (and presumably A. persicus) while the spring season is best for Huso huso and A. stellatus (De Meulenaer and Raymakers, 1996). Autumn is the main season when the sturgeons migrate to the southern Caspian Sea (De Meulenaer and Raymakers, 1996). The draft "Agreement on the Conservation and Utilization of the Biological Resources of the Caspian Sea" in 1995 prohibited sturgeon fishing in the open Caspian Sea except for traditional methods by Iran near its coast within quota limits (Vinogradov in Glantz and Zonn, 1997).

Gill nets used to capture bony fishes, mainly cyprinids, are responsible for an increase in malformations observed in sturgeons in recent years (Mehdizadeh, 1993). Fins are broken or cut, rostrums (snouts) deformed and net fragments embedded in flesh. Gill nets were prohibited in the Caspian Sea off Iran, except for sturgeons, but during the Iran-Iraq War economic necessity brought back gill netting for bony fishes and cooperatives were established (Habibnejaad, 1993). Gill netter cooperatives were changed to kilka or beach seiner cooperatives and by the end of 1993 no gill nets were allowed in the Caspian Sea. However it took 12 years to overcome objections to banning gill nets by fishermen and in parliament. Problems with excess mortality through inappropriate fishing methods are not new. In the period 1925-1930 the total length of long-lines used in the Caspian Sea was 7-8,000 km while sturgeon nets exceeded 10,000 km. Many fish died in unattended nets or tore lose from long-lines, later to die from hook injuries (Sternin and Doré, 1993). The prohibition of the use of gill nets with a less than 12 cm mesh in 1994 by Iran has conserved stocks along the southern coast of the Caspian. Additionally licenses were restricted and fishing co-operatives closed down in order to control the take (Raymakers, 2002).

Iranian fisheries have taken place mainly in the sea and so a lot of immature fish are caught whereas the former Soviet fisheries took place in rivers where only adults were taken (and ideally could be controlled more easily). However state control in Iran has meant, as noted above, better control over the fisheries and more effective conservation, although poaching does occur.

An attempt has been made to raise sevryuga sturgeon in the central Iranian desert 100 km southeast of Yazd (www.iranmania.com, downloaded 13 March 2003 and other news reports) in a 5000 sq m artificial pond, perhaps more an indication of the desperate straits of the sturgeon populations than anything else.

Pourkazemi et al. (2000) examined the phylogenetic relationships of the 5 sturgeon species in Iran using mtDNA. Huso huso and Acipenser nudiventris showed a close evolutionary relationship as did A. gueldenstaedtii and A. persicus. The latter two species apparently diverged about 1 MYA. Birstein and DeSalle (1998) using molecular techniques found that the Ponto-Caspian species of sturgeons dispersed through the Black, Azov, Mediterranean and Aral seas during the Pleistocene 1.5 MYA and later, the A. stellatus-A. persicus lineage originated 6.0-5.5 MYA in the Upper Miocene-Lower Pliocene, the A. gueldenstaedtii lineage and the Ponto-Caspian sturgeons originated 15 MYA in the Middle Miocene, Acipenser originated and diverged 95-65 MYA in the Upper Cretaceous, and Acipenseridae diverged from Polyodontidae, a related family, 200-135 MYA in the Jurassic.

An important, recent literature source on Caspian sturgeons is Holčík (1989) as well as specific works on Iranian sturgeon biology and fisheries by Rostami (1961b), Vladykov (1964) and Mobayen (1968). The fisheries information in these last three works, relating to techniques and stations, is somewhat dated and not detailed here (Raymakers (2002) gives a map showing Iranian fisheries stations). A general account of sturgeons is given in Birstein et al. (1997) and in Hochleithner and Gessner (1999). Billard (2000) is a recent review of reproduction and associated methodologies used on fish farms. CITES (2001) gives an identification guide in English, French and Spanish, with numerous pictures and diagrams. Pavlov et al. (2001) review the types of spawning migrations carried out by sturgeons. There are numerous other popular reports and scientific papers on Caspian Sea sturgeons, not all of which can be cited or analysed here. A Bibliography of Sturgeons is given by Y. Keivany and V. J. Birstein at www.geocities.com/keivany/sturgbibl.html. Various manuscript reports on the biology and rearing of the economically important sturgeons have appeared in Farsi, e.g. Abdolhay (1997), Abdolhay and Baradaran Tahori (1998), Baradaran and Abtahee (1998), Fadaee (1997), Kohneshahri and Azari (1974), Moghim et al. (1996), Nasrichari (1993), Pourkazemi (1996), Shafizadeh and Vahabi (1996), etc. Regular symposia on sturgeons are held and extensive presentations and publications result, e.g. The 5th International Symposium on Sturgeons, papers from it being published in Journal of Applied Ichthyology 22(s1)(2006). These works have not all been summarised here because of expense in obtaining copies and because many papers refer to details of aquaculture physiology and biochemistry.

A general Farsi name for these sturgeons is سگ ماهي (sag mahi = dog fish).

Caviar
خاویار

Further information on the catches of sturgeons and production of caviar in Iran can be found in the Species Accounts below. Iran is the second largest producer of caviar, after Russia, with 20% of the world market valued in excess of $50 million (Khajehpour-Khouei, 2000). Azari Takami et al. (1997b) outline the historical development of the caviar trade and Hosseini Seyed et al. (2008) ranks export goal markets for this commercially important product.

Sturgeon roe or eggs are known as caviar and form an expensive delicacy (Bolourchi, 1997). The word caviar may come from Farsi "kaya-dar", "khay-dar" or "khay-var" meaning "having eggs", from خاگ‌آور or khāgāvar for roe-generator, or from "chav-jar" meaning "a cake of strength or power" or "bread of lovers" in allusion to its reputed aphrodisiac qualities; havyar in Turkish means "fish eggs" but the origin of this word seems in some dispute among etymologists and it may be Greek (Georgacas, 1978; Sternin and Doré, 1993; Bolourchi, 1997).

In addition to sturgeon roe, eggs of other species are eaten in Gilan and Mazandaran, where the meal is known as ashpal. The species include Rutilus frisii, Abramis brama, Salmo caspius and less commonly Cyprinus carpio and Barbus sensu lato spp. The roe can be eaten cured as a condiment or when fresh, grilled, steamed or mixed with eggs and fried to form ashpal kuku, a custard-like dish.

The history of the caviar industry in Iran is a complex subject, variously reported in the popular media and in legend. The Russians are said to have obtained writs from Moslem leaders in the Caucasus in the early nineteenth century to the effect that Moslems could not eat these fishes, leaving the valuable caviar fisheries for Russian fishermen to monopolise (Kayhan International, 1 December 1962). The caviar industry was first granted by the Iranian government to Stepan Martinovitch Lianozoff (or Lionosoff, Lianozov) an Armenian subject of Czarist Russia in 1873 (or 1876 or 1879, accounts vary), regularly renewed and later transferred to his only son George. In 1896 the lease was renewed at an annual cost of 450,000 gold francs (Fortescue, 1920). In one version of events, Martin (the grandson of Stepan) disappeared in 1923, kidnapped while meeting two ravishing Armenian sisters, leaving only a letter ceding his rights in the caviar fishery to the Soviets (Tehran Mossavar Magazine, 18 April 1952; Time, 9 February 1953; L'Illustré, Lausanne, 20 January 1955; Tehran Radio, 6 May 1959). Another version simply has Martin selling his rights to the Soviet Government (Mirfendereski, 2000) or refusing to pay during the vicissitudes of the Russian Revolution. In 1919 another Russian subject, Grigor Petrovic Vanitsof rented the southern Caspian fisheries for 20 years but could not fulfill his obligations. A joint Irano-Soviet company, "Mahi Iran", formed under Soviet pressure on the Iranian government, was given a monopoly of the foreign sale of caviar in 1927 (in 1923 the fisheries of Astara, Anzali and Hasan Kiadeh had been occupied by Soviet troops and declared part of the Soviet fisheries). The Irano-Soviet company was run almost entirely by Soviet technicians and the caviar was marketed as of Russian origin (Kayhan International, 27 June 1959; Saffron, 2002). One part of the Persia/U.S.S.R. agreement banned chemical and explosive uses for capturing fish (Mirfendereski, 2000). The fishery was nationalised in 1953 and administered by the Iranian Fishery Company (Sherkat Shilat). Most of the catch was sold to the former U.S.S.R. (Anonymous, 1961b) and Soviet scientists organised caviar production until the Iranian Revolution in 1979-1980 (Taylor, 1997). Greenspan (1989) details more recent skullduggery.

Keyvanfar (1988) described the preparation of Iranian caviar from the various species and the following is taken from that account. Emadi (1994) and De Meulenaer and Raymakers (1996) also give accounts of this process and the kinds of caviar obtained. Sturgeons are alive or very fresh when brought to the processing plant. They are usually killed by a blow to the head. Sex is determined with an awl-shaped instrument inserted into the cloaca, pulling out some ova. The female is split open along the belly and the eggs and the enveloping adipose and connective tissues removed. The eggs are generally about 10% of the body weight. However, an average beluga of 68 kg can yield 18 kg of caviar in Iran (ca. 26%) (V. D. Vladykov, in litt., 1966), and a 40 kg beluga from Iran yielded 8 kg of caviar (20%) (L'Illustré, Lausanne, 20 January 1955). The largest amount obtained from a beluga was 360 kg of caviar (V. D. Vladykov, in litt., 1966). The other species give an average of 6 kg of caviar in Iran. The eggs are separated from the tissues by breaking the ovaries into pieces by hand and delicately pressing the eggs through a 10 x 10 mm screen. This takes only a few minutes. The eggs are then washed in fresh, cold (8-12°C) water for 30-40 seconds to remove fragments of ovarian tissue. The eggs are separated from the washing water by collecting them on a very fine mesh screen, the process taking 3-4 minutes. This type of washing is not done with A. stellatus because of the fragility of the egg membrane in this species.

The type and quality of the caviar is determined next and they depend on the colour, diameter and membrane strength of the egg. Large eggs with a strong membrane and a clear, grey, dark brown or gold colour are the best and are packed in metal containers. Small eggs with fragile membranes and sombre colouring are second quality and used for pressed caviar or bulk caviar. Pasteurised caviar is made from eggs with weak membranes since the heat solidifies the membranes. Salt is added at a rate of 4-6% to the weight of the eggs, varying with the season and being higher in the warm summer months. The salt is 99.2% sodium chloride and only 1-10 kg of eggs can be salted at one time so that salting is uniform. Boric acid and borax are added in a ratio of 2:3, comprising 20% of the total salt added, to aid in conserving the caviar. Caviar for export to the U.S.A. is exempt from this addition of boric acid and borax and only salt is used, 100 g for each 1 kg of caviar. The salt is mixed delicately with the eggs by hand for 50-250 seconds. A good salting process is essential for the preparation of caviar and is evidenced by the eggs having small white lines on their surface, the membrane becomes stronger and more resistant, the egg proteins become denser and coagulate, the eggs lose their adhesiveness, liquid stops coming from the eggs, and the density of the brine coming from the process increases. When these factors are detected the salting process is stopped. A salting process which is too long removes too much protein from the eggs and causes the eggs to clump together. A process which is too short removes too little water from the eggs and these eggs lose water gradually over several days in their container and become soft and semi-liquid. The eggs are then separated from the brine on a very fine mesh screen.

The U.S. Customs Service produces a description of caviar for the trade community (www.customs.ustreas.gov/imp-exp1/comply/caviar.htm downloaded on 20 July 1999). Caviar is graded on grain size, colour, flavour and firmness. Gold coloured caviar is the rarest and most desirable followed by light grey. Large grains are preferred over smaller ones.

There is a demand for caviar without borax and boric acid and such chemicals as methyl parahydroxy benzoate and propyl parahydroxy benzoate have been examined in Iran as alternatives (Iranian Fisheries Research and Training Organization Newsletter, 7:3, 1995).

Fresh caviar is not salted and requires careful refrigeration; its shelf life is short, a maximum of six weeks. Lightly salted caviar is called "malossol" from the Russian for "little salt", usually a 2.4% content in Iranian caviar which is very good quality compared to some caviar which contains up to 11% salt. The higher the salt content the longer the shelf life. Chilled malossol kept at -2 to 4°C will be edible for up to three months. Pressed caviar is prepared in a similar fashion except the salt content is higher, at 7% in the finished product. It will keep for a long time at 4-8°C. Borax gives a longer shelf life too and is less dangerous to human health than the amount of salt needed to give the caviar an acceptable shelf life. Most caviar consumed world-wide is pasteurised as some countries do not accept caviar with borax and higher salt levels are not acceptable to consumers. Pasteurised caviar has a shelf life of 12-15 months (De Meulenaer and Raymakers, 1996). Caviar should not be frozen or pasteurised as this affects the taste. Good quality caviar must be refrigerated. U.S. packaged caviar also contained tragacanth gum according to labels on jars from the 1960s. Bankehsaz (2009) found that the quality and grade of exported caviar can be maintained if storage time is less than 6 months in a -3°C cold room. Razavilar et al. (2001) found Iranian caviar to have a good microbial condition during processing and storage in Mazandaran.

The caviar is placed in boxes of 0.5 to 2 kg, each box being filled to within 1-2 cm of the lid. Sternin and Doré (1993) give tin sizes of 0.6 and 1.8 kg with a limited amount of 100, 200 and 300 g tins - most caviar is repacked at its destination in 30 g, 50 g, 125 g, 250 g, 500 g and 1 kg tins and jars). The lid is pressed on centrally to exclude as much air as possible and the excess brine is allowed to drain away by stacking the boxes vertically for 1-15 minutes. One further press is carried out manually, the outside of the box is cleaned, and boxes are stacked in piles of five for 20-24 hours in the cold season (October-March) and 12 hours in the warm season. During this period, the pile of boxes is turned over several times to remove the last traces of excess brine. After one last press on the centre of each box to ensure the lid adheres to the eggs and no air remains, the box is sealed hermetically with a ring of rubber. Well-prepared caviar has lost 4-6% of its initial weight, has a salt content of 3-5% and the eggs are separate and non-adhesive. Caviar in this form will keep for a long time at 0-2°C.  Caviar is re-packed in fully airtight tins, slowing down the maturation process for three months after which the caviar deteriorates.

A microbiological analysis of Iranian caviar imported to Turkey has been carried out by Altug and Bayrak (2003) who did not find any pathogenic and toxin producing Salmonella spp. and Clostridium perfringens. Coliforms, bacteria and yeasts showed some high counts, perhaps contamination during production stages.

First quality caviar consists of healthy, non-fragile eggs from one species with a large or medium size. The caviar is dry, of uniform colour - between clear-grey and dark-grey - without odour or abnormal taste. The box is filled within a centimetre of the edge. Second quality caviar has eggs which may be fragile, are of large, medium or small size, their colour varies from clear-grey to black, and they may be damp. Yellowish or brown caviar from A. gueldenstaedtii is acceptable for these two qualities of caviar. Egg size is determined by the cubic centimetres occupied by 100 eggs, e.g. for A. gueldenstaedtii large eggs occupy >1.9 cc, medium 1.4-1.9 cc and small <1.4 cc and for A. stellatus large eggs occupy >1.3 cc, medium 0.9-1.3 cc and small <0.9 cc. Egg sizes are not determined for Huso huso and A. nudiventris. Eggs of the former are much larger than A. gueldenstaedtii eggs while those of the latter are nearly the same size as A. stellatus eggs.

Russian and Iranian caviar tins, beluga, osetra and sevruga (Wikimedia Commons).

Russian and Iranian caviar tins, beluga, osetra and sevruga (Wikimedia Commons).

 

Sturgeon species cannot be readily identified from the size or colour of the eggs making up caviar. Diet, pigmentation of the adult, and age of the fish all appear to influence egg colour. Huso huso eggs are often light to dark grey, Acipenser gueldenstaedtii eggs are blackish to brown or almost golden and A. stellatus eggs are black according to traders. White caviar where the egg has a red spot on it is from albino fish. Light grey beluga and light yellow oscietra caviar are now very rare, in the past being found in only a small proportion of the species population which itself is now in decline (De Meulenaer and Raymakers, 1996). Le Comptoir du Caviar, which markets caviar (www.gourmet-tradition.com/en/comptoir_du_caviar.html, downloaded on 19 March 1999), describes Iranian sevruga as grey to black with fine grains and an iodised taste, Iranian oscietre as grey-black with bronze shades, middle-sized grains, very iodised with a little taste of walnut, and Iranian beluga as very rare, dark or light grey, large grains and a fine and gusty savour.

The single biggest market for caviar is first-class airline passengers. Supermarkets, hotels, restaurants and specialised retailers also market caviar. France consumed the largest amount in the 1990s, about 60-80 t, while Germany consumed 40-50 t. The Shilat packages its product carefully to ensure consumers know the caviar is genuinely Iranian. The large tins in which the caviar is packed keep their contents edible for 12-18 months at -2 to -3°C (the oil content and added salt prevent freezing). These tins are sealed in a piece of net which in turn is sealed on both sides with consecutive numbers, placed in a sealed linen bag and then in a wooden box. Each tin is also marked with the loading station number (where the fish are brought after capture to have their caviar removed) and also the number of the individual fish scratched on the side. Tins are shipped by air in "cooltainers" which have their own refrigeration unit. These large tins are vacuum-packed into smaller ones in packing centres in Europe (Christie, 1995). The main market in 2000 was Japan to which 30% of Iranian production was exported. Permanent markets in Europe are Switzerland, Germany, France, Luxembourg and Spain (I.F.R.O. Newsletter, 26:3, 2001) and the European Union is often the biggest importer of Iranian caviar. Iran was the top exporter of caviar in the year 2000 at 71.5 t valued at $34.4 million (IFRO Newsletter, 28:2, 2001). This is a value increase of 17% although the amount was less than in 1999 at 84.9 t. In 2002, 87% of caviar came from Iran (www.caviar.ru/english/digest.htm, downloaded 12 December 2002) although IRNA (8 December 2002) gives a figure of of almost 50%. The caviar export quota was 50,505 kg for Iran in 2006 (iran-daily.com, downloaded 28 July 2006) or 44.3 t (Iran Daily, 11 September 2006).

Iranian caviar sold in major airports like Heathrow in London comes in several kinds. Caviar House markets imperial, which has large gold grains and was previously reserved for the Shah's family (from Acipenser persicus); beluga, light to dark grey and large grained; royal black consisting of large deep-black grains from a 20-40 year old osetr; "oscietre", which is dark grey-brown to a golden yellowish; classic grey, a pale grey with large grains; and sevryuga, which is dark grey and fine grained. Prices vary with quality and time as shown below (personal observations):-
 
Imperial Beluga Royal Black Oscietre Classic Grey Sevryuga
December 1993
50 g £82 £76 £48 £36 £38 £23
1000 g £1411 £1318 £819 £621 £656 £399
September 1995
50 g £94 £101 £54 £48 £40 £36
1000 g £1640 £1759 £939 £836 £697 £630
September 1997
50 g £89 £88 £53 £47 £39 £34
1000 g £1536 £1540 £912 £812 £680 £590
November 1999
50 g £140 £160 £75 £60 £65 £53
1000 g £2420 £2770 £1312 £1060 £1138 £920
November 2000
50 g £184 £208 £114 £95 £99 £79
1000 g £3541 £3987 £2177 £1818 £1894 £1527
April 2002
50 g £184 £208 £114 £95 £99 £79
1000 g not given not given not given not given not given not given 

The types of caviar listed changed in 2003 as follows:-

 
Imperial XO Beluga Beluga XXL Royal Black Royal Black XL Oscietre Gold Classic Grey
March 2003
50 g £289 £160 £309 £11 6 £197 £119 £125

The types of caviar listed changed in 2004 as follows:-

 
Imperial XO Beluga Beluga XXL Royal Black Royal Black XL Classic Grey Sevruga Oscietre Imperial
September 2004      
50 g £145 £160 £195 £88 £114 £66 £56 £75 £98
100 g £275 £318 £385 £175 £225 £129 £110 £149 £195
200 g £540 £619 £750 £340 £435 £247 £221 £297 £385

The types and weights of caviar listed changed in 2006 as follows for "Prestige Selection":-

 
Beluga Royal Black Classic Grey Sevruga Oscietre Oscietre Gold*
March 2006      
50 g £155 £125 £105 £95 £120 £130
125 g £380 £305 £255 £230 £295 £320
250 g £750 £595 £495 £450 £580 £630

* = golden-coloured eggs from a mature oscietre.

The types and weights of caviar listed changed in 2006 as follows for "Prunier":-

 
Traditional Saint James Great American Paris Heritage
March 2006    
50 g £65 £85 £95 £120 £155
125 g £155 £205 £230 £295 £380
250 g £305 £405 £455 £585 £755
500 g £605 £805 £905 £1165 £1505
March 2007 (500 g not listed)          
50 g £75 £110 (not listed) £120 £170
125 g £185 £270 (not listed) £295 £415
250 g £370 £535 (not listed) £585 £830

The types and weights of caviar listed changed in 2006 as follows for "Private Reserve":-

 
Royal Black XL Imperial XO Beluga XXL
March 2006
50 g £135 £145 £195
125 g £330 £335 £480
250 g £650 £695 £950

Prices for these brands were not listed in March 2007, and were only available on request, indicative of both scarcity and constantly changing prices. Names of the different types of caviar keep changing so it is difficult to track price increases. In November 2008, beluga caviar from Caviar House was selling for £2170 for 250 g and other types had also continued to increase in price, often dramatically. Curiously, in July 2010 under "Caspian Sea caviar" beluga was selling at £1340 per 250 g. In 2011, all available caviar was Russian or farmed.

Taylor (1997) gives prices in Deutschmarks (DM) per kilogramme net weight (no duty paid) for Iranian caviar over 12 years including the approximate "bazaar" or illegal price for smuggled caviar (note also that A. gueldenstaedti probably includes A. persicus):-

 
Year Huso huso A. gueldenstaedti A. stellatus "Bazaar"
1983 540 408 341 200
1984 600 424 400 180
1985 675 465 404 180
1986 650 460 345 200
1987 650 414 325 180
1988 1630 445 310 180
1989 2600 510 345 220
1990 1596 432 304 220
1991 1600 450 337 180
1992 1600 470 345 160
1993 950 435 345 160
1994 950 500 355 80

Taylor (1997) also compares demand from western markets with supply from Iran; for Huso huso demand is 0.2 tonnes while supply is 2.0 t, for A. gueldenstaedti type I.A 2.0 t and 0.5 t, for A. gueldenstaedti type I.B 60.0 t and 40.0 t, for A. gueldenstaedti type II 15.0 t and 10.0 t, for A. stellatus I 100.0 t and 30.0 t, and for A. stellatus type II 100.0 t and 25.0 t.

Friedland (1986) gives a variety of recipes for caviar dishes.

Rehbein (1985) and Keyvanfar et al. (1987) studied soluble caviar proteins of sturgeon species including A. gueldenstaedtii, A. stellatus, A. nudiventris and Huso huso. They were able to distinguish the species on this basis and thus provide a means of detecting fraudulent caviar. Rezvani Gilkolaei (2002) used DNA PCR amplification and RAPD markers to identify caviar, in particular that of the endangered Acipenser nudiventris whose caviar has been substituted for more expensive caviar of A. gueldenstaedtii and A. persicus. Rehbein et al. (2008) tested and reviewed different methods for identifying caviar by species, including DNA, differential scanning colorimetry and determination of stable isotopes. Gessner et al. (2008) were able to distinguish between farmed and wild sturgeons based on fatty acid composition and recommend use of specific fatty acids as additives in the formulated diets of farmed fish. However, Ludwig (2008) reviewed methods of identifying caviar and other sturgeon products and detailed difficulties. No single method met the criteria he established (species-level identification, population identification, wild versus aquaculture, age of caviar). Cost was also a factor. Keyvanfar (1984) was unable to find genetic polymorphism in erythrocytes of the four species listed above using serological techniques.

Genus Acipenser
Linnaeus, 1758

This genus is characterised by a small, transverse mouth (large and crescentic in Huso), by the gill membranes being joined to the isthmus and not to each other (joined to each other and free of the isthmus in Huso), by a rounded or elongate snout, and cylindrical barbels. Bani et al. (2008) give details of brain morphology in Acipenser stellatus and A. persicus that suggest sturgeons have evolved different sensory strategies to cope with life in the deep sea.

There are 16 species in the genus and 4 are reported from Iran.

Ventral view of heads of Huso huso, Acipenser nudiventris, A. gueldenstaedtii and A. stellatus
(A. persicus is similar to A. gueldenstaedtii)

Acipenser baerii
Brandt, 1869

Introduced to the Caspian Sea basin by Soviet authorities (Karpevich and Lukonina, 1971; 1972; McNeil, 1979) but no records from Iran.

Acipenser gueldenstaedtii
Brandt and Ratzeburg, 1833

Common names

چالباش (= chalbash or short head), تاس ماهي (= tas mahi or bald fish; this term includes A. gueldenstaedtii, A. persicus and A. nudiventris for large eggs, in fisheries statistics), تاس ماهي روس (= tasmahi-ye Rus or tasmahi-e-russ), تاس ماهي ايراني (= tas mahi Irani), osiotra, osyetra, سگ ماهي (sag mahi), ماهي خاويار (= mahi-ye kaviar, meaning caviar fish), kaviari rusi.

[russkii osetr or Russian sturgeon in Russian; nere or rus neresi in Azerbaijanian; bekra or bekre balyk in Turkmenian].

Systematics

This species was originally described in part from the Volga, Ural and Terek rivers of the Caspian Sea. Sometimes spelt güldenstädti, but accents on letters are not used in Latin scientific names. Birstein et al. (1997) and Reshetnikov et al. (1997) spell the name gueldenstaedtii, regarding the double "i" ending correct as opposed to the emended single "i" which appears in much recent literature.

Acipenser gueldenstaedti persicus natio kurensis Belyaeff, 1932 was described as the Kura River subspecies but see below under Acipenser persicus. Comparison of serum proteins have shown antigenic characteristics distinguishing Volga and Kura River stocks in the Caspian Sea, matched by morphometric characters.

The fishes identified as A. gueldenstaedtii in Iran may well be almost entirely A. persicus, although this remains to be determined. Consequently data on morphology and biology are confused in many accounts. The distinction of A. persicus is questioned by authors (see below).

Some specimens have strong spines on the scutes and have been described as morpha aculeatus Lovetzky, 1834 although this has no nomenclatural status.

Birstein and Ruban (2004) and Birstein et al. (2005) state that this species has at least three, morphologically indistinguishable, genetic forms in the Caspian Sea. These are the pure form, one similar to A. baerii of Siberia, and one to A. naccarii of the Adriatic, with competing hypotheses to explain this. The most likely hypothesis is that the Caspian forms are closely related to the ancestral forms of the three species, evolving first as subdivisions of the original Caspian Sea population and then moving to different geographical areas when the Caspian was connected to them.

Pourkazemi et al. (1999; Rezvani Gilkolaei, 2000) found two distinctive genotypes and therefore populations of A. gueldenstaedtii in Iranian waters using molecular techniques. This species and A. persicus showed great degrees of similarity in a phylogenetic analysis (Iranian Fisheries Research and Training Organization Newsletter, 14:4-5, 1996; Pourkazemi et al., 2000). The common origin of the two species was about 1 million years ago (Annual Report, 1995-1996, Iranian Fisheries Research and Training Organization, Tehran, p. 61-62, 1997; Pourkazemi et al., 2000).

Key characters

This sturgeon has a short snout (less than 60% of head length) with a rounded tip in contrast to the long snout (>60%) and pointed tip in A. stellatus. Huso huso has an unusual, crescent-shaped mouth and continuous gill membranes forming a fold on the isthmus and A. nudiventris has a continuous lower lip and usually more than 50 lateral scutes. Closely resembling A. persicus, it is distinguished from that species by the short and blunt snout, yellowish-white belly and golden-brown back. A drawing in Vlasenko, Pavlov and Vasil'ev in Holčík (1989) of the two species has a snout length in head length of 4.3 as opposed to 3.2 for Acipenser persicus but figures of snout length in total length overlap for the two species. The interorbital distance is much less in the Persian sturgeon (29.2-30.5% of head length) than in A. gueldenstaedtii (Artyukhin and Zarkua, 1986) but in small specimens examined by me from Iran, some had gueldenstaedtii interorbital distance and persicus snout length. There is a colour plate and line drawings of the heads of the two species from the Black Sea in Birstein et al. (1997:8, 220).

Morphology

The lower lip is interrupted at its centre. Barbels are not fringed, lie nearer the snout tip than the mouth, and do not extend back to the mouth. Sheibani (2003b) described the posterior alimentary canal in this species.

Dorsal fin rays 26-51, anal fin rays 18-35. Gill rakers 15-36. Dorsal scutes 5-19, lateral scutes 21-50 and ventral scutes 6-14. There are rows of smaller star-shaped scutes between the dorsal and ventral rows in some fish, rounded in this species and more triangular in A. persicus. The chromosome number is 2n=250 ± 8 or 2n=247 ± 7 (Klinkhardt et al., 1995).

Sexual dimorphism

Females are larger than males of the same age.

Colour

The back is usually golden-brown but may be olive-grey to dark green, the flanks grey-brown, and the belly yellowish-white or rarely a lemon yellow. Young are blue dorsally and white ventrally.

Size

Attains 160 kg and 2.36 m, perhaps as much as 4 m although not confirmed. In Iran, fish identified as this species (see Systematics) averaged 16-20 kg and 1.4-1.6 m in the 1950s (Farid-Pak, no date). Tsepkin and Sokolov (1971) state that Safid River fish reach 2.42 m. De Meulenaer and Raymakers (1996) give 200 kg and an average length of 2 m.

Distribution

This species is found in the Caspian Sea, particularly in the Volga River basin, as far as Moscow in the past. Very small numbers are caught in the Kura and Astara rivers. Also found in the Black Sea basin. Khodorevskaya et al. (2001) review abundance and distribution in former Soviet waters of the Caspian Sea. It is less common than Acipenser persicus in Iranian waters.

In Iran, it is recorded from the Astara River in the west to the Gorgan River in the east (but see Systematics). Reported recently from such rivers as the Atrak, Gorgan, Gharasu, Tajan, Babol, Haraz, and Safid, the southeast Caspian Sea, southwest Caspian Sea and south-central Caspian Sea (Abbasi et al., 1999; Kiabi et al., 1999; Abdoli and Naderi, 2009). V. D. Vladykov's field notes in the early 1960s reported it from Kopurchal, Khadjenafas, Tazeabad, 12 Bahman, Nevissi, Izadeh and Hasan Kiadeh. Access to many rivers must now be restricted by reduced water flow, construction, weirs, dams, irrigation canals and pollution.

Zoogeography

Presumably a relict of the isolation of the Caspian and Black seas from the Mediterranean-Atlantic.

Habitat

There is no marked seasonal variation in depth distribution in the south Caspian Sea in contrast to the middle Caspian. This species is found over sand or sandy-silt bottoms in a temperature range of 2.3-24.8°C and a salinity range of 6.28-14.34‰ in the sea. It approaches closer to the coast in winter (February) than other sturgeons because it favours colder temperatures. It is found in numbers down to 50 m with only the occasional specimen being caught below this depth (Legeza, 1972; 1973). Brackish water is favoured because of food concentrations. High oxygen concentrations are needed, 6-7 mg/l for adults, although larvae only require a minimum of 1.56 mg/l at 20°C. Reproduction in the Kura ceases when temperatures reach 26°C. Eggs are sensitive to oil concentrations of 0.5-1.0 mg/l. Levin (1997) reports concentrations on the western shelf of the Caspian Sea during winter, as far south as Azerbaijan, at depths of 5-24 m.

Age and growth

Veshchev and Novikova (1986) and others have recently studied the spawning run of this species in the Volga River and found fish from 7 to 39 years old with 87.9% 13 to 27 years old. The spawning population comprises 32 age groups therefore. Males dominate at 63.6%. Males vary in length from 101 to 185 cm and weigh 3 to 38 kg and females from 116 to 200 cm and 9 to 46 kg. Most males begin to reproduce at 11-13 years while females begin at 12-16 years. Growth can be rapid, young-of-the-year reaching as much as 35 cm by autumn. Life span exceeded 48 years in the past. Khodorevskaya et al. (1993) cited in Levin (1997) gives Volga River spawning ages of 8 to 35 years with females 6-8 years older than males. Females have an average weight of 26-29 kg and a length of 136-163 cm; males are 12.0-14.5 kg and 130-134 cm. Females mature at 10 years, 2-3 years later than males. Minimum spawning intervals are 2-3 years for males and 3-4 years for females.

Von Bertalanffy growth parameters in Iranian females are L = 201 cm and K = 0.073 or 192 cm and 0.082 and for males 189 cm and 0.092 depending on the methodology used. Total mortality (Z) was 0.33-0.67 for females and 0.46-0.82 for males, natural mortality (M) was 0.05 for females and 0.06 for males, fishing mortality (F) was 0.62 for females and 0.39 for males, and optimum fishing mortality was (F) 0.21 for females and 0.37 for males (Iranian Fisheries Research and Training Organization Newsletter, 16:4-5, 1997).

Food

This species is primarily a mollusc eater (Polyaninova et al., 1999) but also takes crustaceans such as chironomids and gammarids and small fishes such as gobies (Gobiidae) and Clupeonella caspia. The introduced species of mollusc, Abra ovata, polychaete, Nereis diversicolor, and crab, Rhithropanopeus harrisii are now important diet items at 49.3%, 12.3% and 9.2% respectively in the Caspian Sea. The importance of oligochaetes like Nereis and Enchytraeus albidus in the diet of sturgeon species is recognised in Iran and studies on their ecology have been carried (IFRO Newsletter, 28:3, 2001). In rivers, fingerlings feed on various benthic organisms. Hajimoradloo et al. (2002) examined the diet of juvenile fish taken in beach seines from the Miankaleh peninsula in Golestan and compared it with the diet of A. persicus. The latter favoured cumaceans while A. gueldenstaedtii favoured gammarids. Both species had more empty stomachs in autumn and less in winter. A. gueldenstaedtii had more empty stomachs in all seasons than A. persicus. The food niche width was less in A. gueldenstaedtii and food overlap was highest in winter and lowest in spring.

Reproduction

Spawning migrations in sturgeons are triggered by temperature, daylength and flood discharge. This has been discussed more fully by Barannikova (1972) along with the effects of dams on this complexly timed, hormonal process. In northern rivers the water temperatures are 8-18°C (Artyukhin and Zarkua, 1986). The adult loses 25-30% of its weight after spawning and females are only ready to spawn again after 4-6 years and males after 2-4 years.

The spawning run in the Kura River is complex and four "races" have been recognised (Gerbilskii, 1955; Berg, 1959). These are early and late vernal, spawning in their year of entry, and summer-arriving and autumn-arriving hiemal which overwinter to spawn the following spring. The chief spawning period in the Kura River is from the end of May to the beginning of July (Zakharyan, 1972). (Note that this may in fact apply to A. persicus).

In the Volga River, A. gueldenstaedtii has a run beginning at the end of March or beginning of April at 1-4°C, peaking in July. Migration speed in the Volga is 18.1-22.6 km/day. Eggs are laid on gravel or stone beds at 4-25 m depths and a current velocity of 1-1.5 m/sec. in the Volga River. Some eggs are laid in shallower, flooded areas. Egg incubation is optimal at 9-15°C. The downstream migration of spawned out fish in the Volga begins in the second half of May and peaks in June and July. Levin (1997) summarises the migration of Volga River fish as follows. The small population of the early spring race enters the Volga delta in April-May and migrates upriver for 600-700 km before spawning in May-June at 12-15°C. The late spring race migrates to spawning sites in May-June, spawning in July-August at 19-22°C. In June-July the winter race enters the delta but only migrates upriver in the next summer. In August-October the late winter race enters the river. These winter races overwinter in deep parts of the river and spawn in April-May at 9-13°C.

Volga River sturgeons had a fecundity of 332,900 eggs in one study (Veshchev and Novikova, 1986), elsewhere reported up to 1,165,000 eggs for the Volga. The Safid River sturgeon fecundity is said to be less (this may be A. persicus). Eggs are brownish-grey and ovate, up to 3.3 x 3.8 mm in dimensions. A 150 kg fish yielded 5 kg of caviar (IFRO Newsletter, 29:4, 2001).

The sexual cycle lasts 2-3 years on the Iranian coast and is described by light microscopy in Hedayatifard et al. (2009).

The caviar of this species comprises 4-5 kg on average, making up 16% of the body weight in Iran. In Mazandaran fish enter rivers in autumn, overwinter and spawn in spring (Iranian Fisheries Research and Training Organization Newsletter, 9:6, 1995).

Parasites and predators

Niak et al. (1970) report infestations of the ciliate Trichodina sp. in sturgeons (species unspecified) in breeding ponds in Iran. Golvan and Mokhayer (1973) describe Corynosoma caspicum as a new species from this and other sturgeon species in Iran. Mokhayer and Anwar (1973) report the following parasites from Iranian sturgeons in general. These are the protozoan Trichodina reticulata, the coelenterate Polypodium hydriforme, the trematodes Skrjabinopsolus acipenserinus and S. skrjabini, the cestodes Amphilina foliacea, Bothrimonus fallax and Eubothrium acipenserinum, the adult nematodes Ascarophis ovotrichuria, Cyclozone acipenserina and Cucullanus sphaerocephala, the larval nematodes Contracaecum squalii, Anisakis schupakowi and Eustrongylides excisus, the acanthocephalans Leptorhynchoides plagicephalus, Pomphorhynchus laevis and Corynosoma caspicum, the annelid Piscicola geometra and the crustacean Pseudotracheliastes stellatus. Polypodium hydriforme destroys the eggs of sturgeons, up to 80% of the gonads, rendering reproduction insufficient to maintain the species. Amphilina foliacea causes parasitic castration in sturgeons. Many of the parasites provoke anaemia or block the intestine when numbers are high. Pomphorhynchus laevis is capable of piercing the intestine. Eustrongylides excisus produces stomach abscesses. Ectoparasites take blood but also facilitate attack by bacteria and fungi. On fish farms, Trichodina reticulata can cause high mortalities while having no apparent effect under natural conditions. Parasite numbers are controlled on fish farms by immersing the sturgeons in salty water to remove ectoparasites, by feeding food items known not to be carriers of parasites and avoiding such natural foods and intermediate parasite hosts as amphipods. Mokhayer (1976b) reports gas bubble disease in Iranian sturgeons without specifying the species of sturgeon as well as the monogenetic trematodes Diclobothrium armatum and Nitzschia sturionis, the digenetic trematodes Skrjabinopsolus acipenseris and S. skrjabini, the cestodarian Amphilina foliacea, the cestodes Bothrimonus fallax and Eubothrium acipenserinum, the nematode larvae Anisakis schupakowi, Contracaecum squalii and Eustrongylides excisus, and the nematode adults Ascarophis ovothricuria, Cucullanus sphaerocephala and Cyclozone acipenserina, the acanthocephalans Corynosoma caspicum, Leptorhynchoides plagicephalus and Pomphorhynchus laevis, and the crustacean Pseudotracheliastes stellatus. Hajimoradloo (2002) records the nematode Cystoopsis acipenseris in juveniles at a frequency of 6.42%. Sattari et al. (2002) record Cucullanus sphaerocephalus, Eustrongylides excisus, Skrjabinopsolus semiarmatus, Leptorhynchoides plagicephalus, Anisakis sp. and Corynosoma strumosum, the fauna being similar to other sturgeons because of their piscivorous feeding. Hajimoradloo and Ghorbani Nasrabadi (2003) found the prevalence of metazoan parasites in juveniles of this fish in the southeast Caspian Sea to be 8 species with Anisakis larvae the highest at 13.3%. Pazooki and Masoumian (2004) report on blood parasites form fish caught at Anzali, recording Cryptobia acipenseris and Haemogregarina acipenseris. These parasites caused no pathological effects in the wild fish but can lead to severe infections and cause anaemia on fish farms. Sattari and Mokhayer (2005a; 2005b) recorded the occurrence of parasites in this species from the Iranian southwestern and central coast of the Caspian Sea. The species found were the nematodes Cucullanus sphaerocephalus, Eustrongyloides excisus and Anisakis sp., the acanthocephalans Leptorhynchoides plagicephalus and Corynosoma strumosum, and the digenean trematode Skrjabinopsolus semiarmatus. General conclusions were that the diversity of parasites was less in Iranian waters than in the northern Caspian Sea, perhaps a reflection of the more varied habitat, its productivity and the carbonate ions differing between the two regions. The diversity of parasite seems to have declined over time also, perhaps as a result of unfavourable environmental conditions, particularly in the freshwater ecosystem which limits the waters available for spawning and parasite acquisition. Barzegar and Jalali (2009), in their summary of crustacean parasites of Iranian fishes, recorded Pseudotracheliastes stellatus from this sturgeon.

A wide range of fish species are predators on the eggs of this sturgeon and the young are taken by Silurus glanis, Alosa spp., Huso huso, and gobiids.

Economic importance

Chalbash have been fished in the Caspian Sea for at least 6000 years based on excavations at a Neolithic site on the eastern Caspian coast (Tsepkin, 1986).

This particular species is fished primarily in the months of September and in April-May in Iran. Caviar yield was 4-7 kg per female in the 1950s (Farid-Pak, no date). Yields from 1963 to 1967 of meat (and caviar) were 794.2 tonnes (69.3 tonnes), 918.2 (66.7), 849.0 (71.8), 974.6 (72.8), and 977.1 (75.9) respectively (RaLonde and Walczak, 1970b). A commercial house maintains (1995) that this species comprises 27% of the total catch. These data presumably include or are almost entirely A. persicus in Iran. Spring-caught chalbash produce 2-3 kg of eggs per fish while those caught in the fall have egg weights of 3-4 kg. The former are more suitable for pressed caviar than the higher priced grain caviar made from the larger eggs of fish caught in fall (Vladykov, 1964). Figures for tas mahi (this species plus A. persicus, and also A. nudiventris when eggs are large) average yearly catches in Iran were given by Vladykov (1964) for the period 1927/28-1931/31 to 1957/58-1961/62. Body weight varied from 264,105 kg (36.9% of total sturgeon catch) to 842,050 kg (78.9%) while caviar weight varied from 33,098 kg (69.3%) to 159,931 kg (85.1%) although the lowest percentage share of caviar for any of the five-year periods in tas mahi was 28.6%. The category of tas mahi provided the majority of eggs for caviar up to 1946/1947 (50-89%) but this fell to 29-31% for the period after 1949/1950 in Vladykov's data. Earlier data from Nevraev (1929) listed as A. gueldenstaedtii and A. nudiventris combined for the Astara region of Iran gives catches of 2002 to 9176 individuals for the period 1901-1902 to 1913-1914, for the Safid Rud region 26,721 to 54,257 individuals for the period 1899-1900 to 1913-1914, for the Mazandaran region 4065 to 8818 individuals for 1906-1907 to 1913-1914, and for the Astrabad (= Gorgan) region 2988 to 6044 individuals for 1902-1903 to 1913-1914. The capture fishery for tas mahi (A. gueldenstaedtii, A. persicus and A. nudiventris) was 89%, 4.2% and 6.2% respectively in 1973 but by 1993 had changed to 27%, 69% and 4% due to fingerling production of A. persicus (Abdolhay and Tahori, 1999). The stock of this species in Iranian waters in 2001 was 9.4 million (0.64 million) specimens comprising 12,900 tonnes (2074 t) with a commercial stock of 220 t (223 t) (Ivanov and Kanunin, 2001; figures in parentheses from text which does not agree with table).

Catches of this species in the southern Caspian Sea have declined from 837 t and 602 kg/boat/day in 1971-1972 to 57 t and 0.34 kg/boat/day in 1999. Young fish decreased in the decade prior to this study while older fish dominate at present (Moghim, 2004a). A sharp decrease in sea ranching of fingerlings and a consequent decrease in young fish abundance, will cause a a considerable decline in future catches.

Dry-smoked flesh (balyk) is especially favoured in Russia where this species occupies the first place in catches. Catches in the period 1898-1913 in the northern Caspian reached 10,000 tonnes a year only to decline through overfishing. The ban on sea fishing in 1941, restricting catches to rivers where they could be more closely controlled, led to a rebound of stocks and by 1977 a record catch of 11,980 tonnes was made.

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in aquaculture and as food.

Conservation

Sturgeons generally are conserved by fish farming and release of young and fry, attempting to augment natural populations. Stroganov (1968) reviews Russian fish farming methods. Derzhavin (1923) reported release of 7,620,000 fingerlings in the Safid River of Iran in 1923. Release of unfed sturgeon fry was discontinued in Iran in 1965 as unproductive. A hatchery produced annually 5.5 million sturgeon juveniles at 3-5 g each (McNeil, 1979), comprising the species A. stellatus and A. gueldenstaedtii (the latter presumably includes A. persicus). The Sad-e-Sangar (Dr. Beheshti or Martyr Beheshti) Fish Farm or Hatchery 27 km from Rasht in Gilan produced 14-15 million sturgeon larvae in 1987 and up to 3 million 2-3 g sturgeon are produced annually (Petr, 1987). Fingerling production from four hatcheries in Iran reached a record high of 12 million fish in 1995-1996 and with a new hatchery in the Gorgan region is expected to reach 20 million fingerlings (Abzeeyan, Tehran, 7(6):V, 1996). IRNA reported on 31 August 1998 that 24 million fry had been raised since March of that year, a 15.3% increase over the previous year and 20 million fry are now released each year. The Shahid Rajaee Fish Aquaculture Center at Sari, Mazandaran produces 5.5 million sturgeon fingerlings annually, released in 13 Caspian Sea rivers (IFRO Newsletter, 28:3, 2001). The only species not produced is Huso huso and the most popular is Acipenser persicus for its better quality caviar. The young are fed on daphnia and later oligochaetes (white worms). Fingerlings may be grown to 10-15 cm length before being released in the Safid River about 20 km from the sea to imprint on the river. In 1987 2.28 million fingerlings were released and in 1993 6.5 million from the Beheshti Hatchery. In 1993 a closed system fish culture plant was opened at this hatchery to produce at least 5 million sturgeon fingerlings annually (Abzeeyan, Tehran, 4(9):IV, 1993; see also Anonymous (1993c)). A later report mentions culture of Huso huso in addition to the sturgeon species mentioned above for the Dr. Beheshti Sturgeon Hatchery, and production of fingerlings exceeded 60 million in 1991, the best year from 1973 to 1993 (Abzeeyan, Tehran, 5(3 & 4):IX-X, 1994).

About half a million fingerlings were produced in autumn 1995 in Mazandaran province (Iranian Fisheries Research and Training Organization Newsletter, 9:6, 1995). In 1996, it was expected that 15 million sturgeon fingerlings would be produced from hatcheries, the main species being Acipenser persicus, Acipenser stellatus and Huso huso. Fingerlings would be 3-5 g in weight when released in the Safid River (Abzeeyan, Tehran, 7(2):IV, 1996). The Shaheed Beheshti Fish Propagation and Rearing Complex of Shilat (Iranian Fisheries Company) produced 9 million sturgeon fingerlings in 1997, each 3-5 g, for restocking (Bartley and Rana, 1998b). Eggs are incubated for 5-7 days. Fingerlings are fed on live Artemia, Daphnia and oligochaetes in 2.5 sq m circular tanks from day 15 (60-80 mg) and then in earthen ponds for 50 days to the 2-3 g size. The fingerlings remain in the release river for 10-15 days before entering the Caspian Sea.

The International Sturgeon Research Institute, which opened in 1994 near Rasht, released 22 million fry in 1996-1997 (Bartley and Rana, 1998b). The Institute carries out varies research programmes, e.g. on the histology of the gonads of reared sturgeons which have been found to be the same as sturgeon in nature (Bahmani and Kazemi, 1998).

Abdolhay and Tahori (2006) give fingerling production for this species as:-

Process/Year 2000 2001 2002 2003 2004
Female broodstock captured 81 74 65 0 31
Injected broodstock 29 24 19 0 10
Spawning rate * (%) 89.6 79 66 0 80
Fertilisation rate (%) 70 55.5 49 0 71
Survival rate in incubators (%) 53 53.9 48 0 75.1
Survival rate in tanks (%) 80 70.1 68 0 79
Stocking density in ponds (fish/ha) 88,333 74,580 63,752 0 65,000
Survival rate in ponds (%) 65 79 71 0 65.1
Fingerling production (x 1000) 1327 447 1816 0 617

* Rate of response to hormone injection

An experimental approach to conservation of stocks has been the raising of sturgeon artificially to a size where they produce caviar. The Shahid Beheshti sturgeon aquaculture centre raised a member of this species to 121 cm, 11.5 kg and 8 years of age when it yielded 1.4 kg of caviar (Iranian Fisheries Research Organization Newsletter, 37:2, 2003; Iranian Fisheries Research Organization Newsletter, 40 & 41:4, 2004).

Shevchenko et al. (1999) summarise rearing technology for A. gueldenstaedtii in Iran. Fingerlings are raised on artificial feeds in 1-4 cu m plastic tanks for up to 180 days. A mean mass of 120 g is attained, with a maximum of 300 g. Growth rate of different age groups varied from 1.59 to 0.56% and daily weight gain was from 4.23 to 1.42%. The mean daily increment was affected by stocking density, daily rations, oxygen content, feed quality and maintenance of feeding routine. Falahatkar and Amini (2003) give further details on propagation from broodstocks including maturity duration, oocyte diameter and weight, motility and density of spermatozoids, time taken to reach 4 and 16 cell divisions, incubation duration, fertilisation percentage achieved at each stage, mortality rate during incubation, number of larvae obtained from each broodstock, number of larvae per gramme, weight of each larva, and morphometric parameters and age for each broodstock. Akrami et al. (2005) found cladocerans, copepods and chironomid larvae were secondary prey items of fingerlings in one earthen pond with ostracods occasional prey, while in another pond all these were secondary prey. Condition factor and growth decreased as weight and length of fingerlings increased. Growth was was negatively allometric (b<3).

De Meulenaer and Raymakers (1996) give figures for Iranian hatchery production from 1983 to 1992 as 1.03 to 6.61 million fingerlings (mean 2.9 million) although mature adults are becoming increasingly difficult to catch for stripping of eggs and sperm. These Iranian hatcheries are much smaller than Russian ones which produced about 25 times this number on average annually from the Volga River hatcheries alone.

There is an extensive Russian literature on how to raise sturgeons, e.g. Mil'shtein (1957; 1972), Marti (1972), Barannikova (1987) and Dettlaff et al. (1993). A recent (1984-1986) estimate of this species in the Caspian Sea is 47.7 million fish with 24-28% produced by artificial means.

All sturgeons are particularly threatened on the spawning migration when they concentrate in rivers (Rochard et al., 1990). Sturgeons in the Aras River on the former Soviet-Iranian border, for example, are threatened by dams and water diversion schemes (Zakharyan, 1972). However this is not an annual migration so the populations are not subject to loss every year. The common problems encountered by all Caspian sturgeons are dams and weirs which block reproductive migrations of adults upriver and also of young and adults returning to the sea, water abstraction for irrigation which reduces flow or even dries up a river, degradation of the river bed by extraction of gravel for construction or the change in silt deposits by the filtering effects of dams, increased water clarity enables predators to be more effective changes in the oxygen and temperature regimes caused by water abstraction, retention of water behind dams or untimely release from dams, pollution, attraction of adults into irrigation channels by their strong water flow and changes in the invertebrate fauna on which the young feed in rivers (Vladykov, 1964; Anonymous, 1970c; Whitney, 1979; Rochard et al., 1990). Variations in Caspian Sea levels also had effects (Khodorevskaya et al. 1997). For Huso huso these include lowered accessibility to feeding sites and variations in food abundance which lead to decreases in relative weight gain and to a halving in the number of females. The growth and survival of juvenile Acipenser gueldenstaedtii in the Volga River delta during their first winter is affected by lower water levels.

Stocks in their sea life were fairly safe until trawling was introduced. There are restrictions on trawling in the sea to reduce loss of young sturgeons (Ricker, 1970) and trawling is banned in the territorial waters of Azerbaijan (Markarova and Alekperov, 1989). It has been suggested that the Caspian Sea level should be maintained at -28.5 m or above to retain water productivity on which sturgeons ultimately rely. A 1 m decline in level can reduce fish food supply by 60% and hinders migration to feeding grounds, another 20% loss (Petr, 1987).

The institution of closed seasons for fishing and restrictions on techniques used to limit juvenile catches have been implemented in the former U.S.S.R. The fine for illegal possession of a Huso huso was about £280 in 1977. Fish lifts have been built on the Volga River about 5000 km upstream from the Caspian Sea to transport sturgeon around the Volgograd Dam. The system transports about 10-20% of migrating Huso huso, Acipenser gueldenstaedtii and A. stellatus but is relatively inefficient (Rochard et al., 1990). The poor situation is compounded by the lack of suitable spawning conditions above the dam and by adults having to migrate downstream through the dam's turbines. The turbines have wide blades and rotate slowly so most adults cannot make it through although the young are short enough to survive the transit. Khodorevskaya et al. (1997) summarise the decline in catches of this species after the regulation of the Volga River flow by the Volgograd Dam, built in 1958-1960, which cut off as much as 80% of the spawning grounds.

In Iran baiting hooks with oilcloth or fish was banned in 1952 as this method took large numbers of immature sturgeon (Vladykov, 1964). Some problems however may be intractable such as local consumption of immature fish rather than release or registration in catches. This lack of registration prevents adequate assessment of the catch and effective management suffers (Vladykov, 1964). Iran has recently taken a number of steps to protect the caviar resources including a reduction in the annual catch from 3000 tons (sic, probably tonnes) to 1500 tons, restricting export to the government rather than private companies, combating the illegal caviar trade, and the setting of export quotas and price controls for Caspian Sea countries (Abzeeyan, Tehran, 4(5):VI; 4(7):VI, 1993). Gill netting was prohibited in 1995 (Abzeeyan, Tehran, 6(5, 6):IV-V, 1995). The break-up of the Soviet Union led to smuggling and overfishing in the newly independent countries around the Caspian but Iran was able to stabilize world prices by reducing its caviar exports by 30%. Until 1992 Russian caviar dominated the world market but more recently Iran became the main supplier with income for 1989-1994 twice that of 1979 and 1989 (Abzeeyan, Tehran 5(1 & 2):VII, 1994; Ferguson, 1994). Nevertheless, some authorities believe overfishing by the five Caspian nations, particularly in the sea where immature fish are taken along with adults, will result in the extinction of the sturgeon species there (Los Angeles Times, Part A, page 1, 28 August 1993). An account of the caviar black market in Dagestan is given by Chenciner (1998).

Moghim et al. (no date) note that juveniles of this species are caught in the beach seine fishery for other species in Mazandaran. During 2001-2002, 23,760 seine hauls had a by-catch of 2% for this species among sturgeons captured.

Lelek (1987) and Maitland (1991) report this species as "vulnerable" in Europe because it grows and matures slowly, it is exploited, affected by pollution and killed by river engineering. Critically endangered in Turkey (Fricke et al., 2007). This species showed the greatest decline among Iranian sturgeon species through overfishing of younger age groups and habitat alterations (RaLonde and Walczak, 1970b). Kiabi et al. (1999) consider this species to be vulnerable in the south Caspian Sea basin according to IUCN criteria. Criteria include commercial fishing, medium numbers, habitat destruction, medium range (25-75% of water bodies), absent in other water bodies in Iran, and present outside the Caspian Sea basin. IUCN ranks all stocks as endangered (Vecsei, 2001).

Further work

The main concern with all sturgeon species is maintaining a viable commercial stock. Poaching has caused a decline in the available number of fish which can be used for breeding and moreover more than 30% of breeders do not respond to hormone stimulation (Kokoza et al., 1995). There were 6 times more nets in Azerbaijan waters and 4 times more in the Volga River delta in 1993 than in the 1980s. The legal catch will probably have to be completely prohibited (Ivanov et al., 1995). Efron (1993), for example, describes the "caviar crisis" in the Caspian Sea but problems have long been evident (Anonymous, 1961a). In 1996, 1 t of caviar was seized from smugglers in Gilan and one smuggler was fined 20 billion rials (IRNA, 28 July 1997, www.netiran.com). Maintenance of the stock may only be possible by hatchery production as river regeneration is no longer feasible because of dams. Mortality in Iran for hatchery reared eggs of 2 months age was 30-35%, for larvae 20-40%, and for fingerlings 30-40%, a satisfactory level but this could always be improved on (Petr, 1987). Yearly production of sturgeon fingerlings in government hatcheries in Iran was 1.03 millions in 1983, 1.11 in 1984, 1.13 in 1985, 2.28 in 1986, 3.10 in 1987, 3.16 in 1988, 3.15 in 1989, 4.34 in 1990, 6.60 in 1991, and 3.20 in 1992 (Emadi, 1993a). The 1996 hatchery production of sturgeon was 12.5 million in 1996 (Bartley and Rana, 1998a). A hatchery facility in Gilan covers 136 ha, produces up to 7 million sturgeon fingerlings a year with plans for up to 20 millions, and is said to be the largest and most modern sturgeon hatchery in the world.

The Israelis farm osetra and caviar from this species was on sale at Philadelphia airport at US$75/oz on 19 April 2006.

A detailed comparative study of the morphology of this species and Acipenser persicus in Iran would enable the young and adults to be clearly distinguished as well as stocks within each species as a management tool.

Sources

See under the family account.

Iranian material: Hatchery adults examined at Bandar-e Anzali.

Acipenser nudiventris
Lovetzky, 1828

Common names

شيپ (= ship, šep or sheap), تاس ماهي (= tas mahi, included under this name with A. gueldenstaedtii and A. persicus when eggs are large for fisheries statistics), tass mahi shekam brahne, سگ ماهي (sag mahi), ماهي خاويار (= mahi-ye kaviar, meaning caviar fish), keshdi, shenavar.

[kalamo, kelemo or kulamo, xazar kalamosu, gaya baligi, girt, ag-gyal or bich-nyarya in Azerbaijanian; sip or bekre balyk in Turkmenian; spiny sturgeon, thorn sturgeon, fringebarbel sturgeon, barbel sturgeon, bastard sturgeon].

Systematics

Acipenser nudiventris was originally described from the Aral Sea.

Acipenser schypa Eichwald, 1831 is a synonym. It is credited to Linnaeus by Eichwald but not described by Linnaeus; if this name is available then it is preoccupied by Acipenser schypa Gueldenstaedt, 1772 (Eschmeyer et al., 1996). Note that Holčík (1989) gives the spelling as shypa. Acipenser shipa Lovetsky, 1834 and Acipenser schypa Kessler, 1856 are synonyms. Acipenser schip Eichwald, 1841 is presumably a misspelling. Acipenser shyp Forster, 1767 may have priority but this has not been investigated.

Acipenser nudiventris derjavini Borzenko, 1950 was described as the Caspian Sea subspecies, as the type locality for the nominate subspecies is the Aral Sea, but derjavini is no longer recognised (Holčík, 1989).

A hybrid with Acipenser stellatus is reported from the Safid River (Nedoshivin and Iljin, 1927) and it also hybridises with Huso huso (Berg, 1948-1949).

V. D. Vladykov points out (in litt., 1973) that ship (in Russian) is probably a Turko-Tartar word referring to a hybrid since this species has a snout intermediate in length between that of Acipenser gueldenstaedtii, which is short, and that of Acipenser stellatus, which is long. The Russian word means prickle or thorn and has given rise to the common names for this fish in English of "spiny" or "thorn" sturgeon. Acipenser nudiventris, as its name indicates, has weakly developed or worn ventral scutes so the names spiny or thorn sturgeon are inappropriate. Vladykov recommends "sheap" as the common name to avoid confusion with the word "ship" in Russian (or for that matter in English).

Nucleotide diversity is much lower than other sturgeons in the Caspian Sea, possibly due to a smaller population size. Haplotypes of sturgeons from the Ural River in the north Caspian and Iranian waters were significantly different (Qasemi et al., 2006). Microsatellite studies indicate that there is more than one population in the south Caspian Sea and these populations are different from the Ural River one in the north Caspian Sea (Safari et al., 2007; 2008; 2008).

Key characters

This species has a continuous and thick lower lip, usually more than 50 scutes laterally, fimbriate barbels, and a transverse mouth.

Morphology

The body is deepest at the first dorsal scute. The rostrum is rounded and conical in shape in adults, more spatulate in young. Adults are covered with minute scutes giving a sandpaper texture although visually appearing smooth. Dorsal fin rays 39-57 and anal fin rays 23-37. Dorsal scutes 11-17, lateral scutes 49-74 and ventral scutes 10-17. There are no large plates on the body between the scutes. Scutes lack a hook and even juveniles have this usual feature barely developed. Ventral scutes are lost or absorbed in large adults (hence the scientific name). Gill rakers 24-45. Chromosome number is 2n=118 ± 2 (Klinkhardt et al., 1995) or 2n=118 ± 3 (Nowruzfashkhami et al., 2000). Nourouz Fashkhami et al. (2009) gives details of a method to produce the most metaphase plates.

Sexual dimorphism

Females are larger than males. Abdurakhmanov (1962) reports a greater average number of gill rakers in females, a longer postorbital distance in females, and longer caudal peduncle, pectoral fin, pelvic fin, snout, eye and snout tip to barbel distance in males.

Colour

The back is olive-green, grey-green or grey-blue, fading to a yellowish-white belly. Fins are grey.Juveniles mayhave the same colouration as adults or be almost black dorsally and laterally with a white belly.

Size

Attains 2.21 m and 127 kg. Safid River fish reached 43 kg, weighed when frozen, with the average being 20.1 kg, in 1914-1915 (Nedoshivin and Iljin, 1927).

Distribution

Found in the Black, Caspian and Aral seas and their drainages but extinct in the latter. In the Caspian Sea it is most common in the south, being rare in the Volga River for example. A long residency in fresh water probably accounts for their scarcity since mortality from winter and predators is high. Migrations in the Kura River extended 650 km and in the Aras River 300 km until the Mingechaur Dam was built. Enters the Aras, Astara, Safid, Tajen and Babol rivers in Iran (Derzhavin, 1934; Armantrout, 1980; CITES website, downloaded 5 April 2004). Also reported from Hasan Kiadeh by Derzhavin (1934) and by V. D. Vladykov based on field work notes made in 1962. Rostami (1961) also records this species from several localities on the Safid River. More recent works only report it from the Safid River, the southeast Caspian Sea, southwest Caspian Sea and south-central Caspian Sea (Kiabi et al., 1999; Abdoli and Naderi, 2009) and from the Safid River (Abbasi et al., 1999). Vecsei et al. (2002) consider it as rarely observed in Iran,

Zoogeography

Presumably a relict of the past isolation of the Aral-Caspian-Black seas from the Mediterranean-Atlantic. This species is reported from the Karakum Canal and Kopetdag Reservoir in Turkmenistan by Shakirova and Sukhanova (1994) and Sal'nikov (1995) and may eventually reach the Tedzhen (= Hari) River basin of Iran.

Habitat

A rare species in trawl catches but known from feeding grounds along the eastern coast of the south Caspian Sea (Legeza, 1973). Only 100 fish enter the Kura and the Ural stock, an undammed river, is in the low thousands (Vecsei et al., 2002). This species was never as abundant as other sturgeons because young spent 2-8 years in fresh water where predators abound and food is more limited (Vecsei et al., 2002) As an adult, it favours the areas near river mouths with muddy bottoms. Markarova et al. (1991) state that its main abundance is south of the mouth of the Kura River and that it ascends the Safid Rud to spawn, although in smaller numbers than the Kura River. This species is uncommon in Iranian waters, only 2.5% in numbers and 4% in weight of the Safid River catch in 1914-1915 (Nedoshivin and Iljin, 1927; RaLonde, 1970b), and catches in Azerbaijan are not more than 5% of all sturgeons (Markarova and Alekperov, 1989). It is usually found over mud near shore at 30-60 m.

Age and growth

Maturity is attained 6-13 years in males and begins at 12-22 years in females and most are mature at 14 years. Females grow faster than males. Caspian fish grow faster and larger than those in the Aral Sea. The oldest fish in the Kura River was 35 years (Markarova et al., 1991), and maximum age is 36 years. Growth is rapid with one-year-olds in the Caspian being 23-29 cm long and weighing 40-60 g.

Food

Markarova et al. (1991) found sheap in the south Caspian Sea to eat fishes such as Atherina, Neogobius (presumably including related genera), Benthophilus and Clupeonella, polychaete worms (Nereis), and various crustaceans. Molluscs play a small part in their diet but eggs of other sturgeons and the crab Rhithropanopeus harrisii are very important. The crab, an accidental introduction to the Caspian Sea at the end of the 1950s, comprises 70% by weight of the food taken. Young sheap in the Kura River feed on insect larvae such as caddisflies, dragonflies, mayflies and stoneflies. Hashemyan et al. (2005) found diet in A. persicus, A. stellatus and A. nudiventris in coastal waters of Mazandaran and Golestan at depths less than 20 m to consist of annelids (50.8%), amphipods (41.5%), small fish 4.8%), decapods (2%) and bivalves (0.9%). Fish shorter than 40 cm fed mostly on shrimps, polychaetes and gammarids, 41-80 cm fish fed on shrimps, gammarids, polychaetes, bivalves and smaller fish, while fish greater than 80 cm fed mostly on shrimps and smaller fish.

Reproduction

A spawning migration to rivers occurs year-round but peaks in March-April and in October-December in the Kura River of Azerbaijan (Markarova et al., 1991). The spring run begins at 6.2-13.0°C while the fall run is at 12.0-17.9°C. Males predominate over females by 3-6 times. Spawning occurs in April-May at water temperatures of 10-25°C and normal development occurs between 11.0 and 17.1°C. Eggs are laid on pebbly substrates at current speeds of 1-2 m/sec. Fecundity in sea-caught fish was up to 959,100 eggs (Markarova et al., 1991). Elsewhere egg numbers may reach 1,290,000 with diameters up to 3 mm. Fry soon migrate to the sea. Spawning by individuals is not an annual event but occurs at intervals of 2-3 years for females and 1-2 years for males, allowing for recovery and fattening. Some spent fish may remain in the Kura River for up to 8 years. Halajian et al. (2007) used biopsies to determine sex and sexual maturity stages in 5 and 6 year old fish. Males matured sooner than females. Shalouei and Imanpour (2009) found that spermatozoa were immotile in ovarian fluid because of the high concentration of potassium and osmotic pressure.

Parasites and predators

Niak et al. (1970) report infestations of the ciliate Trichodina sp. in sturgeons (species unspecified) in breeding ponds in Iran. Mokhayer and Anwar (1973) report on parasites of sturgeons including this species (see under Acipenser gueldenstaedtii). Mokhayer (1976b) reports gas bubble disease in Iranian sturgeons without specifying the species of sturgeon as well as the monogenetic trematodes Diclobothrium armatum and Nitzschia sturionis. Sattari et al. (2002) record Cucullanus sphaerocephalus, Eustrongylides excisus, Skrjabinopsolus semiarmatus, Leptorhynchoides plagicephalus and Eubothrium acipenserinum, the fauna being similar to other sturgeons because of their piscivorous feeding. Sattari and Mokhayer (2005a; 2005b) recorded the occurrence of parasites in this species from the Iranian southwestern and central coast of the Caspian Sea. The species found were the nematodes Cucullanus sphaerocephalus and Eustrongyloides excisus, the cestode Eubothrium acipenserinum, the acanthocephalan Leptorhynchoides plagicephalus, and the digenean trematode Skrjabinopsolus semiarmatus. General conclusions were that the diversity of parasites was less in Iranian waters than in the northern Caspian Sea, perhaps a reflection of the more varied habitat, its productivity and the carbonate ions differing between the two regions. The diversity of parasite seems to have declined over time also, perhaps as a result of unfavourable environmental conditions, particularly in the freshwater ecosystem which limits the waters available for spawning and parasite acquisition. Shenavar Masouleh et al. (2006) found hatchery fingerlings to harbour Trichodina sp.

Economic importance

The relative scarcity of this species accounts for it being not more than 1% of the Caspian Sea catch of sturgeons. The highest catch in the Kura River seems to have been 6000 fish in the 1930s. The Iranian catch after the CITES website (downloaded 5 April 2004) was:-

Year 1990 1991 1992 1993 1994 1995 1996 1997 1998 1999 2000
Tonnes 1.9 22.4 19.0 17.5 17.3 15.7 16.6 13.5 19.4 21.0 3.5 (spring only)

Moghim (2004b) records the total Iranian catch as 2% of the total sturgeon composition and it is declining. In 1972 the catch per unit effort was 67 tonnes and 0.5 kg/boat/day but by 2002 it was 15 t and0.09 kg/boat/day.

Conservation

Reduction in flow of the Kura River, the main spawning ground, is four times less than before regulation (5.5 km3/year compared to 20-24 km3/year). Sheap find it difficult to enter the river. Artificial propagation will be the only way to maintain the population. Between 2.9 and 6.2 million young sturgeon were released annually in the Caspian Sea from 1966 to 1971. This situation is mirrored in Iranian rivers such as the Safid Rud. This species is also particularly sensitive to oil pollution when young. There are reports that all but the Ural River population are on the verge of extinction (The Sturgeon Quarterly, 2(2):1, 1994; Vecsei, 2001). It is already extinct in the Aral Sea (DeSalle and Birstein, 1996). This species is protected in Iran since populations along the southern Caspian shore have been greatly reduced and there are not enough mature fish for fish farming (Bartley and Rana, 1998b; Vecsei et al., 2002). However the CITES website (downloaded 5 April 2004, but citing September 2000 data) reports that Iranian hatcheries still obtain some breeders from rivers. CITES also notes that the number of fishing stations for this species in Iran has been decreased by half, use of gillnets for Rutilus spp. prohibited as they take sturgeons too, egg removal by caesarian section instituted, release of fry from a breeding stock of 3000 fish, and lower export quotas instituted.

Abdolhay and Tahori (2006) give fingerling production as:-

Process/Year 2000 2001 2002 2003 2004
Female broodstock captured 15 38 16 50 25
Injected broodstock 14 21 29 32 19
Spawning rate * (%) 86 95.2 78 74 75
Fertilisation rate (%) 80 71.5 73 70 72
Survival rate in incubators (%) 54 61.5 51 49 74
Survival rate in tanks (%) 70 74.7 76 61 66
Stocking density in ponds (fish/ha) 92,100 77,005 56,194 87,986 61,667
Survival rate in ponds (%) 71 60 85 34 20
Fingerling production (x 1000) 1143 1782 1819 1414 1311

* Rate of response to hormone injection

Moghim et al. (no date) note that juveniles of this species are caught in the beach seine fishery for other species in Mazandaran. During 2001-2002, 23,760 seine hauls had a by-catch of 1% for this species among sturgeons captured.

This species is sensitive to pesticides such as diazinon. The LC50 (96 h) was 4.6 mg/l and lowered erythrocyte and lymphocyte counts were recorded with a significant increase in neutrophil counts (Khoshbavar Rostami and Soltani, 2005). Parand Avar et al. (2008) studied the effects of photoperiod during feeding by juveniles on Daphnia. Uptake was higher in dark conditions. Shalouei et al. (2009) studied extenders of spermatozoa motility and Shalouei et al. (2008) the correlation between seminal plasma indices and spermatozoa motility..

Maitland (1991) lists this species as "endangered" in Europe because of the declining population, slowness in growth and maturity, exploitation, and pollution and dams on the spawning migration. Birstein (1993) and the CITES website (downloaded 5 April 2004) also consider it be endangered. Critically endangered in Turkey (Fricke et al., 2007). Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in aquaculture and as food.

Kiabi et al. (1999) and Moghim (2004b) consider this species to be critically endangered in the south Caspian Sea basin according to IUCN criteria while the IUCN gives endangered (Vecsei et al. (2002). Criteria include commercial fishing, few in numbers, habitat destruction, limited range (less than 25% of water bodies), absent in other water bodies in Iran, and present outside the Caspian Sea basin.

See also under A. gueldenstaedtii.

Further work

See under A. gueldenstaedtii.

Sources

See under the family account.

Iranian material: None.

Comparative material: BM(NH) 1879.11.14:56, 1, 255.0 mm total length, U.S.S.R., Tschinas (no other locality data); BM(NH) 1879.11.14:57, 1, 217.0 mm total length, U.S.S.R., Tschinas (no other locality data); BM(NH) 1897.1.25:9, 1, 411.7 mm total length, Romania, Orsova, lower Danube (no other locality data).

Acipenser persicus
Borodin, 1897

Common name

قره برون (= qara burun, kara burun, kareh burun or ghareburun, meaning black nose), تاس ماهي (= tas mahi, this term includes A. gueldenstaedtii), دراكول (= darakul), تيريج (= tirij), تاس ماهي ايراني (= tasmahi-ye Iran), تاس ماهي ايراني (= tasmahi Irani or tasmahi-e-Iran), سگ ماهي (sag mahi), ماهي خاويار (= mahi-ye kaviar, meaning caviar fish), cetra.

[nara, nyarya or njara, neresi, Kur narasi for natio kurensis, or bekra in Azerbaijan; perseya, gunorta perseya, bekre balygy in Turkmenian; kurinskii or persidskii osetr, i.e. Kura or Persian sturgeon in Russian].

Systematics

The type locality of this species is the Ural and Kura rivers.

Regarded as a not distinct from or a subspecies of Acipenser gueldenstaedtii by some authors (see Borodin, 1926; Berg, 1948-1949; Whitehead et al., 1984-1986; Keyvanfar et al., 1987; Keyvanfar, 1988; Ruban et al., 2008) but Luk'yanenko et al. (1974), Artyukhin and Zarkua (1986), Vlasenko, Pavlov and Vasil'ev in Holčík (1989), Keyvanfar and Nasrichari (1999), Pourkazemi et al. (2000), Subbotkin and Subbotkina (2001), Ghorbani and Hajimoradloo (2002), and Gharei et al. (2005) restore it to a full species on meristic, morphological, ecological, caviar proteins, serum proteins, mtDNA, genomic DNA and immunological grounds. And again, Birstein et al. (2005) consider it not to be distinct from Acipenser gueldenstaedtii on the basis of molecular analyses. Ruban et al. (2008) used meristic, morphometric and molecular data to come to the same conclusion. Acipenser gueldenstaedtii has a complex intraspecies structure according to Birstein et al. (2005) and, depending on the rivers and populations sampled for any given studies, conflicting results can arise. For the moment, the taxon A. persicus is retained here as distinct until further resolution of the problem is attained, although given the decimation of populations this may not be possible.

Moghim et al. (2009) report at least 18 groups that segregate spatially and temporally for spawning in the Caspian Sea basin.

The type subspecies is found in the Caspian Sea and Acipenser persicus colchicus Marti, 1940 in the eastern Black Sea. A natio kurensis Belyaeff, 1932 is reported from the Kur River of Azerbaijan within Acipenser gueldenstaedtii persicus. Electrophoretic studies of blood proteins coupled with morphological data indicate that Gorgan and Safid River populations are two geographical races (Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 42, 1996).

Hybrids with Acipenser gueldenstaedtii are reported from the Volga and the Caspian Sea and have been produced artificially (Vasil'eva et al., 2001).

Two syntypes of Acipenser persicus are possibly in the Zoological Institute, St. Petersburg (ZISP, formerly ZIL) (Eschmeyer et al., 1996).

Key characters

This species has long been confused with A. gueldenstaedtii, but can be distinguished by a more elongate, massive and downward curved snout, a white belly and a grey-blue back. A drawing in Vlasenko, Pavlov and Vasil'ev in Holčík (1989) of the two species has a snout length in head length of 3.2 as opposed to 4.3 for Acipenser gueldenstaedtii but figures of snout length in total length overlap for the two species. The interorbital distance is much less in the Persian sturgeon (29.2-30.5% of head length) than in A. gueldenstaedtii (Artyukhin and Zarkua, 1986) but in small specimens examined by me from Iran, some had gueldenstaedtii interorbital distance and persicus snout length.

Morphology

The Persian sturgeon is slender with an elongate and cylindrical body, a long head, and a narrow, medium length (5.6% of total length), massive and usually depressed snout. The snout width near the mouth is 37% of head length. The pectoral fins are relatively small and have only a weak bony ray. There are usually 1-4 rows of smaller, longitudinally arranged, bony plaques between the scutes of the dorsal and lateral rows and sometimes between the lateral and abdominal rows. The barbels are located relatively closer to the snout tip than those of A. gueldenstaedtii. Sheibani and Adib Moradi (2000) described the histology of the pylorus and pyloric caecum in this species and Sheibani and Pahlavan (2003) the developmental histology of the liver and pancreas from fry to fingerling.

Dorsal fin rays 27-51, anal fin rays 16-35 according to Holčík (1989) and 30-49 and 18-32 according to Berg (1948-1949) for Kura River fish. Gill rakers 15-36. Dorsal scutes 7-19, lateral scutes 23-50 and ventral scutes 7-13 according to Holčík (1989) while Berg (1948-1949) gives 5-13, 21-42, and 7-14 respectively for scutes from Kura River fish. Safid River fish have a higher number of lateral scutes than fish from the Kura River.

The chromosome number is 2n>200 (Nowruz Fashkhami, 1996), later amended to 2n=258 ± 4 (Nowruzfashkhami et al., 2000).

Sexual dimorphism

Females are heavier and longer than males of the same age.

Colour

The back is greyish-blue to dark blue, the flanks with a steel-blue sheen, the head is lighter than the back, and scutes are lighter in colour than the background, usually pale yellow in adults but copper-gold in young. The belly is off-white, sometimes slightly yellowish.

Size

Reaches 2.42 m, possibly 2.50 m, and 76 kg, possibly 80 kg. A specimen caught by the Bandar-e Torkeman fishery weighed 63 kg, as opposed to the usual weight of 18-20 kg (Abzeeyan, Tehran, 5(3 & 4):V-VI, 1994). Males migrating into the Volga River typically weigh 20-30 kg and females 30-35 kg (Vecsei and Artyukhin, 2001).

Distribution

Found in the Caspian Sea, migrating to the north but mainly in the south in the Kura River of Azerbaijan and rivers of Iran where it is more common than A. gueldenstaedtii (Ivanov and Katunin, 2001). Also in the eastern Black Sea as a distinct subspecies.

In Iran, it is found from the Astara River in the west to the Gorgan River in the east (Armantrout, 1980), but apparently not the Atrak River on the border with Turkmenistan (Berg, 1936). Distribution includes the Safid River (to Kisom and "Musachayu"), Shalman, Golchan, Langerud, "Djef", "Youssefabad", "Tchontchenan", Dehkah, "Polrud", Sorkhrud, Feridounkenar, Talar, Tajan, Neka, "Palarud", Babol, "Mirerud", and "Ferikhabad" (Kozhin, 1957; Rostami, 1961; Armantrout, 1980). Also reported from Kargan, Kopurchal, Golshan, Larim, Nirroud, Tazeabad, 12 Bahman, Nevissi, Iz Deh, and Hasan Kiadeh by V. D. Vladykov based on field work notes made in 1962. Reported more recently as occurring in the Gorgan, Babol and Aras rivers by Holčík (1989), in the Gorgan, Gharasu, Tajan, Babol, Haraz, and Safid rivers, Gorgan Bay, the southeast Caspian Sea, southwest Caspian Sea and south-central Caspian Sea by Kiabi et al. (1999) and Abdoli and Naderi (2009) and in the Safid River and Anzali Talab by Abbasi et al. (1999). It used to ascend the Aras River but numbers in Iranian reaches were always small. Some literature records of A. gueldenstaedtii may be this species.

Zoogeography

Presumably a relict of the past isolation of the Black-Caspian seas from the Mediterranean-Atlantic.

Habitat

This species predominately inhabits the southern part of the Caspian Sea but does not form dense concentrations. Catches do not exceed 10-20 fish in 30 minutes of trawling. In winter to spring it is concentrated in the eastern coastal region and moves north in summer. In spring, maturing fish are concentrated in the southwest (Legeza, 1973). There is no seasonal variation in depth distribution in the south Caspian Sea in contrast to the middle Caspian. It is found on silty bottoms in the south Caspian Sea, sometimes with a sand admixture, at a temperature range of 4.1-28.0°C and a salinity range in the sea of 8.59-14.2‰. This species is more stenohaline than the others, preferring waters with higher salinity as in typical marine Caspian water and is also more sensitive to lowered oxygen levels (Legeza, 1972). Kazemi et al. (2003) found that osmoregulatory ability and development of chloride cells increased during growth, enabling the fish to transition between fresh and more saline waters. Khodabandeh et al. (2007) found fry transferred from fresh water to 7.5 and 10‰ sea water experienced 100% mortality after one hour acclimation; cortisol treatment increased the ability of fry to withstand these salinities. Ivanov and Katunin (2001) in a trawl survey along the Iranian coast found 14.2 fish/trawl in the west and 6.7 fish/trawl in the east with undersized and juvenile fish in the west at 57 fish/trawl. The higher western catches were attributed to the presence of more rivers, in particular the Safid River. The general abundance of this species was 8.775 million fish.

This species prefers fast rivers for spawning and migrate long distances. In the Volga River they migrate at an average speed of 22.6 km/day. They may remain in fresh water after spawning for a year or more although most return to the sea. These freshwater fish overwinter in deeper holes and feed intensively on fishes, crustaceans and molluscs. Larvae move downstream immediately after hatching. Cultured fingerings can be released safely and optimally into the rivers and estuaries of Iran at an age of 33-35 days after yolk-sac absorption at a weight of 1.8-2.4 g and 6.2-7.5 cm length (I.F.R.O. Newsletter, 30-31:5, 2002).

Bahmani et al. (2001) have shown that broodfish caught by seines in the Safid River were less stressed than fish caught by gillnets in estuaries.

Age and growth

Maximum age for accidental catches in the Caspian Sea off Azerbaijan is 32 years but most (82%) are 14-23 years old. Maturity is attained between 8 and 13 years in the Kura River (Markarova and Alekperov, 1989). Most fish entering the Kura River to spawn are 7-34 years old and the main spawning population is 11-24 years. Mean lengths for Safid River fish are 161 cm for females and 141 cm for males. Females have a faster growth rate than males. Growth rate is faster than for A. gueldenstaedtii and in the Volga size and weight is considerably higher. The numbers of this species and A. gueldenstaedtii in the southern Caspian are about equal. Maximum life span is 48 years. Studies in 2007, however, when 50 stations were sampled in waters less than 10 m deep, found this species to comprise 82.7% of the absolute frequency and 59% of the biomass of the total sturgeon catch. A. gueldenstaedtii was last with 5.5% and 2.3% respectively (Iranian Fisheries Research Organization Newsletter, 51:2, 2007).

Von Bertalanffy growth parameters in Iranian females are L = 225 cm and K = 0.066 or 207 cm and 0.079 and for males 197 cm and 0.084 or 186 cm and 0.105 depending on the methodology used. Total mortality (Z) was 0.24-0.57 for females and 0.40-1.1 for males, natural mortality (M) was 0.04 for females and 0.06 for males, fishing mortality (F) was 0.47 for females and 0.34 for males, and optimum fishing mortality was (F) 0.16 for females and 0.34 for males (Iranian Fisheries Research and Training Organization Newsletter, 16:4-5, 1997). A sample of 31 males and 49 females from the Turkman Sturgeon Fishery Station in 2001 showed sexual maturity at more than 19 years for females and more than 17 years for males (Alavi et al., 2005). Fish taken at 9 fishing stations along the Iranian coast numbering 4689 individuals had a mean length of 139.1 cm for males and 153.4 cm for females, weight s19.95 kg and 29.09 kg respectively and an age of 14.15 years and 16.59 years respectively. The sex ratio was 1:2.2 in favour of females and the majority of females (89.6%) were at level IV maturity. An increase in sexual maturity of females occurred in autumn while males were most mature from June to September (Falahatkar, 2006). Samples taken from the whole Caspian shore of Iran from 2002 to 2004 numbering 11,480 fish had a length range of 90-240 cm and growth parameters L∞ = 230 cm and K = 0.058 year-1 (www.shilat.com, downloaded 28 February 2007).

Food

Diet is composed of molluscs, crustaceans including the introduced crab (Rhithropanopeus harrisii), worms, chironomids and fish such as gobies (Gobiidae) and small herrings (Clupeonella spp.). Fish are a large part of the food of young sturgeon at sea. Azari Takami et al. (1980) found adults to prefer fish, mostly gobies, followed by crustaceans and two clam species Abra ovata and Cerastoderma umbonatum in Iran. The zebra mussel is also eaten as evidenced by a mass of these small clams from the stomach of a 1.6 m, 35 kg female from Nevissi caught on 29 September 1962. Reportedly the food diversity of this species is much less than for Huso huso and Acipenser gueldenstaedtii. Commercial sized fish feed particularly in the northern Caspian Sea (Ivanov and Katunin, 2001). Hashemyan et al. (2005) found diet in A. persicus, A. stellatus and A. nudiventris in coastal waters of Mazandaran and Golestan at depths less than 20 m to consist of annelids (50.8%), amphipods (41.5%), small fish 4.8%), decapods (2%) and bivalves (0.9%). Fish shorter than 40 cm fed mostly on shrimps, polychaetes and gammarids, 41-80 cm fish fed on shrimps, gammarids, polychaetes, bivalves and smaller fish, while fish greater than 80 cm fed mostly on shrimps and smaller fish. Immature A. persicus, less than two years old, from fishing stations off Gilan fed on the benthic invertebrates, namely the polychaetes Hypania sp., Hypaniola sp. and Nereis sp., the cumaceans Pterocuma sp. and Stenocuma sp., the clam Abra ovata, and the crustaceans Paramysis sp. and Gammarus sp. Adults fed mostly on fish (gobies, smelts and herrings). Haddadi Moghadam et al. (2009) studied diet in fish collected in summer and winter in the south Caspian Sea from 2004 to 2006. Food items were fishes (Neogobius sp., Atherina caspia, Clupeonella cultriventris (= caspia) and invertebrates (polychaete worms such as Ampharetidae and Nereis diversicolor; crustaceans such as Gammarus and Paramysis; and the bivalve mollusc Abra ovata). The diet varied with season and size group and was similar to A. stellatus.

The account under A. gueldenstaedtii above gives some comparative details of diet.

Reproduction

This species was long confused with the chalbash, A. gueldenstaedtii, and was thought to be a late spring or early summer spawning population of that species. The spawning run follows that of A. gueldensatedti. Spawning runs are dominated by the spring form and winter fish are very rare (Artyukhin and Zarkua, 1986). Fecundity off Azerbaijan is up to 558,900 eggs (Markarova and Alekperov, 1989) but may reach 840,000 eggs. In the Safid River it attains 375,000 eggs. The eggs are brownish-grey and measure up to 3.8 mm in diameter.

The unusually large specimen caught by the Bandar-e Torkeman fishery gave 22 kg of caviar, almost 35% of the body weight (Abzeeyan, Tehran, 5(3 & 4):V-VI, 1994).

Spawning takes place at 15-25°C, mainly at 17-23°C, at higher temperatures than A. gueldenstaedtii (8-18°C). Spawning sites are gravel, pebble, clay or shell beds, depths are 2-20 m and current speeds 1.0-1.7 m/sec. Catches of what were probably this species in the estuary of the Safid River for the period 1928/29-1936/37 showed strong peaks in April and May with a minor peak in September and October (Vladykov, 1964). The Safid is the main spawning river in Iran (Aslaanparveez, 1993). Spawning takes place in southern Caspian rivers from April to June and again in August to September. There is a 2 month interruption in spawning in the Safid River during summer when water temperatures are 26-30°C. There is a period of at least 2-4 years before this species can spawn again. Incubation takes 3-5 days. Shafizadeh and Parivar (1999) state that most embryos hatch 82-87 hours after fertilisation, most of the yolk is absorbed 6 days after hatching and swimup fry appear from day 7 to 8 at 19-21°C. The timing of passage of fingerlings into the sea after a hatchery release into the Tajan River was found to be 12-72 hours after release with a peak migration at 0-3 a.m. Smaller fingerlings stayed longer in the river before leaving (Ramezani, 2003).

Egg size is positively correlated with larval length and weight, yolk sac volume, hatching time and duration of hatching time, but there was no correlation with mortality during yolk sac absorption or with mortality during the first feeding stage (Nazari et al., 2009). Imanpoor et al. (2009) found the average hydrated egg diameter was 3.64 mm, yolk diameter was 3.26 mm, surface-to-volume ratio was 1.65 and yolk sphere-to-perivitelline space ratio was 0.75, the latter two being very high. The metabolic rate was low and spawning can occur in low-temperature waters.

Asadi et al. (2006) have examined serum biochemical parameters that can be used assessing maturity and managing endangered species.

Parasites and predators

Mokhayer (1976b) reports gas bubble disease in Iranian sturgeons without specifying the species of sturgeon as well as the monogenetic trematodes Diclobothrium armatum and Nitzschia sturionis. Most of the data for parasites and diseases summarised under A. gueldenstaedtii above for Iran may well refer to this species. Soltani et al. (2000) examined parasites of this species in three locations in Gilan and found Cucullanus sphaerocephalus and Skrjabinopsolus semiarmatus had the highest prevalence and intensity. Eustrongylides excisus, Anisakis sp. and Amphilina foliacea were recorded for the first time from this sturgeon and diet was strongly correlated with diversity of parasites. Soltani and Kolbassi (2001) describe the use of different antigens for fingerlings against Aeromonas hydrophila septicaemia. Hajimoradloo (2002) records the nematode Cystoopsis acipenseris in juveniles at a frequency of 5.83%. Hajimoradloo and Ghorbani Nasrabadi (2003) found the prevalence of metazoan parasites in juveniles of this fish in the southeast Caspian Sea to be 10 species with Anisakis larvae the highest at 19.7%. Pazooki and Masoumian (2004) report on blood parasites form fish caught at Anzali, recording Cryptobia acipenseris and Haemogregarina acipenseris. These parasites caused no pathological effects in the wild fish but can lead to severe infections and cause anaemia on fish farms. Gorogi (2006a) recorded the nematode Cucullanus sphaerocephalus, the the digenean Skrjabinopsolus semiarmatus and the acanthocephalan Leptorhynchoides plagicephalus from Iranian waters. Sattari and Mokhayer (2005a; 2005b) recorded the occurrence of parasites in this species from the Iranian southwestern and central coast of the Caspian Sea. The species found were the nematodes Cucullanus sphaerocephalus, Eustrongyloides excisus and Anisakis sp., the cestode Amphilina foliacea, the acanthocephalan Leptorhynchoides plagicephalus, the digenean trematode Skrjabinopsolus semiarmatus, the monogenean trematodes Diclybothrium armatum and Nitzschia storionis and the crustacean Pseudotracheliastes stellatus. General conclusions were that the diversity of parasites was less in Iranian waters than in the northern Caspian Sea, perhaps a reflection of the more varied habitat, its productivity and the carbonate ions differing between the two regions. The diversity of parasite seems to have declined over time also, perhaps as a result of unfavourable environmental conditions, particularly in the freshwater ecosystem which limits the waters available for spawning and parasite acquisition. Shenavar Masouleh et al. (2006) found hatchery fingerlings to harbour Diplostomum spathaceum, Trichodina sp. and Gyrodactylus sp. Ebrahimi and Malek (2007) found the helminths Cucullanus sphaerocephalus, Skrjabinopsolus semiarmatus, Leptorhynchoides plagicephalus and Eustrongylides excisus. Haghparast et al. (2007) found Cucullanus sphaerocephalus and Skrjabinopsolus semiarmatus to have the highest incidence (80 and 55%) in digestive tracts of broodstocks. Masoumzadeh et al. (2007) examined broodstocks and found Cucullanus sphaerocephalus, Skrjabinopsolus semiarmatus, Eubothrium acipenserinum, Corynosoma strumosum, Leptorhynchoides plagicephalus and Amphilina foliacea. Rajabpour et al. (2008) recorded helminth parasites from fish at three coastal stations in the southeast Caspian Sea, namely the nematode Cucullanus sphaerocephalus and the digenean Skrjabinopsolus semiarmatus. Jalilpour et al. (2009) identified a wide range of fungi on eggs and larvae of fish from the Shahid Beheshti Sturgeon Rearing Centre. Bazari Moghaddam et al. (2010) examined larvae and fingerlings in the Shahid Beheshti Hatchery and observed development of parasitism from the ciliate Trichodina reticulata and the digenean trematode Diplostomum spathaceum after release into earthen ponds and the river respectively.

Economic importance

See also under A. gueldenstaedtii where much of the data on this species is subsumed. The average weight of eggs in this species in Iran is 4-6 kg per fish and these eggs are ideal for first grade caviar (Vladykov, 1964). This species has the largest abundance (61.9%), biomass (50%) and catch-per-unit-effort among all Acipenseridae in Iran in both 2003 and 2004 from sampling 85 stations at 2-100 m depths (followed by A. stellatus (Iranian Fisheries Research Organization Newsletter, 38:1, 2004)).

Catches of A. persicus declined in the Safid River after construction of a dam at Manjil which released water for rice farming and held back sediment, both important triggers for attracting spawning sturgeon. In 1962, flow was reduced to 7-10 cu m/sec resulting in water temperatures up to 29°C, destroying insects and crustaceans on which young sturgeon fed and making the river narrow and shallow (Vladykov, 1964). Many fish were attracted into the stronger flow of irrigation canals where they eventually died. Catches of this and other species also declined because of the introduction of the more efficient synthetic fibre gill nets in 1957 (Vladykov, 1964). In Iran this sturgeon is caught both in the sea and in rivers.

Catches in the Safid River in 1930/31-1934/35 peaked at 13,867 fish in April with 10,693 fish in May and 3433 fish in March and an annual total of 32,700 fish (Berg, 1948-1949). Holmes (1845) and Eastwick (1864) reported on fishing for sturgeon in the Safid River. The principal method in the first half of the nineteenth century was to stretch 100 foot (30.5 m) lines across the very shallow, rapid and murky river with 1 yard (0.9 m) lengths of line attached at intervals of about 2 feet (0.6 m). These lengths of line were armed with large hooks which snagged the migrating sturgeon. Sturgeon up to 5 feet (1.5 m) were caught from February to April. At the beginning of February about 100 fish were taken each day, rising to 600-800 at the end of the month, 800-2000 in March and to 3500-3800 per day in April. After May sturgeons had little or no roe. About 125,000 fish were taken annually and sold for their flesh, caviar and isinglass.

Keyvanfar and Nasrichari (1999) state that from an average 2000 t annual catch over 10 years (1980-1990) 25% of meat and 24% of caviar were from this species while 17% of meat and 14% of caviar were from A. gueldensatedtii. This species produces 51% of Iran's caviar production (I.F.R.O. Newsletter, 30-31:5, 2002). Catches in the Kura River from 1974-1978 varied from 90 to 220 tonnes.

Extensive studies have been carried out on this species, either on hatchery specimens to improve their survival or using hatchery specimens as experimental organisms. These studies include rearing using earthworms (Kazerooni Monfared, 1995); ideal stocking densities in tanks (Derakhshandeh Ghazi Mahale, 1997); stress during transport and confinement of brood stock as evaluated using blood samples (Bahmani et al., 2000; Bahmani and Oryan, 2004); on growth performance using Daphnia magna and Artemia nauplii as food for fry (50% Artemia and 50% Daphnia given at 70% larval body weight was the best), and on osmoregulation during restocking (Jabbarzadeh Shiadeh et al., 2000); procedures against infectious diseases using antigens from Aeromonas hydrophila which causes septicaemia (Kalbassi et al., 2000); on effective stocking density of eggs and larvae in incubators and rearing tanks (Mohseni et al., 2000); haematological variables in juveniles and adults at different water temperatures (Pourgholam and Saeidi, 2000); on optimum feeding rate for fingerlings (Yousefpour Pirbazari et al., 2000); on blood parameters for fingerlings in a Gilan fish farm (Shahsavani et al., 2001); a histological study of the intestines (Sheibani and Pousti, 2001); sperm has been cryo-preserved to conserve the gene pool (Vecsei and Artyukhin, 2001); on clove oil having no significant difference with MS222, an anaesthetic used in fish farms (Abtahi et al., 2002; 2003); food and feeding of fingerlings after release and their travel time to the estuary (Kamali and Imanpoor, 2002); the relation between biochemical composition of eggs and their fertilisation rate (Mohammad Nazari et al., 2002); changes in the levels of sex steroids as oocytes developed (Nazari et al., 2002); nutrition in fish ponds where cladocerans and chironomids were staples and copepods and their nauplii were secondary items (Aslan Parviz and Aghaei Moghadam, 2003; Aghaei Moghadam and Aslan Parviz, 2006); the enhancement effect of ozone and physical treatment on the hatching rate of eggs (Ghomi et al., 2003); purification and partial characterisation of serum immunoglobulins (Kalbassi et al., 2003); physiological studies on the liver oxidase system (Karimzadeh et al., 2003); toxicity of the insecticide diazinon to fingerlings (Pazhand et al., 2003); dietary levels of fat and protein effecting growth and chemical composition of fingerlings (Ebrahimi et al., 2004; Mohseni et al., 2007); on sperm motility (Hadi Alavi et al., 2004); the identification of fatty acids in the flesh and the effects of long-term freezing on them (Hedayatifard and Moini, 2004); effect of temperature on fertilisation percentage achieved by broodstock (16.1-18.0ºC was optimal)(Hosseini Najd Gerami and Hajimoradlu, 2004); the effect of the timing of first feeding with live food on growth and survival of larvae (Kordjazi et al., 2004); determination of the 96h LC50 of Saturn, a herbicide, and Malathion, an insecticide, at 0.007 and 10 mg/l respectively (Nezami et al., 2004); histology of the gut from hatching to 56 days (Pahlavan Yali et al., 2004); levels of zinc and copper in muscle tissue and caviar (Sadeghird et al., 2004); reproductive conditions of broodstock and when they should no longer be used (Hosseini Najdegrami et al., 2005); the optimal weight and length for release of fingerlings into rivers and estuaries (1.8-2.4 g, 6.2-7.5 cm, 33-35 days after yolk sac absorption)(Kazemi et al., 2005); the timing of initial feeding in relation to behaviour (negative phototaxis and assumption of a benthic life at 5-6 days post-hatching) and expulsion of the melanin plug (larvae can feed with it present so expulsion cannot be used to determine active feeding)(Kordjazi et al., 2005); sperm density and fertilisation rate (Nazari et al., 2005); the toxic effects on fingerlings of various pollutants such as the oil products phenol and 1-naftol, the herbicide butachlor, and polyaromatic hydrocarbons from oil wells in the Caspian Sea (Nezami et al., 2005; Padjand et al., 2005; Soltani et al., 2006); a macroscopic and microscopic study of the spleen and and associated lymphatic tissue (Sheibani, 2005); evaluation of hydrogen peroxide against malachite green (possibly toxic and teratogenic) for fungal disinfection of eggs showed the former to be superior (Vahabzadeh et al., 2005); antifungal studies on eggs comparing the utility of formalin, malachite green and potassium permanganate in fish farms, the latter being safest for controlling Saprolegnia (Abtahi et al., 2005); sperm studies evaluating ionic composition and osmolality of seminal plasma, sperm density and motility in regard to sperm cryopreservation (Alavi et al., 2006); inulin-like growth factor-I inducing oocyte maturation (Bahrami Kanagar et al., 2006); the micro-cesarean method of extracting eggs from brood stock was better than conventional methods (Feyzbakhsh et al., 2006); the use of rotifers (Brachionus plicatilis) in conjunction with Artemia nauplii as food for larvae (Haddai Moghadam, 2006); use of oxolinic acid bioencapsulated in Artemia urmiana as a means to increase resistance to Aeromonas hydrophila infection in larvae (Hajimoradlou and Agh, 2006); studies on blood serum osmotic and ionic regulation in wild adults and reared juveniles, important in understanding the best use of water with different salinities in commercial rearing of this species (Kazemi et al., 2006); induction of ovulation using glycerin as a solvent for hypophysis powder proved better than physiologic serum (Noroozi et al., 2006); feeding formulated diets to larvae and juveniles in hatchery rearing (Pourali Fashtomi and Mohseni, 2006); establishing blood serum parameters as tools in disease prognosis and control (Shahsavani et al., 2006a, 2006b); the maximum allowable concentration of Safid River sediments as determined in aquaria was 1536.74 mg/l (Yosefi Garakoei et al., 2006); Abedian Kennari et al. (2007) on use of Daphnia magna enriched with cod liver oil as a source of highly unsaturated fatty acid on growth, survival, stress resistance and fatty acid composition of larvae; the effect of stripping frequency on ionic content and osmolality in seminal plasma composition (Alavi et al., 2007); details of sperm morphology in comparison to that of fil mahi (Baradaran Noveyri et al., 2007); comparison of the efficiency of the Yushchenko and Azarakhash incubators, the latter being better in terms of fertilisation percentage, mortality rate, active feeding and survival (Farabi et al., 2007); ability of Artemia urmiana to act as a carrier of oxolinic acid, a drug used to combat infection in fish larvae (Ghorbani et al., 2007); use of probiotic bacillus bioencapsulated with Artemia urmiana nauplii to increase growth of larvae (Jafarian et al., 2007); fatty acid composition in fresh and frozen tissues, concluding cold storage should not exceed 12 months (Moeini and Hedayatifard, 2007); variations in meat quality using dry and mix salting (salt and 1% madder) (Seyfzadeh et al., 2007); propagation efficiency of broodstock from two farms in Mazandaran and Golestan were shown to be different (Yousefian and Farabi, 2007); Askarian et al. (2008) examined the gastrointestinal tract for lactic acid bacteria and found the population levels to be significantly lower than in Huso huso;on amino acids in food pellets increasing consumption (Jafari Shamushaki et al., 2008); fertilising ability of cryopreserved spermatozoa (Alipour et al., 2009); on serum biochemical parameters (Asadi et al., 2009); on the median lethal concentration of suspended sediment from the Safid River, this pecies showing higher tolerance than A. stellatus (Garakouei et al., 2009); isolation of Lactobacillus species, which ferment carbohydrates, from the intestine (Ghanbari et al., 2009); changes in fatty acid composition after freezing and long-term cold storage (Hedayatifard and Keyvan, 2009); use of Artemia urmiana enriched with the essential fatty acid docosahexaenoic acid and its effects on growth, survival and composition of larvae (Hafezieh et al., 2009); the important influence of temperature on hatching time, start of exogenous feeding, growth performance and survival of larvae (Jalali et al., 2009); immunolocalisation of gill chloride cells used in ionic and osmotic regulation (Khoushnoud et al., 2009); varied effects of egg size on length, weight growth and survival of prelarval and early feeding stage (Nazari et al., 2009); recommended use of methyl paraben as a safe preservative in caviar infected with the bacterium Clostridium botulinum (Salmani et al., 2009); regulation of water temperature during the embryonic period, temperatures of 15-18ºC being the upper limit of thermal optima (Soleymani and Karimabadi 2009); fish effects of cooking methods on the physico-chemical and nutritional and digestibility properties of fillets (Alipour et al., 2010); identification of 13 fungal species in cultivated and natural populations (Firouzbakhsh et al., 2010); positive effects of Artemia urmiana enriched with highly unsaturated fatty acids on growth, survival and fatty acids composition of larvae (Hafezieh et al., 2010); the effects of sex steroids on hormonal control of reproduction (Hajirezaee et al., 2010); anaesthetic effects of clove essence (400 p.p.m. and 24ºC was best treatment and for recovery) (Imanpoor et al., 2010); the impact of plasma sex steroids on gonad development (Nazari, 2010); inhibitory effects on lipid oxidation (or rancidity) of ascorbic and citric acids compared with vacuum packaging in frozen fillets; Ghanbari and Jami (2011b) on Lactobacillus species from the guts; etc.

Conservation

See also under A. gueldenstaedtii. Catches in the sea off Iran are made with large seines and gill nets and many juveniles and fish below legal size are taken. Netting of sturgeon along the coast of Iran has been banned and hatchery production in Iran is directed to this species to maintain stocks. Moghim et al. (no date) note that juveniles of this species are caught in the beach seine fishery for other species in Mazandaran. During 2001-2002, 23,760 seine hauls had a by-catch of 54% for this species among sturgeons captured. Moghiem (2003) found that catch-per-unit-effort fluctuated from 2.249 to 2.971 kg over the previous decade, mean length, weight and age declined, the age structure changed with younger fish increasing in numbers, and catches showed an increase. Alavi et al. (2005) found overfishing of females in their sample from the Turkman Sturgeon Fishery Station.

Abdolhay et al. (2006) report on 1062 adults caught in 1998 of which 581 fish were injected with hypophysis extract and produced 22.5 million fingerlings while in 2002, 802 were caught and 538 produced 12.3 million fingerlings.

Hormonal studies are used to select fertile broodstock to ensure effective aquaculture (Mojabi et al., 1999; Safi et al., 1999) and other studies relevant to hatchery success, and thus conservation, are listed above. Nezami et al. (2000) maintain that sea-ranching has restored this species in Iran. Moghim et al. (2001) have used ultrasonography to determine sex and maturity of this species as there are no obvious external sex characteristics. Sex and maturity determination were accurate at 100% and 98.6% respectively, confirmed by necropsy, and thus would prevent the loss of male and immature female fish if the technique were used in the caviar fisheries.

This species is now found in the northern Caspian Sea, the fish being from Iranian stocking programmes (Kottelat and Freyhof, 2007).

Amini (2005) and Abdolhay and Tahori (2006) summarise hatchery production for this species:-

Process/Year 2000 2001 2002 2003 2004
Female broodstock captured 661 591 620 2056 742
Injected broodstock 437 492 528 1288 436
Spawning rate* (%) 81 86.5 410 (sic) 80 85
Fertilisation rate (%) 72 76.1 83 71 75
Survival rate in incubators (%) 56 52.6 75 50 64
Survival rate in tanks (%) 76 76.4 53 67 74
Stocking density in ponds (fish/ha) 84,076 89,131 76,000 97,941 95,661
Survival rate in ponds (%) 56 47.4 56 52 56
Fingerling production 13,711,199 16,278,595 12,301,214 18,388,962^ 17,412,529

* Rate of response to hormone injection; ^ 18,288 in Abdolhay and Tahori (2006)

Studies on heavy metal contamination (Zn, Cu, Cd,, Pb and Hg) of both flesh and caviar showed levels were below the maxima allowed for consumption, based on international standards (Sadeghi Rad et al., 2005; Amini Ranjbar et al. (2003), Amini Ranjbar and Shariat, 2006; Sadeghi Rad et al., 2009).

Lelek (1987) lists this species as endangered. Extinct in Turkey (Fricke et al., 2007). Kiabi et al. (1999) consider this species to be vulnerable in the south Caspian Sea basin according to IUCN criteria. Vecsei and Artyukhin (2001) list it as endangered with the IUCN. Criteria include commercial fishing, abundant in numbers, habitat destruction, widespread range (75% of water bodies), absent in other water bodies in Iran, and present outside the Caspian Sea basin. Mostafavi (2007) lists it as vulnerable in the Talar River, Mazandaran. Kottelat and Freyhof (2007) state that there is likely no natural reproduction in Iranian waters, fish being from artificial stocking programmes.

Further work

Fresh samples of sturgeon from Iranian rivers should be examined systematically and with care to determine if they are indeed this species and not A. gueldenstaedtii. A detailed comparative study of the morphology of this species and Acipenser gueldenstaedtii in Iran would enable the young and adults to be clearly distinguished as well as stocks within each species as a management tool.

Sources

Holcik (1993) and Shariati (1994) give accounts of this species in Farsi. See also under family above.

Iranian material: Hatchery adults examined at Bandar-e Anzali.

Acipenser ruthenus
Linnaeus, 1758

Found in the Caspian Sea basin but no records from Iran proper. Single specimens have been recorded as entering the Kura River of Azerbaijan and fishermen reported one fish from off Soviet Astara in 1929 (Berg, 1948-1949) on the border with Iran. The import of 30,000 fingerlings and 20 male parent stock of this species to Iran for artificial reproduction was envisaged in an agreement with the Russian Research Centre of Commercial Sturgeon Reproduction in 1995 (Iranian Fisheries Research and Training Organization Newsletter, 9:3, 1995). Tatina et al. (2010) studied effects of dietary vitamins C and E on haematological and biochemical parameters in this fish in the breeding centre in Rasht.   Acipenser primigenius Chalikov, 1944 is a hybrid of this species and Acipenser gueldenstaedtii (Eschmeyer et al., 1996). The Farsi name is استرلياد (esterliad). Listed as Endangered in the Volga River (Peterson et al., 2009).

Acipenser stellatus
Pallas, 1771

Common names

ازون برون or اوزون بورون (uzun burun or ozoonboroon = long nose), دراكول (= derakul or darakul); tirij (after Wossugh-Zamani (1991a), meaning shaped like an arrow; see also A. persicus); سوروگا (= sevruga or sevroga), سگ ماهي (sag mahi), ماهي خاويار (= mahi-ye kaviar, meaning caviar fish), puze draz.

[uzunburun, Kur uzunburun for natio cyrensis, ag-balyk, all in Azerbaijanian; tirana in Turkmenian; sevryuga, sevruga or stellate sturgeon (this term also includes A. nudiventris with small eggs for fisheries statistics), yuzhnokaspiiskaya sevryuga or South Caspian stellate sturgeon, both in Russian; star or starred sturgeon].

Systematics

Originally described from the Volga River near Simbirsk.

Acipenser seuruga Güldenstädt, 1772 from the Caspian Sea, Acipenser hellops Pallas, 1814 from the Black and Caspian seas, Acipenser Helops Pallas, 1814 from the Araks River, and Acipenser Ratzeburgii Brandt in Brandt and Ratzeburg, 1833 from the Caspian Sea at the mouth of the Emba River, are synonyms.

Acipenser stellatus stellatus natio cyrensis Berg, 1932 is described from the southern Caspian Sea and tributary rivers but has no taxonomic status as an infrasubspecific rank. Morphologically, this Kura River form is similar to north Caspian members of the species, differing principally in postorbital distance. Growth and fecundity are lower in the Kura form and spawning time is different. M. Poorhazemi (Pourkazemi) finds that A. stellatus is highly polymorphic with more than one population using molecular techniques (Iranian Fisheries Research and Training Organization Newsletter, 14:4-5, 1996). Norouzi et al. (2009) used microsatellite markers to determine that there is more than one population in the south Caspian Sea which has importance in terms of stock management, restocking and conservation. Shabani et al. (2003; 2006) found no significant differences between Volga River and Gorgan, Tajan and Safid River fish of Iran when examining mtDNA. Norouzi et al. (2008) and Norouzi and Pourkazemi (2009) examined the population and genetic structure of this species in Iranian waters using microsatellite markers and found evidence for at least three populations, particularly a separate one in the Safid River, and probably more than one in each river such as the Safid and Gorgan rivers.

A hybrid with Acipenser nudiventris is reported from the Safid River (Nedoshivin and Iljin, 1927). Artificial hybrids with Huso huso have been produced in Mazandaran for aquaculture projects (Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 6, 1996).

Key characters

This sturgeon has a long snout (59-65% of head length) with a pointed tip in contrast to the short snout and rounded tip in A. gueldenstaedtii and A. persicus. The continuous lower lip in A. nudiventris and the large crescentic mouth in Huso huso distinguish these species.

Morphology

The lower lip is interrupted at its centre, barbels are not fringed, are short, and do not reach the mouth but are closer to the mouth than the snout tip.

Dorsal fin rays 38-54 and anal fin rays 20-40; or 40-54 and 22-35 respectively in the Kura for natio cyrensis (Berg, 1948-1949). Dorsal scutes 9-16, lateral scutes 26-43 and ventral scutes 9-14. There are smaller scutes between the main rows. Gill rakers 24-29, usually 25-26 in natio cyrensis. Chromosome number 2n=115 ± 1 (Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 43, 1996), 2n=118 ± 2 or 113 ± 1 (Iranian Fisheries Research and Training Organization Newsletter, 8:5, 1995) or 2n=118 ± 1 (Nowruz Fashkhami, 1996), 2n=114 (Nowruz Fashkhami and Khosroshahi, 1999); 2n=146±6 (Chicca et al., 2002). Sheibani (2003a) described the anterior digestive canal of this species.

Keyvanfar (1986) found a transferrin polymorphism in the serum proteins of this species but not the other Iranian species of sturgeon and Keyvanfar (1988) found several variants corresponding to transferrin in the other species.

Sexual dimorphism

Females are larger than males of the same age; in the Ural River 1.3-1.6 times larger. Head depth and preanal distance differ between sexes in Kura River fish but only when gonads are ripening. Abdurakhmanov (1962) reports a longer anal fin, snout, and snout tip to barbel distance in males, and a longer predorsal length, preanal length, postorbital length and a greater caudal peduncle depth in females.

Colour

The back is dark grey, ash grey or cinnamon brown, almost black in some fish, and fades to a white belly. Flanks are yellowish-white. In small fish, the scutes are lighter than the adjacent body and so are distinctive. Sea fish are darker than river fish. An eyeless specimen, 1.11 m long, caught in Mazandaran was dark black (Abzeeyan, Tehran 4(7):V). The eyes were completely absent and their position on the head was covered with smooth bone.

Size

Attains about 2.21 m and more than 80 kg. Sternin and Doré (1993) cite a specimen of 2.9 m. Iranian captures averaged 1.3-1.4 m and 9-10 kg in the 1950s (Farid-Pak, no date). One of the largest specimens ever caught was 2.18 m long and was taken off the Astara River on the border of Azerbaijan and Iran in 1932. Much larger fish are known from archaeological sites of the 10th-13th centuries on the Terek River, up to 2.7 m (Tsepkin and Sokolov, 1971).

Distribution

Found in the Adriatic, Aegean, Black and Caspian seas and their drainages but the largest populations are in the Caspian. Generally found from the Astara River in the west to the Gorgan River in the east in Iran (Berg, 1948-1949; Kozhin, 1957; Armantrout, 1980) but not the Atrak River on the eastern Caspian border of Iran with Turkmenistan (Berg, 1936). Found in the Safid River at Kisom and the Mirerud (Derzhavin, 1934; Kozhin, 1957). It used to ascend the Aras River but numbers in Iranian reaches were always small (Berg, 1948-1949). The Kura River catch was up to 90% of the sturgeons taken. Rostami (1961) records this species from several localities on the Safid River and from the Golchan, "Djef", Youssefabad, Tchontchenan, Dehkah, Sorkh, Talar, Tajan, and Neka rivers. Also reported from Kargan and Hasan Kiadeh by V. D. Vladykov based on field work notes made in 1962. Reported more recently from the Gorgan, Gharasu, Tajan, Babol, Haraz, and Safid rivers, Gorgan Bay, the southeast Caspian Sea, southwest Caspian Sea and south-central Caspian Sea by Kiabi et al. (1999) and Abdoli and Naderi (2009) and from the Safid River and Anzali Talab by Abbasi et al. (1999).

Zoogeography

Presumably a relict of the past isolation of waters now encompassing the Black-Caspian seas.

Habitat

This sturgeon is found in large concentrations in the eastern coastal region of the south Caspian Sea in August-September with up to 25-30 fish taken in a single trawl, having moved south from northern waters. Ivanov and Katunin (2001) note the densest concentration in the per-estuary zone of the Gorgan River, with catches reaching 26 fish/trawl while along the central part of the Iranian coast catches did not exceed 4 fish/trawl. At the end of winter and particularly in early spring, uzun burun move onto the Iranian shore. Migrations between the Kura River lower reaches, the Safid River and elsewhere are reported. They usually does not descend below 100-130 m except along the southern shore of the Caspian Sea (Legeza, 1973) where they may descend to 300 m. Uzun burun are common only down to 50 m. There is no seasonal variation in depth distribution in the south Caspian Sea in contrast to the middle Caspian. They are often found in surface waters during the day, and retire to the bottom during the night. Uzun burun are found on silt and sand-silt bottoms but will also feed on sand and shell grounds. Temperature range is 4-24°C, in winter 7.5-10.5°C and 11.0-24.0°C in summer and fall, with an absolute range of 2.4-29.5°C. Water temperatures below 6°C are unsuitable for feeding however. Salinity range in the sea is 0.1-14.6‰ and this is the most euryhaline sturgeon in the Caspian Sea. This species is the best swimmer among sturgeons in the Caspian Sea in terms of power to body weight and in the Volga River migration speed averages 110 km/day (although progress is only 17.6 km/day because of the current).

The effects of diazinon on haematological parameters was examined by Khoshbavar Rostami et al. (2005) who also found the LC50 was 4.98 mg/l over 96 hours.

Age and growth

Maximum age for accidental catches in the Caspian Sea off Azerbaijan is 21 years but most are 8-13 years old. Males mature at 11-13 years, the youngest at 7 years, and females at 14-17 years, the youngest at 8 years in the Kura River. Populations in the Kura River and Iranian rivers take the longest time to mature, have a slower growth rate and lower fecundity. Vecsei et al. (2007) give a maturity range of 5-17 years. Like other sturgeons, this species does not reproduce every year and in the Caspian and there is a 3-4 year gap between reproductive periods in any individual. Females live longer than males. Maximum life span is about 41 years. Levin (1997) summarises the Volga spawning population as being age 6-28 years (11-16 years on average) with females 150-152 cm and 11-12 kg and males 128-130 cm and 6-7 kg. Spawning temperature is 16-22°C. The stock on the Iranian coast was estimated at 3.2 million fish weighing 18,500 tonnes with 6.7% of fish mature (Ivanov and Katunin, 2001).

Studies in 2007 along the whole Iranian coast when 50 stations were sampled in waters less than 10 m deep, found this species to comprise 11.8% of the absolute frequency and 38.7% of the biomass of the total sturgeon catch, second after A. persicus (Iranian Fisheries Research Organization Newsletter, 51:2, 2007).

Von Bertalanffy growth parameters in Iranian females are L = 213 cm and K = 0.062 or 188 cm and 0.104 and for males 190 cm and 0.083 or 171 cm and 0.113 depending on the methodology used. Total mortality (Z) was 0.52-1.1 for females and 0.62-1.1 for males, natural mortality (M) was 0.07 for females and 0.08 for males, fishing mortality (F) was 1.03 for females and 0.54 for males, and optimum fishing mortality was (F) 0.42 for females and 0.30 for males (Iranian Fisheries Research and Training Organization Newsletter, 16:4-5, 1997). Samples taken from the whole Caspian shore of Iran from 2002 to 2004 had growth parameters ∞ = 219 cm and K = 0.06 year-1 (www.shilat.com, downloaded 28 February 2007). Yelghi et al. (2007) found maximum age frequencies for fish from the southeastern Caspian Sea were were 9-13 years for male and 12-13 years for females. Brood fishes more than 15 years old formed little of the total catch. The oldest and largest individuals were 17 years and 156 cm for males and 27 years and 178 cm for females. Growth was negative allometric.

Food

Young specimens feed on crustaceans, older fish on chironomid larvae and the oldest specimens on fish (Rostami, 1961b). Azari Takami et al. (1980) found adults to consume gobies (Gobiidae) and kilka (Clupeonella) with the clams Abra ovata and Cerastoderma umbonatum as secondary items in Iran. In the Caspian Sea off Azerbaijan, Zarbalieva (1987) found that the polychaete worm Nereis diversicolor (82.7% by weight) dominated in the diet of sturgeons 20-80 cm long, being replaced by the mollusc Abra ovata (88.6%) at 90-120 cm and by Clupeonella spp. (65.1%) and Abra ovata (31.5%) at 125-140. Sturgeons 50-80 cm long also took the crab Rhithropanopeus harrisii (21.2%). Other foods include Rutilus rutilus (and presumably R. caspicus) Cobitis taenia, mysids, cumaceans, and amphipods. Gobies are generally of lesser importance than clupeids. Hashemyan et al. (2005) found diet in A. persicus, A. stellatus and A. nudiventris in coastal waters of Mazandaran and Golestan at depths less than 20 m to consist of annelids (50.8%), amphipods (41.5%), small fish 4.8%), decapods (2%) and bivalves (0.9%). Fish shorter than 40 cm fed mostly on shrimps, polychaetes and gammarids, 41-80 cm fish fed on shrimps, gammarids, polychaetes, bivalves and smaller fish, while fish greater than 80 cm fed mostly on shrimps and smaller fish. Haddadi Moghadam et al. (2009) studied diet in fish collected in summer and winter in the south Caspian Sea from 2004 to 2006. Food items were fishes (Neogobius sp., Atherina caspia, Clupeonella cultriventris (= caspia) and invertebrates (polychaete worms such as Ampharetidae and Nereis diversicolor; crustaceans such as Gammarus and Paramysis; and the bivalve mollusc Abra ovata). The diet varied with season and size group and was similar to A. persicus.

In rivers, juveniles feed on gammarids, chironomid larvae, mysids and worms. Spawning fish eat little or no food and, having used up much of their fat reserves, return to their feeding grounds in the sea immediately after spawning. This downstream migration varies from 70 to 80 km/day.

Reproduction

The peak migration in Iran is in April. There is also a peak run in fall (September-October) in the Kura River, and probably in Iran too (see below), but it is much less important than the spring run (Berg, 1959). Migrations in the Kura and Safid rivers can be found year round outside these peaks. The spring run in the Kura begins at about 10°C and peaks at 18°C, the runs decline in warmer summer temperatures and the fall run begins as water cools. Water level is also an important factor influencing runs and spawning. Water level fluctuations exceeding 0.2-0.5 m causes spawning to stop as fish migrate to deeper water. Summer and fall run fish do not spawn until the following year. Males arrive on the spawning ground before females and stay up to 6 weeks; females stay only 10-12 days. The Volga run begins in March-April with a peak in May but continues to October-November (Levin, 1997).

Up to 950,000 adhesive eggs are laid although in rivers of the southern Caspian absolute fecundity is lower, 35,400-362,900 eggs in the Kura River for example. Fertility is higher in the Volga compared to the Safid River (Iranian Fisheries Research and Training Organization Newsletter, 17:6, 1997). The spawning period in the Kura River is April-September at 15-29°C. Fish may leap out of the water during spawning and scrape their bodies on the bottom, leaving scratches and bruises. Eggs are deposited over gravel, pebbles, or stones mixed with shell fragments and coarse sand in the river bed or on flooded banks at a current velocity of 0.7-1.8 m/sec. A gravel bottom and a current speed of 1.2-1.5 m/sec are ideal. Eggs are round to ovate, brownish-grey and up to 3.2 mm in diameter. The adult loses 25-30% of its weight after spawning and females are only ready to spawn again after 5-6 years and males after 3-4 years. Spawning occurs at 15-26°C. Incubation takes 44-80 hours at 20-28°C. Young fish descend to the sea at 3-4 months of age but in some populations this occurs immediately after hatching, taking only 12-15 days.

Moghim et al. (2000) have used ultrasonography to determine sex and maturity stage of this sturgeon. Sex determination had a 97.2% accuracy and took 30 seconds or less per fish. This non-invasive technique reduces stress and enables immature females caught at sea to be released.

Parasites and predators

Niak et al. (1970) report infestations of the ciliate Trichodina sp. in sturgeons (species unspecified) in breeding ponds in Iran. Golvan and Mokhayer (1973) record the acanthocephalan Leptorhynchoides plagicephalus and describe a new species, Corynosoma caspicum, from this sturgeon in Iran. The coelenterate Polypodium hydriforme is recorded from the eggs of this sturgeon in the Safid Rud. Mokhayer and Anwar (1973) report on sturgeon parasites in general (see under Acipenser gueldenstaedtii). Mokhayer (1976b) also reports gas bubble disease in Iranian sturgeons without specifying the species of sturgeon as well as the monogenetic trematodes Diclobothrium armatum and Nitzschia sturionis. Larvae of the nematode Anisakis is reported from this species in Iran (Eslami and Mokhayer, 1977). Mokhayer (1989) reports metacercariae of the eye fluke, Diplostomum spathaceum from this species in Iran, which can cause complete blindness and death in commercially important species. Sattari et al. (2001) found the following parasites in fish from the southwest Caspian Sea: Skrjabinopsolus semiarmatus, Leptorhynchoides plagicephalus, Cucullanus sphaerocephalus, Eubothrium acipenserinum, Bothriomonus fallax, Eustrongylides excisus, Aniskais sp., Amphilina foliacea and Corynosoma strumosum. Hajimoradloo (2002) records the nematode Cystoopsis acipenseris in adult fish. Pazooki and Masoumian (2004) report on blood parasites form fish caught at Anzali, recording Cryptobia acipenseris and Haemogregarina acipenseris. These parasites caused no pathological effects in the wild fish but can lead to severe infections and cause anaemia on fish farms. Sattari and Mokhayer (2005a; 2005b) recorded the occurrence of parasites in this species from the Iranian southwestern and central coast of the Caspian Sea. The species found were the nematodes Cucullanus sphaerocephalus, Eustrongyloides excisus and Anisakis sp., the cestodes Eubothrium acipenserinum, Amphilina foliacea and Bothrimonus fallax, the acanthocephalans Leptorhynchoides plagicephalus and Corynosoma strumosum, the digenean trematode Skrjabinopsolus semiarmatus. General conclusions were that the diversity of parasites was less in Iranian waters than in the northern Caspian Sea, perhaps a reflection of the more varied habitat, its productivity and the carbonate ions differing between the two regions. The diversity of parasite seems to have declined over time also, perhaps as a result of unfavourable environmental conditions, particularly in the freshwater ecosystem which limits the waters available for spawning and parasite acquisition. Shenavar Masouleh et al. (2006) found hatchery fingerlings to harbour Diplostomum spathaceum, Trichodina sp. and Gyrodactylus sp. Ebrahimi and Malek (2007) found the helminths Cucullanus sphaerocephalus, Skrjabinopsolus semiarmatus, Leptorhynchoides plagicephalus and Eustrongylides excisus. Rajabpour et al. (2008) recorded helminth parasites from fish at three coastal stations in the southeast Caspian Sea, namely the nematode Cucullanus sphaerocephalus, the digenean Skrjabinopsolus semiarmatus, the acanthocephalan Leptorhynchoides plagicephalus and the cestode Amphilina foliacea. Barzegar and Jalali (2009), in their summary of crustacean parasites of Iranian fishes, recorded Pseudotracheliastes stellatus from this sturgeon.

Predators are most evident on the young and include Silurus glanis and various gobies (Gobiidae) while eggs are taken by Blicca bjoerkna, Pelecus cultratus, Gobio sp., and gobies.

Economic importance

Uzun burun are known from a Neolithic site on the eastern Caspian shore in the former Soviet Union from about 6000 years ago (Tsepkin, 1986).

This sturgeon provided the majority of the caviar produced in Iran according to reports from the 1960s and beginning of the 1970s (Vladykov, 1964; RaLonde, 1970b), 70% of the total catch according to commercial suppliers in 1995. It is reputed to have the tastiest flesh and also the best caviar (Ricker, 1970) but others maintain beluga caviar is the best. Farid-Pak (no date) gives an average yield of 1.5-2.0 kg for each female in the 1950s in Iran. Catch records for the Safid River in 1930-1935 showed that 31.7% of fish were caught in May, 18.1% in April and 9.6% in June, with a small peak in October of 7.9%. Nevraev (1929) records catches of this species varying from 22,278 to 43,593 individuals in the Astara region of Iran for the period 1901-1902 to 1913-1914, for the Safid Rud region 5536 to 12,670 individuals for the period 1899-1900 to 1913-1914, for the Mazandaran region 846 to 1490 individuals for 1906-1907 to 1913-1914, and for the Astrabad (= Gorgan) region 2613 to 5160 individuals for 1902-1903 to 1913-1914. Vladykov (1964) records average yearly catches in Iran of this species (including some A. nudiventris with small eggs) from 1927/28-1931/32 to 1957/58-1961/62 with ranges of 59,291-301,218 kg body weight (9.7-23.8% of the total sturgeon catch; 33.8% in another five-year period when weight was lower than the maximum shown here) and 8246-77,780 kg caviar (10.0-48.2%; total range 9.5-54.5%). RaLonde and Walczak (1970b) summarise yields for the years 1963 to 1967 in Iran of meat and caviar as 385.2 tonnes (100.4 tonnes), 450.8 (99.3), 436.6 (98.9), 564.4 (113.0), and 584.7 (106.5) respectively. Hassan Nia (1995) analysed the stocks of this species for a 61-year period (1927-1987) and calculated projected yields for the period 1988-1992. Actual yields proved to be the same as projected yields. The catch in the northern Caspian Sea reached 13,200 tonnes in the latter half of the 1970s.

This species has not been used as extensively as others for studies on physiology, biochemistry and aquaculture. Some works include Taleban et al. (1998) who studied consumption of this fatty fish and found a reduction in mean serum triglycerides and very low lipoprotein cholesterol, and an increase in high density lipoprotein cholesterol; Pourgholam and Saeidi (2000) investigated haematological variables in juveniles and adults at different water temperatures; Hedayatifard et al. (2003) studied variation in fatty acids composition in cold storage and found the best holding time was three months; Pazhand et al. (2003) on the toxicity of the insecticide diazinon to fingerlings; Sadeghird et al. (2004) examined levels of zinc and copper in muscle tissue and caviar; Padjand et al. (2005) examined the toxic effects on fingerlings of the herbicide butachlor; Hedayatifard and Moeini (2007) determined the levels of fatty acids in fresh and frozen samples and their effects on shelf life; Hedayatifard and Yousefian (2007) looked at shelf life and changes of lipid and fatty acid composition in frozen storage; Mokaremi Rostami et al. (2007) on the effects on juveniles of creosote on mortality rate and blood biochemistry with significant differences from controls; Alipour et al. (2009) on fertilising ability of cryopreserved spermatozoa; Bahmani et al. (2009) on seasonal fluctuations of sex steroids in farmed 7-year-old fish; Asadi et al. (2009) on serum biochemical parameters; Hedayatifard and Aroujalian (2010) on packaging and shelf life; etc.

The use of 2000 p.p.m. potassium sorbate in processing caviar from this species gives a better quality product than caviar without preservatives (Salmani, 1995).

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in aquaculture and aquaria and as food.

Conservation

See also under A. gueldenstaedtii. Lelek (1987) lists this species as vulnerable and Birstein (1993) as intermediate in status. It is now rare in the Safid and Gorgan rivers of Iran because of dam construction, which inhibits the spawning migration, and irrigation control structures near river mouths. The ban on sea fishing in 1962 by Soviet authorities led to an increased abundance of this species. Artificial spawning sites with gravel 3-10 cm in diameter have proved useful in the former U.S.S.R. and stocking is well established with up to 23 million young being released in the Volga area annually in the mid-1970s. However Veshchev (1995) reports that the population of this species in the Volga could be lost, and this doubtless mirrors the situation in other Caspian Sea states including Iran. About 30% of all individuals caught in the Caspian in the late 1980s were hatchery stock (De Meulenaer and Raymakers, 1996). Abdolhay et al. (2006) report on 193 adults caught in 1998 which produced 623,000 million fingerlings while in 2002, 290 breeders were caught and 67 produced 1.3 million fingerlings. Mohseni et al., (2000) have studied effective stocking density of eggs and larvae in incubators and rearing tanks in order to maximise production and avoid various morphological deformities. Moghim et al. (no date) note that juveniles of this species are caught in the beach seine fishery for other species in Mazandaran. During 2001-2002, 23,760 seine hauls had a by-catch of 37% for this species among sturgeons.

Khodorevskaya et al. (1997) summarises the decline of this species in the Volga and Ural rivers. The problems are the same for all sturgeons, namely flow alterations affecting the volume of water on the spawning grounds, reduction in numbers reaching the these grounds through poaching, and increased pollution affecting reproductive efficiency. Studies on heavy metal contamination (Zn, Cu, Cd,, Pb and Hg) of both flesh and caviar in Iran, however, showed levels were below the maxima allowed for consumption, based on international standards (Sadeghi Rad et al., 2005; Abtahi et al. 2007). The median lethal concentration of suspended sediment from the Safid River has been studied by Garakouei et al. (2009); who found this species showed a lower tolerance than A. persicus.

Kiabi et al. (1999) consider this species to be vulnerable in the south Caspian Sea basin according to IUCN criteria. Criteria include commercial fishing, abundant in numbers, habitat destruction, widespread range (75% of water bodies), absent in other water bodies in Iran, and present outside the Caspian Sea basin. Nezami et al. (2000) maintain that despite artificial spawning and fingerling production, restoration of this species in Iran was not very successful. Mostafavi (2007) lists it as vulnerable in the Talar River, Mazandaran. Critically endangered in Turkey (Fricke et al., 2007). Under IUCN and Appendix II of CITES, this species is now endangered (Vecsei et al., 2007).

Artificial breeding has been carried out with this species in Iran using hormones (I.F.R.O. Newsletter, 30-31:4, 2002). In contrast to other sturgeons, this species does not respond well to pituitary injections used to stimulate artificial reproduction. Pourkazemi (2006) examined haematological parameters and found wide fluctuations, with female spawners in particular differing in sexual maturity and physiological state. Although fish do respond to pituitary injections, the oocytes do not follow a normal course to maturity, remaining in the ovary. Oocytes at stage IV had overripe or degenerated oocytes. When overdosed with pituitary extract, ovulation occurred but oocytes were not mature and could not be fertilised. Degeneration of the egg membrane was found in 82% of spawners caught in the wild, presumably due to pollution. Baradaran Tahouri (1994) examined the effects of pond fertilisation on growth. Haddadi Moghaddam et al. (2001) studied the growth rate of this sturgeon in fertilised earthen ponds with added Daphnia. Shahsavani et al. (2001) determined blood parameters for fingerlings in a Gilan fish farm. Bahmani et al. (2006) recommended alleviating stress during capture, handling, transport and confinement, selecting breeders with suitable morphology and correct stage of sexual maturity, and using the hormone GnRH with domperidone as a substitute for pituitary extract. Luteinizing hormone releasing hormone analogue (LHRHa) was also found to be effective at 20.0-31.2 μg/kg body weight (Behmanesh, 2002). Kazemi et al. (2003) give a detailed histological study of the oocytes of this species. Caviar and fingerlings have been produced from farmed breeders (Iranian Fisheries Research Organization Newsletter, 49:3, 2006).). Sexual maturity was stimulated by injection of GnRH and anti-dopamine, eggs were extracted surgically, of which more than 80% hatched successfully using sperm taken by using tubes, and caviar and flesh harvested from one fish was comparable to natural samples.

Abdolhay and Tahori (2006) give fingerling production as:-

Process/Year 2000 2001 2002 2003 2004
Female broodstock captured 101 58 43 70 52
Injected broodstock 43 38 67 42 12
Spawning rate * (%) 60.4 50 49 63 50
Fertilisation rate (%) 55 58.6 51 51 83
Survival rate in incubators (%) 23 49.6 46 44 77
Survival rate in tanks (%) 54 93.5 60 72 77
Stocking density in ponds (fish/ha) 92,500 44,812 68,000 90,000 92,000
Survival rate in ponds (%) 12.2 67.7 38 61 86
Fingerling production (x 1000) 226 820 13,009 196 314

* Rate of response to hormone injection

Moghim et al. (2002) used ultrasonography to determine sex and maturity. This is important in management of endangered species when external sexual dimorphism is not apparent. Accuracy was 97.2% and was around 30 seconds or less per fish.

Further work

See under A. gueldenstaedtii.

Sources

See under the family account. Wossugh-Zamani (1991a) gives an account of this species in Farsi. Derzhavin (1922) and Borzenko (1942) are older works giving details of the biology of this species.

Iranian material: Hatchery adults examined at Bandar-e Anzali.

Comparative material: BM(NH) 1873.4.21:21-23, 2, 99.6-236.6 mm total length, Russia, Black Sea (no other locality data); BM(NH) 1929.8.7:4-5 and BM(NH) 1930.3.21:2, 3, 246.4-308.2 mm total length, Ukraine, Sebastopol, Black Sea (no other locality data).

Genus Huso
Brandt and Ratzeberg, 1833

This genus is characterised by a large and crescentic mouth (small and transverse in Acipenser) and by the gill membranes being joined to each other and free of the isthmus (joined to the isthmus in Acipenser). The snout is short and blunt although Caspian Sea stocks have a longer snout than Black Sea ones. The barbels are flattened laterally and gill rakers are rod-like. There are only 2 species in the genus, one in the Caspian, Black and Adriatic seas and one in the Amur River of eastern Asia.

Birstein and DeSalle (1998) using cytochrome b and 12S and 16S rRNA genes found that Huso may not be distinct from Acipenser. Vasil'eva et al. (2009) using cytogenetic and morphological characters also advocate reverting to the original genus Acipenser for Huso species.

Huso huso
(Linnaeus, 1758)

Common names

فيل ماهي (= fil mahi, filmahi or philmahi meaning elephant fish), beluga, beloga, سگ ماهي (sag mahi, meaning dogfish), ماهي خاويار (= mahi-ye kaviar, meaning caviar fish), mahi kaviar-e bozorg (= big caviar fish).

[bolka, Kur bolkasi for natio kurensis, ag-kulag-nyarya, gyuz'gi-burun in Azerbaijanian; doku (akvalyk) in Turkmenian; beluga in Russian; great, giant or European sturgeon].

Systematics

Acipenser huso was originally described from the Danube and rivers of Russia.

Huso huso caspicus Babushkin, 1942 was described as the subspecies of the Caspian Sea basin (with natio kurensis Babushkin, 1942 from the Kura River (also spelt incorrectly cyrensis and curensis)) but Berg (1948-1949) considered Caspian-Volga populations to be typical and this subspecies description as unnecessary. No types of Huso huso caspicus are known (Eschmeyer et al., 1996).

Huso ichthyocolla Bonaparte, 1846 is a synonym (Eschmeyer et al., 1996) and a nomen nudum (Holčík, 1989). Acipenser brandtii Günther, 1870 from the "Black and Caspian Seas, with their rivers" is a hybrid of Huso huso and Acipenser nudiventris based on Acipenser schypa (in part) of Brandt and Ratzeberg (Berg, 1948-1949; Eschmeyer et al., 1996). M. Pourkazemi in PADECO (2002) considers there are two sub-populations in Iran and Ghadirnejad et al. (2008) using microsatellite loci concluded that there were possibly two populations in the southern Caspian Sea.

Hybrids of Huso with Acipenser have been bred by the Aquaculture Department of the Iranian Fisheries Research and Training Organization (Iranian Fisheries Research and Training Organization Newsletter, 3:3, 1994; Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 6, 1996; Annual Report, 1995-1996, Iranian Fisheries Research and Training Organization, Tehran, p. 41, 1997) and natural hybrids with A. gueldenstaedtii, A. nudiventris and A. stellatus are reported from the Caspian Sea (Berg, 1948-1949).

Key characters

This species is identified by its very large, crescent-shaped mouth (small and transverse in other sturgeons) and the gill membranes being joined as a fold across the isthmus.

Morphology

The greatest body depth is slightly anterior to the middle of the body and large fish appear humpbacked. The lower lip is interrupted at its centre. Barbels are flat posteriorly, reach almost to the mouth and have foliate appendages. Experiments on ablading barbels (clipping one, two and four barbels) in 1+ age fish showed no growth differences with an unclipped control (Abasali Zadeh, 2003). The dorsal scutes are covered with skin in sexually mature fish, lateral scutes are smooth and ventro-lateral scutes hidden beneath the skin.

Dorsal fin rays 48-81 and anal fin rays 22-41. Dorsal scutes 9-17, lateral scutes 28-60 and ventral scutes 7-14. Scutes in adults may be reabsorbed. The skin is covered in small denticles. Gill rakers 16-36.

The chromosome number is 2n=118 ± 3 or 115 ± 1 (Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 43, 1996; Iranian Fisheries Research and Training Organization Newsletter, 8:5, 1995), 2n=116 ± 1 (Nowruz Fashkhami, 1996), 2n=118 ± 2 or 2n=116 ± 4 (Klinkhardt et al., 1995), 2n=117 (Nowruz Fashkhami and Khosroshahi, 1999). Sex chromosomes are absent or weakly differentiated in the genome and DNA markers cannot be used to sex fish; minor surgery has to be used (Keyvan Shokoo et al., 2004; Keyvanshokooh et al., 2007).

Sexual dimorphism

None found in morphometric and meristic characters although females are said to be longer and heavier than males of the same age.

Colour

The back is ash-grey, blue-grey to greenish or dark brown, sometimes black, fading to a white or cream belly. The contrast between the dark back and lighter rest of the body is marked. Young often have a metallic sheen which fades with age. The snout is yellowish.

Size

Attained weights of 1228 kg yielding 246 kg of caviar or 7.7 million eggs (Berg, 1948-1949), even 1600 kg (Farid-Pak, no date), and there are newspaper and other reports of fish 1200 kg and 6 m (Ottawa Citizen 14 May 1986) or even 3200 kg and 9 m but such large fish are not seen today and the largest sizes are probably exaggerations. Modern catches are mostly much smaller than these exceptionally large fish. A recent record with the specimen preserved in the Astrakhan Museum in Russia is given in Sternin and Doré (1993) for a fish from the Volga River in 1989 weighing about 980 kg, 4.3 m long and yielding about 110 kg of caviar (Iran News, 14 July 1998, gives 988 kg, 120 kg of caviar and an age of 60 years, presumably the same fish). A photograph of a 1908 capture at Astrakhan in Stein and Bain (1981) shows a fish weighing about 400 lbs (181.4 kg) containing 200 lbs (90.7 kg) of caviar worth more than $69,000 in 1981. Tsepkin and Sokolov (1971) give some examples of large fish from former Soviet waters. Birstein et al. (1997) consider this species to be the largest freshwater fish.

The mean weight of Caspian Sea fish decreased from 110 kg in the early 1970s to 57 kg in 1991 (De Meulenaer and Raymakers, 1996).

Up to 2.83 m and 450 kg generally in Iran (Azari Takami et al., 1980) but see below for news reports. Belugas up to 960 kg tried to enter the Atrak River in 1836 (Vladykov, 1964). The longest fil mahi caught in Iranian waters is apparently one taken on 23 February 1989 by Turkmen fishermen at Shilat-e Nahee 4 in Mazandaran (see Abzeeyan, Tehran, July 1991, page 3). It had a fork length of 4.5 m, a total weight of 725 kg and a caviar weight of 98.2 kg. This individual was worth U.S.$140,000 (Abzeeyan, Tehran, November 1992, page 57). The heaviest fish from Iran is one reported by Hossein Aimani at 3000 lbs (1360.8 kg) from near Babol in 1973 (www.amarillonet.com/stories/120599/bus_LQ7659.shtml, downloaded 7 March 2000). Mobayen (1968) gives the largest Iranian specimen as 4.2 m and 850 kg. Anonymous (1991a) and Sternin and Doré (1993) cite a fish of 1742 lb (= about 791 kg), 7.5 feet long (= about 2.3 m) and yielding 220 lb (= about 100 kg) of caviar from Iran in 1989, the largest caught for 20 years; this may be the same fish as the previous one as confusion in weights and lengths are common in reports of large fishes. Other large specimens were taken at Mahmudabad, Mazandaran on 28 October 1992, measuring 3.2 m, weighing 430 kg and with 61.2 kg of caviar (Abzeeyan, Tehran, November 1992, page 13), at Bandar-e Torkeman (= Bandar-e Shah) weighing 320 and 410 kg giving 110 kg of caviar for the two fish (Abzeeyan, Tehran 4(1):IIX, 1993), at Bandar-e Torkeman, Mazandaran on 27 March 1993, measuring 4.0 m, weighing 550 kg and with 81 kg of caviar (Abzeeyan, Tehran, 4(2):47, 1993), and in Mazandaran one measuring 3.0 m fork length and 3.4 m total length, weighing 960 kg and yielding 62.5 kg of caviar (Iranian Fisheries Research and Training Organization Newsletter, 5:8, 1994). Newspaper reports in 1996 listed a fish of 500 kg with 54 kg of caviar worth $107,000 and a fish caught in October 1997 at Babol Sar weighed 300 kg, measured 3 m in length and had 45.1 kg of caviar. In 1998, one fish 3.4 m long yielded 43 kg of caviar (Reuters), a fish caught off Bandar-e Torkeman on 2 February measured 3.75 m, weighed 405 kg and yielded 50 kg of caviar (IRNA (Islamic Republic News Agency), 3 February 1998), one caught off Bandar Anzali on 25 October weighed 360 kg, was 3 m long and yielded 24 kg of caviar and "meat" worth 3.6 million rials (IRNA, 26 October 1998), one caught off Nour, Mazandaran on 15 November measured 3.5 m, weighed 450 kg, yielded 53 kg of caviar and was 30 years old (IRNA, 16 November 1998), and one caught off Kianshahr, Gilan weighed 290 kg, was 3.5 m long and yielded 50.6 kg of caviar worth 100 million rials (IRNA, 24 November 1998). In 1999 newspaper reports included one caught off Bandar Anzali weighing 155 kg, carrying 31 kg of caviar worth $12,400 (IRNA, 31 October 1999), one caught off Talesh weighing 120 kg with 23.5 kg of caviar worth 150-200 million rials (IRNA, 5 December 1999), and one caught off Bandar-e Torkman weighing over 405 kg with over 52 kg of caviar worth 500 million rials (IRNA, 14 December 1999). One fish caught near Bandar Anzali in weighed 370 kg and yielded 51 kg of caviar (IRNA, 28 October 2002).

Distribution

Found in the Adriatic, Black and Caspian seas and their drainages. Derzhavin (1934) reported it from the Babol, Sorkh and Gorgan rivers but it was rare in the Safid River, although reported up to Kisom and quite abundant in the sea off its mouth. Nedoshivin and Iljin (1927) record this species from 10 river mouths while A. stellatus and A. gueldenstaedtii are reported from 18; the 10 river mouths are Yusufabad, Musachai, Hasan Kiadeh, Dastak, Safid, Kasumabad, Chalkarud, Sardabrud, Chalus and Kheirud. Kozhin (1957), Rostami (1961) and Armantrout (1980) stated that it enters the Astara, Safid, Babol and Gorgan rivers and the Anzali and Gorgan mordabs. It comprised only 0.5% in numbers and 2.5% in weight of the Safid River catch in 1914-1915 (Nedoshivin and Iljin, 1927). Large numbers were caught in the sea off Gasan-kuli in Turkmenistan near the Iranian border (Berg, 1948-1949). Also reported from Hasan Kiadeh by V. D. Vladykov based on field work notes made in 1962. More recently reported from the Gorgan and Safid rivers, the southeast Caspian Sea, southwest Caspian Sea and south-central Caspian Sea by Kiabi et al. (1999) and Abdoli and Naderi (2009), from the Safid River by Abbasi et al. (1999) and from the Safid, Gorgan and Tedjen rivers. This species was not caught in a survey along the Iranian coast in 2001 (Ivanov and Katunin, 2001). In 2004 there were plans to introduce this species to isolated, natural waters bodies in Fars Province (H. R. Esmaeili, in litt., 2004).

Zoogeography

Presumably a relict of past isolation of the Black-Caspian seas from the world ocean.

Habitat

This sturgeon is found in large concentrations in the eastern coastal region of the south Caspian Sea in all seasons. It is rare in trawl catches, possibly because it has a more pelagic life than other sturgeons. Fil mahi descend to greater depths than other sturgeons, 100-140 m in the Caspian and to 180 m in the Black Sea. There is no seasonal variation in depth distribution in the south Caspian Sea in contrast to the middle Caspian (Legeza, 1972; 1973). Only the young are found in shallow, warm areas. On the spawning migration, this sturgeon usually follows the deepest part of the river.

Most of this sturgeon's life is spent in the sea and it ascends rivers only to spawn. The new-born sturgeon returns to the sea. Farabi et al. (2007) examined salinity tolerance and physiology of juvenile fish in Iran. Only the youngest fish showed mortality on direct transfer from fresh to estuarine and Caspian sea water. Adults are typically found on silty or muddy bottoms in the sea but may be found on shelly and coarse sand at a temperature range of 5.6-29.3°C and depths of 5-140 m. In the southeastern Caspian it remains below 30 m in winter, entering shallower water at depths of 10-20 m in spring as the temperature ameliorates, dispersing throughout the southeastern Caspian in summer and migrating into Iranian waters in autumn (Legeza, 1972; Filippov, 1976). Depth distribution depends in large part on the available food supplies.

Oxygen requirements are high, averaging about 14 mg/l, but they can survive at 2-3 mg/l. Salinities up to 22‰ are tolerated. Feeding occurs over a temperature range of 0.5-30°C and the spawning migration at a range of 6-21°C. The highest densities in the southern Caspian Sea occur at 22-29ºC, feeding in winter at 10-12ºC (Caspian Sea Biodiversity Database, www.caspianenvironment.org).

Age and growth

Males become sexually mature at age 9-16 years and females at 12-22 years, varying with the spawning river. This is a very late maturation age among fishes world-wide. Spawning intervals are 4-7 years for males and 5-7 years for females (Vecsei et al., 2002; see below for other ranges but certainly intervals are long for a fish species). Spring-spawning females (see below) first spawn at 201-209 cm, 50-60 kg and 17 years. Winter-spawning females first spawn at 181-190 cm, 30-39 kg and 16 years. Most spring females are 230-300 cm long, weigh 80-160 kg and are 23-28 years of age. Most winter females are 201-300 cm, 50-160 kg and 17-26 years (Raspopov and Dubinin, 1990). Spawning populations have a complex age structure, the Volga River in 1936 had 50 age groups for example but only 28 in 1964. There has been a trend for spawners to be younger. Average catches in former Soviet waters of the Caspian Sea now weigh only 77 pounds (34.9 kg) each, a decline caused by overfishing (Los Angeles Times, Part A, page 1, 28 August 1993). A life span of 150 years was reputed for this species but the greatest known age for a Caspian fish is 75 years (Berg, 1948-1949). Most Caspian fish are now less than 20 years old and made up of individuals from re-stocking programmes (De Meulenaer and Raymakers, 1996). Raspopov (1993a; 1993b) gives the life cycle of Volga River fish as 56 years, although this is not the maximum age. Kura River sturgeon grow more slowly and mature later than sturgeon from the Volga River. Growth in this species is rapid with 1-year-old fish in the Caspian being 51 cm long and weighing 571 g. Growth is slower in the Caspian than the Black Sea because of the decrease in numbers of Alosa spp., the prime food item. Growth is also slower in the south Caspian than the north (Caspian Sea Biodiversity Database, www.caspianenvironment.org). Hedayatai et al. (2009) were able to correlate weight and length with immature male gonadal stage, but not for females, in work directed to reducing maturation time. Moghim et al. (2008) studied sex ratio along the Iranian coast for the years 1990-2003 and found females dominated at 60-80% of landed fish. Immature females decreased from 71 to 47% of the catch.

Levin (1997) summarises the spawning population of the Volga River over the last 10 years as follows although he notes this population is almost extinct. Rarely spawners enter between August and October and breed after a winter hibernation. Other fish enter from December to May with a peak from February to March. Peak spawning is in May with a downstream migration to the Caspian Sea from June to September. Females, comprising 20-24% of the spawning population, average 236-261 cm and 106-160 kg and are 17-21 years old with fish larger than 400 cm being very rare. Males are 199-204 cm and 48-55 kg and are 11-18 years old. Spawning occurs at 9-11°C.

Farid-Pak (no date) gives approximate weights for Iranian beluga of 75-100 kg and 2.0-2.5 m, and a yield of 17-20 kg of caviar per female. 2608 beluga from Astara in Azerbaijan averaged 168 cm for males and 192 cm for females.

Von Bertalanffy growth parameters in Iranian females are L = 320 cm and K = 0.065 for juveniles, 450 cm and 0.029 for the middle stanza and 533 cm and 0.023 for older fish and for males 270 cm and 0.086 or 302 cm and 0.072, depending on the methodology used. Total mortality (Z) was 0.21-0.67 for females and 0.22-0.75 for males, natural mortality (M) was 0.03 for females and 0.05 for males, fishing mortality (F) was 0.45 for females and 0.33 for males, and optimum fishing mortality was (F) 0.07 for females and 0.16 for males (Iranian Fisheries Research and Training Organization Newsletter, 16:4-5, 1997). Taghavi Motlagh (2001) gives more complete data (on which the previous summary was based) on growth, mortality and yield-per-recruit on this species from 1995 to 1999 in the Iranian Caspian Sea. He concluded that fishing mortality should be stopped. Maximum age in his sample was 46 years.

Food

In contrast to other sturgeons, this species is a pelagic predator as adults. Even sea birds and seals may be eaten. However, the introduced polychaete worm Nereis is now a mainstay of the diet of this species in the north Caspian Sea. Other foods are molluscs, formerly a main food, and small fish such as Rutilus rutilus (and presumably R. caspicus)and gobies (Gobiidae). Fish are the main diet item when large, invertebrates when young. This species needs to find thick concentrations of small or large fishes in order to feed actively; in the north Caspian these are kilka and fish on migration at fishways and in the midde Caspian spawning atherinids and commercial herrings (Polyanina et al., 1999). The fish found by Azari Takami et al. (1980) in Iran were gobies, Cyprinus carpio, Liza, and Rutilus. Gobies are a favourite food item but bivalves and crustaceans are taken if fish are absent. Filippov (1976) notes that large specimens eat sturgeons such as sevryuga, kopur (Cyprinus carpio), mullets (Mugilidae), birds such as coots, and baby seals and because of its pelagic life takes the clupeids Alosa braschnikowii and Clupeonella caspia and also the shrimp Leander adspersus. Crabs are also eaten. The principal food as percent by weight in the southeastern Caspian was Neogobius fluviatilis (= pallasi) (up to 78.1%), gobies accounted for up to 81.2% and fish 81.6-100%. Crustaceans accounted for up to 7.8% and molluscs only up to 0.2%. The cyprinid, Chalcalburnus (= Alburnus) chalcoides, is also eaten (Mageramov and Zarbalieva, 1989).

Reproduction

Roux (1961a) maintained that this species did not reproduce in Iranian rivers but Rudin (1966) said that they inhabited the Safid and Gorgan rivers. The main spawning river was the Volga as 90% of the Caspian stock reproduced there, travelling as far up as the Moskva River. Males arrive at spawning sites before females. Despite their size, these sturgeons may leap out of the water on the spawning run and possibly during spawning. Adhesive eggs are deposited on sandy substrates, with rocky and gravelly bottoms near the bank, in the strong current of mid-river (1.5-2.0 m/sec.). Water temperatures are 9-17°C and eggs develop in 9-10 days (Novikova, 1994; Vecsei et al., 2002). Spawning usually takes place at a depth of 4-15 m, sometimes as deep as 40 m. Weight loss after spawning may reach 50% and females are only ready to spawn again after 5-6 years and males after 3-4 years (4-8 and 4-7 years in Speer et al., 2000). The migration in the Volga River occurs year-round with peaks in spring (<30% of the stock) and autumn. The spring race reach the spawning beds in the same year, reproduce and return to the sea. The winter race, migrating in summer and fall, overwinter in the river and reproduce the following spring. The spring run is in March and April and the winter run in September and October. The chief spawning period in the Kura River is from the end of May to the beginning of June (Zakharyan, 1972) and fish were found as far up as Tbilisi (= Tiflis).

Fecundity reaches, exceptionally, 7,729,700 eggs but does not increase with age for fish of equal length and weight (Raspopov, 1987; Raspopov and Dubinin, 1990). Mean fecundity for the Volga stock was 531,600 eggs. Normal deposition of eggs is 500/sq m in the Volga but densities fell below 5/sq m in the 1980s, as low as 0.2/sq m and with an average of 1.5/sq m (Novikova, 1994). Kura River sturgeon are less fecund than Volga sturgeon. Egg diameter reaches 4.3 mm. Eggs are a dark silver and oval. Larvae hatch in 10-14 days, the yolk sac is absorbed in 10-14 days and feeding larvae move downstream at up to 60 km/day (Vecsei et al., 2002).

Parasites and predators

Niak et al. (1970) report infestations of the ciliate Trichodina sp. in sturgeons (species unspecified) in breeding ponds in Iran. Golvan and Mokhayer (1973) describe a new species of acanthocephalan, Corynosoma caspicum, and also Leptorhynchoides plagicephalus from this sturgeon in Iran. Mokhayer and Anwar (1973) report on sturgeon parasites in general (see under Acipenser gueldenstaedtii). Mokhayer (1976b) reports gas bubble disease in Iranian sturgeons without specifying the species of sturgeon as well as the monogenetic trematodes Diclobothrium armatum and Nitzschia sturionis. Pourgholam (1994) reports the coelenterate Polypodium hydriforme from this species caught on the Babol Sar and Bandar-e Torkeman fishing grounds in Mazandaran. Larvae of the nematode Anisakis simplex and the acanthocephalan Corynosoma strumosum are also reported from this species (Annual Bulletin 1993-94, Iranian Fisheries Research and Training Organization, Tehran, p. 48-49, 1995). Sattari et al. (2002) record Cucullanus sphaerocephalus, Eustrongylides excisus, Skrjabinopsolus semiarmatus, Anisakis sp., Eubothrium acipenserinum and Corynosoma strumosum, the fauna being similar to other sturgeons because of their piscivorous feeding. Gorogi (2006b) recorded the nematodes Cucullanus sphaerocephalus and Anisakis schupakovi, the cestode Eubothrium acipsenserinum and the acanthocephalans Leptorhynchoides plagicephalus and Corynosoma strumosum from Iranian waters. Sattari and Mokhayer (2005a; 2005b) recorded the occurrence of parasites in this species from the Iranian southwestern and central coast of the Caspian Sea. The species found were the nematodes Cucullanus sphaerocephalus, Eustrongyloides excisus and Anisakis sp., the cestode Eubothrium acipenserinum, the acanthocephalan Corynosoma strumosum, the digenean trematode Skrjabinopsolus semiarmatus. General conclusions were that the diversity of parasites was less in Iranian waters than in the northern Caspian Sea, perhaps a reflection of the more varied habitat, its productivity and the carbonate ions differing between the two regions. The diversity of parasite seems to have declined over time also, perhaps as a result of unfavourable environmental conditions, particularly in the freshwater ecosystem which limits the waters available for spawning and parasite acquisition. Shenavar Masouleh et al. (2006) found hatchery fingerlings to harbour Diplostomum spathaceum and Trichodina sp. Barzegar and Jalali (2009), in their summary of crustacean parasites of Iranian fishes, recorded Pseudotracheliastes stellatus from this sturgeon.

The fil mahi is so large that its predators are only effective on young fish. They include Sander lucioperca and Silurus glanis and, needless to say at all sizes, mankind.

Economic importance

This species provides the best caviar according to Borodin (1930). The large eggs fetch a higher price on the American market. Up to 80% (3000 kg in 2002) of the legal beluga caviar export is consumed in the U.S.A. (Hamilton, 2002). A 1227 kg specimen caught in Russian waters in 1924 gave 245 kg of caviar worth £189,350. In the 1990s, a 225 kg fil mahi could yield 22 kg of caviar worth $120,000 (Trickey, 1995). Catches in the Volga region in the 1970s were in the range 740-2650 tonnes and in the 1980s 460-900 t comprising 4.4-12.2% and 3.7-4.4% respectively by weight of the total catch of all sturgeons there. The highest catch in the Caspian Sea was in 1902-1907 (Birstein, 1993). Khodorevskaya et al. (1997) and Khodorevskaya (1999) summarise the decline in catches and make the startling observation that 96.3% of all fil mahi in the Volga River are hatchery reared.

Fil mahi were fished intensively off the Iranian coast in the southeastern Caspian and in 1950 amounted to 38.6% of the total sturgeon catch. During the five-year period 1957/1958 to 1961/1962 fil mahi catches in the Gorgan Division of the Iranian fishery varied between 86-90% of total Iranian catches. The Atrak River estuary area was particularly important for this species. Catches of the oldest age groups has declined and the proportion of young and immature fish has increased. Iranian rivers suitable for this sturgeon were the Safid and the Gorgan but both are now regulated so Iranian stocks are probably maintained by fish reproducing in the rivers of the former U.S.S.R. (Filippov, 1976). Fil mahi cannot be managed by Iranian authorities therefore. However the "Gharasoo" Research Station in Mazandaran is researching the culture and release of fil mahi up to 1 kg (Madbaygi, 1993b) and farming through pen culture in Gorgan Bay (Iranian Fisheries Research and Training Organization Newsletter, 11:6, 1996). Two million "roes" (presumably young fish) were released into the Caspian Sea from Mazandaran prior to 1 June 1995 with a further 2 million to be released later in the year (http://netiran.com/news/IRNA/html/950701IRGG08.html). In 1997, 852 fishermen were fishing for fil mahi on the northern Iranian coast (Anonymous, 1997c).

Farid-Pak (no date) gives the months of September-October and March-April as the most important for the fisheries of this species. Nevraev (1929) gives catch ranges of 109-3100 fil mahi individuals for the Astara region of Iran over the period from 1901-1902 to 1913-1914, for the Safid Rud region 104 to 730 individuals for the period 1899-1900 to 1913-1914, for the Mazandaran region 31 to 491 individuals for 1906-1907 to 1913-1914, and for the Astrabad (= Gorgan) region 688 to 1764 individuals for 1902-1903 to 1913-1914. Vladykov (1964) records average yearly catches in Iran of this species from 1927/28-1931/32 to 1957/58-1961/62 with ranges of 57,820-418,059 kg body weight (5.4-33.0% of the total sturgeon catch) and 2038-32,873 kg caviar (2.6-20.4%). There was an upward trend in caviar production from this species in the 1950s (Vladykov, 1964). RaLonde and Walczak (1970b) summarise yields for the years 1963 to 1967 in Iran of meat and caviar as 572.3 tonnes (40.1 tonnes), 583.5 (47.3), 575.8 (39.1), 458.1 (29.5), and 507.2 (30.0) respectively. A commercial house maintains (1995) that caviar from this species comprises only 3% of the total catch. Taghavi Motlagh (2001) noted a decline in the share of Iranian caviar production from 18% in 1971 to 4% in 2000.

This species has been studied in ponds as breeders are used to produce fingerlings which are then available as experimental fish for chemical and growth studies. Ghorbani et al. (2003) studied the influence of heavy metals on the level of alfa-amylase activity in the digestive tract and found decrease in enzyme activity was not significant. Karimzadeh et al. (2005) studied cytochrome P4501A1, a major isoenzyme in the monooxygenase system which can be induced by polycyclic aromatic hydrocarbon pollutants. Khoshbavar Rostami et al. (2006) studied the effects of polyaromatic hydrocarbons from Caspian Sea oil wells on 8.5 g fingerlings and found these chemicals to seriously affect the fish blood and enzyme systems. Khoshbavar Rostami et al. (2004; 2006) studied the organophosphate diazinon and its deleterious effects on haematological parameters in this sturgeon. Sharifpour et al. (2004) studied the effects of the insecticide endosulfan, sturgeon weighing 3-5 g showing irregular swimming, whirling, convulsions, with other conditions, and eventually death. Endosulfan is highly toxic to beluga fingerlings. Sudagar et al. (20050 examined the addition of betaine and methionine (an important nutrient and an enzyme) to the diet of juvenile beluga. The fish showed improved weight gain, weight gain percentage, specific growth rate, protein efficiency ratio, net protein utilisation, condition factor, survival, and price index at enrichment levels of 0.5% betaine and 1% methionine. Ghorbani et al. (2004) examined the influence of a series of microelements (zinc, nickel, cobalt, manganese, iron and copper) on the level of proteolytic enzymes and alkaline phosphatase activity (used for enzyme inmmunoassays) in the digestive tract of juvenile beluga. Most treatments showed the level of enzyme activity was less than the control. Shahsavani (2002) determined blood parameters of fingerlings from fish farms and found the fish to be healthy. Blood parameters are used to indicate physiological condition and sublethal stress due to endogenous and exogenous changes, hence the need to determine normal values. Askarian et al. (2006) looked at serum osmoregulatory parameters under different light regimes, one form of physical stressor in aquaculture of this endangered species. No differences in serum cortisol levels were found between treatments although elevations of serum cortisol, glucose and triglyceride occurred.in a continuous dark regime. Gafarian et al. (2007) used probiotic bacillus in the feeding of larval sturgeon and found that it positively affected feeding efficiency and levels of carcass nutrient composition. Khoshbavar-Rostami et al. (2007) examined the immune response to Aeromonas hydrophila bacterin. Soulati and Falahatkar (2007) looked at stress response in sub-yearlings exposed to air. Shamloufar et al. (2007) examined the sub-lethal effects of diazinon on haematological indices in juveniles. Akrami et al. (2008) studied the effect of prebiotic inulin levels and found it did not increase growth performance of juveniles. Askarian et al. (2008) examined the gastrointestinal tract for lactic acid bacteria and found the population levels to be significantly higher  than in Acipenser persicus. Hedayati et al. (2008) studied blood indices of fish cultured in brackish water. Hosseini et al. (2008) examined the organochlorine content of four sturgeon species and found fil mahi had four times more than the next highest species (A. nudiventris); generally pollutants had been reduced compared to previous studies but some specimens exceeded guideline levels for food. Soltani et al. (2008) found that 100-200 mg/kg of vitamin C was optimum for rearing this sturgeon. Baghfalaki et al. (2009) carried out studies on seminal plasma indices in order to improve short and long-term storage of semen. Darvish Bastami et al. (2009) found that addition of Daphnia and Artemia extracts had positive outcomes on growth in juveniles. Akbari et al. (2009) studied the use of sperm extenders and found that they prolonged spermatozoa viability in short-term storage and prolonged sperm motility. Ghanbari et al. (2009) isolated Lactobacillus species, which ferment carbohydrates, from the intestine. Jalali et al. (2009) found that Artemia urmiana nauplii on enriched with HUFA and vitamin C and fed to larval sturgeon improves some growth and stress tolerance. Seifzadeh et al. (2009) examined microbial quality of packaged fillets of this sturgeon. Askarian and Kousha (2008) examined food ration on the acute stress response, those receiving a high ration performing better. Alizadeh et al. (2009) studied effects of different diets on energy levels and gonad development for fish reared in inland brackish water, this environment proving suitable. Askarian and Kousha (2009) studied photoperiod in rearing year-old fil mahi evaluated by growth (no effect) and serum parameters (various individual responses to stress. Falahatkar et al. (2009) examined dosages of vitamin C that enhanced immune responses to disease. Sepahdari et al. (2009) found various skin lesions in fish fed a diet containing aflatoxin B1 (a naturally occurring fungal toxin). Sepahdari et al. (2010) found deleterious changes in liver tissues n fish fed a diet containing aflatoxin B1. Ghanbari and Jami (2011b) reported Lactobacillus species from the guts of this species.

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in aquaculture and aquaria, as food and in textbooks.

Conservation

See also under A. gueldenstaedtii. Critically endangered in Turkey (Fricke et al., 2007). Despite loss of 99% of the Volga River spawning beds to dam construction, natural reproduction increased over a recent five-year period, but continues to be dependent on the variable flow-regime (Raspopov and Dubinin, 1990). Novikova (1994) estimated the capacity of the Volga spawning grounds to be 9-11,000 fish. A major problem in the 1990s was poaching. Trickey (1995), referring to Russian stocks, expected a legal harvest of 4400 tonnes with poachers taking twice that amount. This legal and illegal catch is still less than catches of 20 years ago, primarily because of pollution. Birstein (1996) records the catch of the Volga delta hatcheries in 1995 to be only 35 fish, insufficient for artificial reproduction. Natural spawners are taken by poachers. The level of poaching in the Ural River is also high, and this was the only river where some natural reproduction was going on. The fil mahi has effectively stopped reproducing in the Caspian Sea.

Moghim et al. (no date) note that juveniles of this species are caught in the beach seine fishery for other species in Mazandaran. During 2001-2002, 23,760 seine hauls had a by-catch of 6% for this species among sturgeons captured.

Khodorevskaya and Novikova (1995) point out that cooperation among all the Caspian Sea states is needed to maintain this species along with an annual release of at least 20 million young from hatcheries. Fingerlings released per year from 1998 to 2002 range from 6.9 to 12.6 million for all Caspian states (CITES website). Spawning migrations are now seen only in the Volga and Ural rivers, the Kura, Terek and Sulak rivers no longer supporting stocks. The Volga migration was 25,500 fish weighing 2600 t in the early 1970s but has fallen to 11,700 fish weighing 750 t. The commercial catch fell from 2000 t to 500 t. In the Volga River 96.3% of the spawning population consists of hatchery fish although the Ural River maintains a naturally reproducing stock.

Since stocks are maintained mostly by artificial rearing, this sturgeon has been proposed for inclusion in the "Red Book of the U.S.S.R." which forms the basis for measures to protect species (Pavlov et al., 1985; Mina, 1992). Stocks have been increased through rearing and natural reproduction in the Ural River, the number rising from 9.6 million in 1976 to 15.3 million in 1983, so the status of this species was then regarded as acceptable. However Lelek (1987) and Birstein (1993) list this species as vulnerable to endangered. Kiabi et al. (1999) consider this species to be endangered in the south Caspian Sea basin according to IUCN criteria as does IUCN and CITES (Vecsei et al., 2002). The U. S. Fish and Wildlife Service lists it as threatened under the U.S. Endangered Species Act as of 21 October 2004 (http://news/fws.gov/newsreleases, (dated 20 April 2004) and downloaded 22 April 2004) and the Wildlife Service has been petitioned to make it endangered (Speer et al., 2000). Endangered status would stop importation of flesh and caviar to the United States. Suspension of trade in this species from the Black Sea basin by the U.S. Fish and Wildlife Service was instituted in 2005 (Federal Register, 2005) and imports from Iran are banned for political reasons along with other sturgeons. Criteria for the various status assessments include commercial overfishing (fishermen cannot even catch the set quotas), failure of regulatory oversight, few in numbers, habitat destruction, dams preventing spawning migrations, medium range (25-75% of water bodies), absent in other water bodies in Iran, poaching, pollution, diseases due to pollution, and presence outside the Caspian Sea basin. The World Wildlife Federation (WWF) listed this species as number 4 on the top 10 most endangered species in the world (www.extravalue.com/sturgeon.shtml, downloaded 13 March 2000). The species status may be changed to Appendix I on the CITES listing, when international trade in its caviar would be banned (Vecsei et al., 2002). The export quota for this sturgeon in the Caspian Sea 2004 was reduced to 4425 kg although an illegal harvest was still substantial (www.tehrantimes.com, downloaded 14 October 2004).

Illegal fishing from 1990 onward and cessation of hatchery releases will lead to loss of the stock unless an agreement between Caspian states can be reached to protect this species.

The invasion of the ctenophore Mnemiopsis has led to declines in the kilka (Clupeonella spp.) stocks, a prime food of fil mahi (Kideys, 2002).

Caviar from Russian caught fil mahi bought in New York stores has been examined for pollutant content (Boyle, 1994). Three stores carried caviar with 3.17-3.27 parts per million of DDT plus its metabolites DDD and DDE, 410-640 parts per billion of the PCB Arclor 1254, and 2.1-2.8 parts per million selenium. These values are below the U.S. Food and Drug Administration's action levels of 5 parts per million for DDT, 2 parts per million of PCB and 10-50 parts per billion of selenium in drinking water. Nevertheless they are cause for concern.

Various studies have been carried out on the aquaculture of this valuable sturgeon in Iran. Mohseni et al., (2000) have studied effective stocking density of eggs and larvae in incubators and rearing tanks in order to maximise production and avoid various morphological deformities.

Abdolhay and Tahori (2006) give fingerling production as:-

Process/Year 2000 2001 2002 2003 2004
Female broodstock captured 32 29 29 48 16
Injected broodstock 19 14 21 30 9
Spawning rate * (%) 74 71.4 62 65 77
Fertilisation rate (%) 55 65.5 65 54 65
Survival rate in incubators (%) 62 73.4 62 32 72
Survival rate in tanks (%) 80 62 56 100 79
Stocking density in ponds (fish/ha) 82,100 51,639 51,333 52,359 65,448
Survival rate in ponds (%) 73 51.3 67 43 59
Fingerling production (x 1000) 1900 640 24,037 42 146

* Rate of response to hormone injection

Mohseni et al. (2006) studied the best stocking density for rearing juveniles less than one year old weighing 92.09 g on average and one-year-old fish weighing 918.14 g on average. Stocking densities were 1.6, 2.8 and 4.0 kg/m2 for the juveniles and 1.5, 2.5, 3.5 and 4.5 kg/m2 for the older fish. Increased density had a negative impact on growth, body weight, specific growth rate and food conversion ratio in both experiments. Higher concentrations of fishes even had malformed caudal fins and body injuries from increased contact. Recommended stocking densities were 1.5-2.0 kg/m2 for fish up to 90 g and 2.5-3.0 kg/m2 for fish over 900 g.

Cage culture of fingerlings has been carried out in Gorgan Bay starting in 1992. Cages were 3200 sq m with a depth of 2.5 m and contained 11,500 fingerlings. Over 16-17 months average weight increased from 20 g to 1365.5 g, to a maximum of 2200 g. Mean fork length was 58.6 cm. Food in the first phase was a concentrate of ground carp and kilka but in later phases natural foods such as benthos and fry were used. The preliminary results indicate economic feasibility for cage culture (Iranian Fisheries Research and Training Organization Newsletter, 7:4-5, 1995; Annual Bulletin 1993-94, Iranian Fisheries Research and Training Organization, Tehran, p. 46-47, 1995).

Kamali and Farabi (2005) showed that juveniles weighing 20 g or more adapted better to concentrated feed in fibreglass tanks. Mohseni et al. (2004) studying growth rate, food conversion ratio and survival in fingerlings held in fibreglass tanks found these factors to be dependent on higher feeding frequencies (3, 5 and 8 times per day). Akrami et al. (2005) found Cladocera were the primary prey of fingerlings in earthen ponds with chironomid larvae and ostracods secondary prey, and the copepod Cyclops an occasional prey. Condition factor and growth decreased as weight and length of fingerlings increased. Growth was was positively allometric (b>3). Mohseni et al. (2005) found growth of fil mahi was better in fibreglass tanks but later in the rearing process the trend reversed and earthen tanks showed a better condition. Mohseni et al. (2006) examined the effects of feeding rates (1, 2, 3 and 4% of biomass) on various factors for fish weighing an average 867.9 g and fed for 100 days in fibreglass tanks. Increase in feeding ratio directly increased daily food consumption and negatively affected the feeding efficiency, food conversion ratio, specific growth rate and price index. When fish were given 2% of the body weight, one unit of meat was produced from 1.92 units of food. A second trial with feeding rates 0.75, 1.5, 2.5 and 3% took place with fish weighing 2096.1 g and fed for 125 days. Feeding with 0.75% produced one unit of meat per 1.82 units of food consumed. Fatemeh and Armin (2005) studied the effect of photoperiod on growth in one-year-old fil mahi. Extended day length had a positive effect on growth rate, specific growth rate, weight and length, and condition factor. The organophosphate diazinon was studied experimentally by Khoshbavar Rostami et al. (2006) as to its effects on haematological and biochemical factors of the blood serum of this fish. Falahatkar et al. (2006) experimented with various levels of vitamin C as a diet supplement and recommended 200 mg kg-1 during the first weeks of growth and development. Mohsen et al. (2008) found that diets supplemented with L-carnitine improved growth rate, feed utilisation and stimulated protein-sparing effect. L-carnitine is a vitamin-like compound found naturally in fishes and is involved transporting long-chain fatty acids in metabolism. Ahmadifar et al. (2009) found that dietary Ergosan had some positive effects on growth and haematological parameters (Ergosan comprises algines and polysaccharides known to strengthen the full range of natural defence systems in fish).

Nezami et al. (2000) maintain that despite artificial spawning and fingerling production, restoration of this species in Iran was not very successful. Abdolhay et al. (2006) report on 17 adults caught in 1998 of which 10 fish were injected with hypophysis extract and produced 1.08 million fingerlings while in 2002, 29 were caught and 21 produced 2.4 million fingerlings. Azari Takami (1999) cites production of 300-350 kg/ha in 40 days with 106,000 fingerlings produced per 15 females in 40 days with a release weight of 10-15 g. Spawning fish were captured in the sea as they no longer migrated into Iranian rivers and propagation results were not as good as in previous years (420-587 kg/ha in 25 days, 690,000 per 2 females, release weight 5-8 g). About 1 million fingerlings were released into the Caspian Sea. Iranian releases of fingerlings were 687,400 (1988), 406,100 (1999), 1,900,919 (2000), 700,000 (2001), 2,403,794 (2002) with ca. 4 million proposed for 2003 (CITES website). The annual release of fingerlings weighing 3-5 g into the Caspian from Iran is 1-2 million fish and some of these are tagged for future studies (Iranian Fisheries Research Organization Newsletter, 39:1, 2004). In 2001, 8 females and 12-14 males were caught in Gilan, about half of which could be used as broodstock at the Shahid Beheshti hatchery (Raymakers, 2002).

Fingerlings have been raised in fibreglass ponds in brackish and fresh waters in Iran (Iranian Fisheries Research Organization Newsletter, 35:3, 2003; H. Pouralifashtomi in the 5th International Symposium on Sturgeon, Iranian Fisheries Research Organization, 9-13 May 2005, Ramsar; Pouralifashtomi, 2006). Growth was better in brackish water when fed diets containing 45% protein and 12.8% fat. Studies of cultured male fil mahi show that they attain maturity at 8-10 years, earlier than fish in natural habitats, indicative of their potential for caviar production under culture conditions (Iranian Fisheries Research Organization Newsletter, 39:3, 2004).

Cultivation of this species in earthen ponds in the central Iranian desert at Bafqh near Yazd has been carried out. After three months at 24ºC and a salinity of 12.5‰ the fish reached 250 g with a survival rate of 60%, after six months at 16ºC and 11.0‰ the fish weighed 1100 g with a survival rate of 96%. Growth was better during the cold season (Iranian Fisheries Research Organization Newsletter, 34:3; 36:4, 2003).

Further work

See under A. gueldenstaedtii.

Sources

See under family above. Babushkin (1964) gives a general review of the biology and catch of this species.

Iranian material: None.

Comparative material: CMNFI 1986-0147, 1, ca. 305 mm total length, Romania, Black Sea at Sulina (45°09'N, 29°41'E).

Pseudoscaphirhynchus
Nikolskii, 1900

Pseudoscaphirhynchus kaufmanni
(Kessler, 1877)

This species is reported from the Karakum Canal and Kopetdag Reservoir in Turkmenistan by Shakirova and Sukhanova (1994) and Sal'nikov (1995). It may eventually be found in the Tedzhen River and Caspian Sea basins of Iran. No Iranian record.

Anguillidae
 

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Freshwater eels are found world-wide in temperate to tropical waters except for the south Atlantic Ocean and the whole eastern Pacific Ocean. There are 15 species with 1 found in Iran.

The family is characterised by the elongate body; numerous vertebrae; small elliptical scales which are difficult to see casually; a small and elliptical gill opening just in front of the pectoral fin base; very long dorsal and anal fins confluent with a reduced caudal fin; a terminal mouth with the lower jaw projecting a little; small teeth in several rows on the jaws and palate; the dorsal fin origin well behind the pectoral fin level but in front of the anus level; no pelvic fins; and by a suite of osteological characters. The term eel-like is based on the body shape of freshwater eels and includes the muscular slipperiness associated with this fish and its mucus-producing skin.

The life cycle of Atlantic eels was unknown until Johannes Schmidt published his 1922 study based on years of collecting. Where the adults went on their seaward migration and where the elvers ascending rivers came from were a mystery. These eels are catadromous, living in fresh water but migrating to the sea to spawn and die. In the North Atlantic Ocean spawning occurs in the Sargasso Sea. The young eels or leptocephali (= thin head larvae) are distinctive, being transparent and leaf-like. A newspaper can be read through the body of a leptocephalus. In this form they drift to the shores of America and Europe, transform into elvers with the more familiar eel-shape and move into rivers and lakes to feed and grow. Some scientists believe that the European eel is not a distinct species but merely American eels (Anguilla rostrata (Le Sueur, 1817)) which develop in cooler areas of the Sargasso Sea and are carried by different ocean currents to the shores of Europe. Differences between the American and European eels overlap and include such characters as vertebral number which is known to vary with development temperature. Recent studies using mitochondrial DNA (mt DNA) showed no genetic divergence among samples of American eels along 4000 km of North American coastline reflecting a single spawning population. However European eels had a distinct mtDNA genotype and the conclusion to be drawn is that American and European eels have separate spawning sites such that larval dispersal ends up on different continents. The mt DNA differences are marked but do not prove species distinction as this level of distinction is known to occur among fishes which are a single species (though some authorities would argue that these "single" species are themselves complexes of two or more species). However Icelandic eels seem to be hybrids between the two putative species. All other evidence (vertebral and other counts, body proportions, biology, electrophoresis) suggests that the American and European eels are the same species but have different spawning sites.

The biology of eels is based almost entirely on the freshwater phase of their life. Adults in fresh water develop large eyes, the gut degenerates and coloration changes in preparation for the migration to the Sargasso Sea. Adults were only caught in the deep ocean, at nearly 2000 m near the Bahamas, in 1977. The Sargasso spawning ground is deduced from collections of larvae across the Atlantic Ocean - the smallest and youngest larvae are found around the Sargasso Sea. The spawning grounds are at about 400 m, at a 17°C temperature and in saltier water than usual sea conditions according to some authors but since spawning adults have never been caught this remains dubious.

The theory advanced by D. W. Tucker in 1959 maintained that European eels lack the energy resources in their migratory, spawning phase to reach the Sargasso Sea 7000 km from Europe. They are presumed to be following an instinct to head out to sea, dating from an earlier geological age when the Atlantic Ocean was narrower before the separation caused by Continental Drift. All European eels die at sea and Europe is restocked by larvae drifting there spawned from American parents. The American populations are closer to the Sargasso and can make the journey easily. Differences between American and European eels are merely the consequence of different environmental regimes in different parts of the Sargasso. This theory has not found general acceptance but, if true, means that all European eels can be harvested for food without depleting stocks. Eels are valued as food, particularly in Europe and Japan,

Hochleithner (2010) gives a review of eel biology and aquaculture.

Genus Anguilla
Schrank, 1798

Characters of the family also serve for the genus.

Anguilla anguilla
(Linnaeus, 1758)

Common names

marmahi-ye ma'muli (= common snake fish), مارماهي مهاجر (= marmahi mohajer, meaning migrating snakefish), marmahi-ye haghighi.
[rechnoi ugor' or river eel in Russian; European eel].

Systematics

No major synonyms. Muraena anguilla was originally described from Europe.

Key characters

The eel shape is characteristic along with the long and spineless dorsal and anal fins and the absence of pelvic fins. The Caspian lamprey, Caspiomyzon wagneri, has a similar shape but lacks pelvic fins, has seven gill openings in a row behind the eye, and has a round suctorial mouth.

Morphology

The scales are small, elliptical in shape and embedded in the skin. The lateral line is distinct. Some fish in any population may have a broad or a narrow head. Fish approaching sexual maturity develop very large eyes, the olfactory organs atrophy, the lateral line becomes more conspicuous, a tougher and thicker skin develops, and the colour changes as detailed below.

Dorsal fin rays 243-275, anal fin rays 175-249 and pectoral fin rays 15-21. Vertebrae 110-119, usually 114-116. The chromosomes are 2n=38 (Klinkhardt et al., 1995).

The leptocephalus and elver stages are not found in Iranian waters and are not described here (see below under Reproduction).

Sexual dimorphism

At the silver eel stage males are 29-40 cm and females 38-130 cm long.

Colour

Colour is variable but the back is usually grey-brown, olive-brown, brownish-green, yellowish or black and the belly is whitish to yellowish. The dorsal fin is dark, other fins are yellowish. The iris is yellow. This yellow or green eel stage changes to the silver or bronze eel at maturity. The mature fish is darker on the back, has silvery or bronze to coppery flanks and belly and a black pectoral fin.

Size

Attains 2.0 m, but rarely, and 12.7 kg, possibly 14.0 kg. Iranian specimens up to 1.0 m long have been caught near Bandar Anzali (Firouz, 2005).

Distribution

Occasionally caught in Iranian waters (P. Walczak, pers. comm., 1978; Holcík and Razavi, 1992). Holčík and Oláh (1992) report single specimens from the Anzali Mordab (= Talab) and its exit streams and near Bandar-e Anzali. Also reported generally from the southeast Caspian Sea, southwest Caspian Sea and south-central Caspian Sea (Kiabi et al., 1999). Reported from the Safid River and Anzali Talab by Abbasi et al. (1999). Berra (2001) does not show the Iranian distribution because the fish are introduced.

Elsewhere it is common in Europe including the Mediterranean Sea, and east to the Black Sea although few young eels migrate naturally as far as this.

    

Zoogeography

An exotic species in Iranian waters, arriving there through the influence of mankind. This species established itself in the Caspian Sea after the opening of the Volga-Baltic waterway and the introduction of larvae from France and England and was recorded from fresh waters in Azerbaijan by Abdurakhmanov and Kuliyev (1968).

Habitat

Eels are caught by fishermen between Bandar-e Anzali and the mouth of the Safid River in beach seines, in the Anzali Mordab and are probably present in rivers along the Caspian coast. The catch appears to be increasing (Holčík and Razavi, 1992). About 10-40 specimens are caught annually weighing up to 3 kg (Holčík and Oláh, 1992). In Europe freshwater populations show considerable migratory movements in summer and this helps explain their widening distribution in the Caspian Sea basin. However, Abbasi (2005) states that the population has decreased.

Eels will live in almost any kind of water over a wide range of temperatures; warmer waters being preferred as long as oxygen is not low. Elvers flourish in sandy areas where grain size is 0.25 mm or in gravel areas where size is 2 mm or larger, the former for burrowing, the latter for insinuating between. Adults also prefer a substrate that can be burrowed into during the day, emerging at night. The burrows are usually at a 45° angle and the eel sticks its head out at this angle too. Eels show some migratory habits within fresh water, moving between summer and winter areas, over a distance of a few metres to tens of kilometres.

In Europe, the silver eels begin to migrate to the sea in late summer and autumn on their journey to the Sargasso Sea where they arrive the following spring. They travel at about 2 km/hour, particularly at night when the moon is at or a few days after the last quarter and light levels are low. Iranian fish cannot migrate, being constrained by distance and lack of ready access to the open ocean.

Age and growth

Eels generally begin to mature only at sizes above 30 cm long. Females grow much larger than males and usually begin to mature at 54 cm or longer. Maturity is actually attained after leaving European waters en route to the Sargasso Sea. Eel larvae do not all metamorphose at the same age (this can vary from 1 to 6 years) with subsequent effects on age at the same length. In addition, growth varies widely with the habitat and available food supply. Fish of the same length often have very different weights. Life span is up to a reputed 88 years based on a captive specimen.

Food

Eels are principally nocturnal but feed both at night and during the day. Food includes almost any edible item and includes fish spawn, small fishes, and larger dead fish which have a mouthful of flesh torn off by a rapid rotation along the long axis of the eel body. Food includes insect larvae and algae but fishes, worms, crustaceans and molluscs are the most important items in order. In the southern Caspian they have been reported to eat gobies (Gobiidae) and Rutilus sp. in November, suggesting that feeding continues late in the year in contrast to other waters where they dig into sand or silt and hibernate (Abdurakhmanov and Kuliyev, 1968). Eels will lie buried in mud or gravel with just the head projecting, seizing by a sudden strike any food item passing by. Eels will feed on commercially important species such as salmonids and crayfishes. They are reliably reported to even leave the water and enter fields, presumably to feed on slugs and worms.

Reproduction

This has not been observed in the wild but under artificial conditions eels are promiscuous and fertilisation is external. The eel is believed to spawn in the Sargasso Sea at 100-200 m depths off the coast of America after a long migration from Europe. Spawning takes place at the beginning of March. Mature females contain 3 million eggs per 1 kg body weight. The ovary is a rosy-pink because of numerous blood vessels. The pelagic eggs are 1.2 mm in diameter. The eggs develop into a distinctive leptocephalus larva which has a leaf-like shape quite unlike the adult eel. During its leptocephalus phase, the eel drifts on ocean currents and actively swims from the American side of the Atlantic, arriving in Europe in its third summer. It is now fully grown and 7.5 cm long. The larva gradually transforms into the elver at depths of 1000 m off the coast of Europe. The elver is a eel-shaped and transparent and reduces in length and weight during the autumn when it does not feed. The elvers begin to migrate into rivers and lakes in Europe in winter. They are regarded as young eels once they begin to feed and are fully pigmented.

Parasites and predators

There is a heavy toll on elvers which are taken on the migration into rivers and lakes by a wide variety of fish and birds. Adults are eaten by large fishes including larger eels and by birds such as herons and cormorants. A large variety of parasites have been reported from eels.

Economic importance

Not used in Iran for food, probably because its minute scales make it appear scaleless, and in any case the annual catch is only about 40-60 specimens (Holčík and Razavi, 1992). It is of considerable economic importance in Europe where annual catches have reached 22,000 tonnes. The 1981 catch in Turkey, for example, was 374 tonnes. This species is also farmed quite extensively. The flesh has a high fat content and the eel is often smoked for sale.

The blood of this fish is poisonous but the poison is destroyed by cooking. Fresh eel blood should never be ingested; a dog injected with eel serum died within one minute. Symptoms include diarrhoea, bloody stools, nausea, vomiting, frothing at the mouth, skin eruptions, cyanosis, apathy, irregular pulse, weakness, numbness, paralysis, respiratory distress, and death. Severe inflammations will result if the blood touches the eye or tongue.

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in aquaculture and aquaria, as food, for sport and in textbooks.

Conservation

The peculiar migratory behaviour of this species prevents spawning in Iranian waters and all stocks must be replenished through migration from European waters or by artificial introductions. As an exotic species, no conservation measures are required. Critically endangered in Turkey and throughout its range (Fricke et al., 2007).

Further work

The spread of this species in Iran should be tracked and its numbers assessed. Eels can attack commercial fishes trapped in nets and this should be monitored as a potential loss to fisheries. This species has potential commercial importance for fish farming although numbers are too low in Iran to provide adequate supplies to stock a fish farm.

Sources

There is little information on this species in Iran because of its scarcity and general biology is taken from Bertin (1956), Tesch (1973), Sinha and Jones (1975), Deelder (1984), and Hoestlandt (1991).

Clupeidae

Herrings, shads, sardines, pilchards and menhadens are moderate-sized fishes, usually less than 25 cm long, found in warmer marine waters with some species anadromous or permanent freshwater residents. There are about 57 genera and 188 species world-wide (Nelson, 2006), with 8 species in the Caspian Sea and 1 commonly found in Persian Gulf drainages. Some other species are known to enter rivers in southern Iran (see Marine List in Checklists in Introduction). The diversity of this family in the Caspian Sea is seen in the number of subspecies which have been described, rather than in genera. At the species level there are several endemics.

Curiously, the species and subspecies in the Caspian are generally of larger size than their relatives in the Black Sea basin. These observations are attributed to the variable environment in the Caspian Sea over time, with repeated changes in salinity and temperature which the fish could not avoid. Black, Mediterranean and Atlantic species lived under more stable conditions and could, in any case, retreat from lowered temperatures for example. In addition, the Caspian Sea clupeids lacked the competitors which entered the Black Sea from the Mediterranean and Atlantic and some (Clupeonella spp., Alosa caspia) could occupy the pelagic, planktivore niche taken up by other species in the Black Sea. There are no other pelagic fish but these herrings in the stable salinity areas of the Caspian Sea.

These fishes usually have modified scales on the belly forming abdominal scutes with a saw-like edge. Most species have 2, long, rod-like postcleithra. The lateral line is usually absent or on only a few scales. Silvery cycloid scales are easily detached and are found only on the body. The mouth is usually terminal with jaws about equal in length. Teeth are small or absent but gill rakers are long and numerous for sieving plankton. Fins lack spines and there are no barbels. There is no adipose fin. The pectoral and pelvic fins have a large axillary scale. The caudal fin is deeply forked. The eye is partly covered by an adipose eyelid. The flesh is particularly oily and is highly nutritional.

Members of this family often form immense schools in surface waters of the ocean and the Caspian Sea where they feed on plankton. Schooling is an anti-predator device making it difficult for a predator to pick out an individual from a tight mass of fish. There is also a "sentry effect" where awareness is increased by the presence of many fish. The school is maintained by a balance between visual attraction and lateral line stimulus repulsion. Herring can feed on the smaller plankton, less than 300-400 µm, at night by filter-feeding but during the day can also use particulate feeding. In the latter, they select larger plankton using the area temporalis, a specialised ventro-posterior region of the retina which improves vision as herring approach food items from slightly below.

Herring are easily caught and are extremely valuable to commercial fisheries. They are the most important fishes economically, both as food for man and also for many other commercial fish species. Wars have been fought over fisheries for herrings. In one year, members of the herring family made up 37.3% of all fish caught in the world. Some are used for fish meal, as fertiliser and as an oil source. The 1994-1995 catch of clupeids in the Iranian Caspian was 98.3 tonnes by beach seine and 671.5 t by gill nets, a decrease of 200 t in total over the previous year's catch (Iranian Fisheries Research and Training Organization Newsletter, 10:4-5, 1995)(but see later under Clupeonella where catch is much higher). The Caspian Sea shads account for about 35% of total inland production in Iran which was 117,300 t in 1995 (Bartley and Rana, 1998). These fish are used in a high value form as frozen whole consumer packs, as fish meal for poultry and in aquaculture, and in canning (Food and Agriculture Organization, Fisheries Department, 1996).

The catch of "sprats" (Clupeidae) in Azerbaijani waters is near extinction through poor fishery management according to Golub (1992).

Major sources for the biology and systematics of Caspian clupeids remains Svetovidov (1952), now inevitably dated but not yet updated, Whitehead (1985) and Hoestlandt (1991). There has been no recent, careful systematic and taxonomic study of these species in the Caspian Sea basin and extensive new material was not available for examination here.

Genus Alosa
Linck, 1790

The Caspian species of Alosa were formerly placed in the genus Caspialosa Berg, 1915. Svetovidov (1952) synonymised the genus Caspialosa Berg, 1915 with Alosa. There are 5 species in Iranian waters and the Caspian Sea as a whole but numerous subspecies have been described. Alosa species are also found in the Black Sea, Mediterranean Sea and Atlantic Ocean.

Often distinguished by gill raker counts which in any case overlap, the various subspecies are difficult to identify. Morphometric characters are of little help and Zamakhaev (1944) points out that some named taxa are merely different age groups. This problem is commented on further in the Species Accounts.

Caspialosa suworowi (Berg, 1913) (also spelt suvorovi in the literature) has been used for hybrids of various Caspian herrings and is not a valid species (Berg, 1948-1949). The holotype is in the Zoological Institute, St. Petersburg under ZISP 15927 (Svetovidov, 1952; Eschmeyer et al., 1996).

Alosa species are distinguished from sympatric Clupeonella species by larger size (up to 75 cm total length compared to 20 cm), a large mouth, a black spot on the flank behind the operculum and sometimes a row of such spots, an elongate scale or ala at the upper and lower base of the caudal fin, a notch at the mid-line of the upper jaw and by the last two anal fin rays not being elongated.

Caspian Sea species have a laterally compressed belly with 29-36 spiny scutes running from the throat to the anal fin; the dorsal fin origin is closer to the snout tip than the caudal fin base; the dorsal fin lies in a groove formed by enlarged scales; scales are easily detached; the pelvic fin origin lies below or slightly posterior to the dorsal fin origin; teeth are usually present on the jaws, roof of the mouth (on the palatine bone and always on the vomer bone), and on the tongue; the opercular bone is distinctly striated; eggs are demersal, semi-pelagic, and lack an oil globule; gill rakers highly variable in shape and number (18-180); dorsal fin branched rays 11-16, anal fin branched rays 10-21, scales in lateral series 49-60, and vertebrae 43-55.

Afraei Bandpyi et al. (2004) examined Alosa species from Mazandaran and Golestan provinces and found the following distinguishing characters:-

Species Gill rakers Ratio of eye diameter to total length (%)
A. braschnikowii 20-40, mean 30.9 2.9-5.8, mean 4.7
A. caspia 110-125, mean 118.3 5.7-7.5, mean 6.2
A. pontica (= kessleri) 60-73, mean 66.8 4.3-6.5, mean 5.5
A. saposchnikowii 20-48, mean 32.8 6.0-9.3, mean 7.3

The general Farsi name for these fishes is shag mahi or zalun (both in Gilaki).

These herrings migrate from the north Caspian Sea to overwinter in the central and southern parts, returning north in the spring.

Alosa braschnikowii
(Borodin, 1904)

Common names

shagmahi, shagmahi-ye Khazari.

[dolkii siyanayn, Agraxan siyanayi, Sara siyanayi, irikoz siyanak, hasangulu siyanayi, agbas siyanak, all in Azerbaijan; Caspian marine shad, Kurinskaya sel'd or Kura herring, poloschataya sel'd or striped herring, Agrakhanskaya sel'd or Agrakhan herring, bol'sheglazaya sel'd or bigeye herring, dolginskaya sel'd or dolginka herring, belogolovaya sel'd or whitehead herring, Astrabadskaya sel'd or Astrabad herring, sel'd-gonets or driver, zheltospinka or yellow-back, Gasankulinskaya sel'd or Gasan-Kuli herring, kiselevichevskaya sel'd or Kiselevitch herring, Krasnovodskaya sel'd or Krasnovodsk herring, vostochnaya sel'd or eastern herring, obzhorka or glutton, Sarinskaya sel'd or Sara herring, maiskaya sel'd or May herring, Brashnikovskaya sel'd or Brashhnikov's shad, all in Russian].

Systematics

Originally described as Clupea caspio-pontica var. Braschnikowii. Reshetnikov et al. (1997) revert to the original double "i" ending to the specific name. A lectotype from Fort Shevchenko (Aleksandrovsk) is in the Zoological Institute, St. Petersburg (ZISP 13051) and paralectotypes were designated by Svetovidov (1952)(ZISP 13051). Clupea caspio-pontica is an unneeded new name according to Eschmeyer et al. (1996).

Alosa braschnikowii is regarded as a subspecies of Alosa caspia by some authors. Clupeonella leucocephala Berg, 1913 from Sumgait and Gyurgenchai, Azerbaijan is a synonym (as Caspialosa brashnikovi leucocephalia (sic) it is listed as a synonym of C. b. grimmi in Mikhailovskaya (1941)), as is Caspialosa caspia nigra Kisselevitsh, 1923 from the Caspian Sea opposite Dzambai (the material also included specimens of Alosa saposchnikowii) (Whitehead, 1985; Eschmeyer et al., 1996).

Alosa braschnikowii has 9 subspecies in the Caspian Sea (including Alosa curensis (q.v.) the Kura or striped herring), namely agrachanica (Mikhailovskaya, 1941) (author also spelt Mikhaylovsky or Mikhailovsky; dated 1940 in Eschmeyer et al. (1996) here and below but 1941 on the paper itself and in Svetovidov (1952) and Berg (1948-1949); species also spelt agrakhanika in Berg (1948-1949); Caspialosa brashnikovi morpha elata is a synonym according to Mikhailovskaya (1941)), the Agrakhan herring; autumnalis (Berg, 1915), the bigeye herring; braschnikowii (Borodin, 1904) (also spelt brashnikovi in Svetovidov (1952) and Berg (1948-1949)), the dolginka herring; grimmi (Borodin, 1904), the whitehead or Astrabad herring, driver or yellow-back; kisselevitshi (Bulgakov, 1926) (spelt kisselevitschi on the plate in Bulgakov (1926), kisselevitschi in Mikhailovskaya (1941), kisselevitshi in Svetovidov (1952) and Whitehead (1985) and kisselewitschi in Berg (1948-1949)), the Gasan-Kuli or Kiselevitch herring; nirchi (Morosov, 1928)(author also spelt Morosow in Mikhailovskaya (1941) and Morozov in Eschmeyer et al. (1996)) (with Caspialosa brashnikovi kenderlensis Budamshin, 1938 from Kendyrli Bay as a synonym in Svetovidov (1952) and Berg (1948-1949)), the Krasnovodsk herring; orientalis (Mikhailovskaya, 1941), the eastern herring or glutton; and sarensis (Mikhailovskaya, 1941), the Sara or May herring. Caspialosa brashnikovi derzhavini Tarasevich, 1946 described from the Caspian Sea near the Apsheron Peninsula, Azerbaijan may be another subspecies. Caspialosa kiselevitschi morpha elata Morozov, 1928 from the Caspian Sea, Krasnovodsk Bay, Turkmenistan is an infrasubspecific taxon and its availability and validity as a taxon have not been examined (Eschmeyer et al., 1996).

This high number of subspecies is an indication of the populational variation of this shad and not all subspecies may be valid. A modern revision is required to assess this problem. In light of this uncertainty and the lack of adequate sample sizes to determine which of the subspecies occurs in Iranian waters or which taxa are valid, reference is made here mostly to the species level. Additionally, it should be noted that hybrids between the various subspecies, and between this species and other species, do occur to complicate matters even further.

The neotype of Caspialosa brashnikovi agrachanica was designated by Svetovidov (1952) as the specimen described by Berg as Caspialosa brashnikovi m. elata taken in front of the Sulak River mouth, Agrakhan Bay and housed in the Zoological Institute. St. Petersburg under ZISP 7334.

The neotype of Caspialosa braschnikowi autumnalis was designated by Svetovidov (1952) as a specimen 26.9 cm long from the eastern shore of the Caspian Sea at Gasan-Kuli (just north of the Iranian border in Turkmenistan) caught on 8 April 1948 and housed under ZISP 31749.

The lectotype of Clupea curensis from the Kura River estuary is under ZISP 13984 with many paralectotypes as established by Svetovidov (1952) (Eschmeyer et al., 1996).

The neotype of Caspialosa kisselevitshi is also from Gasan-Kuli caught on 30 June-1 July 1926 and was housed in the Faculty of Zoology, Central-Asian State University (Sredne-Aziatskogo Gosudarstvennogo Universiteta), Tashkent.

The neotype of Clupea caspio-pontica var. grimmi was designated by Svetovidov (1952) as a specimen 34.0 cm long found at Ashur-ade (= Ashuradeh) near Astrabad Bay (= Gorgan Bay or Khalij-e Gorgan) on 23 April 1903 is under ZISP 13045.

The neotype of Caspialosa nirchi as designated by Svetovidov (1952) is from the southern part of the Caspian Sea opposite North Cheleken Spit and is under ZISP 31780.

The neotype of Caspialosa brashnikovi orientalis as designated by Svetovidov (1952) is from the southern part of the Caspian Sea opposite Kara-Ashly and is under ZISP 32187.

The neotype of Caspialosa brashnikovi sarensis from Sara Island is under ZISP 32184 as established by Svetovidov (1952).

Key characters

Characterised by a relatively elongate and rounded body likened to a "herring" shape, not as deep as in some related species which are likened to a "shad" shape. Total gill rakers 18-49 and short (about equal to gill filaments in length, sometimes shorter). Teeth are well developed in both jaws.

Morphology

Dorsal fin with 3-5 unbranched and 12-15, mostly 14, branched rays, anal fin with 2-4, usually 3, unbranched rays and 10-20, mostly 18, branched rays. Scales in lateral series 51-54. Teeth are well-developed on the jaws, tongue and roof of the mouth.

The accompanying table summarises characters of the subspecies and is taken from Svetovidov (1952) and Mikhailovskaya (1941) but identification to subspecies should be done with the keys from these works. Some of the characters used in the keys are not in the table as they do show individual variation and are difficult to summarise. An example is the nature of the gill raker (thin, thick, blunt, pointed, bent, straight, curved, branched, broken off, forked, swollen at the tip, etc.); another is the degree of protrusion of the lower jaw.

The subspecies grimmi is quite specialised in association with its benthic mode of life, feeding mostly on gobies (Gobiidae). It has a unique character in the well-developed callus on the tip of the lower jaw which adults acquire from rubbing the jaw on the sea bed while feeding, gill rakers are low in number as fine food is not taken, and the tips are broken off, broadened, and split owing to abrasion, and the rakers on the lower arch are reduced in number so the first raker is far from the tongue base. The subspecies nirchi is similar. In contrast, the subspecies kisselevitshi has a high gill raker count, rakers are pointed and not split at the tips, and the first raker is close to the tongue base. This species lives in surface waters feeding on Clupeonella, Atherina, shrimps, gammarids, and gobies (Gobiidae).

Character / Subspecies

Gill rakers (mostly)

Pectoral fins as % body length

Vertebrae

Head length as % of body length

agrachanica

20-46 (28-33)

13.1-15.6

47-54

22.6-25.2
autumnalis

21-37 (28-30)

16.4-19.9

45-53

26.0-29.2

braschnikowii

24-47 (30-33)

14.3-16.7

48-55

23.5-26.6
curensis

26-54

17.3-18.8

47-52

25.7-26.5

grimmi

18-28 (20-22)

12.9-15.2

45-52

22.9-26.4

kisselevitshi

29-49 (36-40)

13.9-16.8

43-53

24.2-26.9

nirchi

20-31 (23-26)

10.9-14.7

48-52

23.4-26.3

orientalis

20-35 (27-32)

13.5-18.0

45-53

25.0-27.8

sarensis

20-33 (24-27)

14.1-16.2

45-53

23.8-26.6

Sexual dimorphism

None reported.

Colour

The back and top of the head are dark with a green or blue tint and may be grey-green. Some subspecies are paler in colour with a grey or grass-green back and pale flanks, nirchi has a whitish blue-green head, light grey back with a slight greenish tint, and lower jaw and pectoral fins light, while grimmi is also quite pale with a grey-blue back and top of the head and whitish anterior head and pectoral fins. There is a dark spot behind the operculum but no series of spots along the flank in most subspecies, except in rare cases when there may be up to 7, occasionally 12-13. The subspecies grimmi regularly has a row of diffused, grey spots almost merging into a stripe, and nirchi occasionally. Pectoral fins are dark on some subspecies (braschnikowii, sarensis, kisselevitshi), pale or whitish on the others, although there is confusion in the literature over this, perhaps indicative of individual variation (cf. sarensis in Mikhailovskaya (1941)). The back and upper part of the head may become a deep black at spawning. The flanks and belly are silvery.

Size

Attains 50 cm standard length but average lengths are about 27-34 cm.

Distribution

All the Caspian subspecies are found widely distributed in the sea but chiefly in the south in winter, moving north to spawn in spring. The subspecies sarensis is reported from the Lenkoran coast and from southwest of Gasan-Kuli (in Turkmenistan just north of the Iranian border), the subspecies orientalis from Gorgan or Astrabad Bay, autumnalis from coastal waters at Gasan-Kuli, kisselevitshi from Astara and Gasan-Kuli, and grimmi from Astara and Gorgan Bay.

Zoogeography

This species is endemic to the Caspian Sea.

Habitat

In winter this species moves into deeper water towards the Iranian coast. In March it approaches coastal waters (Vetchanin, 1984) including brackish waters but does not enter fresh water. It never enters rivers in the south of the Caspian Sea (Jolodar and Abdoli, 2004). Salinities up to 47.6‰ are survived by this species. Spawning and feeding grounds are in the north Caspian for some populations but others live permanently in the south Caspian Sea and are of smaller size. The subspecies kisselevitshi, for example, lives off Gasan-Kuli in winter at depths below 25 m, not migrating or feeding. In March they move north to feed and then return south to spawn but lives almost entirely as a pelagic species in the southern Caspian Sea. Knipovich (1921) reports this species from depths of 80-98 m in Iranian waters. The density of this species increased from east to west in a 1999-2001 study in Iranian waters (Afraei, 2006). Abdoli and Naderi (2009) list it as from the southwest, southeast and south-central Caspian Sea in Iranian waters.

Age and growth

Maturity is attained at age 2-5 and life span is up to 10 years, although this varies with the subspecies. Most south Caspian forms apparently mature at age 2 according to Svetovidov (1952). Growth rates also vary between subspecies, orientalis being one of the slowest growing herrings in the Caspian Sea and reaching 10 years of age. The catch near Astara of sarensis, for example, is mainly 4-5 year olds but this too varies with the subspecies and also with the year-class strength. Vetchanin (1992) reported on grimmi catches from the southeastern Caspian where the average length was 27.8-28.6 cm and the average weight 294-313 g. There is a tendency for length and weight to fall in catches as the summer progressed, from April to July. Length and weight are less in southern, compared to northern, waters. Afraei (2000) found this species to be the largest Alosa in Iranian waters on average at 395 mm and 760.3 g. Males predominate at 55.8% in Mazandaran and 69.4% in Golestan catches. Six age classes were present (1+ to 6+) with the 2+ class being the most common at 28.9% and 6+ the rarest at 8.9%.

Food

Diet in the southeastern Caspian Sea in winter comprises 85% Clupeonella engrauliformis with some gobies (Neogobius) and shrimps (Vetchanin, 1984). From March to November the diet is dominated by Clupeonella caspia, Atherina boyeri (= caspia) and shrimps. Juvenile Liza saliens, Syngnathus caspius, molluscs, crabs and higher aquatic plants are also recorded along with foreign objects such as rice husks, pieces of wood, foil, polyethylene, etc. This species is a cannibal. The more southerly populations examined favour Atherina boyeri (= caspia) and Neogobius species and some of these populations favour benthic invertebrates. The subspecies grimmi is the most benthic one and takes primarily gobies with some molluscs as well as Clupeonella. Feeding intensity rises sharply after spawning. While some herrings, like Alosa pontica (= kessleri), feed poorly on their migration, this species feeds intensively on its spring migration.

The feeding regime altered after the invasion of the ctenophore, Mnemiopsis leidyi. A shift was observed from 85% Clupeonella engrauliformis to 65% Atherina boyeri (= caspia). Other fishes were also eaten including Clupeonella grimmi, C. caspia, Cyprinus carpio, Liza saliens, as well as Palaemon spp. (Iranian Fisheries Research Organization Newsletter, 49:2, 2006).

Reproduction

Vetchanin (1984) reports spawning of this species in the southeastern Caspian Sea north of Iranian waters to begin in early May, continuing to July as it is intermittent. The subspecies sarensis spawns along the Lenkoran coast from mid-April to the end of June. The subspecies orientalis spawns in Gorgan Bay from the end of March to the beginning of April, spawning schools forming at 17-18°C or higher. The subspecies autumnalis spawns at the same time off Gasan-Kuli near the Iranian border with Turkmenistan. The subspecies grimmi spawns in May-June in Gorgan Bay. The subspecies kisselevitshi has the latest spawning date, June to July and even in August off Gasan-Kuli when temperatures exceed 25°C. Spawning takes place in shallow water (1.8-5.8 m) in the sea over sand or silt bottoms at 15-18°C (some subspecies and populations at 20-22°C, others beginning as low as 11°C), and a salinity of 8-13‰. Fecundity is up to 178,400 eggs, average 66,000 per fish. There is no feeding while spawning. Early maturers, like the south Caspian populations, can reproduce up to 7 times in their life.

Parasites and predators

The Caspian seal, Pusa caspica, is a predator on this species (Krylov, 1984).

Economic importance

The catch for all species of "Caspialosa" in Iran varied between 5337 kg and 419,518 kg for the years 1956/1957 to 1961/1962 (Vladykov, 1964). In the Anzali region the catch for the years 1933/1934 to 1961/1962 varied from 1553 kg to 539,710 kg (Vladykov, 1964).

The catch has been as high as 126,900 centners or 12,690 t in the sea as a whole for the type subspecies alone (1 centner = 100 kg (Svetovidov, 1952)), taken chiefly in spring. Other subspecies were not fished for as extensively although kisselevitshi was the most numerous of the south Caspian forms of Alosa braschnikowii, forming 70% of the drift net catch.

Conservation

Reputedly depleted in Iranian waters. Kiabi et al. (1999) consider this species to be data deficient in the south Caspian Sea basin according to IUCN criteria. Criteria include medium numbers, medium range (25-75% of water bodies), absent in other water bodies in Iran, and present outside the Caspian Sea basin. Extinct in Turkey (Fricke et al., 2007).

Further work

The biology of this species in Iranian waters and the stocks or taxa found there need to be elucidated.

Sources

Iranian material: CMNFI 1970-0581, 5, 226.0-245.0 mm standard length, Gilan, Caspian Sea near Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1979-0431, 1, 297.2 mm standard length, Mazandaran, bazaar at Now Shahr (no other locality data); CMNFI 1980-0126, 1, 245.8 mm standard length Gilan, Caspian Sea near Bandar-e Anzali (37º28'N, 49º27'E); CMNFI 1980-0150, 1, 222.4 mm standard length, Gilan, Safid River estuary (37º24'N, 49º58'E).

Comparative material: BM(NH) 1938.8.2:1, 1, 245.9 mm standard length, Kazakhstan, Caspian Sea, Kaidak Bay (no other locality data); BM(NH) 1938.8.2:2, 1, ca. 337.5 mm standard length, Kazakhstan, Caspian Sea, Kaidak Bay (no other locality data); BM(NH) 1939.2.21:17-18, 2, 285.0-305.2 mm standard length, Caspian Sea (no other locality data); BM(NH) 1939.2.21:19-20, 2, 222.9-273.4 mm standard length, Caspian Sea (no other locality data).

Alosa caspia
(Eichwald, 1838)

Common names

shagmahi-ye shekambozorg (= big belly herring fish), shagmahi-ye chekameh dar, shagmahi-ye darya-ye khazar (= Caspian Sea herring fish), شاه ماهي (= shah mahi, meaning king fish), zalun (in Gilaki), puzanok.

[xazar sisgarini, sara sisgarini in Azerbaijan; Kaspiiskii puzanok or Caspian shad, severokaspiiskii puzanok or North Caspian shad, srednekaspiiskii puzanok or Central Caspian shad, il'mennyi puzanok or il'men shad, Enzeliiskii puzanok or Enzeli (= Anzali) shad, Sarinskii puzanok or Sara shad, Bakinskii puzanok or Baku shad, Astrabadskii puzanok or Astrabad shad, all in Russian].

Systematics

Clupea caspia was originally described in Latin from "Hab. in Caspio mari, meridiem versus" (Caspian Sea, towards the south).

A. caspia has 3 subspecies in the Caspian Sea basin, namely caspia (Eichwald, 1838) (= North Caspian, Central Caspian, Caspian or il'men shad); knipowitschi (Iljin, 1927) with natio knipowitschi (Iljin, 1927) (= Enzeli or Anzali shad) and natio saraica (Svetovidov, 1943) (= Sara or Baku shad); and persica (Iljin, 1927) (= Astrabad shad). The differences between natio knipowitschi and natio saraica are small (e.g. gill rakers 122-166 versus 140-150, both averaging 145; vertebrae 43-49 versus 45-51, both with mostly 47 or 48; growth differences are known, the former grows faster in the first 2 years of life but the latter reaches a greater size) and they probably have no taxonomic significance being simply separate breeding populations. The differences between Alosa caspia caspia natio caspia (the North or Central Caspian shad) and natio aestuarina Berg, 1932 (the il'men shad) were found to be based on geography and growth rate and these names have no taxonomic standing (Svetovidov, 1952). These natio are infrasubspecific ranks and have no validity as names.

Alosa rossica Kessler, 1870 described from the Volga River is a nomen nudum and is this species. Other taxa now considered as synonyms of Alosa caspia are Caspialosa caspia salina Svetovidov, 1936 from Mertvyi Kultuk and Kaidak bays in the northeast Caspian Sea and Caspialosa caspia kaidakensis Kazancheev, 1936 (spelt kajdakensis in Svetovidov (1952)) from Kaidak, the latter being in any case a synonym of the former subspecies. Clupeonella caspia m. elongata Berg, 1913 is also a synonym. Alosa caspica Yakovlev, 1871 is presumably a misspelling.

Knipovich (1921) records a species, Caspialosa enzeliensis Iljin, from the southern Caspian Sea which he places as a subspecies of caspia. I have been unable to locate the original description of this taxon, which presumably is found in the Anzali Mordab of Iran. It is probably an unused manuscript name for what Iljin later described as knipowitschi. As of 15 July 2007, this scientific name is a Googleblat for this page.

The lectotype of Caspialosa knipowitschi is a specimen 21.2 cm long from Anzali in Iran caught on 15 April 1915 and housed in the Zoological Institute, St. Petersburg (ZISP 31892). The lectotype of Caspialosa caspia var. persica is a specimen 147.5 mm long from the Caspian Sea Bay of Asterabad (= Gorgan Bay or Khalij-e Gorgan) north of Ashur-ade (= Ashuradeh) at 36°53'N, 53°55'E caught on 25 April 1904 on the Caspian Expedition of 1904 and housed in the Zoological Institute, St. Petersburg (ZISP 16413). The lectotype of Caspialosa caspia knipowitschi n. saraica is from near Sara Island and is under ZISP 32183. The lectotype of Caspialosa caspia salina is from Mertvyi Kultuk Bay, 10 km west of Cape Kizil-kair and is under ZISP 25813. These taxa were designated by Svetovidov (1952) as none were before or material was not preserved.

Key characters

Characterised by a relatively deep and compressed body likened to a "shad" shape, not as elongate and rounded as in some related species which are likened to a "herring" shape. Total gill rakers 50-180, variously reported as thin or thick, long (obviously longer than the gill filaments), and forming a convex outline on the lower arch. Teeth are poorly developed in both jaws.

Morphology

Dorsal fin with 3-4 unbranched and 12-15 branched rays, anal fin with 3-4, usually 3, unbranched and 15-20 branched rays. Scales in lateral series 49-54.

The characters distinguishing subspecies all overlap widely and are given below after Svetovidov (1952) and Hoestlandt (1991):-

Characters / Subspecies Head length as % of body length Pectoral length as % of body length

Vertebrae

Gill rakers

caspia

25.5-28.1

15.5-18.1

45-52 (49-51)

68-150 (100-140)

knipowitschi

18.3-24.1*

16.0-19.1

43-51 (47-48)

120-180 (130-160)

persica

25.6-27.1

16.5-17.7

45-51 (47-49)

50-120 (60-90)

* The numbers cited in Svetovidov (1952: 256 in the English version) and Hoestlandt (1991: 128) in the keys to subspecies do not agree with the numbers on p. 148 and p. 265 respectively in the species descriptions. The text numbers are used here.

Sexual dimorphism

Females are longer and weigh more than males of the same age.

Colour

The back is blue-green to dark and the flanks silvery. There is a black spot on the flank behind the upper operculum margin and sometimes up to 7 spots extending along the upper flank to a level of the rear of the dorsal fin.

Size

Reaches 28 cm standard length for caspia, to 29.6 cm for knipowitschi, and to 33.8 cm for persica.

Distribution

Found in the Caspian and Black seas. The subspecies caspia is found mostly in the western half of the Caspian Sea basin but is the most widely distributed subspecies, found throughout almost the whole sea. The subspecies knipowitschi is found in the south near Anzali, Astara and the Baku Archipelago, near the northern shore of the Apsheron Peninsula in autumn with a few reaching the Gorgan Mordab in fall and winter; natio saraica is found north of Astara and spawns near Sara Island, natio knipowitschi spawns in the Anzali Mordab. The subspecies persica is found in the southeast, near Gorgan or Astrabad Bay. Holčík and Oláh (1992) report persica from the western basin of the Anzali Mordab (= Talab) and this species is reported from the Safid River and Anzali Talab as subspecies persica and from the Anzali Talab as knipowitschi (Abbasi et al., 1999). Abdoli and Naderi (2009) list it as from the southwest, southeast and south-central Caspian Sea, the Anzali Talab and Gorgan Bay in Iranian waters for both knipowitschi and persica.

Zoogeography

This species is part of a marine fauna encompassing the Black and Caspian seas, surviving in the reduced salinity of the latter.

Habitat

The type subspecies prefers open waters. Caspian shad winter at depths of 30-40 m or more and prefer temperatures not less than 8-11°C. They rise to surface waters in spring, moving north along the western shore of the Caspian Sea in waters of about 9-11°C according to Kushnarenko (1986) while Heckman in Hoestlandt (1991) states that this shad begins to migrate at the end of March at 5-6°C water temperature with a peak at 9-14°C in mid to late-April, ending in early May. Males migrate in large numbers at the beginning and end of the migration, females in the middle (Pushbarnek, 1987) while Heckman in Hoestlandt (1991) states that two waves of migration occur, one usually in late April at 7.6-10.2°C comprised of over 80% males and the second in the first half of May at 10.8-14.0°C comprised of over 70% females. The young, which hatch in the spring, leave the summer feeding grounds before the adults and migrate south before October-November. Adults follow as temperatures fall. Some populations do not migrate north and spend their whole life in the southern Caspian Sea. This subspecies will enter fresh waters to spawn in addition to spawning in the open Caspian Sea. The subspecies knipowitschia prefers water warmer than that of all other Caspian Sea clupeids except for Alosa caspia persica. Its sea movements are not well known but spawning fish favour waters with freshwater input and some fish enter rivers so it is classified as semi-anadromous. This subspecies was common in the Anzali Mordab but is now replaced by persica (Holčík and Oláh, 1992). It is also reported westwards to Astara and eastwards to Gorgan Bay. The winter habitat of persica is unknown. It is semi-anadromous and remains in the southern Caspian Sea near the shore. From spring to fall this subspecies moves northward along the eastern Caspian shore towards Krasnovodsk Bay and westwards to the Anzali Mordab.

Age and growth

Pushbarnek (1987) found shad of the type subspecies up to 7 years of age on the western coast of the middle Caspian Sea. In the spawning population, the predominant sizes and weighs for males were 16-21 cm and 60-130 g and for females 18-23 cm and 70-140 g. Males and females usually mature at 2-3 years although most spawn for the first time at 3 years. Females grow faster than males. Shad may spawn up to four times as the period of sexual maturation may continue for 2-5 years. The age composition of the spawning population is dependent on year-class strength. First spawners constitute 75.9% of 3-year-olds, 41.7% of 4-year-olds and 23.5% of 5-year-olds. The Caspian shad is a slow-growing species compared to A. braschnikowii and A. saposchnikowii, its mean length being 21.2 cm compared to 32.2 cm and 25.6 cm for the two other species respectively (Shubina, 1981). Dmitriev (1947) briefly examined the Anzali, Iran population and found 6 age groups but life span is noted by Heckman in Hoestlandt (1991) to be up to 9 years. Maturity is attained as early as 2 years although most fish appear to mature later as most spawners are 4-5 years old. The subspecies persica is the slowest growing of the shad species in the Caspian Sea, sexually mature fish being 13-21 cm long. Some fish become mature at 2 years of age. Life span is up to 8 years. The populations of both knipowitschi and persica are small compared to caspia. Abbasi and Sabkara (2004c) studied 180 fish from the southeast Caspian Sea coast of Iran and found fork length to be 103-232 mm, mean 158.8 mm, weight 16-130 g, mean 52.2 g and age 2-5 years, mean 2.64 years. Afraei (2000) found this species to be the smallest Alosa in Iranian waters on average at 110 mm and 109 g.

Food

The most intensive feeding period occurs after reproduction, beginning in June and the highest condition factor is found at the end of this summer feeding period. Little food is eaten in winter. Temperature (affecting metabolic rate) and zooplankton biomass (decreases engender competition with Clupeonella engrauliformis and other planktivores) are important factors governing catches of this species (Shubina, 1981). Food is chiefly copepods, more than 70%, with mysids at 20%, but some phytoplankton and small fishes are taken. Food in rivers after spawning is mostly cladocerans and other crustaceans. The above refers to the type subspecies; food of the other two subspecies is assumed to be similar. The southeast Caspian Sea fish studied by Abbasi and Sabkara (2004c) fed on phytoplankton (Rhizosolenia and Sprirogyra) at 4.5%, zooplankton (Foraminifera, Copepoda, Cirripedia, Bivalvia larvae) at 95.0%, and bony fish larvae and eggs at 0.5%. The presence of the ctenophore, Mnemiopsis leidyi, a food competitor reduced the index of fullness and fish growth was reduced. Abdollapour Bereya et al. (2007) studied diet in fish from beach seines and gill nets in Gilan. 98.0% of the stomach contents were zooplankton (ostracods, rhizopods, cladocerans, rotatarians, copepods, cirripedes, mysids, bivalve larvae and bony fish larvae and eggs), 1.8% was phytoplankton (notably Rhizosolenia and Spirogyra), and 0.2% was benhthic items (foraminiferans, sponges, cumaceans, amphipods, insect lavae and palaemonids). Acartia spp. (copepods) at 83.1% and Balanus (cypris larvae of the cirripede) at 12.9% were the most abundant. The zooplankton have declined drastically from predation by Mnemiopsis leidyi, the invasive ctenophore, and the fish have shown a great reduction in the index of fullness and in growth recently.

Reproduction

Most spawning of the type subspecies occurs in the north Caspian Sea near the outflow of the major rivers, particularly the Volga, and the fish overwinter in the south Caspian, migrating between the two areas (Shubina, 1981). This subspecies spawns successively, 3 times within a week. Some fish enter fresh water to spawn. Spawning takes place at the favoured water temperature of 13.8-24.1°C, with mass spawning at 18-22°C, beginning as early as late April or as late as mid-May and continuing to mid- or late June. Most eggs are released in the upper 3 m of the water column. Fecundity reaches 41,000 eggs. The eggs are 1.11-1.38 mm when ripe but unfertilised and 1.92-2.91 mm in diameter when fertilised and are semi-pelagic to demersal. The subspecies knipowitschi spawns in the Anzali Mordab (and probably the "Chemkhala" River to the east of the Safid River) in May and June after a spring migration from the sea, leaving in the fall. Spawning of the subspecies persica takes place in Gorgan Bay and Holčík and Oláh (1992) suspect from catches of mature and spent fish that it also occurs in the Anzali Mordab.

Parasites and predators

The Caspian seal, Pusa caspica, is a predator on this species (Krylov, 1984) and it forms a substantial part of the diet of Silurus glanis in the Anzali Mordab (Holčík and Oláh, 1992). Naem et al. (2002) found the monogenean trematode Mazocraes alosae on the gills of this species in the western branch of the Safid River. Barzegar et al. (2008) record the digenean eye parasite Diplostomum spathaceum from this fish.

Economic importance

The type subspecies was the most important subspecies in the herring family in the Caspian Sea. It is caught off the coasts of Dagestan and Azerbaijan for research purposes and comprises 85% of the clupeid catch (Pushbarnek, 1987), 80-90% of the Caspian commercial catch (Kushnarenko, 1986). During the 1970s it was only 2% of the total Caspian fishery production. These herrings dominated the commercial catch in the Caspian Sea until the 1960s when commercial fishing was banned except on the western coast of the central Caspian. Many young of other commercial species were being killed in the herring fishery, entangled in the gill nets used. Soviet catches have weighed as much as 75,000 t. This fish is fattier than other Caspian Clupeidae, except for Alosa kessleri, up to 18.1% of the body weight. The fat content decreases on the spring migration. The catch of the subspecies knipowitschi is of minor economic importance and had been little exploited when Svetovidov (1952) summarised biology, as the age of captured fish indicated. About 420 tons (sic, possibly tonnes) were caught in the Anzali Mordab in 1933 and 1934, but this may be an error in the report by Vladykov (1964) according to Holčík and Oláh (1992) although Berg (1948-1949) reports 4200 centners for the same period. The fishing season in the mordab began in mid-April and ended in mid-June when spent fish appeared. There appears to be no fishery data on the subspecies persica in the sea. Holčík and Oláh (1992) report catches of persica, which replaced knipowitschi, in the Anzali Mordab from the end of April to the beginning of June but in 1990 this comprised only 5 kg. It is regarded as of inferior quality in Iran. The Caspian shad is the dominant fish catch in the Iranian Caspian, comprising 51,000 t in 1994 rising from nothing a decade earlier (Food and Agriculture Organization, Fisheries Department, 1996). Robins et al. (1991) list this species as important to North Americans. Importance is based on its use as food.

Conservation

The stocks of this species in the Anzali Mordab are likely to increase as the lagoon becomes more saline (Holčík and Oláh, 1992). Kiabi et al. (1999) consider this species to be of least concern in the south Caspian Sea basin according to IUCN criteria. Criteria include abundant in numbers, widespread range (75% of water bodies), absent in other water bodies in Iran, and absent outside the Caspian Sea basin. Extinct in Turkey (Fricke et al., 2007).

Further work

The biology of this species in Iranian waters and the stocks or taxa found there need to be elucidated.

Sources

See under family heading.

Iranian material: CMNFI 1970-0524, 11, 58.7-88.9 mm standard length, Gilan, Caspian Sea at Bandar-e Anzali (37º28'N, 49º27'E); CMNFI 1970-0532, 1, 113.0 mm standard length, Gilan, Caspian Sea near Bandar-e Anzali (37º28'N, 49º27'E); CMNFI 1970-0543A, 1, 85.9 mm standard length, Gilan, Caspian Sea at Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1970-0586, 1, 77.5 mm standard length, Mazandaran, Gorgan Mordab at Ashuradeh-ye Kuchak (36º50'N, 53º56'E); CMNFI 1970-0587, 2, 107.4-108.6 mm standard length, Mazandaran, Babol Sar (36º43'N, 52º39'E); CMNFI 1971-0343, 1, 95.5 mm standard length, Gilan, Langarud at Chamkhaleh (37º13'N, 50º16'E); CMNFI 1979-0430, 1, 118.0 mm standard length, Mazandaran, river east of Now Shahr (36º39'N, 51º31'E); CMNFI 1979-0431, 7, 120.8-155.1 mm standard length, Mazandaran, Now Shahr bazaar (no other locality data); CMNFI 1979-0686, 2, 119.7-126.9 mm standard length, Gilan, Safid River (37º24'N, 49º58'E); CMNFI 1980-0146, 2, 106.9-171.8 mm standard length, Mazandaran, Gorgan Mordab at Ashuradeh-ye Kuchak (36º50'N, 53º56'E).

Comparative material: BM(NH) 1938.8.2:3, 1, 203.8 mm standard length, Caspian Sea (no other locality data); BM(NH) 1939.2.21:22-23, 2, 175.6-179.2 mm standard length, Caspian Sea (no other locality data); BM(NH) 1954.6.24:5-7, 3, 164.1-189.1 mm standard length, Caspian Sea (no other locality data).

Alosa curensis
(Suvorov, 1907)

This species is poorly known and not recorded from Iran but from Kyzylagach Bay of Azerbaijan. It may, in any case, be a subspecies or synonym of Alosa braschnikowii (see Svetovidov (1952) and the Alosa braschnikowii account herein).

Alosa kessleri
(Grimm, 1887)

Common names

shagmahi-ye poshtsiah, shagmahi darya-ye siah, shagmahi-ye moohajer or shagmahi-e-mohajer, zalun (in Gilaki), puzanok.

[Volga siyanayi, garabel siyanak in Azerbaijanian; arkasy gara takgas in Turkmenian; blackback, Caspian anadromous shad; chernospinka or black-spined herring, chernonosik or blacknose, beshenka, zalom, poluzalom, zheleznitsa, veselka, Volzhskaya mnogotychinkovaya sel'd or Volga many-rakered herring, Volzhskaya sel'd or Volga herring, Astrakhanskaya sel'd or Astrakhan herring, all in Russian; Pontic shad, Black Sea herring].

Systematics

Clupea kessleri was originally described from the Volga River delta, Astrakhan. Clupea pontica was originally described in Latin from "Hab. in Ponte Euxino prope Odessam" (= Black Sea near Odessa).

Alosa kessleri was formerly considered as a subspecies of A. pontica. Alosa pontica then had two subspecies in the Caspian Sea, namely kessleri (Grimm, 1887) (chernospinka or black-spined herring, chernonosik or blacknose, beshenka, zalom, poluzalom, zheleznitsa, veselka, blackback), and volgensis (Berg, 1913) (Volzhskaya mnogotychinkovaya sel'd or Volga many-rakered herring, Volzhskaya sel'd or Volga herring, Astrakhanskaya sel'd or Astrakhan herring, zheleznitsa, beshenka, veselka).  Kottelat and Freyhof (2007), Abdoli and Naderi (2009) and Naseka and Bogutskaya (2009) consider Alosa kessleri and A. volgensis to be valid species.

A lectotype of kessleri, 40.1 cm long, was designated from the Volga Delta by L. S. Berg under ZISP 15925 (in the Zoological Institute, St. Petersburg). A lectotype of volgensis, 34.8 cm long, is under ZISP 15926 and is from the Volga River at Chernyi Yar (Svetovidov, 1952). A paralectotype of kessleri is under ZIN 15922.

Caspialosa volgensis bergi Tanasiichuk, 1938 described from the Volga Delta is a synonym of Alosa kessleri (Heckman in Hoestlandt, 1991). Eschmeyer et al. (1996) give author and date for Alosa volgensis bergi as Tanassiychuk, 1940, the variation probably being due to transliteration of a Russian name and to year of actual publication rather than year on the journal.

Caspialosa kessleri infraspecies volgensis imitans Berg, 1948 from the Caspian Sea (see Berg (1948-1949) for further details) is not available because of its infrasubspecific rank (Eschmeyer et al., 1996).

Clupea caspio-pontica Borodin, 1904 is an unneeded new name for these fishes from the Black and Caspian seas (Eschmeyer et al., 1996).

Key characters

Characterised by a relatively elongate and rounded body likened to a "herring" shape, not as deep and compressed as in some related species which are likened to a "shad" shape. Total gill rakers 57-158 in the Caspian Sea, 57-95 in kessleri, 87-158 in volgensis. Rakers are usually longer than the gill filaments in volgensis, shorter in adult kessleri. Teeth are well developed in both jaws in kessleri and can be felt with a finger, poorly developed in volgensis such that they sometimes cannot be felt.

Morphology

Dorsal fin with 3-5 unbranched and 12-16 branched rays, anal fin with 2-4, usually 3, unbranched and 15-21 branched rays. Vertebrae 47-50 in kessleri (also a report of 50-54, both in Svetovidov (1952)), 48-54 in volgensis. Pyloric caeca 21-62. Scales in lateral series 53-56. Gill rakers in adults are thick and often broken off at the tip or near the base in kessleri, unbroken in volgensis. The tips of the gill rakers may be swollen and they are arranged in a straight line. Young fish have long and thin gill rakers with strong lateral spines. Spines are lost with age. Chromosome number is 2n=48 (Klinkhardt et al., 1995).

Sexual dimorphism

None reported.

Colour

The overall coloration is dark with a black back which has a violet tinge in spring in kessleri, light olive green in volgensis. There is dark, sometimes vague, spot on the flank behind the operculum and sometimes a series of spots in kessleri, but these are absent in volgensis. The pectoral fin is black on top. Spawning kessleri become grey or grey-green on the back and flanks with bronze spots on the operculum and flanks. A greenish-yellow circle forms around the eye after spawning.

Size

Reaches 52 cm total length and 2.0 kg for kessleri, 40 cm for volgensis.

Distribution

Found in the Black and Caspian seas and throughout the latter, entering northern rivers to spawn. Abdoli and Naderi (2009) list it as from the southwest, southeast and south-central Caspian Sea in Iranian waters.

Zoogeography

This species is part of a marine fauna encompassing the Black and Caspian seas, surviving in the reduced salinity of the latter.

Habitat

Both subspecies are found in the open sea but kessleri ascends rivers much higher than volgensis which spawns in the delta region. Both subspecies overwinter in the southern Caspian Sea off the Iranian coast and then migrate north to enter the Volga and other northern rivers to spawn. The subspecies volgensis is absent from the southern Caspian in summer. The subspecies kessleri shows a greater affinity than volgensis for cold water.

The subspecies kessleri begins to migrate northward in March and April mostly along the western shore of the Caspian Sea, beginning to arrive in northern waters when temperatures are still below 5°C, most arriving when temperatures are 6-8°C compared to 10-13°C for volgensis. A mass migration into the lower Volga takes place in late April or early May for both subspecies when water temperature reaches 9°C and the peak run begins at 12-15°C, ending at 22°C. The run of volgensis is about 10 days later than that of kessleri and spawning takes place earlier as they do not travel as far upriver. Speed is up to 70 km/day for kessleri and depends on temperature. This fish used to run 2000 km up the Volga River. Sexually immature fish remain in the south and do not migrate. Knipovich (1921) reports kessleri as deep as 235-300 m in Iranian waters. Temperatures up to 25ºC are tolerated.

Age and growth

Males are sexually mature at 3 years and females at 4 years, other reports give 4-5 years for both sexes in kessleri. Many fish die after spawning but some survive to spawn two or three times. Four and five-year- olds dominate on kessleri spawning runs with some older fish also present. Females predominate in older fish making the spawning run. Life span is between 7 and 8 years.

Growth of the volgensis subspecies is slower than in kessleri, which apparently grows faster than any other Caspian clupeid. Life span in volgensis is 7-8 years with females living longer than males. Most spawners are 3-4 years old although in some years 5 year old fish are abundant. Males may mature at 2 years, females later. Most fish spawn again the next year after their first time but some may miss a year. An individual may spawn up to four times during its life.

Yılmaz and Polat (2002) compared scales, vertebrae, otoliths, opercles and subopercles as ageing structures and determined vertebrae to be the most accurate and reliable for a Turkish Black Sea population at Samsun. Six age classes were found.

Food

Cladocerans are the main food item of young kessleri which have a feeding peak at 1800-2200 hours and another at about 0800 hours. Adults in the sea take fishes such as Clupeonella and Atherina with some crustaceans and insect larvae. Clupeonella caspia makes up 92% of the diet of kessleri in the northern Caspian in May, with Sander lucioperca at 6.6% and gammarids at 1.0%. There is said to be little feeding on the spawning run although some fish sampled contained cladocerans, copepods, insects, bryozoans and fish fry.

The food of volgensis is similar to the other subspecies, taking copepods when young and larger items with growth. The main items are copepods, mysids, cumaceans, amphipods and small fishes. This subspecies feeds on the spawning migration.

Reproduction

Spawning in kessleri occurs in rivers from mid-May to mid-August, either the delta or lower reaches when entering in a ripe condition, or as much as 500 km upriver when entering in an unripe condition. Larger fish have spawning grounds further upriver than smaller fish and predominate earlier in the run. The spawning grounds in the Volga River cover a considerable stretch. Spawning usually occurs at 18-20°C between 0300 and 0600 hours or from 1600 hours to sunset. Spawning occurs in the main channel, over shallow sand banks, or in backwaters. Batches of eggs are laid at intervals of several days. Eggs are pelagic as in other Caspian Alosa and develop as they drift downriver near the bottom. At 22.7°C incubation takes about 40 hours. The young fish descend in late summer and early fall. Fecundity in kessleri reaches 344,000 eggs and egg diameter 1.51 mm. Shed eggs are up to 4.1m in diameter. Some fish may return to spawn in total three times.

Spawning of the first batch of eggs in volgensis may occur in the sea with the subsequent 2 batches at 7-10 day intervals in the delta and river. This takes place from mid-May to the beginning of August. Up to 281,000 eggs are shed. Peak spawning occurs at 15-19°C and ends at 25-27°C. Most spawning takes place in the evening between the 1600 and 2200 hours. The young appear in the pre-estuarine area of the Volga River in July and towards October begin to migrate south.

Parasites and predators

The Caspian seal, Pusa caspica, is a predator on this species (Krylov, 1984), larval shad are fed on by other fishes and by various invertebrates, and adults by various fishes and birds.

Economic importance

The subspecies kessleri and volgensis were caught on the spawning run with as much as 5750 t being taken annually pre-World War II. It is the biggest shad in the Caspian Sea. The subspecies kessleri was the most important and valuable herring in the Caspian Sea. Early spring catches were mostly kessleri but as the run of volgensis built up it formed an increasingly significant part of the catch, forming as much as 92% of the total. The catch of volgensis has declined from this period until the 1970s when the fish taken were mostly kessleri. The catch of Alosa pontica (= kessleri) on the North Caspian fishing grounds in 1965-1972 has declined to 2-4% of the 1938-1943 catch. The subspecies volgensis was one of the most important Caspian herrings, 23-29% of the total catch from 1936-1939, as high as 69,100 t in 1939.

The subspecies kessleri is said to be the tastiest Caspian clupeid because of its high fat content, averaging 18.9% of weight along the coast of Azerbaijan, while in volgensis it was 9.6%. Post-spawners of kessleri may have a fat content as low as 0.5%. Catches are processed as canned, salted and pickled fish. Beach seines are used to catch this fish. Akhondzadeh Basteh et al. (2006) found the bacterial pathogen Vibrio haemolyticus in fresh and smoked Alosa kessleri.

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use as food.

Conservation

Stocks in Iranian waters are said to be depleted. The subspecies volgensis was in Category I on the "Red List" of the Russian Republic (Pavlov et al., 1985). Kiabi et al. (1999) consider this species (as A. kessleri) to be data deficient in the south Caspian Sea basin according to IUCN criteria. Criteria include commercial fishing, numbers unknown, range unknown absent in other water bodies in Iran, absent outside the Caspian Sea basin.

Further work

Stocks in Iranian waters need to be assessed and protected if required.

Sources

See under family account.

Iranian material: None available.

Comparative material: BM(NH) 1879.11.14:22-23, 2, 255.9-259.1 mm standard length, Caspian Sea (no other locality data); BM(NH) 1939.2.21:21, 1, 388.6 mm standard length, Caspian Sea (no other locality data).

Alosa saposchnikowii
(Grimm, 1887)

Common names

shagmahi-ye cheshmdorosht, shagmahi, kilka (incorrectly), herring.

[irikoz sisgarin in Azerbaijan; bol'sheglazyi puzanok or bigeye shad, Sapozhnikovskii puzanok or Saposhnikovi shad, all in Russian].

Systematics

The lectotype of Clupea saposchnikowii from the Volga Delta is in the Zoological Institute, St. Petersburg under ZISP 15921 (Berg, 1948-1949; Eschmeyer et al., 1996). The name is often spelt saposchnikovi, in error, or with a single terminal "i"; Reshetnikov et al. (1997) revert to the original spelling of the specific name.

Caspialosa caspia nigra Kisselevitsh, 1923, in part, from the Caspian Sea opposite Dzambai is a synonym with a lectotype in the Zoological Institute, St. Petersburg (ZISP 15938) (Kisselevitsh is also transliterated Kiselevich and Kisselevitz). The material also included specimens of Alosa braschnikowii (Whitehead, 1985; Eschmeyer et al., 1996).

Key characters

Characterised by a relatively deep and compressed body likened to a "shad" shape, not as elongate and rounded as in some related species which are likened to a "herring" shape. The upper and lower head profiles are straight. The upper edge of the lower jaw is straight. Total gill rakers 24-41, short (obviously shorter than the gill filaments), and thick. Teeth are well developed in both jaws.

Morphology

Dorsal fin with 3-5, usually 4, unbranched rays and 12-15, mostly 13, branched rays, anal fin with 2-4, usually 3, unbranched rays and 15-21, mostly 18, branched rays. Lateral series scales 52-55. Vertebrae 47-53. Pyloric caeca 36-59.

Sexual dimorphism

None reported.

Colour

Fish from the southern Caspian Sea are more intensively coloured than those from the north. The back is violet with green sheen, the flank has 4 dark stripes which merge with the dark on the back. There is a spot posterior to the operculum, which may be absent, and there is no series of spots.

Size

Reaches 36 cm total length and 650 g.

Distribution

Found mainly in the north Caspian Sea and the coast of Dagestan but entering Iranian waters. Abdoli and Naderi (2009) list it as from the southwest, southeast and south-central Caspian Sea in Iranian waters.

Zoogeography

This species is endemic to the Caspian Sea.

Habitat

This species spends its whole life in the Caspian Sea and never enters rivers. It favours colder water and is one of the first clupeid species to migrate north in spring, principally along the western coast. Large fish migrate first. Fish first approach the shore of Azerbaijan in mid-March with a mass approach from late March to mid-April. It is less frequently encountered in the southern part of the Caspian Sea, overwintering in the central Caspian and only moving south if winters are cold. A Caspian Sea Biodiversity Database (from www.caspianenvironment.org) has it at 400-600 m in the southern Caspian in cold winters but later states it keeps at 15-32 m. Winter temperatures at which this species is found are 6-7°C. Depths are 25-32 m in winter, more shallow in summer but below 9 m. Knipovich (1921) reports this species in a depth range of 52-77 m in Iranian waters. It tolerates a range of 3-25°C and spawns at salinities of 0.7-11.0‰, although preferring 4.0-7.5‰. The Caspian Sea Biodiversity Database (from www.caspianenvironment.org) estimates a population of 1.1125 billion fish.

Age and growth

Life span is about 9 years and female lengths and weights exceed those of males throughout life. On average, males weigh less than half the weight of females since females carry a heavy egg load. Growth is most intensive in the first two years of life and slows thereafter (Chang, 1972). Males mature at age 2 and females at age 3.

Food

A rapacious fish which takes young herrings and kilka, Atherina and even Benthophilus (Lönnberg, 1900b) as well as large crustaceans such as mysids and gammarids. It is a cannibal. This shad overwinters and feeds in the south Caspian Sea (Chang, 1972).

Reproduction

The spring spawning migration (end of April to end of May) enters the north Caspian Sea and fish are mostly 15-25 cm in body length. Males mature at a younger age than females as evidenced by fish 3-4 years old predominating among females and fish 2-4 years old among males in the north Caspian catch. Spawning takes place in May (peaking in the first 10 days) and most fish are returning for the second time. Spawning temperatures are lower than in Alosa caspia, being only 13-14°C although the peak is at 19-20°C. Spawning occurs in il'mens, the sea where there is a freshwater discharge such as near the Volga River mouth, and in the northeastern sea. Females may spawn up to 6 times and males up to 5 times (Chang, 1972). Spawning takes place in shallow water at 1-6 m depths. Fecundity is up to 318,852 eggs. The young migrate southwards.

Parasites and predators

The Caspian seal, Pusa caspica, is a predator on this species (Krylov, 1984).

Economic importance

An important commercial species in the central and northern Caspian, taken on their way to, and on, the spawning grounds. The fishery in Azerbaijan during 1937 caught fish on average 17 cm long and 62 g in weight, most fish being 2-3 years old. The Caspian catch in the period 1936-1939 reached a peak of 8,800 t annually. Fish are caught with beach seines, stationary nets and drift nets.

Conservation

Stocks in Iranian waters are reputed to be depleted. Kiabi et al. (1999) consider this species to be data deficient in the south Caspian Sea basin according to IUCN criteria. Criteria include numbers unknown, range unknown, absent in other water bodies in Iran, absent outside the Caspian Sea basin.

Further work

The biology of this species in Iranian waters and the stocks or taxa found there need to be elucidated.

Sources

See under family heading.

Iranian material: CMNFI 1970-0531, 15, 49.9-108.7 mm standard length, Mazandaran, Larim River (36º46'N, 52º58'E); CMNFI 1970-0532, 1, 137.4 mm standard length, Gilan, Caspian Sea near Bandar-e Anzali (37º28'N, 49º27'E); CMNFI 1970-0543A, 2, 78.8-80.2 mm standard length, Gilan, Caspian Sea at Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1970-0581, 1, 102.1 mm standard length, Gilan, Caspian Sea near Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1979-0788, 3, 96.0-114.9 mm standard length, Mazandaran at Khadje Nafas (37º00'N, 54º07'E); CMNFI 1980-0136, 3, 107.3-127.6 mm standard length, Mazandaran, Fereydun Kenar River (36º41'N, 52º29'E); CMNFI 1980-0157, 2, 96.6-101.1 mm standard length, Mazandaran, Gorgan River estuary (36º59'N, 53º59'30"E); CMNFI 1980-0908, 1, 77.9 mm standard length, Gilan, Safid River estuary (ca. 37º28'N, ca. 49º54'E).

Comparative material: BM(NH) 1954.6.24:8-10, 3, 150.5-177.0 mm standard length, Caspian Sea (no other locality data).

Alosa sphaerocephala
(Berg, 1913)

Common names

shagmahi-ye Agrakhan.

[kruglogolovyi puzanok or roundheaded shad, Agrakhanskii puzanok or Agrakhan shad, both in Russian].

Systematics

The holotype of Clupeonella sphaerocephala from Agrakhan Bay, at Tyulenii Island, Turali in the northern part of the Caspian Sea is in the Zoological Institute, St. Petersburg under ZISP 15928 with more than 30 paratypes (Eschmeyer et al., 1996).

Key characters

Characterised by a relatively deep and compressed body likened to a "shad" shape, not as elongate and rounded as in some related species which are likened to a "herring" shape. The upper and lower head profiles are obviously rounded. The upper edge of the lower jaw is crescent-shaped. Total gill rakers 25-45, long (equal to or longer than the gill filaments), and thin. Teeth are well developed in both jaws.

Morphology

Dorsal fin with 3-4, usually 4, unbranched rays and 13-15 branched rays, anal fin with 3-4, usually 3, unbranched rays and 17-20 branched rays. Vertebrae 47-51.

Sexual dimorphism

None reported.

Colour

The back is dark with an olive tint, the tip of the snout is occasionally black and the pectoral fins are dark. There is a black spot behind the operculum and occasionally a row of such spots.

Size

Reaches 25 cm.

Distribution

Found in the Caspian Sea including Iranian waters.

Zoogeography

This species is endemic to the Caspian Sea.

Habitat

This species does not enter fresh waters. It is most common along the eastern shore of the northern part of the sea in spring where spawning occurs and along the northern shore of the northern part of the sea in summer. Knipovich (1921) reports this species from Iranian waters in a depth range of 52-77 m.

Age and growth

Unknown.

Food

Unknown, although presumably similar to other shads.

Reproduction

Spawning takes place in the northeastern Caspian from mid-May to the end of June peaking at 18-20°C, most frequently in a salinity of 8-11‰ and in depths around 3-8 m. The young move south in late autumn, as late as November, the last clupeids to leave this area. Fecundity is about 20,000 eggs.

Parasites and predators

The Caspian seal, Pusa caspica, is a predator on this species (Krylov, 1984).

Economic importance

This species is caught only in small numbers.

Conservation

The status of this species is unknown.

Further work

This species is poorly known biologically and studies in Iranian waters should be carried out on its life history.

Sources

Iranian material: None available.

Comparative material: BM(NH) 1954.6.24:11-13, 3, 145.6-162.1 mm standard length, Caspian Sea (no other locality data).

Alosa volgensis
(Berg, 1913)

Recorded from Iranian waters by Kottelat and Freyhof (2007) but presence needs confirmation by specimens.

Genus Clupeonella
Kessler, 1877

This genus is found in the Black and Caspian seas basins with 5 species, 3 of which are in the Caspian Sea and in Iranian waters.

Clupeonella species are distinguished from sympatric Alosa species by smaller size, a small and toothless mouth, adipose eyelids are small or rudimentary, no spots on the flank, no elongate scales (ala) at the base of the caudal fin, no vomerine teeth, the lack of a notch at the mid-line of the upper jaw, and by the last two anal fin rays being elongated.

Species in this genus live entirely in the sea, or in fresh water, or migrate between the two. Eggs are pelagic and have a large oil globule.

The general Farsi name for these fishes is كيلكا (= kilka or kelka, i.e. "sprat", although sprat is erroneous according to Berg (1948-1949) who uses tyulka for these fishes).

The three Clupeonella species have been fished in modern Iran since December 1971 but the commercial catch did not exceed 15,000 tonnes. Earlier catches date back only to 1939 with an annual catch of about 100 t in 1943-1949 exported in a marinated form to the Soviet Union (Alam, no date). Curiously, the abundance of kilka has long been known as Kinneir (1813) records "and herrings are in such abundance, that after a storm, the shores of Ghilan and Mazanderaun are nearly covered with them". Caddy (1984) refers to the kilka fisheries of the Iranian Caspian by the scientific name Sprattus sprattus but this is an error.

Caddy (1984) indicated that there were problems in marketing and utilizing these fishes in Iran even though up to 50,000 t could be caught annually (200,000 t elsewhere in the same article). Their best use was probably as food for predators such as Sander lucioperca, Esox lucius and Salmo caspius. A study by Razavi Sayad (1993b) suggested a ceiling of 100,000 t was possible. The Caspian Sea resources of kilka is estimated at 800,000 t from which 340,000 t can be exploited (Abzeeyan, Tehran, 6(8):IV, 1995).

The catch reached 51,000 t in 1994 from none 10 years previously (Food and Agriculture Organization, Fisheries Department, 1996) and was 36,000 t in 1997-1998 (IRNA, 31 March 1998) and 85,000 t in 1998-1999 (Fazli and Roohi, 2002). The catch for the first 6 months of the Iranian year was 17,000 t, taken by 70 trawlers and a 10% increase over the previous year (IRNA, 20 October 1998). Fishermen in Gilan caught 50,000 t annually in the late 1990s (Tehran Times, 5 September 1999). A reported catch of 56,000 t was made in 1999-2000, a 13% increase over the previous year (IRNA, 27 March 2000). A later estimate expects the kilka catch to reach 66,000 t by the year 2000 (Abzeeyan, Tehran, 5(9):IV, 1995). Fazli (2006a) records that kilka fishing ships discharge their catches at three ports, Babolsar and Amirabad in Mazandaran and Anzali in Gilan. The catch decreased from 28,000 t to 19,600 t in Mazandaran and from 57,000 to 42,600 t in Gilan from 1999 to 2000. The catch per unit effort also decreased from 3900 kg to 2500 kg over the two years. Anchovy kilka dominated the catch but the frequency fell from 85-90% to 76% of the catch and common kilka sharply increased. Common kilka had been caught in spring and summer but in 2000 they were taken in all months. The average length of anchovy kilka declined from 96.3 mm in 1997 to 87.3 mm in 2000 and this was also reflected in the age structure, 5+ and 6+ fish being rare. The presence of the ctenophore, Mnemiopsis leidyi, was thought to be damaging stocks (Fazli and Roohi, 2002). Darvishi et al. (200$) studied dietary overlap between the ctenophore and the anchovy kilka (see below). Fazli (no date) studied kilka catches off Mazandaran in 1996-2000. Fishing occurred at night and lasted 7.78-8.22 hours. The maximum catch at 42.8% was taken in October, November and December with a minimum catch in June. The least annual catch per vessel occurred in 1999-2000 (499,401 kg).

A study utilizing an echo-sounder and a pelagic trawler concludes that the maximum biomasses for the three Clupeonella species in the southern Caspian Sea are in winter (422,300 t) and autumn (326,900 t) while in summer and spring values are lower at 275,100 t and 260,800 t respectively. The population consists of 66.1% anchovy kilka (C. engrauliformis), 18.9% bigeye kilka (C. grimmi) and 15% common kilka (C. caspia) (Iranian Fisheries Research and Training Organization Newsletter, 14:6, 1996). Note that later, the Iranian Fisheries Research and Training Organization Newsletter (17:3, 1997) gives kilka biomass in the southern Caspian Sea as winter 22,300 t, autumn 26,900 t, summer 75,100 and spring 60,800 t, presumably lacking the initial digit, and the percentages of kilka species in the biomass are also wrong. This is corrected in a subsequent newsletter (Iranian Fisheries Research and Training Organization Newsletter, Tehran (18:43, 1997) but the corrected percentage biomasses are given as 66% for C. engrauliformis, 19% for C. caspia (as C. delicatula) and 15% for C. grimmi. It is unclear whether grimmi or caspia is the second most important kilka species. Pourgholam et al. (1996) give a stock assessment for kilkas in 1995-1995 using the hydro-acoustic method.

C. engrauliformis dominates the catch in Iran at 91.8%, followed by C. grimmi at 6.84% and by C. caspia at only 1.35%. The 2+ and 3+ year classes account for 69.95% of C. engrauliformis, 81.06% of C. grimmi and 80.88% of C. caspia catches. Catch rates of kilka on the top ranking 17 fishing grounds of 56 studied range from 800 to 1200 kg per unit effort per hour while traditional grounds have rates of 400-800 kg per unit effort per hour. The kilka are caught by attraction to lights and netting or pumping the catch into specially constructed ships. The kilka fishing fleet of Iran expanded in the 1980s and 1990s. There were 30 active vessels in Mazandaran in 1994, each with a capacity up to 30 tons (sic, probably tonnes here and elsewhere for modern catches) (Abzeeyan, Tehran, 4(10):IV, 1994). The Mazandaran Kilka Cooperative Companies Union had 75 boats in 2000 (Tehran Times, 31 December 2000). Gilan planned to construct 12 fish meal factories each with an annual capacity of 1000 t and 10 kilka canneries also with 1000 ton capacities (Abzeeyan, Tehran, 4(4):III, 1993). Catches off Gilan alone from April 1994 to January 1995 increased 59% compared to the same period in 1993-1994, exceeding 20,000 t (Abzeeyan, Tehran, 6(1):II, 1995). The catch off Mazandaran from March 1994 to March 1995 was 15,400 t, an increase of 10% over the previous year. About 1000 t were processed for human consumption and the rest for fishmeal production (Abzeeyan, Tehran, 6(2):V, 1995). The total kilka catch for Iran has increased to 45,000 t annually and efforts were being made to increase it to 110,000 t (Abzeeyan, Tehran 4(5):IV, 1993). The catch in 1995 was 32,000 t with 64.7% from Mazandaran and 35.3% from Gilan, with the maximum catch occurring in April (Abzeeyan, Tehran, 7(6):II, 1996). Catches declined from 95,000 t in 1999 to 15,497 t in 2003 (Sayyad Bourani et al., 2008). Annual Soviet catches reached 37,000 t in 1956 but this declined to 300-1500 t by the end of the 1970s or 0.2-0.8% of all kinds of tyulka or kilka in the Caspian Sea. Turkmenistan harvested 7660 and 8500 t in 1995 and 1996 although previously almost 45,000 t valued at $22.5 million had been taken before equipment deteriorated (http://bisnis.doc.gov/bisnis/isa/9805fish.htm, downloaded 14 March 2000). Stocks remain large even though kilka are heavily fished.

Kilka are smoked, salted, canned in sauce and oil and marinated according to a traditional recipe and seasoned with fruits, herbs and vegetables (Keivany and Nasrollahzadeh, 1990; www.netiran.com/business.html, downloaded 31 October 2003). Moini and Koochekain (2003) give details of fish sauce production from kilkas using traditional, microbial and enzymatic methods, along with taste tests. Vacuum packaging of fresh, smoked and salted kilka has been investigated in Iran (Annual Report, 1995-1996, Iranian Fisheries Research and Training Organization, Tehran, p. 45-46, 1997) and studies on processing kilkas as fish balls have also been carried out (Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 40, 1996). Koochekian Sabour and Moini (2009) describe investigations on using Iranian kilkas to produce a fermented fish sauce for marketing in Southeast Asian countries. One company markets kilka in a clear package which gives the product a bright and colourful appearance. Kilka have even been made into crackers (Iranian Fisheries Research and Training Organization Newsletter, Tehran, 18:6, 1997; Shojaei, 1998). Kilka have also been made into oil as a by-product of the fish meal industry (Iranian Fisheries Research and Training Organization Newsletter, 27:3, 2001). Omega-3 fatty acids have been extracted from kilka oil under laboratory conditions (Salmani Joloudar et al., 2009). M. Shivazad , H. John Mohammady, A. A. Yousef Hakimi and H. Fazaely (http://iman.ut.ac.ir/news/agr.htm, downloaded 12 December 2004) discuss the use of Clupeonella engauliformis as fish meal in animal nutrition and analyse the protein quality and Faeed et al. (2006) studied spoilage in kilka meal from bacteria and fungi. The Iranian Fisheries Research and Training Organization Newsletter (20:4, 1998) and Rezaei et al. (2003) report on methods of transporting kilka in cold water and crushed ice to processing factories which were better than traditional methods. Salmani et al. (2001) recommend chilled sea water for preservation for human consumption. Motamedzadegan et al. (2009) found that partial hydrolysis of fish myofibrillar proteins using papain improves its functionality. Motalebi et al. (2010) investigated the use of whey protein coating on quality and shelf life of kilkas; it can enhance quality and increase frozen shelf life in fish stored for up to 4 months.

The kilka fisheries are threatened by the comb jelly, Mnemiopsis leidyi, which arrived in the Caspian in 1995 in the ballast water of ships and spread through the entire sea by the year 2000, feeding voraciously on zooplankton. It is now known as the "Caspian monster" despite its small size of 5 cm (Muir, 2001). It doubles in size in one day, reaches maturity in two weeks and then produces 8000 young each day (Muir, 2001). The fisheries collapsed by 50% in a few months, catches by one fisherman falling from being 3-6 t a night to half a tonne. Ghadirneja (2003) reports that C. engrauliformis originally dominated the kilka catch at 85-90% but has dropped to 55% through the impact of the comb jelly which has up to 2285 individuals per cubic metre in the southwest Caspian Sea.  Fazli et al. (2009) describe a multi-species approach for stock management, allowing for the decline of C. engrauliformis and increase in C. caspia in Iranian waters through competition with the ctenophore. The fisheries may recover somewhat after the comb jelly population collapses (Tidwell, 2001b) or if a predator, Beroe ovata, is introduced and can survive in the less saline waters of the Caspian Sea (Muir, 2001). Studies indicate it can survive the brackish Caspian Sea water, feed on the comb jelly and not feed on other plankton (Iranian Fisheries Research Organization Newsletter, 36:35, 2003). The following catch records for the total kilka catch in Mazandaran in tonnes is courtesy of F. Darvishi (pers. comm., 2003) and shows the drastic decline caused by the ctenophore, as well as monthly variations in catches:-


Months/Years 1998 (1377) 1999 (1378) 2000 (1379) 2001 (1380) 2002 (1381) Mean
March-April (Farvardin) 2848 2703 4644 1217 876 2458
April-May (Ordibehesht) 1116 607 972 1422 195 862
May-June (Khordad) 370 763 1819 125 158 647
June-July (Tir) 1392 919 194 425 444 675
July-August (Mordad) 2152 2306 433 614 249 1151
August-September (Shahrivar) 3117 2010 581 528 336 1314
September-October (Mehr) 3103 6184 1785 432 575 2416
October-November (Aban) 4120 3468 2305 3051 1196 2828
November-December (Azar) 3835 3410 2655 993 - 2723*
(2179)
December-January (Dey) 2754 1735 620 1082 - 1548*
(1238)
January-February (Bahman) 3968 1262 2146 1586 - 2241*
(1792)
February-March (Esfand) 2815 1667 1192 1903 - 1894*
(1515)
Total 31,590 28,034 19,648 13,378 4029  

* = averaged over 4 and (5) years.

The species composition of kilkas changed after the introduction of the comb jelly comparing the year 2000 and before with the year 2002 - the common kilka changed from about 1-5% to about 30%, the bigeye from about 10-15% to 0/2% and the anchovy kilka from about 85-90% to about 70% (Iranian Fisheries Research Organization Newsletter, 36:2, 2003). The catch per unit effort (catch per vessel per fishing night) fell from 4 t to 1 t.

In 2004, more than 200 fishing boats had been forced to stop operations. The kilka stock has been reduced from 400,000 t to 80,000 t over the past 4 years and the catch fell by 34,000 t (www.iranmania.com, downloaded 4 October 2004). See also the section on the Caspian Sea basin in the Introduction. Mamedov (2006) gives details of the biology and decline of kilkas in Azerbaijan waters.

The Caspian seal was once a major predator on kilkas but the number of seals has declined on the Kazakhstan and Iranian coasts from 300,000 to 5000 in recent years through DDT pollution, viral infections and food shortages (Hashemi, 2001).

An account on the biology and identification of Caspian kilka in Farsi is given by Emadi (1991) and Fazli (1990), Fazli and Besharat (1998) and Poorgholam et al. (1996) give accounts of biology and catches in Iran in Farsi.

Clupeonella caspia
Svetovidov, 1941

Common names

rizeh keraye (= tiny ?), rizeh kuli, kilka-ye ma'muli or kilka-e-maamooli (= common shad).

[xazar kilkasi in Azerbaijanian; adaty kulke balyk in Turkmenian; Kaspiiskaya tyul'ka or kil'ka (i.e. Caspian tyulka or kilka), tyulka, obyknovennaya tyul'ka (i.e. common tyulka), all in Russian; common kilka, common Caspian kilka, sardelle, Caspian sprat, Black Sea sprat].

Systematics

Formerly identified as Clupea cultriventris, originally described from the northern shore of the Black Sea. Clupea delicatula Nordmann, 1840, described from Odessa market on the Black Sea, is a synonym of C. cultriventris and a lectotype is in the Zoological Museum. St. Petersburg under ZISP 2254 with paralectotypes also under ZISP 2254, as designated by Svetovidov (1952). Clupeonella delicatula caspia Svetovidov, 1941 was considered to be a synonym and was described as from the "Caspian Sea, where it is met with almost everywhere, from very saline parts (Kaydak Bay) to quite fresh. Enters the mouths of the Volga and the Ural rivers, ascending sometimes very far upstream". The holotype is from the Volga Delta and is under ZISP 15883 (Svetovidov, 1952). Kottelat and Freyhof (2007) consider this subspecies to be a a distinct species found in the Caspian Sea with cultriventris restricted to the Black Sea. Reshetnikov et al. (1997) consider recognition of this subspecies as questionable.

The Caspian Sea taxon, Clupeonella caspia, has a lectotype, 152 mm long, designated by Svetovidov (1952) in the Zoological Institute, St. Petersburg (ZISP 15883).

Clupea cultriventris is spelled cultiventris in some parts of Eschmeyer et al. (1996), apparently in error. Three syntypes of Clupea cultriventris may be in the Muséum National d'Histoire Naturelle, Paris under MNHN 3681 (Svetovidov, 1952; Eschmeyer et al., 1996).

Clupea cultriventris var. tscharchalensis Borodin, 1896 from Lake Charkhal in the Ural River basin is variously listed as a variety, morpha or a distinct species (see Svetovidov (1952) and Kottelat and Freyhof (2007)).

mtDNA studies of fish from Mazandaran and from Gilan showed statistically significant differences in haplotype frequencies, indicating genetically different populations (Laloei et al., 2006).

Key characters

This species has a moderately deep body (21-27% of standard length), a short and wide head (interorbital width 17.5% or more of head length), a sharply keeled belly, and pointed pectoral fin tips.

The Caspian subspecies is distinguished from the type subspecies of the Black Sea by having shorter pectoral (15.5-19.0% of standard length) and pelvic fins (8.5-12.5% of standard length), although ranges overlap, a shallower body, and a shallower and shorter head. It also grows faster and is more fatty than the Black Sea subspecies.

Morphology

The dorsal fin has 3-4 unbranched rays, usually 3, followed by 11-14 branched rays and the anal fin has 1-3 unbranched rays, usually 3, and 14-19 branched rays. Scales in lateral series 42-55. There are 24-30 belly scutes and 41-62 (rarely to 64), usually 51 or more, gill rakers. Vertebrae 40-44 (rarely to 45) compared to 44-47 in the anchovy kilka and 46-48 in the bigeye kilka, probably as a result of higher water temperatures during development compared to other kilka species (Prikhod'ko, 1979b).

Sexual dimorphism

Sexual dimorphism is only evident during egg development when the belly of females is swollen.

Colour

The back is blue-green or light-green, the flanks silvery and the belly silvery-white or golden-yellow. Fins are hyaline except the dorsal fin which has a central dark but faint stripe and the caudal fin which is darkish at the base. The iris is black.

Size

Reaches 14.5 cm standard length and 19 g.

Distribution

Found in the Black and Caspian seas, tributary rivers and some adjacent lakes. In Iran, it is reported from sea and also the confluence of the Pasikhan and Pir Bazar rivers of the Anzali Mordab, the Anzali Mordab and its outlets by Holčík and Oláh (1992) and from the Safid River and Anzali Talab (= Mordab) by Abbasi et al. (1999). Abdoli and Naderi (2009) list it as from the southwest, southeast and south-central Caspian Sea in Iranian waters.

Zoogeography

This species is part of a marine fauna encompassing the Black and Caspian seas, surviving in the reduced salinity of the latter.

Habitat

The habitat of this species in the Caspian Sea is the coastal zone of the sea at depths less than 100 m, more usually less than 50-70 m, over a wide range of temperatures (2.6-27.6°C for adults, higher for larvae, and possibly lower temperatures since they are found under ice and probably over 28°C according to some reports), and in fresh and hypersaline waters (to 36‰). The young can develop in water at 16‰. Southern populations live in a more saline habitat than northern and central Caspian populations which are mostly in fresh water. This tyulka may not migrate far but does move between summer-winter feeding and spring-early summer spawning grounds. Large schools are found 0.5-2.0 km from shore at depths of 20-25 m on the eastern coast of the Caspian Sea, descending deeper if water temperatures rise and coming up to about 8 m in autumn as temperatures fall. In winter this species is found at about 30-40 m deep where the temperature range is 7-10°C, warmer than surface waters. Larvae and young remain in shallow coastal areas. Knipovich (1921) reports a fish from a depth range of 235-300 m in Iranian waters but populations at these depths are small (Iranian Fisheries Research and Training Organization Newsletter, 14:6, 1996). The Caspian Sea Biodiversity Database (from www.caspianenvironment.org) states that the largest concentrations are found at 3-7‰ with most intensive spawning at 2-4‰.

It is the most widely distributed kilka and with the other kilka species the most abundant fish in the Caspian Sea (Prikhod'ko, 1979b). Large schools can be found by day but these disperse at night. It overwinters in the southern Caspian Sea and some individuals move north to spawn and feed in April. The Caspian Sea Biodiversity Database (from www.caspianenvironment.org) estimates the population to number 224 billion fish, with 96 billion fish in the south Caspian. The south and north Caspian Sea stocks are about equal in number after a decline in copepod biomass in the north. The relative frequency of this species compared to other kilkas increased after the invasion of Mnemiopsis leidyi, by more than 10% (Fazli, 2006b; Fazli et al., 2006).

Age and growth

Osipov and Kiyashko (2008) found that using otoliths gave more reliable estimates than using scales for ageing. The Caspian subspecies grows faster than the Black Sea subspecies. Together with the sturgeons, this species comprises 82.1% of the fish biomass in the Caspian Sea. Condition in this species is better in winter because of the summer-autumn feeding period after spring spawning compared to C. engrauliformis in the Big Kizil-Agach (= Bol'shoy Kyzylagach or Imeni Kirova) Bay of Azerbaijan (Badalov, 1972). Local populations have differing growth regimes depending on the productivity of these areas (Prikhod'ko, 1979b) and there are great variations on a yearly basis too. Southern populations grow faster than northern ones in their first year. Females grow somewhat faster than males (9.0 g versus 7.3 g average weight along the Dagestan coast for example), and life span is about 6 years. This species is mature there at 1 year and average life span is about 3 years.

Females dominate the population in Iran and sexual maturity is attained usually at age 2 and 2-4 year olds dominate catches but life span is up to 8 years (Iranian Fisheries Research and Training Organization Newsletter, 14:6, 1996; Abtahi et al., 2002). Fazli (2006b) found age classes 0+ to 5+ in Iranian waters with 0+ to 3+ making up 95% of the fish in 1997-1999. In 2000, age classes 0+ and 1+ were reduced in numbers and 2+ to 4+ fish comprised 93.8%. Abtahi et al. (2004) examined fish from the conical net and light catch at Babolsar and found average fork lengths were 69.82 mm, 83.56 mm, 88.38 mm and 88.43 mm while weights were 2.2 g, 4.18 g, 4.77 g and 5.06 g for fishes at maturity stages I, II, II and IV. Fazli et al. (2007) studied this species from 1995 to 2004 in Iranian waters, sampled at landing sites at Amirabad and Babolsar in Mazandaran and Anzali in Gilan. Growth parameters were L = 132 mm, K = 0.259/yr. t0 = -1.285/yr. The instantaneous coefficient of natural mortality was 0.506/yr, the instantaneous coefficient of total mortality (Z) was 1.62/yr and the instantaneous coefficient of fishing mortality varied over 10 years from 0.125/yr to 1.487/yr. Annual survival rate (S) was 0.200/yr. Age at first capture was 2.8 years. The von Bertalanffy growth equation was Lt = 132 (1-e-0.259(t +1.285)). Ages ranged from 1 to 7 years with age groups 2, 3 and 4 dominating at different periods. Mean fork lengths were 59.3, 77.5, 87.4, 97.2, 104.5, 111.9 and 116.8 mm. Females dominated in each month except April, averaging 0.47:1, possibly due to differing attraction to lights used in the fishery. Biomass increased from 16,000 mt in 1995 to more than 41,000 mt in 2002, declining to less than 28,000 mt in 2004. The increase was simultaneous with a sharp decline in anchovy kilka, changes in zooplankton composition and abundance, and especially an increase in zooplankton species favoured by this kilka. Currently this kilka is overfished. Karimzadeh et al. (2010) examined fish from the Babolsar region off Mazandaran and calculated growth parameters as L = 143.5 mm, K = 0.30/yr-1 and t0 = -1.02/yr, instantaneous coefficient of natural mortality was 0.671/yr-1 and the current exploitation rate was estimated as 0.55 and this species is now overfished.

Food

Plankton is the main food and copepods predominate but diet also includes Cladocera, Balanus larvae and clam larvae. The dominant food item is the copepod Eurytemora grimmi, particularly in winter when plankton biomass is lowered in the Bol'shoy Kyzylagach Bay of Azerbaijan. The food of the common kilka is more varied than the other kilka species simply because of its habitat in shallow coastal areas (Badalov, 1972; Prikhod'ko, 1979b). Older fish take larger and faster crustaceans and consume less food in proportion to body size as they grow. The most intensive feeding is in summer and autumn, decreasing in winter and during reproduction. Food is taken during the day. Roushan Tabari et al. (2009) examined fish from a fishing vessel of Mazandaran and found highest feeding activity in April with 280±153 prey items per fish weighing 2.9±1.6 mg. Balanus nauplii and cypris larvae comprised 93% and Acartia 7% at this time with increasing spring temperatures and reproduction, but the copepod Acartia biomass dominated from October to February.

Reproduction

Spawning occurs in January-February in the southern Caspian, later in the north, mainly in depths less than 10 m and where salinity is low to average for the Caspian Sea (Badalov, 1972; Prikhod'ko, 1979b). The largest southern Caspian population spawns near the mouths of the Volga and Ural rivers (Kozlovsky in Hoestlandt, 1991). Spawning is most intensive at 11°C, but occurs at 10-20°C. Spawning is intermittent and lasts from mid-April to July. Peak spawning in Iranian waters of Mazandaran Province is April-May with an average fecundity of 28,240 eggs (Abtahi et al., 2002). Fazli (2006b) recorded mass spawning in Iranian waters in April, continuing on until August. Eggs are released in water 0.5-9.0 m deep at a salinity range of 0.02-15‰, perhaps as high as 29.15‰. Fecundity reaches 60,000 eggs and egg diameter 1 mm, 0.48-1.46 mm for fertilised eggs. Relative fertility is 4-13 times greater than in Alosa species. Holčík and Oláh (1992) consider that it may spawn in rivers entering the Anzali Mordab. The studies of Fazli et al. (2006; 2007) showed that reproduction started in March, peaked in May and finished at the end of August. Half the females were mature at 84.3 mm fork length.

Parasites and predators

Samples of this species from Babol Sar and Bandar Anzali contain the digenean parasites Pseudopentagramma symmetrica and Bunocotyle cingulata, the acanthocephalan Corynosoma strumosum, metacercariae of a Bucephalus species, and larvae of a Contracaecum and an Anisakis species (Iranian Fisheries Research and Training Organization Newsletter, 11:4-5, 1996; Annual Report, 1995-1996, Iranian Fisheries Research and Training Organization, Tehran, p. 28, 1997; Shamsi and Dalimi, 1996; Shamsi et al., 1998). Varshoie et al. (2010) record the helminths Pseudopentagramma symmetrica, Bunocotyle cingulata and Mazocreas alosae in this species from Iranian waters.

Clupeonella species are an important food fish for sturgeons (59.4% by weight of Acipenser stellatus diet in the Middle Caspian), Sander, herrings (Clupeidae) and the Caspian seal (Badalov, 1972; Krylov, 1984) as well as Salmo  caspius and Stenodus leucichthys (Kosarev and Yablonskaya, 1994).

Economic importance

It is caught by attraction to underwater electrical lights (Prikhod'ko, 1979b). The other subspecies is also of major importance in the Sea of Azov. The Caspian subspecies is caught in school seines in spring and purse seines in summer. In Iranian waters this species formed only a small proportion (1.35%) of the total kilka catch in a study by Razavi Sayad (1993b) and Fazli (2006b) gives values of 1.34%, 2.5% and 5.5% for the years 1990-91, 1997-98 and 1998-99 respectively. However, as the anchovy kilka catch declined, this species increased from 13.7% of the total catch in 1999 to 48.9% in 2003 (Sayyad Bourani et al., 2008).

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use as food and as bait.

Conservation

Stocks on the Iranian coast are said to have been depleted but its ecological specialisation on zooplankton means there is comparatively little competition with other fishes. It is probably not in any immediate danger. Kiabi et al. (1999) consider this species to be of least concern in the south Caspian Sea basin according to IUCN criteria. Criteria include commercial fishing, abundant in numbers, widespread range (75% of water bodies), absent in other water bodies in Iran, and present outside the Caspian Sea basin.

Further work

The biology of this species in Iranian waters needs to be elucidated.

Sources

Counts are based in part on Svetovidov (1945a). See also under family heading.

Iranian material: CMNFI 1970-0531, 14, 78.0-88.6 mm standard length, Mazandaran, Larim River (36º46'N, 52º58'E); CMNFI 1980-0146, 7, 79.9-96.2 mm standard length, Mazandaran, Gorgan Bay at Ashuradeh-ye Kuchak (36º50'N, 53º56'E); CMNFI 1993-0146, 3, 80.2-98.2 mm standard length, Mazandaran, Gorgan Bay (no other locality data); CMNFI 1993-0167, 1, 96.6 mm standard length, Mazandaran, Caspian Sea, 10 km offshore (ca. 36º49'N, ca. 52º39'E); CMNFI 1993-0168, 3, 84.9-88.0 mm standard length, Mazandaran, Caspian Sea, 10 km offshore (ca. 36º49'N, ca. 52º39'E).

Clupeonella engrauliformis
(Borodin, 1904)

Common names

rizeh keraye (= tiny ?), kilka-ye anchovy or kilka-e-anchovi.

[ancousabanzar kilka in Azerbaijanian; ancous sekilli kulke balyk in Turkmenian; anchousovidnaya tyul'ka or anchovy-like tyulka, sardelle or sardel'ka, "sardinka" but incorrectly, all in Russian; anchovy kilka, anchovy sprat].

Systematics

No major synonyms. Originally described from Buinak, central part of the Caspian Sea. The lectotype is in the Zoological Institute, St. Petersburg (ZISP 13860) with paralectotypes as established by Svetovidov (1952) (Eschmeyer et al., 1996). Eschmeyer et al. (1996) give the date as 1906 but Reshetnikov et al. (1997) give 1904.

Key characters

This species has a slender body (16-19% of standard length), a short and wide head (interorbital width 16-18.5% of head length), a rounded belly, and pointed pectoral fin tips.

Morphology

Dorsal fin with 3 unbranched and 12-14 branched rays, anal fin with 3 unbranched and 15-19 branched rays. Scales in lateral series 45-49. Vertebrae 44-47, rarely to 48 compared to 41-44 in the common kilka (C. caspia). Gill rakers number 56-67. Belly scutes 23-31.

Sexual dimorphism

None reported.

Colour

The back and head are dark blue with violet, green or olive tints. These colours become brighter or turn black in dead fish. The fins are hyaline except the caudal fin which has a black base and the dorsal fin which has a central dark stripe.

Size

Attains 15.5 cm standard length.

Distribution

Found in the central and southern Caspian Sea, and in Iranian waters the southeast Caspian Sea, southwest Caspian Sea and the south-central Caspian Sea (Kiabi et al., 1999) as well as the Anzali Mordab, Babol Sar Beach and Gorgan Bay (Armantrout, 1980). Abdoli and Naderi (2009) list it as from the southwest, southeast and south-central Caspian Sea, the Anzali Talab and Gorgan Bay in Iranian waters.

Zoogeography

This species is endemic to the Caspian Sea.

Habitat

The anchovy kilka, along with other kilkas, is the most abundant fish in the Caspian Sea forming large concentrations in the central and southern Caspian wherever water depth exceeds 30 m. The anchovy kilka is estimated to be the most numerous kilka at about 77% (Ivanov and Katunin, 2001; Daskalov and Mamedov, 2007). It is generally found in the upper water layers but may descend to 120 m. Nearshore areas, inlets and water of a salinity below 8‰ are avoided. They can tolerate a salinity range of 8-14‰ but the main part of the population is found at 10-12‰ (Fazli et al., 2007). Overwintering takes place in the southern Caspian and the southern part of the central Caspian Sea at 8.5-9.0°C and up to 13.5°C. Schools extend their range into the central and northern Caspian in spring to feed (Prikhod'ko, 1979b). This species has a hibernation period in the south Caspian Sea, a spring migration of part of the population to the central Caspian, a feeding period in the central and south Caspian and an autumn prespawning migration to the south Caspian (Sedov and Rychagova, 1983).

In Iran larvae are found mostly in surface layers at 5-20 m while adults are found in deeper zones. males dominate in winter while females dominate in other seasons. The maximum juvenile density (fish <75 mm), comprising 36% of the population, is seen in the summer (Iranian Fisheries Research and Training Organization Newsletter, 20:7, 1998). Jolodar and Abdoli (2004) state it is most abundant at 100-150 m.

Age and growth

Abundance of young anchovy kilka, and hence future year-class strengths, depends on water temperature in autumn (October-November). Falling water temperatures, in the eastern Caspian for example, are caused by upwelling which brings nutrients to surface waters and promotes growth of plankton on which the kilka larvae feed (Prikhod'ko, 1979a). Females are somewhat larger than males in the spawning areas. Sexual maturity is attained usually at age 2 and 2-4 year olds dominate catches but life span is up to 8 years (Iranian Fisheries Research and Training Organization Newsletter, 14:6, 1996). This species shows the fastest rate of growth in the genus. Of the 8 age classes, 0+, 1+, 2+ and 3+ form 99.91% of the whole population (Iranian Fisheries Research and Training Organization Newsletter, 20:7, 1998). The same study showed that 18.6% of the population matures in the first year of life while 81% matures in the second. The mean age in coastal areas is 2.9 years, slightly higher than that in deep zones below 200 m where 0+ fish are more abundant. The Caspian Sea Biodiversity Database (from www.caspianenvironment.org) gives a population of up to 293 billion fish in the Caspian Sea.

Fazli et al. (2007) and Sayyad Bourani et al. (2008) studied these kilkas from catches with conical liftnets carrying underwater lights in the fisheries of Gilan and Mazandaran in the 1995-2004 period. Fish were aged using the sagittal otoliths. Length and weight ranges were 40-140 mm and 0.4-18.4 g with averages of 94.0 mm and 5.7 g (89.2-100.4 mm from 1999 to 2003 in Sayyad Bourani et al., 2008). The age range was 1-7 years. The dominant age group varied from age 2 to age 4, making up 40.6% to 57.7% of the catch (Fazli et al., 2007) or 5+ years with 4+-5+ making up 84.6% for 1999-2003 (Sayyad Bourani et al., 2008). Growth was high for the first year of life and then gradually decreased. The von Bertalanffy growth equation was Lt = 148(1-e-0.238(t+1.340)) (Fazli et al., 2007, and following data). The sex ratio varied with season and was significantly different from equal at male:female = 0.78:1 for adults. Females were more abundant from January to June and males predominated from September to November. Condition factors differed significantly between years, increasing from 1995 to 1996, being lowest in 1998 and then increasing to 2004, and between months, being lowest in January and February and then increasing in March. 50% of fish were mature at 84.5 mm fork length. Annual survival rate was estimated at 0.32, the instantaneous coefficient of total mortality (Z) was 1.14/year, natural mortality was 0.473/year. Age at first capture was estimated as 2.92 years. The total biomass declined from 186,000 t in 1996 to less than 12,000 t in 2004 and the exploitation rate for 1995-2004 varied between 0.340 and 0.815. Sayyad Bourani et al. (2008) give a K value of 0.598/year and a L of 110.13 mm. Natural, fishing and total mortality coefficients were 0.69, 0.31 and 1 per year respectively and the sex ratio was female:male = 68.2-31.8. These latter results for the 1999-2003 period show how value scan change when subsets of data are used. Fatemi et al. (2009) examined fish taken from commercial vessels in 2007 using lift nets and lights. Age structure ranged from 2 to 7 years and was dominated by the third year class (38.6%). Back-calculation methods were validated using otoliths to determine lengths. Karimzadeh et al. (2010) examined fish from the Babolsar region off Mazandaran and calculated growth parameters as L = 151.9 mm, K = 0.28/yr-1 and t0 = -1.12/yr, instantaneous coefficient of natural mortality was 0.633/yr-1 and the current exploitation rate was estimated as 0.41.

Food

Plankton is the main food and copepods predominate but diet also includes Cladocera, Balanus larvae and clam larvae. The dominant food item is the copepod Eurytemora grimmi, particularly in winter when plankton biomass is lowered (Badalov, 1972). It can make up over 70% of its food. This copepod is more characteristic of the diet of this kilka compared to the other two species and the daily vertical migrations and seasonal movements of the copepod are mirrored by the kilka. The most abundant fish species in the Caspian depends on the most abundant member of the crustacean zooplankton (Prikhod'ko, 1979b). This species feeds in winter, unlike Clupeonella caspia. Bankehsaz (1996) surveys the fluctuation in fat content of this species through the year. Intensive feeding begins in spring as a preparation for spawning (Sedov and Rychagova, 1983). Spawning males show a positive response to light and so feed during the spawning season, while females do not. F. Darvishi (pers. comm., 2003) has demonstrated that the this species has a similar feeding niche as the exotic ctenophore Mnemiopsis leidyi and Esmaili Sari et al. (2002) determined that there is a similar diet in Iranian waters suggesting that a decline in stocks of the fish is the result of competition. Darvishi et al. (2004) studied catches of the anchovy kilka and the ctenophore in the southern Caspian Sea from August 2001 to October 2002. Dietary overlap was >89 in Babolsar samples and >84 in Nowshahr samples using the Schoener Index (presumably 0.89 and 0.84 where 0 is no dietary overlap and 1 is an identical diet). The ctenophore was also feeding on fish eggs but the effect of this was less than competition for food.

Reproduction

Spawning ends in late autumn and winter food requirements are higher than in spring-spawning C. caspia (Badalov, 1972). Areas for spawning in this species are extensive. Spawning is most intensive in July when temperatures are 13-24°C and salinity 8-13‰ although the Caspian Sea Biodiversity Database (from www.caspianenvironment.org) gives peak spawning (70%) as in October-November. Fazli (2006a) gives spawning in Iran as spring and autumn but mass spawning takes place in in autumn. Spawning takes place in the central and southern Caspian along both eastern and western shores both in coastal regions and the open sea from late April to November. Mass spawning takes place at depths of 50-200 m and as a result eggs and larvae are carried over a wide area by the Caspian gyral current at these depths (Prikhod'ko, 1979b). Young hatch mainly in autumn and reach 4.5-8.0 cm at an age of 8-10 months (Prikhod'ko, 1979a). Eggs are up to 1.82 mm in diameter and fecundity reaches 39,900 eggs.

In Iran, 80% of the population spawn in autumn and the remainder in spring. Accordingly the fishery should be closed in October and November (Iranian Fisheries Research and Training Organization Newsletter, 19:5, 1998). The subsequent Iranian Fisheries Research and Training Organization Newsletter (20:7, 1998) states that 89% of the population spawns in autumn with September, at 68.3%, the major month. Fazli et al. (2007) found reproduction to start in June, peaking in October and then declining.

Parasites and predators

Samples of this species from Babol Sar and Bandar Anzali contain the digenean trematode parasites Pseudopentagramma symmetrica and Bunocotyle cingulata, the acanthocephalan Corynosoma strumosum and larvae of the nematode Contracaecum sp. (Iranian Fisheries Research and Training Organization Newsletter, 11:4-5, 1996; Shamsi et al., 1996; Annual Report, 1995-1996, Iranian Fisheries Research and Training Organization, Tehran, p. 28, 1997; Shamsi and Dalimi, 1996; Shamsi et al., 1998). Clupeonella species are an important food fish for sturgeons (59.4% by weight of sevryuga diet in the Middle Caspian), Sander (Percidae) and herrings and the Caspian seal (Badalov, 1972; Krylov, 1984) as well as other fishes. Varshoie et al. (2010) record the helminths Pseudopentagramma symmetrica, Bunocotyle cingulata and Mazocreas alosae in this species from Iranian waters.

Economic importance

This species forms 80-90% of the catches of kilkas in former Soviet waters (Sedov and Rychagova, 1983) and, as noted above, 91.8% of catches in an Iranian study (Razavi Sayad, 1993b; Rezaei et al., 2003). High catches are related to the larger spawning and foraging range of this species compared to other kilkas and to its habitat in the Caspian gyre, an area of increased biological productivity (Prikhod'ko, 1979b). It is caught in former Soviet waters by attraction to underwater electrical lights attached to the middle of the mouth of a fine-mesh conical net or the sides of a fish pump (Ben-Yami, 1976). Fishing is suspended at full moons as the fish are dispersed (Saheli, 1999). Both large and small individuals are taken by these non-selective methods (Prikhod'ko, 1981). Incidental catches include Mugilidae (common), and Alosa spp., Atherinidae and the cyprinid Pelecus cultratus (all occasional) (Ben-Yami, 1976).

It is regarded as a valuable and cheap food resource in Iran where it is canned, made into sausages and surimi, and processed as fish meal (Shamsi et al., 1996; Moeini, 2002; Shabanpour et al., 2002,,2006). The catch per unit effort for funnel nets and midwater trawls is 2321 and 1014 respectively (Iranian Fisheries Research and Training Organization Newsletter, 20:7, 1998). Various studies on its preparation and storage as food have been carried out, e.g. Rezaei et al. (2002; 2003; Moeini et al., 2009).

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use as food and as bait.

Conservation

Prikhod'ko (1981) recommends fishing in deeper waters where larger fish are concentrated to avoid an excessive take of young fish which favour the upper water layers. Stocks in the southern Caspian Sea are said to be depleted. Kiabi et al. (1999) consider this species to be of least concern in the south Caspian Sea basin according to IUCN criteria. Criteria include commercial fishing, abundant in numbers, widespread range (75% of water bodies), absent in other water bodies in Iran, absent outside the Caspian Sea basin. Daskalov and Mamedov (2007) studied commercial catch data in the Caspian Sea generally and found a period of high catches from 1991 to 2000 with high spawning-stock biomass and relatively good recruitment. Catches peaked at 271,400 t, fishing mortality reached 1.8y-1 in 1999 and overfishing occurred. From 2001 to 2004, the stock collapsed, recruitment failed in 2001 and catches fell to 54,300 t in 2005. This was attributed to the spread of the ctenophore Mnemiopsis leidyi, with contributions from overfishing. Fazli et al. (2007) also concur that both overfishing and the invasive ctenophore caused the collapse of stocks. The catch in Iran declined from 71% of the total kilka catch in 1999 to 52.5% in 2003 (Sayyad Bourani et al., 2008).

Further work

The biology of this species in Iranian waters needs to be elucidated.

Sources

Iranian material: CMNFI 1993-0167, 1, 99.5 mm standard length, Mazandaran, Caspian Sea (ca. 36º49'N, ca. 52º39'E); CMNFI 1993-0168, 4, 89.3-107.6 mm standard length, Mazandaran, Caspian Sea (ca. 36º49'N, ca. 52º39'E).

Clupeonella grimmi
Kessler, 1877

Common names

kilka-ye cheshmdorosht (= bigeye kilka).

[irikoz kilka in Azerbaijan; sardelle or sardel'ka, bol'sheglazaya tyul'ka or bigeye tyulka, bol'sheglazaya kil'ka or bigeye kilka, all in Russian; southern Caspian sprat].

Systematics

Clupeonella Grimmi was originally described from the central part of the Caspian Sea. The lectotype is in the Zoological Institute, St. Petersburg under ZISP 10934 as designated by Svetovidov (1952).

Harengula macrophthalma Knipovich, 1921 is a synonym. Four syntypes are in the Natural History Museum, London under BM(NH) 1897.7.5:41-44 (when examined were numbered 42-44, 3 fish, 29.9-33.5 mm standard length in poor condition, September 2007), with many others apparently in the Zoological Institute, St. Petersburg (Eschmeyer et al., 1996).

Key characters

This species has a moderately slender body (17-22% of standard length), a long and narrow head (interorbital width 13-15% of head length), a sharply keeled belly, and rounded pectoral fin tips.

Morphology

Dorsal fin unbranched rays 3-4, usually 3, branched rays 13-15, and anal fin unbranched rays 3, branched rays 14-21. There are 44-49, usually 46-48, vertebrae, more than in the other two kilka species and probably a consequence of the low water temperature larvae develop in. Belly with 26-32 scutes. Gill rakers 42-51.

The bigeye kilka is adapted to life in deeper water having, as its name indicates, big eyes with more rod cells and a weaker retina but also more transparent body tissues than other kilkas.

Sexual dimorphism

None reported except size.

Colour

The back and top of the head are dark.

Size

Reaches 14.5 cm standard length.

Distribution

Found in the Caspian Sea and concentrated in the south including Iranian waters. Abdoli and Naderi (2009) list it as from the southwest, southeast and south-central Caspian Sea in Iranian waters.

Zoogeography

This species is endemic to the Caspian Sea.

Habitat

The bigeye kilka is found further away from the coast than the anchovy kilka at depths over 50-70 m, down to 450 m, with large schools down to 130 m. It does not enter fresh water or low salinity areas, staying well away from the shore. There is a daily vertical migration, avoiding sunlight, and following food items. Larvae live in water temperatures of 5°C. Overwintering occurs in the southern Caspian at temperatures of 9-11°C, a migration to the central Caspian takes place in spring, with a return south in autumn (Prikhod'ko, 1979b).

Age and growth

Sexual maturity is attained usually at age 2, and 2-4 year olds dominate catches, but life span is up to 8 years (Iranian Fisheries Research and Training Organization Newsletter, 14:6, 1996). The female is larger than the male at the same age. Growth is slower than in C. engrauliformis. Males dominate the population (Iranian Fisheries Research and Training Organization Newsletter, 14:6, 1996; Fazli et al., 2005) but this study may have sampled spawning fish (see below).

Fazli et al. (2005) examined fish from the main landing ports (Babolsar, Amirabad and Anzali) found the mean fork length of fish increased from 95.87 mm in 1997 to 105.0 mm in 2000 but then decreased to 102.3 mm afterwards. Over this time period, fork length range became wider with specimens in the upper length classes representing most of the catch. Six age classes were present, 1+ to 6+ years. During 1998-1999, age classes 1+ to 3+ comprised more than 90% of the catch. In 2000, there was a decrease in age classes 1+ and 2+ and an increase in 3+ to 5+ classes. In 2001, age classes 3+ and 4+ decreased and classes 5+ and 6+ increased. The relative frequency of the bigeye kilka has decreased in recent years as a result of the introduction of the ctenophore, Mnemiopsis leidyi, a food competitor and predator on kilka eggs and young. Khorashadizadeh et al. (2006) found fish in the Babolsar area of the Iranian coast to have 5 age classes, dominated by the 4+ class. Fazli et al. (2009) examined changes in the population biology of this kilka over the period 1995 to 2001, attributed to the inavsive ctenophore. The overall sex ratio was 1.65:1 in favour of males, length-weight regressions were W = 0.00922L2.851 for females and W= 0.008021L2.907 for males, indicating a negative growth for both sexes, growth parameters were L = 142 mm, K = 0.28 year-1, and t0 = -1.39 years, the instantaneous coefficient of natural mortality was 0.460 year-1, and the instantaneous coefficient of fishing mortality varied between 0.469 and 0.980 year-1. Biomass increased from 36,900 mt in 1995 to more than 53,500 mt in 1998 but declined to less than 5900 mt in 2001. This was attributed to overfishing and the appearance of the ctenophore, a competitor for zooplankton food.

Karimzadeh et al. (2010) examined fish from the Babolsar region off Mazandaran and calculated growth parameters as L = 148.6 mm, K = 0.46/yr-1 and t0 = -0.18/yr, instantaneous coefficient of natural mortality was 0.881/yr-1 and the current exploitation rate was estimated as 0.26.

Food

Migratory mysids often predominate in the planktonic diet of this species. Fish fry are also eaten. Its foods are less diverse than that of other kilkas because the variety is less in the deeper waters this fish inhabits during the day. The three kilkas share the available habitat and its foods, the common kilka in shallow, coastal waters, the anchovy kilka in the upper layers of the open sea and the bigeye kilka in deeper water of the open sea (Badalov, 1972; Prikhod'ko, 1979b).

Reproduction

Spawning is extended, from January through to September but is most intense in spring and autumn (Prikhod'ko, 1979b). Males predominate in the spawning areas, remaining there while females leave immediately after spawning. Males are mainly at 10-20 m and females at 20-25 m during the spawning season. Water temperatures at 6-13°C and salinity 12.6-13.0‰. Fecundity is 28,300 eggs. In Iranian waters, mature fish ready to spawn are always present in catches in winter and early spring (Fazli et al., 2005). Khorashadizadeh et al. (2006) found fish in the Babolsar area of the Iranian coast to have peak spawning in early January.

Parasites and predators

Samples of this species from Babol Sar and Bandar Anzali contain the digenean parasites Pseudopentagramma symmetrica, Bunocotyle cingulata, the acanthocephalan Corynosoma strumosum, Eustrongylides excisus, and larvae of a Contracaecum and an Anisakis species (Iranian Fisheries Research and Training Organization Newsletter, 11:4-5, 1996; Annual Report, 1995-1996, Iranian Fisheries Research and Training Organization, Tehran, p. 28, 1997; Shamsi and Dalimi, 1996; Shamsi et al., 1998; Shamsi et al., 1998). Varshoie et al. (2010) record the helminths Pseudopentagramma symmetrica, Bunocotyle cingulata and Mazocreas alosae in this species from Iranian waters.

Clupeonella species are an important food fish for sturgeons (59.4% by weight of sevryuga (Acipenser stellatus) diet in the Middle Caspian), Sander (Percidae) and herrings and the Caspian seal. Predators consume 590 million kg of the three kilka species which themselves are the main consumers of zooplankton. Kilkas are a very important element in the life of the Caspian Sea (Badalov, 1972; Prikhod'ko, 1979b; Krylov, 1984). This species is taken to a lesser extent than other Clupeonella species because it is relatively sparse.

Economic importance

The bigeye kilka catch amounts to about 70 million kg a year in former Soviet waters of the Caspian by means of electric light. All three kilka species are caught by using underwater electric lights and fish pumps (Nikonorov, 1964) but in the case of the bigeye the effect is avoidance used to drive it to the bottom where it can be caught. Other kilkas are attracted to the light but the bigeye is a vertical migrator, avoiding sunlight (Prikhod'ko, 1979b). Light-assisted catches of kilkas damages young shad (Alosa) stocks which are an incidental catch (Zakharyan and Teruni, 1979). Catches in Iranian waters are only 6.84% of the total kilka take (Razavi Sayad, 1993b). The relative frequency of the bigeye kilka in Iranian catches was ranked second after anchovy kilka in 1990-1991 at 6.84%, increasing to 12.6% and 21.7% in 1997 and 1998 and then decreasing.

Omega-3 fatty acids from fish oil of this species has been tested as a dietary supplement and was found to relieve symptoms of dysmenorrhoea (Moghadamnia et al., 2010).

Conservation

Stocks in Iranian waters are said to be depleted. Kiabi et al. (1999) consider this species to be of least concern in the south Caspian Sea basin according to IUCN criteria. Criteria include commercial fishing, abundant in numbers, widespread range (75% of water bodies), absent in other water bodies in Iran, and absent outside the Caspian Sea basin.

Further work

The biology of this species in Iranian waters needs to be elucidated.

Sources

Iranian material: CMNFI 1993-0167, 1, 93.0 mm standard length, Mazandaran, Caspian Sea (ca. 36º49'N, ca. 52º39'E); CMNFI 1993-0168, 2, 91.8-94.0 mm standard length, Mazandaran, Caspian Sea (ca. 36º49'N, ca. 52º39'E).

Genus Tenualosa
Fowler, 1934

This genus comprises 5 species found from the Indian Ocean to Indonesia and China. A single species enters rivers of southern Iran. The genus is defined by a series of characters listed below under Key characters. These fishes form part of local, artisanal fisheries throughout their range.

Tenualosa ilisha
(Hamilton, 1822)

Common names

صبور (= sobur, soboor, sobour, sabur, zobur, zabur, zamur or zomur, all variants of the same word), bari, barak; mahi-ye khor kuchiku (= small bone fish, at Abadan from www.abadan.com/abadanhistory.html, 15 March 1998).

[zoboor, soboor, sbour in Arabic; hilsa, Indian shad or river shad; palo, palla or pulla and tikki-palwar in Pakistan].

Systematics

Clupanodon ilisha was originally described from the Ganges estuaries in India. Formerly placed in the genus Hilsa Regan, 1917. Al-Hassan (1982), citing a personal communication from a Mr. Al-Abaychi in 1973, suggests that Shatt al Arab fish are distinct from those in Pakistan on morphometric and meristic grounds but no data have been published. Milton and Chenery (2001) used genetic and otolith chemistry data that provided strong evidence for a distinct stock in Kuwait, compared with stocks from India to Sumatra. Al-Hassan (1999) mentions that people in Basrah can distinguish two kinds of sobur, based on taste. One is the tastier and pricier Shatt al-Arab form and the other is the less desirable estuarine/sea form. This has not been confirmed by systematic studies. Jorfi et al. (2008; 2009) found differences between populations in Iran and Iraq using molecular techniques.

Key characters

This species is distinguished from other Indian Ocean clupeids by the upper jaw with a median notch, the anal fin ray count being less than 30 rays, a terminal mouth (lower jaw not prominent nor flared at the corners), scales in lateral series are not perforated posteriorly, last dorsal fin ray not filamentous, weakly developed lines (the fronto-parietal striae) on top of the head (usually covered by skin and not visible), gill rakers on inner arches straight not curled, a long head 28-32% of standard length, and 30-33 ventral scutes forming a keel along the belly, 15-18 being prepelvic and 11-15 postpelvic (Al-Nasiri and Al-Mukhtar, 1988a, 1988b; Marammazi et al., 1995).

Morphology

Dorsal fin with 4-5 unbranched rays followed by 14-16 branched rays, anal fin with 2-3 unbranched rays followed by 16-20 branched rays, pectoral fin branched rays 12-15 and pelvic fin branched rays 7. Lateral series scales 44-51. Gill rakers are fine and numerous, up to about 275 on the lower arch.

Iranian fish examined by Marammazi et al. (1995) from the Bahmanshir River in Khuzestan have 30-32 total scutes along the belly, 16-18 prepelvic scutes, 13-15 postpelvic scutes, 19-21 dorsal fin rays, 19-24 anal fin rays, 13-15 pectoral fin rays, 8 pelvic fin rays and 44-51 scales.

Sexual dimorphism

None reported.

Colour

The back is grey-blue, bluish to green and the sides are silvery with golden, purplish or pink highlights. The dorsal fin is grey, the caudal fin grey-blue with a silvery tinge and darkened margin, and the anal fin is light blue with some silvery tinges. Paired fins are hyaline. The area behind the gill cover in young fish and many adults have a dark blotch followed by a series of spots or blotches running along the upper flank, for a total of 6-7. The blotches may take the form of bars. The eye is yellow to red. Young have a bronze back, silvery flanks and a caudal fin margined in black.

Size

Attains 60.6 cm total length and 2.49 kg for females and 43 cm and 0.68 kg for males. A sample of 233 moribund fish from the Ashar Canal, a branch of the Shatt al Arab, Iraq examined by Al-Nasiri and Al-Mukhtar (1988a; 1988b) had a total length range of 70-152 mm. Hussain, Jabir and Yousif (1994) record fish migrating to the Shatt al Arab for breeding at 21-38 cm for males and 33-43 cm for females. Mature females in the Shatt al Arab weighed about 0.5-1.1 kg (Jabir and Faris, 1989). Fishes from Kuwait attained 57 cm (Al-Baz and Grove, 1995). Fishes from the Arvand, Bahmanshir, Karun and Dez rivers of Iran were 120-500 mm long (Marammazi et al., 1998; Ghafleh Marammazi et al., 2004).

Distribution

Reportedly found from the Red Sea and Persian Gulf through the Indian subcontinent to the Malayan Archipelago in some general works, or more narrowly from the Persian Gulf to Myanmar. It enters the Shatt al Arab and Tigris River, once as far north as Baghdad (Kanazawa, 1955), but the northernmost distribution today in Iraq is the Hawr al Hammar. Before the construction of dams on the Euphrates the migration was up to "Yaou" and "Meshkhau" and up to Qal`at Salih (31°31'N, 47°16'E) in the Tigris of Iraq (van den Eelaart, 1954).

The lower reaches of the Tigris and Euphrates rivers were connected by a channel to the Khor Al-Zubair in Iraq during 1983. As a consequence the Khor became oligohaline (at less than 10‰) rather than hypersaline (at more than 40‰), becoming an estuary with heavy reed growth. The catch of sobour in the Khor by 1997 exceeded that in the Shatt al Arab and may involve diversion of stocks from the original habitat of the Shatt (Hussain, 1997).

In Iran, it is recorded as far north as the Gargar Shoteit on the Dez River (Marammazi, 1994). Hussain, Jabir and Yousif (in litt., 1995) record this species from the Shatt al Arab in Iraq and the Bahmanshir, Jarrahi, Zohreh and Hilleh rivers in Iran. Marammazi (1994) and Marammazi et al. (1998) report this species from the Arvand, Bahmanshir, Karun and Dez rivers. Ghafleh Marammazi et al. (2004) record it from the Zohreh, Bahmanshir, Arvand and Karun rivers in Iran. It may be found in the Hormuz basin but this has not been verified with specimens.

In the sea, they are found from Bushehr around to Kuwait in coastal waters (Blegvad and Løppenthin, 1944; Hussain, Jabir and Yousif, in litt., 1995).

Zoogeography

Al-Hassan (1982) mentions a study comparing a population of this species from Basrah, Iraq with one from Pakistan and finding significant meristic and morphometric differences, perhaps indicative of distinct stocks.

Habitat

Sobour enter the Shatt al Arab in February and March during high tides and feed there until the fall according to a study by Al-Nasiri and Al-Mukhtar (1988a; 1988b) working on fish taken from the Ashar Canal, Basrah, Iraq. van den Eelaart (1954) reports that most fish enter the Shatt al Arab in April during the last and first phase of the moon and anecdotal reports indicate the end of March to be the peak period of entry. They ascended into the Hawr all Hammar and from there into the Euphrates as well as into the Tigris (van den Eelaart, 1954). Significant numbers were recording as entering the recovering Hawr al Hammar in 2005-2006 (Hussain et al., 2006). Small specimens (50-100 mm) were observed in the east Hawr al Hammar in June 2005 and July 2006 (www.iraqmarshes.org, downloaded 29 August 2005; N. A. Hussain, in litt., 2006). In mid-April sbour were found below the Yaou and Moshkhab regulators which formed the limit of their migration on the Euphrates in the early 1950s. The limit in the Tigris was beyond Amara. The main spawning grounds in the Euphrates were probably somewhere between Shinafiya and Samawa and in the Tigris between Amara and Qalat Saleh.

The last ones leave the Shatt in July and fry are found in the rivers of Iraq at the end of the June. Hussain, Jabir and Yousif (1994) record sobour ascending the Shatt al Arab during March with a continuing migration upstream through April to July for spawning and a return migration to the sea during August to October. Al-Hassan (1993) notes that local people believe that sobour ascend the Shatt al Arab during spring to marshes north of Basrah for spawning, suggesting that they are the fluvial anadromous type. Al-Hassan (1999) considers they migrate to the sea in September-November, when they are landed in Kuwait, and they then migrate to the Iranian coast during December-January. Males and females move upriver in separate groups according to Iraqi fishermen (Al-Hassan, 1999).

Jorfi et al. (2008) suggest, based on molecular studies, that a population in the Persian Gulf chooses the Karun River for spawning and migrates via the Bahmanshir River, while others migrate up the Tigris and Euphrates rivers in Iraq via both the Bahmanshir and the Arvand rivers.

Blegvad and Løppenthin (1944) mention this species on sale at Khorramshahr on 28-29 April. The spawning migration in Iran occurs in spring (I. Sharifpour, in litt., 1991). It is only found in the Zohreh River in spring and summer (Marammazi, 1994).

They may be found in deep water, over 18 m, or in shallows, on their spawning migration. Large concentrations of sobour occur below dams blocking their migration. Young occur in side branches of the Shatt al Arab near food, shelter and the spawning grounds (Hussain, Jabir and Yousif, in litt., 1995).

This species occurs in river estuaries and coastal waters and appears to be restricted to the northern end of the Persian Gulf because this is the only part with large spawning rivers (Hussain, Jabir and Yousif, in litt., 1995). These authors also suggest that an anadromous stock from the Shatt al Arab migrates to warmer waters off Bushehr during January, February and March. At the same time there is a winter decline of Kuwaiti stocks. There may also be a marine stock inhabiting coastal waters of Kuwait since larvae have been found in Kuwait Bay during June and November and catches are made in the Bay year round.

Biogenic and anthropogenic sources were noted for the hydrocarbons in this species from the Shatt al Arab; n-alkanes attained 31.11 µg/g and hydrocarbons 10.91 µg/g, the highest for the fish species studied (Al-Saad et al., 1997). The fat content of this shad is a factor in these high levels (Al-Saad, 1990).

Hussain (1997) notes that the changing conditions in the Khawr az Zubayr, which became oligohaline from hypersaline after it was connected to the Tigris-Euphrates basin by the Shatt al Basrah Canal. In 1994 fishermen began catching sbour in the Khawr az Zubayr and by 1997 the numbers caught exceeded the catch in the Shatt al Arab.

Migrations in the Indus River of Pakistan (Islam and Talbot, 1968) may last over 7 months and the migration up the Ganges River extends over 1287 km. Fish may move as much as 70.8 km in one day and may jump out of the water on the migration.

Age and growth

Al-Nasiri and Al-Mukhtar (1988a; 1988b) give a length-weight relationship of W = 3.9 x 10-6 L3.16 or log W = 3.16 log L-5.4 for fish aged at 0+ from the Ashar Canal at Basrah. The mean condition factor was 0.87. Fishes in the Shatt al Arab are in age groups 5 to 6 for the period May to August (Hussain et al., 1991). In contrast, a later study on the Shatt al Arab fish showed there are 5 age groups and the second and third age groups dominate in catches (Hussain, Jabir and Yousif, 1994). In this latter study, Shatt al Arab fish mature at 25 cm for males and 33 cm for females, similar to an Iranian study (see below). The length-weight relationship was log W = -4.7074 + 3.0479 log L for females and log W = -4.5802 + 3.0193 log L. Condition factor gradually increased with length groups in males, peaking at 32-33 cm followed by a sharp decline while females had a nearly stable condition factor from 34 to 43 cm. Mohammed et al. (2001) gave a von Bertalanffy growth equation as L = 60.47 cm and a condition factor of 0.32, slower growth than in Indian and Bangladesh populations and probably maturing later.

Amodeo (1956) gives lengths of 25 to 35 cm for fish caught in the Shatt al Arab on their spawning migration. Young grow rapidly, 4.3 cm in October-November. Most fish on the migration in the Indus River were in age groups 3 and 4. Life span is up to an estimated 7 years with maturity as early as 1 year. Jawad et al. (2004) found haematocrit level to increase with body length up to 40 cm after which it decreased, males showed higher levels than females, and levels were higher pre-spawning than during spawning and increased slightly post-spawning, a general correlation with fish activity.

Al-Baz and Grove (1995) studied fish taken from Kuwait fish markets. Females dominated the catch, male:female ratio being 1:2.4, perhaps because the sexes moved in different schools. The smallest mature female was 34.4 cm and 50% of the females are mature at 41.5 cm. They estimated natural mortality (M) based on von Bertalanffy growth parameters (L and K) and mean annual water temperature as log M = -0.0066 -0.279 log L+ 0.6543 log K + 0.4634 log T. The length-weight relationship was W = 0.011 L2.983 for males and W = 0.007 L3.104 for females. Growth in the sexes follows different patterns. Five age groups were detected using otoliths and fish were fully recruited to the fishery at 3 years of age. von Bertalanffy growth parameters were L = 52.70 cm and condition factor (K) = 0.28 per year while using Allen's method they were L =52.50 cm and condition factor (K) = 0.36 per year Growth curves were given. Annual total mortality was estimated to be 1.2 using the K value of 0.36. A fishing mortality was calculated to be 0.8 per year.

In the Bahmanshir River, Iran most fish are 4-5 years old. The minimum total length and age at maturity are 26.2 cm, 200 g and 2 years for males and 32.18 cm, 450 g and 3 years for females. Von Bertalanffy growth parameters in Iranian females are L = 57.78 cm and K = 0.282 and in males 46.37 cm and 0.252 (Marammazi, 1995; Iranian Fisheries Research and Training Organization Newsletter, 12:5, 1996; Annual Report, 1995-1996, Iranian Fisheries Research and Training Organization, Tehran, p. 53-54, 1997).

Hashemi et al. (2009) studied fish landed at Hendigan and Abadan and recorded L as 42.81 cm, K was 0.9, M was 1.37, F was 2.41, Z was 3.78 and E was 0.64. Y'/R was 0.048 and B'/R was 0.19, exploitation rate (U) was 0.61, annual stock at the beginning of the year (P) was 7615 t, annual standing stock (b) was 1927 t and MSY was 3642 t. The stock was overfished.

Hashemi et al. (2010) studied 9317 fish from landings at Abadan and Hendijan. Size range was 20-39 cm. The von Bertalanffy growth parameters were L = 43.32 cm, K = 0.78 yr-1, Φ' was 3.16 and t0 was -0.18. Mortality rates were M = 1.29 and Z = 4.53, and fishing mortality (F) was 3.24 yr-1. The exploitation rate (E) was 0.72 and the stock was overfished. Values of the sizes where the probability of capture was 50% (L50) and 100% (L100) were 22.3 and 28.5 cm TL respectively. Fish were recruited to the fishery at a mean size of  L100 = 22.3 cm. The relative yield per recruit (Y'/R) was 0.062, relative biomass per recruit (B'/R) was 0.12 and exploitation rate (U) was 0.76. The values for annual catch, total annual stock, standing stock and maximum sustainable yield were 4645 t, 6635.71 t, 1433.64 t and 3274.19 t respectively. The fishing pressure must be reduced from 3.24 yr-1 to about 0.97 yr-1for this population to be adequately managed.

Roomiani and Jamili (2011) examined fish landed in Iran from a northern Persian Gulf fishery. Growth was isometric. Maximum total length was 43 cm and weight 949 g. von Bertalanffy growth parameters were  L = 42.74 cm total length, K = 0.77 and t0 = -0.21 years-1. Total mortality (Z) was 2.55 years-1, natural mortality was 0.75 years-1, fishing mortality was 1.8 years-1, and exploitation rate (E) was 0.7 years-1, and paarmeters indicate overfishing. Maximum sustainable yield was calculated to be 2653 t.

Food

The Ashar Canal study found them to feed on phytoplankton such as dinoflagellates and diatoms and on zooplankton, mainly copepods, as well as their own young. The sieve-like gill rakers are used to strain out planktonic organisms without selection. Presence of some sand grains indicates that feeding can occur on the river bed. Feeding intensity may decrease or cease on the spawning migration and is very high after spawning. The Bahmanshir fish feed principally on copepods and diatoms. Shatt al Arab juveniles feed mostly on filamentous algae and diatoms with some organic matter, fish eggs and zooplankton while adults have empty stomachs on the spawning migration (Hussain, Jabir and Yousif, in litt., 1995). In the Indus River, the newly hatched larvae and juveniles graze for five to six months in fresh waters before they migrate to the sea (http://www.jang-group.com/thenews/feb2003-daily/18-02-2003/business/b2.htm , downloaded 18 February 2003).

Reproduction

The spawning migration depends on the flood regime of the rivers. Turbid water and fast current are probably stimulants to egg deposition. The sbour depends on river-edge vegetation for egg deposition. Spawning grounds in Iraq are probably located near the beginning of the side branches of the northern sector of the Shatt al Arab, 120 km from the sea (Hussain, Jabir and Yousif, 1994). This species is gonochoristic (Blaber et al., 1997). Males may ascend the river before females but females become dominant in Indian populations. Males dominate in March in the Shatt al Arab and the sex ratio reaches equilibrium in the spawning months of May-July (elsewhere in the same communication spawning is given as June to August) (Hussain, Jabir and Yousif, 1994; Jawad et al. (2004). Spawning may occur more than once in a season in India. This has not been demonstrated for Iran but could occur. The gonadosomatic index for fishes in the Iraqi Shatt al Arab indicates peaks in March-May and July-August, suggesting two spawnings (Hussain et al., 1991) although a later report (Hussain, Jabir and Yousif, 1994) gives spawning as June to July and July to August as evidenced by two modes of juveniles found in September. Sex ratio is equal during this period. All females entering the Shatt al Arab were mature with smallest female being 33.0 cm long. Males less than 25.0 cm were immature, the population reaching 100% maturity at 31-32 cm (Hussain, Jabir and Yousif, 1994). The Kuwait fish studied by Al-Baz and Grove (1995) indicated spawning between May and July with a peak in June.

Fecundity in the Indus River population was estimated to be up to 2,917,000 eggs per female, egg diameters reached 0.89 mm, and the hatching takes place in about 23 to 26 hours (http://www.jang-group.com/thenews/feb2003-daily/18-02-2003/business/b2.htm, downloaded 18 February 2003). Estimates for the Hooghly River of India reach 13,230,500 eggs per female (Al-Hassan, 1993). Fecundity in the Shatt al Arab ranges between 444,960 and 1,616,560 eggs for fish 33.0-41.5 cm total length although 2 fish 37.3 and 2 fish 39.0 cm total length had a range in egg numbers of 109,000-233,840, showing that great variations in fecundity occur between individuals; possibly some fish had partially spawned before capture (Jabir and Faris, 1989). This latter study gave a relationship between absolute fecundity and total length as F = 1.3699 L3.6681 and log F = 0.1367 + 3.6681 log L and between fecundity and weight F = 302.8214 W1.2087 and log F = 2.4812 + 1.2087 log W. Fecundity increased significantly with body weight, ovary weight and total length. Relative fecundity (ova/gramme body weight) varied from 737 to 1721, mean 1216.

Hatching can occur within one day at an average temperature of 23°C. Eggs, larvae and young are found on the spawning grounds but with growth the young move into estuarine and foreshore areas during winter months. Hussain, Jabir and Yousif (1994) record the appearance of juveniles from the northern Shatt al Arab from June to November. Adults return to their original habitat in the sea after spawning. There is some evidence for freshwater resident populations in India which migrate upriver to spawn but do not descend to the sea.

The Bahmanshir fish are thought to spawn from April to July. Only adults enter the Bahmanshir (Iranian Fisheries Research and Training Organization Newsletter, 12:5, 1996). Absolute fecundity of fish from the Arvand, Bahmanshir, Karun and Dez rivers ranges from 374,892 to 1,954,144 eggs for total lengths of 380 to 500 mm respectively and is related to age. Ova with diameters 0.64-0.795 mm were released spontaneously in a study of this fish in Khuzestan province, in several batches along its migration route (Ghafleh Marammazi et al., 2004). Spawning begins on entry to the Bahmanshir and Arvand rivers in Khuzestan in April, continuing to September and the end of their migration at the cities of Shushtar and Dezful higher upriver. Males enter these rivers first in March, followed by females in April (Ghafleh Marammazi et al., 2004).

Parasites and predators

None reported from Iran other than nematode larvae by Ebrahimzadeh and Nabawi (1975) for fish from the Karun River.

Economic importance

The Ashar Canal study cites 996,308 kg reaching the Ashar fish market from October 1975 to June 1977 (see also Sharma, 1980). The catch landed at Fao on the Shatt al Arab estuary of Iraq was 6576 t in 1990-1991 (L. A. J. Al-Hassan, in litt., 1995; however this seems much too high although the estimate is from the Food and Agriculture Organization). This species forms the most important commercial fishery in the Basrah region of southern Iraq, average catches being 491.086, 319.661 and 267.988 t in 1977, 1978 and 1979 respectively (sic, Jabir and Faris, 1989). There is a drift-net and stake-net ("hadra") fishery in the sea by Kuwait in Kuwait Bay and around Falaikah Island (Al-Baz and Grove, 1995).

The fishing season on the Tigris-Euphrates is March to August with a peak in April, or late April to early June (Jabir and Faris, 1989) or to November (Ali et al., 1998). van den Eelaart (1954) gave the fishing season for this species as March-August (peaking in April) in rivers, and March-May (peaking in April) in Hawr al Hammar, Iraq. Fish are caught at the mouth of the Shatt al Arab as they enter the river with stationary gill nets, drifting gill nets, in "mailan" and "odda" traps from March to August. The catch averaged 150-180 kg per ten odda and in March 1953 the total catch at the mouth of the Shatt al Arab was about 25,000 kg (Amodeo, 1956). Large fish are only caught in the summer (Al-Hassan, 1999).

The catch at Abadan from February to November in 1943 was about 401.42 t and from January to June about 336.67 t (Pillay and Rosa, 1963). This species is seen on markets at Ahvaz, Khuzestan in November but these are sea-caught fish. Marjan Iran Company was selling 600-800 g fish for U.S.$1.40/kg, 800-1000 g fish for U.S.$1.60/kg, 1000-1200 g fish for U.S.$1.70/kg, and 1200 g and larger fish for U.S.$1.80/kg in August 2003 (http://groups.yahoo.com/groups/hilsa/message/25). The catch in Khuzestan province in 2000 was 2688 t (Ghafleh Marammazi et al., 2004) and  in 2006 was 4989.83 t (about 15% of Khuzestan's total commercial fish landing) (Roomiani and Jamili, 2011). The catch in Khuzestan Province in 2008 was 4645 t (Hashemi et al., 2010).

These fish are caught with traps, weirs, gill nets and other devices in rivers on the spawning migration. They are excellent eating until spawning occurs after which they lose their flavour. However this species has been implicated in clupeotoxic poisoning. Hindi et al. (1996a) give the chemical composition of flesh of this species as 66.41% moisture, 12.12% fat, 18.72% protein and 1.98% ash, indicating a valuable food fish characterised as fatty. Hindi et al. (1996b) give chemical indices for assessing fish freshness according to the month of capture and marketing (pH 6.06, total volatile nitrogen bases 15.32 mgN/100g fish, thiobarbituric acid 1.35 mg, and free fatty acids 1.33%). Salari and Sadough (2009) compared heavy metal (Cd, Pb, Cu, Co, Ni) content in muscle, liver and gill tissues of fish from the Karun River and found levels less than those considered dangerous in Iran.

In Pakistan, the Indus River fishermen number between 8,000 and 9,000. Jafri (1994) reviews the Indus fishery which had yields up to 2694 mt. It is the most important Indo-Pacific shad species. The failure of the Indus River fishery in 2003 through drought resulted in Iranian fish being flown to Pakistan for marketing there at rupees150-400 per piece (www.jang-group.com/thenews/feb2003-daily/18-02-2003/business/b2.htm, downloaded 18 February 2003).

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use as food, in aquaculture and in textbooks.

Conservation

Hussain, Jabir and Yousif (in litt., 1995) report a decline in catches over the previous two decades in the Shatt al Arab. Al-Nasiri and Al-Mukhtar (1988a; 1988b) mention that fish enter the polluted Ashar Canal, a side tributary of the Shatt al Arab, during high tide when waters are diluted. A low tide in October resulted in severe oxygen depletion and fish suffocated. Das et al. (1977) found samples from the Ashar fish market in Basrah to be contaminated with hydrocarbons, emitting a kerosene smell and being unfit for human consumption. Al-Saad (1990) found petroleum hydrocarbon residues to be high in Khawr az Zubayr fish at 40.6 μg/g as this species is one that accumulates fat. Evidently, overfishing and pollution are major factors in the conservation of this species, to which must be added variations in freshwater flow and quality from the marshes and Tigris-Euphrates through human processes.

Further work

The migratory habits and ecological requirements of this food fish need to be examined in more detail for Iranian waters.

Sources

Some aspects of the biology of this species were based on Pillay and Rosa (1963) and Al-Hassan (1993) writing mostly on Indian and Pakistani populations. Specimens on markets in Ahvaz, Khuzestan examined.

Iranian material: CMNFI 1991-0153, 1, 243.3 mm standard length, Khuzestan, Zohreh River (no other locality data).

Comparative material: BM(NH) 1875.1.14:11-13, 3, 118.8-135.8 mm standard length, Iraq, Tigris River (no other locality data); BM(NH) 1920.3.3:178-182, 6, 103.3-132.4 mm standard length, Iraq, Basra (30º30'N, 47º47'E); BM(NH) 1989.1.13:1-3, 3, 53.9-59.9 mm standard length, Iraq, Khawr az Zubayr (no other locality data); BM(NH) 1989.1.13:4-5, 2, 66.6-69.8 mm standard length, Iraq, Khawr az Zubayr (no other locality data).

Chanidae
 

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This family contains only one species. It is characterised by a compressed and oblong body; small, toothless and terminal mouth; non-protractile upper jaw; lateral pouches on the posterior part of the branchial chamber forming an epibranchial organ; presence of intermuscular bones; first 3 vertebrae specialised in structure; presence of an alarm substance; swimbladder present; gill membranes united and not attached to the isthmus; caudal fin deeply forked; the dorsal and pelvic fins opposite and small to moderate; cycloid scales; and a distinct lateral line.

Genus Chanos
Lacepède, 1803

Characters for the only species in the genus and family are summarised under the family.

Chanos chanos
(Forsskål, 1775)

Common names

khameh mahi (= literally cream fish but probably meant as milkfish).

[sheem in Arabic; milkfish, salmon herring, giant herring].

Systematics

No major synonyms. Mugil chanos was originally described from Jidda on the Red Sea.

Key characters

The milkfish resembles members of the family Clupeidae but is distinguished by a low number of branchiostegal rays (4 as opposed to 6-7), the presence of a lateral line, and the absence of scutes along the belly.

Morphology

The mouth is small and lacks teeth. There is a notch on the upper jaw in the mid-line into which a lower jaw protuberance fits. The large eyes have an adipose eyelid. The intestine is very long with many folds. The lower part of the oesophagus has a "gizzard', an area with longitudinal folds.

Lateral line scales 70-92, with 3-11 on the tail fin this latter count varying widely between authors. Total dorsal fin rays 13-17 including usually 2-6 unbranched rays, branched dorsal fin rays 9-14, usually 11-12; anal fin unbranched rays 2-3, branched rays 6-10, usually 7-8; pectoral rays 14-18, usually 15-16 and pelvic branched rays 10-12. Gill rakers and pyloric caeca very numerous. Gut long and complexly coiled. Vertebrae 42-46. Chromosome number 2n=32, low compared to other primitive teleosts (Klinkhardt et al., 1995).

Meristic values for Iranian specimens are:- lateral line scales 70(1), 71(2), 72(1), 74(2), 75(1), 76(2) or 79(2); unbranched dorsal fin rays 4(11); branched dorsal fin rays 11(7) or 12(4); unbranched anal fin rays 3(11); and branched anal fin rays 7(10) or 8(1); pelvic fin branched rays 9(1), 10(4) or 11(6); vertebrae 42(2) or 43(9).

Sexual dimorphism

None reported.

Colour

The overall colour is silvery with bluish or olive tints dorsally. The flank may have golden tints. The top of the head is yellowish-olive, the sides and ventral surface bright silvery to whitish. The snout is a light brown. The dorsal and caudal fins are colourless to grey, yellowish or brown with dark margins. The anal and pelvic fins are white, the anal with a dark margin. The peritoneum is black. The iris is silvery.

Size

Up to 1.85 m and 18.6 kg, although Bagarinao (1994) cites 1.5 m and 14 kg as more reasonable.

Distribution

Found from the African coast, Red Sea and Persian Gulf to the southern Pacific coast of the U.S.A. and to Peru, and north to Japan and south to Australia. Reported to penetrate 100 km up the Shatt al Arab (McKinnon and Vine, 1992).

In Iran, this species is reported from the Baghu River, Hormozgan near Bandar-e Abbas in 1976, found dead in brackish water about 14 km by river from the sea (Coad, 1981b). Also reported from the lower Mand River in Bushehr Province by M. Rabbaniha (pers. comm., 1995). Abdoli (2000) illustrates it as entering several rivers around the Straits of Hormuz including the Minab and Kul rivers. Salehi (1999) records this species from estuaries in Hormozgan and Makran including the "Khoor-Chel", Shur, "Takhtenze", Tiab, Gask, Heylaru, "Gorginee" and Gabrik rivers.

Zoogeography

The milkfish is unusual in being one of the few Indo-West Pacific fish species found also in the eastern Pacific, although the mechanism of dispersal is uncertain. Larvae are probably incapable of crossing the 6500 km wide East Pacific Barrier but adults could do so as they can cruise at 2 km/h. However adults have not been caught on the high seas.

Habitat

Usually found in littoral waters of the ocean, rarely entering rivers, but it has been acclimatised to freshwater ponds. It becomes sluggish at temperatures below 15°C and dies at about 12°C but can survive temperatures of 41°C. It seems to prefer waters warmer than about 20°C. The Iranian specimens (Coad, 1981b) were caught at 16°C in a shallow, muddy, backwater and probably died through exposure to colder temperatures in the main river. Persian Gulf temperatures can fall below 15°C in winter.

It is resistant to salinity changes and can survive in fresh and hypersaline water (0-158 p.p.t.) as well as waters low in oxygen. The dorsal and upper caudal fin lobes may cut the surface of the water and milkfish are often mistaken for sharks. Milkfish are solitary or found in small schools.

Age and growth

Life span is at least 12 years based on pond specimens but large adults in the sea undoubtedly live much longer. Growth in the sea is poorly known and most data refer to pond-reared populations. Fish at sea are 20-43 cm long after 1 year, a growth rate considerably less than in ponds. Maturity is reached at 3-5 years in the wild but as long as 8-10 years in ponds.

Food

Young and adult milkfish will feed on surface scum, taking it in with a smacking noise. Benthic and epiphytic organisms are also taken in ponds with the body inclined at an angle of about 30°. Cyanobacteria, benthic diatoms, foraminiferans, filamentous green algae, detritus, clams, snails, worms, some crustaceans, and fish eggs and larvae are taken in from the sea floor. Other reports have this species feeding on plankton but this may be detritus fallen to the sea floor from surface layers

Reproduction

There are two spawning seasons in India, March to June and September to November but individuals may only spawn once a year. Spawning occurs in clear water of sheltered, sandy bays within about 6 km of shore, away from river mouths, i.e. saline water, and is probably triggered by rising temperatures in spring (25-30°C) and high tides. These conditions give water deep enough to avoid eggs being eaten by corals and other benthic organisms yet close enough to shore for larvae to reach their preferred inshore habitats. Eggs are fertilised while floating in surface waters. Fecundity is reported to reach 7.3 million eggs but this is not based on large fish and fecundity could be considerably more. Egg diameters are up to 1.25 mm when fertilised. Ribbon-like larvae migrate to coastal areas, metamorphose and may enter creeks and estuaries. About 4 weeks later the young leave coastal waters and spread out in waters where there are adequate supplies of benthic and planktonic food. Some may remain in estuarine areas for 4 years before returning to the sea. In February-March, and again in October in some populations, the adult migration to inshore waters for spawning takes place. Adults leave coastal waters after spawning.

Parasites and predators

Young are eaten by a wide variety of predators in nearshore waters as indicated by the high egg production and adult survival rate.

Economic importance

This is the most important tropical marine fish used in aquaculture with a history dating back 500 years. Milkfish are raised in brackish or freshwater ponds throughout Asia, based on larvae captured in shore waters. 1.35 billion larvae were caught in the Philippines in 1974. The Philippines, Indonesia and Taiwan produce about 330,000 tonnes of milkfish per year. Their wide tolerance of environmental variables and herbivorous diet (rice bran and pelleted foods in captivity) combined with fast growth make them a success in aquaculture. The fish are marketed at 200-300 g. Adults form part of fisheries aimed at other species.

McKinnon and Vine (1992) report that this species is sold in the fish market at Basrah, Iraq. In the Persian Gulf they can be caught by set nets, gill nets, traps and hook-and-line (Carpenter et al., 1997).

Milkfish have been cultured in concrete ponds at Tiyab, Hormozgan from March to October. Fry were caught in local estuaries such as the Shur River 30 km east of Bandar Abbas. They were fed commercial carp food pellets and after 7 months weighed 450 g, or on poor protein food 130 g after 11 months with no growth in the cold season (Annual Report, 1995-1996, Iranian Fisheries Research and Training Organization, Tehran, p. 40, 1997; Forughi-e-Fard and Gharibnia, 1998; Fourooghi-e-Fard, 2000). They have also been cultured at Tiyab with Indian white shrimp (Penaeus indicus) and mean weight of shrimps was found to be higher than in monoculture (Foroughifard, 2001).

Milkfish have been reported as being ciguatoxic (intermittently poisonous through feeding on toxic food) (Bagnis et al., 1970).

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use as food and as bait, in aquaculture and in textbooks.

Conservation

None required as this species is probably an accidental visitor to Iranian fresh waters.

Further work

The culture of this species in southern Iran could be developed further.

Sources

Biology was based on Schuster (1960) and Bagarinao (1994).

Iranian material: CMNFI 1979-0142, 11, 70.5-98.5 mm standard length, Hormozgan, Baghu River (27º17'N, 56º28'E) (Coad, 1981b).

Cyprinidae

This family contains by far the most species in the Iranian freshwater ichthyofauna and is divided into two files Abramis to Cyprinus (here), and Garra to Vimba (see both in Contents).

The carp or minnow family is one of the most widespread and speciose families of fishes in the world, certainly the most speciose in fresh water and possibly the largest family of vertebrates (the Gobiidae may be the first). The family is found in North America, Eurasia and Africa. Other common names in English for species include barbels, breams, roaches, snow trouts, bitterlings, shiners, daces, chubs, barbs, "sharks", among many others. There are about 220 genera and over 2420 species (Nelson, 2006), about 8.5% of the world's fishes. In Iran, the family is represented by about ?32 native genera (interpretations of genera differ between authors) and at least ?73 species (with more to be described) found in all the major drainage basins. 

The minnow or carp family is comprised of small to very large fishes (1 cm and up to 3 m, with some of the largest members in Iran) characterised by throat or pharyngeal teeth in 1-3 rows, with a maximum of 8 teeth in a row, tooth counts and form are often characteristic of the genus or species, no jaw teeth, body form various from fusiform to compressed, lips are usually thin and not sucker-like (but can show hypertrophy), the upper jaw is bordered by the premaxillae bones and usually protrusible, barbels are absent or present in 1-3 pairs (not more than 2 pairs in Iranian species), body covered in cycloid scales, in some species easily lost, while the head is scaleless, no adipose fin, the anterior 4 vertebrae are modified for conduction of sound from the air bladder to the ear and are known as the Weberian apparatus, pelvic fins are abdominal in position, no pyloric caeca, air bladder usually present and well-developed, connected to the gut by a duct, and not enclosed in a bony capsule, no true stomach, branchiostegal rays always 3 in number, no true spines in the fins although in some the last unbranched dorsal fin ray (at the front of the fin) may be thickened and spine-like and in Cyprinus and Carassius the last unbranched anal ray is also thickened. The primitive chromosome number is 2n=50 but polyploidy is common and seen in Cyprinus, Carassius and in the schizothoracines. Collares-Pereira (1994) argues that the polyploid condition (e.g. 2n=100) is primitive or plesiomorphic.

There are 2-4 unbranched rays (including rudimentary ones) in the dorsal and anal fins followed by the more numerous branched rays (the last two branched rays are counted as one). The first pectoral and the first pelvic fin ray are unbranched and not included in counts. Pharyngeal teeth lie on a modified, fifth gill arch which can be seen or probed behind the shoulder girdle, just inside the gill opening. The arch has to be removed with dissecting equipment to count the teeth. Tooth counts are presented as a formula such as 2,5-4,1 which indicates 2 teeth in the outer left row and 4 on the inner right row. Teeth may be lost from major or minor rows so variant formulae are given after the principal one. A horny pad on the underside of the basioccipital bone of the skull is used to masticate the food against. Tooth form varies with the food - molar-shaped teeth are used to crush molluscs, flat but grooved surfaces for grinding plant food and sharp edged teeth for slicing various invertebrate foods.

Two subfamilies, the Alburninae and Leuciscinae, are paraphyletic but together seem to form a monophyletic group with a radiation about 20 million years ago, based on allozyme, cytochrome b, 16S rDNA and mitochondrial control region data from European cyprinids (Hänfling and Brandl, 2000; Gilles et al., 2001). These two subfamilies contain many Iranian genera. Zardoya and Doadrio (1999) analysed the cytochrome b nucleotide sequence of a variety of cyprinids, mostly European, and found support for two subfamilies Cyprininae (including barbins) and Leuciscinae (including cultrins, tincins, gobionins, phoxinins and alburnins + leuciscins). The origin of cyprinids is estimated at 38.9MYA and the separation of Cyprininae and Leuciscinae at 27.7MYA. They also found the phylogenetic utility of barbel possession to be limited as they were acquired independently in the two subfamilies. The number of rows of pharyngeal teeth were a more reliable phylogenetic marker, at least at the generic level. Perea et al. (2010) using mitochondrial and nuclear DNA give details of major cladogenetic events in the leuciscin lineages in the circum-Mediterranean, involving genera and species found in Iran.

Chen and Mayden (2009) investigated the major clades of cyprinids using a multiple nuclear gene approach and tentatively recommended elevation of certain subfamilies to family. This is not in general use at this writing and the Cyprinidae is retained as a single family here.

Durand et al. (2002) using cytochrome b DNA of Cyprinidae conclude that the the Middle East is an important interchange area for this freshwater ichthyofauna rather than a centre of speciation. The Middle East leuciscine cyprinids have Europe as an important Palearctic influence consistent with the Lago Mare dispersion while the the cyprinine cyprinids show three highly divergent lineages, namely one shared with the Euro-Mediterranean area (Barbus/Luciobarbus), a relict of the Lago Mare dispersion, one shared with Africa (Carasobarbus/Varicorhinus subgenus) and one with Asia (Garra). The Lago Mare dispersion occurred during a salinity crisis in the Mediterranean Sea 5.5 MY ago in the Late Miocene when freshwater fish were able to disperse through oligohaline or fresh water in the Paratethys Sea to reach the Middle East (Bianco, 1990). Some data of Durand et al. (2002) conflict with this scenario - the Carasobarbus clade that includes Barbus (= Tor) grypus shows a separation divergence later than the salinity crisis in the Pliocene when no migration route was available. But note that some authors place Barbus grypus in the Indian genus Tor and that evidently more work needs to be done on its relationships and on those of other species that have no evident Euro-Mediterranean relatives, but whose origins may well lie in the Oriental Region.

Other Middle Eastern cyprinid genera are regarded by Durand et al. (2002) as relicts of older colonization waves and show an eastern influence consistent with an Asian origin of the family Cyprinidae. Cyprinion has no sister species in the Euro-Mediterranean area and has been isolated in the Middle East since before the salinity crisis, 7.8-8.8 MY ago. Cyprinion may have entered the Middle East during the colonization event that isolated the genera Barbus sensu lato and Schizothorax in the European and Asian basins respectively. The divergence of these species is similar in time to the radiation of the Leuciscinae supposedly centred in Siberia based on fossil records. Siberia was probably an important dispersion centre for both Leucicinae and Cyprininae at that time. Otero (2001) describes a ?Barbus sp. (sic) from the Lower Miocene of Saudi Arabia showing an early date for the entry of cyprinids to the Afro-Arabian Plate.

Some species may enter brackish water but the family is primarily a freshwater one. Carps have extremely sensitive hearing via the Weberian apparatus and this is thought to account for their success. Carps produce an "alarm substance" when injured. This chemical stimulates other carps to flee and hide, another useful adaptation. Carps are remarkable for changes they undergo during the spawning season. Some fish, which are usually silvery, develop bright reds and yellows. Nuptial, pearl or breeding tubercles develop on the head, scales and fin rays often in distinct patterns, and there are swellings of the head or fin rays in some species. These changes are most apparent in males. Tubercles and swollen rays are used to clasp females during the spawning act. Generally males have longer pectoral fins than females. Tubercles are also used to fight other males and defend and clean nests. Colour attracts females for mating. Nest building males are larger than females, the reverse of the situation in most fishes where egg-bearing females are the largest. Not all species build nests and some simply broadcast eggs over weed, gravel or sand. Fractional spawning is common in carps. This is a prolonged spawning season which ensures no single batch of eggs is lost to unfavourable, temporary environmental changes such as floods. Carps are mostly omnivores, feeding on small crustaceans, insects and some minute plants but some specialise in eating large plants, or other fishes. Diet is reflected in pharyngeal tooth shape as mentioned above. Gut length is important too. A long intestine indicates a reliance on plant material which takes longer to digest. A simple, s-shaped gut is found in insectivorous fish. A black peritoneum is thought to protect gut bacteria from damaging light. The bacteria aid in breaking down the strong cell walls of plants. Size and shape of the mouth are also indicative of diet. Carps are found in many diverse habitats from swift, cold streams to warm bogs. These are schooling fishes, especially when young.

Carps play an important role in fresh waters as food for other fishes and some species are commercially important as bait fish, as sport fish or as food in Asian countries. Raising minnows as bait and as forage fish for sport fish is a big business in the U.S.A. They are an important element in the commercial aquarium trade and certain species are used in experimental studies by scientists. Cyprinids were also important in the past, sacred fish ponds being reported from Mesopotamia in 3000 B.C., and in Iran today cyprinids associated with mosques and shrines are "sacred". A general review of Eurasian cyprinids is given by Bănărescu and Coad (1991).

Carp family members are particularly important in Iran in aquaculture. The "Chinese carps" (Cyprinus carpio or common carp, also native to Iran, Ctenopharyngodon idella or grass carp, Hypophthalmichthys molitrix or silver carp, and to a lesser extent Hypophthalmichthys nobilis or bighead carp) are the main species used in warmwater culture in almost all the provinces of Iran. Common, grass and silver carps are processed into fish fingers in Iran (Iranian Fisheries Research Organization Newsletter, 25:1, 2000). Danesh-e-Khoshashi (1998) describes facilities and methods used for spawning Chinese carps in Gilan Province. The production of Chinese carp fingerlings has been relinquished to the private sector in Iran. The silver carp catch increased from none in 1989 to 24,720 t in 1994 (Food and Agriculture Organization, Fisheries Department, 1996). Chinese carp production peaked in 2006 at more than 77000 t according to Salehi (2009) who also reviews carp farming costs. Chinese carp fingerling production was 22.7 million in 1996 (Bartley and Rana, 1998a). Stakei (1999) studied nutrients, BOD and COD in manured polyculture ponds with Chinese carps. A review of world cyprinid culture, with special reference to the Chinese carps, is given by Billard (1995).

Rana and Bartley (1998a) give details of carp aquaculture in Iran. They note that silver carp production increased 11% per year between 1991 and 1996 and bighead carp 7%. Most carp production occurs in the provinces of Gilan, Mazandaran and Khuzestan and is a private sector enterprise. Carp broodstock is selected based on head size, colour and gill structure (surface and shape). Adults are replaced after 3-4 years. Circular concrete tanks are used for spawning and have egg collecting and incubation devices which reduce handling to the minimum. The young carp are grown to market size in ponds or complex fish farms. In 1994, there were 2583 registered farms with a water surface area of about 8000 ha. Organic and inorganic fertilizers are used along with supplementary foods. Fertilizers include urea (135-1500 kg/ha/yr), ammonium phosphate (80-575 kg/ha/yr) and manure (3-10 tonnes/ha/yr). Supplementary diets include a variety of grains (100-6000 kg/ha/yr) or, for intensive monoculture of common carp, high protein pellets (30-40%). Fingerlings are stocked in March-April at a density of 2000-6000 per hectare and sold between November and February. Production is 1.6-5.5 tonnes/ha. Cultivated carps are susceptible to fungal infections as detailed by Ebrahimzadeh et al. (2000) for the Safid River Fish Farm Centre where 31 species of fungi were isolated and Firouzbakhsh et al. (2005) where 39 fungal species were identified from gill lesions in common, silver and grass carp on five fish farms in Mazandaran.

Rice fields in Iran are now being considered for fish culture. Experimental production of 300-500 kg per hectare of "carp seed" (presumably young fish) an 750-1000 kg of fish and ducks in the autumn after the paddy is harvested (Iranian Fisheries Research Organization Newsletter, 22:2, 2000). In the early 1970s intensive carp culture yielded only half the profits of rice culture (Carl Bond archives, Oregon State University, Corvallis).

Experiments in the Caspian region for artificial propagation of Aspius aspius and Barbus (= Luciobarbus) brachycephalus to enhance stocks and for farming Rutilus frisii and Abramis brama using mono- and polyculture along with Chinese carps have been carried out (Iranian Fisheries Research and Training Organization Annual Report, 1992-93; Annual Bulletin 1993-94, Iranian Fisheries Research and Training Organization, Tehran, p. 77-78, 1995). There are about 3000 fish farms producing over 98% of the cultured fish in the country. Yearly production of all cultured fish has increased from 4753 tonnes in 1985 to 45,134 t in 1990. Production of carps in government hatcheries has risen as follows: 2.19 million fingerlings in 1983, 5.04 million in 1984, 12.84 million in 1985, 20.83 million in 1986, 19.05 million in 1987, 50.00 million in 1988, 50.80 million in 1989, 97.70 million in 1990, 58.00 million in 1991, and 50.00 million in 1992. In addition private sector production probably equals these figures (Emadi, 1993a). Polyculture of common, bighead and silver carp has been tried in Iran (Kamaly, 1991). Fish were stocked in four 200 sq m ponds at three densities in polyculture (2700, 3750 and 4750 by species) and at one density in monoculture (9500) fish per hectare. Bighead and silver carp attained a mean weight of 526 and 498 g in polyculture and common carp averaged 343, 190 and 100 g in the same culture but only 13.6 g in monoculture. The growth rate in summer averaged 94.4, 93.7 and 76.1% for silver, bighead and common carp in polyculture and 71.9% for common carp in monoculture. Pen culture in the Caspian Sea has been investigated for Cyprinus carpio and the various Chinese carps (Iranian Fisheries Research and Training Organization Annual Report, Tehran, 1992-93). Semi-artificial breeding of grass, silver and bighead carps has been carried out in Iran (Iranian Fisheries Research and Training Organization Newsletter, 6:3-4, 1994; Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 39, 1996). Hormone injections were used to induce breeding of fish held in a round trough for spawning with a rectangular egg collection trough and a round egg hatching trough. Spawning occurred within 6.5-12.5 hours of injection. The percentage of hatched larvae in this semi-artificial method was higher than a control artificial method where eggs are kept in incubators. The increase was 6% for grass carp, 33.72% for silver carp and 16.7% for bighead. Active larvae increased from 180,000 to 450-500,000 for grass carp, from 157,000 to 400-450,000 for silver carp and from 680,000 to 970,000 for bighead carp. Additionally female breeder mortality was 3.37% less for grass carp and 45.19% less for silver carp.

Many carp species can be caught on hook and line by various angling techniques but outside the larger rivers of Khuzestan and the Caspian shore this hobby is not much pursued. Even small species and specimens can give some sport on light tackle such as worm baited hooks including Luciobarbus barbulus, Carasobarbus luteus, Alburnus mossulensis, Cyprinion macrostomum and Garra rufa among others.

Fingerlings of Labeo rohita, an Indian carp, were imported to Gilan in Iran in 2004 to enrich the diversity of cultured fish and increase protein production. There is always the potential for escapes and establishment of this exotic.
 


Genus Abramis
Cuvier, 1816 
 

The bream genus comprises 4 species found in Europe, Asia Minor and the Caspian and Aral Sea basins. There are 2 species in Iran but see also Blicca and Vimba.

The genus is characterised by a strongly compressed and deep body, a scaleless keel between the vent and pelvic fins, a scaleless groove on the back in front of the dorsal fin but not behind the fin, pharyngeal teeth in 1 row, compressed and with a groove on the grinding surface, dorsal fin short and spineless, anal fin long to very long, and lateral line decurved.

Durand et al. (2002) studying cytochrome b data concluded that this genus is not monophyletic since A. ballerus and A. sapa are placed basal to a group of species including A. brama, Blicca bjoerkna, Vimba species, Acanthalburnus microlepis and Acanthobrama.

.Abramis brama
(Linnaeus, 1758)

Abramis brama, aqaurium fish. Wikimedia Commons.

Common names

سيم (sim or seam = silver), ماهي سيم (= mahi-ye sim, meaning silver fish).

[capag, chakag, chapakh or chipakh, all in Azerbaijan; gundogar tarany (topi) in Turkmenian; vostochnyi leshch or Oriental bream in Russian; common, bronze, eastern or carp bream].

Systematics

Cyprinus Brama was originally described from Sweden.

Abramis brama orientalis Berg, 1949 is reported for the Caspian and Aral Sea basins but Koshara and Izyumov (1991) restricted this subspecies to the Aral Sea with the type subspecies in the Caspian Sea basin. They did not examine any Iranian material. Kozhara and Mironovskiy (1988) using numbers of pores in the seismosensory canals for samples taken over a wide range of this species identified 8 population groups but did not recognise subspecies. Some earlier works also indicate that no subspecies exist (see Reshetnikov et al., 1997).

Caspian material reportedly has more gill rakers, fewer vertebrae and fewer scales than the type subspecies from the Baltic Sea basin (Berg, 1948-1949) but further study over the whole range of the species is needed to clarify the situation, analyzing for clines. The Iranian populations are referred to the type subspecies for the moment. The type locality of this subspecies is the Aral Sea at Muinak and Lake Yaskhan in the Uzboi.

Khara et al. (2007; 2007) compared fish from the Anzali wetland and the Caspian Sea, and the Caspian Sea and Aras Dam, both meristically and morphometrically. Significant differences were noted in particular for morphometric characters in the former comparison and morphometrically and meristically in the second. These differences were attributed to differing habitats and environmental conditions. Ghasemi et al. (2007) used 5 microsatellite loci in comparing Iranian and Azeri bream and found Iranian stocks have reduced genetic variability attributed to inbreeding and genetic drift. Khara et al. (2009) compared fish from the Anzali Wetland the southern coast of the Caspian Sea in Iran and the southwest coast in Azerbaijan using mtDNA. The greatest genetic diversity was found in Azerbaijan which was significantly different from the Iranian samples, which were not themselves significantly different.

Abramis brama bergi Grib and Vernidub, 1935 (preoccupied by Abramis sapa bergi Belyaeff, 1930 according to Eschmeyer et al. (1996)) was originally described from the Aral Sea at Muinak and is also found in the Uzboi Valley of Turkmenistan, north of the Iranian border (Berg, 1948-1949). It was replaced by Abramis brama orientalis.

A syntype of Cyprinus brama is in the Natural History Museum, London as a skin under BM(NH) 1853.11.12:147 (Eschmeyer et al., 1996).

Artificial hybrids with Rutilus frisii kutum and Rutilus rutilus (may involve R. caspicus) have been bred in Iran (Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 39-40, 1996).

Key characters

The scaleless keel on the belly, deep body, high number of branched rays in the anal fin (22-30), modally 9 branched dorsal fin rays, and uniserial pharyngeal teeth are key characters.

Morphology

The mouth is small but highly protrusible. There is a strong dorsal ridge anterior to the dorsal fin. Dorsal fin with 3 unbranched and 8-10, usually 9, branched rays, anal fin with 3 unbranched and 22-30 branched rays. Lateral line scales 48-60. The lateral line is moderately decurved. Scales are regularly arranged, sheathing the anal fin base. Scales have numerous fine circuli but only relatively few posterior and even fewer anterior radii. In a fish about 6 cm long there are as few as 8 total radii. The focus is almost central and the anterior scale margin is wavy. There is a pelvic axillary scale. The ventral keel between the pelvic fin bases and the anal fin is well-developed. Gill rakers number 18-30 and are short, reaching the raker below when appressed. They are strongly tuberculate on the inner surface. Vertebrae 38-47, usually 42-44 in the Caspian populations (lower counts in literature may not include 4 Weberian vertebrae). The chromosome number is 2n=50-52 (Klinkhardt et al., 1995). The chromosome number based on fish from the Iranian coast of the Caspian Sea is 2n = 50 with the number of arms NF = 82 and the karyotype being 8 pairs of metacentric, 8 pairs of submetacentric and 9 pairs of acrocentric chromosomes (Nahavandi et al., 2001).

Pharyngeal tooth formula modally 5-5, with variants of 6-5 (2.2-4.8%), 5-4 (2.2-4.4%) and 4-5 (8.6%) for collections from the Caspian and Aral seas basins in former Soviet waters (Vasil'yeva and Ustarbekov, 1991). Other variants are summarised in Tadajewska (1998). Teeth bear a small hook at the tip in the main row and have long, narrow and flat crowns. In young fish, the hook is more pronounced and the crown has a few tubercles or a series of serrations. The gut is s-shaped with a small anterior loop.

Khar et al. (2007) compared this species from the Caspian Sea and the Anzali wetland and found significant morphometric, but not meristic, differences, attributing this to habitat conditions.

Meristic values for Iranian specimens are:- dorsal fin branched rays 9(12) or 10(1); anal fin branched rays 24(3), 25(3), 26(1), 27(4) or 28(2); pectoral fin branched rays 16(7) or 17(6); pelvic fin branched rays 8(13); lateral line scales 49(2), 50(2), 51(3), 52(3), 54(1) or 55(2); total gill rakers 23(3), 24(2), 25(6), 26(1), or 27(1); total vertebrae 44(12) or 45(1); and pharyngeal teeth 5-5(12).

Sexual dimorphism

Males bear tubercles on the head, body and fins. Scale tubercles appear singly, in pairs or occasionally as 3 per scale. There is some evidence of differences in gill raker counts between the sexes but sometimes the males have higher mean counts and sometimes the females. Abdurakhmanov (1962) reports eye diameter, greatest body depth and predorsal distance to be greater in females and dorsal fin base length, pectoral and pelvic fin lengths and interorbital width to be greater in males from Azerbaijan.

Colour

In Dagestan, the resident form is darker in colour than the semi-anadromous form (Shikhshabekov, 1969). Overall colour is silvery. The iris is silvery with a little grey pigment on the upper part. The dorsal and caudal fins are pale grey, almost transparent, to a greyish-blue, the pectoral fins may be grey or colourless, and pelvic fins are colourless. All fins except the pectorals have black tips. Large fish are a dark olive-green on the back and bronze on the flanks and old fish may have all fins black. The peritoneum is silvery to light brown in preserved fish.

Size

Attains 90.0 cm total length and 11.55 kg, possibly 100.0 cm and 16.4 kg.

Distribution

Found from the British Isles across Europe north of the Pyrenees and Alps eastwards to the Black, Caspian and Aral sea basins although not in western Transcaucasia. In Iran it is found from the Astara to the Atrak rivers in the whole Caspian Sea basin (Kozhin, 1957) including the Anzali Mordab, its outlets and tributaries and the Siah-Keshim Protected Region (Holčík and Oláh, 1992; Riazi, 1996; Kiabi et al., 1999), the Safid River (Abbasi et al., 1999), Gorgan Bay (Derzhavin, 1934), and freshened areas of the Caspian Sea. It s also found in the Aras Dam (Khara et al. 2007).

This species is also recorded from the Karakum Canal and Kopetdag Reservoir in Turkmenistan (Shakirova and Sukhanova, 1994; Sal'nikov, 1995) and may eventually reach Iranian waters in the Tedzhen (= Hari) River basin where it has been reported by Aliev et al. (1988).

    

Zoogeography

This species is part of a northern European and northern Southwest Asian fauna whose zoogeographical history has not been researched. It origins may lie in a Danubian or Sarmatian fauna.

Habitat

The bream prefers still water and is low in numbers even in rivers with weak current. Abundant littoral vegetation and a very muddy bottom are favoured in lakes for reproduction and feeding respectively. It retreats to deeper water in winter, forming schools numbering in the many thousands, packed densely together (Muus and Dahlstrøm, 1999). It is more numerous in the Anzali Mordab along the Caspian coast of Iran (Jolodar and Abdoli, 2004).

This species can tolerate high temperatures of 33-34°C in southern areas like Iran for a time but above 28°C growth rate decreases. Adults can live in a salinity of 12.9‰, perhaps 14‰, and eggs may be fertilised at a salinity of 10.2‰. However preferred levels are 2-4‰. Salinity and water level changes have significant effects on abundance in this species. Population densities vary markedly in both fresh and brackish water populations.

Bream living in the Caspian Sea basin are semi-migratory. They feed in the brackish sea but spawn and winter in the lower reaches and deltas of large rivers. A spring migration up rivers begins with ice melt or warmer temperatures in the sea and after spawning the fish return to disperse and feed in the sea. In the fall the fish migrate into the deeper parts of river deltas. In Russian parts of the Caspian they are found at depths not exceeding 4-5 m but Knipovich (1921) reports them at 14.6-16.5 m, possibly deeper, in the Iranian Caspian Sea.

There were spring and winter migrants in the southwestern Caspian including the Anzali Mordab (A. M. Shukolyukov in Berg, 1948-1949). The spring bream had a longer snout, deeper head, lower body, lower dorsal and anal fins, and more scales. The spring bream entered the Mordab for spawning only while the winter bream overwintered in bottom pools. Changing conditions in the mordab environment in the late 1980s and the 1990s may have altered this migration. Riazi (1996) reports that this species migrates into the Siah-Keshim Protected Region of the Anzali Mordab.

Age and growth

The resident form in Dagestan is slightly inferior in length (2-3 cm), weight and age to the semi-anadromous form (Shikhshabekov, 1969). In Dagestan, the resident form becomes sexually mature at 3 years for females and 2 years for males at lengths of 23-26 cm and weights of 200-240 g while the semi-anadromous form matures at 4 years and a length of at least 25-28 cm and a weight of 250-300 g. In Uzbekistan females mature at lengths ranging from 10.5 to 27 cm in different reservoirs, usually at age 3 (Kamilov, 1994). Maturity is attained at a younger age in southern waters generally in this species and this probably applies in Iran. The maturity range is 2-10 years with males often maturing a year earlier than females. Females predominate in the older age groups.

Maximum age exceeds 32 years although in southern waters the maximum age does not exceed 15 years. Semi-migratory bream of the Caspian Sea have a fast growth rate and a short life cycle, reaching 37.5 cm standard length by age 8.

Most fish examined by Razivi et al. (1972) from commercial catches in Iran were 3-6 years old, 25.6-39.8 cm long and weighed 249-950 g. Over a three year period there was a decline in average age. Young and immature fish formed most of the catch in 1998-1999 when one-year-old fish comprised 20.3% and two-year-old fish 37.3%. The average length, weight and age for 1998-1999 were 22.5 cm, 212.2 g and 2.4 years. The rate of recruitment was 4.6% in 1991 and 2.7% in 1992 (Saiad Borani, 2001). Abdolmalaki (2005b) studied Caspian Sea fish from Iran and found mean fork length, weight and age to be 21.7 cm, 191 g and 2.72 years, respectively. The length-weight relationship was W = 0.2312L2.9 and von Bertalanffy growth parameters were Lt = 45[1-exp-0.125(t  2.768)], and the instantaneous rate of total (Z), natural (M) and fishing mortality (F) were 0.92 year-1, 0.28 year-1 and 0.64 year-1, respectively. The exploitation rate (E) was 0.7. Biomass was calculated as 46.362 t and the maximum sustainable yield was 14.99 t.

Food

Young fish feed on zooplankton. Adults use a strong sucking power and a tube-like snout to feed on invertebrates and detritus in mud. This sucking action leaves evident "bream pits" in soft mud, depressions about 10 cm across. In the northern Caspian Sea food items include Cumacea, Corophiidae, the clams Adacna (69% by weight) and Monodacna, Tendipedidae (= Chironomidae), Polychaeta, Gammaridae, Mysidae, and Oligochaeta. When overcrowded or in turbid conditions, plankton may be eaten in addition to the normal foods (Muus and Dahlstrøm, 1999). Large specimens may feed on small fishes. A specimen from the Langarud, Gilan, 158.6 mm standard length, contained chironomids.

Reproduction

Bream enter the Kura River from December to February with a peak in January (Berg, 1959) and travel some distance upriver. These fish have an average length of 31.1 cm and an average weight of 633 g. Length and weight in Azerbaijan vary from 25.4 to 31.9 cm and 306 to 681 g. Bream enter the Anzali Mordab, the main spawning area in the southern Caspian, in the first half of March until the beginning of May. Males precede females on the spawning ground by about 3 days and males outnumber females by about 3 to 1. Spawning begins in the first half of April in shallow water and lasts until mid-May. Fecundity in Dagestan reservoirs reaches 191,000 eggs (Shikhshabekov, 1969), in Uzbekistan reservoirs 772,000 eggs (Kamilov, 1994) and a maximum elsewhere of 941,000 yellowish eggs is reported. Bream spawn repeatedly with different partners and although most bream spawn only once a year, multiple spawnings are known. Spawning occurs in masses over a period of 2-3 days triggered by temperatures of 12-13°C or above. The commonest spawning temperature for the species overall is 16-18°C. Spawning is most intensive at night in some populations while others show late morning and late afternoon peaks. There is much splashing of the water by their tails and the noise can be heard some distance away although the fish are easily scared into deeper water by any noise like human voices. Males probably defend territories, attracting females and scaring other males away. There can be up to 2.3 million eggs per sq m however, suggesting that many fish may spawn in the same area. Eggs are deposited in quiet water, most commonly at depths of 20-80 cm, and they adhere to aquatic plants or flooded land plants. Eggs are up to 1.9 mm in diameter.

Parasites and predators

Jalali and Molnár (1990a) record the monogenean Dactylogyrus zandti from this species in the Safid River. Sattari and Faramarzi (1997) record Caryophyllaeus fimbriceps from 28% of bream in the Anzali lagoon. Naem et al. (2002) found the following parasites on the gills of this species from the western branch of the Safid River, namely the monogenean trematodes Dactylogyrus zandti and D. wonderi. Masoumian et al. (2005) report the protozoan parasites Ichthyophthirius multifilis and Trichodina perforatafrom this species in the Aras Dam in West Azarbayjan. Jalali et al. (2005) summarise the occurrence of Gyrodactylus species in Iran and record G. elegans from fish in the Safid River. Sattari et al. (2004, 2005) survey this species in the Anzali wetland, recording Raphidascaris acus larvae. Pazooki et al. (2007) record various parasites from localities in West Azarbayjan Province, namely Ligula intestinalis, Digrama sp., Argulus foliaceus and Caryophyllaeus laticeps. Sattari et al. (2007) record the cestode Caryophyllaeus fimbriceps, the digenean Diplostomum spathaceum and the monogeneans Dactylogyrus extensus and Gyrodactylus sp. in this species in the Anzali wetland of the Caspian shore and also mention that the monogenean Diplozoon sp. is also known from this species in the Iranian Caspian Sea. Barzegar et al. (2008) record the digenean eye parasite Diplostomum spathaceum from this fish. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Lernaea sp. on this species.

The Caspian seal, Pusa caspica, is a predator on this species (Krylov, 1984). Various predatory fishes take bream including Huso huso, Perca fluviatilis, Sander lucioperca, Aspius aspius and Silurus glanis but this is comparatively rare especially when bream exceed 20 cm in length. Birds such as grebes, herons, divers and cormorants are also predators.

Economic importance

This species is an important food fish being both tasty and of large size. In addition it can live out of water for some time and thus remain fresh while being transported to market.

Nevraev (1929) gives catches for various fishing regions in Iran in the early twentieth century. For the Anzali region from 1901-1902 to 1913-1914 the catch was 2283 to 419,117 individuals, for the Safid River region from 1908-1909 to 1917-1918 the catch was 17,195 to 474,200 individuals (rising steadily but falling in 1917-1918) with no fish reported in the years 1899-1900 to 1907-1908 and in 1918-1919, and in Astrabad (= Gorgan) region from 1900-1901 to 1912-1913 the catch was 20,600 to 1,381,500 individuals with no clear trend, the catches varying markedly from year to year. The commercial catch in Iran from 1956/1957 to 1961/1962 varied from 0 to 158 kg (Vladykov, 1964), from 1965/66 to 1968/69 varied from 0 to 29 tonnes (Andersskog, 1970) and from 1963 to 1967 from 0.5 to 16.0 tonnes (with no reported catch in the first 3 years)(RaLonde and Walczak, 1970b). The catch in the Bandar-e Anzali region from 1933/34 to 1961/62 varied between only 2 kg and over 1394 t with some years reporting no catches. Holčík and Oláh (1992) report a catch of 34 kg in the Anzali Mordab for 1990 and for the period 1932-1964 catches ranged from none to 1133.5 tonnes annually. The total catch of the Northern Shilot (Fisheries Company) from 1965/66 to 1968/69 varied between 13 and 74 t (RaLonde and Walczak, 1972). There are obviously wide variations in annual catches and/or in reporting statistics. The general trend is one of decline in catches with large fish being caught and the average stock size being lowered, resulting in a decreased spawning success. This species has a deep body and immature fish are easily caught. The catch in the Anzali Mordab was important until the end of the 1940s but had virtually disappeared by the 1980s (Petr, 1987). Abdolmalaki (2005b) gives a total catch of 17 t for the 2000-2001 fishing season, only 0.1% of the commercial catch in Iranian coastal waters of the Caspian Sea. In contrast, the total catch for Iranian waters was estimated at 26.3 tons of which 15.4 tons was from beach seines; most fish were immature and undersized (Abdolmalaki, 2006a).

In former Soviet waters of the Caspian Sea, the age composition in commercial catches was 2-10 years, with the great majority being 3-5 years old. Trawls, seines, pound nets and gill nets are used in the northern Caspian Sea to catch the bream with 60-70% being taken in spring. Spawning and breeding farms were established in the former Soviet Union to rear young fish. Catches in the Volga-Caspian and Ural regions has been as high as 344,900 centners, prior to 1930, and in the Aral Sea in 1931 the catch was 115,200 centners.

Mono- and poly-culture of this species has been carried out in Iran (Annual Bulletin 1993-94, Iranian Fisheries Research and Training Organization, Tehran, p. 77-78, 1995). Polyculture comprised 70% Abramis brama, 20% silver carp (Hypophthalmichthys molitrix) and 10% grass carp (Ctenopharyngodon idella) and gave a greater yield than monoculture. From an average initial weight of 30 g, fish attained averages of 188 or 211 g in monoculture (average 200 g) and 221 or 278 g (average 250 g) in polyculture with maximum weights of 300 or 580 g at the end of two one-year periods. Water temperatures were 9-33°C (Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 38-39, 1996; Danesh-e-Khoshashi, 1997).

A state supported stocking programme has released about 70-80 million fingerlings into the Anzali Mordab, all descended from a single pair mating 8 years ago (Rana and Bartley, 1998a; 1998b). These fish are intolerant of low oxygen and so perform poorly under pond conditions. Stocks may be imported from Azerbaijan in the future (Rana and Bartley, 1998b). The release of 70.46 million fry in 1992-1993 to 1998-1999 period has not been successful in restoring the stocks in Iran. Stock depletion was attributed to improper fishing methods, pollution, destruction of spawning grounds, presence of predatory Esox lucius and Silurus glanis in fry stocking areas, and lack of necessary arrangements in regard to artificial spawning (Saiad Borani, 2001).

The roe or eggs of this species have been implicated in poisoning (Halstead, 1967-1970; Coad, 1979b) and should be avoided (see under the genus Schizothorax for more information on egg poisoning). Fish should be carefully cleaned in the spawning season to remove the eggs and ensure against contamination of flesh. Severe cases of egg poisoning in other species have resulted in death.

This species has been used in Iran for experimental studies, e.g. on the toxicity and LC50 of phenol and 1-naphthol (Shariati et al., 2004).

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use as food and in aquaculture.

Conservation

The subspecies has been proposed for inclusion in the "Red Book of the U.S.S.R." which forms the basis for measures to protect species (Pavlov et al., 1985).

RaLonde and Walczak (1970b) reported that 90% of the bream caught in Iran in 1970 were immature and the stock was in danger of extinction. About 19-20% of commercial catches in the Volga region are from hatchery raised stock (Petr, 1987) and it was thought that stocking could help this species in Iran. During the 1980s and 1990s there were practically no catch figures for this species in Iran. Artificial propagation began in 1986 on an experimental basis and 6 million fish were released (Ghenaat Parast, 1993). In 1992-1993 (an Iranian calendar year), 2.4 million fingerlings were released into the Anzali Mordab and nearby rivers, a 100% increase over the previous year (Abzeeyan, Tehran, 4(2):VI, 1993). Total production in government hatcheries for 1990 was 0.66 million fingerlings, in 1991 2.28 million and in 1992 5.3 million fingerlings (Emadi, 1993a). Fingerling production was 11.217 million in 1995 and 8.5 million in 1996 (Bartley and Rana, 1998a; 1998b). In 1999-2000, 20 million juveniles were released (Iranian Fisheries Research Organization Newsletter Newsletter, 23:4, 2000). From October to March 2000, 14 million juveniles raised in the Shahid Ansari aquaculture and breeding centre in Gilan were released into the Caspian Sea and neighbouring water bodies (Iranian Fisheries Research Organization Newsletter, 26:2, 2001). Illegal fishing and non-standard nets threaten the stocks (Annual Report, 1995-1996, Iranian Fisheries Research and Training Organization, Tehran, p. 19-20, 1997). Billard and Cosson (2002) give an annual production of 15 million alevins.

Ramin (1997) details studies on the artificial breeding of this species in Iran, based on 38 brooders, with the goal of saving it from extinction. Gonadotropic hormone extracted from the pituitary of the common carp was used to induce brooders. One or two doses at 5-6 mg/kg body weight gave optimum stripping of eggs at 18°C. Fertilisation rate was 75-95% and hatching rate was 75-85%. Incubation took nearly 4 days at 18-21°C. The grey, pink or yellow eggs numbered 9142-60,050 per spawner with a swelled diameter of 1.0-1.2 mm. The yolk sac was absorbed after 72 hours and newly hatched larvae were 2.9-3.7 mm long.

Khara et al. (2009) (see above) carried out their molecular study in order to determine sources for broodstock to increase genetic diversity after losses from overfishing, pollution and loss of spawning regions.

Kiabi et al. (1999) consider this species to be vulnerable in the south Caspian Sea basin according to IUCN criteria. Criteria include commercial fishing, sport fishing, few in number, habitat destruction, limited range (less than 25% of water bodies), not present in other water bodies in Iran, and present outside the Caspian sea basin. Nezami et al. (2000) consider this species to be endangered because of overfishing, habitat destruction and spawning groundn degradation.

Further work

Stocks should be carefully monitored on a continuing basis and efforts made to resurrect this commercial species.

Sources

The chief literature summary for earlier works is Backiel and Zawisza (1968) although little apparently refers to the Caspian basin populations and even less to those of the Iranian shore. Nevertheless this work gives a general overview of biology and general comments above are based on it.

Iranian material: CMNFI 1970-0542, 4, 75.4-173.7 mm standard length, Gilan, Old Safid River estuary (37°23'N, 50°11'E); CMNFI 1970-0543A, 1, 70.0 mm standard length, Gilan, Caspian Sea at Hasan Kiadeh (37°24'N, 49°58'E); CMNFI 1971-0343, 1, 158.6 mm standard length, Gilan, Langarud at Chamkhaleh (37°13'N, 50°16'E); CMNFI 1980-0127, 3, 166.1-170.1 mm standard length, Gilan, Caspian Sea near Hasan Kiadeh (37°24'N, 49°58'E); CMNFI 1980-0142, 1, 160.6 mm standard length, Gilan, Nahang Roga River (no other locality data); CMNFI 1980-0906, 3, 105.6-176.0 mm standard length, Gilan, Caspian Sea basin (no other locality data).

Abramis sapa
(Pallas, 1814)

Common names

سابا, سبا (saba, from the species name), سيم (sim = silver), ماهي سيم كند پوزه (= mahi sim kondpuzeh, meaning bluntsnout silver fish).

[pori or poru, both in Azerbaijan; tarashka, taran' and rybets, erroneously in Azerbaijan; yuzhnokaspiiskaya beloglazka or South Caspian white-eye bream in Russian; white-eye bream, southern white-eye bream, Danube bream].

Systematics

Cyprinus Sapa was originally described from the Sura, Samara and Kinel' rivers in the Volga River basin. No types known.

May be placed in the genus Ballerus Heckel, 1843 (see Hensel (1978), Shcherbukha (1973), Howes (1981), Bogutskaya (1986) and Bogutskaya and Naseka (2004) for various opinions). The nominate subspecies was described from the Volga River and tributaries.

The subspecies reported from the southern Caspian Sea basin is Abramis sapa bergi Belyaev, 1929, described from the Kura River in Azerbaijan. Eschmeyer et al. (1996) date this subspecies to 1930 although the article is dated 1929. Recognition of subspecies is disputable (Reshetnikov et al., 1997).

Key characters

The scaleless keel on the belly, deep body, very high number of branched rays in the anal fin (31-44), modally 8 branched dorsal fin rays, and uniserial pharyngeal teeth are key characters.

Morphology

Dorsal fin with 2-3, usually 3, unbranched and 7-9, usually 8, branched rays, anal fin with 3 unbranched and 31-44, mostly 34-38 branched rays, pectoral fin branched rays about 15 and pelvic fin branched rays about 8. Lateral line scales 42-55, mostly 51-52, regularly arranged over the body. Scales bear numerous very fine circuli, an almost central focus, numerous to few posterior radii (quite variable between scales of similar size) and few to none anterior radii. The anterior scale margin is wavy. A pelvic axillary scale is present. There is an evident, scaleless keel on the belly between the pelvic fin bases and the anal fin. Gill rakers 18-25, short, reaching the raker below or almost the second raker when appressed. Vertebrae 45-48. Pharyngeal teeth 5-5, with elongate, narrow and flattened, concave or rounded crowns below a hooked tip. The gut is s-shaped with a small anterior loop. The chromosome number is 2n=50 (Klinkhardt et al., 1995).

Belyaev (1929) for Kura River fish gives lateral line scale counts as 48(3), 49(6), 50(24), 51(50), 52(54), 53(16) or 54(7) and anal fin branched rays as 32(1), 33(5), 34(22), 35(32), 36(38), 37(47), 38(25), 39(9), 40(5), 41(2) or 42(1). This subspecies is distinguished from the type form in the Black Sea (Don River) by fewer lateral line scales and anal fin branched rays, a longer snout, smaller eyes, less deep body, lower dorsal fin, shorter anal fin, and longer postorbital length.

Sexual dimorphism

Unknown.

Colour

The Caspian subspecies has a dark back with a bluish tint, flanks and belly are silvery, fins are a greyish-white and sometimes have a black margin, and the iris is silvery. The peritoneum is dark brown in preserved fish.

Size

Attains 41 cm and 0.8 kg.

Distribution

Found in the basins of the Black, Caspian and Aral seas. Reported from the gut of a Silurus glanis in the Anzali Mordab (Derzhavin, 1934) but not found in recent years (Holčík and Oláh, 1992). Other reports are from the lower Safid River at Hasan Kiadeh (Belyaev, 1929; Derzhavin, 1934) and in the Aras River at Karadonly (Berg, 1948-1949).

    

Zoogeography

This species is part of a northern European and northern Southwest Asian fauna whose zoogeographical history has not been researched.

Habitat

This species feeds in brackish water but spawns and overwinters in the lower reaches of rivers. It is commonest along the western shore of the middle and southern Caspian Sea.

Age and growth

Females are 28-29 cm long on average, maximum 39 cm, while males are about 24 cm, maximum 30 cm (Belyaev, 1929). Males and females mature at 2-3 years and life span is 5 years in Azerbaijan (Abdurakhmanov, 1962).

Food

Food items include small molluscs, crustaceans and insect larvae as well as some plant fragments and detritus. Young feed on zooplankton.

Reproduction

A migration into rivers, particularly the Kura, occurs in winter when temperatures fluctuate from 5 to 10°C (Belyaev, 1929). The run begins in November and peaks in January. The Kura migration was once over 700 km from the mouth. Spawning occurs in rivers with gravel bottoms or dense vegetation from April to May. Fecundity reaches about 150,000 eggs with diameters up to 1.8 mm. Eggs adhere to stones or plants.

Parasites and predators

Eaten by Silurus glanis (Derzhavin, 1934).

Economic importance

Up to 1-2 million fish were caught in the Kura at spawning (Belyaev, 1929). The annual average catch in Azerbaijan in 1931-1935 was 1,860,000 fish weighing 6200 centners.

Conservation

The subspecies A. sapa bergi has been proposed for inclusion in the "Red Book of the U.S.S.R." which forms the basis for measures to protect species (Pavlov et al., 1985). It has always been very rare in Iran and its absence from the Anzali Mordab may be due to loss of spawning grounds (Holčík and Oláh, 1992). Lelek (1987) considers this species to be rare to vulnerable in Europe.

Further work

The status of this species in Iran should be assessed by field surveys. It is apparently quite rare and was not caught during two collecting trips along the Caspian shore in the 1970s. It is recorded only from two localities in Iran in 1929 and 1934.

Sources

Iranian material:- None available, based on literature reports.

Comparative material:- CMNFI 1986-0458, 2, 209.0-211.7 mm standard length, Germany, Danube River (48º58'N, 12º18'E); BC 59-301, 2, 136.3-154.2 mm standard length, Ukraine, Tisa, Danube drainage (no other locality data).

Genus Acanthalburnus
Berg, 1916

This genus contains only 2 species, both found in Iran. Berg (1948-1949) characterises it as similar to Alburnoides but with the last unbranched dorsal fin ray thickened into a spine which is strong basally but becomes thinner and flexible on about the last third of the ray length. Pharyngeal teeth are in 2 rows as opposed to 1 row in Abramis. Durand et al. (2002) include this genus in the Abramis clade based on cytochrome b data while Perea et al. (2010) using mitochondrial and nuclear DNA propose synonymy with Acanthobrama.

Acanthalburnus microlepis
(De Filippi, 1863)

Common names

مرواريد ماهي لب نازك (= morvaridmahi-e labnazok), kuli.

[garagas or taxta balig, both in Azerbaijan; chernobrovka and napota in Russian; blackbrow bleak].

Systematics

Abramis microlepis was described from the "Kur, presso Tiflis" (= Kura River near Tbilisi, Georgia) and the holotype is in the Istituto e Museo di Zoologia della R. Università di Torino under MZUT N.673 (Tortonese, 1940; Eschmeyer et al., 1996).

Alburnus punctulatus Kessler, 1877, described from the Kura River at Tiflis (= Tbilisi) and Borzhom, Georgia, is a synonym. A syntype of Alburnus punctulatus from the St. Petersburg Museum, 84.6 mm standard length, from "R. Kura, Tiflis" is in the Natural History Museum, London (BM(NH) 1897.7.5:34).

Alburnus Brandtii is apparently a manuscript name for this species first reported without a formal description in Brandt (1880) and listed as "Alburnus Brandtii n. sp. 1 ex. Tschaldyr" and attributed to K. Kessler in the account of the travels of Professor A. F. Brandt in Transcaucasia (see Kavraiskii, 1897). Bogutskaya (1997b) lists it as a nomen nudum.

Alburnus microlepis of Kamensky (1901), which is Acanthalburnus microlepis, should not be confused with Alburnus microlepis Heckel, 1843, a distinct species described from Aleppo (= Haleb, Syria).

Key characters

This species is distinguished from A. urmianus by having more lateral line scales, more anal fin branched rays, fewer gill rakers and gill raker morphology according to Saadati (1977). Gill raker counts are the same but scale and anal fin ray counts are generally higher with some overlap. Gill raker morphology does not appear to differ in the fish examined by me. Distribution is the easiest separating factor. Both species are distinguished from other cyprinids in Iran by the dorsal fin spine, 2 rows of pharyngeal teeth, and fin ray and scale counts.

Morphology

Dorsal fin with 3 unbranched and 7-9, usually 8, branched rays, anal fin with 2-4, usually 3, unbranched and 12-19, usually 15-17, branched rays. Pectoral fin branched rays 12-17 and pelvic fin rays 7-9. Lateral line scales 60-87. There is a large pelvic axillary scale. Scales at the base of the anal fin are somewhat enlarged and may be vertically elongate, forming a sheath. The scale focus is sub-central anterior with fine but not numerous circuli and very few posterior radii (less than 10 main radii in the largest fish seen).

Gill rakers 6-12 and sickle-shaped (Saadati, 1977) but this count presumably includes only lower arch rakers. Total gill rakers 10-14, short and only reaching the adjacent raker when appressed. The rounded raker has a triangular flap on its internal surface with the tip of the rounded raker projecting. The raker tip may be squarish or even forked in larger fish. The inner edge of the flap is finely tuberculate. Vertebrae 40-45. Pharyngeal teeth 2,5-5,2 with variants 2,5-5,1, 1,5-5,2, 1,5-5,1, 3,5-5,2, 2,5-4,2, 2,5-4,1, 2,4-5,1, 2,4-4,1 1,5-4,1, 1,5-4,0, 1,4-5,2, 1,4-5,1 and 2,6-5,2. The teeth are hooked at the tip with an elongate flat area below and the largest tooth may be strongly serrated. The posteriormost major row tooth may be almost vertically above the fourth tooth rather than posterior to it. The last unbranched dorsal fin ray is thickened in its lower two-thirds but the last third is thin and flexible. There is an obvious scaleless keel from the pelvic fins to the vent on the belly mid-line. The mouth is oblique and subterminal in adults and most young, oblique and terminal in some young. The gut is relatively short with anterior and posterior loops.

Both males and females, as well as young, may have fine tubercles distributed over the head and especially well-developed ventrally and even on the lips. Belly and lower flank scales have fine tubercles concentrated at the base of the exposed scale, some lining the scale margin. Fine tubercles line the dorsal and ventral surfaces of the pectoral and pelvic fins concentrated on rays but also on membranes, in a single file or variably dispersed.

Meristic values for Iranian specimens are:- dorsal fin branched rays 7(1) or 8(52); anal fin branched rays 14(4), 15(23), 16(24) or 17(2); pectoral fin branched rays 14(3), 15(36), 16(11) or 17(3); pelvic fin branched rays 8(53); lateral line scales 60(1), 62(2), 63(4), 64(6), 65(11), 66(6), 67(7), 68(4), 69(3), 70(5), 72(1) or 73(2); total gill rakers 10(1), 11(4), 12(18), 13(22) or 14(8); total vertebrae 43(2), 44(10) or 45(13); pharyngeal teeth modally 2,5-5,2(33) with variants 2,5-5,1(6), 1,5-5,2(3) or 1,5-5,1(2).

Sexual dimorphism

Unknown.

Colour

The back and upper head are olive-green to green and the upper flank has a golden sheen. Flanks below are silvery and the abdomen is silvery-white. There is a dark and wide stripe (about orbit diameter) on the flank, not always evident in fresh fish. Above the dark stripe is a narrow golden stripe, about one-third orbit diameter. Dorsal and caudal fins have black tips while paired fins can have a reddish or orange base. The peritoneum is brown with dark blotches or speckles.

Size

Reaches 25 cm.

Distribution

Found in the Kura River of Azerbaijan as far down as Mingechaur but not the lower reaches. In Iran it is found in the Caspian Sea basin including the Aras River shared with Azerbaijan and Iran, as far down as Karadonly, and in the Qarasu a tributary of the Aras. Reported from the Safid River basin (Abbasi et al., 1999; Kiabi et al., 1999; Abdoli, 2000; Jolodar and Abdoli, 2004; Abdoli and Naderi, 2009) and in the Anzali Talab drainage. Records from the middle Agi Chai or Talkheh River near Tabriz and the Zarrineh River of the Lake Orumiyeh basin are presumably of A. urmianus (Abdoli, 2000).

     

Zoogeography

The genus and its two species are restricted to the Caspian Sea basin and the adjacent Lake Orumiyeh basin and are presumably derived from a common ancestor related to the Alburnoides-Alburnus lineage.

Habitat

This species inhabits both rivers and lakes.

Age and growth

Females mature at 2 years (Abdurakhmanov, 1962). Spawning probably occurs in the spring judging from fish caught on 31 January which had developing eggs. Türkmen et al. (2001) found fish to 7 years of age in the upper Aras River in Turkey, with three-year-old fish dominant, and also gave length-weight and length-age relationships. Females attained a greater age and size than males.

Food

Food includes aquatic insects, crustaceans and snails, and detritus.

Reproduction

Fecundity is up to 19,060 eggs and egg diameter to 1.87 mm. In Armenia maturity is reached at the end of the second year or beginning of the third year at 80-120 mm and spawning takes place in late April to early May and may continue to late August (Pipoyan and Arakelyan, 1999). In the Turkish Aras, maturity for both sexes began at age 2 years, with all fish mature at 4 years, and spawning started in early May and continued to the end of July. Fecundity reached a mean value of 9705 eggs and egg size reached 1.65 mm (Türkmen et al., 2001).

Parasites and predators

None reported from Iran.

Economic importance

None.

Conservation

Kiabi et al. (1999) consider this species to be conservation dependent in the south Caspian Sea basin according to IUCN criteria. Criteria include sport fishing, few in numbers, habitat destruction, limited range (less than 25% of water bodies), absent in other water bodies in Iran, and absent outside the Caspian sea basin.

Further work

The biology of this species has not been investigated and its population biology is also unknown.

Sources

Type material: See above, syntype of Alburnus punctulatus (BM(NH) 1897.7.5:34).

Iranian material: CMNFI 1970-0522, 2, 55.1-71.3 mm standard length, Gilan, Safid River at Astaneh Bridge (37º16'30"N, 49º56'E); CMNFI 1970-0536, 4, 70.9-109.3 mm standard length, Gilan, Siah River estuary (36º53'N, 49º32'E); CMNFI 1970-0538, 1, 70.7 mm standard length, Gilan, Qezel Owzan River (ca. 36º44'N, ca. 49º24'E); CMNFI 1970-0583, 11, 39.0-79.9 mm standard length, Gilan, Nahang Roga River (37º28'N, 49º28'E); CMNFI 1979-0454, 8, 37.7-64.7 mm standard length, Zanjan, Qezel Owzan River at Gilavan (36º47'N, 49º08'E); CMNFI 1979-0455, 7, 50.2-123.3 mm standard length, Markazi, Manjil Dam (36º45'N, 49º17'E); CMNFI 1979-0695, 15, 71.6-112.7 mm standard length, Gilan, Safid River at Manjil Bridge (36º46'N, 49º24'E); CMNFI 1980-0116, 1, 75.5 mm standard length, Gilan, Safid River at Astaneh Bridge (37º16'30"N, 49º56'E); CMNFI 2007-0087, 4, mm standard length, Azarbayjan-e Khavari, Qareh Su (38º22'N, 48º19'E).

Comparative material: CMNFI 1980-0807, 2, 138.2-143.8 mm standard length, Turkey, Ölçek Suyu (no other locality data); CMNFI 1986-0007, 1, 132.2 mm standard length, Turkey, Kars River (ca. 41º00'N, ca. 43º00'E).

Acanthalburnus urmianus
(Günther, 1899)

 Common names

None.

Systematics

Originally described in the genus Abramis Cuvier, 1816, Berg (1948-1949) placed this species in the genus Alburnoides Jeitteles, 1861 but Saadati (1977) places the species in this genus.

The type material in the Natural History Museum, London comprises 2 specimens, 54.9-58.6 mm standard length, from the Urmi River (BM(NH) 1899.9.30:116-117), 1 specimen, 111.7 mm standard length, from the Ocksa River (BM(NH) 1899.9.30:118) (these three fish being labelled paralectotypes, with 118 being the lectotype, by P. M. Bănărescu in 1980), and 8 specimens, 50.5-111.7 mm standard length, from the Ocksa River (BM(NH) 1899.9.30:119-126), these being syntypes. Günther (1899) refers to the type series as "Five specimens from the Gader Chai and two small ones from the Urmi River; the largest is only 144 millim. long" so there is some confusion over this material.

Key characters

This species is distinguished from A. microlepis by having fewer lateral line scales, fewer anal fin branched rays, more gill rakers and gill raker morphology according to Saadati (1977). Gill raker morphology does not appear to differ in the fish examined by me. Gill raker counts are the same but scale and anal fin ray counts are generally lower with some overlap. Distribution is the easiest separating factor.

Morphology

Dorsal fin with 3 unbranched and 7-9, usually 8, branched rays, anal fin with 3 unbranched and 10-13 branched rays. Pectoral fin branched rays 14-16 and pelvic fin branched rays7-8. Lateral line scales 50-68. Scales bear only a few posterior radii and have a subcentral anterior focus. A pelvic axillary scale is present. Gill rakers 10-14, short not quite or just reaching the adjacent raker when appressed; rounded with a projected tip and distinct from its congener according to Saadati (1977) but closely resembling the structure seen in A. microlepis according to my observations (see above under A. microlepis). Pharyngeal teeth usually 2,5-5,2 or 2,5-4,2 with variants 2,4-5,2, 1,5-4,2 or 2,4-4,2. Posterior teeth are hooked at the tip, anterior teeth being rounded, and have no, slight, moderate or even strong serrations. There is a narrow and slightly concave surface below the tip. Some fish have the anterior margin of the concave surface higher than the posterior margin, but this is variable and in some teeth the condition is the reverse. The ventral keel extends from the anus to the base of the pelvic fins and is fleshy from half way to the whole length. The intestine is an elongate s-shaped with a small anterior loop. Total vertebrae 41-43.

Meristic values for Iranian specimens are:- dorsal fin branched rays 7(1), 8(20) or 9(1); anal fin branched rays 10(1), 11(4), 12(11), or 13(6); pectoral fin branched rays 14(4), 15(17) or 16(1); pelvic fin branched rays 7(3) or 8(19); lateral line scales 50(1), 52(2), 53(2), 55(1), 56(1), 57(2), 59(2), 60(3), 61(3), 62(2), 63(1), 64(1) or 68(1); total gill rakers 10(1), 11(2), 12(6), 13(8), or 14(5); pharyngeal teeth 2,5-5,2(1), 2,5-4,2(1), 2,4-5,2(1) or 2,4-4,2(1); total vertebrae 41(5), 42(7) or 43(2).

Sexual dimorphism

Male fish bear tubercles but fully tuberculate fish have not been examined. One male, 94.7 mm standard length, had a single row of tubercles on anterior pectoral fin rays.

Colour

Overall colour is silvery with a greenish-olive back and flanks with numerous minute brown pigment spots which are crowded above the lateral line to form an inconspicuous darker stripe along the whole side. The dorsal, caudal and pectoral fins have a light to evident speckling of melanophores on the rays and membranes but are almost immaculate in preserved specimens. Larger fish have pigment proximally on the anterior anal fin rays. The peritoneum is silvery but densely speckled with melanophores.

Size

Reaches 15.6 cm standard length, almost 20 cm in total length.

Distribution

This species is endemic to the Lake Orumiyeh basin, apparently in southern and western tributaries (Günther, 1899) although records of A. microlepis from the middle Agi Chai or Talkheh River near Tabriz are presumably of A. urmianus (Abdoli, 2000).

    

Zoogeography

The closest relative and only congener, Acanthalburnus microlepis, is found in the Caspian Sea basin. Connections between the Lake Orumiyeh basin and the Caspian Sea basin have been suggested by Saadati (1977), an early one in the Pliocene to early Pleistocene resulting in endemic species and a later one in the late Pleistocene resulting in species which are the same as the Caspian or only subspecifically distinct. This species presumably dates from the earlier connection (but see the Lake Orumiyeh drainage basin account for more details).

Habitat

Details of habitat requirements are unknown but is has been collected in both river and lakes.

Age and growth

Fish are mature at 14.4 cm. This species is relatively fast-growing, short-lived species with males attaining 6+ years and females 7+ years in the Kazemi Dam on the Zarrineh River (Abdoli et al., 2008). The von Bertalanffy growth curve was estimated as K = 0.427 in males and 0.506 in females, indicating that females grow faster. The sex ratio was 598♂:912♀ and there were no significant differences between males and females in the linear length-weight relationships.

Food

Diet is generally unknown and guts examined were empty except for a few plant and crustacean remains.

Reproduction

Reproductive data is unknown although this species probably spawns in the spring as do most members of this family.

Parasites and predators

Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Lernaea sp. and  Ergasilus sp. on this species.

Economic importance

None.

Conservation

This species is known only from the type series and a few other specimens in museum collections. Its status is unknown.

Further work

Field work should be carried out to determine the habitat requirements, ecology and numbers of this uniquely Iranian fish.

Sources

Type material: See above (BM(NH) 1899.9.30:116-117,  BM(NH) 1899.9.30:118, BM(NH) 1899.9.30:119-126).

Iranian material: CMNFI 1979-0093, 2, 127.5-130.5 mm standard length, Azarbayjan-e Bakhtari, Lake Qowpi (36º57'N, 45º52'E); CMNFI 2007-0098, 1, 156.3 mm standard length, Azarbayjan-e Bakhtari, river south of Mahabad (ca. 36º42'N, ca. 45º41'E); CMNFI 2007-0101, 1, 129.3 mm standard length, Azarbayjan-e Bakhtari, Tata'u River south of Miandow Ab (ca. 36º54'N, ca. 46º07'E); CMNFI 2007-0105, 1, 90.8 mm standard length, Kordestan, Zarineh River basin south of Saqqez (ca. 36º06'N, ca. 46º20'E); USNM 205904, 1, 84.7 mm standard length, Azarbayjan-e Bakhtari, Nazlu-chay near Rezaiyeh (37º40'N, 45º05'E); USNM 205934, 2, 94.5-141.9 mm standard length, Azarbayjan-e Bakhtari, Lake Qowpi (36º57'N, 45º52'E);  uncatalogued, 4, 105.1-134.9 mm standard length, Azarbayjan-e Bakhtari, Zarineh River (no other locality data);

Genus Acanthobrama
Heckel, 1843

Howes (1981) placed Acanthobrama Heckel, 1843 in the genus Rutilus Rafinesque, 1820 on osteological grounds but most other authors retain Acanthobrama as a distinct genus (Coad, 1984a; Krupp, 1985c; Eschmeyer, 1990; Bănărescu, 1992b) based on the scale, keel and anal fin characters listed below. Durand et al. (2002) include this genus in the Abramis clade based on cytochrome b data. The genus Trachibrama Heckel, 1843 is a lapsus (Krupp and Schneider, 1989).

This genus is characterised by a compressed, deep body of small to moderate size, no barbels, relatively small scales with reduced numbers of radii, a fleshy keel between the base of the pelvic fins and the vent, the last unbranched dorsal fin ray is thickened, spine-like and smooth, and the anal fin is long (9-22 branched rays). Pharyngeal teeth are usually in a single row on each arch. Gut short.

There are 8 species endemic to Southwest Asia with 1 found in southwestern Iran (Goren et al., 1973; Coad et al., 1983; Krupp, 1985c).

Acanthobrama marmid
Heckel, 1843

Common names

كلاش پا (= kalashpa), شبه ساردين (shebeh sardin = pseudo-sardine or resembling sardine), شبه نازي (= shebeh nazy), mahi sim nama (= bream-like fish).

[semnan arrez; samnan areed; arath (Rahemo et al., 2006); marmid, marmid handscherli (= marmid armed with a dagger), marmid abbiad (= white marmid), marmid asphar (= yellow marmid) or marmid mablue (= swallowing or devouring marmid) at Aleppo, arrhada (= dove, lion!) at Mosul (all these latter Arabic names after Heckel (1843b; 1846-1849a), the conflicting names for arrhada included, and are probably antiquated; Tigris bream].

Systematics

Acanthobrama Arrhada Heckel, 1843, Acanthobrama cupida Heckel, 1843, Acanthobrama marmid morpha elata Berg, 1949 and Acanthobrama marmid orontis Berg, 1949 are synonyms.

The type locality for Acanthobrama Marmid is "Gewässern bei Aleppo", for Acanthobrama arrhada "in Mossul", and for Acanthobrama cupida "in Aleppo" according to Heckel (1843b) and "Flusse Kueik bei Aleppo" in Heckel (1846-1849a). The type locality of Acanthobrama marmid morpha elata is Lake Balikli, 12 km from Erzurum, 8 km from the Karasu River, upper Euphrates, in Turkey. The type locality of Acanthobrama marmid orontis is the upper Euphrates region according to Eschmeyer et al. (1996) (but this is an error, see below).

Details on the syntypes of this species and its synonyms arrhada and cupida in the Naturhistorisches Museum Wien are given by Krupp (1985c). Eight syntypes of marmid measuring 41-144 mm standard length are listed from Mosul (in contrast to Heckel's papers where the type locality is Aleppo), the number of fish agreeing with the catalogue in Vienna. These 8 fish are under NMW 55334. Eschmeyer et al. (1996) do not list these fish as types and the card index in Vienna in 1997 concurs. A further 15 fish are listed by Krupp from the Quwayq near Aleppo: 1 fish, 102 mm standard length (NMW 55342 - not in the 1997 card index; but the following NMW fish are listed), 2, 113-139 mm standard length (NMW 55345), 2, 86-121 mm standard length (NMW 55346), 2, 98-126 mm standard length (NMW 55347), 2, 113-132 mm standard length (NMW 55348), 2, 114-138 mm standard length (NMW 79068), and 4 fish in the Senckenberg Museum Frankfurt, 82-112 mm standard length (SMF 543, formerly NMW). Eschmeyer et al. (1996) list NMW 55345-48 (8), NMW 79068 (2), SMF 543 (4) and in the Rijksmuseum van Natuurlijke Historie, Leiden RMNH 2537 (4) and RMNH 2539 (2) (both formerly NMW) as the type series.

Two syntypes of A. arrhada from Mosul, 85-92 mm standard length, are in the Senckenberg Museum Frankfurt (SMF 411, formerly NMW) (F. Krupp, pers. comm., 1985; 85.7-89.0 mm standard length) while 2 others are in the Naturhistorisches Museum Wien, ca. 150 mm standard length (NMW 55335) and 141 mm standard length (NMW 55336) (Krupp, 1985c). However, the Vienna catalogue lists 6 specimens of A. arrhada and in addition to the above material there is also NMW 55334 (8 fish) tagged as syntypes so there is some confusion in what constitutes the type series. Two possible syntypes are in the Rijksmuseum van Natuurlijke Historie, Leiden (RMNH 2538) (Eschmeyer et al., 1996).

Krupp (1985c) records syntypes of A. cupida, 151 mm standard length, (NMW 55340) and 152 mm standard length (NMW 55341). The Vienna catalogue lists 4 A. cupida which agrees with Heckel's description although I observed only NMW 55340 (1 fish), NMW 55341 (1) and also NMW 55342 (1). Eschmeyer et al. (1996) list NMW 55340-43 (1, 1, 1) as syntypes but the numbers indicate 4 fish. The card index in Vienna in 1997 also lists 55505 (5 fish), one of which is designated as the lectotype.

The 2 syntypes of Acanthobrama marmid orontis are in the Zoological Institute, St. Petersburg under ZISP 6720 from "L. Antioch, 1884, Lortet" according to Berg (1949). This subspecies is distinguished only by larger scales from the typical form but the 2 syntypes examined by me had lost their scales and were difficult to count; one seemed to have a count around 64, not as low as 54-55 recorded by Berg (1949). Krupp (1985c) examined type material and new specimens from the Orontes and found them not to differ from A. marmid from the Quwayq and Tigris-Euphrates basins. He accordingly synonymises Acanthobrama marmid orontis with the type subspecies.

Karaman (1972) considered Acanthobrama arrhada to be a subspecies of A. marmid rather than a synonym based on an unusually strongly ossified spiny dorsal fin ray in the former. Since A. marmid was described from Aleppo (= Halab, Syria) and A. arrhada from Mosul, the synonomy of these two taxa may warrant re-examination.

The fish reported from the Tigris River basin of Iran by Nümann (1966) as Xenocypris macrolepidotus was this species (Zoologisches Institut und Zoologisches Museum, Hamburg catalogue number ZMH H2700 examined by me). Saadati (1977) thought it a new species of Acanthobrama but I disagree.

A hybrid with Chalcalburnus (= Alburnus) mossulensis was reported from the Hawr al Hammar in southern Iraq by Krupp et al. (1992).

Key characters

The characters of the genus distinguish this species from all other Iranian cyprinids.

Morphology

Mouth nearly horizontal to oblique, equal or lower jaw slightly behind the upper. The belly has a fleshy keel where the ventral scales do not meet along the mid-line between the pelvic base and the anus. The last unbranched dorsal fin ray is a thickened, stiff and smooth spine, the rigid part varying from 15 to 26% of standard length. The spine may be strong for much of its length and then abruptly become thin and flexible or it may taper gradually to a flexible tip. Some small fish lack an enlarged dorsal fin spine.

Lateral line scales 53-72, scales above the lateral 10-14, scales between the pelvic fin and lateral line 4-7. There is a pelvic axillary scale. Radii are restricted to the posterior field on scales and are few in number. The focus is subcentral anterior to almost central. Dorsal fin with 3 unbranched rays and 7-9 branched rays. Anal fin unbranched rays 3, branched rays 13-22. Pectoral fin branched rays 12-18, pelvic fin branched rays 7-9. Total vertebrae 38-43 (38(3), 39(3), 40(7), 41(5), 42(7) or 43(1) combining Iranian and Iraqi material).. Gill rakers short with a basal swelling, 2-4 on the upper arch, 0-1 at the flexure and 9-12 on the lower arch. Total rakers 12-17. The rakers reach the one below or to its further base end when appressed. Pharyngeal teeth usually 5-5, with the anterior tooth compressed and bluntly pointed, the remainder bevelled with a cutting edge and a hooked tip. The two anterior teeth are more rounded than the others although the second one may have a slight hook and is bevelled. Tigris River basin fish may have 1-2 teeth in a second row. The gut is an elongate s-shape with a large anterior loop in larger fish. The diploid chromosome number is 2n=50, with the karyotype consisting of 8 metacentric, 13 submetacentric and 4 pairs of subtelocentric to acrocentric chromosomes. The karyotype is nearly identical to other Eurasian leuciscine cyprinids (Gaffaroğlu et al., 2006)..

Different body forms occur in slow-flowing and fast-flowing waters. In the former habitat fish have a deep body, often humped behind the head, while in the latter the body is more streamlined (Karaman, 1972). It seems that A. marmid is founded on the humped form and A. arrhada and A. cupida on the streamlined one.

Meristic values for Iranian specimens are:- dorsal fin branched rays 7(1) or 8(8); anal fin branched rays 13(2), 14(2), 15(4) or 17(1); pectoral fin branched rays 13(2), 14(3), 15(3) or 18(1); pelvic fin branched rays 7(1), 8(7) or 9(1); lateral line scales 54(1), 55(2), 56(2), 58(1), 59(1) or 63(1); total gill rakers 12(1), 14(6) or 17(1); pharyngeal teeth 5-4(1) or 5-5(7); and total vertebrae 38(3), 39(1), 41(2), 42(2) or 43(1).

Sexual dimorphism

Fine tubercles are found over the top, sides and bottom of the head in males. Tubercles line the first, unbranched pectoral fin ray irregularly with up to 2 branching rows. Very fine tubercles are found on the adjacent membrane and on the lower pectoral fin surface. Tubercles line the pelvic fin rays in branching rows. The lower caudal fin rays are lined with tubercles. Anterior upper flank scales, all belly scales and lower caudal peduncle scales have their margin lined with tubercles, the peduncle with some tubercles on the mid-scale and the belly with a concentration on the scale base.

Colour

The overall colour is silvery to whitish with the head and back reddish-brown. The flanks can be greyish to blackish from numerous melanophores. There may be a well-developed mid-flank stripe or it may be poorly developed or evident only posteriorly. The pelvic fins are bright red, the pectoral and anal fins less red and the dorsal and caudal fins reddish proximally and black distally. Fin colours may be more orange or yellow than red. All fin rays and membranes have melanophores and these can be quite concentrated such that some fish have dark fins. Young fish in preservative have numerous, distinctive, small to minute, rounded, square or oblong patches of pigment in 1-3, irregular, mid-flank rows. Peritoneum black, silvery with a dorsal concentration of melanophores or with widely scattered melanophores so it appears silvery.

Size

Reaches 20.8 cm (Berg, 1949)

Distribution

This species is found in the Tigris-Euphrates basin of Turkey, Syria, Iraq and Iran, the Quwayq (= Kueik) and Orontes rivers, and possibly the Amik Lake and the Bardan suyu (= stream) near Tarsus (Ladiges, 1960; - Krupp (1985c) suggests these latter should be checked). In Iran it is found in the Tigris River basin including the upper reaches of the Karkheh, the Qara Su, and in marshes such as the Hawr Al Azim.

    

Zoogeography

The majority of species are found in the Levant which once had connections to the Tigris-Euphrates basin (Krupp, 1985c).

Habitat

Hussain et al. (1997) report this species to be dominant in the small fish assemblages in the Shatt Al-Arab near Basrah, Iraq at 70.8% of 14,084 fish caught. It favours side branches off the Shatt al Arab, presumably to avoid predators which are found in deeper water. Younis et al. (2001b) noted that this species dominated in the polluted and disturbed environment of a dockyard on the Shatt al Arab. This was one of the most abundant species in the recovering marshes of southern Iraq in 2005-2006 (Hussain et al., 2006) and is also known from large rivers and dams. Also recorded from the Hawr al Azim marsh in Iran.

Age and growth

Al-Nasiri and Salman (1977) studied this species in the Little Zab River, Iraq. Their largest specimen was 13.7 cm. They described length-weight relationships and condition factors but some important length groups were missing from their samples. Condition factor showed a gradual decrease with increasing length and the means for actual and calculated weights were 1.141 and 1.118 respectively. Relative condition factor was 1.0009. Younis et al. (2001) examined three populations of this species in the Shatt al Arab, Iraq and found the 0+ age group to be represented by fish 2.1-11.0 cm long and 1+ age group by fish 8.3-14.1cm. The length-weight relationship was W = -3.821 L2.32. Four age groups with a length range of 4-19 cm were found in the Qarmat Ali River of southern Iraq, with maturity in the first year (Saud, 1997).

Ünlü et al., (1994) examined a population of this species in the Tigris River, Turkey and gave figures for growth in length and in weight. Females grew faster and are larger in size than males at the same age, particularly for age groups III and IV. Condition factor for males was 1.554 and for females 1.550. They found 5 age groups with age group III dominant for both sexes. Overall sex ratio was 1.83 females:1 male. Sexual maturity was attained by 75% of females and 85% of males in the second year of life and all fish in age group III were mature.

Food

Heckel (1843b) suggests that they are ravenous feeders based on the name "swallowing marmid". Gut contents are crustaceans, insects, and plant and gastropod shell fragments in Iranian specimens. Younis et al. (2001a; 2001b) found Shatt al Arab, Iraq fish to be detritivores, having organic detritus as the dominant gut content, followed by phytoplankton (blue-green algae and diatoms), small crustaceans (ostracods, cyclopoids, cladocerans), and aquatic plants, with dominance varying by month. Gut contents were crustaceans, insects, and plant and gastropod shell fragments in fish from Iran examined by me. In a study of the recovering Hammar Marsh, Iraq, diet was 70.77% insects and 9.81% algae with diatoms, plants, crustaceans and snails at less than 10% each, in the Hawr al Hawizah 66.4% insects and 14.1% algae, with amounts of diatoms and various crustaceans being less than 10% each, and in the Al Kaba'ish (= Chabaish) Marsh 62.7% insects and 17.7% algae with diatoms, plants and various crustaceans at less than 10% each (Hussain et al., 2006).

Reproduction

Younis et al. (2001) found most females to be ripe in March and July samples, and some were spent. Well-developed testes are noted in fish caught on 16 May in Turkey and 7 July near Ravansar, Kermanshahan indicating either a prolonged breeding season or local variations.

Ünlü et al., (1994) report spawning in May to late June for their Tigris River, Turkey population. They cite data for a Keban Dam population (on the Euphrates River in Turkey) where the spawning season is extended and runs from April to August. Egg diameter exceeds 1.2 mm and egg numbers reach 8125, and elsewhere may reach 11,000 eggs. In the Qarmat Ali River in southern Iraq, fecundity reached 1759-9293 eggs.

Parasites and predators

None reported from Iran.

Economic importance

None in Iran. In the early 1990s in Iraq, this species was used for human consumption and for fish meal (Younis et al., 2001).

Conservation

This species is rarely reported from Iranian waters and its status needs to be assessed through further field work. Endangered in Turkey (Fricke et al., 2007).

Further work

Additional field work is required to secure more materials and assess conservation status and biology.

Sources

Type material: See discussion above. Syntypes of Acanthobrama marmid (NMW 55345, NMW 55346, NMW 55347, NMW 55348, NMW 79068, SMF 543); syntypes of Acanthobrama marmid orontis (ZISP 6720), syntypes of A. arrhada (SMF 411, NMW 55335, NMW 55336, NMW 55334); syntypes of A. cupida (NMW 55340, NMW 55341, NMW 55342, NMW 55505).

Iranian material: CMNFI 1979-0287, 2, 89.9-92.1 mm standard length, Kermanshahan, spring near Ravansar (ca. 34º42'N, ca. 46º40'E); CMNFI 1979-0360, 1, 40.6 mm standard length, Khuzestan, Karkeheh River canal (31º40'N, 48º35'E); CMNFI 1979-0377, 2, 28.5-34.6 mm standard length, Khuzestan, Karkheh River (ca. 32º57'N, ca. 47º50'E); CMNFI 1979-0384, 1, 23.1 mm standard length, Khuzestan, Ab-e Shur drainage (32º00'N, 49º07'E); CMNFI 1991-0154, 1, 113.6 mm standard length, Khuzestan, Hawr-al-Azim (ca. 31º45'N, ca. 47º55'E); CMNFI 1993-0128, 1, 113.6 mm standard length, Kermanshahan, Sarab-e Sabz `Ali Khan (34º25'N, 46º32'E); CMNFI 2007-0114, Kermanshahan, Qareh Su basin (ca. 34º28'N, ca. 46º54'E); ZMH H2700, 1, 145.0 mm standard length, Kermanshahan, Gharasu-Gamasiab-Seymarreh (Qareh Su, Gav Masiab and Simareh rivers, no other locality data); uncatalogued, 1, 101.7 mm standard length, Kermanshahan, sarabs near Kermanshah (no other locality data).

Comparative material: BM(NH) 1931.12.21:22-25, 4, 65.7-84.6 mm standard length, Iraq, Mosul (ca. 36º20'N, ca. 43º08'E); BM(NH) 1974.2.22:1084-1091, 7, 105.1-118.3 mm standard length, Iraq, Najab Bazar (no other locality data); BM(NH) 1974.2.22:1094, 109.3 mm standard length, Iraq, Great Zab River at Aski Kalak (36º16'N, 43º39'E); BM(NH) 1971.4.2:7, 96.5 mm standard length, Iraq, River Tigris near Mosul (ca. 36º20'N, ca. 43º08'E); BM(NH) 1974.2.22:1078-1083, 6, 105.2-122.8 mm standard length, Iraq, Najab Bazar (no other locality data); BM(NH) 1974.2.22:1092, 109.5 mm standard length, Iraq, Najab Bazar (no other locality data); CMNFI 1987-0017, 3, 83.8-108.3 mm standard length, Iraq, Hawr al Hammar (no other locality data); BM(NH) 1920.3.3:147-156, 15, 29.5-102.0 mm standard length, Syria, Ouadi Khneizer (no other locality data); BM(NH) 1968.12.13:108-112, 1 (of 5), 112.6 mm standard length, Syria, Ouadi Khneizer, Khabour (no other locality data) (collections amalgamated as BM(NH 1968,12.13:105-341 seem to include the preceding and following collectiosn, 224 (7 as alizarin specimens), 24.1-69.7 mm standard length); BM(NH) 1968.12.13:113-118, 6, 56.5-117.4 mm standard length, Syria, River Euphrates at Houreira (no other locality data); ZSM 26136, 5, 55.3-80.3 mm standard length, Syria, Assad Reservoir, Euphrates basin (no other locality data); CMNFI 1980-0810, 2, 114.8-118.3 mm standard length, Turkey, Göksu in Tigris River basin (no other locality data); CMNFI 1980-1036, 1, 101.5 mm standard length, Turkey, Keban Dam on Murat Nehri near Elâzığ (no other locality data).

Genus Alburnoides
Jeitteles, 1861

This genus is found in Europe, Asia Minor and Central Asia with ?11 species, with 6 reported in Iran.

The riffle minnows are similar in appearance to the genus Alburnus but have smooth rather than serrated pharyngeal teeth. Arguably this distinction is insufficient to warrant a separate genus but it is retained here as this has not been investigated in depth and the genus has widespread usage. Certainly it is not uncommon to find individuals of Alburnus hohenackeri lacking serrations on their pharyngeal teeth.

Pharyngeal teeth in Alburnoides are in 2 rows with strongly hooked tips but unserrated, scales of medium size, no groove before the dorsal fin, a keel behind the pelvic fins is usually scaleless but may be wholly scaled, short dorsal and moderate to long anal fin, last dorsal fin unbranched ray thickened, decurved lateral line often with a characteristic spotting pattern above and below each pore, and gill rakers short and few.

Alburnoides bipunctatus (Bloch, 1782) was the name applied to most populations across Europe and the Middle East from France north of the Alps eastwards to the Black, Caspian and Aral Sea basins but ongoing research is revealing a greater diversity (Bogutskaya and Coad, 2009; Coad and Bogutskaya, 2009).

A record of A. bipunctatus from a qanat at Hormak (29°58'N, 60°51'E) in the Sistan basin by Saadati (1977) is probably an error of labelling or sorting. It is not mentioned in the collector's (R. J. Behnke) original field notes nor in a typed version. Also this species was not collected there by me.  

Records of parasites for fish identified as "A. bipunctatus" in Iran are as follows:

Jalali and Molnár (1990a) record the monogeneans Dactylogyrus alatus and D. chalcalburni from this species in the Zayandeh Rud. Gussev et al. (1993b) also reports the latter species and locality. The monogenean Diplozoon paradoxum is recorded from this species in the Tajan River, Mazandaran (Iranian Fisheries Research and Training Organization Newsletter, 6:7, 1994). Shamsi et al. (1997) report Clinostomum complanatum, a parasite causing laryngo-pharyngitis in humans, from this species. Masoumian and Pazooki (1998) surveyed myxosporeans in this species in Gilan and Mazandaran provinces, finding Myxobolus ellipsoides. Masoumian et al. (2005) report the protozoan parasites Ichthyophthirius multifilis, Trichodina perforata and Chilodonella, sp. from this species in water bodies in West Azarbayjan. Mortazavi Tabrizi et al. (2005) record Ligula intestinalis in this species from the Sattarkhan Dam in East Azerbaijan. Pazooki et al. (2005) record Trichodina  perforata from this species in waterbodies of Zanjan Province. Pazooki et al. (2006) record the monogeneans Dactylogyrus vistulae, Gyrodactylus sp. and Paradiplozoon sp. from this fish in Zanjan Province. Mehdipoor et al. (2004) record the monogeneans Dactylogyrus alatus, D. chalcalburni and D. pulcher in the Zayandeh River.

To be assigned: CMNFI 1970-0522, 22, 40.4-80.3 mm standard length, Gilan, Safid River at Astaneh Bridge (37º16'30"N, 49º56'E); CMNFI 1970-0536, 3, 71.9-89.6 mm standard length, Gilan, Siah River estuary (36º53'N, 49º32'E); CMNFI 1970-0546, 3, 57.1-69.4 mm standard length, Gilan, Safid River canal (no other locality data); CMNFI 1970-0551, 1, 108.4 mm standard length, Gilan, Ghaleh River near Fowman (37º13'N, 49º19'E); CMNFI 1970-0583, 16, 40.7-87.3 mm standard length, Gilan, Nahang Roga River (37º28'N, 49º28'E); CMNFI 1971-0327A, 6, 59.3-81.0 mm standard length, Gilan Shafa River (37º35'N, 49º09'E); CMNFI 1979-0239, 2, 57.1-79.3 mm standard length, Markazi, Nam River near Firuzkuh (35º43'N, 52º40'E); CMNFI 1979-0439A, 4, 53.4-72.2 mm standard length, Gilan, Shafa River (37º35'30"N, 49º05'30"E); CMNFI 1979-0440, 11, 53.7-88.6 mm standard length, Gilan, Lomir River (37º37'N, 49º02'30"E); CMNFI 1979-0441, 4, 52.4-55.7 mm standard length, Gilan, river 14 km south of Hashtpar (37º42'N, 48º58'E); CMNFI 1979-0445, 1, 70.6 mm standard length, Gilan, stream 10 km south of Astara (38º21'N, 48º51'E); CMNFI 1979-0453, 2, 45.8-65.1 mm standard length, Zanjan, Zanjan River (37º06'N, 47º56'E); CMNFI 1979-0454, 6, 39.6-56.0 mm standard length, Zanjan, Qezel Owzan River at Gilavan (36º47'N, 49º08'E); CMNFI 1979-0483, 2, 93.0-98.6 mm standard length, Mazandaran, Chashmeh River (37º23'30"N, 55º51'30"E); CMNFI 1979-0493, 11, 51.1-82.8 mm standard length, Mazandaran, Tajan River drainage (36º19'N, 53º23'E); CMNFI 1979-0695, 74, 34.1-71.1 mm standard length, Gilan, Safid River at Manjil Bridge (36º46'N, 489º24'E); CMNFI 1980-0116, 19, 41.1-70.3 mm standard length, Gilan, Safid River at Astaneh Bridge (37º16'30"N, 49º56'E).

check all species correctly transposed from Bogutskaya and Coad?

add holotype meristics to paratypes frequency

As A. bip from Atrak, Gorgan Gharasu, Tajan, babol, Jaraz, sardab, Aras, Tonekabon, Pol-e Rud, Safis and Anzali Talab (Abdoli and Naderi, 2009).

Alburnoides eichwaldii
De Filippii, 1863

Common names

خياطه (= khayataeh) or ماهي خياطه (= mahi khayateh, tailor or tailoress fish, possibly from lateral line pattern like stitches), لپك (= lapak in Mazandaran), پرك (= parak in Gilaki), sima, kuli.

[gijovcu in Azerbaijan; vostochnaya bystryanka or oriental bystranka, zakavkazskaya bystryanka or Transcaucasian bystranka, Armyanskaya bystryanka or Armenian bystryanka for A. b. armeniensis, all in Russian; spirlin, riffle minnow or riffle bleak in general].

Systematics

Cyprinus bipunctatus was originally described from the Weser River in Germany. Alburnus Eichwaldii De Filippi, 1863, described from the "Kur presso Tiflis" (= Kura River near Tbilisi, Georgia), was regarded as a Caspian Sea basin subspecies of Alburnoides bipunctatus but Bănărescu (1991) briefly stated that it cannot be distinguished from Alburnoides bipunctatus fasciatus (Nordmann, 1840) of the Black Sea basin. Holčík and Jedlička (1994) considered that the observed variation is clinal and subspecies are not warranted. Reshetnikov et al. (1997) also consider subspecies as disputable. There is another nominal subspecies in the Aras River drainage of Armenia, Alburnoides bipunctatus armeniensis Dadikyan, 1972, from the rivers Arpa, Vorotan, Vedi, Marmarik, Kasakh and their tributaries, now regarded as a synonym of eichwaldii (Bogutskaya and Coad, 2009).

Bogutskaya and Coad (2009) resurrect A. eichwaldii, and it is present in at least in the western part of the Caspian Sea basin, west of the Safid River.

A syntype of Cyprinus bipunctatus described from the Weser River, Germany is in the Museum für Naturkunde, Universität Humboldt, Berlin (ZMB 3357) (Eschmeyer et al., 1996).

Two syntypes of Alburnus eichwaldi from "Tiflis" are in the Naturhistorisches Museum Wien under NMW 55516 and 4 syntypes are in the Istituto e Museo di Zoologia della R. Università di Torino under MZUT N.677 (Tortonese, 1940; Eschmeyer et al., 1996).

Syntypes of Alburnoides bipunctatus armeniensis are in the Zoological Institute, St. Petersburg under ZISP 37502.

Key characters

See B and C 2009 here and below

The pigmentation along the lateral line is distinctive. Total gill raker counts (5-12) are much less than in Alburnus hohenackeri (16-29, usually 20 or more) which has similar general scale and fin rays counts.

A. eichwaldii differs from the related A. gmelini by having fewer branched anal-fin rays (11-14, modally 12-13, vs. 13-16, modally 14-15; means, 12.2 and 14.3, respectively) and a larger number of total vertebrae (mean 41.3 vs. 40.6, statistically different).

Morphology

see tables for counts?

The original diagnosis of A. eichwaldii gave the following characters: the body is deep, its length exceeds the depth in four times; eye large; dorsal-fin rays branched rays 8; branched anal-fin rays 12; scales in the lateral series 50, 11 scales above and 7 scales below lateral line. The original description may be added to by the following combination of characters: the caudal fin lobes are moderately rounded, the fin is not deeply forked; the ventral keel is commonly scaleless but may be variably scaled (up to completely scaled); the head is commonly deep and the snout is slightly to markedly rounded; the upper jaw is slightly protruding over the lower jaw; the tip of the mouth cleft is slightly below the level of the middle of the eye or at about the lower margin of pupil; the number of dorsal-fin rays is 8, rarely 7 or 9; the number of branched anal-fin rays is (10)11-14 with the modal range of 12-13; pharyngeal teeth are commonly 2.5-4.2 and other variants with four teeth in the longer row of the right ceratobranchial, also, less frequently, 2.5-5.2 or 2.5-5.1; the number of total lateral line scales 44-56 (Dadikyan, 1972, 1973, gives 39-56, averaging 48.7, in A. bipunctatus armeniensis); gill rakers 6-10; the number of total vertebrae is (38, 39)40-43 with a mode of 41; predorsal vertebrae are (12)13-15 with a mode of 14; the number of abdominal vertebrae is (18)19-22 with a mode of 21, and that of caudal vertebrae is 19-22 with a mode of 21; the caudal region is commonly one vertebra shorter than, equal to the abdominal region or one vertebra longer than the abdominal region, and the difference between the abdominal and caudal numbers varies from +3 to -1 with a mode of 0; and the most common vertebral formulae are 21+21, 21+20 and 20+21.

Dorsal fin with 2-3 unbranched and 6-10, usually 8, branched rays, anal fin with 2-3 unbranched and 10-18 branched rays, usually 12-13 (but see below for Iran). Lateral line scales 41-58. Gill rakers 5-12, usually 7-10. Vertebrae 37-44. Pharyngeal teeth 2,5-4,2, rarely 2,5-5,2, 2,4-5,2, or 1,5-4,2, with variants being 1,5-4,1, 2,5-4,3, 2,3-4,2, 2,4-4,2, 1,5-4,0, and 1,2,5-4,3. The chromosome number is 2n=50 (Klinkhardt et al., 1995).

Meristics in Iranian specimens from the Caspian Sea: dorsal fin branched rays 7(6), 8(121) or 9(3); anal fin branched rays 11(1), 12(26), 13(61), 14(32), 15(9) or 16(1); pectoral fin branched rays 12(3), 13(23), 14(71), 15(24) or 16(9); pelvic fin branched rays 6(3), 7(116) or 8(11); lateral line scales 43(4), 44(5), 45(25), 46(29), 47(23), 48(13), 49(7), 50(10), 51(6) 52(5), 54(1) or 55(2); total gill rakers 6(7), 7(35), 8(57), 9(30) or 10(1); pharyngeal teeth 2,5-4,2(14), 2,5-5,2(1) or 2,4-5,2(2); and total vertebrae 37(1), 38(1), 39(4), 40(49), 41(32) or 42(2).

The chromosome number is 2n=50 and Nazari et al. (2009) give further details.

Sexual dimorphism

Abdurakhmanov (1962) reports pelvic fin length greater in males and snout length greater in females for this species in Azerbaijan.

Colour

?There is a characteristic pigmentation along the lateral line with a small spot above, and another below, the lateral line opening on each scale. This only appears in preserved material as live fish are an overall silvery colour. It can be absent, mostly in lake forms. The flank has a blue-grey stripe wider than the eye diameter. Above the lateral line there may be a series of 5-9 black lines formed of triangular blotches and 3-5 similar lines below the lateral line. The back and head are dark olive, almost black, dark green or dark brown. The flank above the lateral line may have purple iridescent tints. The flanks can be a golden yellow. The belly and lower head are pearly-white. The dorsal and caudal fins have some grey pigment or may be dark grey. The bases of the pectoral, pelvic and anal fins have orange to red pigmentation which is not well developed in young. The extent and intensity of this pigment is variable between fins, although in some fish it is equally developed in all these fins.

Size

?

Distribution

Found in river drainages of the southwestern Caspian coast from the Samur (according to Berg, 1948-1949) down to rivers of the Lenkoran’. The Aras River basin also harbours this species.    

Zoogeography

?This species shows considerable variation over its range from Europe to southern Iran. Dadikyan (1973) demonstrated variability in this species in a mountainous region of Armenia within the Aras River basin. Up to 10 characters could be used to distinguish populations within the same river but taken at different altitudes. Populations at similar altitudes but in different rivers (and habitat types, e.g. rushing rocky streams compared to a bog) also varied but the characters were not necessarily the same as those distinguishing altitudinal variants within one river. Local conditions, such as temperature and flow regime, may govern the characters at any one site. Gene flow may play a part as fish are carried downstream by heavy rainfall. Populations living within the same river are presumably more closely related than populations in different river systems but may show more differences than populations at similar altitudes but which have had no gene flow for long periods. These factors complicate designation of subspecies in this species and accurate analysis requires large series of specimens.

Habitat

?This species inhabits small streams and is less frequent in the main flow of large rivers. In Iran, it is one of two most abundant species in Caspian rivers along with Capoeta capoeta (Iranian Fisheries Research and Training Organization Newsletter, 19:4, 1998). It prefers well-oxygenated water, low in pollution, with hard stream beds. In laboratory experiments with European specimens, Bless (1996) found that reproduction requires a stream velocity of 0.4 ms-1 and a gravel substrate with a diameter of 2-15 cm which allows interstitial flow.

Age and growth

In Azerbaijan, maturity is attained at 1-2 years and life span is 3 years (Abdurakhmanov, 1962).

Food

Food is taken from the bottom or from the water surface, the former being mostly insect larvae and the latter terrestrial organisms which fall on the water. Abdoli (2000) lists Simuliidae, Plecoptera, Ephemeroptera, Chironimidae and Trichoptera. Diatoms are also found in gut contents (Abdurakhmanov, 1962).

Reproduction

?Spawning takes place in spring (April-June) at 13-15.6°C and adhesive eggs are laid on sand or gravel in fast-flowing water. Fecundity reaches 6496 eggs and egg diameter 2.16 mm (Abdurakhmanov, 1962). Bless (1996) reports multiple spawning over a period of 15 weeks in laboratory conditions.

Parasites and predators

Barzegar et al. (2008) record the digenean eye parasite Diplostomum spathaceum from this fish (as A. bipunctatus).

Economic importance

Unknown. A. bipunctatus is listed as important to North Americans (Robins et al., 1991). Importance is based on its use as bait and in textbooks. It is also a known feeder on the larvae of the malaria-carrying mosquito.

Conservation

Lelek (1987) considers A. bipunctatus to be vulnerable to endangered in Europe through pollution and eutrophication. It is listed as Vulnerable in Turkey (Fricke et al., 2007). Kiabi et al. (1999), examining Iranian material, consider A bipunctatus to be of least concern in the south Caspian Sea basin according to IUCN criteria. Criteria include abundant in numbers, habitat destruction, widespread range (75% of water bodies), present in other water bodies in Iran, and present outside the Caspian Sea basin. These assessments may apply to the current taxon.

Further work

Other populations in Iran related to this taxon are under study (2009).

Sources

Iranian material: CMNFI 2007-0090, ?

Comparative material: See Bogutskaya and Coad (2009).

Alburnoides idignensis
Bogutskaya and Coad, 2009

 

?
Description of holotype. A ventral keel
between the pelvics and the anal fin is scaleless
along about 1/2 of its length. There is a
pelvic axillary scale and scales extend over
the proximal bases of the anal fin forming
a sheath. The upper body profile is convex,
similar to the lower profile. The caudal
fin lobes are rounded, the fin is shallowly
forked. The snout is markedly rounded,
stout. The mouth is small, between terminal
and subterminal; the tip of the mouth
cleft is on a level of the lower margin of the
pupil.


Dorsal fin rays are 3 unbranched and 8½
branched, anal fin rays are 3 unbranched
and 12½ branched, branched pectoral fin
rays are 14, pelvic fin branched rays are 7.
The anal fin origin is somewhat in front of a
vertical from the posterior end of the dorsal
fin base. Total lateral line scales number 45
and those to posterior margin of hypurals
44, scales around caudal peduncle 15, scales
above lateral line to dorsal fin origin are 9,
scales below lateral line to anal fin origin
are 5, scales below lateral line to pelvic fin
origin are 4, and midline predorsal scales
are 19. Pharyngeal teeth 2.5-4.2. Gill rakers
number 7, they are short and stubby, the
longest touching the adjacent one when appressed.
Total vertebrae are 38, comprising
19 abdominal and 19 caudal vertebrae. Predorsal
vertebrae number 11.

Description of paratypes.
The body is moderately compressed,
relatively thick. The caudal fin lobes are
rounded, the fin is shallowly forked. The
ventral keel between the pelvics and anal
fin is variably scaled: completely scaleless
(4), scaled along about ¼-1/3 of its length
(11), scaled along ½ of its length (7), scaled
along about 2/3 of its length (3) or completely
scaled (4). The anal fin origin is in
front of a vertical from the posterior end
of the dorsal fin base. The snout is moderately
stout, rounded. The mouth is almost
horizontal, its position is between terminal
and subterminal; the tip of the mouth cleft
is between a level of the lower margin of the
pupil and a lower margin of the eye. The
junction of the lower jaw and the quadrate
is on about a vertical through the anterior
margin of the pupil.



Dorsal fin unbranched rays 3, branched
dorsal-fin rays 6½ (1), 7½ (2) and 8½ (10)

(7.7, 0.63). Anal fin unbranched rays 3,
branched anal-fin rays 10½ (1), 11½ (8)
12½ (4) (11.2, 0.60). The dorsal fin outer
margin is truncate to markedly convex
and the anal fin outer margin is clearly
concave. Pectoral fin branched rays 12(2),
13(5), 14(4), 15(2) (13.5, 0.97), pelvic fin
branched rays 6(1), 7(12) (6.9, 0.28).
Pharyngeal tooth counts are 2.5-4.2
(20), 2.4-4.2 (5), 2.5-4.1 (2), 2.5-4.3 (2),
1.5-4.2 (1). The lateral line is complete with
none, 1 or 2 unpored scales at the posterior
end of the lateral series; total lateral line
scales 41(3), 42(2), 43(1), 44(4), 45(1),
46(2) (43.3, 1.80); lateral line scales to the
margin of hypurals 39(1), 40(3), 41(2),
42(1), 43(3), 44(3) (41.9, 1.77). Scales
around caudal peduncle 12(1), 13(-), 14(3),
15(5), 16(-), 17(4) (15.2, 1.52); scales between
dorsal fin origin and lateral line 8(1),
9(11), 10(1) (9.0, 0.41); scales between anal
fin origin and lateral line 4(5), 5(7), 6(1)
(4.7, 0.63); scales between pelvic fin origin
and lateral line 4(10), 5(3) (4.2, 0.44), and
predorsal scales 17(1), 18(2), 19(6), 20(1),
21(2), 22(1) (19.3, 1.38). Total gill rakers in
the outer row on first left arch number 6(2),
7(3), 8(7), 9(1) (7.5, 0.88). Total vertebrae
38(1), 39(11), 40(1) (39.0, 0.41).
Other characters as in holotype.
Paratypes bear pigmentation above and
below the lateral line pores, forming a pale
line margined with dark although this is
obscured by background pigment on the
caudal peduncle. A mid-flank stripe is diffuse
posteriorly and fades anteriorly. A thin
dark stripe at the junction of the hypaxial
and epaxial muscles masses is evident
but also fades anteriorly. The pigment on
scales above and below the lateral line can
be strongly or weakly expressed, forming
stripes, but can be absent. The back is dark
and obscures a predorsal and postdorsal
stripe. A series of strong melanophores is
present on the inner margin of the pectoral
fin unbranched ray. Most fins lack much
pigment, the dorsal fin pigment lining the
rays being the strongest apart from that
noted on the pectoral fin.
Summarized data for the paratypes and
additional material of A. idignensis material
(excluding holotype).
Dorsal fin unbranched rays 3, branched
dorsal-fin rays 6½ (1), 7½ (10), 8½ (50),
9½ (1); among 46 radiographed specimens
6½ (1), 7½ (10), 8½ (35) (7.7, 0.49) (Table
1). Anal fin unbranched rays 3, branched
anal-fin rays 9½ (1), 10½ (2), 11½ (29),
12½ (23), 13½ (6), 14½ (1); among 46
radiographed specimens 9½ (1), 10½ (2),
11½ (23), 12½ (16) (11.3, 0.67). The dorsal
fin outer margin is truncate to markedly
convex and the anal fin outer margin is
slightly concave. Pectoral fin branched rays
12(2), 13(20), 14(23), 15(15), 16(2), pelvic
fin branched rays 6(3), 7(58), 8(1).
Total lateral line scales 41(4), 42(8),
43(2), 44(10), 45(14), 46(10), 47(7), 48(3),
49(2), 50(1), 51(1); lateral line scales to
the margin of hypurals 39(2), 40(7), 41(5),
42(4), 43(13), 44(9), 45(10), 46(7), 47(2),
48(2), 49(1). Scales around caudal peduncle
12(1), 13(-), 14(11), 15(21), 16(14),
17(12), 18(3); scales between dorsal fin
origin and lateral line 8(3), 9(33), 10(21),
11(5); scales between anal fin origin and
lateral line 4(20), 5(32), 6(8), 7(2); scales
between pelvic fin origin and lateral line
3(2), 4(16), 5(29), 6(15), and predorsal
scales 17(2), 18(6), 19(19), 20(12), 21(12),
22(7), 23(3), 24(1). Total gill rakers in the
outer row on first left arch number 6(7),
7(14), 8(32), 9(8), 10(1).
Vertebral counts given below were calculated
in 46 specimens. Total vertebrae
number (37)38-40 with a mode of 39 (39.0,
0.65) (Tables 2 and 4). Predorsal vertebrae
number 11-13(14) (12.2, 0.4) (Tables 2 and
5). Abdominal vertebrae number (18)19-20
(19.5, 0.55) (Tables 3 and 5). Caudal vertebrae
number (18)19-20 (19.5, 0.55) (Tables
3 and 6). The vertebral formulae are 20+19
(16), 19+20 (14), 20+20 (8), 19+19 (6),
19+18 (1), and 18+20 (1). Thus, the mean
difference between abdominal and caudal
counts varies between +3 and -2 averaging
0 (0.0, 0.88) (Tables 3 and 6).


see tables for counts

Common names

شبه زوري (shebeh zury = resembling zury) in Khuzestan for Alburnoides spp..

Systematics

The holotype (CMNFI 2007-0118) is a male, 106.8 mm TL, 89.2 mm SL from Kermanshahan, Bid Sorkh River between Sahneh and Kangavar, Gav Masiab River drainage, ca. 34°23´N, 47°52´E; 1976 and paratypes (CMNFI 2007-0118A) number 13, 33.5-90.0 mm SL, same data as holotype. The species is named for the Tigris River which was called Idigna in Sumerian (Akkadian: Idiklat; biblical: Hiddekel; Arabic: Dijlah; Turkish: Dicle).

Key characters

This species is distinguished by a combination of characters which includes an unbranched pectoral fin ray strongly lined with melanophores on its inner margin; an eye of an average size, the orbit diameter larger than the snout length and markedly smaller than the interorbital width; caudal fin lobes rounded and fin shallowly forked; a variably scaled ventral keel though most commonly scaled along about 1/3-2/3 of its length; a deep head with a markedly rounded,
stout snout; a small mouth which is between terminal and subterminal; a tip of the mouth cleft on a level from the lower margin of the pupil; commonly 8 branched dorsal-fin rays; 10-12(13-14) branched anal-fin rays; 41-49(50-51) total lateral line scales (39-49 scales to posterior margin of hypurals); commonly 2.5-4.2 or 2.4-4.2 pharyngeal teeth; (37)38-40, with a mode of 39, total vertebrae; 11-13(14) predorsal vertebrae, (18)19-20 abdominal vertebrae; (18)19-20 caudal vertebrae; a caudal vertebral region most commonly one vertebra shorter or one vertebra longer than the abdominal region; the most common vertebral formulae are 20+19 and 19+20, and the difference between  the abdominal and caudal counts averaging 0.

Morphology

Sexual dimorphism

The following characters were significantly different between sexes (p<0.05). Greater in females: head width, postorbital distance, pelvic fin origin to anal fin origin distance. Greater in males: head length, pectoral fin length in pectoral fin origin to pelvic fin origin distance, and pelvic fin length in pelvic fin origin to anal fin origin distance.

Colour

The lateral line is delineated by some darker pigment above and below but not as strongly as in the A. petrubanarescui holotype and obscured by background pigmentation on the caudal peduncle. Some pigment on the flank scales above and below the lateral line give the impression of stripes but is not strongly developed.A mid-flank stripe is not developed. A thin dark stripe separates the epaxial and hypaxial muscle masses. The back is dark and obscures a predorsal and postdorsal stripe. The fins are mostly immaculate, with some melanophores lining the rays of the dorsal and pectoral fins in particular. The unbranched pectoral fin ray is strongly lined with melanophores on its inner margin. The peritoneum is silvery with fine melanophores and some spots.

Size

?

Distribution

This species is known from some upper reaches of tributaries of Karkheh [Qareh Su] River in the Zagros Mountains. The Karkheh drains into the Tigris just below its confluence with the Euphrates.

Zoogeography

?

Habitat

This species was captured in the Sarab Dowrah River at an altitude of 1370 m, in clear water at 19°C, with pH 6.8, the shore was bushy, some plants were present in the water, and the river had a stony bed. Other
species recorded together with this species were Barbus lacerta, a “Nemacheilus” sp., Alburnus mossulensis, Capoeta aculeata, Cyprinion macrostomum and Garra rufa.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

Unknown.

Conservation

See under A. eichwaldii.

Further Work

This recently described species needs its conservation status and biology investigated.

Sources

Comparative material: CMNFI 1979-0278, 5, 43.3-52.8 mm SL, Lorestan, Sarab Dowrah River in Kashkan River drainage, 30 km from Khorramabad (33°34´N, 48°01´E); CMNFI 2007-0075, 36, 38.1-72.1 mm SL, Hamadan, Qareh Su River system, Malayer River at bridge 5 km from Malayer (ca. 34°17´N, 48°47´E); CMNFI 2007-0115, 8, 43.3-62.7 mm SL, Kermanshahan, stream in Karkheh River system north of Kermanshah (ca. 34°34´N, 46°47´E).

Alburnoides namaki
Bogutskaya and Coad, 2009

 

see tables for counts

Common names

None.

Systematics

The holotype, CMNFI 1979-0461, is a female, 91.2 mm SL from Hamadan, qanat at Taveh, 35°07´N, 49°02´E. Paratypes are under  CMNFI 1979-0461A, 188, 27.2-96.9 mm SL, same data as the holotype. The species is named for the Namak Lake. Namak means salt in Farsi.

Key characters

 

Morphology

Sexual dimorphism

The following characters were significantly different between sexes (p<0.05). Head depth, body depth, head width, orbit diameter, and predorsal length were greater in females while pectoral fin length, pelvic fin length, longest dorsal fin ray length, pectoral fin length in pectoral fin origin to pelvic fin origin distance and pelvic fin length in pelvic fin origin to anal fin origin distance were greater in males. One male bore tubercles lining scale margins and sparsely on the top and sides of the head. Tubercles are strongest on scales of the caudal peduncle. The anal-fin rays bear tubercles which follow the branching of the distal rays. Tubercles are present on the dorsal, pectoral and pelvic fin rays but are less developed than those on the anal fin.

Colour

harmonise within?

The lateral line is somewhat darker than the surrounding flank but there are no strong spots or dark outline to canal. Some pigment on flank scales above and below the lateral line give a faint impression of stripes. A mid-flank stripe is only weakly apparent. A predorsal and postdorsal stripe is present on the back. The fins are mostly immaculate, with some melanophores lining the rays of the dorsal and pectoral fins. The flanks were a golden-yellow, belly white, back dark green, base of paired and anal fins orange, other fins hyaline in life. Some paratypes bear strong pigmentation above and below the lateral line pores, forming an evident pale line margined with dark. A broad mid-flank stripe can be well developed or weakly expressed and, on the caudal peduncle, obscures the lateral line pigment pattern. However, the lateral line pattern can be weak and this can be seen over the anal fin where the flank stripe does not extend down to the decurved lateral line. The pigment on scales above and below the lateral line (and below the mid-flank stripe) can be strongly or weakly expressed, and in the former case it appears as a series of thin, discontinuous stripes. Some fish have a series of strong melanophores on the inner margin of the pectoral fin unbranched ray. Dorsal fin membranes may be dusky and lack pigment lining the rays. The peritoneum is silvery with a few melanophores.

 

Size

 

Distribution

    

    

Zoogeography

 

Habitat

Habitat data is based only on the collection data. Altitude was 1640 m, water temperature 15.5 °C, pH 6.0, conductivity 1.2 mS, qanat stream width 1.5 m, maximum depth 75 cm, vegetation in water encrusting, shore grassy, gravel and mud bottom, medium current, and water clear in parts, others cloudy and polluted, The species was collected with Capoeta buhsei.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

Unknown.

Conservation

See under A. eichwaldii.

Further Work

This recently described species needs its conservation status and biology investigated.

Sources

Type material: See above.

Comparative material: CMNFI 2007-0121, 3, 28.0-74.8 mm SL, Hamadan, stream in Qareh Chay basin north of Razan, ca. 35°25´N, 49°02´E; CMNFI 2007-0074, 4, 33.1-41.8 mm SL, Markazi, Qareh Chay, 32 km west
of Arak, 34°03´N, 49°21´E;  ZMH 4183 (7, ).?

Alburnoides namaki
Bogutskaya and Coad, 2009

Diagnosis. The species is distinguished by
a combination of characters which includes
the lack of strong spots or dark outline to the
lateral line canal; a small eye, the orbit width
about equal to the snout length but markedly
smaller than the interidth; caudal
fin lobes rounded and fin shallowly forked; a
sharp scaleless ventral keel behind the pelvic
fins along the abdomen to the anus; a deep
head with a stout snout which is markedly
rounded; a tip of the mouth cleft on the level
below the lower margin of the eye; commonly
8½ branched dorsal-fin rays; 10-13½,
commonly 11-12½, branched anal-fin rays;
(43)44-50(52) total lateral line scales (42-51
scales to posterior margin of hypurals); 2.5-
4.2 pharyngeal teeth (or other variants with
four teeth on the right ceratobranchial);
commonly 39-41 total vertebrae; 11-13(14),
commonly12-13, predorsal vertebrae; 19-
20(21) abdominal vertebrae; 19-21 caudal
vertebrae; a caudal vertebral region most
commonly equal to the abdominal region;
and the most common vertebral formulae are
20+20, 20+19 and 19+20.
Description of holotype. A ventral keel
between the pelvics and the anal fin is completely
scaleless. There is a pelvic axillary
scale and scales extend over the proximal
bases of the anal fin forming a sheath. Dorsal
fin rays are 3 unbranched and 8½ branched,
anal fin rays are 3 unbranched and 12½
branched, branched pectoral fin rays are 13,
pelvic fin branched rays are 6. The anal fin
origin is on a vertical from the posterior end
of the dorsal fin base. Total lateral line scales
number 50 and those to posterior margin of
hypurals 51, scales around caudal peduncle
16, scales above lateral line to dorsal fin origin
are 12, scales below lateral line to anal
fin origin are 6, scales below lateral line to
pelvic fin origin are 7, and midline predorsal
scales are 25. Pharyngeal teeth 2.5-4.3. Gill
rakers number 7, they are short and stubby,
the longest touching the adjacent one when
appressed. Total vertebrae are 40, comprising
20 abdominal and 20 caudal vertebrae.
Predorsal vertebrae number 13.
The upper body profile is convex, similar
to the lower profile. The snout is markedly
rounded, stout. The mouth is small, almost
subterminal; the tip of the mouth cleft is
on a level from the lower margin of the eye.


Description of paratypes. The body is
compressed. The ventral keel between the
pelvics and anal fin is completely scaleless,
very sharp and prominent in all specimens.
The anal fin origin is below the posterior
end of the dorsal fin base. The snout is short
and markedly rounded in smaller and larger
individuals. The mouth is almost subterminal,
with the tip of the mouth cleft on a level
of the lower margin of the eye or below. The
junction of the lower jaw and the quadrate
is on about a vertical through the middle of
the eye.



Dorsal fin unbranched rays 3, branched
dorsal-fin rays 7½ (2), 8½ (48), 9½ (8)
(8.1, 0.41). Anal fin unbranched rays 3,
branched anal-fin rays 10½ (5), 11½ (14),
12½ (29), 13½ (9), 14½ (1) (11.8, 0.88)
(see also Tables 1 and 4 for data based on a
set of another 48 specimens which were radiographed).
The dorsal fin outer margin is
truncate to markedly convex and the anal
fin outer margin is slightly concave. Pectoral
fin branched rays 12(6), 13(33), 14(17),
15(2) (13.3, 0.69), pelvic fin branched rays
6(7), 7(51) (6.9, 0.33).
Pharyngeal tooth counts are 2.5-4.2
(20), 2.4-4.2 (5), 2.5-4.1 (2), 2.5-4.3 (2),
1.5-4.2 (1). The lateral line is complete
with none or 1 unpored scales at the posterior
end of the lateral series; total lateral
line scales 43(1), 44(3), 45(3), 46(11), 47(12), 48(16), 49(8), 50(3), 51(-), 52(1)
(47.3, 1.68); lateral line scales to the margin
of hypurals 42(1), 43(4), 44(5), 45(12),
46(15), 47(10), 48(9), 49(1), 50(-), 51(1)
(46.1, 1.70). Scales around caudal peduncle
14(2), 15(9) 16(13), 17(19), 18(14), 19(1)
(16.6, 1.17); scales between dorsal fin origin
and lateral line 9(4), 10(29), 11(24), 12(-),
13(1) (10.4, 0.70); scales between anal fin
origin and lateral line 4(10), 5(37), 6(10),
7(1) (5.0, 0.65); scales between pelvic fin
origin and lateral line 4(3), 5(31), 6(22),
7(2) (5.4, 0.65), and predorsal scales 18(1),
19(4), 20(13), 21(16), 22(13), 23(4), 24(3),
25(4) (21.3, 1.58). Total gill rakers in the
outer row on first left arch number 5(1),
6(14), 7(26), 8(14), 9(3) (7.0, 0.90).
Vertebral counts were calculated for 48
specimens (including holotype). Total vertebrae
number 39-40(41) (39.7, 0.59) (Tables
2 and 4). Predorsal vertebrae number
(11)12-13(14) (12.2, 0.54) (Tables 2 and
5). Abdominal vertebrae number 19-21with
a mode of 20 (19.8, 0.52) (Tables 3 and
5). Caudal vertebrae number 19-21 (19.9,
0.58) (Tables 3 and 6). The vertebral formulae
are 20+20 (in 21 specimens), 19+20
(10), 20+19 (8), 19+21 (3), 20+21 (3), and
21+19 (2). Thus, the mean difference between
abdominal and caudal counts varies
between +3 and -2 with a mode of 0 (-0.1,
0.92) (Tables 3 and 6).
Many
scales are regenerated in various fish from
this collection, perhaps indicating a traumatic
life.


Comparative remarks. Alburnoides namaki
sp. n. differs from all the congeners primarily
by having a combination of a sharp
scaleles keel, a short markedly rounded
snout, an almost subterminal mouth and a
low number of predorsal vertebrae (modally
12). In tree diagrams based on combined
data (Figs 3-6) A. namaki is clustered
together with A. varentsovi sp. n. from the
northern slope of Kopetdag. Alburnoides
namaki which shares with A. varentsovi sp.
n. (and A. idignensis sp. n.) the lowest number
of predorsal vertebrae (modally 12) is
distinguished by a shallowly forked caudal
fin with rounded lobes (vs. clearly forked,
with pointed lobes), a small, almost subterminal
mouth with the tip of the mouth cleft
on a level from the lower margin of the eye
or below (vs. oblique and terminal, the tip
of the mouth cleft on a level from the middle
of the eye or slightly above), a sharp and
commonly completely scaleless ventral keel
(vs. commonly partly scaled).
 

 

Common names

None.

Systematics

Key characters

 

Morphology

Sexual dimorphism

Colour

Size

 

Distribution

    

    

Zoogeography

 

Habitat

 

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

Unknown.

Conservation

 

Further Work

 

 

Alburnoides nicolausi
Bogutskaya and Coad, 2009

 

?see tables for counts, check agaisnt etxt


Diagnosis. The species is distinguished
by a combination of characters which includes
an eye of an average size, the orbit
diameter larger than the snout length and
smaller than the interorbital width; caudal
fin lobes rounded and fin shallowly forked;
a variably scaled ventral keel though most
commonly scaled only along about 1/3 of
its length or scaleless; a deep head with a
moderately stout snout which is slightly
pointed; a tip of the mouth cleft on the level
about the lower margin of the pupil, commonly
7½ branched dorsal-fin rays; 8-11½
branched anal-fin rays; (43)43-47(48-50)
total lateral line scales (42-48 scales to posterior
margin of hypurals); commonly 2.5-
4.2 or 2.4-4.2 pharyngeal teeth; commonly
39-40 total vertebrae; 12-13 predorsal vertebrae;
19-20(21) abdominal vertebrae;
18-20 caudal vertebrae; a caudal vertebral
region most commonly one vertebra shorter
than the abdominal region; and the most
common vertebral formulae are 20+19,
19+20 and 20+20.

Dorsal fin rays are 3 unbranched and 7½
branched, anal fin rays are 3 unbranched
and 10½ branched, branched pectoral fin
rays are 13, pelvic fin branched rays are 7.
The anal fin origin is slightly behind a vertical
from the posterior end of the dorsal fin
base. Total lateral line scales number 47 and
those to posterior margin of hypurals 45,
scales around caudal peduncle 17, scales
above lateral line to dorsal fin origin are 10,
scales below lateral line to anal fin origin
are 5, scales below lateral line to pelvic fin
origin are 5, and midline predorsal scales
are 19. Pharyngeal teeth 2.5-4.2. Gill rakers
number 8, they are short and stubby, the
longest touching the adjacent one when appressed.
Total vertebrae are 38, comprising
20 abdominal and 18 caudal vertebrae. Predorsal
vertebrae number 12.


Description of paratypes.

 


Dorsal fin unbranched rays 3, branched dorsal-fin rays 7½ (52) and 8½ (7) (7.1, 0.33). Anal fin unbranched rays 3, branched anal-fin rays 8½ (2), 9½ (13), 10½ (32), 11½ (12) (10.0, 0.68) (see also Tables 1 and 4 for 42 radiographed specimens). The dorsal
fin outer margin is commonly truncate,
slightly convex or slightly concave, and the
anal fin outer margin is truncate or only
slightly concave. Pectoral fin branched rays
11(1), 12(30), 13(25), 14(3) (12.5, 0.63),
pelvic fin branched rays 6(6), 7(53) (6.9,
0.30).
Pharyngeal tooth counts are 2.5-4.2
(24), 2.4-4.2 (5), 2.4-5.2 (1). The lateral
line is complete with none, 1 or 2 unpored
scales at the posterior end of the lateral series;
total lateral line scales 42(2), 43(11),
44(15), 45(6), 46(13), 47(7), 48(2), 49(2),
50(2) (45.0, 1.82); lateral line scales to the
margin of hypurals 41(3), 42(14), 43(11),
44(11), 45(8), 46(9), 47(-), 48(2), 49(1)
(43.9, 1.83). Scales around caudal peduncle
13(1), 14(9), 15(30) 16(14), 17(5) (15.2,
0.87); scales between dorsal fin origin and
lateral line 8(2), 9(30), 10(25), 11(2) (9.5,
0.62); scales between anal fin origin and lateral
line 3(1), 4(27), 5(28), 6(3) (4.6, 0.62);
scales between pelvic fin origin and lateral
line 4(21), 5(37), 6(1) (4.7, 0.51), and predorsal
scales 18(4), 19(13), 20(18), 21(10),
22(8), 23(4), 24(2) (20.4, 1.49). Total gill
rakers in the outer row on first left arch
number 5(1), 6(1), 7(33), 8(22), 9(2) (7.4,
0.67).
Vertebral counts given below were calculated
in 42 specimens. Total vertebrae
number 38-40 with a mode of 39 (38.9,
0.58) (Tables 2 and 4). Predorsal vertebrae
number 12-13 (12.6, 0.50) (Tables 2 and 5).
Abdominal vertebrae number 19-21with
a mode of 20 (19.8, 0.53) (Tables 3 and 5).
Caudal vertebrae number 18-20 (19.1, 0.68)
(Tables 3 and 6). The vertebral formulae are
20+19 (in 18 specimens), 19+20 (9), 20+18
(6), 20+20 (4), 19+19 (3), 21+18 (1), and
21+19 (1). Thus, the mean difference between
abdominal and caudal counts varies
between +3 and -1 with a mode of 1 (0.6,
1.08) (Tables 3 and 6).
 

 



 

 

Common names

شبه زوري (shebeh zury = resembling zury) in Khuzestan.

Systematics

The holotype (CMNFI 1979-0281) is a female, 75.0 mm SL, Lorestan, stream in Simareh River drainage, 5 km south of Nurabad (34°03´30´´N, 47°58´30´´E) and paratypes (CMNFI 1979-0281A) comprise 164 specimens, 21.3-65.0 mm SL, same data as holotype. The species is named after a Latin male name Nicolaus, a derivative of the Greek Nikolaos (victory of the people), a compound name composed of the elements nikē (victory) and laos (the people); a Russian name Nikolay and an English name Nicholas, the names of, respectively, Nina Bogutskaya’s elder son and Brian Coad’s son, are also derivatives from Nicolaus.

Key characters

This species differs from all the congeners primarily by having a combination of commonly 7 branched dorsal-fin rays, 8-11 branched anal-fin rays, and 38-40, modally 39, total vertebrae.

Morphology

The body is moderately compressed, relatively thick. The upper body profile is convex similar to the lower profile. The snout is only slightly rounded, almost pointed. The mouth is oblique, slightly below than terminal; the tip of the mouth cleft is slightly below a level of the lower margin of the pupil. The junction of the lower jaw and the quadrate is on about a vertical through the middle of the eye. The caudal fin lobes are rounded, the fin is shallowly forked. A ventral keel between the pelvics and the anal fin is not sharp and is variably scaled: completely scaleless (9), scaled along about ¼-1/3 of its length (9), scaled along  half of its length (6), scaled along about two-thirds of its length (4) or completely scaled (2).  There is a pelvic axillary scale and scales extend over the  proximal bases of the anal fin forming a sheath. The anal fin origin is somewhat behind a vertical from the posterior end of the dorsal fin base.
 

Sexual dimorphism

The following characters were larger in females: pectoral fin origin to pelvic fin origin distance, pelvic fin origin to anal fin origin distance, prepelvic fin length, and mouth width while the following were larger in males: caudal peduncle length, pectoral fin length, pelvic fin length, longest dorsal fin ray length, longest anal fin ray length, pectoral fin length in pectoral fin origin to pelvic fin origin distance, and pelvic fin length in pelvic fin origin to anal fin origin distance.

Colour

?harmonise

The lateral line is delineated by some darker pigment above and below but not as strongly as in A. petrubanarescui holotype and obscured by background pigmentation on the caudal peduncle. Some pigment on the flank scales above and below the lateral line is weak or irregular and an impression of stripes is not very evident. The mid-flank stripe is weak and diffuse, fading anteriorly under the dorsal fin. The back is dark but predorsal and postdorsal stripes are evident. The fins are mostly immaculate, with some melanophores lining rays of the dorsal and pectoral fins in particular. The unbranched pectoral fin ray is lined with melanophores on its inner margin, but not as strongly as in some other samples.

Paratypes can bear strong pigmentation above and below the lateral line pores, forming an evident pale line margined with dark, or this pattern may be quite faint. The mid-flank stripe is weak or diffuse and fades anteriorly. A thin line of pigment can be evident separating the hypaxial and epaxial muscle masses, fading anteriorly. The pigment on scales above and below the lateral line (and below the mid-flank stripe) can  be obvious and form a series of thin, discontinuous stripes, or it can be absent. Some fish have a series of strong melanophores on the inner margin of the pectoral fin unbranched ray. The peritoneum is silvery with fine melanophores and some spots.

Size

Distribution

The species is known only from its type locality, a stream in the Simareh River drainage at Nurabad. The Simareh (Seymareh) flows into the Karkheh (Qareh Su) River which enters the Hawr al Hawizeh (Hawr al Azim) on the Iran-Iraq border (Tigris River drainage).          

Zoogeography

In tree diagrams (Bogutksaya and Coad, 2009) based on combined data, this species it is clustered together with another Tigris River basin species, A. idignensis. ??

Habitat

Habitat data is based on collection data. Fish were collected at 2000 m altitude, 19°C water temperature, clear water, pH 6.8, forested shore, stony river bed, moderate amounts of aquatic plants, and no other species taken.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

Unknown.

Conservation

See under A. eichwaldii.

Further Work

This recently described species needs its conservation status and biology investigated.

Sources

Alburnoides petrubanarescui
Bogutskaya and Coad, 2009

 

?see tables for counts, check against etxt

Holotype. CMNFI 1970-0558, female, 109.1 mm TL, 88.8 mm SL; Iran, Azarbaijan-e Bakhtari, Qasemlou Chay, Orumiyeh [Urmia] Lake basin, ca. 37°21´N, 45°09´E; 27 June 1962; coll. V.D. Vladykov. Paratypes. CMNFI 1970-0558A, 51, 28.7-87.3 mm SL, counts and measurements on 29 fish 33.6-87.3 mm SL; same data as holotype.

Diagnosis. The species is distinguished by a combination of characters which includes a small eye; the orbit width about equal to the snout length but markedly smaller than the interorbital width; caudal fin lobes rounded and the fin shallowly forked; a scaled ventral keel behind the pelvic fins along the abdomen to the anus, a deep head with a stout snout which is markedly rounded; a tip of the mouth cleft on the level below the lower margin of the eye; commonly 7½ (less frequently 8½) branched dorsal-fin rays; 8-10½, commonly 9½, branched anal-fin rays; 44-51 total lateral line scales (42-49 scales to posterior margin of hypurals); 2.5- 4.2 pharyngeal teeth (or other variants with four teeth on the right ceratobranchial); commonly 40-41 total vertebrae; 13-14 predorsal vertebrae; 20-22, commonly 21, abdominal vertebrae; 19-20 caudal vertebrae; a caudal vertebral region most commonly
shorter than the abdominal region; and the most common vertebral formulae are 21+19 and 21+20.
Description of holotype. The caudal fin lobes are rounded and the fin is shallowly forked. A ventral keel between the pelvics and the anal fin is smooth and completely scaled. There is a pelvic axillary scale and scales extend over the proximal bases of the anal fin forming a sheath. The upper body profile is convex, similar to the lower profile. The snout is markedly rounded, stout. The mouth is small, subterminal; the tip of the mouth cleft is on a level below the lower margin of the eye. The body depth enters SL 3.3 times, HL enters 4.3, predorsal length 1.8, caudal peduncle depth 7.7, caudal peduncle length 4.1, length of longest dorsal fin ray 5.2, and length of longest anal fin ray to scale sheath 6.8. Orbit diameter enters HL 3.5 times, snout length enters 3.6, and interorbital width 2.6. Pectoral fin length enters pectoral fin origin to pelvic fin origin distance 1.3 times, and pelvic fin length
enters pelvic fin origin to anal fin origin distance 1.2 times. Dorsal fin rays are 3 unbranched and 7½ branched, anal fin rays are 3 unbranched and 9½ branched, branched pectoral fin rays are 13, pelvic fin branched rays are 7. The anal fin origin is on a vertical from the posterior end of the dorsal fin base. Total lateral line scales number 46 and those to posterior margin of hypurals 45, scales around caudal peduncle 15, scales above lateral line to dorsal fin origin are 9, scales below lateral line to anal fin origin are 5, scales below lateral line to pelvic fin origin are 6, and midline predorsal scales are 21. Pharyngeal teeth 2.5-4.2. Gill rakers number 7, they are short
and stubby, the longest touching the adjacent one when appressed. Total vertebrae are 41, comprising 21 abdominal and 20 caudal vertebrae. Predorsal vertebrae number 13.
The peritoneum is silvery with fine melanophores and some large spots. The lateral line is clearly delineated by darker pigment above and below. Some pigment on flank scales above and below the lateral line give the impression of stripes. A mid-flank stripe is evident. The back is dark and obscures a predorsal and postdorsal stripe. The fins are mostly immaculate, with some melanophores lining the rays of the dorsal and pectoral
fins. The unbranched pectoral fin ray is strongly lined with melanophores on its inner margin.

Description of paratypes.
The body is compressed but relatively thick. The ventral keel between the pelvics and anal fin is not sharp and is completely covered by scales in all specimens. The anal fin origin is below the posterior end of the dorsal fin base. The snout is short and markedly rounded in smaller and larger individuals. The mouth is subterminal, with the tip of the mouth cleft on a level below the lower margin of the eye. The junction of the lower jaw and the quadrate is on about a vertical through the anterior eye margin. The following characters were significantly different between sexes (p<0.05). Greater in females: postorbital length, predorsal length, pectoral fin origin to pelvic fin origin distance, pelvic fin origin to anal fin origin distance. Greater in males: HL, pectoral fin length in pectoral fin origin to pelvic fin origin distance, and pelvic fin length in pelvic fin origin to anal fin origin distance. Dorsal fin unbranched rays 3, branched dorsal-fin rays 7½ (19) or 8½ (10) (7.3, 0.48). Anal fin unbranched rays 3, branched anal-fin rays 8-10½ (9.3, 0.64, including holotype) (Tables 1 and 4). The dorsal fin outer margin is truncate to markedly convex and the anal fin outer margin is slightly concave. Pectoral fin branched rays 13(16), 14(12), 15(1) (13.5, 0.57), pelvic fin branched rays
6(5), 7(24) (6.8, 0.38) Pharyngeal tooth counts are 2.5-4.2 (18), 2.4-4.2 (4), 2.5-4.1 (5), 1.4-4.1 (1), 1.5-4.0 (1). The lateral line is complete with none or 1 unpored scales at the posterior end of the lateral series; total lateral
line scales 43(1), 44(3), 45(2), 46(8), 47(5), 48(6), 49(3), 50(1) (46.7, 1.73); lateral line scales to the margin of hypurals 42(1), 43(3), 44(4), 45(5), 46(6), 47(6), 48(3), 49(1) (45.6, 1.76). Scales around caudal peduncle 14(2), 15(7) 16(10), 17(9), 18(-), 19(1) (16.0, 1.09); scales between dorsal fin origin and lateral line 9(7), 10(18), 11(4) (9.9, 0.62); scales between anal fin origin and lateral line 4(5), 5(23), 6(1) (4.9, 0.44); scales between pelvic fin origin and lateral line 3(1), 4(8), 5(20) (4.7, 0.55); predorsal scales 20(1), 21(10), 22(14), 23(3), 24(1) (21.8, 0.83). Total gill rakers in the outer row on first left arch number 6(3), 7(18), 8(8) (7.2, 0.60). Vertebral counts given below include holotype. Total vertebrae number (39)40-41(42) (40.5, 0.63) (Tables 2 and 4). Predorsal vertebrae number 13-14 with a mode of 13 (13.4, 0.50) (Tables 2 and 5). Abdominal vertebrae number 20-22 with a mode of 21 (21.0, 0.41) (Tables 3, 5). Caudal vertebrae number 19-20(21) (19.5, 0.57) (Tables 3 and 6). The vertebral formulae are 21+19 (in 12 specimens), 21+20 (10), 22+19 (3), 20+19 (1), and 21+21 (1). Thus, the caudal vertebral region is shorter than the abdominal region, rarely equal to it (in 3 specimens), the mean difference between abdominal and caudal counts being +1.4 (std 0.77) (Tables 3 and 6). Other characters as in holotype.
Most paratypes bear strong pigmentation above and below the lateral line pores, forming an evident pale line margined with dark. The broad mid-flank stripe is well-developed. The pigment on scales above and below the lateral line (and below the midflank stripe) form a series of thin, discontinuous stripes. Some fish have a series of strong melanophores on the inner margin of the pectoral fin unbranched ray. The lateral line over the pectoral and pelvic fins can be wavy rather than a smooth decurved line.

Comparative remarks. Alburnoides petrubanarescui sp. n. differs from all the congeners primarily by having a combination of a scaled keel, the lowest number of branched anal-fin rays (modal value 9½ vs. 10½ and more), and the highest value of the difference between the abdominal and vertebral counts. A completely scaled keel is a character shared by A. petrubanarescui sp. n., A. oblongus distributed in the lower reaches of the Syr Darya and Alburnoides sp. from Pulvar (Kor River drainage). However, A. petrubanarescui sp. n. is distinguished from the two other species of this group by having fewer branched dorsal-fin rays (commonly 7½ vs. 8½) and fewer branched anal-fin rays (8-10½ vs. 10-12½). Besides, A. petrubanarescui sp. n. is clearly different from A. oblongus by having larger scales (43-50 total lateral line scales vs. 50-56), 2.5-4.2 and 2.4-4.2 pharyngeal teeth (vs.
2.5-5.2 or 1.5-5.1), fewer gill rakers (6-9 vs. 10-13), a truncate or rounded margin of the dorsal fin (vs. concave). A. petrubanarescui sp. n. differs from Alburnoides sp. from Pulvar, besides some other characters, by fewer dorsal-fin branched rays (commonly 7½ vs. 8½), fewer anal-fin branched rays (8-10½, commonly 9½, vs. 10-12½, commonly 11½) and 21+19 or 21+20 vertebrae (vs. 20+20 or 20+21) the difference between abdominal and caudal counts averaging +1.4 (vs. -0.3).
Etymology. The species is named after the late Petru Bǎnǎrescu, a great freshwater ichthyologist who contributed significantly to our knowledge of fishes of Eurasia.

Distribution. This species is described from a river in the Orumiyeh [Urmia] lake basin and we suppose that it may be an endemic species to the Orumiyeh lake basin.

Habitat data for the type locality (June 1962): water 18 °C, fast current in stream, pebbles and sand bottom, shore grassy, much aquatic plant life, caught with dipnet, other species included Alburnus atropatenae, Barbus
lacerta, “Nemacheilus” sp.

Common names

None.

Systematics

Key characters

Morphology

Sexual dimorphism

Colour

Size

Distribution

    

    

Zoogeography

Habitat

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

Unknown.

Conservation

See under A. eichwaldii.

Further Work

This recently described species needs its conservation status and biology investigated.

Sources

Alburnoides qanati
Coad and Bogutskaya, 2009

Common names

None.

Systematics

The female holotype is in the Canadian Museum of Nature, Ottawa, under CMNFI 1977-0509, 81.5 mm TL, 65.0 mm SL, Fars, at source and along stream of a qanat at Naqsh-e Rostam, Pulvar River system (29°59’30”N, 52°54’00’’E). Paratypes are under CMNFI 1977-0510, 168 (not 178 as in type description) specimens, 24.9-72.5 mm SL, same data as holotype. The species was named after the famous qanat system which taps groundwater to support human survival in desert regions and, incidentally, a habitat for fishes.

Key characters

The species is distinguished by a combination of characters which includes a large eye, the orbit width exceeding both the snout length and the interorbital width, a scaled ventral keel behind the pelvic fins along the abdomen to the anus, commonly 43-47 lateral line scales to posterior margin of hypurals, 2.5-4.2 pharyngeal teeth, commonly 8 branched dorsal fin rays, 10-12 branched anal fin rays, 40-41 total vertebrae, an d the caudal vertebral region equal or longer then the abdominal region (vertebral formulae 20+20 or 20+21).

Morphology

The body is markedly compressed. The upper body profile is convex or, in larger specimens, slightly to markedly straightened while the lower profile is considerably convex. The ventral keel between the pelvics and anal fin is not sharp and is completely covered by scales in all specimens but four possessing a short scaleless portion of keel (about half of keel length) just in front of the anus. The dorsal fin outer margin is truncate to slightly rounded and the anal fin outer margin is truncate to slightly concave. The anal fin origin is behind the posterior end of the dorsal fin base. A pelvic axillary scale is present and the anal fin base is proximally overlain by flank scales. The snout is short and slightly pointed. The mouth is terminal to upturned, with the tip of the mouth cleft on a level from slightly above the middle of the eye to the upper margin of the pupil. The mouth cleft is always turned upward, never horizontal, the lower jaw slightly to moderately projecting relative to the upper jaw, and the junction of the lower jaw and the quadrate is on about a vertical through the anterior eye margin. The lateral line is decurved and only the last few scales are elevated and on the mid-caudal peduncle.

Dorsal fin unbranched rays commonly 3, 4 in 3 specimens only, dorsal fin branched rays 7(3) or 8(28), anal fin unbranched rays 3, anal fin branched rays 10(3), 11(22), 12(6), branched pectoral fin rays 13(4), 14(20) or 15(7), pelvic fin branched rays 7(30).  The lateral line is complete with none, 1 or 2 unpored scales at the posterior end of the lateral series. Lateral line scales to posterior margin of hypurals 41(1), 42(1), 43(5), 44(6), 45(3), 46(8), 47(5) 48(1) or 49(1),  scales above lateral line to dorsal fin origin 9(10), 10(18) or 11(3), scales below lateral line to pelvic fin origin 3(4), 4(20) or 5(7), and scales below lateral line to anal fin origin 4(17), 5(13) or 6(1). Total scale radii 8(1), 9(1), 10(4), 11(8), 12(20), 13(17), 14(16) 15(12), 16(7), 17(3) or 18(1) (13.2, 1.91). Scale radii are restricted to the posterior field encroaching laterally, circuli are eccentric and the focus is anteriorly located. Total gill rakers in the outer row on first left arch 6(4), 7(4), 8(21) or 9(1); gill rakers are very short and widely spaced, not touching the adjacent raker when appressed. Total vertebrae including 4 Weberian vertebrae and last complex centrum 40(14) or 41(17), abdominal vertebrae (including intermediate ones; precaudal vertebrae auctorum) 20 (29) or 21(12), predorsal vertebrae (anterior to first dorsal pterygiophore) 13(24) or 14(6), and caudal vertebrae 20(18) or 21(13). The vertebral formula is 20+20(16), 20+21(12) or 21+20(2). Thus, the caudal vertebral region most commonly (in 93% of examined specimens) is equal to or slightly longer then the abdominal region, the mean difference between abdominal and caudal counts being -0.3. Pharyngeal tooth counts are 2.5-4.2 in 10 fish examined with one additional fish being a variant with 2.4-4.0. Teeth are hooked at the tip and not serrated below it. The gut shape is a simple “S” with an occasional specimen showing a slight flexure to the left of the anterior loop. The general topography of cephalic sensory canals and numbers of pores is typical of most Alburnoides, as described by Bogutskaya (1988). The supraorbital canal is not lengthened in its posterior section and has 7-11, commonly 8-10 pores, with 2-4 (3 in 90%) and 5-7 (6 in 73%) canal openings on the nasal and frontal bones, respectively. The infraorbital canal has 10-15 pores (13 in 38%, 12 in 30%) with 4 (93%) or 5 canal openings on the first infraorbital. The preopercular-mandibular canal is complete, with 11-17, modally 13-16, pores (14 in 38%) with (3)4-6 (5 in 77%) and 7-10 (8 in 62%) canal openings on the dentary and preoperculum, respectively. The supratemporal canal is complete, with (4)5-7 (7 in 54%) pores.

Sexual dimorphism

Head length is longer in males than in females. Pectoral fin length and pelvic fin length are also longer in males.

Colour

Pigmentation of the holotype in 5% formalin consisted of a dark lateral line dividing the hypaxial and epaxial muscle masses and a weakly developed stripe of black pigment on mid-flank prominent posteriorly on the caudal peduncle but fading over the pectoral fin and often interrupted anteriorly. The lateral line pores were lined by pigment dorsally and ventrally. A mid-dorsal line was apparent before the dorsal fin, weakly developed behind the fin. The fins were mostly hyaline with some black pigment lining the fin rays of the dorsal and caudal fins, the dorsal rays of the pectoral fins and the anterior rays of the anal fin.

Overall colouration is silvery with the bases of the pectoral, pelvic and anal fins pink in life. An orange line parallels the anal fin base and the lateral line, lying midway between the two. The ventral surface of the head between the dentaries may be yellow-orange and similarly coloured spots may be found on either side of the dorsal mid-line extending along the whole body. Faint yellow spots occur in rows along the flanks also. Pigmentation in preserved fish is as described for the holotype although the lateral stripe is weakly-developed in some specimens, the mid-flank band of spots of black pigment may be variably developed, and the lateral line may be clearly or only faintly edged by pigment. The peritoneum is rarely dark brown but usually is white-grey to light brown with black spots.

Size

Attains 72.5 mm standard length.

Distribution

Known originally from the Pulvar River drainage of the Kor River basin in southern Iran but also recorded from Harat in the Sirjan basin at 30º01.196'N, 54º20.33'E (material from H. R Esmaeili, 2011). 

Zoogeography

This is the southernmost Alburnoides species and may have entered the Kor River basin by headwater capture from the Tigris-Euphrates River basin.

Habitat

The qanat stream in the Pulvar River basin at 15.00 hours on 6 October 1976 had clear and colourless water, a temperature of 21°C, pH 6.8, conductivity 0.475 mS, the current was slow to medium, stream width was about 2 m and maximum depth was up to 1 m, the shore was grassy, plant life in the stream consisted of encrusting and submergent types, and the stream bed was gravel and mud. The Harat locality was at an altitude of 1585 m, pH 8.17, dissolved oxygen 7.25 mg/l, conductivity 816 mS and temperature 22.9-23.3°C.

Type locality, mouth of qanat and stream origin. Photo: Brian W. Coad.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

None.

Conservation

The numbers and wider distribution of this species should be researched as it is known from only two localities.

Further Work

See under Conservation. Biology is unknown.

Sources

Type material: See above.

Other material: CMNFI 1979-0060, 4, 21.0-35.4 mm SL, Fars, spring and irrigation channel, 7 km north of Sa’adatabad (30°06’N, 53°12’E).

Alburnoides taeniatus
(Kessler, 1874)

Reported from the Tedzhen River basin (Aliev et al., 1988), Karakum Canal, Kopetdag Reservoir and Uzboi lakes (Shakirova and Sukhanova, 1994; Sal'nikov, 1995) in Turkmenistan on the northeastern border of Iran. It may eventually reach the Caspian Sea basin and the Tedzhen (= Hari) River basin of Iran. No Iranian record.

Genus Alburnus
Rafinesque, 1820

The bleaks and shemayas are found in Europe and the northern parts of Southwest Asia with about 38 species (depending on definitions of the taxa). There are 5 species in Iran. Records of Alburnus orontis Sauvage, 1882 from Iran by Armantrout (1969), Banarescu (1977) and Wossughi (1978) are in error (Krupp, 1985c). Chalcalburnus Berg, 1933 is now regarded as a synonym of Alburnus Rafinesque, 1820. There have been numerous variant views of this synonymy. Bogutskaya (1990) considers Chalcalburnus to be distinct but later, Bogutskaya (1997b; Bogutskaya et al., 2000; Bogutskaya and Naseka, 2004), synonymises it with Alburnus. Reshetnikov et al. (1997) retain Chalcalburnus as a distinct genus as does Eschmeyer in "Catalog of Fishes" (downloaded, 10 August 2007). Banister (1980) points out that the distinction of the genus from Alburnus is based on the relative lengths of the ventral keel and the relative thickness of the last unbranched dorsal fin ray, characters which he views with suspicion in the absence of other corroborating evidence.

This genus is characterised by an elongate, compressed, moderately deep body of small to moderate size, a terminal mouth, no barbels, scales of moderate size, pharyngeal teeth in 2 rows (2,5-5,2 or 2,5-4,2) with hooked tips and usually serrations (often absent), short dorsal fin without a thickened ray, a long anal fin, long and relatively numerous gill rakers, a fleshy keel between the base of the pelvic fins and the vent (the naked part usually not reaching as far forward as the pelvic fin bases in species formerly placed in Chalcalburnus), and a light to brown or black peritoneum. Some authors consider the genus Alburnoides to be synonyms of Alburnus (e.g. Saadati (1977)) while others disagree (e.g. Bogutskaya (1990)). These genera are treated separately here to accord with common usage in Southwest Asia, a conservative measure when there are conflicting opinions.

Jalali et al. (2002) and Jalali and Barzegar (2006) record several parasites from an undescribed Chalcalburnus species in Lake Zarivar, namely Ichthyophirius multifilis, two species of Argulus, a Trichodina species, Dactylogyrus alatus, Diplostomum spathaceum, Myxobolus molnari and Ligula intestinalis. Masoumian et al. (2007) record the myxosporean parasite Myxobolus saidovi from Alburnus maculatus (sic) in the Zayandeh River and Mehdipoor et al. (2004) record the monogenean Dactylogyrus alatus from Alburnus maculatus (sic), also in the Zayandeh River..

Alburnus doriae de Filippi, 1865 has a type locality of "dintorni di Schiraz" but fish resembling this species have not been caught there in late twentieth and early twenty-first century collections. Krupp (1985c) refers 5 specimens from the type series of Alburnus doriae to his Alburnus sellal and 2 specimens to Squalius lepidus. The lectotype (MZUT N.720 or MZUT P1110) of Alburnus doriae is stored in the Istituto e Museo di Zoologia della R. Università di Torino (122.0 mm standard length as measured by me) and 5 paralectotypes (MSNG C.E. 9102) of this nominal species are in the Museo Civico di Storia Naturale di Genova (Tortonese, 1934; 1940; 1961), only one of which is A. doriae (109.1 mm standard length as measured by me). Eschmeyer's "Catalog of Fishes" (downloaded 10 August 2007) has 6 specimens in MSNG C.E. 9102, 5 not this species and gives a locality as probably south of Shiraz.

It seems probable that the fish were collected north of Shiraz, presumably in a Tigris River basin stream based on the other species included in the jar (although Alburnus sellal is more likely to be A. mossulensis, q.v.). These materials may, however, have been mixed and the type locality of this nominal species is obscure.

The species of Alburnus in the Zagros Mountains north of Shiraz and west of Esfahan are currently under investigation and final species identities cannot be given at present. Note that materials identified by Coad (1982d: Alburnus maculatus; 1985: A. doriae) as Leuciscus lepidus were in error. An illustration of A. doriae is given above based on the type material.

Small fishes and members of the genus Alburnus are called kuli in Farsi. In Gilan, kuli are eaten with their heads on and are said to full of phosphorus, conferring open-mindedness, intelligence and sophistication on the Gilanis.

Alburnus atropatenae
Berg, 1925

Common names

None.

Systematics

The type series is the material called Alburnus filippii by Günther (1899) from "Sujbulak and Superghan near the mouth of the Nazlu Chai" as noted in Berg (1925). This material is in the Natural History Museum, London under BM(NH) 1899.9.30:127, syntype, 1 specimen, 89.7 mm Sl, Azarbayjan-e Bakhtari, Superghan near the mouth of the Nazlu Chai (Sopurghan on the Nazlu Chay is at 37°45'N, 45°12'E); BM(NH) 1899.9.30:128-30, syntypes, 3, 70.7-96.3 mm Sl, Azarbayjan-e Bakhtari, Tatawa Chai near Sujbulak (the Tata'u Chay or Simineh River is not close to Saujbulagh or Mahabad at 36°45'N, 45°43'E so the exact locality of this collection is unclear).

These syntypes bear an external label, apparently in A. Günther's handwriting, listing these fish under the name "brevianalis" which is crossed out and filippii substituted. It appears that Günther originally intended to describe them as distinct and subsequently changed his mind.

Berg's (1925) material was not found in a search of the collections of the Zoological Institute, St. Petersburg (ZISP) in November 1993. Eschmeyer et al. (1996) give the following data: Syntypes: (46) ZIL (ZIL being the old acronym for ZISP) but this material is presumably comparative specimens mentioned by Berg (1925).

Coad and Holčík (1999) demonstrated variation between three populations isolated by the salt Lake Orumiyeh but considered this variation as insufficiently different to warrant taxonomic distinction. Nonetheless, the analysis demonstrated that the three populations have diverged in a measurable manner, presumably through geographical isolation, although ecological factors may have played a part as one sample was from a lacustrine rather than a riverine environment.

Key characters

This species is distinguished from its relatives in the former genus Chalcalburnus (having a short, naked ventral keel) by a combination of characters:-

Species

Total gill rakers

Branched anal fin rays

Pored scales in lateral line

Peritoneum colour

atropatenae

11-16

9-12

46-63

black

chalcoides

18-31

12-19

54-74

light brown

mossulensis

11-18

10-14

58-89

brown to black

tarichi (Lake Van, Turkey)

26-29

9-11

65-82

light brown

Morphology

Dorsal fin rays branched 7-9, modally 8, after 3 unbranched rays, anal fin branched rays 9-12 after 3 unbranched rays, pectoral fin branched rays 13-16 and pelvic fin branched rays 7-8. Lateral line scales 46-63. There is a pelvic axillary scale. The scale focus is slightly anterior or central and there are relatively few anterior and posterior radii about equal in number. The exposed fleshy keel in front of the anus is about 1-4 scales lengths, usually 2, long. Gill rakers lanceolate but short, less than half eye width, reaching between the first and second adjacent rakers or touching the second when appressed, total numbering 11-16. Pharyngeal teeth are hooked at the tip and usually bear a few, large serrations on the larger major row teeth or more rarely have no serrations, apparently size independent. The posteriormost major row tooth may be dorsal rather than posterior to the tooth ahead of it. Tooth counts are usually 2,5-4,2. The gut is an elongate s-shape, sometimes with an anterior loop to the left. Total vertebrae 41-43.

Meristic values for Iranian material: dorsal fin branched rays 7(2), 8(102) or 9(1); anal fin branched rays 9(5), 10(49), 11(45) or 12(6); pectoral fin branched rays 13(7), 14(44), 15(41) or 16(13); pelvic fin branched rays 7(17) or 8(88); lateral line scales 46(4), 47(5), 48(12), 49(15), 50(13), 51(15), 52(14), 53(5), 54(5), 55(5), 56(2), 58(6) or 63(1); total gill rakers 11(12), 12(30), 13(35), 14(16), 15(7) or 16(2); pharyngeal tooth counts 2,5-4,2(54), 2,4-4,2(2), 2,4-5,2(1), 2,5-5,2(1), 1,5-4,2(1) or 2,5-3,2(1); and total vertebrae 41(4), 42(12) or 43(3).

Sexual dimorphism

Male specimens have small scattered tubercles on the top of the head with fewer tubercles on the side of the head. Tubercles are variably distributed on the head depending on the specimen, or even be different on each side of a single fish. A distinct row may parallel the upper lip, another row may follow the upper eye margin, a patch may be present between the nostril and the upper lip, and there may be tubercles between the mouth and the eye. Very small tubercles line the scale margins on the back, flank and belly and belly scales have a fine row of tubercles on the scale base. Tubercles line the rays of the pectoral, dorsal, pelvic and anal fins and weakly on the caudal fin, the rows branching with the fin rays.

Colour

The back is a dark olive brown to grey, with a narrow stripe. The flank has a dark stripe, as wide as the pupil of the eye, extending onto the head as far as the eye and back to the middle of the caudal fin. The stripe is black to dark green. The flank above the stripe is often lighter in contrast to the darker back and accentuates the distinctiveness of the stripe. The flank below this stripe, the belly and the lower head are silvery, and the stripe is clearly set off from the lower flank. The front of the lower jaw is dark and some of this pigment extends into the floor of the mouth. The iris is silvery on the lower half and dark above. The dorsal fin is faintly pigmented grey along its rays, the caudal fin is grey and the other fins are colourless. Melanophores are present on the dorsal and caudal fin rays and the anterior rays of the pectoral, pelvic and anal fin rays. The nostrils may be dark. The peritoneum is black.

Size

Reaches 21.8 cm.

Distribution

This species is endemic to the Lake Orumiyeh basin and is recorded from the Kazim-chai, Ozband River, Talkheh, Zarrineh and Tatavi rivers (Günther, 1899; Berg, 1925; Abdoli, 2000).

Zoogeography

Lake Orumiyeh was formed during the late Pliocene-Pleistocene, lies at 1275-1295 m, and may well have had a Pleistocene connection to the Caspian Sea basin although this is in dispute (Scharlu, 1968; Schweizer, 1975). Pleistocene shorelines from 30 to 115 m above the present level have been confirmed, and the lake covered twice its present area, but this would not permit an external discharge. Berg (1940) reports benches at levels of about 1800 m, 1650-1550 m and 1500-1360 m, which may represent shorelines, and a level of about 1570 m would have had an outlet to the Aras River basin through the Kara-tepe Pass in the northwest and across the plain near the city of Khvoy. Saadati (1977) suggests two connections with the Caspian Sea, an early one in the Pliocene to early Pleistocene resulting in endemic species and a later one in the late Pleistocene resulting in species which are the same as the Caspian or only subspecifically distinct. A. atropatenae may have its origin in the earlier transgression.

Habitat

Unknown.

Age and growth

Unknown.

Food

Gut contents are insects, crustaceans and worms. Filamentous algae are also present, possibly as accidental inclusions.

Reproduction

Fish captured 25-27 June carried mature eggs.

Parasites and predators

None reported from Iran.

Economic importance

Unknown.

Conservation

Biology is poorly known and numbers and habitat requirements would have to be examined for a conservation assessment.

Further work

The biology of this species requires a detailed study.

Sources

Type material. See above, Alburnus atropatenae (BM(NH) 1899.9.30:127, 1899.9.30:128-30).

Iranian material: CMNFI 1970-0557, 26, 17.9-31.6 mm standard length, Azarbayjan-e Bakhtari, Shaher Chay (ca. 37º27'N, ca. 44º55'E); CMNFI 1970-0558, 8, 25.0-88.7 mm standard length, Azarbayjan- e Bakhtari, Qasemlu Chay (ca. 37º21'N, ca. 45º09'E); CMNFI 1970-0559, 48, 31.4-85.2 mm standard length, Azarbayjan-e Bakhtari, Baranduz Chay (37º25'N, 45º10'E); CMNFI 1979-0785, 11, 72.6-123.8 mm standard length, Azarbayjan-e Bakhtari, Shaher Chay (ca. 37º27'N, ca. 44º55'E); CMNFI 1979-0786, 26, 65.0-92.2 mm standard length, Azarbayjan-e Khavari, Guru Lake (37º55'N, 46º24'E); CMNFI 2007-0096, 1, 54.7 mm standard length, Azarbayjan-e Bakhtari, Qasemlu River in Baranduz Chay basin (ca. 37º25'N, ca. 45º10'E); CMNFI 2007-0097, 2, 42.0-54.9 mm standard length, Azarbayjan-e Bakhtari, Baranduz Chay basin (ca. 37º16'N, ca. 45º08'E); CMNFI 2007-0103, 6, 43.3-73.3 mm standard length, Kordestan, Zarrineh River basin (ca. 36º18'N, ca. 46º16'E); CMNFI 2007-0105, 6, 67.3-112.1 mm standard length, Kordestan, Zarrineh River basin (ca. 36º06'N, ca. 46º20'E); OSU 8122, 2, 73.1-83.5 mm standard length, Azarbayjan-e Bakhtari, Shaher Chay (ca. 37º27'N, ca. 44º55'E); USNM 205904, 2, 73.0-82.6 mm standard length, Azarbayjan-e Bakhtari, Nazlu Chay (37º40'N, 45º05'E); uncatalogued, 1, 81.6 mm standard length, Azarbayjan-e Bakhtari, Haladj River near Mahabad (ca. 36º45'N, ca. 45º43'E) (Coad and Holčík, 1999).

?check against Iraq book

Alburnus caeruleus
Heckel, 1843

Common names

None.

Systematics

The type locality is Aleppo (= Halab), Syria and material is held inthe Naturhistorisches Museum Wien. Syntypes are listed in Eschmeyer et al. (1996) as NMW 16688 (4, 65.7-86.6 mm standard length as measured by me), NMW 55511-13 (2, 64.5-75.4 mm standard length, 2, 61.2-71.1 mm standard length, 2, 74.1-77.4 mm standard length), 57161 (3, 59.6-71.1 mm standard length) and additionally ?RMNH 2656 [ex NMW] (4); SMF 100 [ex NMW] (4, 61.3-75.7 mm standard length. See also below.

Key characters

Distinguished from its relatives by ? fewer scales along the lateral line (45-58 compared to 60-89) and a deeper body (2.9-3.5 in standard length compared to 4.0-5.1).

Morphology

Dorsal fin with 3 unbranched and 8-9 branched rays, usually 8, anal fin with 3 unbranched and 13-18 branched rays, mostly 14-16, pectoral fin rays 12-15 and pelvic fin rays 7-8. Lateral line moderately to strongly decurved, scales 43-58. Scales lack radii on the anterior field. The naked ventral keel is obvious. Pharyngeal teeth hooked at tip and deeply notched or serrated below. Modally 2,5-4,2, with variants 2,5-5,2, and 2,5-4,1.Total gill rakers 10-13, just reaching past adjacent raker when appressed. Total vertebrae 39 (Bogutskaya et al., 2000). The body is relatively deep with a slight nuchal hump, 2.9-3.5 times in standard length. The gut is s-shaped.

Sexual dimorphism

Males have tubercles on the lower jaw, the sides and dorsal surface of the head and on flank scales. Tubercles are evident on the pectoral fin and appear as traces on the pelvic fins.

Colour

Back blackish, flanks silvery. Horizontal stripe along flank sky-blue, more diffuse in larger fish but very evident in smaller ones. Flanks, even lower flanks, and head heavily speckled. The lateral line may bear pigment spots above and below each pore but the stitched effect is not as marked as in some Alburnoides species. Fins generally yellowish, dorsal, anal and pelvic fins apically black to sky blue. The membranes of the dorsal and anal fins are heavily pigmented while the rays are clearer. This pigmentation is more evident anteriorly on small fish but in both large and small fish fins appear dark, especially when the fins are collapsed. On the anal fin, some fish have dark pigment on all membranes, others, even large fish, have less pigmentation distally on the posterior membranes. In larger fish, the pectoral and pelvic fins have dark membranes, the pigmentation fading on the smaller rays. The pectoral and pelvic fins can be orange. In some specimens the edge of the caudal fin is quite dark. The peritoneum is brown to black.

Size

Attains 86.9 mm standard length.

Distribution

Found in the Tigris-Euphrates and Quwayq River systems. The Orontes (= Asi) River is not a locality (Krupp, 1985c). In Iran, it is recorded by Keyvan Abbasi (Iranian Fisheries Research Organization Newsletter, 57:2, 2009) from the Gamasiab and Doab rivers (34º22'16"N, 47º54'51"E at 1412 m altitude and 34º27'11"N, 47º39'34"E at 1322 m) and, given the fishing effort, were quite rare (0.02% of fishing sites, 8 individuals). It may be more widely distributed than museum and literature records suggest.

Zoogeography

The relationships of this species zoogeographically have not been studied.

Habitat

Khalifa (1989) reported this species as widely distributed in rivers and ponds, and it is also found in streams, dams and reservoirs in Iraq. Epler et al. (2001) found it to be the third most dominant species of fish in the Iraqi lakes Habbaniyah, Tharthar and Razzazah, comprising 8.7% of all fish collected.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Large eggs were visible in fish from Syria caught on 19 May, suggesting spring spawning.

Parasites and predators

Unknown in Iran.

Economic importance

None.

Conservation

This species is poorly known and documented in Iran so its conservation status is unknown.

Further work

The biology, distribution and conservation status of this species needs investigation in Iran.

Sources

Type material:- Syntypes NMW 16688, NMW 55511, NMW 55512, NMW 55513, NMW 57161, SMF 100.

?see Excel file where some fish re-identified

Comparative material:-  BM(NH) 1931.12.21:21, 86.9 mm standard length, Mosul, Mesopotamia but this has 17 rakers. ; BM(NH) 1974.2.22:83, 1, 67.9 mm standard length, Iraq, Sirwan, Tigris River near Faish Khabour (no other locality data); ZMB 3364 (possibly syntypes as marked from Vienna Museum), 4, 55.6-65.8 mm standard length, Syria, Aleppo (= Halab); SMF 28638, 14, 69.1-100.7 mm standard length, Syria, Euphrates River, Deir ez zor (35º31'N, 39º54'E); SMF 28678, 3, 59.0-98.8 mm standard length, Syria, Euphrates River upstream Deir ez zor (35º31'N, 39º57'E); SMF 28698, 4, 84.4-105.6 mm standard length,  Syria, Euphrates River, downstream Baath Lake (35º55.723'N, 39º00.572'E); SMF 28712, 3, 51.8-59.3 mm standard length,  Syria, Euphrates River Raqqa to Halebye-Zalebye (35º36.083'N, 39º00.572'E to 53º50.029'N, 39º20.797'E); BM(NH) 1968.12.13:124-135, 40.1-50.7 mm standard length, Syria, Tigris River at Ain Diwar (?); BM(NH) 1968.12.13:147-154, 4, 38.5-71.1 mm standard length, Syria, Quwayq River at Behourte (?).

Alburnus chalcoides
(Güldenstaedt, 1772)

Common names

شاه كولي (shah kuli or shah kooli in Gilaki; kuli is widely used for any small fish and may derive from kul which can mean any pond or sheet of water) or شاه ماهي  (= shah mahi menaing royal fish or king fish in the sense of the best or most important fish); mahi shah kuli; كاس كولي (= kas-e kuli, meaning cup or bowl fish?); aslak in Mazandaran, siah kole (= presumably siah kuli, black fish), safid kuli (= white fish).

[samayi, schamay or schumai, Lankaran samayisi for A. chalcoides longissimus, Kur samayisi for A. chalcoides, all in Azerbaijan; Iranskaya shemaya or Iranian shemaya, Lenkoranskaya shemaya or Lenkoran shemaya, shemaya or shamaya in Russian; Caspian shemaya; bleak, Danube bleak].

Systematics

Cyprinus chalcoides was originally described from the Terek, Sulak and Cyrus (= Kura) rivers, Russia.

Cyprinus clupeoides Pallas, 1776 from the Caspian Sea, Terek and Kura rivers (also spelt clupoides in error), possibly Leuciscus albuloides Valenciennes, 1844 from "rivières de Perse", Alburnus longissimus Warpakhovskii, 1892 from the Geoktapinka River, Lenkoran District, Azerbaijan and Alburnus latissimus Kamenskii, 1901 from the mouth of the Kura River, Azerbaijan are synonyms. Since Alburnus latissimus occurs with Alburnus chalcoides in the Kura River, its status is necessarily equivocal.

Chalcalburnus chalcoides iranicus Svetovidov, 1945 was described as the subspecies of the Iranian shore of the Caspian Sea basin and Alburnus chalcoides longissimus Warpakhovskii, 1892 as the subspecies of the Lenkoran in Azerbaijan neighbouring Iran. Coad (1996b) examined the types of iranicus and longissimus and found them not to be distinguishable. The latter name has priority but both these nominal subspecies, and latissimus, are most probably not distinct from the type subspecies. They were founded on small samples from relatively homogenous spawning populations. Variation may be clinal or related to local temperature and other environmental variables. A very large series of specimens would be necessary to define this.

The Caspian Sea species may be Alburnus chalcoides chalcoides with a distinct subspecies, Alburnus chalcoides mento (Heckel, 1836), in the Black Sea basin although up to 13 subspecies are named from Anatolia and the basins of the Black, Caspian and Aral seas.

The type material of Chalcalburnus chalcoides iranicus is in the Zoological Institute, St. Petersburg (ZISP 31231, holotype (see below), and 3 paratypes 142.0-199.9 mm standard length), the type locality being "a small stream near the hospital near Shahi, Talar River basin" on labels in the Zoological Institute, St. Petersburg and "a small river in the vicinity of town Shakhi (basin of the river Talar, running into the Caspian Sea west of the Gorgan Bay" (Svetovidov, 1945b). Shahi or Qa'emshahr is at 36°28'N, 52°53'E. Svetovidov (1945b) lists the holotype as a female of total length 263.5 mm and body length 226 mm but the holotype in ZISP is 216.7 mm standard length (Coad, 1996b).

The type material of Alburnus longissimus is in the Zoological Institute, St. Petersburg (ZISP 8653, 2 syntypes, 164.8-185.9 mm standard length, from "Fl. Geoktapinka" (Lenkoran). The locality is probably near Prishib at 39°08'N, 48°36'E (Coad, 1996b). ZISP 8654 (6 fish, 121.2-164.4 mm standard length) from the type locality are listed as types in Berg (1911-1914) but not in the ZISP catalogue. Also an A. longissimus syntype from St. Petersburg is in the Natural History Museum, London from "R. Geotapinka" (BM(NH) 1891.10.7:28).

Bagherian and Rahmani (2007; 2009) examined two populations, from the Haraz River and the Shirud, morphometrically. The males and the females between the two populations were different, but this was attributed to environmental factors. Truss analysis separated the two populations. Rahmani et al. (2007) were able almost to separate the two populations using meristic characters. Rahmani et al. (2006) were able to separate populations from the Gazafrud and Haraz rivers using morphometric characters but not meristic ones. Rahmani et al. (2009) used the 18S rRNA gene and found populations from the Haraz, Shirud and Gazafrud rivers were homogenous.

A hybrid of Alburnus chalcoides and Vimba vimba persa was reported from the Safid River (Petrov, 1926) and a hybrid between Leuciscus (= Squalius) cephalus and Alburnus chalcoides is reported from Turkey (Ünver and Erk'akan, 2005; Ünver et al., 2008).

Key characters

The short, naked ventral keel, usually 8 branched dorsal fin rays, distribution, and the characters in the table under A. atropatenae can be used to identify this species.

Morphology

Lateral line scales 54-74. The dorsal and ventral scale margins are parallel or rounded and the anterior margin is wavy or has a pronounced central protuberance. The posterior scale margin can be rounded and more or less smooth or rounded and finely crenulate. Crenulation may be related to size or sexual maturity but is not always evident even in spawning males. Circuli are numerous and fine, radii are few and present on the anterior and posterior fields (a few fish had some scales with no anterior radii), and the focus is slightly subcentral anterior. There is a well-developed pelvic axillary scale. The ventral keel is only naked near the vent and rarely may be scaled along its entire length although Kottelat and Freyhof (2007) have an exposed keel of 8-12 scale lengths, up to 80% of the anus to pelvic fin base distance. Dorsal fin with 2-3, usually 3, unbranched and 7-9, usually 8, branched rays, anal fin with 3 unbranched and 12-19 branched rays, pectoral fin branched rays 13-16, and pelvic fin branched rays 7-9. Ginzburg (1936b) gives counts of 13(7), 14(34), 15(52), and 16(7) for anal fin rays from Iranian material, modally different from my counts below (possibly the last two rays were counted separately but variation between samples is also possible). Gill rakers 18-31, serrated medially and elongate, reaching the second or third adjacent raker when appressed. Total vertebrae 43-45. Pharyngeal teeth 2,5-5,2, more rarely 2,5-5,1, 2,5-5,3, 2,5-5,4, or 3,5-5,3. Teeth are elongate, slender, curved inward, strongly hooked at the tip and strongly serrated with serrations on the anterior margin of the long, narrow and concave grinding surface. The most posterior main row tooth may lie medial to the second tooth. The swimbladder is pointed posteriorly (rounded in Alburnus hohenackeri and A. filippii). The gut is an elongate s-shape. Total vertebrae 41-45.

Meristics in Iranian specimens: dorsal fin branched rays 7(3), 8(55) or 9(2); anal fin branched rays 12(1), 13(4), 14(33), 15(19) or 16(3); pectoral fin branched rays 13(4), 14(9), 15(34) or 16(13); pelvic fin branched rays 7(2), 8(57) or 9(1); lateral line scales 54(1), 55(2), 56(2), 57(5), 58(8), 59(5), 60(14), 61(7), 62(5), 63(6), 64(2), 65(1), 66(1) or 67(1); total gill rakers 18(1), 19(5), 20(12), 21(15), 22(14), 23(9), 24(3) or 25(1); pharyngeal teeth 2,5-5,2(30), 2,5-4,2(1), 2,4-5,2(1) or 2,5-5,3(1); and total vertebrae 42(2), 43(9), 44(32) or 45(7).

Sexual dimorphism

Abdurakhmanov (1962) reports the eye diameter and anal fin base to be larger in males on average for fish from the Kura River basin in Azerbaijan. Iranian males taken in July have small tubercles scattered on top of the head and fine tubercles lining the anterior flank scales. Females are larger than males (Bagherian and Rahmani, 2007)

Colour

The overall colour is metallic silvery and the back is a contrasting olive-green. The iris is bright silver. There is no dark band along the sides. The dorsal and caudal fins are greyish and the other fins colourless to whitish. The peritoneum is light brown but with numerous melanophores in contrast to the dark peritoneum in A. mossulensis.

Size

Reaches 45.0 cm and 1.5 kg. Shemaya on the Kura River of Azerbaijan are larger than those in the south Caspian, up to 36 cm as opposed to 29 cm.

Distribution

Found from central Europe to the basins of the Black, western and southern Caspian and Aral seas. It is recorded from the entire southern coast of the Caspian Sea and its rivers, including the Atrak, Gorgan, Gharasu, Tajan, Babol, Haraz, Sardab, Aras, Tonekabon, Pol-e Rud and Safid rivers, the Anzali Talab, Gorgan Bay, southeast, southwest and south-central Caspian Sea (Derzhavin, 1934; Kozhin, 1957; Svetovidov, 1945b; Holčík and Oláh, 1992; Shamsi et al., 1997; Abbasi et al., 1999); Kiabi et al., 1999; Abdoli, 2000; Bagherian and Rahmani, 2007; 2009; Patimar et al., 2010; Abdoli and Naderi, 2009).

Alburnus chalcoides aralensis Berg, 1926 is reported from the Karakum Canal in Turkmenistan (Shakirova and Sukhanova, 1994; Sal'nikov, 1995) and may eventually be found in the Tedzhen River and Caspian Sea basins of Iran.

Zoogeography

A widespread species with numerous nominal subspecies which have not all been fully investigated. It presumably originated as part of a Danubian or Sarmatian fauna and the subspecies have become isolated in parts of this former basin.

Habitat

Young are rheophilous (Abdurakhmanov, 1975). A migration to piedmont and montane zones used to occur before dams and weirs obstructed movements. Some populations are landlocked while others are semi-anadromous. Knipovich (1921) reports this species from depths of 23.8-25.6 m in the Iranian Caspian Sea. Kottelat and Freyhof (2007) record a tolerance of 14‰ salinity. Riazi (1996) reports that this species is native (resident) to the Siah-Keshim Protected Region of the Anzali Mordab. Shape differences found by Bagherian and Rahmani (2007) in two Iranian rivers were attributed to the Haraz River having a muddy estuary, a shallow slope to the bottom, high turbidity and low water flow in contrast to the Shirud which was sandy with high water flow and high clarity. The latter population developed a more slender body due to increased resistance to water flow.

Age and growth

Life span is 5 years with a theoretical limit of 6.5 years in Azerbaijan (Abdurakhmanov, 1975) and at least 5 years in Iran (Holčík and Oláh, 1992) and Turkey (Tarkan et al., 2005). Sexual maturity is attained at 3 years of age in Azerbaijan and growth is most rapid at an age of 2 years, decreasing thereafter because of high natural mortality (Abdurakhmanov, 1975). The fishes on the spring spawning run in the Anzali Mordab are 10.5-29.0 cm standard length, average 14.0 cm, and 2-5 years old with most (63%) fish in age group 3. Males are mature at 2-4 years and females at 3-5 years. Growth is high during the first 3 years of life and then declines (Holčík and Oláh, 1992). Karimpour et al. (1993) found the Anzali Mordab population to be smaller than the Kura River population but the mordab fish showed greater growth after maturation. The spawning migration into the mordab begins in March and peaks in May and at the beginning of June. Length range was 10.0-24.0 cm, average 16.2 cm with a mean weight of 64.7 g. Age composition was 2-5 years with 3-year-olds comprising 62.5% of the fish. Females formed 57% of the migrating fish. Rahmani (2008) investigated this species in the Haraz and Shirud rivers and found maximum length and weight in a 5- (sic) year old female at 251 mm and 96 g, the most abundant age groups were 2+ and 3+ years for males and females respectively, males in the Shirud population were heavier and longer on average in younger ages while differences in females were not significant, and females of the Shirud population has isometric growth while Haraz fish had positive allometry. The von Bertalanffy growth parameters were Lt = [405.9 (1-e-0.1(t+1.54))] for males in Haraz and Lt = [442.6 (1-e-0.1(t+1.43))] for females in Haraz, and  Lt = [359.5 (1-e-0.145(t+1.002))] for males in Shirud and Lt =[405.9 (1-e-0.1(t+1.54))] for females in Shirud. Females had a higher L while K values for males were relatively higher in the two rivers. Rahmani et al. (2009) found growth was better in the Shirud compared with other populations because this river had desirable biological parameters for immigration. Patimar et al. (2010) compared fish from the Siah and Gorgan rivers and found a five-year life cycle, with negative allometric growth for Siah males and positive allometric growth for Siah females and for both sexes in the Gorgan, and sex ratios were unbalanced in favour of females in both rivers. The von Bertalanffy growth parameters were Lt = [370.08 (1-e-0.15(t+0.70))] for males in Siah and Lt = [432.52 (1-e-0.11(t+1.21))] for females in Siah, and  Lt = [371.79 (1-e-0.14(t+0.96))] for males in Gorgan and Lt = [436.10 (1-e-0.11(t+1.34))] for females in Gorgan.

Food

Holčík and Oláh (1992) report a feeding migration in July to September in the western basin of the Anzali Mordab. Gut contents include diatoms and algae, dragonfly larvae, and copepods (Abdurakhmanov, 1962). Iranian fish had plant fragments, sand grains, crustaceans, insect remains and chironomid larvae in gut contents.

Reproduction

This species is an intermittent spawner with three batches of eggs, only two of which are laid at an interval of 18-19 days. Fecundity reaches 54,700 eggs in Azerbaijan but this is less than that of diadromous populations. Egg diameter is up to 1.9 mm. Spawning takes place in the second half of July to the end of August at water temperatures of 18-25°C in the Mingechaur Reservoir in Azerbaijan. Eggs are laid on rocky bottoms in 15-20 cm of water after a migration into streams or on rocky grounds of reservoirs (Abdurakhmanov, 1962; 1975; Elanidze, 1983). There is a spawning migration into the Kura River from October to April, peaking in December-January, with spawning taking place in spring in the upper reaches (Berg, 1959). In Lake Tuş, Turkey spawning occurred in May-June, egg numbers reached 20,971 and average egg diameter 1.05 mm (Balık et al., 1996).

Svetovidov (1945b) considers that Iranian populations (his iranicus subspecies) spawn nearly throughout the year since fish having ripe sex products were caught in both July and February and young were found along the Iranian coast throughout the year. Spawning takes place in the sea, in areas such as Gorgan Bay, and in the lower reaches of rivers. Khaval (1998) reports a spawning migration into the Safid River despite construction, sand removal and pollution. Holčík and Oláh (1992) report a migration into the Anzali Mordab for spawning in late February to early April (but see above; possibly a confusion between the migration at an earlier date than the spawning act). Karimpour et al. (1993) give an absolute fecundity of 6630 eggs in the Anzali Mordab population while mean relative fecundity is 140 eggs/g of body weight. Iranian fish have 1.5 mm eggs as early as 13 March (fish standard length 213.2 mm) and 1.7 mm eggs on 4 June (fish length 154.6 mm) while eggs are only 1.3 mm on 15 July (fish length 142.8 mm). Larger fish may mature and spawn earlier than younger fish. Rahmani et al. (2009) found a peak gonadosomatic index for males in May and for females in early June in the Shirud. Average fecundity was about 3900 eggs with diameter reaching 1.17 mm. Patimar et al. (2010) in their study of Siah and Gorgan River fish found spawning between April and July in the Siah and March and June in the Gorgan, peaking in May in both rivers. Absolute fecundity was up to 38,340 eggs, mean 8426 eggs in the Siah  and up to 17,263 eggs, mean 4215 eggs in the Gorgan. Relative fecundity was up to 599 eggs/g, average 212 eggs/g of body weight in the Siah and up to 696 eggs/g, average 112 eggs/g in the Gorgan. Mean egg diameters were 1.40 mm in the Siah and 1.27 mm in the Gorgan. These differences in life history (see also Age and growth above) were attributed to differing habitat characteristics.

Parasites and predators

Molnár and Jalali (1992) report the monogeneans Dactylogyrus minor, D. alatus and D. vistulae from this species in the Ghasemlu River, an inland watershed, with the latter species also in the Safid Rud. They also describe a new species of monogenean, Dactylogyrus holciki, from this species in the Beshar River of the Persian Gulf drainage, possibly confusing this Caspian Sea basin cyprinid with A. mossulensis. Molnár and Jalali (1992) also record the monogenean Dactylogyrus chalcalburni from the Safid and Zayandeh rivers, although this Caspian Sea basin cyprinid does not occur in the latter locality, possibly again confusing the same species as noted above. Shamsi et al. (1997) report Clinostomum complanatum, a parasite causing laryngo-pharyngitis in humans, from this species. Masoumian and Pazooki (1998) surveyed myxosporeans in this species in Gilan and Mazandaran provinces, finding Myxobolus pseudodispar. The helminths Pentagramma symmetrica and Mazocea alaosa are recorded from the guts of Chalcaburnus tarichi (sic, presumably A. chalcoides) from the Anzali wetland (Ataee and Eslami, 1999; www.mondialvet99.com, downloaded 31 May 2000). Naem et al. (2002) found the following parasites on the gills of this species from the western branch of the Safid River, namely the monogenean trematodes Dactylogyrus chalcalburni and Gyrodactylus sp.. Sattari et al. (2004, 2005) surveyed this species in the Anzali wetlands, recording Anisakis sp. Maleki and Malek (2007) examined fish from the Shirud in the Caspian Sea basin and recorded the digeneans Posthodiplostomum cuticola, Diplostomum spathaceum, Clinostomum complanatum and Allocreadium sp.Sattari et al. (2007) record the nematode Anisakis sp., the digenean Diplostomum spathaceum and the monogenean Dactylogyrus extensus in this species in the Anzali wetland of the Caspian shore. Miar et al. (2008) examined fish in Valasht Lake and the Chalus River, Mazandaran and found the metazoan Argulus foliaceus. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Argulus foliaceus on this species.

Economic importance

The shemaya was a valuable edible fish on the Kura River of Azerbaijan with catches as high as 500 centners (1 centner = 100 kg) prior to construction of the Kura dam. The catch for Azerbaijan in 1933 was 1950 centners or 2,029,000 fish. Catches in the Mingechaur Reservoir formed by the dam were 133 centners in 1972 (Abdurakhmanov, 1975). Reputedly delicious eating (Lönnberg, 1900b). They are fished for on the spawning run when fatty. In Iran they are caught by cast nets in the inlets and outlets of the Anzali Mordab in spring on the spawning run and by gill nets in the western basin on the feeding migration. Holčík and Oláh (1992) report a catch of 956 kg in the Anzali Mordab in 1990 but catches in recent years may have been confused with the exotic Hemiculter leucisculus (Holčík and Olah, 1990).

Conservation

Holčík and Oláh (1992) report a decline in the numbers of this species owing to damming of rivers where it used to spawn. Kiabi et al. (1999) consider this species to be near threatened in the south Caspian Sea basin according to IUCN criteria. Criteria include commercial fishing, sport fishing, abundant in numbers, habitat destruction, widespread range (75% of water bodies), absent in other water bodies in Iran, and present outside the Caspian Sea basin. Mostafavi (2007) lists it as near threatened in the Talar River, Mazandaran. Endangered in Turkey (Fricke et al., 2007). 

This species has been artificially bred without hormones on the Shirrud with a fertilisation rate of 90-98%. Hatching took 6 days and the hatching rate was 57% (I.F.R.O. Newsletter, 36:4, 2003). On the Tajan River, induction of ovulation has been carried out using LRH-Aa with metoclopramide and carp pituitary extract (Yousefian et al., 2008). Fertilisation rate was 83%, hatching rate 90% and survival of larvae 81%. Shirvani and Jamili (2009) found excessive levels of cadmium and lead in this fish from regions of Anzali where oil ship traffic was highest. Daei et al. (2009) reported on the effects of cadmium and lead on the iron solute in blood.

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in aquaculture and as food. Lelek (1987) classifies this species as vulnerable to endangered in Europe.

Further work

The various subspecies should be examined using molecular techniques and numbers of this species in Iranian rivers monitored for conservation management.

Sources

The types of C. chalcoides iranicus are included in the meristic data for Iranian specimens.

Type material: See above, Chalcalburnus chalcoides iranicus (ZISP 31231), Alburnus longissimus (ZISP 8653, ?8645, BM(NH) 1891.10.7:28).

Iranian material: CMNFI 1970-0531, 4, 64.5-74.9 mm standard length, Mazandaran, Larim River (36º46'N, 52º58'E); CMNFI 1970-0553, 2, 101.9-163.1 mm standard length, Gilan, Sowsar Roga River (37º27'N, 49º30'E); CMNFI 1971-0327A, 2, 54.5-116.9 mm standard length, Gilan, Shafa River (37º35'N, 49º09'E); CMNFI 1979-0081, 7, 77.8-106.5 mm standard length, Mazandaran, Caspian Sea 3 km west of Chalus (36º41'N, 51º24'E); CMNFI 1979-0434, 4, 47.3-154.6 mm standard length, Mazandaran, Shir River (36º51'N, 50º49'E); CMNFI 1979-0435, 1, 170.5 mm standard length, Gilan, stream 10 km west of Ramsar (36º57'N, 50º37'E); CMNFI 1979-0437, 2, 164.5-175.6 mm standard length, Gilan, Safid River 2 km west of Astaneh (37º16'30"N, 49º56'E); CMNFI 1979-0438, 12, 114.9-158.9 mm standard length, Gilan, Gholab Ghir River (37º27'N, 49º37'E); CMNFI 1979-0439, 2, 156.6-173.2 mm standard length, Gilan, Anzali Mordab (ca. 37º27'N, ca. 49º25'E); CMNFI 1979-0441, 1, 109.8 mm standard length, Gilan, river 14 km south of Hashtpar (37º42'N, 48º58'E); CMNFI 1979-0443, 1, 159.6 mm standard length, Gilan, river 34 km north of Hashtpar (38º06'N, 48º53'E); CMNFI 1979-0445, 1, 114.9 mm standard length, Gilan, stream 10 km south of Astara (38º21'N, 48º51'E); CMNFI 1979-0455, 1, 88.5 mm standard length, Zanjan, Qezel Owzan River at Gilavan (36º47'N, 49º08'E); CMNFI 1979-0474, 1, 141.0 mm standard length, Mazandaran, Tajan River (36º34'N, 53º05'E); CMNFI 1979-0686, 23, 25.5-111.0 mm standard length, Gilan, Safid River (37º24'N, 49º598'E); CMNFI 1979-0788, 48, 35.2-74.7 mm standard length, Mazandaran, Gorgan River at Khadje Nafas (37º00'N, 54º07'E); CMNFI 1980-0120, 17, 56.4-69.5 mm standard length, Mazandaran, Babol River at Babol Sar (36º43'N, 52º39'E); CMNFI 1980-0123, 2, 97.0-106.4 mm standard length, Gilan, Safid River around Dakha (no other locality data); CMNFI 1980-0126, 3, 182.1-213.2 mm standard length, Gilan, Caspian Sea near Bandar-e Anzali (37º28'N, 49º27'E); CMNFI 1980-0132, 7, 18.7-142.8 mm standard length, Gilan, Safid River at Kisom (37º12'N, 49º54'E); CMNFI 1980-0142, 2, 135.0-187.2 mm standard length, Gilan, Nahang Roga River (37º28'N, 49º28'E); CMNFI 1980-0908, 3, 45.4-155.2 mm standard length, Gilan, Safid River estuary (ca. 37º28'N, ca. 49º54'E).

Alburnus filippii
Kessler, 1877

Common names

كولي كورا (= kuli-ye Kura), ماهي مرواريد or مرواريد ماهي (= mahi morvarid or morvarid mahi, meaning pearl fish).

[Kur kumuscasi in Azerbaijan; Kurinskaya ukleika or Kura bleak, ukleika filippi or Filippi's bleak, both in Russian].

Systematics

The lectotype of Alburnus Filippii as designated by N. Bogutskaya is in the Zoological Institute, St. Petersburg (ZISP 2926) and is from "Fl. Kura pr. Tiflis", Acad. Brandt, 1867, 75.3 mm standard length. Paralectotypes are ZISP 2925, 13 fish, same data as lectotype, 43.0-84.4 mm standard length, ZISP 2914, 2 fish, "Fl. Kura pr. Borshoma", Acad. Brandt, 1867, 83.6-87.6 mm standard length, and ZISP 50412, 16 fish, "Reka Kura Tiflis", Acad. Brandt, 1867, 60.6-88.6 mm standard length. A syntype, 57.3 mm standard length, is in the Natural History Museum, London from Tiflis (BM(NH) 1897.7.5:33, formerly in ZISP).

Alburnus filipii var. Kessler in Brandt, 1880 from the Tchaldyr Lake is also this species.

Knipovich (1921) reports a Caspian basin species Alburnus philippii Kessler which is presumably a misspelling of filippii. The specific name is sometimes spelt filippi, which is incorrect.

Abdurakhmanov (1962) compares a sample from the Kura River basin with one from the Kendalanchaya in the Aras River basin of Azerbaijan and finds 15 characters are significantly different on average. Fish from the Kura have a longer head, greater dorsal and anal fin heights, and longer pectoral, pelvic and upper and lower caudal fin lobes while fish from the Aras have more scales in the lateral line, a deeper head, body and caudal peduncle, and a longer anal fin base, pectoral-pelvic fin distance and snout, and a greater interorbital width. No taxonomic status is assigned these two populations.

A hybrid with Alburnus charusini hohenackeri (= Alburnus hohenackeri) was reported by Petrov (1926) from the Safid River and the Kumbashinka in Lenkoran.

Key characters

This species is distinguished from its relative (Alburnus chalcoides, also with a long, naked ventral keel) by having modally 7 branched dorsal fin rays and generally lower anal fin ray counts although these do overlap (10-21, usually 12-16 in Iran for alburnus; 9-13, usually 10-12, for filippii). See also table under A. atropatenae.

Morphology

Dorsal fin with 3 unbranched and 6-8, usually 7, branched rays, anal fin with 2-3, usually 3, unbranched and 9-13 branched rays, usually 10-12. Pectoral fin branched rays 12-16, pelvic fin branched rays 6-8, usually 7. Lateral line scales 46-64. Scales have a wavy anterior margin, an overall vertical oval shape, sometimes tapering to a rounded posterior point and sometimes more rounded, few anterior and posterior radii, and a subcentral anterior focus. There is a pelvic axillary scale. The naked ventral keel usually extends more than half way from the anal papilla to the pelvic fin insertion but is often completely scaled, notably in fish from the Safid River basin. The scaled keel runs from the papilla to the pelvic fin base. Gill rakers 12-21, reaching the second or third adjacent raker when appressed. Pharyngeal teeth 2,5-5,2 (but see below for Iranian specimens) with variants 2,5-5,1, 1,5-5,2, 1,5-5,1, 2,5-4,2, 2,4-5,2, 2,5-4,1, 2,4-4,2, 1,5-4,1, 1,4-5,1, and 1,5-4,2. Teeth are stongly hooked and strongly serrated. Serrations are on the anterior margin of each tooth. The degree of hook and serration development varies individually and does not seem to be size related. Some fish have little development of either character. The area below the hook is an elongate, flat to concave surface. Vertebrae number 38-43. The swimbladder has a rounded end in contrast to the pointed end in Alburnus chalcoides. The gut is an elongate s-shape with a small anterior loop. The chromosome number is 2n=50 and Nazari et al. (2009) give further details.

Meristic variation in Iranian specimens: dorsal fin branched rays 6(1), 7(44) or 8(5); anal fin branched rays 9(1), 10(19), 11(24), 12(5) or 13(1); pectoral fin branched rays 12(3), 13(19), 14(20), 15(7) or 16(1); pelvic fin branched rays 6(3), 7(42) or 8(5); lateral line scales 46(1), 49(1), 50(5), 51(5), 52(4), 53(12), 54(5), 55(2), 56(5), 57(6), 58(1), 60(2) or 63(1); total gill rakers 12(4), 13(8), 14(19), 15(10), 16(6) or 17(3); pharyngeal teeth 2,5-4,2(10), 2,4-5,2(2), 2,4-4,2(2), 2,5-5,2(1), 1,5-4,2(2), 1,5-5,2(1), 1,5-4,1(1) and 1,4-5,1(1); and total vertebrae 38(2), 39(8), 40(18), 41(9) or 42(1).

Sexual dimorphism

Males and females have moderate-sized tubercles widely scattered on the top of the head, on the snout and lining the lower edge of the jaw. Much smaller tubercles are scattered among the ones on top of the head.

Colour

The back is brown, flanks silvery and the belly white. A characteristic dark streak, as wide as the eye, runs along mid-flank. Fins are hyaline. The peritoneum is brown or light with large scattered melanophores.

Size

Reaches 17.0 cm standard length.

Distribution

Found only in the Caspian Sea basin from the Kura River of Azerbaijan to the Safid River of Iran including headwaters in Turkey, Armenia and Iran at altitudes over 3000 m. It is distributed from the upper to the lower reaches of the Aras (Qareh Su) and Safid (Qezel Owzan) rivers in Iran and in the Anzali Talab (Holčík and Oláh,1992; Abbasi et al., 1999; Kiabi et al., 1999; Abdoli and Naderi, 2009).

    

Zoogeography

The relationships of this species with other Alburnus needs to be examined. It presumably originated as part of a Sarmatian fauna, isolated in the Caspian Sea.

Habitat

Primarily a freshwater species, this minnow may be found in the brackish outlets of the Anzali Mordab (Holčík and Oláh, 1992). Jolodar and Abdoli (2004) note that it is found more in upstream waters than A. alburnus.

Age and growth

Life span is about 5 years with maturity at 1 year for males and 2 years for females.

Food

Gut contents include plant remains, mayflies and algae (Abdurakhmanov, 1962). Iranian specimens contain insect remains, a few crustaceans and sand grains. One sample from the Qareh Su north of Ardebil had been feeding on water beetles (Hydrophilidae) but also spiders and scarab beetles (Euoniticellus sp.) indicating food is also taken from the surface.

Reproduction

Eggs number up to 14,210 and diameters up to 1.51 mm. May is the principal spawning month in Azerbaijan (Abdurakhmanov, 1962). Male fish caught on 6 June in Iran had tubercles scars on top of the head while female fish from another locality (Zanjan River) taken on 8 June had mature eggs measuring 1.2-1.3 mm. Spawning probably occurs in May and June in Iran, depending on local conditions.

Parasites and predators

Jalali et al. (2005) summarise the occurrence of Gyrodactylus species in Iran and record G. elegans in the Beheshtabad River. Mortazavi Tabrizi et al. (2005) record Ligula intestinalis and Bothriocephalus acheilognathi in this species from the Sattarkhan Dam in East Azerbaijan. Undoubtedly food for various predatory fishes. Pazooki et al. (2005) record Trichodina  perforata from this species in waterbodies of Zanjan Province. Pazooki et al. (2006) record the monogeneans Dactylogyrus vistulae and Gyrodactylus sp. from this fish in Zanjan Province.

Economic importance

None.

Conservation

Kiabi et al. (1999) consider this species to be of least concern in the south Caspian Sea basin according to IUCN criteria. Criteria include medium numbers, habitat destruction, medium range (25-75% of water bodies), absent in other water bodies in Iran, and absent outside the Caspian sea basin. Vulnerable in Turkey (Fricke et al., 2007).

Further work

The biology of this species needs investigation.

Sources

Type material: See above, Alburnus Filippii (ZISP 2926, 2925, 2914, 50412, BM(NH) 1897.7.5:33).

Iranian material: CMNFI 1970-0538, 8, 34.9-61.8 mm standard length, Gilan, Qezel Owzan River (ca. 36º44'N, 49º24'E); CMNFI 1970-0552, 1, 50.1 mm standard length, Gilan, Sowsar Roga River (37º27'N, 49º30'E); CMNFI 1979-0448, 1, 70.9 mm standard length, Azarbayjan-e Khavari, Ahar Chay 8 km from Ardabil (38º18'30"N, 48º22'E); CMNFI 1979-0452, 2, 52.4-54.9 mm standard length, Azarbayjan-e Khavari, Qezel Owxan River 6 km from Mianeh (37º23'N, 47º45'E); CMNFI 1979-0453, 9, 43.7-73.3 mm standard length, Zanjan, Zanjan River (37º06'N, 47º56'E); CMNFI 1979-0455, 17, 42.8-62.5 mm standard length, Markazi, Manjil Dam (36º45'N, 49º17'E); CMNFI 1979-0695, 3, 61.3-63.5 mm standard length, Gilan, Safid River (36º46'N, 49º24'E); CMNFI 2007-0081, 1, 51.0 mm standard length, Zanjan, Zanjan River near Soltaniyeh (ca. 36º27'N, ca. 48º45'E); CMNFI 2007-0082, 11, 41.2-59.6 mm standard length, Zanjan, Zanjan River basin near Zanjan (ca. 36º36'N, ca. 48º32'E); CMNFI 2007-0087, 6, 55.7-83.1 mm standard length, Azarbayjan-e Khavari, Qareh Su north of Ardebil (38º22'N, 48º19'E); CMNFI 2007-0107, 3, 41.1-42.3 mm standard length, Kordestan, Qezel Owzan River basin near Bijar (ca. 35º54'N, ca. 47º20'E).

Alburnus hohenackeri
Kessler, 1870

Marun River, courtesy of Kai Borkenhagen

Common names

ماهي مرواريد or مرواريد ماهي (= mahi morvarid or morvarid mahi, meaning pearl fish), kuli (= general term for small fish), كولي ايراني (= kuli-ye Irani).

[simali gafgaz kumuscasi for A. c. charusini or zagafgaziya kumuscasi for A. c. hohenackeri, both in Azerbaijan; ukleika or bleak, zakavkazskaya ukleika or Transcaucasian bleak, persidskaya ukleika or Persian bleak, sefidrudskaya ukleika or Safid River bleak, all in Russian; Caucasian bleak (as A. hohenackeri)].

Systematics

Alburnus Hohenackeri was originally decsribed  from Karabakh, Azerbaijan, on the Kura River. The taxon in Iran was formerly included within the wide-ranging species Alburnus alburnus (Linnaesus, 1758). Cyprinus Alburnus was originally described from Europe.

Synonyms are Alburnus charusini Herzenstein in Zograff and Kavraiskii, 1889 described from the Kamysh-Samarskie lakes between the Volga and Ural rivers in Kazakhstan, Alburnus alasanicus Kamenskii, 1901 from the Alasan, Alazan' or Alazani River, a left bank Kura River tributary in Georgia, Alburnus lucidus var. macropterus Kamenskii, 1901 described from the Alazan' River, Alburnus alburnus charusini natio elata Petrov, 1926 from the Prorva River (lower reaches of the Terek River), the Sulak River and the Divichi Liman, western Caspian Sea, Alburnus striatus Petrov, 1926 from "Kizil-Agachskogo Zaliva" (Kizil-Agach Bay, Turkmenistan) and "Astrabadskogo Zaliva" (= Astrabad or Gorgan Bay, Iran), and Alburnus alburnus natio dagestanicus Petrov, 1930 (sic) but later in the same paper given, and probably originally meant, as A. a. charusini n. dagestanicus) described from the "Kaukasusküste des Kaspische Meeres".

Alburnus pseudospirlinus Petrov, 1926 from "Novaya Rechkaya (nizov'ya Sefid-Rud)" (= new stream, lower Safid River) is a hybrid of this species and Alburnoides bipunctatus (sic) (Berg, 1948-1949). A hybrid with Alburnus filippi was described from the Kumbashinka River in the Lenkoran and from the Safid River (Petrov, 1926).

The holotype of Alburnus hohenackeri is in the Zoological Institute, St. Petersburg (ZISP 2839). The holotype of Alburnus charusini is in the Zoological Museum of Moscow State University under MMSU P.1314. Four fish as listed as questionable syntypes under MMSU P.1812 by Svetovidova (1978) although according to Eschmeyer et al. (1996) the original says P.1314 with a unique holotype only.

This species was recognised as Alburnus charusini in Iran but characters overlap with Alburnus alburnus, a highly variable species (Gäsowska, 1974). In any case hohenackeri has priority over charusini. Literature sources conflict on the correct name. Petrov (1926; 1930) refers to Alburnus alburnus hohenackeri Kessler, 1877 for fish in northern Iran with natio persicus Petrov, 1926 in the Safid River, natio dagestanicus Petrov, 1930 in the Dagestan area of Azerbaijan and natio kumbashensis Petrov, 1926 from the Kumbashinka River and Lake Ol'khovskoye in the Lenkoran area of Azerbaijan. Natio are not recognised by the Zoological Code of Nomenclature (Ride et al., 1985). Liška and Pivnička (1985) refer southern and southeastern populations of this species to Alburnus alburnus albidus Costa, 1838, and this would include the Iranian populations. These fish are separated from the type subspecies by having 39-47 lateral line scales, most frequently 42-44 (44-54, most frequently 47-50 in A. a. alburnus), branched anal fin rays 10-17, most frequently 13-15 (14-21, most frequently 16-19), and head length as % of body length 22-27 most frequently 23-25 (19-25, most frequently 21-23). N. Bogutskaya (pers. comm., 1995) and Reshetnikov et al. (1997) refer Iranian fish to Alburnus alburnus hohenackeri as there is a definite character break at the Terek River separating northern populations from southern ones. Petrov (1930) came to a similar conclusion on the name of the Iranian populations in his study as noted above. Aburakhmanov (1962) too refers the taxon hohenackeri to fish found in the Kura and Aras rivers and in rivers of the Lenkoran coast (and presumably the Iranian coast) while his charusini are north of the Apsheron Peninsula. Bogutskaya and Naseka (2004) and Kottelat and Freyhof (2007) recognise A. hohenackeri as a distinct species.

Key characters

This species can be confused with Alburnoides eichwaldii which has similar scale, fin ray and pharyngeal counts. A key distinction is the total gill raker count of 16-29 (usually 20 or more) in this species as opposed to 5-12, usually 7-10 in Alburnoides. Alburnus rakers are more than twice as long as those in Alburnoides and, being more numerous, are crowded on the arch without the large gaps between individual rakers which characterises Alburnoides. Modal dorsal fin branched ray count of 8 separates it from A. filippii. ?separation from other Alburnus in Caspian

Morphology

Dorsal fin branched rays 7-9, usually 8, after 2-4 unbranched rays, anal fin branched rays 10-21 after 3-4 unbranched rays (note that anal fin count will be a narrower range if A. hohenackeri is recognised as distinct from a widespread A. alburnus (see Iranian counts below). Pectoral fin branched rays 11-16 and pelvic fin branched rays 6-9. Lateral line scales 36-55. Scales bear both anterior and posterior radii with a few curved radii in the lateral fields. The focus is subcentral anterior and circuli are numerous and fine. The naked ventral keel is often wholly or partially covered by scales. Gill rakers 15-29, elongate reaching the third, or rarely second, below when appressed. Vertebrae 36-46. Pharyngeal teeth 2,5-5,2 with variants 2,5-5,1, 2,5-5,3, 1,5-5,2, 1,5-5,1, 2,5-4,2, 2,4-5,2, 2,4-5,1, 2,4-4,2, 1,5-4,2, 2,5-4,1, 1,5-4,1, 1,4-4,1. The elongate and narrow teeth bear a strongly hooked tip and have evident serrations in most specimens although some lack them entirely. The gut is an elongate s-shape with a small anterior loop. The posterior end of the swimbladder is rounded (pointed in Alburnus chalcoides). The chromosome number is 2n=50-52, generally 50 (Klinkhardt et al., 1995).

The natio persicus from the Safid River has dorsal fin branched rays 7-9, anal fin branched rays 12-16 and lateral line scales 40-45. Fish from the Kura-Aras basin and Lenkoran (hohenackeri) have anal fin branched rays 10-15, lateral line scales 38-48, pharyngeal teeth 2,5-5,2, total gill rakers 16-25 and total vertebrae 37-42 (courtesy of N. Bogutskaya, Zoological Institute, St. Petersburg).

Meristics for Iranian fish including Petrov's (1930) counts of dorsal and anal branched rays and lateral line scales for Safid River fish are:- branched dorsal fin rays 7(7), 8(76) or 9(8); branched anal fin rays 12(6), 13(37), 14(28), 15(16) or 16(2); branched pectoral fin rays 12(2), 13(18), 14(17) or 15(3); branched pelvic fin rays 7(11) or 8(29); lateral line scales 39(2), 40(8), 41(10), 42(28), 43(13), 44(9), 45(7), 46(1), 47(1), 48(1), or 50(1); total gill rakers 19(1), 20(2), 21(18), 22(7), 23(5), 24(4) or 25(3); pharyngeal teeth 2,5-5,2(13), 2,5-4,2(11), 2,5-4,1(1), 2,4-5,2(2) or 2,4-4,2(1); and total vertebrae 37(2), 38(24), 39(20), 40(7) or 41(1).

Sexual dimorphism

Tubercles line the edge of each scale and in single file line the rays of all fins. Fine tubercles cover the whole head.

Colour

The overall colour is bright silvery with the posterior scale margins grey on the upper flank. The back is dark blue to olive or bluish-green and is sharply distinct from the lighter flanks. The mid-line of the back has a narrow dark line. The lateral line and the area above it have some pigmentation, concentrated along the lateral line itself, but there is no dark stripe or it is only faintly developed and is bluish or greyish. Above this stripe is an iridescent golden-green stripe only visible at a certain angle. The bluish or greyish stripe is more evident in preserved material. The belly and lower head surface are pearly-white. The iris is silvery with a yellow ring along the outer eye rim but very little around the pupil. The upper part of the iris may have some dark pigment. The dorsal and caudal fins have dark rays and transparent membranes but may be a dirty yellow. Membranes may have some pigment, particularly on the dorsal fin. The upper anterior edge of the pectoral fin has a little dark pigment while the rest of the fin is colourless to grey or orange. Some fish have a yellow base to the pectoral fin. The pelvic and anal fins are usually colourless, although the anal rays may have some grey or there may be some yellow, orange or red on the fin generally. The caudal fin tip is dark grey.

In preserved fish, most flank pigment is above the lateral line. Lateral line scales have pigment both above and below the pore so the pore stands out. This is not as distinctive as in some Alburnoides spp.. A mid-dorsal stripe is more evident in smaller fish and is obscured by the generally darker back and upper flank pigmentation in larger fish. The peritoneum is a light silvery with scattered melanophores. A flank stripe may be developed although not as strongly as in Alburnus filippii; the stripe is more a darker area along the muscle mass divide between a lighter upper flank and lower flank.

Size

Reaches 20 cm.

Distribution

Found from England through Europe and east to the Caspian Sea basin or narrowly the western and southern Caspian Sea basin as A. hohenackeri. It is reported from the Aras River (including the upper reaches of its tributary, the Qara Su) to the Atrak River along the Caspian coast of Iran including the Anzali Talab and Gorgan Bay, and the Gorgan, Gharasu, Tajan, Babol, Haraz, Sardab, Tonekabon; Pol-e Rud and Safid rivers (Derzhavin, 1934; Holčík and Oláh, 1992; Kiabi et al., 1999; Abbasi et al., 1999; Abdoli and Naderi, 2009). Also widely introduced across western, central and eastern Iran, including in the Ab-e Sirvan in the upper Diyala River, Lake Zarivar, in the Zayandeh River of the Esfahan basin, in the Kalshur, Jajarm and Qareh Su of northeastern Dasht-e Kavir basin, and in the Hamun Kushk, and Kahak and Sistan dams of the Sistan basin, and possibly in Minab (= Esteghlal) Dam (A. Abdoli, pers. comm., 1995; J. Holčík, in litt., 1996;.Abdoli, 2000; Ghorbani Chafi, 2000; A. Afzali, pers. comm., 2002; Esmaeili et al., 2010).

    

Zoogeography

This is a widespread species showing great morphological variability over its range, sometimes recognised as taxa. Zoogeographical relationships of these taxa and of the species to other Alburnus have still to be worked out.

Habitat

This species is found in open waters of lakes along the shore or in slow rivers, avoiding turbid conditions and heavy vegetation. There was a mass mortality, presumed to be of this species, on the Babol Sar beach on 24 June 1963 (USNM 270909). It is found more abundantly at river estuaries along the Iranian Caspian shore than Alburnus filippii (Jolodar and Abdoli, 2004).

Age and growth

Maturity is attained at 3 years and life span is up to 9 years. In more northern waters, most spawning males are 3+ and 4+ years while females are 5+ and 6+ years. Iranian populations probably have a similar structure but the age groups would be lower. Mature males averaged 9.7 cm and females 10.5 cm in one study in Russia (Berg, 1948-1949).

Food

Food is planktonic crustaceans, benthic crustaceans such as amphipods, flying insects which land on the water surface, aquatic insects such as backswimmers (Notonectidae), algae, diatoms, and fish eggs and fry. It is an important prey item for other fishes.

Reproduction

Spawning in Europe takes place from April to July in shallow water over a hard bottom. June is the main spawning month in Azerbaijan judging by egg diameters and condition factors (Abdurakhmanov, 1962). Older fish spawn first. Water temperature is usually at 15-16°C or more. Spawning takes place in 3-6 stages at intervals of 9-11 days. The eggs adhere to stones, branches or vegetation. Fecundity is up to 10,000 eggs and egg diameter to 1.4 mm. Incubation lasts about 1 week. Iranian specimens had 1.1 mm diameter eggs in a sample caught on 11 June and mature males were collected on 10 July. Specimens collected in September showed egg resorption while those taken in December had small, developing eggs and those taken in April with better developed eggs. The specimens were small and spawning probably occurs in July for these fish and possibly June for larger ones.

Parasites and predators

Molnár and Jalali (1992) record the monogeneans Dactylogyrus parvus, D. alatus and D. chalcalburni from Alburnus charusini on the Safid Rud.

Gussev et al. (1993b) report the monogenean, Dactylogyrus chalcalburni, from this species in the Zayandeh Rud but this fish does not occur there. The parasite may have been found in Alburnus mossulensis. Shamsi et al. (1997) report Clinostomum complanatum, a parasite causing laryngo-pharyngitis in humans, from this species. Barzegar et al. (2008) record eye parasites from this fish including the digeneans Diplostomum spathaceum and Tylodelphys clavata.

Some European populations of Sander lucioperca feed almost exclusively on this species. Spent adults are known to eat their own eggs.

Economic importance

The scales contain silvery crystals of guanine which are extracted and used to make essence d'orient (or pearl essence) for artificial pearls. About 5000 fish are required for 100 g of essence. Schools in the lower Don River of the Black Sea number up to 10 million fish weighing 30 tonnes. This abundant species is of indirect commercial importance as food for more valued fishes but it has also been used as food for humans.

Conservation

Kiabi et al. (1999) consider this species to be of least concern in the south Caspian Sea basin according to IUCN criteria. Criteria include abundant in numbers, habitat destruction, widespread range (75% of water bodies), and present in other water bodies in Iran. Endangered in Turkey (Fricke et al., 2007). 

Further work

The biology of this species needs investigation, especially in relation to habitats and other fish species where it has been introduced by accident.

Sources

Iranian material: CMNFI 1970-0510, 8, 44.5-72.1 mm standard length, Gilan, Golshan River (37º26'N, 49º40'E); CMNFI 1970-0580, 27, 33.9-56.1 mm standard length, Mazandaran, river near Iz Deh (36º36'N, 52º07'E); CMNFI 1970-0589, 21, 22.5-67.9 mm standard length, Gilan, Safid River (37º12'N, 49º54'E); CMNFI 1971-0343, 1, 63.5 mm standard length, Gilan, Langarud at Chamkhaleh (37º13'N, 50º16'E); CMNFI 1979-0265, 30, 61.6-90.4 mm standard length, Gilan, head of Anzali Mordab at Abkenar (37º28'N, 49º20'E); CMNFI 1979-0432, 22, 34.4-54.3 mm standard length, Mazandaran, Sardab River branch (36º41'N, 51º22'E); CMNFI 1979-0435, 1, 51.9 mm standard length, Gilan, stream 10 km west of Ramsar (36º57'N, 50º37'E); CMNFI 1979-0480, 6, 14.4-64.3 mm standard length, Mazandaran, Gorgan Rver at Gonbad-e Kavus (37º15'30"N, 55º09'E); CMNFI 1980-0122, 41, 29.8-59.0 mm standard length, Mazandaran, Nerissi River (36º38'N, 52º16'E); CMNFI 1980-0147, 5, 44.3-61.5 mm standard length, Gilan, Lashtenesha River (37º21'N, 49º52'E).

Alburnus mossulensis
Heckel, 1843

Common names

شاه كولي (shah kuli = king fish), shah kuli-ye jonubi (= southern king fish), شاه ماهي (= shah mahi, meaning king fish), shah kuli mosulenzis.

[simnan , semnan or samnan, semnan tuyel; sink, or zurri at Mosul (zurri also used for Chondrostoma regium according to Heckel (1846-1849a), but is also used for Aphanius spp., Gambusia and any small fishes or large fishes when young; all in Arabic; Mosul bleak].

Systematics

Leuciscus maxillaris Valenciennes, 1844 from "rivières de Perse", probably Alburnus capito Heckel, 1843 from "Gebirgsflüssen Kurdistans" (mountain streams of Kurdistan in Heckel (1843b) or "Gebirgsbache in Kurdistan" in Heckel (1846-1849a)), Alburnus Iblis Heckel, 1849 described from the "Gegend um Persepolis oder den Gewässern des Araxes" (= probably the Pulvar (= Sivan) River near Persepolis and the Kor River, both in Fars), Alburnus Schejtan Heckel, 1849 described from the "Araxes bei Persepolis", Alburnus caudimacula Heckel, 1849 described from the "Flusse Kara-Agatsch und bei dem Dorfe Geré (= Qarah Aqaj or Mand River, Fars; possibly near Kereft, 29°01'N, 52°52'E), and Alburnus megacephalus Heckel, 1849 described from the "Araxes" are synonyms (e.g. according to Berg (1949)). The type locality of Alburnus mossulensis is the "Tigris bei Mossul" according to Heckel (1843b).

Saadati (1977) considers Alburnus caudimacula to be a distinct species found in the Mand River of Fars based on head length being longer (but the ranges overlap) and a shorter scaleless keel (which is individually variable in these fishes according to my observations).

A subspecies, Alburnus mossulensis delineatus Battalgil, 1942, is reported from Diyarbakir on the Tigris River in Turkey.

A hybrid with Acanthobrama marmid was reported from the Hawr al Hammar in southern Iraq by Krupp et al. (1992) who also note that A. mossulensis is probably a synonym of Alburnus sellal Heckel, 1843, a species originally described the Quwayq River at Aleppo. However, they retained mossulensis as a distinct species because of colour differences and the difficulty of obtaining fresh material of sellal in its polluted habitat at Aleppo in Syria (see Vesiland (1993) for habitat photograph). Heckel (1846-1849a) differentiates mossulensis from sellal by the former being more slender and elongate, the pelvic, dorsal and anal fins are more anterior so the caudal peduncle is more elongate, the eyes are larger and lower on the head, and there is a lead-coloured stripe separating the upper third of the body from the lower part. Berg (1949) considers that A. mossulensis may be nothing more than a subspecies of A. sellal. A Principal Components Analysis on the types of mossulensis and sellal using 32 morphometric and meristic characters showed some separation between the two taxa and a Discriminant Function Analysis separated most, but not all, specimens. The evidence is not conclusive for separation or synonymy and the taxa are left as distinct in this work.

If mossulensis is a synonym of sellal, then the nominal taxa Alburnus hebes Heckel, 1843, Alburnus microlepis Heckel, 1843 and Alburnus pallidus Heckel, 1843, all from the Kueik (= Quwayq) River at Aleppo (Heckel, 1843b), would have to be added to the synonymy of sellal as indicated by Berg (1949), Krupp (1985c) and Eschmeyer's "Catalog of Fishes" (downloaded 1 September 2007). The 3 syntypes of Alburnus hebes seen by me in the Naturhistorisches Museum Wien were 58.8-156.5 mm standard length (NMW 17558-17560) (but Eschmeyer et al. (1996) list NMW 55523 for these syntypes, and the card index had this number in 1997; possibly they were renumbered). One of these fish is designated as the lectotype. The holotype of Alburnus microlepis (NMW 55655) measures 119 mm standard length (Krupp, 1985c). The holotype of Alburnus pallidus (NMW 55720) measured 76.6 mm standard length.

Krupp (1985c) gives details on the syntypes of Alburnus sellal held at the Naturhistorisches Museum Wien. Six syntypes of A. sellal, 124-140 mm standard length, are under NMW 55665 (2 fish, 137.2-141.3 mm standard length, my measurements) and NMW 55666 (4, 126.9-142.7 mm standard length), and 3, 110-152 mm standard length, are under NMW 55664 (1, 110.5 mm standard length) and 55667 (2, one of which is designated as the lectotype, 140.7-155.4 mm standard length). Eschmeyer et al. (1996) list NMW 55664-67 as having 1, 2, 4, and 2 fish in each number in the series and also 2 syntypes (RMNH 2666) in the Rijksmuseum van Natuurlijke Historie, Leiden from NMW. The catalogue in Vienna lists 8 specimens of A. sellal.

The syntypes of A. mossulensis are under NMW 55656 (2 fish, 111.2-118.4 mm standard length, my measurements), NMW 55717 (2, 83.0-89.4 mm standard length), and NMW 55718 (2, 101.9-131.5 mm standard length). Two syntypes of Alburnus mossulensis are in the Senckenberg Museum Frankfurt (SMF 402, formerly NMW) (F. Krupp, pers. comm., 1985; 80.1-102.7 mm standard length). Eschmeyer et al. (1996) also list NMW 77723 (2, 90.4-135.4 mm standard length) and 1 possible syntype in the Rijksmuseum van Natuurlijke Historie, Leiden (RMNH 2644). The catalogue in Vienna lists 6 specimens of A. mossulensis, with one specimen from NMW 77723 as the lectotype.

Seven syntypes of Alburnus iblis are in the Naturhistorisches Museum Wien under NMW 55524 and measure 91-165 mm standard length (Kähsbauer, 1964; 92.9-172.3 mm standard length by my measurements). One of these fish is designated as the lectotype. The catalogue in Vienna lists 8 specimens in one column and 38 in the adjacent column.

Two syntypes of Alburnus megacephalus are under NMW 55627 and measure 160-162 mm standard length (Kähsbauer, 1964; 162.9-166.1 mm standard length by my measurements); 2 specimens are listed in the Vienna catalogue. One of these fish is the lectotype.

Fifteen syntypes of Alburnus caudimacula are under NMW 55506 and measure 38.5-118.4 mm standard length; the catalogue in Vienna lists 8 specimens in one column and what appears to be 26 specimens in the adjacent column although this may be 20 fish with 6 set aside for A. schejtan. The Rijksmuseum van Natuurlijke Historie, Leiden has 4 syntypes under RMNH 2654, formerly in NMW (Eschmeyer et al., 1996).

Five syntypes of Alburnus capito measure 48.7-101.9 mm standard length (NMW 55505) although the catalogue in Vienna only lists 4 fish.

Four syntypes of Alburnus schejtan measure 71.7-112.6 mm standard length (NMW 22281) and one of these is designated as the lectotype, 2 syntypes measure 104.5-112.3 mm standard length (NMW 55663), 2 syntypes measure 91.8-100.0 mm standard length (NMW 55719), and 2 syntypes measure 81.6-94.4 mm standard length (NMW 55721).

Two syntypes of Leuciscus maxillaris, 165-166 mm total length, are stored in the Muséum national d'Histoire naturelle, Paris (as 13954 according to Fang (1942) or as A.3954 according to Bertin and Estève (1948), M. L. Bauchot, in litt., 1982, and my observations). Fang (1942) regards maxillaris as a distinct species in Alburnus. My measurements were 136.7-136.9 mm standard length.

Krupp (1985c) refers 5 specimens from the type series of Alburnus doriae to his Alburnus sellal and 2 specimens to Leuciscus (= Squalius) lepidus.

Bianco and Banarescu (1982) felt that their samples showed clinal variation from northwest to southeast, with numbers of anal fin branched rays, lateral line scales and gill rakers gradually decreasing. Their fish from the upper Tigris River basin in Turkey not far from Mosul (the type locality) and from the Pulvar River (Kor River basin of Fars) form one subspecies while those from the Mand and Kul River basin draining to the Persian Gulf in Fars are a distinct subspecies. Available names for the former subspecies include capito, iblis, schejtan and megacephalus, the latter requires a new name according to Bianco and Banarescu (1982). The Tigris-Kor sample could be A. mossulensis mossulensis and the Mand-Kul sample A. mossulensis caudimacula (see above). However, Bianco and Banarescu (1982) are correct to point out that variation in this species has not been fully examined, local environmental conditions such as temperature can affect scale counts and the problem of the relationship of A. sellal remains to be resolved. They found in 7 specimens of sellal that scale counts at 71-77 (in contrast to 66-70 in Berg (1949)) overlapped with mossulensis counts. Berg's (1949) and my counts are very wide for A. mossulensis, suggesting that local environment may govern meristic characters as widely demonstrated for fishes. Subspecies recognition requires much further work as Bianco and Banarescu (1982) acknowledge by not proposing a new name for the Mand-Kul fish.

Key characters

The short, naked ventral keel, usually 8 branched dorsal fin rays, distribution, and the characters in the table under A. atropatenae can be used to identify this species.

Morphology

Dorsal fin with 3 unbranched and 7-9 branched rays, anal fin with 3 unbranched and 10-14 branched rays. Pectoral fin branched rays 13-18, pelvic fin branched rays 7-9. Lateral line scales 58-89. Gill rakers 11-18. Pharyngeal teeth 2,5-4,2, with hooked tips and serrated edges to the crowns. Variants include 2,5-5,2, 3,5-5,3 and 2,5-5,3. Populations vary sympatrically in total vertebral counts: 40-43 and 42-45; and in abdominal counts 20-22 and 22-24 (Bogutskaya et al., 2000). The karyotype of fish from the Kızılırmak River in Turkey was 2n=48 (Gül et al., 2000) but this species does not occur in this area.

Meristics for Iranian specimens:- branched dorsal fin rays 7(16), 8(303) or 9(13); branched anal fin rays 10(20), 11(200), 12(104) or 13(8); pectoral fin branched rays 13(2), 14(30), 15(110), 16(134), 17(50) or 18(6), branched pelvic fin rays 7(30), 8(288) or 9(18); lateral line scales 58(1), 59(-), 60(2), 61(1), 62(5), 63(3), 64(9), 65(8), 66(9), 67(12), 68(11), 69(8), 70(18), 71(17), 72(26), 73(21), 74(28), 75(20), 76(19), 77(26), 78(28), 79(21), 80(15), 81(10), 82(5), 83(1), 84(2), 85(2), 86(2), 87(1), 88(1), or 89(1); and total gill rakers 11(7), 12(54), 13(111), 14(101), 15(46), 16(11), 17(1) or 18(1).

Sexual dimorphism

Unknown but males do develop tubercles in the breeding season.

Colour

Overall colour is silvery. The back is a bluish- or reddish-brown, bluish-black or blackish. A dark, lead-coloured stripe runs along and above the mid-flank and has a width about the same as the eye diameter. The stripe may only be evident posteriorly. Scales above the lateral line have fine melanophores at their base. Lateral line scales can have pigment spots above and below the tube near the base of each scale, but this is not as marked as insome Alburnoides spp. The dorsal, anal and caudal fins are margined with black, the latter the darkest. There may be a black spot at the caudal fin base and the first pectoral fin ray may be black dorsally. The pectoral, pelvic and anal fins are yellowish at their base. Pelvic and anal fins may be reddish. The peritoneum is brown but may be thickly speckled with black-brown spots and thus appear almost black.

Size

Reaches about 22 cm (Ergene, 1993).

Distribution

Found in the Tigris-Euphrates basin and adjacent basins. In Iran it is recorded from the Tigris River, Gulf, Lake Maharlu, Kor River and upper reaches of the Hormuz basins (M. Hafezieh, pers. comm.; Berg, 1949; Bianco and Banarescu, 1982; Abdoli, 2000) and questionably from the Esfahan basin (Abdoli, 2000). Records also include the Shapur and Dalaki rivers in the Gulf basin (Gh. Izadpanahi, pers. comm., 1995) and the upper Mand including Qara Agaj reach and Shur tributary, Shur tributary to Dasht-e Palang; upper Zohreh, Marun and Jarrahi, upper Karun and Khersan, Dez, whole middle to upper Karkheh basin (Simarreh, Qarasu, Gav Masiab)(Abdoli, 2000).

Zoogeography

Its former position in the genus Chalcalburnus indicates a relationship with fishes occurring in the Black-Caspian seas basin.

Habitat

This species is found in streams, rivers, lakes, reservoirs and marshes. Al-Habbib (1981) has demonstrated experimentally for specimens taken from the Aloka River, north of Mosul, Iraq that this species can survive temperatures in the range of about 1.25-36.2°C when acclimated (fish were identified incorrectly as Chalcalburnus chalcoides). Epler et al. (2001) found it to be the second most dominant species of fish (identified as A. sheitan) in lakes Habbaniyah, Tharthar and Razzazah in Iraq, comprising 10% of all fish collected. This was one of the most abundant species in the recovering marshes of southern Iraq in 2005-2006 (Hussain et al., 2006).

Age and growth

Jawad (2004) used eye lens diameter for ageing the young (up to age 3) of this species from the marshes north of Basrah. Ergene (1993) studied the growth of this species in the Karasu of Turkey and found 4 age groups, and mentions 5 age groups for another Turkish study. Mean fork length is 118.2 mm, 131.0 mm, 145.2 mm and 163.3 mm respectively. Condition factors for these age groups were 0.87, 0.85, 0.84 and 0.86. Türkmen and Akyurt (2000) also working on this species in the Karasu River found age groups 1 to 6 with age group 3 the most abundant. The mean condition factor for males and females was 1.023 and 1.047 respectively. Age-length, age-weight (von Bertalanffy equations) and length-weight relationships were also calculated as lt = 20.41[1-e-0.2485 (t+1.47)], lt = 21.59[1-e-0.1978 (t+2.13)], W = 80.77 (1-e-0.2485 (t+1.47)2.828, W = 103.63 (1-e-0.1978 (t+2.13)3.082, LogW = -1.796 + 2.828 LogFL (r = 0.943) and LogW = -2.097 + 3.082 LogFL (r = 0.946) respectively. Length and age at first maturity were 1.26 years and 9.24 cm for males and 1.81 years and 9.65 cm for females in the Karasu River, Turkey; age group 7 was the oldest recorded (Yıldırım et al., 2007).

Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 76 Iranian fish measuring 3.15-8.14 cm standard length. The a-value was 0.0197 and the b-value 2.903 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases).

Food

Younis et al. (2001b) found Shatt al Arab, Iraq fish feeding on phytoplankton (algae and diatoms) at 44%, followed by organic detritus at 36.7% (33% in a table), and arthropods at 3.1%, It had a dietary overlap of 89% with Barbus (= Carasobarbus) luteus in May, the highest in the study. In a study of the recovering Hammar Marsh, Iraq diet was 67.95% insects and 14.34% algae with diatoms, plants, crustaceans and fish at less than 10% each, in the Hawr al Hawizah 66.2% insects and 19.2% algae, with amounts of diatoms and crustaceans being less than 10% each, and in the Al Kaba'ish (= Chabaish) Marsh 73.7% insects and 13.1% algae with diatoms, plants and crustaceans at less than 10% each (Hussain et al., 2006).

Reproduction

Berg (1949) reports a female 15.5 cm long with mature eggs. Qarmat Ali River, Iraq fish had a fecundity of 1926-11,779 eggs (Saud, 1997). Yıldırım et al. (2007) examined this species in the Karasu River of Turkey and found a male:female sex ratio of 1:1.08, not significantly different from 1:1, a fecundity range of 3012 to 11,427 eggs, significant correlations between fecundity and fork length, total weight, age and gonad weight, and a spawning season from June to August when water temperature attained 15ºC.

Parasites and predators

Molnár and Jalali (1992) describe a new species of monogenean, Dactylogyrus holciki, from this species in the Beshar River of the Persian Gulf drainage. Gussev et al. (1993b) report the monogenean, Dactylogyrus chalcalburni, from Alburnus alburnus in the Zayandeh Rud but this fish does not occur there. The parasite may have been found in Alburnus mossulensis. González-Solís et al. (1997) report Rhabdochona denudata, Contracaecum sp. larvae and Proleptinae larvae (Nematoda) from this species in the drainage of Lake Maharlu and Contracaecum sp. larvae in the drainage of the Kor River, both in Fars. Jalali et al. (2005) summarise the occurrence of Gyrodactylus species in Iran and record G. sp. from the Beshar River of the Tigris basin in a Chalcalburnus sp., presumably this species. Barzegar and Jalali (2006) report parasites in this species from Kaftar Lake as Lernaea cyprinacea and Diplostomum spathaceum. Barzegar et al. (2008) also record the digenean eye parasite Diplostomum spathaceum from this fish. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Lernaea cyprinacea on this species.

Economic importance

This species has been used in the preparation of fish meal in Iraq.

Conservation

An abundant species where studied, it appears to be under no threat in Iran. Endangered in Turkey (Fricke et al., 2007).

Further work

Its taxonomic status in relation to its Levant relative remains unresolved and the relation between lowland and mountain populations in Iran needs careful analysis. Its biology in Iran has yet to be studied in detail.

Sources

Type material: See above, Alburnus capito (NMW 55505), Alburnus caudimacula (NMW 55506), Alburnus hebes ((NMW 17558-17560 or NMW 55523), Alburnus iblis (NMW 55524), Alburnus megacephalus (NMW 55627), Alburnus microlepis (NMW 55655), Alburnus mossulensis (NMW 55656, 55717, 55718, 77723, SMF 402), Alburnus schejtan (NMW 22281, NMW 55663, 55719, 55721), Alburnus sellal (NMW 55664, 55665, 55666, 55667), Leuciscus maxillaris (MNHN A.3954).

Iranian material: CMNFI 1977-0510A, 44, 35.7-154.6 mm standard length, Fars, Pulvar River tributary (29º59'30"N, 52º54'E); CMNFI 1979-0025, 87, 19.1-138.2 mm standard length, Fars, Kor River at Marv Dasht (29º51'N, 52º46'30"E); CMNFI 1979-0027, 24, 59.8-105.0 mm standard length, Fars, Chehel Cheshmeh (ca. 29º43'N, ca. 52º04'E); CMNFI 1979-0028, 55, 19.1-122.6 mm standard length, Fars, Kor River drainage (no other locality data); CMNFI 1979-0036, 22, 82.3-115.1 mm standard length, Fars, Shapur River at Shapur (29º47'N, 51º35'E); CMNFI 1979-0047, 7, 41.4-78.2 mm standard length, Fars, Ab-e Paravan spring (ca. 29º34'N, ca. 52º42'E); CMNFI 1979-0054, 17, 39.8-95.6 mm standard length, Fars, Shur River tributary (28-29º58-03'N, 52º34-35'E); CMNFI 1979-0061, 51, 32.9-131.1 mm standard length, Fars, Pulvar River tributary (30º04'N, 53º01'E); CMNFI 1979-0067, 55, 11.1-107.9 mm standard length, Fars, qanat at Zarqan (ca. 29º46'N, ca. 52º43'E); CMNFI 1979-0070, 44, 35.0-98.5 mm standard length, Fars, Pulvar River at Naqsh-e Rostam (29º59'N, 52º54'E); CMNFI 1979-0071, 12, 65.3-104.3 mm standard length, Fars, qanat 23 km from Pol-e Khan (ca. 30º00'N, ca. 52º38'E); CMNFI 1979-0073, 26, 50.0-93.3 mm standard length, Fars, Mand River (ca. 29º42'30"N, ca. 52º01'30'E); CMNFI 1979-0074, 39, 23.8-94.0 mm standard length, Fars. Mand River (29º41'N, 52º06'E); CMNFI 1979-0117, 16, 33.4-130.0 mm standard length, Fars, Pulvar River at Naqsh-e Rostam (29º59'N, 52º54'E); CMNFI 1979-0128, 9, 43.2-102.5 mm standard length, Fars, Shur River (28º51'N, 52º31'E); CMNFI 1979-0154B, 1, 46.9 mm standard length, Fars, stream at Koorsiah village (28º45'30"N, 54º24'E); CMNFI 1979-0155, 2, 56.2-64.7 mm standard length, Fars, spring at Gavanoo village (28º47'N, 54º22'E); CMNFI 1979-0156, 11, 49.0-74.4 mm standard length, Fars, qanat at Rashidabad (28º47'N, 54º18'E); CMNFI 1979-0157, 53, 31.8-86.6 mm standard length, Fars, qanat at Hadiabad (28º52'N, 54º13'E); CMNFI 1979-0158, 13, 73.5-108.9 mm standard length, Fars, qanat between Now Bandegan and Qaziabad (28º54'N, 53º53'30"E); CMNFI 1979-0160, 22, 32.4-106.0 mm standard length, Fars, spring at Arteshkhadeh Pomp (29º09'N, 53º37'E); CMNFI 1979-0272, 11, 40.5-130.0 mm standard length, Lorestan, river at Nokhor (ca. 33º40-47'N, ca. 48º28-45'E); CMNFI 1979-0278, 4, 75.5-88.2 mm standard length, Lorestan, Kashkan River drainage (33º34'N, 48º01'E); CMNFI 1979-0279, 3, 68.7-91.4 mm standard length, Lorestan, Khorramabad River (33º37'N, 48º18'E); CMNFI 1979-0282, 19, 40.2-131.3 mm standard length, Lorestan, river at Nurabad (34º05'N, 47º58'E); CMNFI 1979-0284, 30, 73.1-98.3 mm standard length, Kermanshahan, Qareh Su drainage (34º16'N, 46º48'30"E); CMNFI 1979-0285, 4, 124.7-136.8 mm standard length, Kermanshahan, Qareh Su drainage (34º26'N, 46º37'E); CMNFI 1979-0289, 2, 125.3-142.1 mm standard length, Kermanshahan, Diyala River drainage (34º28'N, 45º52'E); CMNFI 1979-0290, 4, 146.9-171.4 mm standard length, Kermnanshahan, Diyala River drainage (34º31'N, 45º35'E); CMNFI 1979-0348, 4, 68.0-78.8 mm standard length, Fars, 2 km from Pol-e Berengie (ca. 29º28'N, ca. 52º32'E); CMNFI 1979-0352, 2, 88.5-93.9 mm standard length, Khuzestan, Jarrahi River drainage (30º33'30"N, 48º48'E); CMNFI 1979-0499, 3, 104.8-133.6 mm standard length, Fars, irrigation ditch on road to Dariush Dam (30º04'30"N, 52º36'E); CMNFI 1979-0500, 2, 112.2-116.5 mm standard length, Fars, Pulvar River at Naqsh-e Rostam (29º59'N, 52º54'E).

Comparative material: BM(NH) 1981.4.13:9-11, 3, 64.3-72.8 mm standard length, Aloka River near Mosul (no other locality data); CMNFI 1980-815, 2, 88.9-107.2 mm; CMNFI 1980-1036, 2, 11.6-145.8 mm standard length, Turkey, ?check this sample for gill raker count ;

Alburnus zagrosensis
Coad, 2009

Common names

None.

Systematics

The holotype (CMNFI 1979-0248) is a male 81.6 mm SL from Chahar Mahall va Bakhtiari, stream, 3 km east of Boldaji, upper Karun River basin (31°55’N, 51°05’E); paratypes are CMNFI 1979-0248A, 128 ♂ 48.4-89.5 mm SL same locality as holotype and CMNFI 1979-0246, 49, 52.5-92.6 mm SL, Chahar Mahall va Bakhtiari, stream, 8 km west of Boldaji, upper Karun River basin (31°57’30”N, 50°59’E). The species is named for the Zagros Mountains of western Iran where it was collected at altitudes over 2300 m.

Key characters

This species is distinguished from other Iranian and Tigris-Euphrates basin Alburnus by such characters as high lateral line scale count (67-83; 64 or lower in alburnus, atropatenae, caeruleus, doriae, filippii), low anal fin branched ray count (9-10; usually 12 or higher in alburnus, caeruleus, chalcoides), and low total gill raker count (12-14; 16 or higher in alburnus, chalcoides). Other characters include a ventral keel almost absent to almost complete, a high frequency of 7 branched dorsal fin rays (35.3%), a high frequency of 7 branched pelvic fin rays (62.7%), a total vertebral count mode at 42, absence of a prominent mid-flank stripe, and small size (females mature at 88.5 mm SL).

Morphology

The body is elongate and a vertical oval in cross-section, somewhat compressed but not deep. The upper and lower profiles are a gentle arch. The snout is short and pointed. The mouth is slightly superior, almost terminal, oblique, and extends back to a level with the mid-nostril. The mouth tip is on a level with the upper half of the eye. The anal fin origin is posterior to the dorsal fin insertion. The dorsal fin margin is rounded and the anal fin margin is straight. The lateral line is decurved and is on the midline of the body only on the posterior half of the caudal peduncle. There is a pelvic axillary scale. The naked ventral keel is flanked for six scale rows anterior to the anal fin.

Dorsal fin with 3 unbranched rays and 7(18) or 8(33) branched rays, anal fin with 3 unbranched rays and 9(29) or 10(22) branched rays, pectoral fin with 14(5), 15(25), 16(15) or 17(6) branched rays, pelvic fin with 7(32) or 8(19) branched rays, lateral line scales to hypural fold 67(1), 69(2), 70(3), 71(2), 72(6), 73(6), 74(2), 75(4), 76(6), 77(7), 78(3), 79(3), 80(2), 81(3), or 83(1), scales around caudal peduncle 20(1), 21(10), 22(18), 23(11), 24(10) or 25(1), predorsal scales 32(1), 33(5), 34(1), 35(8), 36(8), 37(5), 38(9), 39(3), 40(6), 41(4) or 42(1), scales between lateral line and dorsal fin origin 12(1), 13(2), 14(22), 15(17), 16(8), or 18(1), scales between lateral line and anal fin origin 5(6), 6(37) or 7(8), scales between lateral line and pelvic fin origin 4(6), 6(25), 6(15) or 7(5), total gill rakers 12(20), 13(19) or 14(12), short and usually reach past the anterior base of the adjacent raker when appressed, abdominal vertebrae 21(14), 22(34) or 23(3), caudal vertebrae 19(17), 20(32) or 21(2), and total vertebrae 40(4), 41(19), 42(27) or 43(1).

From 0 to 5 scales at the end of the lateral line are not pored. The belly is rounded and has a naked keel in front of the anal fin, varying in extent. The naked keel may be flanked by one to 11 scales, or from almost no keel to a keel extending from the anal fin almost to the pelvic fins base. Scales from below the dorsal fin have fine but not numerous circuli, a subcentral anterior focus and few radii on the anterior and posterior fields with no radii on the lateral fields. Pharyngeal teeth number 2,5-4,2 (6), 2,5-4,1(2) or 1,5-4,2(2). Teeth are hooked at the tip and have serrated edges to the narrow grinding surface below the tip. The gut is an elongate S-shape with a small anterior loop to the left.

Sexual dimorphism

Small tubercles are present on the mature male, on the sides and top of the head, on the dentaries, as fine ones lining the scale margins, most evident on caudal peduncle scales, and on the upper pectoral fin rays. Significant differences between males and females were found in interorbital width (wider in males), caudal peduncle depth (deeper in males), and dorsal, anal, pectoral and pelvic fin lengths (longer in males).

Colour

Fins are mostly immaculate with only slight traces of melanophores. The body is darker dorsally and the upper flank is much darker than the lower flank with an abrupt transition in pigmentation between the two halves along the midline. The anterior flank has a less defined transition with some thin bars extending downward for a short distance. The mid-flank dark pigmentation forms a weak stripe, well-defined by its ventral edge above the anal fin level but the dorsal edge is less well-defined and merges with the upper flank pigmentation. The end of the caudal peduncle has a broad, fan-shaped pigmentation. A broad stripe is present predorsally and postdorsally. The predorsal stripe may be separated into two thin stripes, one on each side of the mid-line with a very thin stripe between them. After 32 years in preservative, this pigment pattern is less evident and the division between the epaxial and hypaxial muscle masses is marked by an apparent thin line of dark pigment as the most prominent feature. The peritoneum is silvery with numerous scattered melanophores grading to fish with continuous melanophores and dark brown overall.

Size

Attains 92.6 mm standard length.

Distribution

Known only from the type locality in the upper Karun River basin.

Empty square = holotype, black square = paratypes, of A. zagrosensis.

Black circles = A. mossulensis.

Zoogeography

This species is related to A. mossulensis which is found in same basin (Tigris-Euphrates) and may have evolved from an isolated population in the upper Karun River.

Habitat

The type locality was at an altitude of 2360 m, water temperature was 22°C at 1545 hours, pH was 6.2 and conductivity 0.45 mS. The stream width was 3-4 m, depth 1 m maximum, the bottom was muddy and the water cloudy. Current was slow, the shore grassy and there was a moderate amount of aquatic vegetation, mostly Myriophyllum. Other species caught were the cyprinid Chondrostoma regium and the tooth-carp Aphanius vladykovi.

The second locality was at an altitude of 2380 m, water temperature was 25°C at 1450 hours, pH was 6.2 and conductivity 0.4 mS. The stream width was 1-2 m, depth 1 m maximum, the bottom was stones and mud and the water muddy. Current was fast, the shore grassy and there was some aquatic vegetation. Some of this material was caught while leaping up a raceway. Other species caught were the cyprinids Chondrostoma regium and a Capoeta sp.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

None.

Conservation

Although known from only two localities in the upper Karun River basin of Iran, this species may not be threatened other than by water abstraction and pollution.

Further work

Distribution, conservation and biology of this species needs investigation.

Sources

See Coad (2009).

Type material: See above.

Genus Aspidoparia
Heckel, 1847

This Oriental genus has only 2 species, one of which enters southeastern Iran. Mirza (2000) proposes that the members of the genus Aspidoparia be placed in a new subfamily, Aspidoparinae. Cabdio Hamilton, 1822 is presumably a senior synonym of Aspidoparia but has not been extensively used in literature on these species so Aspidoparia is retained here.

It is characterised by an elongate and almost cylindrical body with a rounded abdomen, the head has a broad ring of suborbital bones, the mouth is small and inferior, the roof of the mouth has a papillose nodule, the lower jaw has a sharp, crescentic bony edge, no barbels, pharyngeal teeth in 2-3 rows, dorsal fin short, anal fins short to moderate, scales moderate in size, lateral line decurved and running on the lower half of the caudal peduncle, and the gut is long and coiled.

Aspidoparia morar
(Hamilton, 1822)

Common names

None.

[waspi or common chilwa in Pakistan].

Systematics

No relevant synonyms. This species was originally described from the Yamuna and Tista rivers, India. No types are known (Eschmeyer et al., 1996).

Key characters

The suborbital ring of bones is large and distinctive, being almost as deep as the eye, and this feature is unique in southeastern Iranian cyprinids.

Morphology

The snout is short and rounded and overlaps the upper lip. The mouth is small, ventral and transverse. The lower jaw is straight with a slightly horny cutting edge and no lip. The dorsal fin origin is over or slightly behind the pelvic fin origin. The dorsal fin margin is straight to very slightly emarginate and the anal fin is emarginate.

Dorsal fin unbranched rays 2-3 (the first unbranched ray is very small, usually 3 rays are present but not discernible) and branched rays 6-8, anal fin unbranched rays 2 and branched rays 8-10, pectoral branched rays 9-16, usually 12 or more, and pelvic fin branched rays 7-8. Lateral line scales 36-45. Scales have few anterior and more numerous but not many posterior radii. There is a pelvic axillary scale and several elongate and overlapping scales in the pectoral axil. Gill rakers are very short, not touching the adjacent one when appressed, difficult to count at the fleshy ends of each arch, and numbering about 17-25. Pharyngeal teeth 2,4,5-5,4,2 in the literature but the main row count of 4 teeth observed here differs. The main row teeth have large, oval to oblong flattened crowns. The gut is a very elongate s-shape with a small anterior loop. Total vertebrae 36-37. The chromosome number is 2n=48 (Klinkhardt et al., 1995).

Meristic values for Iranian specimens are:- dorsal fin branched rays 6(1) or 7(18); anal fin branched rays 8(4) or 9(15); pectoral fin branched rays 9(1), 12(5), 13(11) or 14(2); pelvic fin branched rays 7(19); lateral line scales 36(1), 37(4), 38(1), 39(5), 40(4), 41(2), 42(1) or 45(1); scales above the lateral line 7(10) or 8(9); scales below the lateral line to the anal fin 3(1), 4(12) or 5(6); scales between the lateral line and the pelvic fin 4(13), 5(5) or 6(1); predorsal scales 17(1), 18(3), 19(2), 20(2), 21(3), 22(3), 23(2) or 24(2); caudal peduncle scales 15(1), 16(5), 17(6), 18(5), 19(1); total gill rakers 17(1), 18(2), 19(2), 21(2), 22(4), 23(2) or 25(1); pharyngeal tooth count 2,4,4-4,4,2(5), 2,4,4-4,4,1(1), or 2,3,4-4,4,2(1), and total vertebrae 36(3) or 37(9).

Sexual dimorphism

Unknown.

Colour

Back light brown to brown-green with the flanks very silvery to silvery-yellow and the belly lighter. There is a golden stripe along the flank. Fins are a distinct dark yellow. The caudal fin may be yellow to orange and paired fins a very light orange-yellow. Preserved fish have immaculate fins except for the caudal fin which has some melanophores lining the rays, a broad stripe along the midline of the back, and fine melanophores on the back and upper flank. Some fish have small, dark dots on the back and upper flank. The peritoneum is black.

Size

Attains 20 cm (Malhotra and Munshi, 1985).

Distribution

This species is reported from the Makran and Mashkid River basins in Pakistan (Mirza, 1992) and eastwards to Thailand. The Iranian distribution encompasses the Mashkel (= Hamun-e Mashkid) and Makran basins, the latter westwards to the Straits of Hormuz (Kiabi and Abdoli, 2000).

    

Zoogeography

The species and genus reaches its westernmost limit of distribution in southeastern Iran. Barriers to further dispersal are unknown but it may be limited by temperature, habitat availability and poor recent connections between streams in the Makran and the southern deserts of Iran.

Habitat

This species favours streams with slow current.

Age and growth

A female, 9.8 cm total length, from Iran had mature eggs (Berg, 1949).

Food

This minnow is a carni-omnivore and a voracious feeder (Bhattacharjee and Dasgupta, 1988). Iranian specimens contained no discernible food items in their guts.

Reproduction

Spawning occurs from February to April in India (Malhotra and Munshi, 1985), Iranian specimens caught in December were not mature suggesting a later spawning season.

Parasites and predators

Jalali et al. (2000) describe two new species of monogeneans, Dactylogyrus yousefpouri and D. mobedii, from this species in the Bahu Kalat River of Baluchestan.

Economic importance

Not of any economic importance in Iran but it is eaten in India.

Conservation

Although known from only a few localities in southeastern Iran, this species may not be threatened other than by water abstraction and pollution.

Further work

The biology of this species, which is at its westernmost range limit in Iran, is unknown. There are some minor differences in characters with literature reports, particularly in pharyngeal tooth count, but sample sizes do not permit an adequate comparison for this wide-ranging species.

Sources

Iranian material: CMNFI 1979-0316, 1, 22.1 mm standard length, Baluchestan, stream in Sarbaz River drainage (26º48'N, 61º02'E);CMNFI 1979-0322, 7, 42.3-86.3 mm standard length, Baluchestan, Dashtiari River (ca. 25º45'N, ca. 61º26'E); CMNFI 1979-0333, 7, 17.7-69.5 mm standard length, Baluchestan, Mashkid River (ca. 27º05'N, ca. 63º12'E); CMNFI 1979-0334, 10, 22.8-62.0 mm standard length, Baluchestan, Mashkid River (27º04'N, 62º54'E): OSU 8123, 5, 45.7-50.6 mm standard length, Baluchestan, Srabaz River (no other locality data).

Comparative material: BC55-61, 2, 67.0-68.2 mm standard length, India, Barakar River near Tillya Dam (no other locality data).

Genus Aspiolucius
Berg, 1907

Aspiolocius esocinus
(Kessler, 1874)

Recorded from the Karakum Canal in Turkmenistan (Sal'nikov, 1995) and may eventually be found in the Tedzhen (= Hari) River basin of Iran. No Iranian record.

Genus Aspius
Agassiz, 1832

The asps comprise 2 species found in Europe and Southwest Asia. Both species are found in Iran.

This genus is characterised by an elongate, rounded and large body, small scales, a large mouth with the lower jaw projecting, lower jaw with a tubercle fitting into a notch in the upper jaw, no barbels, pharyngeal teeth in 2 rows, pointed and hooked, gill rakers short and wide apart, short dorsal fin without a thickened ray, a long anal fin, a scaled keel behind the pelvic fins, and gill slits very wide such that the branchiostegal membranes attach under the posterior end of the eye. Perea et al. (2010) using mitochondrial and nuclear DNA propose the synonymy of this genus with Leuciscus Cuvier, 1816.

Aspius aspius
(Linnaeus, 1758)

Common names

ماش ماهي (= mash mahi, not apparently meaning pea fish as the Farsi could indicate), khasham.

[hasam or khasham in Azerbaijan; krasnogubyi zherekh or redlip asp in Russian; Caspian asp, South Caspian asp].

Systematics

Cyprinus Aspius was described originally from lakes of Sweden.

Cyprinus Rapax Leske, 1774 described from Leipzig, Germany, Cyprinus taeniatus Eichwald, 1831 described from the Kura River at Mingechaur, Aspius erytrostomus Kessler, 1877 (sic, sometimes spelt erythrostomus or erithrostomus) described in part from the Caspian Sea and Kura River, Azerbaijan and from the Aral Sea and lower part of the Amu Darya, Uzbekistan, and Aspius transcaucasicus Warpakhovskii, 1895 from the Lenkoran River and Lake Bussadagny, Azerbaijan, are synonyms. Aspius aspius taeniatus (Eichwald, 1831) is the subspecies found in the Caspian Sea.

The types of Cyprinus aspius described from Swedish lakes are unknown (Eschmeyer et al., 1996).

Eschmeyer et al. (1996) give Aspius transcaucasicus Varpakhovskii, 1896, although Berg (1948-1949) gives 1895; possibly the volume year is 1895 but the work did not appear until 1896. Varpakhovskii is a variant spelling in transliteration from the Russian. Syntypes of this synonym are in the Zoological Institute, St. Petersburg under ZISP 10488 (2) and ZISP 10497-48 (sic, in Eschmeyer et al. (1996) but should read 10497-98 with 5 and 2 specimens respectively (Kottelat, 1997)).

Key characters

The subspecies of the southern Caspian Sea is distinguished from the type subspecies of Europe and the northern Caspian Sea since the latter has lower lateral line scale counts of 64-76 as opposed to 62-105, lips never bright red, anal fin branched rays usually 13 instead of 12 (but see Iranian fish below), and height of dorsal fin usually longer than distance from snout tip to posterior edge of preopercle. Characters of the genus and distribution serve to separate it from other cyprinids in Iran.

Morphology

Dorsal fin branched rays 7-10, usually 8, after 2-3, usually 3, unbranched rays, and anal fin branched rays 11-16, usually 12 (but see below), after 3-4, usually 3, unbranched rays, pectoral fin branched rays 14-17 and pelvic fin rays 7-9. Lateral line scales 62-105. The scales have a central focus, fine circuli and few posterior and anterior radii. There is a pelvic axillary scale. There is a scaled keel behind the pelvic fins. The lower jaw tip projects and fits into a notch in the upper jaw. Gill membranes are narrowly attached to the isthmus, almost under the posterior eye margin. Gill rakers 8-11, very short and club-shaped, almost reaching or not reaching half way to the raker below when appressed. Pharyngeal teeth usually 3,5-5,3, sometimes 2,5-5,3 or with 6 teeth in the main row, teeth elongate, compressed and obviously hooked. Gut an elongate s-shape. Vertebrae 49-51. The chromosome number is 2n=50-52 (Klinkhardt et al., 1995).

Meristic values for Iranian specimens are:- dorsal fin branched rays 8(6); anal fin branched rays 13(6); pectoral fin branched rays 18(3); pelvic fin branched rays 8(3); lateral line scales 68(1), 72(2), 73(1), 74(1) or 75(2); total gill rakers 8(1) or 9(2); pharyngeal teeth 3,5-5,3(3); and total vertebrae 50(2) or 51(1).

Sexual dimorphism

Unknown.

Colour

The overall colour is silvery with the back a blackish-olive or greenish-grey. The iris is silvery with a narrow golden circle around the pupil and a little grey pigment on the upper half. Lips are silvery with a little grey over the upper one. Both lips and iris are often bright red. The dorsal and caudal fins are grey and the other fins are transparent without pigment. Fins may be tinged reddish. Peritoneum silvery to brown.

Size

Reportedly attains 1.2 m and 20.0 kg, possibly over 30.0 kg. The largest of 12,000 fish from the lower Kura River was 77 cm total length, males averaged 61 cm and females 64 cm. The average weight of 105,500 fish caught in 1927-1929 was 2.72 kg, females 2.93 kg (based on 1500 fish), males 2.34 kg and the heaviest fish was 5.5 kg (Berg, 1948-1949).

Distribution

Found from the Rhine and north of the Alps in Europe to the drainages of the Black, Caspian and Aral seas including their southern shores.

This species has been reported from Astara to Gorgan Bay in rivers and marshes and in the Caspian Sea of Iran (Nedoshivin and Iljin, 1929; Derzhavin, 1934; Berg, 1948-1949; Abbasi et al., 1999; Kiabi et al., 1999; Abdoli, 2000; Abdoli and Naderi, 2009). Formerly reported from the Anzali Mordab but no longer present (Holčík and Oláh, 1992) although reported from the Siah-Keshim Protected Region of the Anzali Mordab by (1996). Found also in the Aras River Dam (Jolodar and Abdoli, 2004).

Also recorded from the Uzboi lakes, Karakum Canal and Kopetdag Reservoir in Turkmenistan (Shakirova and Sukhanova, 1994; Sal'nikov, 1995) and may eventually appear in the Tedzhen (= Hari) River basin in Iran.

    

Zoogeography

The closest relative of this species lies to the south and indicates a connection between Euro-Mediterranean and/or Black-Caspian-Aral seas basins.

Habitat

In the waters of Dagestan, asp begin to migrate upriver in October, peaking at the end of November and the beginning of December. They overwinter in deep holes, emerging in early spring as rivers flood and move to the spawning grounds. These grounds include river channels, open lake areas with substantial flow and only rarely places weakly overgrown with very coarse submerged vegetation such as reeds and rushes. After spawning the asps return to the Caspian Sea (Shikhshabekov, 1979). Knipovich (1921) reports this species from depths of 14.6-16.5 m, and possibly deeper, in the Iranian Caspian Sea. Riazi (1996) reports that this species is native (resident) to the Siah-Keshim Protected Region of the Anzali Mordab.

Age and growth

Life span in the Volga delta is 7-8 years with the bulk of the population mature at 6 years (Ali, 1974). In the waters of Dagestan life span is 8 years with maturity at 4 years. Mature males and females are 41-58 cm long and weigh 840-2800 g (Shikhshabekov, 1979). Growth is more rapid in the Kura River of Azerbaijan than in other rivers in the former Soviet Union. Fish taken from commercial catches in Iran are mostly 3-6 years old, 38.1-56.7 cm long and weigh 631-2241 g (Razivi et al., 1972) or 3-6 years and 33-63 cm total length (Holčík and Oláh, 1992). Growth is rapid in the latter report, fish reaching 1 kg during the fourth year of life. Maximum life span may be 15 years.

Food

This species is a solitary predator on other fishes such as gobies (Gobiidae) and silversides (Atherinidae), frogs and even ducklings. An Iranian specimen had the remains of a large crustacean in its gut. Young feed on plankton initially but start to take the fry of fishes at 2-3 months. There is little feeding on the spawning migration.

It may catch other fishes by plunging into shoals at the surface and may leap out of the water as a result. Abdoli (2000) reports Scardinius erythrophthalmus, Atherina boyeri and Blicca bjoerkna as food items in Iran. Surface insects are also eaten.

Reproduction

The spawning season in Gilan is mid-February to late March at 10-13°C with an incubation period of 9-10 days (Hoseinie, 1995).

Spawning is non-intermittent and the period is short (10-15 days) in Dagestan (Shikhshabekov, 1979). Fecundity reaches 483,500 eggs in the south Caspian Sea and maximum egg diameter in the Volga delta is 1.7 mm (Ali, 1974). In Hoseinie's (1995) study of artificial propagation of this species in Iran, large or swollen eggs number 117-277 per gram, and egg diameters 2.0-2.2 mm. Absolute fecundity reaches 264,248 eggs. Abdurakhmanov (1962) gives a maximum fecundity of 342,000 eggs and a maximum egg diameter of 2.4 mm for Azerbaijan populations. Females with ripe eggs are found between mid-April and mid-May at water temperatures of 4-12.2°C, optimally 9-11°C. Up to 20% of Volga asp females do not spawn annually. Eggs develop while between or adhering to stones on the river bed. Young migrate downriver from June to August at age 3-4 months and 5-10 cm length.

Parasites and predators

Molnár and Jalali (1992) record the monogenean Dactylogyrus tuba from this species in the Safid Rud. Masoumian et al. (2005) report the protozoan parasite Chilodonella, sp. from this species in the Aras Dam in West Azarbayjan. Masoumian et al. (2002) investigated parasites from this fish in the Aras and Mahabad dams in northwest Iran and found the protozoan Myxobolus dispar. Sattari (2004) records the presence of the nematode, Eustrongylides excisus, in the body cavity. This parasite can damage muscles in commercial species and render them unsuitable for sale. Sattari et al. (2002, 2004, 2005) and Sattari (2004) records the presence of the nematode, Eustrongylides excisus. This parasite can damage muscles in commercial species and render them unsuitable for sale. Pazooki et al. (2007) recorded various parasites from localities in West Azarbayjan Province, including Argulus foliaceus from this species.

The Caspian seal, Pusa caspica, is a predator (Krylov, 1984).

Economic importance

This fish is taken in Iran as food but comprises only a small portion of the catch. Nevraev (1929) reports catches of 267 to 2429 fish for the period 1914-1915 to 1917-1918 in the Anzali region. Holčík and Oláh (1992) record the catch in the Anzali region for 1969-1970 and 1970-1971 as 45.2 t and 36.1 t respectively, these being 84% and 69% of the total Iranian catch. In 1921-1930 the annual catch in the lower Kura River averaged 249,000 fish and in 1936 for Azerbaijan the catch weighed 8100 centners and numbered 300,000 fish.

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use as food and in sport. The flesh is white and tasty but rather tough.

Conservation

Recruitment in this species is low in Iran because water is taken from the summer spawning streams for irrigation purposes. Spawning success is therefore limited. Larvae of spring spawners are lost when they enter irrigation channels and become stranded in fields (Razivi et al., 1972). Holčík and Oláh (1992) consider the decline in this species to be due to indiscriminate catching of sexually immature fish and, in the Anzali Mordab at least, environmental changes. The Pol-e Astaneh Fish Farm has studied propagation of this species (Keivany and Nasrollahzadeh, 1990) and Hoseinie (1995) demonstrates that artificial propagation is possible. It has also been raised to marketable size in ponds through artificial feeding with ground kilka and a rice product (Annual Bulletin 1993-94, Iranian Fisheries Research and Training Organization, Tehran, p. 81-82, 1995). The Shahid Beheshti hatchery on the Safid River breeds this species (Raymakers, 2002).The asp is bred in the Varvarinsk Hatchery and releases up to 1.5 million yearlings are made into the Kura River, with plans for 8-10 million releases (Kosarev and Yablonskaya, 1994).

Lelek (1987) classifies this species as vulnerable to endangered in Europe. Vulnerable in Turkey (Fricke et al., 2007). Kiabi et al. (1999) consider this species to be data deficient in the south Caspian Sea basin according to IUCN criteria. Criteria include commercial fishing, habitat destruction, limited range (less than 25% of water bodies), absent in other water bodies in Iran, and absent outside the Caspian Sea basin.

Further work

The distribution and abundance of this species in Iranian waters needs investigation as it is sensitive to environmental changes.

Sources

Iranian material: CMNFI 1970-0526, 2, 236.8-246.1 mm standard length, Gilan, Safid River below Astaneh (37º19'N, 49º57'30"E); CMNFI 1980-0494, 1, 319.6 mm standard length, ? Gilan, Caspian Sea basin (no other locality data); ZISP 3917, 1, 402.0 mm standard length, Gilan, Anzali (no other locality data).

Aspius vorax
Heckel, 1843

Common names

shelej, shalaj, sholge, sholgeh.

[shillik, shillig, shiliq, shelej, shalaj; bu aliawi, abu elawi; called "snake" by American soldiers in Iraq because of the name asp being familiar as the snake that killed Cleopatra; kaschschasch (= voracious) from Heckel (1843b); all in Arabic; Tigris asp].

Systematics

The type locality for this species is the "Tigris bei Mossul" according to Heckel (1843b). Krupp (1985c) reports, and I have examined, a syntype held in the Naturhistorisches Museum Wien under NMW 76776, 261.4 mm standard length. The catalogue in Vienna in 1997 also lists NMW 76785 as a type and this specimen is also 261.4 mm standard length. Eschmeyer et al. (1996) lists a dried skin as a syntype under NMW 16527. The catalogue in Vienna lists 4 fish in spirits and 2 fish stuffed.

Banister (1980) suggests that this species may be close to Aspius aspius, perhaps a clinal variant, since the Caspian Sea basin subspecies, A. a. taeniatus (67-90) has scale counts intermediate between European populations of A. aspius (65-74) and A. vorax (93-105) (Banister's figures). However this may be more apparent than real as there is considerable overlap and frequency distributions are not given. There was insufficient material on hand from Iran to investigate this character in more detail.

Key characters

Characters of the genus coupled with distribution serve to identify this species.

Morphology

The head is long and tapers anteriorly. The mouth is oblique and elongate reaching to the anterior half of the eye. The lower jaw projects and has a symphysis knob fitting into an upper jaw notch. There is a hump as the back rises abruptly after the head. The gill opening is large and extends forward to the posterior eye margin level. Fins are more falcate than in the line illustration when partially collapsed.

Dorsal fin with 2-3 unbranched and 7-9, usually 8, branched rays. Al-Nasiri et al. (1975) give a range of 8-11 (probably 7-10 using my system of counting) dorsal fin rays with a strong mode at 9 (i.e. 8) for 271 fish taken from the Basrah fish market from January to June. Anal fin with 2-3 unbranched and 9-13 branched rays. Al-Nasiri et al. (1975) give a range of 10-13 (9-12, 10 modally but high frequencies at 11 too. Pectoral fin branched rays 16-18 (14-18, modally 16, in Al-Nasiri et al. (1975)), pelvic fin branched rays 8-9, usually 8. Lateral line scales 82-110, lateral line low on the flank anteriorly, rising to the midline of the caudal peduncle. There is a pelvic axillary scale. Scales have a few radii on the posterior field only, a central focus and numerous, fine, concentric circuli. Pharyngeal teeth 3,5-5,3 with variants 2,5-5,3 and 2,5-5,2, long, compressed and hooked at the tip. Gill rakers 9-14, reaching base of adjacent raker when appressed but widely spaced and not developed anteriorly. Some rakers do reach the adjacent one when appressed in some fish. Al-Nasiri et al. (1975) give a range of 11-13 gill rakers with a strong mode at 12. Total vertebrae 51-53 (Al-Nasiri et al. (1975) give 37 as a count which cannot be reconciled with my counts). The gut is an elongate s-shape.

Meristic values for Iranian specimens are:- dorsal fin branched rays 8(4); anal fin branched rays 10(1) or 11(3); pectoral fin branched rays 16(1) or 17(3); pelvic fin branched rays 8(4); lateral line scales 96(1), 98(1) or 100(1); total gill rakers 11(1), 12(2) or 13(1); pharyngeal teeth 3,5-5,3(3); and total vertebrae 51(3) or 53(1).

Sexual dimorphism

Unknown.

Colour

The back is greenish to blackish but overall colour is silvery-grey or silvery-white. Fins are said to be all pale yellow in live fish but are dark in some preserved specimens. A photograph of one freshly caught specimen showed reddish pectoral, pelvic and anal fins, with the dorsal fin greenish, similar to the back and flanks. Another freshly caught specimen was overall silvery, with a brownish-green back, fins overall grey with some yellowish tinges The peritoneum is black to brown.

Size

Reaches over 55.0 cm total length and 6.0 kg in Iraq (van den Eelaart, 1954; Herzog, 1967; Shafi and Jasim, 1982; Bartel et al., 1986) and 1.5 m and 60 kg in the Euphrates (Gruvel, 1931; if identification is correct). The Suq al-Shouykh Marsh in April 2005 contained specimens larger than 65.0 cm (www.iraqmarshes.org, downloaded 29 August 2005) and fish in Baghdad palace ponds were estimated to reach 36-40 inches (91-1.02 m) and 15-20 pounds (6.8-9.1 kg) (http://members.cox.net/flybox/FishingUpdate.htm, downloaded 9 January 2006).

Distribution

This species is found in the Tigris-Euphrates and the Orontes River basins in the Middle East. In Iran it is recorded from the lower reaches of rivers in the Tigris River basin including the Bahmanshir River and also such marshes as the Hawr al Azim (Marammazi, 1995).

    

Zoogeography

This is one of several species that has a sister taxon in the Euro-Mediterranean and/or Black-Caspian-Aral seas basin, indicating north-south connections in the past.

Habitat

van den Eelaart (1954) studied this species in Iraq and found that it lives in rivers, lakes and marshes in both open and vegetated areas and remains in shallow water even in summer. It also occurs in smaller water bodies such as ponds. From spring to fall it is found mainly in marshes and lakes. The barrages at Hindiyah and Kut blocked the upstream migration of this species (Mahdi, 1962). Lakes at Camp Slayer in Baghdad contain this species and, in the shallows, the larger fish chase smaller fish and smaller species leaving v-shaped wakes with the tail fin exposed. Smaller fish leap out of the water to escape the shillik (http://members.cox.net/flybox/FishingUpdate.htm, downloaded 9 January 2006).

Age and growth

Shafi and Jasim (1982) made observations on the biology of this cyprinid in Habbaniyah Reservoir, Iraq. They report 8 age groups with most rapid growth in summer months when water temperatures are above 25°C. Growth in weight is about 160.1 g per year to the fourth year of life and about 331 g per year afterwards. Condition factor was 0.74-1.18 with a mean of 1.0, stable values probably related to piscivory. The length-weight relationship was W = 0.0123 x TL3.0601. The von Bertalanffy equation for growth was lt = 91.0[1-e-0.122 (t-0.25)]. Ali et al. (1986) found the condition factor to range from 0.05 to 1.09 (mean 0.73) and also gave the chemical composition and calorific value. This species had a higher fat content than Barbus (= Carasobarbus) luteus with which it was studied. Al-Dabical and Al-Daham (1995) studied growth in the first year of life in fish from the Shatt al Basrah Canal, Iraq and gave the length-weight relationship as loge W = -12.458 + 3.077 loge L and the growth equation as Lt = 104.118 (1-e -0.0121 (t - 87.871)). Epler et al. (2001) found the oldest age groups to be 5+, 6+ and 7+ in Iraqi lakes Razzazah, Habbaniyah and Tharthar respectively. The mean condition factor was 0.88, 0.76 and 0.87 in lakes Habbaniyah, Tharthar and Razzazah respectively. The von Bertalanffy parameters were for Lake Tharthar L (cm) = 145.5, K = 0.0803, t0 = -0.3269, W (g) = 32099 and n = 3.2249. These indicate rather uniform growth rates, as L is relatively high and K very low. Results were considered more reliable than an earlier study by Jasim (1980) which used inappropriate methods. Annual survival in Lake Tharthar for fish 2.6-5.5 years was 62.0% (Szczerbowski et al., 2001). Productivity was low based on chemical and limnological studies, limiting fish production.

Food

This minnow is piscivorous, feeding almost entirely on fish when adult according to Iraqi studies (Shafi and Jasim, 1982), although aufwuchs may also be found in gut contents. It is mainly a mid-water and benthic feeder with limited predation on surface water organisms (Hussein and Al-Kanaani, 1991). Hussein et al. (1991) examined diet in the Garma Marshes, Iraq and found aquatic insects and crustaceans to be important in young shillig in both summer and winter, with molluscs and fish less important. Even in large shillig, fish were outranked by aquatic insects and in winter by crustaceans as well. Molluscs were a minor food. Shillig rejected certain molluscs while taking others, attributed to variations in shell thickness and a attachment strength to substrates. Liza abu is an important food fish (Al-Shamma'a and Jasim, 1993). Hussein and Al-Kanaani (1989; 1991; 1993) examined the diet of this species in the Al-Hammar Marsh and found a gradually reduced feeding intensity towards the winter months, a highest fullness index in May and lowest in January, and a diet governed by food accessibility and availability. Crustaceans, fish and aquatic insects are the main food items in descending order of importance, with fish most important when using a percentage ranking index in large shillig and even in small shillik by volume. Benthic molluscs were the third most important food for young shillik after crustaceans and fish. In a study of the recovering Hammar Marsh, Iraq, diet was 80.0% fish and 20.0% insects, in the Hawr al Hawizah 47.4% fish and 29.4% insects with shrimps, other crustaceans, algae, diatoms, plants and snails at less than 10% each, and in the Al Kaba'ish (= Chabaish) Marsh 73.0% fish and 16.8% insects with shrimps, other crustaceans, algae and plants at less than 10% each (Hussain et al., 2006). Fish are the main diet item of large shillik and there is a gradual shift from small- to large-sized prey as the shillik grows (Salman et al., 1994). Frogs, molluscs and aquatic plants and algae were also found in stomach contents, with frogs being important to large shillik in terms of prey volume. Plants may be accidental inclusions taken when seizing prey in weed beds. The fish eaten in descending order of importance were Liza abu, Gambusia affinis (sic, probably G. holbrooki), Garra rufa and Cyprinus carpio. The main crustacean eaten was Metapenaeus affinis along with decapods and amphipods. The gill rakers are widely spaced, indicative of a piscivorous diet (Salman et al., 1993) and the gut is a short s-shape, about equal to fish standard length, also indicative of a piscivorous diet (Salman et al., 1994). Hussain and Ali (2006) examined feeding relationships among fishes in the Al-Hammar Marsh and found this species to be a carnivore, 41.9% of the diet being crustaceans, 10.0% insects and 34.1% fishes. Epler et al. (2001) studied the diet of this species in Lake Tharthar, Iraq and found year old shillik to be eating oligochaetes, tendipedids and plants material with only fish in 2-7 year old shillik. Dietary coincidence with bizz was high in Lake Tharthar, 96.1%.

Reproduction

Shafi and Jasim (1982) record possible spawning in January at 10°C in Iraq with a fecundity up to 74,509 eggs, a mean of 1157 eggs/g body weight and egg diameter of about 1.1 mm. van den Eelaart (1954) found this species in deep parts of Iraqi rivers in December-January, entering marshes and lakes in February to spawn at the end of February and the beginning of March. Spawning takes place on gravel beds, the same as those used by Barbus (= Luciobarbus) xanthopterus, but also on plants. Epler et al. (2001) studied reproduction in Iraqi lakes Tharthar and Habbaniyah and found males to achieve maturity in the third year of life at 44.2 cm and females in the fourth at47.2 cm. Spawning occurred in February at 13-14ºC. Fecundity was 92,000 eggs/kg body mass.

Parasites and predators

Jalali and Molnár (1990a) record the monogeneans Dactylogyrus pulcher and D. mokhayeri from this species in the Dez River. Moghainemi and Abbasi (1992) record a wide range of parasites from this species in the Hawr al-Azim in Khuzestan. Mortazaei et al. (2000) record an infection rate of 6.6% with the worm Neoechinorhynchus tylosuri in Khuzestan marshes. Farahnak (2000) and Farahnak et al. (2002) record Contracaecum sp. and Anisakis sp. from this fish in Khuzestan Province. Barzegar et al. (2008) record the digenean eye parasite Diplostomum spathaceum from this fish. It is eaten by Silurus triostegus. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Argulus sp., Ergasilus sp., Ergasilus sieboldi, Lernaea sp., Lamproglena sp. and Lamproglena compacta on this species.

Economic importance

Sharma (1980) reports that shillik were an important fish species at the Basrah, Iraq fish market, accounting for 68,948 kg from October 1975 to June 1977, although this is an order of magnitude less than for the three most important species. Its potential for fish farming may be limited by its small gill area which makes it unfit to maintain gas exchange in oxygen-poor water (Salman et al., 1991). Kassim et al. (1998) found locally-raised Scenedesmus acutus algal cultures at 0.5*106 cell/ml with baker's yeast at 0.05 g/L to be the best formula for raising the rotifer Brachionus calcyflorus as live food for shillik larvae. Growth rate was, however, higher on an artificial diet of boiled eggs and soybean meal (52%) compared to 48%, in contrast to common carp (q.v.).

van den Eelaart (1954) gave the fishing season  in Iraq for this species as December-February (peaking in January) and February and June-November (peaking in February and July-August).

Foreign soldiers in Iraq during 2005 regularly caught this species on angling gear using spoons and streamer flies, e.g. www.carpecapio.com, downloaded 26 August 2005.

Conservation

Few specimens have been caught in Iran and deposited in museums. This may reflect rarity or inadequate collection methods. It was commonly caught by American soldiers in Iraq in 2004 as evidenced by emailed photographs sent to me for identification and is an important food fish in Iraq. Detailed surveys using appropriate equipment are needed to assess its distribution and status in Iran. Vulnerable in Turkey (Fricke et al., 2007).

Further work

Its distribution and status in Iran need so be studied as does its distinction from Aspius aspius.

Sources

Scale counts were taken also from Banister (1980).

Type material: See above (NMW 76776 and NMW 76785).

Iranian material: ZMH 2516, 259.9 mm standard length, Kermanshahan, Karasu-Gamasiab-Seymarreh (no further locality data); uncatalogued, 3, 105.6-282.5 mm standard length, Khuzestan, Hawr al Azim and Dez River, (no further locality data). 

Comparative material: NMW 91020, 1, 170.6 mm standard length, Iraq, Shatt-al-Arab, Basrah (30°30'N, 47°47'E); BM(NH) 1920.3.3:127-146, 28, 69.8-284.7 mm standard length, Iraq, Basrah (30°30'N, 47°47'E); BM(NH) 1920.10.8:1, 1, 182.3 mm standard length, Iraq, Tigris River (no other locality data); BM(NH) 1931.12.21:11, 1, 250.2 mm standard length, Iraq, Mosul (36°20'N, 43°08'E); BM(NH) 1972.3.16:1, 1, 112.1 mm standard length, Iraq, Dokan Lake (no other locality data); BM(NH) 1973.5.21:189-190, 2, 166.2-192.0 mm standard length, Iraq, Shatt-al-Arab (no other locality data); FMNH 51242, 1, 322.6 mm standard. length, Iraq, Halfaya east of Amara (31°49'N, 47°26'E); uncatalogued, 1, 200.8 mm standard length, Iraq, Hawr al Hammar (no other locality data). BM(NH) 1968.12.13:182, 1, 251.7 mm standard length, Syria, Cheria River, tributary to the Orontes River (no other locality data); NMW 90366, 1, 309.0 mm standard length, Turkey, Cermik on the Euphrates River (39°09'N, 39°27'E); NMW 90807, 1, 214.8 mm standard length, Turkey, Devegeçidi Çayi, Tigris River basin (no other locality data);

Genus Barbus
Cuvier and Cloquet, 1816

The barbels, genus Barbus sensu lato, are found in Europe, Southwest Asia and Africa and comprise about 800 species with 15 formerly recognised in Iran. Only a single species is now assigned to this genus.

This genus included a wide variety of species and was something of a catchall, serving to cover groups of species which have not been satisfactorily defined as distinct genera to general acceptance. Some authors recognise genera not recognised by others or regard these genera as subgenera - this necessarily affects the species count above. Characters in Southwest Asian species include a rounded or compressed body of moderate to very large size, large to very small scales (lateral line scale count range is at least 26-103), no scale sheath around the anal fin, scales have moderate to high numbers of radii and numerous fine circuli, the presence of barbels in most species, usually 2 pairs, often 1 pair and sometimes none (and individually variable within species), lips variably developed from thin to thick and fleshy, the lower lip sometimes with a well-developed median lobe (and lip development individually variable within species), the last unbranched ray in the short dorsal fin (usually 7-8 branched rays but sometimes more) is thickened and spine-like and may bear teeth or be smooth, a short anal fin, usually with 5 branched rays (but some have 6), pharyngeal teeth in 3 rows with hooked or spoon-shaped tips but sometimes heavy and massive or molariform, gut short, peritoneum white to brown or black, and colour usually brown without distinctive markings in the form of stripes, bands or spots (Luciobarbus subquincunciatus is an exception).

Bănărescu and Bogutskaya in Bănărescu and Bogutskaya (2003) restrict Barbus to tetraploid species with scales having divergent striae. These species have 7-8, occasionally 9, branched dorsal fin rays, 5 branched anal fin rays, papillose lips and two pairs of barbels. This then excludes species placed in Carasobarbus, Kosswigobarbus, Mesopotamichthys and Tor (see below). Two groups of species can be distinguished in this restricted Barbus according to Bănărescu and Bogutskaya in Bănărescu and Bogutskaya (2003), namely those with 5 pharyngeal teeth in the main row and a papillose lower lip separated from the chin by a groove and those with 4 pharyngeal teeth in the main row and a lower lip without papillae and continuous with the chin, this latter group being formerly recognised as the genus Luciobarbus Heckel, 1843. The European/Caucasian member(s) of Barbus sensu.stricto in Iran is lacerta and of Luciobarbus (treated as a subgenus in Bănărescu and Bogutskaya in Bănărescu and Bogutskaya (2003)) are brachycephalus and capito.

Berrebi and Tsigenopoulos in Bănărescu and Bogutskaya (2003) and Tsigenopoulos et al. (2003) review Barbus using molecular markers. They include Barbus cyri (a subspecies of B. lacerta according to some authors) and B. lacerta in the subgenus Barbus, their Northern Mediterranean Group, and B. brachycephalus, capito, esocinus, longiceps, mursa, mystaceus, pectoralis, rajanourum, subquincunciatus, xanthopterus and probably barbulus, kersin, sheich and scincus in the subgenus Luciobarbus, their Southern Group. Levin (2004) studied phenetic relationships of 7 Caucasian taxa and concurred with the division into Barbus and Luciobarbus. See under the species Kosswigobarbus kosswigi for a discussion about the genus/subgenus Kosswigobarbus.

The genus Barbus sensu lato Cuvier and Cloquet, 1816 has been split into a number of genera which are now finding general acceptance. Names used in Southwest Asia include Tor Gray, 1834 sensu Karaman, 1971, Labeobarbus Rüppell, 1836, Systomus McClelland, 1838, Luciobarbus Heckel, 1843, Barynotus Günther, 1868 (preoccupied), Aspiobarbus Berg, 1932, Bertinius Fang, 1943 (and Bertinus Banister, 1980, a misspelling), Bertinichthys Whitley, 1953 (an unneeded replacement of Bertinius), Mesopotamichthys Karaman, 1971, Carasobarbus Karaman, 1971 and Kosswigobarbus Karaman, 1971. Labeobarbus is generally considered to be a synonym of Tor, species of which are found mostly in the Oriental Realm, with only Tor grypus in Iran being a member of the genus Tor (Karaman, 1971; Ekmekçi and Banarescu, 1998). Bertinius is regarded as a synonym of Luciobarbus in Bănărescu and Bogutskaya in Bănărescu and Bogutskaya (2003). A summary table of generic and/or subgeneric names is given below:-

Species

Original genus

Proposed genus or subgenus

barbulus

Barbus

Luciobarbus

brachycephalus

Barbus

Luciobarbus

capito

Cyprinus

Luciobarbus

esocinus

Luciobarbus

Luciobarbus

grypus

Barbus

Tor

kersin

Barbus

Luciobarbus

kosswigi

Cyclocheilichthys

Kosswigobarbus

lacerta

Barbus

Barbus

luteus

Systomus

Carasobarbus

mursa

Cyprinus

Luciobarbus

pectoralis

Barbus

Luciobarbus

sharpeyi

Barbus

Mesopotamichthys

sublimus

Barbus

Kosswigobarbus

subquincunciatus

Barbus

Luciobarbus

xanthopterus

Luciobarbus

Luciobarbus

There are also conflicting views on the validity and synonymy of several nominal "Barbus" species. An extensive comparison of these views is not given here (see, for example, Myers (1960), Karaman (1971), Almaça (1983, 1984a, 1984b, 1986, 1990, 1991, 1992, 1994), Krupp (1985c), Howes (1987), Doadrio (1990), Eschmeyer (1990), Berrebi (1995), Berrebi et al. (1996), Tsigenopoulos and Berrebi (2000)). Karaman's studies have not found general acceptance. Author's views conflict, even when examining the same material. Problems include:- the low number of specimens examined (Almaça (1984a; 1986) for example, examined 11 nominal taxa relevant to Iran in detail but averaged only about 6 specimens per taxon, often from a single locality or outside Iranian waters); a wide range in size of individuals of species being compared making age related changes difficult to assess (denticles in the dorsal fin are often lost with age, barbels are shorter, body shape changes, etc); the possibility of sexual dimorphism; possible variation between populations; ecomorphs being recognised as genera (e.g. Luciobarbus was recognised by having 4, as opposed to 5, teeth in the outer pharyngeal tooth row; Bertinius is founded on this condition and development of molar teeth for crushing molluscs - but this may have risen independently in response to an ecological opportunity (see Krupp (1985c)); paedomorphosis and independent origins from a generalised form in different sites (Mina et al., 2001), and the lack of a wide range of new material. An adequate resolution of the systematics of the Barbus sensu lato species in the Tigris-Euphrates basin in particular would require extensive collections of new material from type localities and from the whole basin and comparison of this material with the extant types. Not all types are extant and some that do exist are in poor condition. If this were not complication enough, "Barbus" species are prone to hybridisation with other "Barbus" species and even other genera, further confusing the resolution of the issue. Almaça (1990) cites a hybridization rate of 5.5-6.0% in "Barbus" of the Iberian Peninsula, higher under changed ecological conditions such as the building of dams.

The status of Bertinius longiceps persicus Karaman, 1971 described from the "Karun b. Ahvaz, Persien" (= Karun River at Ahvaz, Khuzestan) on a single specimen is uncertain (lateral line 56-58, gill rakers 22, subterminal mouth, very short barbels, head somewhat higher and suddenly narrowing compared to the type subspecies of the Jordan and Orontes basins, acuminate snout, dorsal fin margin concave). It is not "Barbus" longiceps (F. Krupp, in litt., 1986). The holotype is in the Zoologischen Instituts und Zoologischen Museums der Universität Hamburg (ZMH H2509).

The roe or eggs of species in these genera have been implicated in poisoning (Halstead, 1967-1970) and should be avoided (see under the genus Schizothorax for more information on egg poisoning). Fish should be carefully cleaned in the spawning season to remove the eggs and ensure against contamination of flesh. Severe cases of egg poisoning in other species have resulted in death. Sykes (1927) however, in his account of the travels of Sir John Chardin in Persia (first published in 1686) quotes "Barbel.... the Spawn of them especially is dangerous, being a certain and a violent Vomit, by Reason that the Sun never shines on that Fish, and that it breeds in raw Waters; or because they take it with the Nux Vomica or the Vomiting Nut". Najafpour and Coad (2002) report a case of roe poisoning from eggs of Carasobarbus luteus.

Barbels are found in running water of streams and rivers although some may inhabit ponds, springs and lakes. Most show migrations for spawning. A species called soleymani, possibly a "Barbus" species, was considered to be on the verge of extinction in the Gav Masiab River of the Tigris River basin, through pollution, overfishing, dam building, aquaculture, and introduction of exotics (IranMania.com, 29 December 2006). "Barbus" species in Khuzestan are thought to be the intermediate hosts of Heterophyidae flukes found in humans and carnivores (Massoud et al., 1981).

Kazeraani (1994) gives a short account of Iranian "Barbus" species in Farsi. The common names in Farsi for these fishes generally are سس ماهي (= sos, ses or sas mahi, meaning unknown) and زرده پر (= zardehpar), zardek or zardak and ourange or ourenge (in reference to yellow or orange colorations, probably of the fins).

The origin and movements of "palaearctic" or Euro-Mediterranean "Barbus" species in Southwest Asia have been examined by Banarescu (1976; 1977) and Almaça (1984b; 1988; 1990) and these works should be consulted for further details. These works are not cladistic analyses but groupings of species based on morphological similarities and may be subject to criticism on this account.

The origin of the genus "Barbus" according to these authors lies in East Asia and reached the Euro-Mediterranean region by a Siberian route. "Barbus" became extinct in northern East Asia, Siberia and northern Europe when the climate cooled during either the Pliocene or the Quaternary. Europe was colonised during the Oligocene and it is from Europe through Anatolia that Southwest Asia received many of its "palaearctic" "Barbus". This route of entry probably did not occur before the Pliocene because the Syrian-Iranian Sea, the last connection between the Tethys Sea and the Indian Ocean, blocked passage of primary freshwater fishes into what is now Iran and adjacent regions although a connection between a Balkan-Aegean-Anatolian landmass and Iran was possible during the early Miocene (20-17 MYA). A marine transgression 16.8-11.8 MYA flooding the eastern Paratethys and the rise of mountain barriers led to independent evolution of "Barbus" in the Balkan-Aegean-Anatolian landmass and in the Iranian Plateau. During the late Miocene the eastern marine connection of Paratethys closed (11.8-10.5 MYA) allowing an exchange of "Barbus" between Iran and Anatolia, continuous from that time. The Paratethys became an intracontinental sea, the Sarmatian Sea, with a basin encompassing the present Black, Caspian and Aral seas and neighbouring low-lying areas (Bianco, 1990). The Sarmatian Sea freshened as large rivers entered it during the late Miocene and Pliocene, facilitating dispersal of freshwater fishes. A second route of entry for "Barbus" to northern Iran was via southwestern Siberia and the Aral Sea basin during the early to middle Oligocene. Bănărescu and Bogutskaya in Bănărescu and Bogutskaya (2003) agree on an east Asian origin for "Barbus", dispersing across Siberia and western Asia. The group split into two branches, one forming Barbus sensu stricto and using a dispersal route north of the Ponto-Caspian basin and reaching western Europe and another (Luciobarbus) dispersing across the present-day Mediterranean Sea (see above in discussion of Berrebi and Tsigenopoulos in Bănărescu and Bogutskaya (2003) and Tsigenopoulos et al. (2003) for listing of nominal taxa relevant to Iran in these branches or groups).

A recent overview of "Barbus" systematics restricts the genus to Europe, Southwest Asia and Northwest Africa (Berrebi et al., 1996). Barbus sensu stricto is recognised as a lineage which shares morphological characters, has an ancestral tetraploid origin of 2n=100, and has similar karyotypes, biochemical markers and parasites. Genetic studies indicate four groups of species, namely West European and Ponto-Caspian, Iberian, Northwest African and Levantine. Iberian barbels are found in Spain and Portugal and along within the Northwest African barbels share no species with Iran. The West European and Ponto-Caspian barbels include B. brachycephalus, B. capito and B. mursa, and the Levantine barbels include B. barbulus, B. cyri, B. esocinus, B. lacerta, B. pectoralis, B. rajanorum, B. scincus, B. subquincunciatus and B. xanthopterus. The authors make no comments on the validity of these nominal species and only B. brachycephalus has been examined in detail for karyotypes and/or nuclear markers. This work is continuing and the authors advocate various methods. They note that accurate descriptions of many taxa are lacking and that morphology is still the fastest and most cost-efficient way to identify species. Accurate identification is the foundation for all other studies.

Machordom and Doadrio (2001), using ATPase 6 and 8 and cytochrome b, found differentiation in "Barbus" capito and "B". brachycephalus in the Plio-Pleistocene. A clade of the subgenus Luciobarbus was found for species from the Caucasus (as above), Greece and North Africa compared to the Iberian Peninsula, isolation having occurred after the Messinian salinity crisis 5.5 MY ago when the Iberian Peninsula broke away from Africa.

Berrebi et al. (1996) recommend that Barbus-like species which cannot be allocated to a clearly defined genus should be placed in a genus called `Barbus', surrounded by single quotation marks, until the systematic position is elucidated. In the text of Freshwater Fishes of Iran double quotation marks (") are used for accounts that referred to Barbus in the old sense, including all or part of the the species listed here (see table above).

Barbus lacerta
Heckel, 1843

 

Chame Saqez River, Kordestan, Lake Orumiyeh basin, courtesy of A. Mahjoor Azad

Chame Saqez River, Kordestan, Lake Orumiyeh basin, courtesy of A. Mahjoor Azad (image rotated)

Chame Saqez River, Kordestan, Lake Orumiyeh basin, courtesy of A. Mahjoor Azad (image rotated)

Chame Saqez River, Kordestan, Lake Orumiyeh basin, courtesy of A. Mahjoor Azad (image rotated)

Caspian Sea basin, courtesy of A. Mahjoor Azad

Above photographs courtesy of Atabak Mahjoor Azad

Common names

blizem, bellizem, سس ماهي (= sos or sas mahi), زرده پر (= zardehpar), orenge, sos mahi Kura.

[Kur sirbiti in Azerbaijan; murtsa, murza, muruza, muruz in Transcaucasia generally; mursa in Armenia; shabout moraqqat in Arabic in Iraq; karrid or karad achmar (red frill or shag, probably from the colour and the long barbels) and karrid asrak (= blue shaggy one) according to Heckel (1843b) in Arabic in Aleppo; Kurinskii usach or Kura barbel in Russian].

Systematics

Howes (1987) places this species in Barbus sensu stricto. Karaman (1971) assigns many taxa as subspecies of Barbus plebejus Bonaparte, 1832 (dated correctly 1839 in Eschmeyer et al. (1996), see Bianco (1995a) for details), found throughout Europe and Southwest Asia. Bianco (1995a) considers that Barbus plebejus is restricted to Adriatic drainages of Italy and Croatia. Valiallahi (2006) considers B. plebejus to be present in Iran and distinct from B. lacerta based mainly on body shape, the relative head length, the body depth and the fourth dorsal fin ray. Barbus plebejus kosswigi Karaman, 1971 is described as new from the "Oberer Teil des Tigris-Systems" and "Hamam suyu, Beytusebab-Hakkari" (upper Tigris River basin in Turkey). Almaça (1991) considers it an ecophenotype of his Barbus plebejus scincus since two subspecies of the same species cannot live in the same river basin. Barbus plebejus kosswigi is a secondary homonym of Cyclocheilichthys (= Kosswigobarbus) kosswigi according to Kottelat (1997).

Barbus plebejus ciscaucasicus Kessler, 1877 is from the western drainages of the Caspian Sea south to Dagestan but only Barbus plebejus lacerta Heckel, 1843 is found in Iran. It is recognised here as a full species since its relationships to European and other taxa cannot be determined on material available for this study. Bianco and Banarescu (1982) place specimens from the Aras River near Maku, which are probably this species, in Barbus cyclolepis cyri De Filippi, 1865.

Almaça (1981; 1983; 1984a; 1984b, 1986) gives lacerta specific status, distinguishing it from Barbus plebejus by the strong denticulations on the last dorsal fin unbranched ray, lower denticle density, number of scales in transverse rows, shorter head and pectoral fin, longer snout, lower body, the decrease in height of the branched dorsal fin rays is gradual and the profile of the fin is straight, unusual in Barbus with a strongly denticulated dorsal spine. Almaça recognises two subspecies from Iranian drainages:- lacerta from the Tigris-Euphrates basin (and Aleppo) and cyri from the southern Caspian Sea basin. Berg (1948-1949) also refers Caspian Sea basin specimens to Barbus lacerta cyri but in Berg (1949) has cyri from the Tigris River basin too. Saadati (1977) suggests that Lake Orumiyeh basin Barbus lacerta are a distinct subspecies based on higher scale counts there (72-89) than in the Caspian Sea basin. However, B. lacerta as recognised has a wide range in scale counts (see below) and counting methods can differ to include or not supernumerary scales in the lateral line and small scales at the caudal fin base. Fishes resembling B. lacerta from the Namak Lake basin have higher scale counts than Caspian Sea specimens although sample size is too small for a definitive study. Berg (1948-1949) notes that his B. lacerta cyri is subject to extremely wide variations in such characters as body depth, fin and barbel lengths, dorsal spine denticle numbers (even absent in some very large fish) and lateral line scale counts, among others. A large series of specimens would be needed to resolve these problems, allowing for size and sexual variation, new character discoveries and consistent methodologies. Molecular studies might be helpful.

Barbus Lacerta was described from the "Flüssen Kueik bei Aleppo" (Heckel, 1843b).

The following species are synonyms. Barbus Scincus Heckel, 1843 described from "Aleppo" and later from the "Flusse Kueik bei Aleppo" in Heckel (1846-1849a), Barbus cyri De Filippi, 1865 described from the "Kur presso Tiflis" (= Kura River near Tbilisi, Georgia) (including Barbus cyri var. tiflissica Kamenskii, 1899 described from the "Kura bei Tiflis" and Barbus cyri var. chaldanica Kamenskii, 1899 described from the "Andshigan-tschai unweit Chaldan"), Barbus caucasicus Kessler, 1877 from the Kura and Araks rivers and tributaries, Azerbaijan, Barbus toporovanicus Kamenskii, 1899 described from the "Toporavan See" (= Lake Paravani or Taparavani at 41°26'N, 43°48'E, in the upper Kura River basin of Georgia), bortschalinicus Kamenskii, 1899 described from the "schwarze Flüsschen (Das schwarze Flüsschen fällt in die Bortschala, rechter Zufluss des Chram, Nebenfluss der Kura)(tschernaja rjetschka)", Georgia, Barbus sursunicus Kamenskii, 1899 described from "Sursuna in dem Flüsscheu (sic) Kara-tschai, Nebenfluss der Kura, oder ihrem Zuflusse, erbeutet in einer Höhe von ca 3200', zwischen den Seen Tschaldyr-göll und Tuman-göll, dass kleinere aus dem Flüsschen Abastuman-tschai" (Azerbaijan; later in the same article this species is spelt zurzunicus), Barbus armenicus Kamenskii, 1899 described from the "See Tschaldyr-göll, 6522' und den Kars-tschai" (Sildir Gölü and the Kars-chai, Turkey), and Barbus angustatus Kamenskii, 1899 described from the "Kura, bei Borshom". Barbus toporovanicus first appeared in Kamenskii (1887) as a variety of Capoeta fundulus (see Capoeta capoeta). Type localities from Kamenskii (1899) are, obviously, taken from the German text; there is also an accompanying and preceding Russian text with localities in Latin and Russian which are very similar, although in some cases abbreviated.

Heckel (1843), the original describer, recognised Barbus scincus as close to his Barbus lacerta but with a shorter head, sharply decurved forehead, small mouth, and small eyes, all characters not easily quantified without detailed analysis. Berg (1949) placed it in the synonymy of lacerta. Berg's view is followed here; others are described by Almaça (1983; 1984a, 1986) who favours placing scincus as a subspecies of Barbus plebejus as noted above.

The problem with the conclusions above remains, as pointed out earlier, the lack of new material.

Four syntypes of Barbus lacerta are in the Naturhistorisches Museum Wien (NMW 54227), 1 syntype is in the Senckenberg Museum Frankfurt (SMF 3471, formerly NMW), and 1 syntype is in the Museum für Naturkunde, Universität Humboldt, Berlin (ZMB 3236, formerly NMW, 110.3 mm standard length, examined February 2006; F. Krupp, pers. comm., 1985; Eschmeyer et al., 1996; Bogutskaya in Bănărescu and Bogutskaya, 2003). The Vienna card catalogue in 1997 lists one of NMW 54227 as the lectotype. The Vienna catalogue lists 6 specimens. Bogutskaya in Bănărescu and Bogutskaya (2003) designates 54227-1, 181.6 mm standard length, as the lectotype.

Syntypes of Barbus scincus from "Aleppo", the type locality in Heckel (1843b), are reported in the Naturhistorisches Museum Wien by Almaça (1986) and were also examined by me (NMW 22272, 2 specimens, 97.6-146.7 mm standard length, in poor condition and NMW 54526, 1 specimen, 158.8 mm standard length, designated as a lectotype by F. Krupp, 31 October 1984). Eschmeyer et al. (1996) also list NMW 54525 as a syntype and this fish measured 124.2 mm standard length and had been dried at some point before it was examined by me. The Vienna catalogue lists 4 specimens and the card catalogue in 1997 lists these 4 fish with NMW 54526 as "? lectotype" (sic).

Tortonese (1940) and Eschmeyer et al. (1996) list the holotype of Barbus cyri as in the Istituto e Museo di Zoologia della R. Università di Torino (MZUT N.690).

The lectotype of Barbus armenicus, as established by Berg (1948-1949:Fig. 451), is in the Zoological Institute, St. Petersburg under ZISP 5198 with 3 paralectotypes (Eschmeyer et al., 1996).

The lectotype of Barbus sursunicus is in the Zoological Institute, St. Petersburg under ZISP 14740 as established in Berg (1948-1949:fig. 451).

Abdurakhmanov (1962) compares fish from the Aras and Kura river basins and the Lenkoranchai. Lenkoran fish have fewer scales, longer head length and depth, greater maximum body depth, greater anal fin height, longer pelvic and ventral fins, a longer lower caudal fin lobe, a shorter caudal peduncle length, a smaller eye, and a shorter interorbital width than Kura and Aras fish; Lenkoran fish have a longer predorsal distance, greater caudal peduncle depth, and greater dorsal fin height than Kura fish though Aras fish are the same; Lenkoran fish have the dorsal fin base and postorbital distance less than in Aras, but not Kura, fish. No taxonomic distinction is made for these variations.

Key characters

The spotting on the body is characteristic.

Morphology

The mouth is moderate in size, with moderate to thick tuberculate lips. The median lobe of the lower lip is not developed, being small to absent; however the lip does have a central area which is thicker and distinct from the lips laterally in small fish. Bogutskaya in Bănărescu and Bogutskaya (2003) gives illustrations of lower lip development and variations in head shape. Males were thought to have a straight head profile while in females the profile falls steeply in front of the nostrils but Bogutskaya in Bănărescu and Bogutskaya (2003) found some males with a hump on the snout. Morphology is quite variable. Barbels are thick, the anterior one not extending past the nostril level and the posterior one reaching or exceeding the preopercle level.

Dorsal fin with 3-5, usually 4-5, unbranched rays followed by 7-9, usually 8, branched rays, anal fin with 3 unbranched rays followed by 4-6, usually 5, branched rays. Pectoral fin branched rays 13-19 and pelvic fin branched rays 7-8. Lateral line scales 49-87. Scales are a horizontal oval to rectangular in shape with the anterior margin bearing a central protuberance, and sometimes a wavy form. Radii are numerous on all scale fields around a subcentral anterior focus with few to moderate numbers of circuli (as scales are small). Scales may be irregularly arranged on the flank because of their small size giving different counts depending on whether smaller scales are included in the lateral line count. There is a pelvic axillary scale. Gill rakers 5-13, short and just reaching the one adjacent when appressed. Rakers may not develop on the anterior arch giving a wide range in counts. Vertebrae 39-45. Pharyngeal teeth 2,3,5-5,3,2 with variants 2,3,5-5,3,1, 1,3,5-5,3,2, 1,3,5-5,3,1, 2,3,4-5,3,2, 2,3,5-4,4,2, 2,4,5-4,4,4 and even 1,2,3,5-5,3,2,1. The fourth inner row tooth is usually the largest, slightly larger, or slightly smaller in some, than the third. The fifth inner row tooth is blunt and other teeth are hooked or pointed. Teeth may be slightly serrated and there is a short concave surface below the hook. The last unbranched ray of the dorsal fin is moderately to strongly developed, varying between individuals and populations, with denticle density high (up to 65) along three-fifths to two-thirds of its length. Denticle extent appears to be quite variable. Denticles are proportionately larger in small fish. The tip of the last unbranched ray is thin and flexible. Denticles may be absent in large fish. The gut is elongate with about 2 anterior and 1 posterior loops.

Meristics in Iranian fish: dorsal fin branched rays 8(36); anal fin branched rays 5(36); pectoral fin branched rays 14(5), 15(7), 16(16) or 17(8); pelvic fin branched rays 7(8) or 8(28); lateral line scales 53(1), 55(1), 56(3), 59(3), 60(3), 63(3), 64(3), 65(2), 66(1), 67(1), 69(3), 70(1), 72(1), 74(2), 76(2), 79(1), 82(3), 85(1) or 87(1); total gill rakers 6(1), 7(9), 8(10), 9(7), 10(6), 11(1) or 13(1); pharyngeal teeth 2,3,5-5,3,2(18), 1,3,5-5,3,1(1), 2,3,4-5,3,2(1) or 2,3,4-4,3,2(1); and total vertebrae ?.

Sexual dimorphism

Females have shorter barbels than males (Berg, 1948-1949) and females have longer anal and ventral fins (Bogutskaya in Bănărescu and Bogutskaya, 2003). Tubercle development in males caught on 25-26 June consists of minute tubercles thickly developed on the head top, sides and ventrally, lining the margin of anterior belly scales but also 1-2 tubercles in mid-scale, on anterior flank scales numbering 1-4 becoming 1 tubercle on more posterior scales although most mid-flank scales lack tubercles. Lower flank and lower caudal peduncle scales bear a tubercle. Back scales have a unique tuberculation consisting of a line rather than a rounded tubercle. The line lies centrally on the scale and extends from the margin part way along the exposed scale. Behind the dorsal fin the back scales have the central line and one on each side radiating back and up and back and down. Tubercles on the dorsal, caudal and anal fins are small and follow the fin branching. they are weak to absent on the pectoral and pelvic fins but are found on the first unbranched pectoral ray in two rows. Males are a dark gold dorsally and all fins slightly reddish with a gold iridescence when spawning (Bogutskaya in Bănărescu and Bogutskaya, 2003). Spawning females have reddish ventral and anal fins.

Colour

The overall colour is yellowish to olive-grey (possibly bluish according to Heckel (1846-1849a)) with numerous, regular dark-brown to black spots on the back, upper flank and dorsal and caudal fins or irregular mottling. The spots may form a stripe in young fish. In general appearance, fish may be quite light or almost blackish as pigmentation level varies individually. The back is olive-brown to light or reddish-brown and the flanks silvery to yellowish. The belly and lower head surface are white. The iris is dark to silvery with a narrow silver-golden ring. Barbels are white. The dorsal fin bears dark spots and extended lines of dark pigment on the rays and membranes. These are not clearly arranged as bars. The margin of the caudal fin is dark in some fish and there may be a band on mid-fin. The caudal fin is often speckled with dark spots which do not form clear bars. The pectoral fin has dark spots and there are odd dark spots on the pelvic and anal fins. The peritoneum is a light brown with dense but spaced melanophores.

Size

Reaches 37.5 cm and 460 g, possibly to 550 g.

Distribution

This species is found in the Tigris-Euphrates, Quwaiq and Caspian Sea basins as well as some internal basins of Iran. In Iran, it is recorded from the Caspian Sea basin in the Aras River and its tributary the Qareh Su, from the Astara to the Atrak rivers including the Anzali Mordab, the upper Safid River drainage in the Qezel Owzan and Shahrud, in Tajan, Babol, Haraz, Sardab, Aras, Tonekabon, Pol-e Rud and Safid rivers, in the Lake Orumiyeh basin in the middle to upper Talkheh River, Nazlu Chai, Tatavi and Zarrineh rivers, the Tigris River basin, and the Esfahan basin (Dopolan River)(Günther, 1899; Laptev, 1934; Berg, 1949; Holčík and Oláh, 1992; Shamsi et al., 1997; Abbasi et al., 1999; Kiabi et al., 1999; Ghorbani Chafi, 2000; Abdoli, 2000).

Zoogeography

Almaça (1991) considers that this species arose from the first wave of colonisers to enter West Asia from South Europe but is more recent in origin than such Barbus (= Luciobarbus) species as esocinus and xanthopterus originating from southwestern Siberia.

Habitat

This species is found in fresh waters and is not migratory. It avoids muddy bottoms and prefers sandy or stony substrates (Solak, 1977; Bogutskaya in Bănărescu and Bogutskaya, 2003). These habitats are rich in benthos, cool, with rapid currents and well-oxygenated; however it may congregate in slow waters where temperatures reach 26°C.

Age and growth

Solak (1989a) examined a population of this species in the Aras River in Turkey and found up to 5 age groups. Abdurakhmanov (1962) records 5 years as life span in Azerbaijan. Çalişkan et al. (1999) also found 5 age groups in Çıldır Lake, Turkey (for Barbus plebejus, probably this species). Fish in age group 2 dominated and the largest fish attained 320 mm and 550 g. Maturity is attained at 2 years for males and 3 years for females (Bogutskaya in Bănărescu and Bogutskaya, 2003).

Food

Plant remains, crustaceans such as amphipods and insect remains such as chironomids and dragonfly larvae have been found in gut contents. Abdoli (2000) lists Plecoptera, Ephemeroptera and Chironomidae. Algae is also consumed (Bogutskaya in Bănărescu and Bogutskaya, 2003).

Reproduction

Eggs number up to 19,680 and a diameter of 2.3 mm (Abdurakhmanov, 1962; Bogutskaya in Bănărescu and Bogutskaya, 2003). Spawning may occur 2-3 times in a season judging by oocyte sizes in mature ovaries and occurs from the end of April to August, varying with locality, once temperatures reach 14°C, ceasing if the temperature exceeds 20°C (Bogutskaya in Bănărescu and Bogutskaya, 2003). Small Iranian specimens (130.7-157.7 mm standard length) have eggs of 1.0 mm diameter and 1.1 mm on 9 July and 11 May respectively. Larger eggs were noted in a fish caught on 9 July (1.7 mm). The spawning season is probably spring for large fish.

Parasites and predators

Molnár and Jalali (1992) record the monogeneans Dactylogyrus carpathicus and D. linstowi from Barbus plebejus, presumably this species, in the Safid Rud. Shamsi et al. (1997) report Clinostomum complanatum, a parasite causing laryngo-pharyngitis in humans, from Barbus barbus plebejus, presumably this species. Masoumian et al. (2003) record Myxobolus valdogeli while Pazooki et al. (2003) and Pazooki (2006) record Rhabdochona hellichi, Bothriocephalus gowkongensis, Pseudocapillaria tomentosa, Allocreadium isoporum and Paradiplozoon homoion, all reports from fishes captured in the Tajan and Zarem rivers of Mazandaran. Pazooki et al. (2005) record Trichodina perforata from this species in waterbodies of Zanjan Province. Pazooki et al. (2006) record the monogeneans Dactylogyrus goktschaicus and Gyrodactylus sp. from this fish in Zanjan Province. Barzegar et al. (2008) record the digenean eye parasite Diplostomum spathaceum from this fish. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Ergasilus sp. and Lernaea sp. on this species.

Economic importance

Not commercially important although it does provide sport in mountain areas of the former U.S.S.R.

Conservation

Kiabi et al. (1999) consider this species to be near threatened in the south Caspian Sea basin according to IUCN criteria. Criteria include sport fishing, medium in numbers, habitat destruction, widespread range (75% of water bodies), present in other water bodies in Iran, and present outside the Caspian Sea basin. Mostafavi (2007) lists it as near threatened in the Talar River, Mazandaran. Endangered in Turkey (Fricke et al., 2007).

Further work

The various populations of this species require more detailed study, especially with molecular methods, to determine their taxonomy.

Sources

Type material: ?

Iranian material: CMNFI 1970-0559, 9, 39.7-114.3 mm standard length, Azarbayjan- eBakhtari, Baranduz Chay (ca. 37º25'N, ca. 45º10'E); CMNFI 1979-0271, 2, ? mm standard length, Lorestan, Kashkan River drainage (33º39'N, 48º32'30"E); CMNFI 1979-0289, 1, 131.6 mm standard length, Kermanshahan, Diyala River drainage (34º28'N, 45º52'E); CMNFI 1979-0449, 2, 85.7-92.2 mm standard length, Azarbayjan-e Khavari, river 18 km from Khalkhal (ca. 37º42'N, ca. 48º27'E); CMNFI 1979-0452, ?, ? mm standard length, Azarbayjan-e Khavari, Qezel Owzan River 6 km from Mianeh (37º23'N, 47º45'E); CMNFI 1979-0468, 7, 30.9-96.1 mm standard length, Mazandaran, Haraz River (36º14'N, 52º22'E); CMNFI 1979-0493, 3, ? mm standard length, Mazandaran, stream in Tajan River drainage (36º19'N, 53º23'E); CMNFI 1979-0557, ?, ? mm standard length, (); CMNFI 1979-0558, ?, ? mm standard length, (); CMNFI 1979-0559, ?, ? mm standard length, (); CMNFI 1979-0785, 2, 115.7-134.8 mm standard length, Aazrabayan-e Bakhtari, Shaher Chay (37º27'N, 34º55'E); CMNFI 1979-0786, 1, 84.1 mm standard length, Azarbayjan-e Khavari, Guru Lake (37º55'N, 46º42'E); CMNFI 1993-0125, 1, 83.1 mm standard length, Kermanshahan, Sarab-e Nilufar (34º24'N, 46º52'E); CMNFI 1993-0126, 2, 157.7 mm standard length, Kermanshahan, Sarab-e Yavari (34º28'N, 46º56'E); CMNFI 1993-0128, 1, 130.7 mm standard length, Kermanshahan, Sarab-e Sabz 'Ali Khan (34º25'N, 46º32'E); CMNFI 1993-0136, 1, ?105.5 or 108.2 mm standard length, Mazandaran, Sardabrud (36º39'42"N, 51º22'36"E); CMNFI 2007-0086, 1, 164.4 mm standard length, Azarbayjan-e Khavari, Qareh Su basin near Nir (ca. 38º02'N, ca. 48º00'E); CMNFI 2007-0087, 2, ? mm standard length, Azarbayjan-e Khavari, Qareh Su north of Ardebil (38º22'N, 48º19'E); CMNFI 2007-0088, 2, ? mm standard length, Azarabyjan-e Khavari, Qareh Su east of Lari (38º30'N, 48º03'E); CMNFI 2007-0093, 1, ? mm standard length, Azarbayjan-e Bakhtari, Qotur River south of Khvoy (38º30'N, 44º58'E); CMNFI 2007-0095, 4, 25.9-73.3 mm standard length, Azarbaijan-e Bakhtari, Shahr Chay southwest of Orumiyeh (ca. 37º27'N, ca. 44º56'E); CMNFI 2007-0096, 1, ? mm standard length, Azarbayjan-e Bakhtari, Qasemul River in Baranduz Chay basin (ca. 37º25'N, ca. 45º10'E); CMNFI 2007-0097, 1, ? mm standard length, Azarbayjan-e Bakhtari, Barunduz Chay basin south of Orumiyeh (ca. 37º16'N, ca. 45º08'E); CMNFI 2007-0098, 2, 193.1-227.4 mm standard length, Azarbayjan-e Bakhtari, river south of Mahabad (ca. 36º42'N, ca. 45º41'E); CMNFI 2007-0099, 2, 28.9-132.1 mm standard length, Azarbayjan-e Bakhtari, Kalwi Chay west of Mahabad (ca. 36º35'N, ca. 45º25'E); CMNFI 2007-0100, 1, ? mm standard length, Azarbayjan-e Bakhtari, Kalwi Chay near Piranshahr (ca. 36º44'N, ca. 45º10'E); CMNFI 2007-0103, 3, 43.6-63.7 mm standard length, Kordestan, Zarineh River basin north of Saqqez (ca. 36º18'N, ca. 46º16'E); CMNFI 2007-0104, 2, 54.6-71.2 mm standard length, Kordestan, Zarineh River basin south of Saqqez (ca. 36º12'N, ca. 46º18'E); CMNFI 2007-0105, 2, ? mm standard length, Kordestan, Zarineh River basin south of Saqqez (ca. 36º06'N, ca. 46º20'E); CMNFI 2007-0106, 1, 99.1 mm standard length, Kordestan, Qezel Owzan River basin near Divandarreh (ca. 35º52'N, ca. 47º05'E); CMNFI 2007-0107, 1, 64.6 mm standard length, Kordestan, Qezel Owzan River basin near Bijar (ca. 35º54'N, ca. 47º20'E); CMNFI 2007-0117, 1, ?66.4 mm standard length, Kermanshahan, Gav Masiab basin near Sahneh (ca. 34º24'N, ca. 47º40'E); CMNFI 2007-0117, 1, ?67.2 mm standard length, Kermanshahan, Gav Masiab near Sahneh (ca. 34º24'N, ca. 47º40'E); CMNFI 2007-0118, 1, ? mm standard length, Kermanshahan, Bid Sorkh River between Sangeh and Kangavar (ca. 34º23'N, ca. 47º52'E); USNM 205931 2, 93.0-115.4 mm standard length, Azarbaijan-e Bakhtari, Baranduz River south of Orumiyeh (37º25'N, 45º05'E); ZMH 2634, 1, 130.5 mm standard length, ?, Haraz River.

Comparative material: BM(NH) 1974.2.22:1236, 1, 113.8 mm standard length, Iraq, Karrid Achmar (no other locality data); BM(NH) 1974.2.22:1327-1328, 2, 121.0-129.9 mm standard length, Iraq (no other locality data); BM(NH) 1974.2.22:1349-1350, 2, 63.1-83.0 mm standard length, Iraq, Qizillja River, Lesser Zab and Serokani near Diana, Rowanduz, Greater Zab (mixed sample); BM(NH) 1974.2.22:1351, 1, 146.8 mm standard length, Iraq, Karrid Asrak (no other locality data).

Genus Barilius
Hamilton, 1822

The members of this genus are found from Pakistan to Thailand with one species in the Tigris-Euphrates and adjacent basins. Their systematics is still poorly understood and there may be about 25 species.

This genus is characterised by a compressed but slender and small body, having small to moderate sized scales, a decurved lateral line, running for example on the lower part of the caudal peduncle, lateral line complete, incomplete or absent, a short dorsal fin and a long anal fin, no fin spines, a moderate and terminal mouth, barbels absent or in 1 or 2 pairs, short gill rakers, pharyngeal teeth in 3 rows, and usually with dark bands or spots on the flank.

These fishes are found mostly in mountain streams although some are lowland species.

Barilius mesopotamicus
Berg, 1932

Common names

None.

[sboura iraqia in Arabic, Mesopotamian minnow].

Systematics

The holotype, 44 mm total length and 35.4 mm standard length, is in the Zoological Institute, St. Petersburg (ZISP 23955) and is decoloured. The collection date is given by Berg (1949) as 16.IV.1914, as 3.IV.1914 in the ZISP catalogue and 5.IV.1914 in the jar. The first two dates are probably correct, one old style and one new style. The type locality is "Stromgebiete des Tigris, in (Siaret) Seid-Hassan, an der persisch-türkischen Grenze, unter 33°20'n. Br., 46°20'ö. L. Seid-Hassan liegt am Flusse Gawi, welcher sich mit dem Kundschian (Gundschian)-tschai vereinigt; der letztere mündet in den Tigris". Seyyed Hasan (33°06'N, 46°11'E) lies on a tributary of the Kanjan Cham River near the Iranian town of Mehran on the Iran-Iraq border. The tributary is presumably the Gawi River.

Howes (1980) stated that this species has apomorph characters shared with species assigned to Leucaspius Heckel and Kner, 1858 but this seems unlikely on general morphological grounds (Coad, 1982b) and Bianco and Banarescu (1982) and Liao et al. (2011) concur, the latter also incorporating molecular evidence. It resembles other Barilius in having barbels (none in Leucaspius), a lateral line low on the body (short and mid-body), broad suborbital bones, and flank bars (none) while Leucaspius is unique in having in females a fold of skin in the shape of two, large, rounded papillae around the genital opening. Bianco and Banarescu (1982) state that this species may be generically distinct from South Asian Barilius but do not diagnose a new genus. Bănărescu and Coad (1991) and Bănărescu (1992b) state that its position and biogeographical affinities are uncertain. Berg (1949) considers it closer to Indian species of the genus Barilius than to African ones.

Key characters

The only member of its genus in Iran, this species is easily identified by the pigment pattern, low lateral line, broad suborbital bones and the barbels.

Morphology

The lower jaw bears a small symphysial knob. The mouth is slightly subterminal, oblique and elongate with the mouth corner under the anterior half of the eye. A well-developed barbel has its origin just anterior to the level of the nostril above the upper lip and lies in a groove between the upper lip and the beginning of the suborbital bone series. This barbel can be absent or minute in some fish (females from Habbaniyah, Iraq (Coad and Krupp, 1983)). In addition to these maxillary barbels, a second pair of barbels have their origin slightly above the posterior edge of the mouth in 8 out of 259 fish examined. They are usually rudimentary but may reach 10.7% of head length. Barbels are difficult to see in smaller fish without magnification. The suborbital bone series is large.

Dorsal fin unbranched rays 2-3, usually 3, branched rays 7-9; anal fin unbranched rays 2-3, usually 3, branched rays 10-14, branched pectoral fin rays 11-15 and branched pelvic fin rays 6-8. Lateral line scales 42-58. Lateral line incomplete or complete, rarely terminating at the pectoral fin level. Lateral line decurved and parallel to the ventral body profile from the pelvic fin origin to the caudal peduncle, being 2-3 scales above this profile. On the caudal peduncle the lateral line is below the mid-line while scales on the caudal fin posterior to the hypural plate are perforated in the mid-line. Pectoral and pelvic axillary scales present. Scales are regularly arranged over the whole body but are not strongly imbricate, particularly on the belly and back anterior to the dorsal fin. Anterior flank scales are oval with subcentral anterior focus and a moderate number of circuli. Radii are found principally on the posterior and lateral fields. Anterior field radii are usually absent although 1-2 radii may occasionally be found. Scale radii based on 5 anterior flank scales from 5 fish (40.7-50.7 mm standard length) number 5-11 primary radii, 0-13 secondary radii and 5-23 total radii. Total gill rakers 7-14. Gill rakers are short and rounded, reaching to or part way to the raker below when appressed. Total vertebrae 38-41. Pharyngeal teeth usually 4,5-5,4, often 4,5-5,3 (25% of 20 fish examined), or more rarely in three rows 1,3,5-5,3,1 or 1,4,5-5,4,1. Teeth are hooked at the tip, slender and have a concave grinding surface below the tip. The gut is a simple s-shape.

Meristic values for Iranian specimens are:- branched dorsal fin rays 7(2), 8(32) or 9(2); branched anal fin rays 10(2), 11(19), 12(13) 13(1) or 14(1); branched pectoral rays 11(1), 12(8), 13(23), 14(2), or 15(1); pelvic fin rays 6(1), 7(33) or 8(2); scales in lateral series 42(1), 43(3), 44(3), 45(2), 46(6), 47(6), 48(1), 49(3), 50(4), 51(4), 52(2) or 54(1); total gill rakers 7(4), 8(6), 9(6), 10(7), 11(6), 12(1), 13(1) or 14(1); pharyngeal teeth 4,5-5,4(8), 4,5-5,3(3) or 1,3,5-5,3,1(1); and total vertebrae 39(8), 40(20) or 41(5).

Sexual dimorphism

Unknown.

Colour

Overall colour is a brilliant silver with a golden-yellow glimmer, with the back darker and having a thin median stripe. Scales are highly deciduous and leave a silvery smear on the hand. The flanks have 6-11 roundish dark, grey-green spots, not clearly apparent in live fish. In preserved fish the spots are brown. A median dorsal stripe is variably developed. Fins are lightly pigmented, most melanophores being on the rays rather than the membranes. The anal and paired fins are almost entirely hyaline. The caudal fin may show one or two irregular bars running parallel to the posterior margin. The peritoneum is light to silvery but bears scattered melanophores which give a greyish tinge in preserved fish. Some fish from Iraq (Habbaniyah stream) lacked, or had weakly expressed, flank spots.

Size

Reaches 50.7 mm standard length.

Distribution

This species is found in the Tigris-Euphrates basin, including its Iranian part and the adjacent Gulf basin (Berg, 1932; 1949; Bianco and Banarescu, 1982). Abdoli (2000) also records this species from the Jarrahi, the lower Karun, the lower Dez, the Zohreh, the lower half of the Helleh, and the middle and lower Mand rivers.

Zoogeography

This species is found in the Tigris-Euphrates basin of Turkey, Syria, Iraq and Iran. It does not appear to be common in Turkey, at least in the upper reaches of this basin there, nor in upper reaches of Iranian rivers. The distribution in the Dalaki River of Iran is outside the modern Tigris-Euphrates basin. It is presumably a relict of the late Pleistocene when the Tigris-Euphrates flowed down a drained Gulf receiving tributaries now isolated by the post-Pleistocene rise in sea level (Coad and Krupp, 1983).

Habitat

Found in both running and still water, from small streams only 1 m wide and irrigation ditches to major rivers more than 200 m across. Current is slow to fast but generally an obvious flow is apparent. However one specimen was collected in a fish pond near Ahvaz (ZSM 25701). The collection localities in Iran are all at low altitudes and no fish were taken in Zagros Mountain streams and rivers. Collections were made over mud and pebble substrates in shallow streams or at river margins. The species may also occur at the surface in mid-river but no collections confirm this supposition. Capture temperatures were 12-24°C and conductivity 0.45-10.5 mS. Salinity in drying pools of 20 cm depth in Syria where this species was caught in March had Cl-1 = 390 mg/l and a salinity of 1.5‰ (Coad and Krupp, 1983).

Age and growth

Unknown.

Food

Gut contents include winged insects (Coleoptera, Heteroptera, Thysanoptera and Diptera) and spiders, suggestive of surface feeding (Coad and Krupp, 1983). Abdoli (2000) also reports Hymenoptera, Brachycera and Culicidae.

Reproduction

Most fish were collected in January when eggs were small but developing suggestive of spring spawning. Al-Rudainy (2008) gives an absolute fecundity of about 200 eggs for Iraq.

aParasites and predators

None reported from Iran.

Economic importance

None.

Conservation

This fish is found in suitable habitats of large rivers and in small ditches and does not appear to be in need of conservation. Vulnerable in Turkey (Fricke et al., 2007).

Further work

Molecular or detailed osteological analyses might reveal its relationships to taxa from the Oriental region.

Sources

Type material: See above (ZISP 23955).

Iranian material: CMNFI, 1979-0120, 3, 19.3-50.7 mm standard length, Bushehr, Dalaki River near Konar Takhteh (29º28'N, 51º21'E); CMNFI 1979-0357, 1, 27.6 mm standard length, Khuzestan, Karkheh River drainage (31º34'N, 48º12'E); CMNFI 1979-0363, 11, 21.4-30.2 mm standard length, Khuzestan, Karkheh River (31º52'N, 48º20'E); CMNFI 1979-0365, 7, 20.0-34.4 mm standard length, Khuzestan, Doveyrich River drainage (32º25'N, 47º36'30"E); CMNFI 1979-0367, 1, 34.2 mm standard length, Khuzestan, Meymeh River (32º44'30"N, 47º09'30"E); CMNFI 1979-0368, 29, 21.6-41.9 mm standard length, Khuzestan, Karkheh River (32º24'30"N, 48º09'E); CMNFI 1979-0372, 2, 30.7-33.1 mm standard length, Khuzestan, Dez River near Chogha Zanbil (ca. 32º02'N, ca. 48º30'E); CMNFI 1979-0377, 3, 28.0-39.4 mm standard length, Khuzestan, Karkheh River (ca. 32º57'N, ca. 47º50'E); CMNFI 1979-0378, 7, 31.9-42.4 mm standard length, Khuzestan, stream tributary to Karkheh River (ca.32º48'N, ca. 48º04'E); CMNFI 1979-0380, 10, 25.3-41.0 mm standard length, Khuzestan, stream tributary to Dez River (ca. 32º10'N, ca. 48º35'E); CMNFI 1979-0381, 7, 24.3-31.2 mm standard length, Khuzestan, stream west of Shushtar (ca. 32º10'N, ca. 48º35'E); CMNFI 1979-0382, 4, 25.9-30.8 mm standard length, Khuzestan, Karun River at Shushtar (32º03'N, 48º51'E); CMNFI 1979-0383, 8, 28.6-34.8 mm standard length, Khuzestan, Ab-e Shur drainage (31º59'30"N, 49º06'E); CMNFI 1979-0384, 3, 26.8-40.8 mm standard length, Khuzestan, Ab-e Shur drainage (32º00'N, 49º07'E); CMNFI 1979-0392, 3, 35.0-39.3 mm standard length, Khuzestan, Zard River (ca. 31º32'N, ca. 49º48'E); CMNFI 1979-0396, 35, 25.1-48.8 mm standard length, Khuzestan, Kheyrabad River (30º32'N, 50º23'30"E); ZSM 25701, 1, 36.5 mm standard length, Khuzestan, fishpond near Ahvaz (no other locality data); ISSB uncatalogued, 1, 48.7 mm standard length, Bushehr, Helleh River (ca. 29º20'N, ca. 51º15'E) (Coad and Krupp, 1983).

Comparative material:- BM(NH) 1974.2.22:1256-1267, 11, 33.7-46.2 mm standard length, Iraq, stream between Lake Habanniyah and Euphrates River (ca. 33º22'N, 43º34'E); BM(NH) 1968.12.13:217-220, 4, 18.5-47.4 mm standard length, Syria, Euphrates River at Mayadine (35º01'N, 40º27'E); BM(NH) 1968.12.13:221-236, 16, 30.8-42.4 mm standard length, Syria, Tigris River at Ain Diwar (37º17'N, 42º11'E); SMF 16442, 5, 28.2-35.9 mm standard length, Syria, Nahr Balikh at Jisr Shanine (36º03'N, 39º06'E); SMF 16443, 63, 17.0-34.9 mm standard length, Syria, Nahr Balikh at Jisr Shanine (36º03'N, 39º06'E); ISSB uncatalogued, 4, 32.8-34.4 mm standard length, Turkey, Batman Suyu (ca. 37º55'N, ca. 40º15'E) (Coad and Krupp, 1983).

Genus Blicca
Heckel, 1843

Shutov (1969) places this genus and species in the genus Abramis Cuvier, 1817 on the basis of literature data as does analyses by Shcherbukha (1973) and Howes (1981). Hensel (1978) and Tadajewska (1998) also place this genus in Abramis on the basis of the lateral line system structure, pharyngeal teeth, scale and dermal bone morphology along with data on ecology, behaviour, ontogenesis, osteology and parasitofauna. Hänfling and Brandl (2000) consider Blicca a junior synonym to Abramis based on allozyme data. In contrast, Bogutskaya (1986) using skull morphology reaffirms its generic status.

The white bream genus contains a single species found from Europe to the Caspian Sea basin including Iran.

The genus is characterised by a deep and strongly compressed body; scales absent on the back behind the dorsal fin thus forming a narrow groove; a scaleless keel between the vent and the pelvic fins; pharyngeal teeth in 2 rows; a small, oblique and subterminal mouth; moderate number of gill rakers; scales of moderate size; a short and spineless dorsal fin and a long anal fin; and a light peritoneum.

Blicca bjoerkna
(Linnaeus, 1758)

Blicca bjoerkna, Wikimedia Commons.

Common names

simparak or seamparak (= silver scales, possible meaning since parak is a small feather), سيم نما (sim nama or mahi sim nama, meaning silvery-like fish or sim-like fish in reference to Abramis brama).

[yastigarin in Azerbaijan; Zakavkazskaya gustera or Transcaucasian white bream, Armyanskaya gustera for A. b. derjavini, all in Russian; silver bream, white bream, flat bream].

Systematics

Cyprinus Björkna was originally described from Lake Mälar, Sweden.

Cyprinus Blicca Bloch, 1782 described from lakes in Germany, Cyprinus gibbosus Pallas, 1814 described from the Sura and Volga rivers and Blicca argyroleuca Heckel, 1843 are synonyms. It appears that the latter taxon is first described in Heckel's work on fishes of Syria, but in the section devoted to classification based on the pharyngeal teeth of cyprinids; the taxon is later described from Europe in Heckel and Kner (1858) and is not a Southwest Asian species. Syntypes of Blicca argyroleuca are in the Naturhistorisches Museum Wien under NMW 16901 (2 fish), NMW 54918 (6), NMW 54919 (4) and NMW 54920 (1) (Eschmeyer et al., 1996). The spelling bjorkna is incorrect (Eschmeyer et al., 1996).

The Caspian Sea basin subspecies is Blicca bjoerkna transcaucasica Berg, 1916, described from the lower reaches of the Kura River, Araks, Lenkoran District, Transcaucasia. It is distinguished by "somewhat" fewer rays in the anal fin (17-21) and "a tendency to have" fewer lateral line scales (40-45) than in the type form which mostly has 21-22 anal fin rays and 45-48 lateral line scales (Berg, 1948-1949). Abdurakhmanov (1962) expands these ranges to 17-22 and 40-48 respectively but gives low means (± standard error) for 100 fish from Azerbaijan of 19.88±0.13 and 43.56±0.05 respectively. This may be a valid subspecies but the possibility of clinal variation has not been examined.

Blicca bjoerkna derjavini Dadikyan, 1970 is described from the "Sevdzhur River, (tributary of Araks River, in Armenian SSR) and the canal and lake system connected with it". It is distinguished from transcaucasica by lower mean number of branched dorsal fin rays and branched anal rays, a higher mean lateral line scale count, and various morphometric characters.

Key characters

The scaleless ventral keel, postdorsal groove, long anal fin, lateral line scale count and small and oblique mouth are characteristic.

Morphology

Dorsal fin with 3 unbranched and 7-10 branched rays, usually 8, anal fin with 3 unbranched and 16-24 branched rays. Pectoral fin branched rays 14-16, pelvic fin branched rays 7-9. Lateral line scales 40-55. Scales have numerous fine circuli, an almost central focus, a wavy anterior margin and a crenulate posterior margin, and few primary anterior and posterior radii, as few as 2 in each field (there may be numerous secondary radii which do not reach the focus). There is a pelvic axillary scale. Gill rakers 12-21, touching the adjacent raker when appressed. Vertebrae 37-43. Pharyngeal teeth 2,5-5,2 with variants 2,5-5,1, 1,5-5,2, 1,5-5,1, 2,5-4,2, 2,5-4,1, 1,5-4,1, 3,5-5,2, and 3,5-5,3 (among others, see below and Tadajewska (1998)), weakly hooked (strongly hooked in young), compressed, concave below the tip and smooth (anterior tooth margin serrated in young). In young fish, the first major row tooth may be medial to the second tooth rather than in line. Tadajewska (1998) gives details of tooth development. The intestine is s-shaped with a small anterior loop. The chromosome number is 2n=50 (Klinkhardt et al., 1995; Pourkazemi et al., 2010).

Meristic values for Iranian specimens are:- dorsal fin branched rays 8(49) or 9(1); anal fin branched rays 17(3), 18(15), 19(21), 20(9) or 21(2); pectoral fin branched rays 14(15), 15(24) or 16(7); pelvic fin branched rays 7(1), 8(47) or 9(2); lateral line scales 41(2), 42(10), 43(9), 44(13), 45(10), 46(5) or 47(1); total gill rakers 13(2), 14(25), 15(18), 16(4) or 18(1); pharyngeal teeth 2,5-5,2(3), 2,5-5,1(4), 1,5-5,2(4), 2,5-4,1(1), 2,5-5,0(1), 0,5-5,2(1), 1,5-5,1(1), 1,5-5,0(2), 1,5-4,1(2) or 2,4-4,1(1); and total vertebrae 38(8), 39(33) or 40(12).

Sexual dimorphism

Breeding males have fine tubercles on the top of the head, operculum and lining the exposed scale margins on the flank. There are occasionally tubercles in mid-scale. Small tubercles are found on the pectoral fin rays, 1-3 rows on the unbranched ray, 1-2 on the first branched ray and usually 1 on the other rays, branching to follow the branching rays. Other fins bear fine tubercles following the fin rays. Larger tubercles are found in clumps on the scales overlapping the anal fin base. Tubercles are absent from the belly. Fine unculi are present on the snout, under the eye and between the tubercles on the head generally as well as on the underside of the pectoral fin.

Colour

The back is a bluish-green and the rest of the body silvery. The pectoral and pelvic fins are orange-red with grey tips. The peritoneum is silvery with scattered melanophores.

Size

Reaches 54.5 cm and 2.25 kg.

Distribution

Found from England through Europe north of the Alps and Pyrenees to the Caspian Sea basin. Apparently it does not penetrate to the higher reaches of even major rivers like the Kura and Aras. In Iran it is found from the Aras River (including its middle reaches in Iran) to the Atrak River in the Caspian Sea basin including the Gorgan, Tajan, Babol, Haraz, Sardab, Tonekabon, and Safid rivers, the Anzali Talab (Derzhavin, 1934; Holčík and Oláh, 1992; Nejatsanatee, 1994; Abbasi et al., 1999; Kiabi et al., 1999; Abdoli, 2000; Abdoli and Naderi, 2009).

Zoogeography

This species is part of a northern European and northern Southwest Asian fauna whose zoogeographical history has not been thoroughly researched. The relationships with similar genera are reviewed under the genus.

Habitat

This species is found in the shallows of warm lakes with heavy vegetation and in the slower reaches of rivers including river estuaries in Iran (Jolodar and Abdoli, 2004). It overwinters in deeper water. There was a mass mortality of this species on the Babol Sar beach on 24 June 1963 (USNM 271217).

Age and growth

Growth is slow with maturity attained at 3-5 years and 10-12 cm. Some males may mature at 2 years. Females are much larger than males of the same age. Life span is up to 16 years. Stunted populations comprising large numbers of individuals develop where predators are absent.

Food

Food items include insect larvae such as chironomids, worms and molluscs, and some vegetation. This is a euryphagous species. Young fish feed principally on copepods and cladocerans. Even adults will feed on plankton and it is less of a bottom feeder than Abramis brama.

Reproduction

Spawning in the Volga delta takes place about the beginning of May at around 11°C water temperature but may run from the end of April to the middle of July in the Volga generally. Spawning in the Aras flood plain occurs in the middle of April. Generally spawning occurs later than in Abramis brama and Rutilus rutilus but may overlap and infertile hybrids result. Shallow weedy areas are preferred. Each female is pursued by several males. Fecundity reaches 109,000 eggs and egg diameter 1.44 mm. Eggs adhere to plants or stones on the bottom. There can be 3 spawnings at intervals of 10-11 days when water temperatures are at least 16-17°C. Batch spawning shows much individual variation as well as varying between localities and by year at the same locality.

Parasites and predators

Khara et al. (2006a) record the eye fluke Diplostomum spathaceum for this fish in the Amirkalayeh Wetland in Gilan. Khara et al. (2008) found the eye parasite Diplostomum spathaceum in this fish from Boojagh Kiashar Wetland in Gilan. Barzegar et al. (2008) record the digenean eye parasite Diplostomum spathaceum from this fish. Tajbakhsh et al. (2010) report the nemtode Philometra rischta from fish in the Anzali wetland. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Lernaea sp. on this species.

The Caspian seal, Pusa caspica, is a predator on this species (Krylov, 1984) as are a variety of other fishes such as perch (Perca fluviatilis) and pike-perch (Sander sp.).

Economic importance

Holčík and Oláh (1992) report a catch of 144 kg in the Anzali Mordab in 1990.

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in aquaria and in textbooks.

Conservation

Lelek (1987) classifies this species as intermediate in Europe (liable to be transferred to vulnerable or rare categories if their habitat deteriorates further). Kiabi et al. (1999) consider this species to be of least concern in the south Caspian Sea basin according to IUCN criteria. Criteria include sport fishing, abundant in numbers, habitat destruction, widespread range (75% of water bodies), absent in other water bodies in Iran, and present outside the Caspian Sea basin.

Further work

The biology of this species needs study in Iran.

Sources

CMNFI 1970-0510, 1, 56.0 mm standard length, Gilan, Golshan River (37º26'N, 49º40'E); CMNFI 1970-0522, 4, 40.0-62.6 mm standard length, Gilan, Safid River at Astaneh Bridge (36º16'30"N, 49º56'E); CMNFI 1970-0532, 6, 30.0-63.2 mm standard length, Gilan, Caspian Sea near Bandar-e Anzali (37º28'N, 49º27'E); CMNFI 1970-0553, 4, 62.1-80.1 mm standard length, Gilan, Sowsar Roga River (37º27'N, 49º30'E); CMNFI 1970-0579, 2, 52.6-56.9 mm standard length, Gilan, Old Safid River estuary (37º23'N, 50º11'E); CMNFI 1970-0580, 31, 31.8-86.3 mm standard length, Mazandaran, river near Iz Deh (36º36'N, 52º07'E); CMNFI 1970-0582, 1, 70.9 mm standard length, Mazandaran, Aliabad Reservoir (36º56'N, 54º50'E); CMNFI 1970-0585, 39, 32.4-52.5 mm standard length, Gilan, Nahang Roga River (37º28'N, 49º28'E); CMNFI 1970-0587, 36, 34.6-55.4 mm standard length, Mazandaran, Babol Sar (36º43'N, 52º39'E); CMNFI 1979-0470, 2, 44.5-51.2 mm standard length, Mazandaran, stream west of Alamdeh (36º35'N, 51º43'E); CMNFI 1979-0472, 30, 38.7-69.6 mm standard length, Mazandaran, stream west of Mahmudabad (36º37'N, 52º12'E); CMNFI 1979-0685, 3, 63.1-67.1 mm standard length, Gilan, Safid River (37º24'N, 49º58'E): CMNFI 1980-0117, 1, 80.0 mm standard length, Gilan, Golshan River (37º26'N, 49º40'E); CMNFI 1980-0122, 15, 38.7-45.3 mm standard length, Mazandaran, Nerissi River (36º38'N, 52º16'E); CMNFI 1980-0149, 6, 60.1-63.7 mm standard length, Gilan, Chabak River (37º21'N, 49º50'E).

Genus Capoeta
Valenciennes, 1842

The genus Capoeta has a wide distribution in Southwest Asia and contains about 20 species of which 7 occur in Iran. Its affinities are uncertain and may lie with the European Barbus/Aulopyge group or with Cyprinion and its southern and east Asian relatives (Karaman, 1971; Howes, 1982; Krupp, 1985c; Bănărescu, 1992b).

Varicorhinus Rüppell, 1836 (as used for Southwest Asian cyprinids) is a synonym of Capoeta Valenciennes, 1842 (see Karaman (1969) for further details: Capoeta is distinguished from Varicorhinus of Africa since it has a denticulate last unbranched dorsal fin ray (as opposed to smooth), very small to medium-sized scales (large), lachrymal bone narrow and covering only a small part of the upper side of the rostrum (large and covering most of the rostrum), suborbital bones narrow and long (short and wide), posterior maxillary process not extending back to a level with the centre of the jugal (extends back to a level of the centre of the suborbitals), lower jaw long (short). Scaphiodon Heckel, 1843 has been used for Capoeta and Cyprinion species in Southwest Asia. The nomenclatural status of this genus is reviewed by Bănărescu in Bănărescu (1999).

This genus Capoeta is characterised by a compressed to rounded and moderately elongate body, small to moderately large scales (lateral line counts 37-99), scales at the anal fin base and anus not usually enlarged (sometimes variably enlarged as is the case with certain cyprinids), an inferior, transverse mouth, the lower jaw with a sharp, horny sheath, barbels absent or in 1 or 2 pairs, dorsal fin short (usually 7-9 branched rays) with the last unbranched ray thickened and bearing serrations (serrations sometimes reduced to absent), anal fin short (usually 5 branched rays), gill rakers short, moderate to numerous, pharyngeal teeth in 3 rows with spoon-shaped and truncate tips, a very long and coiled gut (ca. 7-10 times body length), mostly of uniform colour, and a black peritoneum.

The general name for the members of this genus in northern Iran is سياه ماهي (= siah mahi, meaning black fish) while in the south they are called twiny or touyeni and even gel cheragh (= mud-eater, mud-grazer). The name Capoeta is derived from the Armenian and Georgian name for female Capoeta capoeta packed with eggs, namely "Kapwaeti". Other general names for members of this genus shol khar, ghel khar or choul khar, all variant spoken intonations meaning mud eater.

The origin of Capoeta in Southwest Asia follows the same route as the genus Barbus (q.v.).

CMNFI 1977-0510A, 4, mm standard length, Fars, qanat at Naqsh-e Rostam (29º59'30"N, 52º54'E); CMNFI 1979-0026, , mm standard length, Fars, Shapur River at Shapur (29º47'N, 51º35'E); CMNFI 1979-0027, , mm standard length, Fars, Chehel Chashmeh (ca. 29º43'N, ca. 52º02'E); CMNFI 1979-0036, 2, 83.9-118.3 mm standard length, Fars, Shapur River at Shapur (29º47'N, 51º35'E); CMNFI 1979-0043, , mm standard length, Fars, qanat behind Sarvestan (29º16'N, 53º14'E); CMNFI 1979-0044, , mm standard length, Fars, qanat at Mian Jangal (29º09'N, 53º27'E); CMNFI 1979-0053, 6, 47.3-79.5 mm standard length, Fars, Shur River tributary (ca. 28-29º58-03'N, ca. 52º34-35'E); CMNFI 1979-0054, 16, 35.8-127.9 mm standard length, Fars, Shur River tributary (ca. 28-29º58-03'N, ca. 52º34-35'E); CMNFI 1979-0057, , mm standard length, Fars, Shapur River 4 km from Shapur (29º49'N, 51º34'E); CMNFI 1979-0058, 6, 75.6-115.3 mm standard length, Fars, jube over Shapur River at Shapur (29º47'N, 51º35'E); CMNFI 1979-0059, 2, 45.0-50.4 mm standard length, ID, more than one species? Fars, Pulver River 8km south of Sivand (30º01'30"N, 52º57'E); CMNFI 1979-0061, , mm standard length, ID, more than one species? Fars, stream tributary to Pulvar River (30º04'N, 53º01'E); CMNFI 1979-0063, 2, 201.0-206.7 mm standard length, Fars, qanat under Sa'adi's Tomb, Shiraz (29º37'N, 52º35'E); CMNFI 1979-0067, , mm standard length, Fars, qanat at Zarqan (ca. 29º46'N, ca. 52º43'E); CMNFI 1979-0068, , mm standard length, Fars, qanat 12 km from Shiraz on Esfahan road (ca. 29º43'N, ca. 52º34'30"E); CMNFI 1979-0073, 5, 28.9-86.6 mm standard length, Fars, Mand River beyond Chehel Chashhmeh (ca. 29º42'30"N, ca. 52º01'30"E); CMNFI 1979-0074, , mm standard length, Fars, Mand River backwater (29º41'N, 52º06'E); CMNFI 1979-0075, , mm standard length, Fars, Mand River at Pol-e Kavar (29º11'N, 52º41'E); CMNFI 1979-0079, 2, 120.7-149.9 mm standard length, Fars, Mand River 5 km above Band-e Bahman (ca. 29º12'N, ca. 52º38'E); CMNFI 1979-0079, 1, 159.7 mm standard length, Fars, Mand River 5 km above Band-e Bahman (ca. 29º12'N, ca. 52º38'E); CMNFI 1979-0090, , mm standard length, Esfahan, Gav Khuni (ca. 32º21'N, ca. 52º49'E); CMNFI 1979-0093, 1, 73.9 mm standard length, (); BWC 76-77 check CMNFI # CMNFI 1979-0109, 1, 91.1 mm standard length, Fars, Mand River ar Shahr-e Khafr (28º56'N, 53º14'E); CMNFI 1979-0109, 1, 103.4 mm standard length, Fars, Mand River ar Shahr-e Khafr (28º56'N, 53º14'E); CMNFI 1979-0111, 10, 8.7-54.6 mm standard length, Fars, stream 21-22 km from Shiraz (29º37'30"N, 52º21'E); CMNFI 1979-0113, , mm standard length, Fars, qanat under Sa'adi's Tomb (29º37'N, 52º35'E); CMNFI 1979-0114, , mm standard length, Fars, Mand River at road bridge (29º41'N, 52º06'E); CMNFI 1979-0115, 4, 154.4-172.6 mm standard length, qanat under Sa'adi's Tomb (29º37'N, 52º35'E); CMNFI 1979-0125, 1, 137.8 mm standard length, Bushehr, Dalaki River near Dalaki (ca. 29º28'N, ca. 51º21'E); CMNFI 1979-0128, 16, 34.6-108.6 mm standard length, Fars, Shur River (28º51'N, 52º31'E); CMNFI 1979-0128, 18, 17.2-135.3 mm standard length, Fars, Shur River (28º51'N, 52º31'E); CMNFI 1979-0129, , mm standard length, Fars, spring 2 km north of Farrashband (28º54'N, 52º04'E); CMNFI 1979-0130, 5, 44.4-93.3 mm standard length, Fars, Shur River 4 km west of Firuzabad (28º51'N, 52º32'E); CMNFI 1979-0131, 58, 25.5-140.0 mm standard length, Fars, Mand River tributary (28º38'N, 52º49'E); CMNFI 1979-0132, 23, 51.1-74.4 mm standard length, Fars, Mand River tributary (28º35'N, 52º58'E); CMNFI 1979-0154B, 6, mm standard length, Fars, upper Shur River drainage near Darab (28º45'30"N, 52º24'E); CMNFI 1979-0155, 7, 36.2-80.5 mm standard length, Fars, spring at Gavanoo (28º47'N, 54º22'E); CMNFI 1979-0156, 3, 54.6-122.9 mm standard length, Fars, qanat at Rashidabad (28º47'N, 54º18'E); CMNFI 1979-0157, , mm standard length, Fars, qanat at Hadiabad (28º52'N, 54º13'E); CMNFI 1979-0158, , mm standard length, Fars, qanat over Qasook River (28º54'N, 53º53'30"E); CMNFI 1979-0159, 87, 23.1-167.3 mm standard length, Fars, qanat at Qaziabad (ca. 28º54'N, ca. 53º43'E); CMNFI 1979-0160, 4, 66.3-138.4 mm standard length, Fars, Arteshkadeh Pomp spring (29º09'N, 53º37'E); CMNFI 1979-0161, 29, 33.2-88.3 mm standard length, Fars, qanat on Neyriz to Shiraz road (29º10'30"N, 53º41'E); CMNFI 1979-0162, 9, ?-88.3 mm standard length, Fars, qanat behind Sarvestan (29º16'30"N, 53º14'E); CMNFI 1979-0163, 1, 73.8 mm standard length, ?Fars, neighbourhood of Shiraz (no other locality data); CMNFI 1979-0164, 1, 49.4 mm standard length, ?Fars, neighbourhood of Shiraz (no other locality data); CMNFI 1979-0165, 7, 30.0-96.6 mm standard length, Kerman, qanat at Ahmadabad (30º32'N, 55º38'E); CMNFI 1979-0166, 67, 37.1-123.1 mm standard length, Kerman, qanat at Hassanabad-e Nuq (30º43'N, 55º50'E); CMNFI 1979-0168, , mm standard length, Kerman, qanat at Shahabad (29º07'N, 58º16'E); CMNFI 1979-0169, , mm standard length, Kerman, qanat 10 km from Mahan (30º08'30"N, 57º17'E); CMNFI 1979-0170, , mm standard length, Kerman, qanat at Baghin (30º12'N, 56º48'E); CMNFI 1979-0171, , mm standard length, Kerman, qanat at Bardesir (29º56'N, 56º34'E); CMNFI 1979-0187, , mm standard length, Hormozgan, stream and pools at Sar Khun (27º23'30"N, 56º26'E); CMNFI 1979-0191, , mm standard length, Fars, stream 10 km east of Furg (ca. 28º16'N, ca. 55º18'E); CMNFI 1979-0192, , mm standard length, Fars, qanat 2 km east of Rostaq (28º26'30"N, 55º04'E); CMNFI 1979-0195, , mm standard length, Fars, jube on road to Fasa (ca. 28º54'N, ca. 53º53'30"E); CMNFI 1979-0198, , mm standard length, Fars, stream at Tadovan (28º47'N, 53º24'30"E); CMNFI 1979-0199, 6, 70.8-102.1 mm standard length, Fars, qanat 18 km from Jahrom (ca. 28º23-25'N, ca. 53º31-40'E); CMNFI 1979-0202, , mm standard length, Fars, Mand River (29º01'N, 53º00'E); CMNFI 1979-0203, , mm standard length, Fars, qanat at Dudej (29º33'N, 52º59'E); CMNFI 1979-0204, , mm standard length, Fars, qanat on road to Kharameh (29º33'N, 52º59'E); CMNFI 1979-0205, 12, 45.9-200.5 mm standard length, Fars, jube at Runiz-e Pa'in (29º12'N, 53º42'E); CMNFI 1979-0206, , mm standard length, Fars, qanat on road to Kharameh (29º12'N, 53º40'E); CMNFI 1979-0207, 12, 24.2-83.7 mm standard length, Fars, jube 22 km from Neyriz (29º16'N, 54º28'E); CMNFI 1979-0208, 6, 39.9-130.4 mm standard length, Fars, qanat 47 km from Neyriz (ca. 29º11'N, ca. 54º40'E); CMNFI 1979-0209, 60, 43.6-138.9 mm standard length, Kerman, qanat at Kuch Kuluh (29º25'N, 56º03'E); CMNFI 1979-0211, 63, 33.2-94.3 mm standard length, Kerman, river on road to Baft (29º19'N, 56º12'E); CMNFI 1979-0212, 73, 26.0-99.1 mm standard length, Kerman, qanat on road to Baft (29º14'N, 56º17'E); CMNFI 1979-0213, 5, 51.4-60.2 mm standard length, Kerman, stream in Kharan River drainage (29º15'N, 56º25'E); CMNFI 1979-0214, , mm standard length, Kerman, qanat pool on road to Baft (ca. 29º15'N, ca. 56º28'E); CMNFI 1979-0215, 15, 39.7-125.9 mm standard length, Kerman, Kharan River drainage (29º14'N, 56º37'E); CMNFI 1979-0216, 11?, 51.1-65.8 mm standard length, Kerman, qanat 9 km from Baft (ca. 29º13'N, ca. 56º42'E); CMNFI 1979-0217, 15, 39.7-125.9 mm standard length, Kerman, Kharan River drainage (ca. 28º59'30"N, ca. 56º51'30"E); CMNFI 1979-0221, , mm standard length, Kerman, Halil River drainage (28º51'N, 57º52'E); CMNFI 1979-0241, , mm standard length, Fars, Shapur River at Shapur (29º47'N, 51º35'E); CMNFI 1979-0243, , mm standard length, Esfahan, Zayandeh River at Falavarjan (32º33'N, 51º31'E); CMNFI 1979-0246, , mm standard length, Shahrestan-e Bakhtiari va Chahar Mahall, upper Karun River drainage (31º57'30"N, 50º59'E); CMNFI 1979-0251, , mm standard length, Esfahan, stream 1 km east of Daran (32º59'N, 50º26'E); CMNFI 1979-0251, , mm standard length, Esfahan, stream 1 km east of Daran (32º59'N, 50º26'E); CMNFI 1979-0255, , mm standard length, Markazi, Bar River drainage 2 km west of Shahabiyeh (33º51'30"N, 50º23'E); CMNFI 1979-0269, 1, 125.0 mm standard length, Lorestan, Dez or Karkheh River drainage (no other locality data); CMNFI 1979-0271, , mm standard length, Lorestan, Kashkan River drainage (33º39'N, 48º32'30"E); CMNFI 1979-0272, , mm standard length, Lorestan, river at Nokhor (33º40-47'N, 48º28-45'E); CMNFI 1979-0273, 7, 66.7-137.6 mm standard length, Lorestan, Kashkan River drainage (33º26'N, 48º19'E); CMNFI 1979-0274, 3, 28.9-141.8 mm standard length, Lorestan, Kashkan River drainage (33º27'N, 48º11'E); CMNFI 1979-0276, , mm standard length, Lorestan, Kashkan River drainage (ca. 33º19'N, ca. 47º53'30"E); CMNFI 1979-0277, 2, 116.2-133.4 mm standard length, Lorestan, Kashkan River drainage (33º30'N, 47º59'30"E); CMNFI 1979-0278, 3, 93.5-114.7 mm standard length, Lorestan, Kashkan River drainage (33º34'N, 48º01'E); CMNFI 1979-0279, 1, 126.0 mm standard length, Lorestan, Khorramabad River 16 km from Nurabad (33º37'N, 48º18'E); CMNFI 1979-0279, 5, 115.6-155.8 mm standard length, Lorestan, Khorramabad River 16 km from Nurabad (33º37'N, 48º18'E); CMNFI 1979-0280, 3, 104.7-107.7 mm standard length, Lorestan, Kashkan River drainage (33º43-47'N, 48º12-15'E); CMNFI 1979-0282, 6, 110.3-130.3 mm standard length, Lorestan, Seymareh River drainage at Nurabad (34º05'N, 47º58'E); CMNFI 1979-0283, 2, 113.7-125.0 mm standard length, Kermanshahan, Qareh Su drainage (34º21'N, 47º07'E); CMNFI 1979-0285, 3, 125.5-148.0 mm standard length, Kermanshahan, Qareh Su drainage (34º26'N, 46º37'E); CMNFI 1979-0286, , mm standard length, Kermanshahan, Ravansar River at Ravansar (34º43'N, 46º40'E); CMNFI 1979-0287, 2, 128.2-136.1 mm standard length, Kermanshahan, Chashmeh Javari 2 km from Ravansar (ca. 34º42'N, ca. 46º40'E); CMNFI 1979-0288, 62, 37.6-153.7 mm standard length, Ilam and Poshtkuh, Gangir River at Sarab Ewan (33º50'N, 46º18'E); CMNFI 1979-0289, , mm standard length, Kermanshahan, Diyala River drainage (34º28'N, 45º52'E); CMNFI 1979-0290, , mm standard length, Kermanshahan, Diyala River drainage at Qasr-e Shirin (34º31'N, 45º35'E); CMNFI 1979-0291, , mm standard length, Kermanshahan, Diyala River drainage (34º24'N, 45º37'E); CMNFI 1979-0306, , mm standard length, Kerman, qanat on road to Baft (29º13'N, 54º33'E); CMNFI 1979-0307, 5, 50.9-73.4 mm standard length, Kerman, river at Sartal 6 km from Baft (ca. 29º17'N, ca. 56º38'E); CMNFI 1979-0308, 67, 20.5-246.9 mm standard length, Kerman, river 44 km from Baft (29º02'N, 56º50'E); CMNFI 1979-0309, , mm standard length, Kerman, Fahraj River at Azizabad (28º57'N, 58º42'E); CMNFI 1979-0315, 2, 53.5-65.5 mm standard length, Baluchestan, Bampur River 2 km from Karevandar (27º51'N, 60º46'E); CMNFI 1979-0315, 34?, 53.7-85.1 mm standard length, (); note two collections? CMNFI 1979-0337, , mm standard length, Baluchestan, stream near Kanowak (ca. 28º40'N, ca. 60º48'E); CMNFI 1979-0341, 14, 27.2-75.9 mm standard length, Kerman, Tahrud west of Bam (29º23'N, 57º52'E); CMNFI 1979-0343, , mm standard length, Fars, lake near Deh Bid (ca. 30º32'N, ca. 52º49'E); CMNFI 1979-0411, 7, 42.2-76.5 mm standard length, Hormozgan, Minab River past Rudan (27º24'N, 57º12'E); CMNFI 1979-0419, 1, 62.2 mm standard length, Fars, stream 7 km from Rostaq (28º29'N, 55º01'E); CMNFI 1979-0420, 6, 57.1-150.6 mm standard length, Fars, Rudbar River at Bahregan (30º11'N, 52º03'E); CMNFI 1979-0422, , mm standard length, Boyer Ahmadi-ye Sardsir va Kohkiluyeh, stream in Yasuj valley (30º36'N, 51º34'E); CMNFI 1979-0424, , mm standard length, Fars, stream on Yasuj to Nurabad road (30º18'N, 51º30'30"E); CMNFI 1979-0425, , mm standard length, Fars, Haft Barm-e Kudian (29º49'N, 52º02'E); CMNFI 1979-0426, , mm standard length, Esfahan, qanat at Abbasabad-Natanz (33º36'N, 51º49'E); CMNFI 1979-0458, 2, 90.7-108.4 mm standard length, Markazi, Khar River 6 km north of Ab-Garm (35º47'N, 49º20'E); CMNFI 1979-0460, 3, 54.4-65.0 mm standard length, Hamadan, stream 16 km south of Asadabad (34º39'N, 48º05'E); CMNFI 1979-0462, , mm standard length, Markazi, Mazdaqan River (35º06'30"N, 49º40'30"E); CMNFI 1979-0466, , mm standard length, Esfahan, qanat at Meymeh (33º27'N, 51º10'E); CMNFI 1979-0484, , mm standard length, Khorasan, stream 22 km west from Bojnurd (37º28'N, 56º44'E); CMNFI 1979-0497, 3, 49.8-113.0 mm standard length, Fars, Mand River at Band-e Bahman (29º11'N, 52º40'E); CMNFI 1979-0497, 7, 102.2-132.0 mm standard length, Fars, Mand River at Band-e Bahman (29º11'N, 52º40'E); CMNFI 1979-0499, , mm standard length, Fars, ditch 32 km from Kor River bridge (30º04'30"N, 52º36'E); CMNFI 1979-0501, 6, 34.1-110.9 mm standard length, Fars, Mand River at Kavar (29º11'N, 52º41'E); CMNFI 1979-0502, , mm standard length, Fars, Haft Barm-e Kudian (29º49'N, 52º02'E); CMNFI 1993-0126, , mm standard length, Kermanshahan, Sarab-e Yavari (34º28'N, 46º56'E); CMNFI 2007-0030, , mm standard length, Baluchestan, stream near Eskelabad (28º35'N, 60º48'E); CMNFI 2007-0031, , mm standard length, Baluchestan, headwaters of Bampur River (27º51'N, 60º46'E); CMNFI 2007-0037, , mm standard length, Kerman, Hosseinabad and Gamatabad qanats at Bam (29º06'N, 58º21'E); CMNFI 2007-0038, , mm standard length, Kerman, Mehtiabad qanat (29º06'N, 58º21'E); CMNFI 2007-0039, , mm standard length, Kerman, Tahrud River (ca. 29º23'N, ca. 57º53'E); CMNFI 2007-0040, , mm standard length, Kerman, Qahariz qanat at Jupar (30º04'N, 57º08'E); CMNFI 2007-0041, , mm standard length, Kerman, qanat at Baghin (30º12'N, 56º48'E); CMNFI 2007-0042, , mm standard length, Kerman, qanat at Negar (29º52'N, 56º50'E); CMNFI 2007-0043, , mm standard length, Kerman, qanat at Emamzadeh Sultan (ca. 29º40'N, ca. 56º45'E); CMNFI 2007-0044, , mm standard length, Kerman, Qal'eh-ye Askar stream (ca. 29º28'N, ca. 56º38'E); CMNFI 2007-0045, , mm standard length, Kerman, Kharan River drainage at Baft (29º14'N, 56º38'E); CMNFI 2007-0047, , mm standard length, Kerman, qanat at Hoshun (29º14'N, 56º19'E); CMNFI 2007-0048, , mm standard length, Kerman, qanat at Hasanabad (ca. 28º50'N, ca. 55º50'E); CMNFI 2007-0049, , mm standard length, Hormozgan, upper Kol River basin at Hajjiabad (ca. 28º19'N, ca. 55º55'E); CMNFI 2007-0063, , mm standard length, Fars Mand River tributary outside Jahrom (28º36'N, 53º37'E); CMNFI 2007-0065, , mm standard length, Fars, Barm-e Dalak (ca. 29º35'N, ca. 52º38'E); CMNFI 2007-0066, , mm standard length, Fars, qanat under Sa'adi's Tomb, Shiraz (29º37'N, 52º35'E); CMNFI 2007-0067, , mm standard length, Fars, Sivan River (ca. 30º02'N, ca. 52º57'E); CMNFI 2007-0068, 5, 59.0-89.6 mm standard length, Fars, qanat 4 km south of Abarqu (ca. 31º07'N, ca. 53º14'E); CMNFI 2007-0069, , mm standard length, Yazd, qanat at Zarej (ca. 31º58'N, ca. 54º17'E); CMNFI 2007-0070, , mm standard length, Yazd, qanat at Ardakan, (32º19'N, 53º59'E); CMNFI 2007-0073, , mm standard length, Esfahan, Zayandeh River at Tanderan (32º47'N, 51º02'E); CMNFI 2007-0075, , mm standard length, Hamadan, Malayer River south of Malayer (ca. 34º17'N, ca. 48º47'E); CMNFI 2007-0076, , mm standard length, Markazi, Malekabad qanat (34º05'N, 49º53'E); CMNFI 2007-0083, , mm standard length, Azarbayjan-e Khavari, Qaranqu River basin west of Sar Eskand Khan (ca. 37º25'N, ca. 46º55'E); CMNFI 2007-0084, , mm standard length, Azarbayjan-e Khavari, Talkheh River basin west of Sarab (ca. 37º56'N, ca. 47º19'E); CMNFI 2007-0091, , mm standard length, Azarbayjan-e Khavari, Zilber Chay basin west of Marand (38º30'N, 45º23'E); CMNFI 2007-0100, , mm standard length, Azarbayjan-e Bakhtari, Kalwi Chay near Piranshahr (ca. 36º44'N, ca. 45º10'E); CMNFI 2007-0108, , mm standard length, Kordestan, Qeshlaq River basin north of Sanandaj (ca. 35º33'N, ca. 47º08'E); CMNFI 2007-0109, , mm standard length, Kordestan, Qeshlaq River basin south of Sanandaj (ca. 35º16'N, ca. 47º01'E); CMNFI 2007-0110, , mm standard length, Kordestan, Yuzidar River basin (ca. 35º05'N, ca. 46º56'E); CMNFI 2007-0115, , mm standard length, Kermanshahan, Qareh Su basin north of Kermanshah (ca. 34º34'N, ca. 46º47'E); CMNFI 2007-0116, , mm standard length, Kermanshahan, Gav Masiab River basin west of Sahneh (ca. 34º28'N, ca. 47º36'E); CMNFI 2007-0117, , mm standard length, Kermnashahan, Gav Masiab River basin near Sahneh (ca. 34º24'N, ca. 47º40'E); CMNFI 2007-0117, , mm standard length, Kermnashahan, Gav Masiab River basin near Sahneh (ca. 34º24'N, ca. 47º40'E); CMNFI 2007-0122, , mm standard length, Markazi, Khar River basin south of Takestan (ca. 35º56'N, ca. 49º30'E); USNM 200308, 2, 37.5-47.3 mm standard length, Lorestan, Ab-e Khorramabad (33º30'N, 48º13'E); ? damascina USNM 205933, 5, 97.5-142.4 mm standard length, Baluchestan, Karavandar Creek (no other locality data); ? damscina USNM acc. 303854, 14, 29.1-44.6 mm standard length, Fars, Lake Arzhan (29º36'N, 51º59'E). ? damscina USNM acc. 303854, 4, 55.7-99.0 mm standard length, Fars, pool east of Sangkar (29º48'N, 53º29'E); ?damascina

Capoeta aculeata
(Valenciennes, 1844)

 

Flank scale                                                            Left pharyngeal arch

Common names

shum (= unlucky or inauspicious, possible meaning), سياه ماهي (= siah mahi), زرده پر (= zardehpar), siah mahi aculeata.

Systematics

Chondrostoma aculeatum was originally described from "eaux douces de la Perse".

Scaphiodon macrolepis Heckel, 1849 described from the "Confluenten des Araxes bei Persepolis" (probably the Pulvar (= Sivan) River, Fars near Persepolis) and Varicorhinus bergi Derzhavin, 1929 described in Latin from "Keredsh flumen propea Teherane, Persia septentrionalis" (Karaj River near Tehran, northern Iran) are synonyms.

Six syntypes (MNHN 2357) of Chondrostoma aculeatum in poor condition are stored in the Muséum national d'Histoire naturelle, Paris (Bertin and Estève, 1948; Coad and Krupp, 1994). They measure 86-179 mm standard length (Coad and Krupp, 1994) or 105-210 mm total length (Bertin and Estève, 1948). The largest specimen is designated as the lectotype.

Two syntypes of Scaphiodon macrolepis are in the Naturhistorisches Museum Wien under NMW 55896 and measure 100-175 mm standard length (Kähsbauer, 1964). Two other fish are marked as syntypes under NMW 51653 and are from Persepolis collected by Th. Kotschy. The catalogue in Vienna lists 4 fish and the 1997 card index agrees these 4 fish are the syntypes.

Types of Varicorhinus bergi are unknown (Eschmeyer et al. (1996)).

Berg (1949) considers both aculeata and macrolepis as distinct species although very close, the latter distinguished from the former by a deeper body and a shorter head. Karaman (1969) and Bianco and Banarescu (1982) place both aculeata and macrolepis in Capoeta capoeta; Karaman does suggest that macrolepis could belong in aculeata. Saadati (1977) considers aculeatus not more than subspecifically distinct from macrolepis, not realising the former has priority.

Key characters

This species differs from all others in the genus Capoeta in the lower number of lateral line (93% of 314 fish had range of 39-48) and caudal peduncle scales (90% of 303 fish had a range of 16-20). Capoeta capoeta, a related species, usually has 54 or more lateral line scales and 20 or more caudal peduncle scales.

Morphology

Dorsal fin with 3-5, modally 4, unbranched and 7-9, modally 8, branched rays. The last dorsal fin unbranched ray is thickened and serrated, the denticles being long and narrowly spaced but not strongly developed. Distally this spiny ray is flexible. Smaller fish have proportionately larger and more extensive denticles than larger fish. The extent of denticles from the base distally varies between about two-thirds and three-quarters. Anal fin with 3 unbranched and 5-6, modally 5, branched rays, pectoral fin with 14-21 branched rays, and pelvic fin with 7-10 branched rays.

Lateral line scales 36-52. Caudal peduncle scales 13-23. Scale shape is squarish with shallowly rounded to straight dorsal and ventral margins, sharp corners anteriorly, and a large to moderate central protuberance on the anterior margin. Radii are most numerous on the posterior field but even there are few, relatively few laterally and few anteriorly. Circuli are very fine but break into coarser "bubbles" on the posterior field. The focus is subcentral anterior. The pelvic fin axillary scale varies greatly in size.

The mouth is slightly arched or even straight in ventral view. The horny edge to the lower jaw is usually well-developed but may be lost in preserved specimens. Gill rakers number 16-25 and are short, reaching past the first or second raker when appressed. Rakers are thick and usually hooked at their tips. Pharyngeal teeth are modally 2,3,4-4,3,2 (in 10 fish). Major row teeth are spatulate with a wide crown in large fish. Total vertebrae number 39-44. The gut is extremely elongate with numerous anterior and posterior coils.

Meristic characters in Iranian fish are: dorsal fin branched rays 7(50), 8(255) or 9(4); anal fin branched rays 5(177) or 6(1); pectoral fin branched rays 14(2), 15(3), 16(12), 17(52), 18(123), 19(79), 20(27) or 21(6); pelvic fin branched rays 7(23), 8(183), 9(102) or 10(6); lateral line scales 36(1), 37(5), 38(8), 39(15), 40(25), 41(48), 42(56), 43(38), 44(40), 45(23), 46(22), 47(15), 48(10), 49(1), 50(4), 51(2) or 52(1); scales around the caudal peduncle 13(1), 15(5), 16(48), 17(52), 18(73), 19(64), 20(37), 21(12), 22(5) or 23(6); total gill rakers 16(3), 17(13), 18(40), 19(49), 20(62), 21(43), 22(43), 23(27), 24(16) or 25(6); and total vertebrae 39(1), 40(26), 41(90), 42(103) or 44(16).

Sexual dimorphism

Males have moderately large tubercles on the anal fin rays following the ray branching (2-4 tubercles on last 4 branched anal rays), small tubercles on the lowest caudal fin ray, very fine tubercles on top of the head, larger tubercles on the side of the head, largest on the snout below the eye and nostril as far as the mouth, connecting across the snout, and numbering 1-5 moderately large tubercles on flank scales variously arranged on each scale and best developed on the posterior part of the body.

Colour

The back is almost entirely black to green-brown or olive-green, the upper flank is brownish, the belly and lower flank are yellow up to the lateral line, only the belly centre being white. The flanks are generally silvery in live fish. Some fish have small black spots on the sides and fins. Preserved fish have pigment on the posterior, exposed margin and so are outlined on the flank. The sides of the head are golden-brown. Flank spots may be in 5 longitudinal rows above, and 2 rows below, the lateral line. Some populations have fish with spots and mottles on the body and fins but these are probably occasioned by a parasitic infestation. Fins are often reddish-brown to pink although pelvic and anal fins may be yellowish-green and the dorsal and caudal fins very light to hyaline. Preserved fish have pigment on the rays and membranes of fins without any distinctive pattern. The dorsal and caudal fins are darker than the lower fins. The iris is golden to orange. The peritoneum is black.

Size

Reaches 23.4 cm standard length.

Distribution

This species is found in the Tigris River, Namak Lake, Dasht-e Kavir, Kerman-Na'in, Esfahan, Kor River basins (Rainboth, 1981; Bianco and Banarescu, 1982; Ghorbani Chafi, 2000). Abdoli (2000) maps this species from theKerman-Na'in basin generally; the upper Kal Shur, Jajarm and Jovein rivers in the Dasht-e Kavir basin; the middle and upper Shur and Abhar, Qareh Chai and Qom rivers in the Namak Lake basin; the Zayandeh and Shur rivers in the Esfahan basin; the Jarrahi and Marun, upper Karun and Khersan, Dez, Karkheh, Simarreh and Kashkan rivers in the Tigris River basin.

Zoogeography

Saadati (1977) suggests that this species moved eastward to basins on the plateau during more pluvial periods from the Tigris River basin. See also above under genus.

Habitat

Unknown in detail.

Age and growth

Unknown.

Food

Gut contents include filamentous algae, plant fragments and diatoms with large amounts of sand. This species has been seen turning belly up to feed (field notes for specimens from Jajarm, Khorasan).

Reproduction

Reproduction has not been studied in this species. Specimens from the Khorramabad River contained eggs 1.5 mm in diameter on 6 July and some seemed to be reabsorbing eggs. Spawning presumably takes place in late spring and summer.

Parasites and predators

Barzegar et al. (2004) examined this species for parasites in fish from the Beheshtabad river in Chahar Mahall va Bakhtiari Province and found Dactylogyrus lenkorani, Gyrodactylus sp. and Myxobolus sp. Masoumian et al. (2007) record the myxosporean parasite Myxobolus cristatus from this species in the Zayandeh River. Mehdipoor et al. (2004) record the monogeneans Dactylogyrus chramuli, D. lenkorani and D. gracilis in the Zayandeh River. Barzegar and Jalali (2006) report parasites in this species from Kaftar Lake as Lernaea cyprinacea and Trichodina sp. Barzegar et al. (2008) record the digenean eye parasites Diplostomum spathaceum and Tylodelphys clavata from this fish. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Lernaea cyprinacea on this species.

Economic importance

None.

Conservation

This species is widely distributed in Iran and does not appear to be in need of conservation but its biology and habitat requirements are unknown. Kamali-Far et al.(2009) have used carp pituitary extract in an attempt to induce spawning in this species. Hatchery production could then be used to supplement natural stocks. However, the attempt was unsuccessful. Note that the identity of the species used in this study needs verification judging from the photograph in the paper.

Further work

The biology of this species needs study as does its habitat requirements and conservation needs.

Sources

Type material: See above, Chondrostoma aculeatum (MNHN 2357).

Iranian material: CMNFI 1979-0025, 2, 65.3-68.1 mm standard length, Fars, Kor River near Marv Dasht (29º51'N, 52º46'30"E); CMNFI 1979-0059, 155, 22.9-67.4 mm standard length, Fars, Pulvar River 8 km south of Sivand (30º01'30"N, 52º57'E); CMNFI 1979-0061, 6, 28.6-64.9 mm standard length, Fars, stream tributary to Pulvar River (30º04'N, 53º01'E); CMNFI 1979-0069, 1, 28.7 mm standard length, Fars, qanat at Naqsh-e Rostam (29º59'30"N, 52º54'E); CMNFI 1979-0070, 16, 25.9-60.0 mm standard length, Fars, Pulvar River near Naqsh-e Rostam (29º59'N, 52º54'E); CMNFI 1979-0090, 2, 153.6-160.5 mm standard length, Esfahan, Gav Khuni (ca. 32º21'N, ca. 52º49'E); CMNFI 1979-0116, 49, 24.3-52.1 mm standard length, Fars, Kor River near Marv Dasht (29º51'N, 52º46'30"E); CMNFI 1979-0117, 14, 34.4-44.1 mm standard length, Fars, Pulvar River at Naqsh-e Rostam (29º59'N, 52º54'E); CMNFI 1979-0252, 3, ?, mm standard length, Markazi, jube at Baqerabad (34º55'N, 50º50'E); CMNFI 1979-0253, 5, 40.4-103.7 mm standard length, Markazi, stream in Qareh Chay drainage (34º52'N, 50º49'E); CMNFI 1979-0270, 1, 121.8 mm standard length, Lorestan, Kashkan River draiangae outside Khorramabad (33º26'N, 48º19'E); CMNFI 1979-0271, 1, 52.1 mm standard length, Lorestan, stream in Kashkan River drainage (33º39'N, 48º32'30"E); CMNFI 1979-0273, 28, 51.4-104.5 mm standard length, Lorestan, stream in Kashkan River drainage near Khorramabad (33º26'N, 48º19'E); CMNFI 1979-0274, 6, 20.6-59.2 mm standard length, Lorestan, stream in Kashkan River drainage (33º27'N, 48º11'E); CMNFI 1979-0275, 1, 50.9 mm standard length, Lorestan, Kashkan River 2 km from Ma'mulan (33º25'N, 47º58'E); CMNFI 1979-0279, 18, 41.1-129.9 mm standard length, Lorestan, Khorramabad River (33º37'N, 48º18'E); CMNFI 1979-0282, 7, 99.2-130.8 mm standard length, Lorestan, river at Nurabad (34º05'N, 47º58'E); CMNFI 1979-0283, 2, 125.2-186.3 mm standard length, Kermanshahan, Qareh Su near Kermanshah (34º21'N, 47º07'E); CMNFI 1979-0343, 1, 146.6 mm standard length, Fars, lake near Deh Bid (ca. 30º32'N, ca. 52º49'E); check ID? CMNFI 1979-0365, 1, 25.0 mm standard length, Khuzestan, stream in Doveyrich River drainage (32º25'N, 47º36'30"E); CMNFI 1979-0396, 9, 32.5-58.7 mm standard length, Khuzestan, Kheyrabad River 20 km from Behbehan (30º32'N, 50º23'30"E); CMNFI 1979-0427, 2, 100.5-112.2 mm standard length, Markazi, Cheshmeh Fin at Fin (33º57'N, 51º24'E); checkID? CMNFI 1979-0428, 17, 25.9-104.5 mm standard length, Markazi, stream 3 km south of Sen Sen (34º13'N, 51º16'E); checkID? CMNFI 1979-0458, 9, 48.5-117.8 mm standard length, Markazi, Khar River 6 km north of Ab-garm (35º47'N, 49º20'E); CMNFI 1979-0460, 1, 77.6 mm standard length, Hamadan, stream 16 km south of Asadabad (34º38'N, 48º03'E); checkID? CMNFI 1979-0463, 8, 97.9-135.3 mm standard length, Markazi, Qareh Chay (34º53'N, 50º24'E); checkID? CMNFI 1979-0464, 1, 74.2 mm standard length, Markazi, qanat at Kheyrabad (34º08'N, 50º00'E); CMNFI 1979-0465, 18, 35.7-58.3 mm standard length, Markazi, Qom River (34º18'30"N, 50º32'E); CMNFI 1979-0500, 2, 92.4-98.6 mm standard length, Fars, Pulvar River near Naqsh-e Rostam (29º59'N, 52º54'E); checkID? CMNFI 1980-0156, 27, ? mm standard length, Markazi, Karaj River (35º47'N, 50º58'E); CMNFI 1993-0154, 1, mm standard length, Markazi, Sharra River near Far (34º03'N, 49º20'E); checkID? CMNFI 1993-0156, 1, mm standard length, Markazi, Sharra River (34º03'N, 49º21'E); checkID? CMNFI 2007-0006, 9, 59.9-127.2 mm standard length, Khorasan, spring in Qareh Su basin south of Garmeh (ca. 36º58'N, ca. 56º15'E); CMNFI 2007-0007, 8, 59.4-79.3 mm standard length, Khorasan, stream supplemented by qanats, Kal-e Tangeh (ca. 36º59'N, ca. 56º29'E); CMNFI 2007-0008, 2, 72.1-84.3 mm standard length, Khorasan, qanat at Jajarm (36º57'N, 56º23'E); CMNFI 2007-0009, 18, 35.9-108.1 mm standard length, Khorasan, qanat at Amirabad (ca. 36º31'N, ca. 56º45'E); CMNFI 2007-0010, 11, 80.8-123.1 mm standard length, Khorasan, qanat at Haresabad (36º07'N, 57º37'E); CMNFI 2007-0011, 12, 34.1-85.4 mm standard length, Khorasan, Kalshur River south of Neyshabur (36º05'N, 58º43'E); CMNFI 2007-0071, 10, 70.4-156.9 mm standard length, Esfahan, qanat at Mohammadiyeh, Na'in (32º51'N, 53º06'E); CMNFI 2007-0074, 29, 50.6-100.7 mm standard length, Markazi, Qareh Chai west of Arak (34º03'N, 49º21'E); CMNFI 2007-0075, 16, 29.3-152.5 mm standard length, Hamadan, Hamadan, Malayer River 5 km from Malayer (ca. 34º17'N, ca. 48º47'E); CMNFI 2007-0076, 5, 56.1-97.4 mm standard length, Markazi, Malekabad qanat east of Arak (34º05'N, 49º53'E); CMNFI 2007-0078, 8, 37.6-102.8 mm standard length, Markazi, Qom River (ca. 34º18'N, ca. 50º32'E); CMNFI 2007-0117, ?, mm standard length, Kermanshahan, Gav Masiab River basin near Sahneh (ca. 34º24'N, ca. 47º40'E); CMNFI 2007-0119, ?, mm standard length, Kermanshahan, Gav Masiab River basin near Kangavar (ca. 34º31'N, ca. 48º03'E); CMNFI 2007-0120, 15, 29.0-165.5 mm standard length, Hamadan, Ab Chay near Hamadan (ca. 34º49'N, ca. 48º29'E); CMNFI 2007-0122, 12, 35.0-77.6 mm standard length, Markazi, Khar River basin south of Takestan (ca. 35º56'N, ca. 49º30'E); BM(NH) 1934.10.29:2, 1, 84.0 mm standard length, Markazi, Tehran (no other locality data); BM(NH) 1958.11.7:1-6, 6, 25.6-89.9, Khorasan, Jajarm (36º57'N, 56º23'E); BM(NH) 1975.1.17:255-258, 4, 103.0-161.0 mm standard length, Esfahan, Esfahan (no other locality data); MNHN 1960-611, 2, 127.0-144.0 mm standard length, Markazi, Jajrud east of Tehran (ca. 35º45'N, ca. 51º42'E) USNM 205932, 3, 78.5-159.4 mm standard length, Markazi, stream southwest of Tehran (35º34'N, 51º03'E); ZMH 5905, 2, 57.0-70.0 mm standard length, ?, Jafar Abad qanat (?); ZSM 25703, 1, 76.3 mm standard length, ?, Khorramabad River (no other locality data).

Capoeta barroisi
Lortet, 1894

Common names

siah mahi-ye Dasht-e Arzhani (Arzhan Plain black fish).

[tela barroisi in Arabic; spotted barb, Tigris barb].

Systematics

Subspecies are Capoeta barroisi persica Karaman, 1969 described from "See Zariwar, Mariwan, 120 km westlich v. Sannadaj" (Lake Zaribar near Marivan, Kordestan in the Tigris River basin) and Capoeta barroisi mandica Bianco and Banarescu, 1982 from the "Mand River near Dasht-e-Arzhan" of Fars Province. Krupp (1985c) considers both these to be synonyms of the nominal subspecies, C. b. barroisi.

The subspecies persica is distinguished from the type subspecies by having a more horseshoe-shaped mouth, 8 branched dorsal fin rays, 18 gill rakers, blackish pectoral, pelvic and anal fins, few but very large black spots on the body, a shorter anal fin and a longer pectoral fin, and a deep body, based on a single specimen. Krupp (1985c) considers the characters of mouth form and colour to fall within the range of the nominal subspecies (and by implication the other characters too). Özuluğ and Freyhof (2008) found it difficult to reach a conclusion on the taxonomic status of this subspecies on the basis of a single specimen which could be abnormal.

C. barroisi mandica differs from the type subspecies (C. barroisi barroisi) and C. b. persica in number of scales (61-68 in mandica (58-68 in types examined by me), 69-82 in barroisi, 78-79 in persica), number of gill rakers (21-24 in mandica (22-27 in types examined by me and apparently number is related to size of fish), 27-31 in barroisi, 18 in persica), from barroisi in having usually 8 branched dorsal fin rays (barroisi has 9 but persica also has 8), and from persica by a straight mouth (also straight or transverse in barroisi, arched in persica). Krupp (1985c) considers the scale counts to be within the lower range of the nominal subspecies, gill raker counts and mouth position do not differ from the nominal subspecies, and the dorsal fin ray count of 8 is seen in the subspecies mandica. Krupp observes that meristic and morphometric characters are extremely variable in widely distributed Capoeta species. Özuluğ and Freyhof (2008) examined 5 juvenile specimens from the Mand River and consider the subspecies mandica to be a valid species. Widespread taxa like Capoeta species are prime candidates for  molecular analyses which might help resolve conflicting views on - a single widespread, variable species versus several distinct species.

Berg (1949) considers this species to be close to C. damascina, differing by having a stronger spine in the dorsal fin, hardly an invariant character. Saadati (1977) considers that C. barroisi of Karaman is in fact C. damascina. Turan (2008) using mt 16S rDNA concluded on this evidence that C. barroisi was a subspecies of C. damascina although other genetic markers should be used for a more reliable assessment.

Syntypes of Capoeta barroisi are in the Musée Guimet d'Histoire Naturelle, Lyon (MGHN 3492, 316 mm standard length, from the Orontes near Antakya in Turkey collected by E. Chantre and MGHN 3493, 278 mm standard length, from Buhairat Hims in Syria collected by Th. Barrois) (Krupp, 1985c).

The holotype of C. barroisi persica is in the Zoologischen Instituts und Zoologischen Museums der Universität Hamburg (ZMH H4119, 185.2 mm standard length, Daryacheh-ye Zaribar, 35°32'N, 46°08'E, IV. 1968, W. Nümann (Bianco and Banarescu, 1982; Krupp, 1985c; examined and measured by me).

The holotype of C. b. mandica, 106.9 mm standard length, is in the Istituto di Zoologia dell'Universitá di L'Aquila, Italy (IZA 7890), with 95 paratypes from the same locality in IZA 7891 (now numbering 84 fish measuring 34.2-84.9 mm standard length) and 5 paratypes in the Institutul de Stiinte Biologice, Bucuresti, Romania (ISBB 3123), these 100 specimens having a standard length of 34-86 mm. Six paratypes of mandica are in the Canadian Museum of Nature, Ottawa under CMNFI 1982-0366 (from IZA 7891).?lengths

Key characters

The dorsal fin branched ray count of usually 9 rays is characteristic for the type subspecies but not nominal Iranian ones. Gill raker counts, a head length greater than the dorsal fin spine length and the irregular brownish markings on a silvery-white body are also distinctive.

Morphology

Dorsal fin with 3-4 unbranched and 8-10 branched rays, usually 9 in the type subspecies but 8 in the subspecies mandica, anal fin with 3 unbranched and 5 branched rays. Pectoral fin branched rays 15-16, pelvic fin rays 7-8. Lateral line scales 61-82, 58-68 in the subspecies mandica. Scales on the belly in front of the pelvic fins are small and may not be imbricate. Pharyngeal teeth 2,3,4-4,3,2. Gill rakers 18-31, reaching the second adjacent raker when appressed. The last dorsal fin unbranched ray is very strong, but narrows distally, and bears large denticles or serrations on three-quarters of its length. The snout in the holotype of the mandica subspecies has a depression in front of the nostrils.

ZMH6086 D8, A5 P15, V7, ll 58, gr 24, flanks speckled dorsally, mouth a gentle arch almost straight, large denticles in damaged D spine.

Sexual dimorphism

Tubercles in males are found from eye to eye around the snout with fine tubercles sparse on the top of the head. Most flank scales have a single, centrally-placed tubercle as do scales on the caudal peduncle. There is a single row of tubercles on the last three anal fin rays. There are some weak tubercles on the side of the head.

Colour

There are numerous, small, distinctive brown to black spots on the head, flank and dorsal and anal fins. The back and upper flank are silvery-white with slate to violet overtones and the belly is yellowish. The holotype of the subspecies mandica has light specking and mottling on the upper flank and back. All fins have some speckling on the rays and membranes but no clear rows of spots. Smaller fish (paratypes) have darker and bigger speckles than the holotype which extend lower on the flank.

Size

Reaches 31.6 cm standard length.

Distribution

The subspecies C. barroisi barroisi is found in the Ceyhan, Orontes, Quwayq and Tigris-Euphrates basins, C. b. persica only in Lake Marivan or Zaribar of the Tigris River basin of Iran, and C. b. mandica in the Mand River of the Gulf basin. Krupp (1985c) includes Iranian Gulf drainages for the type subspecies. Abdoli (2000) has Jarrahi and lower Karun in the Tigris River basin and the middle and lower Hilleh, lower Mand, and lower Dasht-e Palang rivers in the Gulf basin.

Zoogeography

Taxa in Iran are remote from the type locality of this species and may be indicative of the links between the Levantine fauna and the former tributaries of the Tigris-Euphrates basin in southern Iran. See also above under genus.

Habitat

Unknown.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

None reported from Iran.

Economic importance

None.

Conservation

Nothing is known of biology and a conservation assessment cannot be made. Endangered in Turkey (Fricke et al., 2007).

Further work

The biology of this species needs study as a prerequisite for a conservation assessment. The relationships of nominal Iranian taxa to material from the type locality need further study using molecular techniques.

Sources

Type material: Capoeta barroisi mandica (IZA 7890, 7891, CMNFI 1982-0366 (from IZA 7891)) and C. barroisi persica (ZMH H4119).

Iranian material: ZMH 6086, 1, 73.6 mm standard length, Fars, Shur Fluβ, zufluβ von Mand-Fluβ (= C. b. mandica) P. Bianco ?check this against his paper.

Comparative material: BMNH 1974.2.22:1853-1856, 3, 48.9-60.2 mm standard length, Iraq, Kaliasan near Sulaymaniyah

Capoeta buhsei
Kessler, 1877

Common names

shamshiri (= sword-like), mahi sibili (= moustached fish, from Karaj Lake).

Systematics

Varicorhinus nikolskii Derzhavin, 1929 described in Latin from the "Keredsh flumen" (= Karaj River near Tehran) is a synonym. Saadati (1977) places Capoeta buhsei in Capoeta damascina.

The 2 syntypes of Capoeta Buhsei, 200.7-211.4 mm standard length, are in the Zoological Institute, St. Petersburg (ZISP 2330) and were collected "iz Persii" (= from Persia) by Dr. Buhse in 1849. The 11 syntypes of Varicorhinus nikolskii have not been located (Eschmeyer et al., 1996).

Key characters

This species is distinguished by its low total gill raker count of 9-17, mean 12.4, modes 12 and 13, lower arch rakers 7-10 (cf. C. damascina which has 17-25 total rakers, mean based on ? specimens), the absence of a keel in front of the dorsal fin, the mouth structure, and by a very weak, unserrated or barely serrated dorsal fin spine in large fish (cf. C. damascina).

Morphology

Dorsal fin unbranched rays 3-4, branched rays 7-9, anal fin unbranched rays 3, branched rays 5., pectoral fin branched rays 14-19, and pelvic fin branched rays 7-9. Lateral line scales 72-99. Scales are found on the back and on the belly. A pelvic axillary scale is present. Scales have parallel dorsal and ventral margins, a rounded posterior margin and an anterior margin with a rounded central protuberance. Radii are found on all fields including a few long and curved ones on the lateral fields. The focus is subcentral anterior and circuli are numerous and fine. Gill rakers 9-19 in literature (but see below), including some counts probably for the lower arm only, and reach the second raker below when appressed but only the next raker in small fish. Pharyngeal teeth in the main row are spatulate, the crowns flat, narrow and curved. Tooth counts are 2,3,4 or 5-5 or 4,3,2. The fifth tooth in either row is small and variably present. This may be size related although the fish examined here were all relatively small and showed no clear trend. The gut is elongate with several long coils.

The mouth is large and a shallow horseshoe-shape with the horny lower jaw layer weakly developed but the lower lip corners in particular fleshy and well-developed. The last unbranched dorsal fin ray is weak with the distal half thin and flexible. Denticles are found on the basal third to two-thirds or more of the ray, their extent and size variable but usually small, weak and less extensive in large fish, while larger and more extensive in the smallest fish. For fish 48.9-174.0 mm standard length extent of dorsal fin spine serrations in spine length is 0.3-0.8, mean 0.6. The lower lip is apparent and finely ridged. The upper lip and snout are covered with unculi which occur also over the head but more widely spaced out. The upper lip unculi are densely concentrated and are broader than other head ones. Unculi are also on the lower head surface and belly scales back to the pelvic fins, and on the anal, pectoral and pelvic fin rays and membranes.

Meristics are as follows: dorsal fin branched rays 8(35) or 9(3); anal fin branched rays 5(38); pectoral fin branched rays 14(1), 17(20), 18(11) or 19(6); pelvic fin branched rays 7(1), 8(5), 9(31); lateral line scales 72(2), 73(1), 75(3), 76(4), 77(2), 78(1), 79(7), 80(4), 81(5), 82(2), 83(2), 84(2), 86(2) or 91(1); total gill rakers 9(2), 10(2), 11(4), 12(12), 13(11), 14(4), 15(2) or 17(1); pharyngeal teeth 2,3,4,-4,3,2(7), 2,3,4-5,3,2(6), 2,3,5-4,3,2(5), 2,3,5-5,3,2(1), 1,3,4-4,3,2 (1) or 2,3,4-4,3,1(1); and total vertebrae 43(1) and 44(6) (USNM 20593 and the syntypes).

Sexual dimorphism

One male specimen measuring 94.6 mm standard length bears large tubercles on anal fin rays, fine tubercles scattered on the head, on the back and upper flanks one tubercle per scale at the scale centre but not on every scale, all along the lateral line at one tubercle per scale, and below the lateral line only in the area above the anal fin.

Colour

Overall colour is brownish in preservative without spots or any distinctive markings. The back is dark. The peritoneum is dark brown to black in preserved fish.

Size

Reaches 25.7 cm.

Distribution

This species is endemic to the Namak Lake basin of Iran (Derzhavin, 1929; Wossughi, 1978; Abdoli, 2000). Abdoli (2000) questionably maps it from the Esfahan basin. A report from Lake Zaribar, Kordestan (Abzeeyan, 5(5):III, 1994) is presumably a mis-identification and records from springs of Kul River basin near Darab in the Hormuz basin (Bianco and Banarescu (1982) and the Hamun-e Jaz Murian basin (Vossoughi, 1998) are also questionable.

Zoogeography

An endemic of an interior Iranian basin, its zoogeographical relationships to other Capoeta have not been resolved. See also above under genus.

Habitat

Unknown.

Age and growth

Unknown.

Food

Gut contents include aquatic insect larvae and masses of filamentous algae, suggesting that aufwuchs is an important diet item.

Reproduction

Generally unknown but fish caught on 5 June measuring 121.3-132.6 mm standard length have small eggs, perhaps because this size of fish is not mature. A 174.0 mm standard length caught in January has larger eggs than those from the June fish. A male fish caught on 5 May and measuring 146.6 mm standard length has mature testes.

Parasites and predators

Williams et al. (1980) report the helminths Khawia armeniaca (a cestode) and Acanthocephalorhynchoides cholodkowskyi (an acanthocephalan) from this species in the Zayandeh River at Esfahan.

Economic importance

Unknown.

Conservation

The conservation status of this species has not been determined by field studies and assessments can only be done from museum collections.

Further work

The distribution and numbers of this species in the Namak Lake basin should be examined by field studies to determine the population status. This basin is mostly in Markazi (= Central) Province which contains Tehran and a very large human population with great demands on limited water resources. It is probably not under any immediate threat but is an Iranian endemic.

Sources

Type material: See above, Capoeta buhsei (ZISP 2330).

Iranian material: CMNFI 1970-0588, 19, 42.4-128.9 mm standard length, Markazi, Karaj Lake (35º57'N, 51º06'E); CMNFI 1979-0094, 2, 143.1-174.0 mm standard length, Markazi, Karaj Lake (35º57'N, 51º06'E); CMNFI 1979-0266, 2, 52.4-54.3 mm standard length, Esfahan, spring at Nowqan (ca. 33º10'N, ca. 50º05'E); CMNFI 1979-0458, 1, 94.2 mm standard length, Markazi, Khar River (35º47'N, 49º20'E); CMNFI 1979-0459, 2 ?check fish, only 1 in catalogue, 27.0-31.6 mm standard length, Hamadan, stream 2 km south of Razan (35º22'N, 49º02'E); CMNFI 1979-0461, 1, 54.1 mm standard length, Hamadan, qanat at Taveh (35º07'N, 49º02'E); CMNFI 1979-0495, 1, 42.5 mm standard length, Markzai, Nam River west of Firuzkuh (35º43'N, 52º40'E); CMNFI 1980-0154, 71, 12.0-34.9 mm standard length, Markazi, Karaj River below village (35º47'N, 50º58'E); CMNFI 1980-0156, 27, 32.4-54.3 mm standard length, Markazi, Karaj River below village (35º47'N, 50º58'E); CMNFI 1993-0151, 1, 146.4 mm standard length, Markazi, Sharra River near Far (34º03'N, 49º19'E); CMNFI 1993-0152, 2, 121.3-132.6 mm standard length, Markazi, Sharra River near Khosbijan (34º07'N, 49º23'E); CMNFI 1993-0153, 2, 104.3-138.9 mm standard length, Markazi, Sharra River near Emarat (33º52'N, 49º36'E); CMNFI 1993-0154, 1, 124.0 mm standard length, Markazi, Sharra River near Far (34º03'N, 49º20'E); CMNFI 2007-0074, 3, ? mm standard length, Markazi, Qareh Chay (34º03'N, 49º21'E); CMNFI 2007-0078, 5, ? mm standard length, Markazi, Qom River (ca. 34º18'N, ca. 50º32'E): check ID? CMNFI 2007-0079, 14, ? mm standard length, Zanjan, Abhar River basin (ca. 36º16'N, ca. 49º08'E); CMNFI 2007-0120, , mm standard length, Hamadan, Ab Chay (ca. 34º49'N, ca. 48º29'E); CMNFI 2007-0121, 3, 82.5-141.5 mm standard length, Hamadan, Qareh Su basin north of Razan (ca. 35º25'N, ca. 49º02'E); CMNFI 2007-0122, , mm standard length, Markazi, Khar River basin south of Takestan (ca. 35º56'N, ca. 49º30'E); USNM 20593, ?, ? mm standard length, (); ZMH 2632, 1, 148.2 mm standard length, Dojodje (); ZMH 2633, ?, ? mm standard length, above Latian ().

Capoeta capoeta
(Güldenstaedt, 1773)

Capoeta capoeta heratensis from Wikimedia Commons.

Common names

tilkhos, سياه ماهي (= siah mahi, meaning black fish), sang lisak (= rock snail?); soru (= slippery) in the Dalaki and Shapur river basins.

[gara balig or Lankaran xramulyasi for C. c. gracilis, Kur xramulyasi for C. capoeta, both in Azerbaijan; khramulya, capoeta, kapuit, kaput (all apparently derived from local names in Georgia and Armenia, namely khramuli and kapweti); Lenkoranskaya khramulya or Lenkoran khramulya, Kurinskaya khramulya or Kura khramulya, Zakaspiiskaya khramulya or Transcaspian khramulya (also marinka is used locally for the Transcaspian khramulya subspecies but this is an error), Araksinskaya khramulya or Araks khramulya, all in Russian; Transcaucasian barb; khramulia; kersin handscherli at Aleppo, in Arabic].

Systematics

Cyprinus capoeta was originally described from Tbilisi, Georgia.

Cyprinus fundulus Güldenstaedt, 1787 from the Caspian Sea, Cyrus River (and Capoeta fundulus Valenciennes, 1842), Scaphiodon asmussii Keyserling, 1861 from "Warme Quelle bei Sultan Karaul, 8 Meilen nordöstlich von Herat" (now in Afghanistan, formerly in Persia), Scaphiodon gracilis Keyserling, 1861 from "Wasserleitung bei Gaes, einige Meilen von Isphahan", Scaphiodon heratensis Keyserling, 1861 described from the "Heri-Rud, ein Fluss bei Herat" (now in Afghanistan, formerly in Persia), Capoeta Hohenackeri Kessler, 1877 from Caucasia (probably lower Kura and Araks rivers, Azerbaijan), Capoeta (Scaphiodon) Steindachneri Kessler, 1872 and Capoeta Steindachneri var. platylepida Kessler, 1872 both from the Zeravshan River, Uzbekistan, and probably Capoeta gibbosa Nikol'skii, 1897 described in Latin as from "Bochsani in Persia orientale" are synonyms. Capöeta Guldenstädtii De Filippi in Tortonese, 1940 from "F. Arasse, Erzerum (Anatolia)" is Capoeta capoeta but it is a manuscript name and is not available (Tortonese, 1940; Eschmeyer et al., 1966); 2 syntypes are in the Istituto e Museo di Zoologia della R. Università di Torino (MZUT N.729).

Eschmeyer et al. (1996) have the date of Cyprinus capoeta as 1772; the type locality is Tiflis, Caspian Sea: they also have Cyprinus fundulus authored by Pallas, 1814 although Berg (1948-1949) has Güldenstädt as the author.

A hybrid of Capoeta capoeta heratensis and Schizothorax pelzami is reported from the northern Kopetdag in Turkmenistan (Starostin, 1936).

Capoeta hohenackeri Kessler, 1877 described from tributaries of the Kura and Aras rivers has a high lateral line scale count in the original description (78) and might be a mislabelled Capoeta tinca (Heckel, 1843) from Black Sea drainages of Georgia and Turkey rather than the Caspian Sea basin.

Capoeta capoeta gracilis is the subspecies of much of Iran and Capoeta capoeta heratensis (figure above) is the subspecies from the Tedzhen River basin (Berg, 1949). The former usually has one pair of barbels, the latter two pairs (but see below). Bianco and Banarescu (1982) limit C. c. gracilis to basins between the Safid River and the Atrak while C. c. capoeta is found in the Kura-Aras basin. Holčík and Jedlička (1994) consider that the two subspecies gracilis and heratensis do not exist but that the taxon C. capoeta exhibits clinal variation.

Bănărescu in Bănărescu (1999) limits C. capoeta gracilis to the Lake Orumiyeh basin and the Safid River in Iran (and the lower Kura River of Azerbaijan) while his C. capoeta aff. gracilis (an unnamed subspecies related to C. capoeta gracilis) is found along the rest of the Iranian Caspian shore. However his material was limited (and did not include any from Esfahan, the type locality of gracilis) and the analysis is based on lateral line scale counts only. Bănărescu in Bănărescu (1999) also states that C. capoeta sevangi de Filippi, 1865 is the subspecies of the Araxes River basin, presumably including Iran, distinguished from the type subspecies, C. capoeta capoeta of the Kura River basin, by having the dorsal fin margin straight or slightly convex as opposed to slightly to moderately notched. This character does not seem to be significant for such wide ranging and variable populations, which he admits in one case at least (Kura River at Mingechaur), show differences between samples from the same locality at different times.

Abdurakhmanov (1962) compares fish from the Kura River basin (presumably C. c. capoeta) with fish from the Lenkoranchai and Bilyashchai in Azerbaijan (C. c. gracilis) and finds that the latter have fewer dorsal fin rays on average, greater head length and depth, smaller eye, longer snout and postorbital distance, greater body depth and caudal peduncle depth, a shorter postdorsal distance, a shorter dorsal fin base, lesser dorsal fin height, a longer anal fin base, a greater pectoral-pelvic distance and a shorter pelvic-anal fin distance.

Dadikyan (1986) refers to Varicorhinus capoeta araxensis subsp. nov. from the Aras River basin in Armenia.

Günther (1899) points out that the considerable morphological variation shown by these fishes has resulted in numerous specific names and that it is difficult to assess them without a large comparative series and better information on localities. Berg (1948-1949) also indicates that the various subspecies are very close to each other and that their distributions are not clearly isolated.

C. c. heratensis shows major variations in body form, sometimes called morpha elata with a deep body and morpha elongata with a shallow and elongate body. These are not taxonomically significant but simply ecomorphs and all intermediates between the two extremes can be found. The deep-bodied form probably formed part of the fishes described as asmussii (Berg, 1948-1949). Reshetnikov and Shakirova (1993) list Capoeta heratensis as a full species.

Samaee et al. (2006) showed differences in morphometry between fish from six rivers along the Iranian Caspian shore with an overall assignment of individuals to a group of 88.6%. The morphometric data were mirrored by molecular data. Differences in morphometry were attributed to environmental and habitat conditions (temperature, turbidity, food availability and water depth and flow) but molecular data indicated a genetic basis, presumably through lack of gene flow between the river populations. Samaee et al. (2009) examined morphological variation with this species in the Shirud of the south Caspian Sea basin. There were no significant differences in meristic characters but morphometric characters varied and could be used to distinguish 5 groups. AnvariFar et al. (2011) compared fish from above and below the Shahid Rajaei Dam (built in 1995) and found the two populations to be morphologically different.

Records of Capoeta capoeta from the Tigris River basin at least are probably Capoeta damascina with low scale counts (F. Krupp, in litt., 1986).

Wossughi (1978) described, in a dissertation, a subspecies from the Namak Lake basin (from "Tschmeh Jafar Abad bei Araq") but this work may not be published in the sense of the International Code of Zoological Nomenclature (Ride et al., 1985). In any case, the holotype is Capoeta aculeata and the other material comprises 21 Leuciscus (= Squalius) cephalus orientalis and 4 Capoeta aculeata (F. Krupp, in litt., 1984). The type material, all female, is stored in the Zoologischen Instituts und Zoologischen Museums der Universität Hamburg (holotype, 132 mm standard length under ZMH 5946, and 2 paratypes, 115-121 mm standard length, under ZMH 5947).

Bianco and Banarescu (1982) described C. capoeta intermedia from the Mand River in Fars but this is referred to C. damascina here (q.v.).

The types of Capoeta gibbosa are in the Zoological Institute, St. Petersburg (ZISP 11104) but have dried at some point. Their locality is given by Nikol'skii (1897) in Latin as "Bochsani in Persia orientali. 4.VII.96 (2)". This may be Bozjani at 35°48'N, 59°36'E. Berg (1949) considers that this nominal species is close to C. capoeta gracilis but is distinguished by body proportions (longer caudal peduncle and a longer head) but it is founded on only 2 specimens, hardly an adequate sample.

Types of Scaphiodon asmussii, S. gracilis and S. heratensis were not kept ? phrasing

Key characters

Berg (1948-1949) and Abdurakhmanov (1962) separate C. c. capoeta from C. capoeta gracilis, both of which may occur in Iran, by the following key:-

1(2). Dorsal fin emarginate above; lateral line scales usually 55-59; dorsal fin spine strong with numerous denticles; back behind occiput and particularly in front of the dorsal fin strongly compressed.....C. c. capoeta

2(1). Dorsal fin truncated in adults; lateral line scales usually 47-58; dorsal fin weak; back behind occiput not or only weakly compressed; radii on scales with minute recesses .....C. c. gracilis

Morphology

Dorsal fin unbranched rays 3-5, branched rays 7-10, usually 8-9; anal fin unbranched rays 2-4, branched rays 5; pectoral fin branched rays 15-20; and pelvic branched rays 7-9. Lateral line scales 46-70. In the subspecies gracilis the scales are said to be somewhat larger than in the type form, 47-58, mostly 48-50, and in heratensis the range is given as 50-60 by Berg (1948-1949). Scales are regularly arranged over the body. There is a pelvic axillary scale. Scales have a wavy anterior edge, few anterior and posterior radii and an almost central focus. Gill rakers 16-30, lower counts may refer to lower arm rakers only and total counts in the range 25-30 are probably more typical. Vertebrae 42-47. Pharyngeal teeth 2,3,4-4,3,2 or 2,3,5-4,3,2 with a hooked tip, spatulate below on posterior teeth while anterior teeth are conical. The last unbranched dorsal fin ray is strong with denticles along one half to two-thirds of its length (less strong in gracilis than in the type form). The number of barbels is variable - in fish from Uzbekistan 2 barbels (126 fish or 58.6%), 3 barbels (15 or 7.0%) or 4 barbels (74 or 34.4%)(Amanov, 1970). The subspecies heratensis is characterised by having 4 barbels but this is probably variable in Iran as in Uzbekistan. Six specimens from the Hari River basin of Iran all had 4 barbels.

Levin et al. (2005) found gracilis and heratensis (and steindachneri) to be oligovertebrate with 41-45 vertebrae, modes 42 to 44, compared to the multivertebrate type subspecies capoeta and sevangi with 45-48 vertebrae, mode 46. Morphometry and longevity also differ between these two groups and it was assumed they belong to different phyletic lines.

The Lake Sevan, Armenia subspecies (sevangi) has 2n=150 and is closer to "Barbus" than to African Varicorhinus, a genus in which Southwest Asian Capoeta were once placed (Krysanov, 1999). C. capoeta from the Safid River, the Shahrud in Rudbar and the Madarso River in Golestan National Park also have 2n=150, NF=230-234 (Pourali et al., 2000; Pourali Darestani et al., 2006). C. c. umbla from the Tigris River of Turkey had 2n=150, possibly hexaploid, with 43 meta-submetacentric chromosomes, 32 pairs of subtelo-acrocentric chromosomes with NF=236 (Kiliç Demirok and Ünlü, 2001).

Günther (1899) points out that this species shows considerable morphological variation, even in fish caught at the same place and time. The mouth can vary from straight to a gentle crescent to a distinct crescent, e.g. in three fish from the Nazlu Chai. There are also variations in dorsal fin spine development and the crown of the head can be flattened or convex.

Meristics for Iranian material: ?

Sexual dimorphism

The snout in males has 2-4 rows of tubercles and tubercles are present on scales and the rays of the anal fin. Abdurakhamanov (1962) reports that caudal peduncle length and lower caudal lobe length are longer in males while anal fin height, pelvic-anal fin distance, postorbital distance and interorbital width are greater in females.

Colour

The back is dark grey or green to brownish and the flanks light, silvery or silvery-grey, or yellowish. There may be several large black spots or blotches on the flank. The belly and lower head surface are pearly-white to dirty yellow. Scales are darkly pigmented. The operculum has a broad, yellow-gold spot. The iris is silvery, somewhat darker or yellow-golden above, or golden overall with traces of grey. The front of the dorsal fin and the margin of the caudal fin are black, and the rest of these fins are grey or yellowish-grey with some pink. The black margin to the caudal fin may be best developed on the upper and lower lobes compared to the posterior margin. The pectoral, pelvic and anal fins are grey with some pink or may be an overall pale pinkish. The peritoneum is black.

Size

Reaches 38 cm standard length and 3.5 kg (Amanov, 1970). This species reaches 43 cm in the Aras River basin of Iran (possibly C. capoeta sevangi, see above; A. Abdoli, pers. comm., 1995), 55 cm in Lake Sevan as C. capoeta sevangi (Bănărescu in Bănărescu, 1999) and 43.5 cm fork length and 1.23 kg in Çıır Lake, Turkey as C. capoeta capoeta.

Distribution

The type species is found in the Kura River basin of Azerbaijan with some Aras River basin fishes very similar. It is not known if the fishes from the Aras River basin in Iran belong to the type species or to gracilis. The subspecies Capoeta capoeta gracilis (known to Russian ichthyologists as the Lenkoran khramulya) was described from near Esfahan and is recognised as the one found over much of Iran including the Caspian Sea basin from the Astara to the Atrak including the Atrak, Gorgan, Gharasu, Tajan, Babol, Haraz, Sardab, Aras, Tonekabon, Pol-e Rud and Safid rivers, the Anzali Talab and Gorgan Bay (Derzhavin, 1934; Bianco and Banarescu, 1982; Holčík and Oláh, 1992; Kiabi et al., 1994; Roshan Tabari, 1997; Shamsi et al., 1997; Abbasi et al., 1999; Kiabi et al., 1999; Abdoli, 2000; Abdoli and Naderi, 2009), the Lake Orumiyeh basin including the Urmi River or Shaher Chai, Nazlu Chai and Talkheh, Tatavi and Zarrineh rivers (Günther, 1899; Abdoli, 2000) and Mahabad Dam (Abdi, 1999; www.mondialvet99.com, downloaded 31 May 2000), Marmisho Lake, Azerbaijan (Shiri et al., 2009), the Dasht-e Kavir basin (Saadati, 1977), the Gulf basin in the Zohreh, Shapur, Dalaki, Helleh, Shur rivers and the upper and lower Mand River as Capoeta capoeta intermedia ((Gh. Izadpanahi, pers. comm., 1995; M. Rabbaniha (pers. comm., 1995; Abdoli, 2000), the Esfahan basin including the Dopolan River, Gav Khuni marsh (Keyserling, 1861), the Tigris River basin in the Regab River near Kermanshah, Nashad River in the Divadarreh region, Kordestan and near Borujerd, Lorestan (records need verification by specimens), the Tedzhen River basin including the Jam and Kashaf rivers (Berg, 1949). This variable species is also recorded from the Dasht-e Kavir, Bejestan. Sirjan and  Namak Lake basins (Esmaeili et al., 2011?).

The subspecies Capoeta capoeta heratensis (the Transcaspian khramulya) is found in the Tedzhen or Harirud basin of Iran and eastwards including the Kashaf River (Abdoli, 2000). This subspecies is also recorded from the Karakum Canal and Kopetdag Reservoir in Turkmenistan (Shakirova and Sukhanova, 1994; Sal'nikov, 1995) and may eventually reach Iranian waters in the Caspian Sea basin.

Zoogeography

Saadati (1977) suggests that this species entered the Dasht-e Kavir basin from either the Tigris River basin, the Hari River basin or the Caspian Sea basin. See also above under genus.

Habitat

The habitat of this species in the Surkhandar'ya of Uzbekistan is backwaters and channels with weak current and silt beds as well as reservoirs (Amanov, 1970). In Iran, it is one of two most abundant species in Caspian rivers along with Alburnoides bipunctatus (= eichwaldii) (Iranian Fisheries Research and Training Organization Newsletter, 19:4, 1998). Ghasemi and Mustafayev (2008) found this species in the Aras River was the most dispersed and had the highest frequency (56.6%) of 17 species collected. It was found to be resistant to environmental changes, such as flooding, in the Madarsoo River in Golestan (Rezaei et al., 2008). Günther (1899) found that Capoeta capoeta placed in saline Lake Orumiyeh water died in 3.5 minutes.

Age and growth

Life span is over 8 years and catches in Uzbekistan are dominated by fish 3-4 or 4-5 years old. Growth is fastest in the first two years of life (Amanov, 1970). Life span in Azerbaijan is over 6 years (Abdurakhmanov, 1962), in northern Anatolia 6 years where vertebrae followed by scales were the best structures for aging (Polat and Işik, 1995), while in Georgia life span exceeds 9 years (Elanidze, 1983), and in Lake Gotchka, Armenia 10 years but only 4+ years in Lenkoran (Bănărescu in Bănărescu, 1999). Günther (1899) reports on a male fish from Ula in the Lake Orumiyeh basin which was only 12.5 cm long yet a sexually mature male, perhaps an instance of a dwarf form. Canbolat et al. (1999) found life span to be over 9 years in Çıldır Lake, Turkey for Capoeta capoeta capoeta. Fish aged 6 years dominated at 31.5% and 61.7% of the sample was female.

In Madarsoo Stream of Golestan National Park, this species had age groups 0-10 years and growth parameters were Lt = 229.67 mm and K = 0.54 for males, 327.95 mm and 0.18 for females (Koohestan Eskandari, 2003). Rezaei et al. (2007) also examined this fish population in the Madarsoo after two floods in 2001-2002. Growth parameters were L = 249 mm, K= 0.22 and t0 = -0.30 for males and  L = 306 mm, K= 0.21 and t0 = -0.38 for females. Length-weight relationships were W = -4.48 + 3.03TL for males and W = -4.59 + 3.0551TL in females, showing good feeding condition and positive isometric growth. Males were smaller than females as they matured earlier. Male to female ratio was 1.5:1, significantly different. Age range was 1+ to 5+ for males and 2+ to 8+ years for females. The dominant age was 2+ years.  Length was greatly decreased compared to previous studies and the population was younger, attributed to the floods.

Gholizadeh et al. (2009) studied a population in the Zarrin-Gol Stream in Golestan amd found age group 0+ was the most common at 59% and age groups 3+ and 4+ were the least common at 1%. Instantaneous growth of fish at age 3+ was much lower than younger age groups. The length-weight relationship was W = 0.00003xL2.822 and the von Bertalanffy equation was Lt = 223.8 (1-exp[0.185(t+1.8)].

Maku Dam lake in West Azarbayjan has an estimated 9.4-10.7 tonnes of this species with a maximum sustainable yield of 4.5-4.8 t (Saiad Bourani and Ghaninejad, 2004). Average length of this population was 23.9 cm, weight was 1626.8 g and age was 2.6 years. Most fish were 3+ years old and 5+ fish were at a minimum. Infinite length and the growth coefficient were computed as 35.6 cm and 0.39 per year. Total mortality was 0.74, natural mortality 0.37 and fishing mortality 0.37. The Yasalegh Stream in the Gorgan River basin had a male to female ratio of 1:0.54, a maximum weight of 71.2 g for males and 119.4 g for females, and age range of 0-3 years, von Bertalanffy growth equations of Lt = 190(1-exp{-0.462 [t+1]} for males and Lt = 230(1-exp{-0.472 [t+0.742]} for females, and weight growth was isometric (b = 3.052 for males and 3.050 for females). Tilabad River fish had an age structure of 0-4 years and the Talar River 2-4 years, similar to Yasalegh Stream but differing from the Madarsoo Stream. The fish in the latter stream had better living conditions in a national park, no pollution, no fishing, no competition from exotic carps, no other human disturbances, no environmental stress and no food shortages. Patimar et al. (2009) found b values ranging from 2.647 (male at Chekchai) to 2.964 (females at Madarsu) indicating negative allometric growth for fish in the Gorgan River basin. They interpreted this variation to the species' response to different habitat conditions.

Food

Food is mainly detritus and ooze, with some higher plants and small amounts of blue-green algae, which is digested in an intestine almost 7 times longer than the body (Amanov, 1970). Small benthic invertebrates may also be included, such as chironomids and molluscs (Bănărescu in Bănărescu, 1999). In Maku Dam lake, this species is a detritivore consuming Chrysophyta from the phytoplankton and Cyclotella a diatom, from the benthos as well as Chironomidae and Ephemeroptera (Valipour, 2004). In the Talar and Yasalegh rivers of the eastern Caspian Sea basin, 27 genera of phytoplankton were identified in the diet, with Chrysophyta being dominant, but with some differences between older and younger fish in the species consumed (Mostafavi and Abdoli, 2005).

Reproduction

Sexual maturity in Uzbekistan is at ages 2-4 years and lengths of 15-20 cm or in some populations at 10-14 cm. Some fish mature as dwarfs before age 1 and Berg (1948-1949) reports males 8.4 cm long of the subspecies gracilis can be mature. Spawning may take place at any time during the period from March to September (Berg,1948-1949) and is intermittent with the first spawning accounting for up to 85% of the eggs and the subsequent two spawnings for the remainder. The yellow eggs have a diameter up to 2.2 mm in the first spawning, up to 0.75 mm in the second and 0.65 mm in the third. Fecundity is up to 86,800 eggs. Eggs are laid at 50-60 cm on sand and stone beds and in water temperatures of 16-23°C (Amanov, 1970). Fecundity in the Kura River may reach 93,861 at 36-40 cm but this is for C. capoeta capoeta and fecundity for C. capoeta sevangi is less (Bănărescu in Bănărescu, 1999). Eggs are shed in running water and on lake shores, and eggs are covered by sand or small stones.

Rezaei et al. (2007) found no change in reproductive characteristics after floods in the Madarsoo Stream population. Mean fecundity was 3116 eggs and the maximum gonadosomatic index was in June.

Parasites and predators

Williams et al. (1980) report the helminths Khawia armeniaca (a cestode) and Acanthocephalorhynchoides cholodkowskyi (an acanthocephalan) from this species in the Zayandeh River at Esfahan. Molnár and Jalali (1992) record for this species the monogeneans Dactylogyrus chramulii, D. gracilis and D. lenkorani in the Safid Rud, D. chramulii and D. lenkorani in the Beshar River of the Persian Gulf drainage, D. gracilis and D. lenkorani in the Zayandeh Rud, D. lenkorani in the Tonekabon and Tajan rivers of the Caspian Sea drainage and the Kor River drainage of Fars, and D. pulcher from the Safid, Tajan, Tonekabon and "Ghasemlu" rivers of the Caspian Sea basin and the Jajrud of the Namak Lake basin. Shamsi et al. (1997) report Clinostomum complanatum, a parasite causing laryngo-pharyngitis in humans, from this species, the highest rate in 9 species examined. Malek (1993) and Malek and Mobedi (2001) report Clinostomum complanatum from this species in Mazandaran, the Shiroud. Up to 60 parasites per fish are recorded, with female fish having the highest infestation (the later study showing no difference between male and female fish), infestation decreasing with increase in body length, and parasites being concentrated in the gill cavity and pharynx.

Masoumian and Pazooki (1998) surveyed myxosporeans in this species in Gilan and Mazandaran provinces, finding Myxobolus musayevi and M. samgoricus. The crustacean parasite Tracheliastes polycolpus is reported from the fins of this species in the Mahabad Dam reservoir (Abdi, 1999; www.mondialvet99.com, downloaded 31 May 2000). Jafari et al. (2001) isolated the acanthocephalan Dendronucleata dogieli from fish in the Zarrineh River, West Azerbaijan. Masoumian et al. (2002) investigated parasites from this fish in the Aras and Mahabad dams in northwest Iran and found the protozoan Myxobolus musayevi which is also recorded from this fish in the Tajan River in Mazandaran. Mokhayer et al. (2002) report Acanthocephalorhynchoides cholodkowskyi (Quadrigyridae) from the midgut and Tracheliastes polycolpus (Lernaeopodidae) on the fins of this fish in Golestan National Park, with more parasites on male fish and differences by season and station. Naem et al. (2002) found the following parasites on the gills of this species from the western branch of the Safid River, namely the protozoan Trichodina sp. and the monogenean trematode Dactylogyrus lenkorani. Mirhasheminasab and Pazooki (2003) list Ergasilus peregrinus, Tracheliastes polycolpus and Lernaea cyprinacea from this species in Mahabad Reservoir, the latter being the most dangerous parasite. Rohei Aminjan and Malek (2004) found 9 parasite species in fish from the Shiroud, namely the trematodes Clinostomum complanatum, Diplostomum spathaceum, Posthodiplostomum cuticola, Allocreadium sp., the monogeneans Dactylogyrus pulcher, D. lenkorani, Gyrodactylus mutabilitas and the nematodes Rhabdochona fortunatowi and Capillaria sp. Masoumian et al. (2005) recorded the protozoan parasites Ichthyophthirius multifilis, Trichodina perforata, Chilodonella, sp., Amphileptus branchiarum, Tetrahymena pyriformis, Apiosoma sp., and Vorticella sp. from this species in water bodies in West Azarbayjan. Araghi Soureh and Jalali Jafari (2005) recorded Dactylogyrus gracilis, D. charmulii, D. lenkorani and D. kendalanicus from this species in the Mahabad River of the Lake Orumiyeh basin, the latter species being a new record for Iran. Pazooki et al. (2007) recorded various parasites from localities in West Azarbayjan Province, namely Diplostomum spathaceum, Ligula intestinalis, Digrama sp., Rhabdochona hellichi, Argulus foliaceus, Allocreadium isoporum, Lamprolegna compacta, Myxobolus cristatus and M. musajevi. Pazooki et al. (2005) record Tracheliastes longicollis, Lamprolegna compacta, Neoechinorhynchus rutili, Capillaria sp., Myxobolus musajevi, M. cristatus, Trichodina perforata, Chilodonella piscicola, Ichthyophthirius multifilis and Ichthyobodo necatrix from this species in waterbodies of Zanjan Province. Pazooki et al. (2006) record the monogeneans Dactylogyrus chramuli, D. gracilis, D. lamellatus, D. lenkorani, D. pulcher and Gyrodactylus sp. from this fish in Zanjan Province. Masoumian et al. (2007) record the myxosporean parasite Myxobolus musajevi from this species in the Zayandeh River. Miar et al. (2008) examined fish in Valasht Lake and the Chalus River, Mazandaran and found the metazoan Myxobolus saidovi. Maleki and Malek (2007) examined fish from the Shirud in the Caspian Sea basin and recorded the digeneans Posthodiplostomum cuticola, Diplostomum spathaceum, Clinostomum complanatum and Allocreadium sp. Barzegar et al. (2008) record the digenean eye parasite Diplostomum spathaceum from this fish. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Argulus foliaceus, Ergasilus sp., Lamproglena compacta, Lernaea sp., Tracheliastes longicollis and Tracheliastes polycolpus on this species.

Economic importan

The subspecies Capoeta capoeta heratensis is a food fish in Uzbekistan (Amanov, 1970) and C. capoeta sevangi and C. capoeta capoeta are commercially important in Lake Gotchka, Armenia and eastern Georgia and Azerbaijan respectively (Bănărescu in Bănărescu, 1999). It is also used in sport fishing in Iran (Samaee et al., 2006). Shiri et al. (2009) report a case of ichthyotoxism after eating fried eggs of this species. Nausea resulted after one minute, and the victim was hospitalised with severe chest pains. No vomiting occurred as this was the only food eaten and symptoms appeared rapidly. Raw consumption should be avoided and even cooked fish or inadequately cleaned fish can be dangerous.

Conservation

Kiabi et al. (1999) consider this species to be of least concern in the south Caspian Sea basin according to IUCN criteria. Criteria include sport fishing, abundant in numbers, habitat destruction, widespread range (75% of water bodies), present in other water bodies in Iran, and present outside the Caspian Sea basin.

Further work

The relationships of the various subspecies need study to determine if they are in fact good species.

Sources

Type material: See above, Capoeta gibbosa (ZISP 11104).

Iranian material: CMNFI 1970-0512, 3, mm standard length, Gilan, Shalman River (37º08'N, 50º15'E); CMNFI 1970-0514, 17, ? mm standard length, Gilan, Shafa River estuary (37º55'N, 49º09'E); CMNFI 1970-0516, 6, ? mm standard length, Gilan, Lemir River (38º14'N, 48º52'30"E); CMNFI 1970-0519, 2, ? mm standard length, Gilan, Chelvand River (ca. 38º18'N, ca. 48º52'E); CMNFI 1970-0520, 2 ?7 on data sheet, 94.0-100.9 mm standard length, Gilan, Astara River (ca. 38º25'N, ca. 48º52'E); CMNFI 1970-0521, 3, ? mm standard length, Gilan, Safid River near Lulaman (no other locality data); CMNFI 1970-0522, 10, ? mm standard length, Gilan, Safid River at Astaneh bridge (37º16'30"N, 49º56'E); CMNFI 1970-0525, 5, 92.8-146.1 mm standard length, Gilan, Safid River near Mohsenabad (ca. 37º22'N, ca. 49º57'E); ?see data sheets 78, 22.1-160.4 CMNFI 1970-0526, 8, ? mm standard length, Gilan, Safid River 6 km below Astaneh bridge (37º19'N, 49º57'30"E); CMNFI 1970-0531, 7, 60.2-84.6 mm standard length, Mazandaran, Larim River (36º46'N, 52º58'E); CMNFI 1970-0536, 2, ?5 on data sheets 101.4-125.4 mm standard length, Gilan, Siah River estuary (36º53'N, 49º32'E); CMNFI 1970-0538, 1, ?5 on data sheet 95.4 mm standard length, Gilan, Qezel Owzan River near Manjil (36º44'N, 49º24'E); CMNFI 1970-0557, 3, ? mm standard length, Azarbayjan-e Bakhtari, Shaher Chay (no other locality data); CMNFI 1970-0559, 6, 83.9-125.4 mm standard length, Azarbayjan-e Bakhtari, Baranduz Chay (ca. 37º25'N, ca. 45º10'E); CMNFI 1970-0568, 9, ? mm standard length, Gilan, Caspian Sea at Kazian beach (ca. 37º29'N, ca. 49º29'E); CMNFI 1970-0577, , mm standard length, Gilan, Caspian Sea at Astara (ca. 38º26'N, ca. 48º53'E); CMNFI 1970-0583, 8, 34.1-93.9 mm standard length, Gilan, Nahang Roga (37º28'N, 49º28'E); CMNFI 1979-0589, , mm standard length, Gilan, Safid River opposite Kisom (37º12'N, 49º54'E); CMNFI 1979-0242, 27, 25.6-107.0 mm standard length, Fars, river at Izadkhvast (31º31'N, 52º07'E); check ID? CMNFI 1979-0249, 33, 66.4-114.2 mm standard length, Esfahan, stream at Dizaj (31º55'N, 51º30'E); check ID? CMNFI 1979-0429, 1, ? mm standard length, Mazandaran, Chalus River (36º34'N, 51º23'E); CMNFI 1979-0432, 1, ? mm standard length, Mazandaran, Sardab River branch (36º41'N, 51º22'E); CMNFI 1979-0434, 1, ? mm standard length, Mazandaran, Shir River (36º51'N, 50º49'E); CMNFI 1979-0435, 1, ? mm standard length, Gilan, stream 10 km west of Ramsar (36º57'N, 50º37'E); CMNFI 1979-0433, 1, 115.2 mm standard length, Mazandaran, stream 18 km west of Chalus (36º42'N, 51º15'E); CMNFI 1979-0438, 2, 142.4-144.8 mm standard length, Gilan, Gholab Ghir River (37º27'N, 49º37'E); CMNFI 1979-0441, 1, 121.9 mm standard length, Gilan, river 14 km south of Hashtpar (37º42'N, 48º58'E); CMNFI 1979-0443, 1, ? mm standard length, Gilan, river 34 km west of Hashtpar (38º06'N, 48º53'E); CMNFI 1979-0444, 1, ? mm standard length, Gilan, Chubar River (38º11'N, 48º52'30"E); CMNFI 1979-0446, 1, ? mm standard length, Gilan, Astara River (38º26'30"N, 48º51'E); CMNFI 1979-0449, 2, ? mm standard length, Azarbayjan-e Khavari, river 18 km from Khalkhal (ca. 37º42'N, ca. 48º27'E); CMNFI 1979-0451, 30, 35.8-97.3 mm standard length, Azarbayjan-e Khavari, Qezel Owzan River (ca. 37º30'N, ca. 47º57'E); CMNFI 1979-0452, 1, 79.7 mm standard length, Azarbayjan-e Khavari, Qezel Owzan River 6 km from Mianeh (37º23'N, 47º45'E); CMNFI 1979-0453, 24, 36.1-111.1 mm standard length, Zanjan, Zanjan River (37º06'N, 47º56'E); CMNFI 1979-0469, 2, 56.6-76.2 mm standard length, Mazandaran, river 36 km west of Alamdeh (36º37'30"N, 51º35'E); CMNFI 1979-0474, 1, ? mm standard length, Mazandaran, Tajan River (36º34'N, 53º05'E); CMNFI 1979-0475, 1, 86.4 mm standard length, Mazandaran, stream on road to Bandar-e Shah (36º46'N, 54º00'E); CMNFI 1979-0480, 2, ? mm standard length, Mazandaran, Gorgan River at Gonbad-e Kavus (37º15'30'N, 55º09'E); CMNFI 1979-0481, 3, 101.9-188.0 mm standard length, Mazandaran, stream 3 km west of Ghalahleekesh (37º18'30"N, 55º31'E); CMNFI 1979-0482, 2, ? mm standard length, Mazandaran, river 2km west of Ghalahleekesh (37º19'30'N, 55º31'E); CMNFI 1979-0483, 4, 121.6-160.5 mm standard length, Mazandaran, river 28 km west of Dasht (37º23'30"N, 55º51'30"E); CMNFI 1979-0485, 3, 71.2-99.1 mm standard length, Khorasan, stream 28 km west of Bojnurd (37º33'N, 57º04'E); CMNFI 1979-0486, 66, 17.5-97.8 mm standard length, Mazandaran, stream in Atrak River draiange (37º44'N, 56º18'E); CMNFI 1979-0487, 20, ? mm standard length, Mazandaran, spring 2 km from Maraveh Tappeh (37º54'N, 55º58'E); CMNFI 1979-0488, 9, 29.7-140.4 mm standard length, Mazandaran, Atrak River at Maraveh Tappeh (37º55'N, 55º57'30"E); CMNFI 1979-0489, 78, ? mm standard length, Mazandaran, stream 13 km from Maraveh Tappeh (37º50'N, 55º53'E); CMNFI 1979-0490, 14, 21.0-108.4 mm standard length, Mazandaran, stream in Gorgan River drainage (ca. 37º39'N, ca. 55º42'E); CMNFI 1979-0491 2, ? mm standard length, Mazandaran, Gorgan River northeast of Kalaleh (ca. 37º33'N, ca. 55º44'E); CMNFI 1979-0492, 25? check jar, 9.3-183.4 mm standard length, Mazandaran, river in Gorgan River drainage (37º05'N, 55º15'E); CMNFI 1979-0695, 13, ? mm standard length, Gilan, Safid River at Manjil Bridge (36º46'N, 49º24'E); CMNFI 1997-0003, , mm standard length, (); CMNFI 1980-0116, 1, ? mm standard length, Gilan, Safid River at Astaneh (37º16'30"N, 49º56'E); CMNFI 1980-0120, , mm standard length, Mazandaran, Babol River at Babol Sar (36º43'N, 52º39'E); CMNFI 1980-0121, , mm standard length, Gilan, Shafa River estuary (37º35'N, 49º09'E); CMNFI 1980-0123, , mm standard length, Gilan, Safid River (ca. 37º22'N, ca. 49º57'E); CMNFI 1980-0141, , mm standard length, Gilan, Lisar River estuary (37º59'N, 48º56'E); CMNFI 1991-0163, , mm standard length, Mazandaran, Ramian River (36º58'N, 55º07'E); CMNFI 1993-0138, 1, mm standard length, Khorasan, Bazangan Lake (36º18'N, 60º27'E); CMNFI 2007-0014, 4, 39.4-99.1 mm standard length, Khorasan, pool in Kuh-e Sang Park, Mashhad (ca. 36º18'N, ca. 59º36'E); CMNFI 2007-0086, 6, ? mm standard length, Azarbayjan-e Khavari, Qareh Su basin near Nir (ca. 38º02'N, ca. 48º00'E); CMNFI 2007-0087, 1, ? mm standard length, Azarbayjan-e Khavari, Qareh Su north of Ardebil (38º22'N, 48º19'E); CMNFI 2007-0088, 5, ? mm standard length, Azarbayjan-e Khavari, Qareh Su east of Lari (38º30'N, 48º03'E); CMNFI 2007-0089, 4, ? mm standard length, Azarbayjan-e Khavari, Ahar Chay at Ahar (38º28'N, 47º03'E); CMNFI 2007-0093, 13, ? mm standard length, Azarbayjan-e Bakhtari, Qotur River south of Khvoy (38º30'N, 44º58'E); CMNFI 2007-0094, 6, ? mm standard length, Azarbayjan-e Bakhtari, Nazlu River north of Reza'iyeh (ca. 37º42'N, ca. 45º04'E); checkID? CMNFI 2007-0095, 2, ? mm standard length, Azarbayjan-e Bakhtari, Shahr Chay southwest of Reza'iyeh (ca. 37º27'N, ca. 44º56'E); checkID? CMNFI 2007-0096, 5, ? mm standard length, Azarbayjan-e Bakhtari, Qasemul River in Baranduz Chay basin (ca. 37º25'N, ca. 45º10'E); checkID? CMNFI 2007-0098, 2, ? mm standard length, Azarbayjan-e Bakhtari, river south of Mahabad (ca. 36º42'N, ca. 45º41'E); CMNFI 2007-0099, 1, ? mm standard length, Azarbayjan-e Bakhtari, Kalwi Chay west of Mahabad (ca. 36º35'N, ca. 45º25'E); checkID? CMNFI 2007-0101, 1, ? mm standard length, Azarbayjan-e Bakhtari, Tata'u River south of Miandow Ab (ca. 36º54'N, ca. 46º07'E); CMNFI 2007-0102, 4, ? mm standard length, Azarbayjan-e Bakhtari, Zarineh River near Miandow Ab (ca. 37º00'N, ca. 46º07'E); CMNFI 2007-0103, 9, ? mm standard length, Kordestan, Zarineh River basin north of Saqqez (ca. 36º18'N, ca. 46º16'E); CMNFI 2007-0104, 4, ? mm standard length, Kordestan, Zarineh River basin south of Saqqez (ca. 36º12'N, ca. 46º18'E); CMNFI 2007-0105, 7, ? mm standard length, Kordestan, Zarineh River basin south of Saqqez (ca. 36º06'N, ca. 46º20'E); CMNFI 2007-0106, 9, ? mm standard length, Kordestan, Qezel Owzan River basin near Divandarreh (ca. 35º52'N, ca. 47º05'E); CMNFI 2007-0107, 10, ? mm standard length, Kordestan, Qezel Owzan River basin near Bijar (ca. 35º54'N, ca. 47º20'E); ZSM 24500, 6, 24.2-31.0 mm standard length, Khorasan, stream near Bojnurd (no other locality data); uncatalogued, 2, 75.8-85.5 mm standard length, Khorasan, Hari River at Sarakhs (36º32'N, 61º11'E).

Capoeta damascina
(Valenciennes, 1842)

Common names

sardeh (= cold one, probably zardeh = yellow one, is more correct and appropriate based on yellow-tinged flank) or سياه ماهي (= siah mahi, meaning black fish) in the Caspian basin; siah mahi damascina; tu'ini (meaning unknown) or gel cheragh (= mud-eater, mud-grazer) or tu'ini gelkhorak (= mud-eater, mud-grazer) in Khuzestan; qezel ala (red spots) in Chahar Mahall but mistakenly.

[twena, toyoueni or toueni, bertin or bartin, tin, zardah masih, tela shami, tela Damascus; kollur, kellur, kollur hadjiari (= the pilgrim or migrating kollur), kellur dischileki (= the strawberry-coloured kollur), kollur achmar (= the red kollur) and kollur aschkar (= the brown kollur), all at Aleppo; all in Arabic; Mesopotamian barb].

Systematics

Gobio damascinus was described from the "fleuve de Damas" (= river of Damascus, Syria).

Synonyms are Scaphiodon capoeta Heckel, 1843 (non sensu Güldenstädt, 1773) described from "Aleppo", Scaphiodon fratercula Heckel, 1843 described from "Gewässern von Damascus", possibly Scaphiodon Umbla Heckel, 1843 described from the "Tigris bei Mossul", Scaphiodon socialis Heckel, 1843 described from "Um Damascus" (= around Damascus) (Heckel, 1843b) and later more completely from the "Orontes" (Heckel, 1846-1849a) (placed in Scaphiodon Capoëta of Heckel by Steindachner (1864)), Scaphiodon peregrinorum Heckel, 1843 described from "Aleppo" and later from "Fluss Kueik bei Aleppo", Chondrostoma syriacum Valenciennes, 1844 from Abraham's River at the foot of Mount Sinai, Egypt (the correct locality is probably in the Jordan River basin (Coad and Krupp, 1994)), Scaphiodon Amir Heckel, 1849 described from the "Araxes" (= Kor River, Fars), Scaphiodon niger Heckel, 1849 described from the "Araxes oder Benth-Amir" (= Kor River, also known as the Bandamir River), Scaphiodon Saadii Heckel, 1849 described from the "Quellen des Saadi" (Sa`di at 29°37'N, 52°35'E, now within the city of Shiraz) and the "Nähe von Persepolis" (= probably the Pulvar (= Sivan) River near Persepolis, Fars), Scaphiodon chebisiensis Keyserling, 1861 from "Wasserleitung in Chebis" (= canal in Chebis, probably Khabis or Shahdad at 30°25'N, 57°42'E in Kerman), Scaphiodon rostratus Keyserling, 1861 from "Wasserleitungun in der Umgegend von Jezd. Das abgebildete Exemplar stammte aus Meibut" (= canals in the vicinity of Yazd. The specimen drawn originated from Meibut, probably Meybod at 32°14'N, 54°01'E), Barbus belayewi Menon, 1960 (Menon and Yazdani (1968) date this species as 1960, presumably the 1956 edition of the journal was delayed) from the "Tigris, Baghdad, Iraq", and Capoeta capoeta intermedia Bianco and Banarescu, 1982 (non Capoeta intermedia Temminck and Schlegel, 1846 = Acheilognathus lanceolata (Temminck and Schlegel, 1846) (see Boeseman, 1947)) described from the "Mand River near Akbar, southern Iran".

The synonymy of Barbus belayewi is suggested by F. Krupp (in litt., 1986) and W. Rainboth (pers. comm., 1986). The synonymy of S. fratercula is pointed out by Berg (1949) since the species was founded on low lateral line scale counts, a variable character in C. damascina, and on a larger orbit but Heckel's comparison was between fish of greatly differing size and no allowance was made for allometry.

Karaman (1969) places damascina in Capoeta capoeta as a subspecies and umbla as another subspecies. Berg (1949) and Saadati (1977) recognise umbla as a distinct species. The latter is distinguished from the former by a higher scale count (87-99), higher dorsal fin branched rays (9-10), longer dorsal fin, longer caudal fin (shorter than or equal to head length in C. damascina), a markedly transverse mouth, and a weaker dorsal fin spine. Saadati (1977) considers fratercula to be a distinct species from the Tigris and Mand rivers in Iran based on scale count (58-66), more gill rakers (20-22), and a more serrated dorsal fin spine; or a subspecies of Capoeta capoeta based on a close similarity in scale counts, average number of gill rakers, and the dorsal fin origin being anterior to that of the pelvic fins. He also considers that Scaphiodon niger from the Kor River of Fars is possibly a synonym of fratercula. Krupp (1985c) considers the synonymy of C. damascina and C. capoeta as extremely doubtful after examining topotypic material.

Bianco and Banarescu (1982) recognise Capoeta saadi as a distinct species based on an arched mouth rather than transverse as in most subspecies of Capoeta capoeta, with a lightly developed horny cover on the lip, a feebly ossified dorsal fin spine, 13-17 gill rakers, modally 8 dorsal fin branched rays, 53-76 lateral line scales and 24-28 scales around the caudal peduncle. However they do point out the extreme variability in scale counts, for example, from fish taken in the same locality and even between opposite sides of the same fish (5 more scales on one side than the other!). Designation of subspecies on such variable characters is difficult and would require very large series and multivariate analysis techniques. Bianco and Banarescu (1982) regard C. c. intermedia as intermediate between C. c. umbla and their C. c. macrolepis on the basis of scale counts, gill raker counts, smaller transverse mouth than in umbla and a rather light colouration.

Capoeta damascina, with a wide distribution and wide variation in morphology, must be regarded as a species complex until detailed analyses can be carried out. Final resolution of the species composition of this complex may well require extensive material for molecular analyses, as well as re-examination of types and topotypic material over the whole range of the taxon. Samaee and Patzner (2011) examined fish from 6 river systems in Iran morphometrically and were able to distinguish distinct groups. However, as they point out, much more work needs to be done to determine if this variation is genetic differentiation or phenotypic plasticity, or a combination of the two.

The syntypes of Gobio damascinus are in the Muséum national d'Histoire naturelle, Paris (MNHN 4494, 2 specimens, 169-179 mm standard length, Damascus, Syria, Bové, MNHN 3948, 1, 289 mm standard length, Nahr Barada, Syria and MNHN A.3947, 1, 169 mm standard length, Syria) (Krupp, 1985c). Bertin and Estève (1948) give 200-210 mm total length for MNHN 4494 and 330-390 mm total length for MNHN 3947, 3948 and A.789. Eschmeyer et al. (1996) list MNHN 4494 as the lectotype (as designated by Krupp and Schneider (1989) although this collection comprises two fish) with MNHN 3947 (1, dry) and MNHN 3948 (1, dry) and possibly MNHN A.789 (1) as paralectotypes. The latter is listed as a syntype in Bertin and Estève (1948) although the localities listed in this article "Fl. Jourdain, à Damas (Syrie)" is obviously an error on geographical grounds.

Syntypes of Scaphiodon capoeta are in the Naturhistorisches Museum Wien under NMW 51650 (1 fish), NMW 51831 (1), and NMW 55845-55846 (2). Heckel (1843) lists 2 specimens in his description.

The holotype of Chondrostoma syriacum is in the Muséum national d'Histoire naturelle, Paris under MNHN 1945 (Eschmeyer et al., 1996).

The holotype of Capoeta capoeta intermedia is in the Istituto di Zoologia dell'Universitá di L'Aquila, Italy (IZA 7892) and is 92.5 mm standard length, collected by P. Bianco and S Zerunian, 27/5/1976. There are 62 paratypes (IZA 7893) from the same collection as the holotype measuring 36-87 mm standard length and 13 paratypes uncatalogued in the Institutul de Stiinte Biologice, Bucuresti, Romania (ISBB) measuring 68-86 mm standard length (Bianco and Banarescu, 1982). Another paratype under IZA 7894 measures 105.5 mm standard length was examined by me. A paratype of Capoeta capoeta intermedia from the Mand River in Fars is in the Zoologischen Instituts und Zoologischen Museums der Universität Hamburg (ZMH 6090, 83.2 mm standard length) (Wilkens and Dohse, 1993; examined by me), one paratype from the Mand is in the California Academy of Sciences, San Francisco (CAS 48113), one paratype from the Mand is in the United States National Museum, Washington (USNM 227935), and 6 paratypes are in the Canadian Museum of Nature, Ottawa under CMNFI 1982-0367 (formerly IZA 7893).

The holotype of Scaphiodon fratercula was taken from "Gewässern von Damascus", the syntypes of Scaphiodon umbla from the "Tigris bei Mossul", the types of Scaphiodon socialis from "Um Damascus" (but listed as "Orontes" in the catalogue in Vienna, possibly in confusion as this part of the catalogue has been overwritten), and the types of Scaphiodon peregrinorum from "Um Aleppo" according to Heckel (1843b) and "Fluss Kueik bei Aleppo" according to Heckel (1846-1849a).

Two syntypes of Scaphiodon niger are in the Naturhistorisches Museum Wien under NMW 51655 with standard lengths of 140.4 and 188.5 mm (another syntype is under NMW 51654 (232.7 mm), and a fourth under NMW 51656 as seen by me; all 4 are listed as syntypes in the 1997 Vienna card index). Eight syntypes of Scaphiodon amir are under NMW 61472 and measure 42-59 mm standard length and there are also 6 fish under NMW 46081 (138.1-282.3 mm standard length); however the card index in 1997 lists only NMW 46081 (6) and 16508 (1, dried). Fifteen syntypes in the catalogue (18 seen by me and in the Vienna card index in 1997) of Scaphiodon saadii from Sa`di are under NMW 51666 (Eschmeyer et al. (1996) have 52666, apparently in error) and measure 58-123 mm standard length (18.3-123.8 mm standard length when measured by me) with a further 4 syntypes from Persepolis under NMW 55900 measuring 84-114 mm standard length (Kähsbauer, 1964; not in the 1997 card index). There is also 1 syntype (RMNH 3166) in the Rijksmuseum van Natuurlijke Historie, Leiden from NMW (Eschmeyer et al., 1996).

The catalogue in Vienna lists no fish opposite the name S. niger, 6 and 2 fish in one column and 5 in the adjacent column for Scaphiodon amir (cf. above), 10 fish in one column and 10 in the adjacent column for S. saadii (cf. above).

A dried syntype of C. umbla is in the Senckenberg Museum Frankfurt (SMF 6777, formerly NMW) (F. Krupp, pers. comm., 1985; ca. 262.3 m standard length), 2 syntypes are in Naturhistorisches Museum Wien (NMW 55932-55933) and another syntype is under NMW 55934. Eschmeyer et al. (1996) also lists NMW 79373-74, both dried. The catalogue in Vienna lists 2 fish in spirits and 2 fish stuffed and the card index in 1997 lists as syntypes NMW 55932-33 and 79373-74 (dried).

Eschmeyer et al. (1996) note that there are no types of Scaphiodon fratercula in the Naturhistorisches Museum Wien.

Two fish are labelled as syntypes of Scaphiodon socialis in the Naturhistorisches Museum Wien (NMW 55855) which agrees with Heckel's text although the catalogue lists only 1 specimen. Eschmeyer et al. (1996) state that there are no types at NMW presumably after Krupp and Schneider (1989) who state that NMW 55670 (1 fish), 55843 (2) and 55855 (2) are not types.

The types of Scaphiodon peregrinorum number 6 according to the catalogue in the Naturhistorisches Museum Wien and may comprise all or part of NMW 51658 (1), NMW 51659 (1), NMW 51660 (1), NMW 51661 (1), NMW 51662 (1), NMW 51663 (1) NMW 51664 (3), and NMW 51665 (1), all labelled as from "Kueik" and possibly RMNH 2681 (3) in the Rijksmuseum van Natuurlijke Historie, Leiden from NMW (Eschmeyer et al., 1996).

The types of Barbus belayewi are in the Zoological Survey of India, Calcutta, the holotype being ZSI F1046/2 and a paratype ZSI F1047/2 (Menon, 1960; Menon and Yazdani, 1968).

Types of Scaphiodon chebisiensis and Scaphiodon rostratus were not kept.? phrasing

Key characters

The mode of 9 dorsal fin rays, small scales, and the presence of large black blotches often distinguish this species from other Capoeta in Iran.

Morphology

Dorsal fin with 3-5 unbranched rays and 8-10 branched rays (Krupp (1985c) gives frequency distributions for his material from Turkey, Syria, Lebanon, Israel and Jordan as 8(52), 9(144) and 10(4)), anal fin with 3 unbranched and 5-6 branched rays (5(179), 6(21) after Krupp (1985c)), pectoral fin branched rays 15-20, and pelvic fin branched rays 8-10. Lateral line scales 60-99. Gill rakers 17-25 (Saadati (1977) gives 9-21!; Krupp (1985c) 12-18 for the lower arm of the arch, Berg (1949) up to 23 on the lower arm). Pharyngeal teeth 2,3,4-4,3,2, often 2,3,5-5,3,2, with spoon-shaped crowns. The mouth is usually horseshoe-shaped, seldom transverse. The last unbranched dorsal fin ray is moderate to strong with denticles along two-thirds of its length. Heckel (1846-1849b) distinguished his Scaphiodon amir and S. niger by the dorsal fin denticles being horizontal or perpendicular to the spine, not hooked downward as in related species. Berg (1949) did not attach any significance to this character, finding it in small fish from the Sarhadd of Baluchestan and from Jordan.

The karyotype for fish in the Tigris River basin of Turkey identified as Capoeta capoeta umbla is 2n=150, possibly hexaploid (Kılıç Demirok and Ünlü, 2001) and of fish identified as C. damascina from the Wadi Karak, Jordan 2n=148-150, indicating a hexaploid species (Gorshkova et al., 2002).

Body form is highly variable as are scale counts between populations and even within populations when large series are examined (Krupp, 1985c). Subspecific designations can only be valid if very large series from the whole range of the species are compared.

Meristic values for Iranian specimens are :

IZA7892, 7894 (2 fish) D8, A5, P15-15, V7, ll 62-64, gr 24

Sexual dimorphism

Males develop breeding tubercles around the snout and the posterior body on both sides of the lateral line (Khalaf, 1987).

Colour

The back is dark brown or brownish to olive or blue-grey, the flanks silvery with some yellowish tinges, sometimes golden, or yellow-brown or reddish-brown above the lateral line, silvery below. The belly is white to yellowish. Cheeks are golden. Dark brown or black spots numbering up to 20 may be scattered irregularly on the flanks. Fins are reddish-brown, yellowish or grey and may be hyaline. The caudal and pectoral fins may be very dark compared to other fins. The pectoral and pelvic fins may a light pink tinge. The cartilaginous edge to the lower jaw is bright yellow to red-yellow. The peritoneum is black. Some fish may be very black with only the underside of the head and belly yellowish-white (specimens described by Heckel (1846-1849b) as Scaphiodon niger; however since these fish "decompose quickly in the commonly used ethyl alcohol concentrations", they may have been poorly preserved and the black colouration resulted from partial decomposition).

Size

Attains 35.3 cm standard length, about 45.0 cm total length and 0.5 kg.

Distribution

Found from Turkey, Syria, Lebanon and Israel to Iran. In Iran, it is reported from the Tigris River including the the Regab River in Kurdistan and the Selakhor River near Borujerd, the Jarrahi, Marun, Karun, Kuhrang, Bazoft and Khersan rivers, throughout the Dez and Karkheh basins to their uppermost reaches; Lake Zarivar; Esfahan including the Dopolan and Zayandeh rivers, Dasht-e Kavir including the Jajarm and upper Kal Shur rivers, Namak Lake including the Karaj, Shur, Abhar, Qareh Su and Qom rivers; Kor River, Lake Maharlu, Gulf including the Zohreh River and its Kheirabad tributary and the Mand River near Akbar and its Shur (Dasht-e Palang) River tributary, Kerman-Na'in, Dasht-e Lut, Sirjan, Hormuz including the middle to upper Hasan Langi, Kul and its Shur River tributary, and Hamun-e Jaz Murian basins (Lovett, 1873; Nikol'skii, 1899; Berg, 1949; Kähsbauer, 1964; Spillman, 1972; Armantrout, 1980; Rainboth, 1981; Bianco and Banarescu, 1982; Abdoli, 2000; Ghorbani Chafi, 2000; R. Mehrani, pers. comm., 2000; Jalali et al., 2005; Esmaeili et al., 2011?).

Zoogeography

Its relationships with other Capoeta species is generally unclear, as is the status of isolated populations some of which have been named. The larger zoogeographical relationships of this and other Capoeta species remain uncertain. See also above under genus.

Habitat

Unknown in detail.

Age and growth

All males are mature at 18 cm and all females at 20 cm in Khalaf's (1987) study in the Lebanon. In Lake Kinneret, Israel, Stoumboudi et al. (1993) found that fish longer than 25 cm have developed gonads, occasionally males mature between 16 and 25 cm as did females between 20 and 25 cm. Khalaf et al. (2002) found 6 age classes (1+ to 6+) in the Nahr el Khalb, a Lebanese stream. Maximal growth was in July and August and minimal growth between December and February. Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 40 Iranian fish measuring 5.23-19.87 cm standard length. The a-value was 0.0282 and the b-value 2.890 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases).

Food

Khalaf (1985) and Spataru and Gophen (1986) examined the food of this species in Lebanon and Israel respectively. Benthic diatoms and filamentous algae are the main foods. Some other algal species and some zoobenthic organisms are present along with large quantities of mud. The species is classified as a phytobenthophagous fish, one that takes its food from bottom sediments. Leaf remains have also been found in gut contents. Abdoli (2000) lists variety of insects: Chironomidae, Formicidae, Epididae, Empididae, Tipulidae, Tabanidae, Simuliidae, Hydroptilidae, Grouvellinus, Elmis, Hydropsyche, Heptagenia, Baetis and hydracariens.

Reproduction

Al-Rudainy (2008) gives sexual maturity at 2-3 years in Iraq with spawning in May, absolute fecundity up to 53,000 eggs and relative fecundity up to 7300 eggs/g body weight, and average egg diameter 1.48 mm.

Khalaf (1987) examined the reproductive cycle in this species for Lebanese waters. Spawning begins in May and ends in July. Eggs number up to 5,138 and egg diameters are up to 2.2 mm. In marked contrast, Stoumboudi et al. (1993) found that gonad weights are greatest in January in Lake Kinneret, Israel, 4 months earlier. This may be evidence of different temperature regimes or populational variation. Fishelson et al. (1996) confirm that this species migrates in winter, December to February in the upper Jordan River of Israel, the process being initiated by rainfall and flooding and a decrease in temperature to 16-18°C. The gonadosomatic index is highest in February and the final months of reproduction are March to May. Lake dwelling fish aggregate and swim up streams as far as 25 km and altitudes of 400-900 m, fattening and ripening at the spawning site. They can jump rapids on this migration (and in Iran large fish cornered in small streams will jump over seine nets!). The females excavate a shallow nest in which to deposit adhesive eggs, up to 4.5 mm in diameter. Dozens of nests are found close together and sand and gravel stirred up by the excavation covers adjacent nests. After spawning the adults return downstream to the stream mouth and lake.

Parasites and predators

Dollfus (1970) describes a new cestode Coelobothrium monodi from this species at "Nasratabad", possibly from the Dasht-e Lut basin. Jalali et al. (1995) describe two new species of monogeneans, Dactylogyrus rohdeianus and D. capoetae, from fish caught in the "Chaghalnandi" River, a Karkheh River tributary north of Ahvaz. González-Solís et al. (1997) report the nematodes Rhabdochona denudata and Rhabdochona fortunatowi from this species in the Mand River, Fars. O. M. Amin (pers. comm., 1998) has identified the acanthocephalan Acanthocephalorhynchoides cholodkowskyi from specimens collected in the Mand River west of Shiraz, Fars. Jalali et al. (2002) and Jalali and Barzegar (2006) record Trichodina pediculus, Dogielius molnari, Gyrodactylus sp., Dactylogyrus carassobabrbi and D. lenkorani from this species in Lake Zarivar. Barzegar et al. (2004) examined this species for parasites in fish from the Beheshtabad river in Chahar Mahall va Bakhtiari Province and found Dactylogyrus lenkorani, Gyrodactylus pulcher, Dactyolgyrus sp., Allocreadium isoporum and Myxobolus molnari. Mehdipoor et al. (2004) record the monogeneans Dactylogyrus lenkorani and D. pulcher in Zayandeh River fish. Barzegar and Jalali (2006) report a parasite in this species from Kaftar Lake as Dactylogyrus lenkorani. Masoumian et al. (2007) record the myxosporean parasites Myxobolus samgoricus and M. varicorhini from this species in the Zayandeh River. Barzegar et al. (2008) record the digenean eye parasite Diplostomum spathaceum from this fish. Nazari Chamak et al. (2010) found the following myxozoan parasites in the genus Myxobolus: buckei, cristatus, karelicus, musajevi, samgoricus, suturalis and varicorhini in fish from the Halil River, Kerman. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Lernaea cyprinacea on this species.

Economic importance

This species is of no economic importance in Iran although in Israel annual catches in Lake Kinneret have been as high as 29 tonnes or 7% of the total fishery (Spataru and Gophen, 1986). Heckel (1846-1849b) reports that this species was "greatly appreciated as food fish by the local people" in the Kor River basin, Fars (as his Scaphiodon amir). Samaee and Patzner (2011) mention that it is fished recreationally in Iran.

The eggs are reputedly poisonous and this is said to account for the low population of introduced Oncorhynchus mykiss in Gahaar Lake, Lorestan (R. Mehrani, pers. comm., 2000).

Conservation

A widely distributed species, probably not in need of conservation. Endangered in Turkey (Fricke et al., 2007).

Further work

The relationships of isolated and named taxa under this species, and the relationships of this species to other Capoeta, would benefit from molecular analyses.

Sources

Type material: See above, Capoeta capoeta intermedia (IZA 7892, 7894, CMNFI 1982-0367 (formerly IZA 7893), ZMH 6090) P. G. Bianco. ? check this last speciemne; Scaphiodon amir (NMW 61472, 46081); Scaphiodon niger (NMW 51655, 51654, 51656); Scaphiodon saadii (NMW 51666).

Iranian material: CMNFI 1993-0154, 1, ? mm standard length, Markazi, Sharra River near Far (34º03'N, 49º20'E); CMNFI 1995-0020, , mm standard length, (); CMNFI 1995-0021, , mm standard length, (); CMNFI 1997-0004, , mm standard length, ();

Comparative material: BM(NH) 1974.2.22:1856, 227.1 mm standard length, Iraq, Mosul (36º20'N, 43º08'E); BM(NH) 1934.9.5:3-5, 6, 14.8-45.5 mm standard length, Rawanduz River, Razanok.

Capoeta fusca
Nikol'skii, 1897

Common names

سياه ماهي (= siah mahi, meaning black fish).

Systematics

The 2 syntypes, listed in Latin as from "Mondechi in Persia orientali", are in the Zoological Institute, St. Petersburg (ZISP 11108) and measure 121.9-172.9 mm standard length. Berg (1949) gives the locality in Russian as "Mondekhi, northern periphery of the Bajistan Salt Desert in southeast Khorasan". This locality is possibly Mandehi or Miandehi at 34°53'N, 58°38'E. Nikol'skii (1897) lists a series of specimens in Latin, presumably all of which he regarded as types, sic:- "11108. Mondechi in Persia orientali. 12.IV.96 (2). 11109. Persia orientalis. 1896. (6). 11110. Persia orientalis. 1896. (5). 11111. Persia orientalis. 1896. 11112. Kuss in Persia orientali. 6.IV.96.", the last two lacking number of specimens. Berg (1949) gives 20 specimens for 11109, 6 specimens for 11110, and 1 specimen for each of the last two. Catalogue dates in ZISP for all these are 26.IV.96, presumably new style, while Berg (1949) gives new style dates 24.IV.1896 for the first and 18.IV.1896 for the last (and this last is 26.IV.1896 in the catalogue). Only ZISP 11108 specimens are regarded as syntypes by Berg (1949). Berg (1949) also points out the confusion over the date when Zarudnyi, the collector, was at "Kuss" (= Khusf at 32°46'N, 58°53'E) given by Nikol'skii as 6.IV.96 old style but on this date Zarudnyi was at "Kiaz-khak" near Asadabad (35°38'N, 59°21'E) south of Mashhad and only reached Khusf on 8 (or 20 new style).VI.96. This is not particularly critical in this instance but serves to point out the difficulties of reconciling literature, field notes, catalogues, and jar labels.

Capoeta nudiventris Nikol'skii, 1897 is a synonym. The syntypes are in the Zoological Institute, St. Petersburg (ZISP 11106) according to Berg (1949) and comprise 3 fish 92.4-121.5 mm standard length. Berg (1949) gives the type locality as "Zeride near Bajistan in southeast Khorasan, 30.IV.1896" (the date in the jar is 26.IV.1896). Nikol'skii (1897) lists 3 collections all from "Saride in Persia orientali. 18.IV.96." with numbers 11105, 11105 (presumably an error for 11106), and 11107 and 6 (actually 7 in the jar and according to Berg (1949)), 3, and 5 specimens respectively. Berg (1949) lists the 5 specimens under 11107 as from "Chakhak in the Al'kor region between Bajistan and Birjand. 9.V.1896", presumably at 33°17'N, 58°54'E. These 5 fish are 37.0-55.2 mm standard length, collected on 25.IV.1896 in the ZISP catalogue and not listed as types in the jar, nor in the catalogue, nor in Berg (1949). The 7 fish in ZISP 11105 measure 46.8-75.3 mm standard length, are from the same locality listed under ZISP 11106 in Berg (1949) and are listed as types in the ZISP catalogue, though not in Berg (1949). Judging from the labels and catalogue sheets, the types are probably from Sarideh at 34°22'N, 58°14'E and comprise 11105 and 11106.

Rainboth (1981) places both fusca and nudiventris in the genus Schizocypris on the basis of the enlarged scales around the vent and anal fin base, a condition reported on by Berg (1949) also but not considered by this latter author to warrant inclusion of these fish in Schizocypris.

Key characters

The strong mode of 7 branched dorsal fin rays, distribution, and the relatively low scale count aid in identifying this species.

Morphology

Dorsal fin with 3 unbranched rays and 7-8, strong mode at 7, branched rays, anal fin with 3 unbranched and 5 branched rays, pectoral fin branched rays 14-20 and pelvic fin branched rays 7-9. Lateral line scales 40-62, mostly 47-56. Scales are found regularly arranged over the whole body and are enlarged around the anus and anal fin base. There is a pelvic axillary scale. Scales are oval and have a subcentral, markedly anterior focus, numerous radii on all fields and moderate numbers of circuli. Gill rakers 11-20, short and touching the raker below when appressed. The mouth is horseshoe-shaped. The last unbranched dorsal fin ray is weak with only a few fine denticles along the basal half. The pharyngeal teeth are very spatulate up to the tip but are thick. There is an occasional trace of a fifth tooth in the major row but all the fish examined had only 4 strongly developed main row teeth. The gut is very elongate with several anterior and posterior loops.

Some populations or individuals may show a very light belly extending up onto the lower flank rendering scales hard to see. Capoeta nudiventris was apparently founded on specimens like this. Some scales low on the flank are incompletely imbricate and deeply embedded in the skin. Berg (1949) in examining the types of fusca and nudiventris found the extent of the scales ventrally to be the same and nudiventris is not naked on the lower flank and belly.

Meristics for Iranian specimens:- dorsal fin branched rays 7(77); anal fin branched rays 5(77); pectoral fin branched rays 14(1), 15(1), 16(8), 17(23), 18(26), 19(13) or 20(5); pelvic fin branched rays 7(8), 8(64) or 9(5); lateral line scales 46(4), 47(6), 48(8), 49(10), 50(10), 51(9), 52(9), 53(9), 54(9), 55(2) or 56(1); total gill rakers 13(1), 14(11), 15(25), 16(26), 17(11), 18(1), or 20(1); pharyngeal teeth 2,3,4-4,3,2(20); and total vertebrae 40(9), 41(42), 42(20) or 43(4).

Sexual dimorphism

Unknown but males presumably bear large tubercles in the breeding season.

Colour

The back and flanks are dark while below the lateral line the body can be very light. The dorsal, anal and caudal fin membranes are dark. Young fish may have a mid-lateral stripe as wide as the eye ending in an indistinct dark blotch on the caudal fin base. The peritoneum is dark brown to black.

Size

Reaches 21.5 cm total length (Johari et al., 2009).

Distribution

This species is found in eastern Iran in the Tedzhen River (including Kashaf River), Dasht-e Kavir, Bejestan, Dasht-e Lut and Sistan basins in rivers, springs and qanats, some of the latter not easily located on maps (Nikol'skii, 1899; Berg, 1949; Abdoli, 2000). A record from the "Schalman Rud" presumably in the Caspian Sea basin is most probably an error (Wossughi, 1978). Johari et al. (2009) record this species from the Ghoorghoori, Asafshad, Mardan Shah, Gazdmoo and Afin rivers in Qae'nat province and in 44 qanats of Birjand County in eastern Iran.

--- Zirkhuch may be Zir-e Kuh at 32°48'N, 59°50'E ? check this and if reasonable search for Zir and add in all text - Zirkhuch is in eastern or southeastern Khorasan for sure

Zoogeography

Saadati (1977) considers that this species entered eastern Iran from the west via the Namak Lake basin. See also above under genus.

Habitat

Karaman (1969) considers that this species shows the greatest adaptation among Capoeta species to desert life: an elongate and low body, scaleless belly in many individuals, weak spiny dorsal fin ray, reduced number of dorsal fin rays, short dorsal fin which can easily lie flat against the body, and the mouth structure. Johari et al. (2009) studied 10 qanats in Birjand County and found the following ranges: 3.8-24.9ºC, 0-6.3 p.p.t. salinity, 7.7-8.5 pH, 3.8-1164 μS, 6.3-13.8 mg/l dissolved oxygen, 0.31-11.5 mg/l nitrate, 0-0.8 mg/l nitrite, 0.04-0.29 mg/l ammonia, 185-750 mg/l total hardness, 2.17-815 total dissolved solids, 25-410 mg/l calcium, 0-100 mg/l magnesium, 0.16-340 mg/l sulphate, 2.3-27 mg/l potassium, 0.01-0.14 mg/l chlorine and 0.2-0.95 mg/l phosphate. No mortalities were noted in fish kept in salinities up to 10 p.p.t. for 120 hours, but higher levels started to show progressive mortalities. As salinity increased, fish became darker and dead fish were almost black. The fish exhibited schooling behaviour both in aquaria and in their natural environment.

Age and growth

Johari et al. (2009) found a total length/weight relationship of body weight = 0.01010 x TL2.9477 for 600 fish  from 10 qanats in Birjand County. Patimar and Mohammadzadeh (2011) examined fish from the Shadmehr qanat in south Khorasan and found a maximum age of 5+ years, negative allometric growth for males and isometric for females, males grew faster than females, and von Bertalanffy growth models Lt=18.74(1-e-0.33(t+0.473)) for males and Lt=22.35(1-e-0.32(t+0.333)) for females.

Food

Gut contents of the few fish examined contained fragments of large plants including large seeds, filamentous algae and sand grains. Johari et al. (2009) found this species to be herbivorous based on relative gut length and to be relatively gluttonous based on gut vacuity index. Large plants and filamentous algae made up 86.8% of the food but molluscs, aquatic insects and frog eggs were secondary foods. Feeding was highest in December and January before spawning and in August and September when presumably productivity was greatest. In March to May, the spawning season, feeding was reduced.

Reproduction

Fish caught in April and May have mature eggs along with some immature eggs, indicating that spawning may occur in stages. Fish caught in November have small but obvious and developing eggs. Johari et al. (2009) found the reproduction period began in March and lasted until the latter part of May based on the gonadosomatic index. Patimar and Mohammadzadeh (2011) found a sex ratio of 1:2.42 in favour of females for their south Khorasan fish, with reproduction in the qanat between May and August with the gonadosomatic index highest for males in June and for females in July. egg diameters attained 2.05 mm, maximum fecundity attained 22,773 eggs and relative fecundity up to 583 eggs/g.

Parasites and predators

Black spots on the head and fins (syntypes of nudiventris as noted by Nikol'skii (1897)) were probably encysted larvae of trematodes (Berg, 1949). Johari et al. (2009) found the trematode Clinostomum in various body parts and their qanat fishes showed lordosis and scoliosis.

Economic importance

This species has been studied in aquaria for the toxicity of lead acetate (Omidi et al., 2009). Toxicity decreased with increase in water hardness, qanat water with a high water hardness (310 mg L-1) showing low toxicity. It will feed on mosquito larvae under aquarium conditions and could have been a better candidate for combating malarial mosquitos than the exotic and deleterious Gambusia holbrooki.

Conservation

A widely distributed species apparently able to survive in a wide range of desert habitats, it may not be in need of conservation.

Further work

Biology in mostly unknown and would help confirm the impression that it is not in need of conservation.

Sources

Type material: See above, Capoeta fusca (ZISP 11108) and Capoeta nudiventris (ZISP 11105, 11106).

Iranian material: CMNFI 2007-0005, 7, 27.8-84.2 mm standard length, Semnan, spring at Nardin (ca. 37º03'N, ca. 55º47'E); check ID? CMNFI 2007-0015, 8, 60.1-85.6 mm standard length, Khorasan, qanat at Khalaj (ca. 34º54'N, ca. 58º52'E; CMNFI 2007-0016, 8, 85.5-171.4 mm standard length, Khorasan, qanat and jube at Bidokht (ca. 34º21'N, ca. 58º46'E); CMNFI 2007-0017, ?, ? mm standard length, Khorasan, qanat at Dasht-e Bayaz (ca. 34º02'N, ca. 58º47'E); CMNFI 2007-0018, 15, 21.7-92.4 mm standard length, Khorasan, Shur River (ca. 33º52'N, ca. 59º41'E); CMNFI 2007-0019, 9, 32.7-141.3 mm standard length, Khorasan, qanat between Esfideh and Abbasabad (ca. 33º29-39'N, ca. 59º38-46'E); CMNFI 2007-0020, 23, 43.7-115.1 mm standard length, Khorasan, qanats at Marak and Rabi'an (ca. 32º55-58'N, ca. 59º26-27'E); CMNFI 2007-0021, 16, 24.8-56.3 mm standard length, Khorasan, Shah Abbas qanat in Asadabad (32º55'N, 60º01'E); CMNFI 2007-0022, 6, 56.7-112.1 mm standard length, Khorasan, qanat pool at Mud-e Dahanab (32º43'N, 59º31'E); CMNFI 2007-0023, 6, 82.5-113.1 mm standard length, Khorasan, qanat at Sarbisheh (32º34'N, 59º48'E); BM(NH) 1958.11.7:1-6, 6, 26.1-90.9 mm standard length, Khorasan, near Jajarm (no other locality data).

Capoeta trutta
(Heckel, 1843)

Common names

tu'ini (and variant spellings in transliteration such as touyeni, tuyeni, tuini or too'ini) in Khuzestan (meaning unknown); tu'ini gelkhorak in Khuzestan (see C. damascina for meaning); shir mahi (= milk fish), barg bidy or barg-e bidi (= willow leaf, perhaps from shape and colour), berzem.

[twena, hemira, tela morqat, tela moraqqat; ethra at Mosul (Heckel (1843b), or takal handscherli (takal = soft or flexible presumably from its small scales, handscherli = armed with a dagger or knife from the dorsal fin spine) at Aleppo (Heckel, 1843b), all in Arabic; trout barb].

Systematics

Rainboth (1981) places this species in Schizocypris on the basis of enlarged scales forming a split to encompass the urogenital region and a bare to partially bare mid-dorsal strip anterior to the dorsal fin. However the schizothoracine fishes are quite different (see accounts for Schizothorax, Schizopygopsis and Schizocypris) and this placement is not accepted here.

The type localities of Capoeta Trutta as given by Heckel (1843b) are "Gewässern bei Aleppo" and the "Tigris bei Mossul". The syntypes are in the Naturhistorisches Museum Wien according to Krupp (1985c) as follows: NMW 55935-37, 55942, 6 specimens 94-274 mm standard length from Mosul, NMW 55926, 55928, 55940-41, 7, 68-192 mm standard length from Aleppo, and in the Senckenberg Museum Frankfurt (SMF 2567 (formerly NMW), 1, 407 mm standard length, from Mosul and SMF 923 (formerly NMW), 1, 175 mm standard length, from Aleppo. Four other syntypes are under NMW 55939, 1 other syntype under NMW 55938 and a dried syntype under NMW 58875. Eschmeyer et al. (1996) lists similar material with the numbers of fish under each catalogue number detailed thus: NMW 55926 (1), NMW 55928 (2), NMW 55935-37 (2, 2, 1), NMW 55939-42 (4, 1, 3, 1), possibly 1 fish in the Rijksmuseum van Natuurlijke Historie, Leiden (RMNH 3164, formerly NMW), 1 syntype in the Senckenberg Museum Frankfurt (SMF 923, formerly NMW) and 1 syntype SMF 2567 (formerly NMW), and 1 dried syntype from the Museum für Naturkunde, Universität Humboldt, Berlin (ZMB 8789; not located in February 2006). The catalogue in Vienna lists only 5 specimens although the card index in 1997 lists NMW fish as syntypes in agreement with Eschmeyer et al. (1996).

Key characters

The combination of small scales, transverse mouth, dorsal and anal fin branched ray counts, the very strong last unbranched dorsal fin ray (longer than head length - usually strong but rarely weak), and the colour pattern identifies this species.

Morphology

Dorsal fin with 3-5 unbranched rays followed by 7-9, usually 8, branched rays, anal fin with 2-3 unbranched rays followed by 5 branched rays, pectoral fin branched rays 14-18, and branched pelvic fin rays 5. Hanel et al. (1992) found 23-31 denticles or teeth on the serrated dorsal fin ray, the largest near the centre of the ray length. Scales in lateral line 68-90, scales above lateral line 15-18 and scales below lateral line 10-17. The back anterior to the dorsal fin is compressed and lacks scales except near the occiput. Scales have a protruding anterior margin but are otherwise rounded, anterior and posterior radii, fine circuli and a subcentral anterior focus. There is a pelvic axillary scale. Gill rakers 23-33, on the lower arm 18-25 (with lowest counts in smallest fish). The rakers reach the second raker below when appressed. Pharyngeal teeth 2,3,4-4,3,2. Teeth are broadly spoon-shaped at the tip, with narrow cusps and stems such that they are quite fragile. A frequency distribution of counts was not taken because of this fragility. Total vertebrae 43-46. The mouth is inferior and transverse with a strong horny cover to the lower jaw. The gut is very elongate with numerous anterior and posterior loops. The karyotype of fish from the Tigris River of Turkey is 2n=150, possibly hexaploid, with 35 meta-submetacentric chromosomes, 40 pairs of subtelo-acrocentric chromosomes with NF=220 (Kılıç Demirok and Ünlü, 2001).

Meristics for Iranian specimens:- branched dorsal fin rays 8(34); branched anal fin rays 5(34); branched pectoral fin rays 14(1), 15(8), 16(18), 17(6) or 18(1); branched pelvic fin rays 5(34); lateral line scales 68(2), 69(1), 70(1), 71(4), 72(5), 73(3), 74(5), 75(2), 76(3), 78(3), 79(1), 80(1), 81(1), 83(1) or 84(1); total gill rakers 22(1), 24(5), 25(4), 26(3), 27(7), 28(8), 29(3), 30(2) or 31(1); and total vertebrae ?more 43(1), 44(6), 45(3) or 46(2) - NMC 79-269, 367, 384, 269, 268, Behnke 231 done.

Sexual dimorphism

Males bear a single tubercle on each flank scale, sometimes 2 tubercles, positioned about the middle of the exposed scale or nearer the posterior edge. The head has small and widely scattered tubercles on the top and sides and large tubercles around the snout from eye to eye below the nostril level. Large tubercles occur in single files on the anal and dorsal fin rays, particularly the posterior rays, becoming apparent on the more anterior rays as tuberculation develops more highly.

Colour

The head and body and the dorsal fin (and sometimes the caudal fin) are covered with small, distinctive black spots, often c- or x-shaped. Spots are apparent through the silver flank colour. Some fish in Khuzestan lack spots but transitional specimens from fully spotted through weakly spotted to immaculate are found. Colour is brownish to yellowish or olive-green on the back with silvery-white flanks and the belly lighter, white with silvery tints. Some fish are very pale almost whitish. Upper flank scales in particular are outlined with dark pigment. The eye is orange above or mostly silvery. Lower fins are orange to yellow at the base and blackish distally, or may be orange to yellow overall. The dorsal and caudal fins are grey or hyaline. The lower rays of the caudal fin have a slight orange-yellow tint. The peritoneum is dark brown to black.

Size

Attains at least 45.8 cm total length. Heckel (1843b) gives 1 Schuh 8 Zoll, or 52.7 cm.

Distribution

Found in the Quwayq, Orontes and Tigris-Euphrates basins including the Iranian portion of the latter (Berg, 1949; Marammazi, 1995) and the Gulf basin in the Zohreh River.

upper Mand ? to check on maps

Zoogeography

Its relationships with other Capoeta species is generally unclear, as are the larger zoogeographical relationships of this and other Capoeta species. See also above under genus.

Habitat

Marammazi (1994) considers this species to be stenohaline but nonetheless more widely distributed than stenohaline Barbus (= Mesopotamichthys) sharpeyi in the Zohreh River which drains to the northern Persian Gulf.

Age and growth

The majority of the population studied by Ünlü (1991) in the Tigris River in Turkey are in age groups 2 and 3 although males live to age 7 and females age 10. Females are usually longer and heavier than males of the same age. Males comprise 41.26% and females 58.74% of this population. In a stream in the Euphrates River drainage of Turkey, Gul et al. (1996) found fish to live for 8 years with 60-90% of the fish in age groups 1 to 3. Females comprised 53.3% and males 46.7% of the population. Kalkan (2008) studied a population in the Karakaya Dam lake on the Turkish Euphrates River. Maximum age was 7 years, age groups 4 and 6 were mostly females whereas age group 3 was mostly male, age-length, age-weight and length-weight formulae were given, and the average growth condition factor was 1.30 for females and 1.28 for males.

Food

Gut contents include diatoms, green algae and large amounts of sand.

Reproduction

Spawning in both the Tigris and Euphrates rivers in Turkey took place in May-June. Males mature at age 2 and females at age 3 in both rivers. Ripe egg size in the Tigris varied between 1.33 and 2.11 mm and egg numbers between 4713 and 18240. Ripe eggs in the Euphrates attained 1.04 mm and the maximum number of eggs per gramme of gonads was 666. Fish from Khuzestan had well-developed eggs on 30 January while adult fish taken on 7 July were not in reproductive condition.

Parasites and predators

Molnár and Jalali (1992) report the monogenean Dactylogyrus pulcher from this species in the Dez River of Khuzestan. Gussev et al. (1993a) describe a new species of monogenean from this species in the Dez River, Dactylogyrus microcirrus. Baska and Masoumian (1996) describe two new species of Myxosporea from fish caught in the Karun River at Ahvaz, Myxobolus molnari taken from the gills and Myxobolus mokhayeri taken from between the soft rays of the fins. The latter species is named after Dr. Baba Mokhayer, an internationally renowned Iranian professor. The new species are of minor pathological importance as the infections are of low intensity and prevalence. Masoumian and Pazooki (1999) list Myxobolus molnari and M. mokhayeri from this species from localities in Khuzestan. Peyghan et al. (2001) record Neoechinorhynchus sp. and Rhabdocona sp. from fish from Khorramabad rivers. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Lernaea sp. and Tracheliastes polycolpus on this species.

Economic importance  

Duman and Duman (1996) give the nutritional value of Capoeta trutta from Keban Dam Lake in Turkey but this fish is little used in Iran. However, Peyghan et al. (2001) report that is is an economically important species with a good market value in the Khorramabad region.

Conservation

This species does not appear in need of conservation but its biology is too poorly known in Iran to be certain. Kalkan (2008) recommended prohibition of fishing in Turkey during March-August and fish under 22.62 cm should not be retained.

Further work

The biology of this species and its relationships to other Capoeta species needs work.

Sources

Type material: ?

Iranian material:- CMNFI 1979-0020, ?, ? mm standard length, (); CMNFI 1979-0268, 3, 115.7-141.2 mm standard length, Lorestan, between Nowqan and Khorramabad (no other locality data); CMNFI 1979-0269, 2, 114.1-144.1 mm standard length, Lorestan, between Nowqan and Khorramabad (no other locality data); CMNFI 1979-0367, 2, 29.7-54.1 mm standard length, Khuzestan, Meymeh River 11 km north of Dehloran (32º44'30"N, 47º09'30"E); CMNFI 1979-0368, 8, 36.3-67.9 mm standard length Khuzestan, Karkheh River (32º24'30"N, 48º09'E); CMNFI 1979-0376, 1, 55.2 mm standard length, Khuzestan, river tributary to Karkheh River (32º48'30"N, 48º04'30"E); CMNFI 1979-0384, 1, 218.4 mm standard length, Khuzestan, Ab-e Shur drainage (32º00'N, 49º07'E); CMNFI 1991-0153, 2, 153.8-217.2 mm standard length, Khuzestan, Zohreh River (no other locality data); CMNFI 1995-0020, ?, ? mm standard length, (); CMNFI 1995-0021, ?, ? mm standard length, (); CMNFI 1995-0030, ?, ? mm standard length, (); CMNFI 2007-0100, 1, 136.7 mm standard length, Azarbayjan-e Gharbi, Kalwi Chay near Piranshahr (ca. 36º44'N, ca. 45º10'E); CMNFI 2007-0109, 11, 61.3-167.4 mm standard length, Kordestan, Qeshlaq River basin north of Sanandaj (ca. 35º33'N, ca. 47º08'E); CMNFI 2007-0110, 3, 96.6-160.3 mm standard length, Kordestan, Yuzidar River basin (ca. 35º05'N, ca. 46º56'E); CMNFI 2007-0113, 1, 74.8 mm standard length, Kermanshahan, Razavar River 35 km northwest of Kermanahah (ca. 34º25'N, ca. 47º01'E); CMNFI 2007-0116, 1, 95.9 mm standard length, Kermanshahan, Gav Masiab west of Sahneh (ca. 34º28'N, ca. 47º36'E); CMNFI 2007-0117, 2, 153.8-217.2 mm standard length, Kermanshahan, Gav Masiab near Sahneh (ca. 34º24'N, ca. 47º40'E); ZMH 2511, 1, 319.0 mm standard length, Kermanshahan, Karasu-Gamasiab-Seymarreh (no other locality data).

Comparative material: BM(NH) 1931.12.21:8, 1, 113.5 mm standard length, Iraq, Mosul (36º20'N, 43º08'E); BM(NH) 1968.12.13:376-390, 15, 35.6-123.3 mm standard length, Syria, Euphrates River at Mayadine (?); BM(NH) 1974.2.22:1374-1377, 4, 66.3-91.2 mm standard length, Iraq, Baghdad (33º21'N, 44º25'E); BM(NH) 1974.2.22:1382, 1, 86.1 mm standard length, Iraq, Baghdad (33º21'N, 44º25'E); BM(NH) 1974.2.22:1388-1389, 2, 259.4-273.3 mm standard length, Iraq, Tigris River at Samarra (?).

Genus Capoetobrama
Berg, 1916

Capoetobrama kuschakewitschi
(Kessler, 1872)

This species is reported from the Karakum Canal of Turkmenistan (Shakirova and Sukhanova, 1994; Sal'nikov, 1995) and may eventually be found in the Tedzhen River and Caspian Sea basins of Iran. No Iranian record.

Genus Carasobarbus
Karaman, 1971

?

Some of the past literature on this genus appeared under Barbus (q.v.)

 

Carasobarbus luteus
(Heckel, 1843)

Common names

حمري (= hemri), himri; sangal or zangol (= blackish, used at Kermanshah, J. Valiallahi, pers. comm., 2001); lab matiki (= from lipstick by professional fishermen at Kermanshah in reference to red lips, from J. Valiallahi, pers. comm., 2001).

[himri, hamria, hamra, binni hamour, binni hamri, bunni himri, binni, binni shifatha, beni asphar (= yellow son), beni abjad (= white son), beni hamra (= red or yellow son), zuri or bartema, all in Arabic; golden barb, yellow barbel].

Systematics

Heckel (1843b) gives localities for the types of Systomus luteus as "Orontes", and "Tigris", and in the next sentence at "Aleppo" and "Mossul". Two syntypes were examined in the Naturhistorisches Museum Wien under NMW 54250 (but see below). Krupp (1985c) records a 301 mm standard length syntype from Aleppo formerly in the Naturhistorisches Museum Wien, now in the Senckenberg Museum Frankfurt as SMF 6784. Eschmeyer et al. (1996) list the following syntypes: NMW 10827 (1 fish), NMW 54247 (2), NMW 54248 (1), NMW 54249 (1), NMW 54253 (2), NMW 54254 (3), NMW 54255 (2), NMW 54520 (2), NMW 80043 (2) and possibly 2 syntypes in the Rijksmuseum van Natuurlijke Historie, Leiden (RMNH 2463, formerly NMW) as well as the syntype in Frankfurt. The catalogue in Vienna seems to list 5 specimens but this part of the catalogue is overwritten and difficult to interpret. The card index in 1997 lists NMW 53680a (1 fish, the lectotype), 53674 (1), 53675 (1) and 53676 (1) as the syntype series.

Systomus albus Heckel, 1843 from the "Tigris" and "Orontes" and Systomus albus var. alpina Heckel, 1849 are synonyms.

Systomus albus var. alpina was described from the "Flusse Kara-Agatsch und den Alpenseen Pire-San und Deria Kaserun" (= Qarah Aqaj River and Lake Famur, Fars; Pire-San being Parishan and Deria Kaserun being Lake Kazerun, both other names for Lake Famur) (Heckel, 1846-1849b). Krupp (1985c) records 4 syntypes of alpina from Shiraz (sic), Th. Kotschy as NMW 53679 (2 fish) and NMW 53681 (2). NMW 53678 (5 fish, 27.6-60.8 mm standard length), NMW 53679 (2 fish, 63.8-70.5 mm standard length), and NMW 53681 (2 fish, 79.6-93.3 mm standard length) are from the "Kara Agatsch bei Schiraz"; and NMW 53682 (2 fish, 201.7-203.7 mm standard length) are from the "Alpenseen Pire-san und Deria Kaserun": all are possibly syntypes of Systomus albus var. alpina although the catalogue in Vienna lists 5 fish under this name in one column and 4 fish in smaller writing in the adjacent column. The card index in 1997 lists syntypes under NMW 53678 (5), 53679 (2), 53681 (2) and 53682 (2, one of which is the lectotype). Eschmeyer et al. (1996) list 2 fish in the Rijksmuseum van Natuurlijke Historie, Leiden (RMNH 2464) as possible former NMW types of this taxon.

A dried specimen of Systomus albus from Mosul collected by Th. Kotschy may be a syntype (NMW 59485). Eschmeyer et al. (1996) gives the syntypes of this species as NMW 53674 (1), NMW 53675 (1), NMW 53676 (1), NMW 53677 (1), NMW 53680 (1), NMW 91400 (1, dry) and SMF 812 (1), formerly NMW. Krupp (1985c) records the syntype of albus in the Senckenberg Museum Frankfurt under SMF 812 as being 84 mm standard length. The Vienna catalogue lists 4 fish under Systomus albus but the card index in 1997 lists the same NMW fish as Eschmeyer et al. (1996) as above with NMW 53680 as lectotype.

Barbus parieschanica Wossughi, Khoshzahmat and Etemadfar, 1982 is presumably also from Lake Famur or Parishan judging by the name and is a synonym (note that the species name is first spelt parschanica on page 23 in the abstract in Farsi and on page 44 in the English abstract but in the text species description (page 34) and in the table (page 37) it appears as parieschanica, and this is presumably the intended correct spelling). The species locality in the text is "Noorabad of Mamasany". ?

Saadati (1977) refers to a new and undescribed Cyprinion species from Lar in southern Iran but the fish are Carasobarbus luteus.

Günther (1874) placed this species in Barynotus Günther, 1868, a genus with the type species from West Africa. Barynotus is preoccupied in Coleoptera and was replaced by Barbellion Whitley, 1931 (Eschmeyer, 1990). Most authors place the species in Barbus although Karaman (1971) erected a new genus for it, Carasobarbus; and Krupp (1985c) also synonymises Carasobarbus with Barbus. Bănărescu (1997) and Ekmekçi and Banarescu (1998) recognise Carasobarbus as a valid genus however. Borkenhagen et al. (2011) recognise C. luteus as a single, generalist species tolerating a wide variety of habitats.

A group of related species share characters with this species (see also under Kosswigobarbus kosswigi). Carasobarbus may be the generic names for certain members of the group.

Key characters

This species is characterised by a low scale count, smooth last unbranched dorsal fin ray, one or two pairs of barbels, and 10 branched dorsal and 6 branched anal fin rays.

Morphology

Dorsal fin with 4 unbranched rays followed by 9-11, usually and modally 10, branched rays. The last unbranched dorsal fin ray is smooth, thickened, sharp-edged and spine-like. Anal fin with 3 unbranched rays followed by 5-7, usually and modally 6, branched rays. Pectoral fin branched rays 13-17 and pelvic fin branched rays 7-9, usually 8. Lateral line scales 23-36. There is a pelvic axillary scale. There are moderate to many anterior field radii and many posterior field radii and occasionally few lateral radii. The focus is central to subcentral anterior, the anterior scale margin is wavy and the exposed part of the scale is coarse. The concealed part of the scale has numerous fine circuli. Total gill rakers 7-14, reaching the adjacent raker when appressed, sometimes forked at the tip and with spinules on the anterior side. Pharyngeal teeth usually 2,3,5-5,3,2, with the anterior 2-3 teeth rounded and heavier than the posterior teeth. Variants may have 2,3,4 or 1,3,5 (Borkenhagen, 2005). Posterior teeth are hooked at the tip and the grinding surface below the tip is irregular with a protuberant knob which may be striated. The gut is elongate with both posterior and anterior loops.

The mouth is terminal to subterminal and lips are weakly developed. There is one pair of short and thin barbels at the corner in most descriptions. Number and frequencies for 130 fish are 2 barbels (47 or 36.2%), 3 barbels with left anterior present (7 or 5.4%), 3 barbels with right anterior present (5 or 3.8%), or 4 barbels (71 or 54.6%). However, this sample is 112 fish or 86.2% from Fars and Hormozgan. Fish from these provinces, at such localities as the lower Mand River and the Sar Khun oasis north of Bandar Abbas consistently have a high frequency of 4 barbels (58.9%), and with 3 barbel counts included 68.8%, than fish from the Tigris River basin. Even the 18 fish from the Tigris River basin had 5 fish with 4 barbels so, at least in the eastern part of this species range, 4-barbelled fish are not rare.

Body form varies with habitat (Ali, 1982a), there being lake and river forms as with many other cyprinid species.

Iranian specimens have the following meristics: dorsal fin branched rays 9(7), 10(102) or 11(7); anal fin branched rays 5(3) or 6(114); pectoral fin branched rays 14(12), 15(44), 16(48) or 17(13); pelvic fin branched rays 7(9), 8(107) or 9(1); lateral line scales 23(2), 24(10), 25(20), 26(22), 27(28), 28(16), 29(14), 30(4) or 31(1); total gill rakers 8(6), 9(24), 10(40), 11(28), 12(12), 13(3) or 14(2); pharyngeal teeth 2,3,5-5,3,2(19), 2,3,4-5,3,2(4) or 2,3,5-4,3,2(2); and total vertebrae 36(8), 37(53), 38(70), 39(25) or 40(1).

Sexual dimorphism

A 12.7 cm specimen from the Mand River has tubercles on the dorsal, anal, caudal, pectoral and pelvic fins, most strongly on the anal fin rays. Fine tubercles cover the top and sides of the head. A 20 cm fish from the same collection lacked tubercles. Another fish from the lower Mand River (128.5 mm standard length) also has fine tubercles on the upper flank scales as well as the head and fin rays. Ali (1982) reports no sexual dimorphism for Iraqi fish.

Colour

The back and upper flank is dark brown, greenish black or grey-green fading to a whitish or silvery belly all overlain by an orange to yellowish tinge. On the upper flank, scale bases are black-brown with a light blue-grey margin. There is a dark stripe along the mid-line of the back and a dark mid-lateral stripe. Fins are greyish to lime-green, reddish-yellow or orange, becoming blackish distally. The pectoral and pelvic fins tend to be more orange than the anal and caudal fins which are more a faint lime-green. The lips are orange. The eye rim is yellow-green. The peritoneum is black. Small fish have a collection of melanophores at the mid-base of the caudal fin forming a spot-like structure.

The fish described by Heckel (1846-1849b) as Systomus albus var. alpina were also painted live and had a lead-grey body, light brown at the head and reddish-white on the belly. Each scale was black-brown at the base and light blue-grey at the margin, particularly on the upper flank. All fins were blackish and the eyes orange-red.

Size

Attains 38 cm calculated maximum length and 501 g (Ahmed, 1982) or 750 g (Borkenhagen, 2005). Heckel (1843b) gives 17 Zoll for Systomus albus (= 44.8 cm).

Distribution

This species is found in the Orontes and Quwayq rivers and the Tigris-Euphrates basin. In Iran, it is found in the Tigris River basin including the Hawr Al Azim marsh, the Gulf basin including the Helleh, Dalaki, Shapur, Mand and Dasht-e Palang rivers and Lake Famur, the Lake Maharlu basin, the Hormuz basin and the Kor River basin (Wossughi, 1978; Bianco and Banarescu, 1982; Gh. Izadpanahi, pers. comm., 1995; M. Rabbaniha, pers. comm., 1995; Abdoli, 2000). The record from the Kor River basin (Abdoli, 2000) needs confirmation with specimens.

Zoogeography

Karaman (1971) considers that the closest relatives of this species were to be found in India and southern Asia.

Habitat

van den Eelaart (1954) reports that this species in Iraq is a resident in still water and the slower sections of rivers and is the main fish in canals. In summer it goes to the deeper basins of marshes and remains in the shade of plants. It tolerates warm water but does not go into open waters. Al-Hassan and Muhsin (1986) record this species from the Khor al Zubair in southern Iraq where annual temperature range is 12-30°C and annual salinity change is 28-47‰. The fish appear unaffected by these conditions while Heteropneustes fossilis is moribund. Mohamed et al. (1993) report Barbus (= Carasobarbus) luteus from 2 km southward of Fao, Iraq in a pure marine habitat (temperature 13-35°C and salinity 30-47‰). The fish were caught in April which is the flood season.

Age and growth

Ahmed et al. (1984) studied the reproductive cycle of this species in the Hawr al Hammar in southern Iraq near Basrah. Maturity is attained at a minimum of 11.2 cm for females and 12.2 cm for males, at age 1+. The largest fish are 26.0 cm and age 6. Barak and Mohamed (1983) also found 6 age groups for fish from the Garma Marshes, Iraq. Ahmed (1982) studied a population in Tharthar Reservoir about 65 km northwest of Baghdad and found 7 age groups. This study has the fastest growth of Iraqi populations. Khalaf et al. (1988) worked on a population in a flooded gravel pit about 50 km north of Baghdad in Iraq and found fish up to age group 7+. Growth is greatest in the first year (67 mm) and averaged only 22.5 mm in the following years. Growth is slow in consequence of high salinity (3-6% (sic)) and poor food resources. Mohamed et al. (1993) report fish up to 7 years of age in a marine setting in Iraq, Epler et al. (1996) up to 5+ years in fresh and salty Iraqi lakes. Biro et al. (1988) found fish up to age group 8+ in the Diyala River, Iraq. Al Hazzaa and Hussein (2007) describe larval development and growth in the laboratory using fish from a Syrian hatchery. Gökçek and Akyurt (2008) found fish up to 9 years of age in the Turkish Orontes River and give growth parameters for this population. Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 34 Iranian fish measuring 3.20-16.80 cm standard length. The a-value was 0.0232 and the b-value 3.036 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases).

Food

Naama and Muhsen (1986) examined feeding periodicities in this species in the Hawr al Hammar, Iraq. Food is mainly detritus, aquatic plants and algae taken throughout the night and day. Barak and Mohamed (1982) studied food habits in the Garma Marshes, near Basrah, Iraq and found this fish to contain principally aquatic plants, the broken and fragmented leaves and stems of Vallisneria in particular. Diatoms and other algae as well as shrimps, chironomid larvae, gastropods and cladocerans are important foods. Invertebrates are about eight times more important in fish smaller than 30 cm than in larger fish. Plant parts are more important, almost twice as much, in larger fish than smaller. Mohamed et al. (1993) report plant remains to be dominant and fish eggs in lesser quantities in a marine setting in Iraq. Epler et al. (1996) found plants to dominate in fish from fresh and salty Iraqi lakes, although not to the same extent as in Barbus (= Mesopotamichthys) sharpeyi where 95.7-100% of the diet was plants. Tendipedids, worms, detritus and fish were also found in B. luteus.

Khoshzahmat et al. (1981) found that this species did not eat molluscs in Lake Perishan (= Famur), near Kazerun in Iran and assume its diet is aquatic plants.

Reproduction

Spawning in the Hawr al Hammar starts in April and after July no fish are found in a partially spent phase. Eggs are yellow to orange in colour and testes white. The eggs attain 1.86 mm in diameter and number up to 38,433 for the oldest fish. Bhatti and Al-Daham (1978) and Al-Daham and Bhatti (1979) report a spawning season of May-July (peak June-July) for a lower Euphrates River, Iraq population, perhaps as a result of cooler temperatures outside the shallow marshes where warmer temperatures cause an earlier development of gonads. Epler et al. (1996) report spawning in June/July in freshwater Iraqi lakes, earlier in a saline lake. Iranian fish have well-developed eggs in May.

Parasites and predators

Bykhovski (1949) reports a new species of monogenetic trematode, Dactylogyrus persis, from this species in the Karkheh River, Iran. Ebrahimzadeh and Nabawi (1975) list species in the nematode genus Philometra, the protozoan genera Myxosoma and Trypanosoma, the trematode genera Dactylogyrus and Gyrodactylus and the nematode species Camallanus lacustris as well as various unidentified cestodes, trematodes, acanthocephalans and hookworms, from this species in the Karun River. Jalali and Molnár (1990a) records two monogenean species, Dactylogyrus spp., from this species in the Dez River. Molnár and Jalali (1992) describe a new species of monogenean, Dogielius persicus, from this species in the Dez and Karun rivers of Khuzestan. Gussev et al. (1993b) describe a new species, Dactylogyrus carassobarbi, from this species in the Dez River, Khuzestan, the specific name being founded on a misspelling of the genus name Carasobarbus. Masoumian et al. (1994) describe a new species of Myxosporea from the gills of this species in the Karun River, Khuzestan, namely Myxobolus persicus, and later (Masoumian et al., 1996) another new species of Myxosporea, Myxobolus nodulointestinalis, in the gut lining of this species and also from rivers of southwestern Iran. Molnár et al. (1996) report additional new species from this fish in Khuzestan, namely Myxobolus iranicus in the spleen and Myxobolus mesopotamiae in connective tissue of the caudal and pectoral fins. Molnár and Pazooki (1995) record philometrid nematodes from this species in the Karun River, and these are presumed to be a new species.

Masoumian and Pazooki (1999) list Myxobolus persicus, M. karuni, M. sharpeyi, M. nodulointestinalis, M. mesopotamiae and M. iranicus from this species in various localities in Khuzestan. Jalali et al. (2005) summarise the occurrence of Gyrodactylus species in Iran and record G. sp. from Dez River fish. Farahnak et al. (2002) record Anisakis sp. from this fish in Khuzestan Province.

González-Solís et al. (1997) report Proleptinae larvae (Nematoda) from this species in the drainage of Lake Maharlu, Fars. The definitive host is a predatory fish, possibly Mastacembelus mastacembelus, not yet recorded from this basin.

Moghainemi and Abbasi (1992) record a wide range of parasites from this species in the Hawr al-Azim in Khuzestan. Mortazaei et al. (2000) record an infection rate of 1.6% with the worm Bothriocephalus opsariichthydis in Khuzestan marshes. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Argulus sp., Ergasilus sp., Ergasilus sieboldi and Lernaea sp. on this species.

Economic importance

An important food fish in southern Iraq and Iran (Al-Daham and Bhatti, 1979; Ahmed, 1982). Sharma (1980) reports that hamri were the fourth most important fish species at Basrah fish market, accounting for 267,570 kg from October 1975 to June 1977. Heckel (1846-1849b) reports that they "reach a good size and are very tasty" in Lake Famur, Fars.

In some parts of Southwest Asia this species is regarded as &qut;sacred" kept and bred in special pools where fishing is forbidden (Tortonese, 1934).

The eggs of this species are poisonous (Najafpour and Coad, 2002). A kebab made of about one-quarter of an ovary was eaten. Toxic effects were dizziness, abdominal pain, vomiting, diarrhoea, bitter taste, dryness of mouth, intense thirst, and faintness. One victim as hospitalised for two days and his stomach pumped while a second victim recovered after one day's rest.

Conservation

Vulnerable in Turkey (Fricke et al., 2007). It is a common species in Iranian freshwaters but no detailed conservation assessment has been made.

Further work

The biology of this species in Iran needs study along with its conservation status. Specimens from Fars show differences in body form from those in Khuzestan and this could be investigated.

Sources

Type material: ?

Iranian material: CMNFI 1979-0023, 17, 58.3-161.4 mm standard length, Fars, neighbourhood of Shiraz (no other locality data); CMNFI 1979-0024, 1, 61.5 mm standard length, Fars, neighbourhood of Shiraz (no other locality data); CMNFI 1979-0026, 2, ? mm standard length, Fars, Shapur River (29º47'N, 51º35'E); CMNFI 1979-0047, 1, ? mm standard length, Fars, Ab-e Paravan (ca. 29º34'N, ca. 52º42'E); CMNFI 1979-0076, 1, ? mm standard length, Fars, Barm-e Shur (29º28'N, 52º41'30"E); CMNFI 1979-0087, 1, ? mm standard length, Khuzestan, Karun River at Ahvaz (31º19'N, 48º42'E); CMNFI 1979-0125, 1, ? mm standard length, Bushehr, Dalaki River near Dalaki (ca. 29º28'N, ca. 51º21'E); CMNFI 1979-0129, 26, ? mm standard length, Fars, spring about 2 km from Farrashband (28º54'N, 52º04'E); CMNFI 1979-0135, 19, ? mm standard length, Fars, Mand River tributary (28º08'N, 53º10'E); CMNFI 1979-0154B, 3, 160.7-258.6 mm standard length, Fars, stream channels at Koorsiah (28º45'30"N, 54º24'E); CMNFI 1979-0155, 2, ? mm standard length, Fars, spring at Gavanoo (28º47'N, 54º22'E); CMNFI 1979-0156, 6, ? mm standard length, Fars, qanat at Rashidabad (28º47'N, 54º18'E); CMNFI 1979-0157, 1, ? mm standard length, Fars, qanat at Hadiabad (28º52'N, 54º13'E); CMNFI 1979-0160, 2, ? mm standard length, Fars, spring at Arteshkkadeh Pomp (29º09'N, 53º37'E); CMNFI 1979-0163, 1, 84.9 mm standard length, Fars, neighbourhood of Shiraz (no other locality data); CMNFI 1979-0164, 6, 56.6-91.1 mm standard length, Fars, neighbourhood of Shiraz (no other locality data); CMNFI 1979-0187, 31, ? mm standard length, Hormozgan, stream and pools at Sar Khun oasis (27º23'30"N, 56º26'E); CMNFI 1979-0206, 3, 24.4-25.1 mm standard length, Fars, qanat near Runiz-e Pa'in (29º12'N, 53º40'E); CMNFI 1979-0240, 3, ? mm standard length, Fars, Parishan Lake (ca. 29º31'N, ca. 51º50'E); CMNFI 1979-0304, 5, ? mm standard length, Fars, Parishan Lake (ca. 29º31'N, ca. 51º50'E); CMNFI 1979-0347, 2, ? mm standard length, Fars, Pol-e Berengie (29º27'30"N, 52º32'E); CMNFI 1979-0352, 7, ? mm standard length, Khuzestan, marsh in Jarrahi River drainage (30º33'30"N, 48º48'E); CMNFI 1979-0358, 1, 23.7 mm standard length, Khuzestan, pond southeast of Bostan (31º37'N, 48º07'E); CMNFI 1979-0360, 8, ? mm standard length, Khuzestan, canal branch of Karkheh River (31º40'N, 48º35'E); CMNFI 1979-0364, 6, ? mm standard length, Khuzestan, river at Abdolkhan (31º52'30"N, 48º20'30"E); CMNFI 1979-0371, 7, ? mm standard length, Khuzestan, stream in Karkheh River drainage (32º05'N, 48º19'E); CMNFI 1979-0687, 7, 124.8-154.1 mm standard length, Fars, Shiraz bazar (no other locality data); CMNFI 1979-0789, 4, ? mm standard length, Fars, Lake Parishan (29º31'N, 51º48'E); CMNFI 1991-0154, 1, ? mm standard length, Khuzestan, Hawr al Azim (ca. 31º45'N, ca. 47º55'E); CMNFI 1993-0126, 1, ? mm standard length, Kermanshahan, Sarab-e Yavari (34º28'N, 46º56'E); CMNFI 1993-0127, 1, ? mm standard length, Kermanshahan, Sarab-e Maran (34º44'N, 46º51'E); CMNFI 2007-0060, 2, ? mm standard length, Fars, Chashmeh Ab-e Shirin near Lar (ca. 27º41'N, ca. 54º17'E); CMNFI 2007-0111, 1, ? mm standard length, Kermanshahan, Alvand River near Sar-e Pol-e Zahab (ca. 34º36'N, ca. 45º56'E); ZSM 21861, 5, 172.0-217.2 mm standard length, Khuzestan, Dez River at Harmaleh (31º57'N, 48º34'E).

Comparative material: CMNFI 1987-0017, 3, 97.3-143.9 mm standard length, (); BM(NH) 1934.9.5:6, 1, 117.3 mm standard length, Kurdistan, Ain al Hamra, Shithatha (); BM(NH) 1973.6.21:194, 1, 203.4 mm standard length, Iraq, Shatt al Arab (); BM(NH) 1974.2.22:1338, 1, 134.9 mm standard length, Iraq, Najab Bazar (); BM(NH) 1974.2.22:1346, 1, 108.7 mm standard length, Iraq, Tigris River near Faish Khabour (); BM(NH) 1986.2.14:4-7, 4, 98.6-146.6 mm standard length, Iraq, Baghdad (33º21'N, 44º25'E).

Genus Carassius
Nilsson, 1832

The goldfishes comprise 2-3 species found in Europe, northern Asia and the Far East. Eschmeyer (1990) and Kottelat (1997) comment on the authorship of Carassius. One species is now common in Iran.

These fishes are characterised by a stout and compressed body, last unbranched dorsal and anal fin rays finely serrated, long dorsal and short anal fin, mouth small and terminal, lips thick and fleshy, no barbels, pharyngeal teeth in 1 row and molariform but compressed, numerous gill rakers, and scales large.

Carassius auratus
(Linnaeus, 1758)

Carassius auratus from Wikimedia Commons.

Carassius auratus from British Fresh-Water Fishes by Rev. William Houghton (1879).

Common names

mahi-ye talaee or mahi-ye talai (= gold fish) or اوشين (ooshin or oushein) in Khuzestan; kapur safid by anglers in Khuzestan at Ahvaz; kopur-cheh (= small carp) or كاراس (= karas, karass or karaz) in Mazandaran; kopur cheky (= by the job carp?), kopur chekeh (= drop carp?); ماهي حوض (= mahi-ye howz or mahi-e-hoz, meaning pond or pool fish), mahi-ye howz-e noqrehi (= silvery pond fish, for silvery form), mahi-ye howz-e talaee (= golden pond fish for orange form).

[samak zahabi, buj-buj in Nasiriyah; samti; yayabash in Basrah; karseen in Baghdad; carp thahabi, all in Iraqi Arabic; serebryanyi karas or silver crucian carp in Russian; goldfish for auratus, Prussian carp for gibelio].

Systematics

Cyprinus auratus was originally described from China and Japanese rivers.

Pelz (1987) discusses the scientific name of the goldfish and its confusion with Carassius carassius. All diploid goldfish of western Europe are Carassius auratus auratus (from introductions, presumably including releases and escapes in Iran) and all triploid goldfish are C. auratus gibelio from eastern areas. Goldfish do not appear to be native to Iran but Iranian specimens are sometimes referred to Carassius auratus gibelio (Bloch, 1782) known as the Prussian carp, European goldfish or silver crucian carp. Berg (1948-1949) considers the familiar pet "goldfish" to be a domesticated form of the Prussian carp. However these fish probably have a number of origins - from aquarium stock and from China. Kottelat (1997) tentatively recognises Carassius gibelio (Bloch, 1782) as a species native to eastern Central Europe, and Kottelat and Freyhof (2007) map gibelio as the introduced species in the Caspian Sea basin of Iran. Vasil'eva and Vasil'ev (2000) state that fish named in the literature as Carassius auratus gibelio from Europe, Siberia and eastern Asia are triploids and are not a valid subspecies of C. auratus s.s. They consider C. gibelio to be a distinct species as long as it has a unique and ancient origin rather than arising de novo, and as long as the type specimens are triploids. Szczerbowski in Bănărescu and Paepke (2002) recognises C. a. auratus and C. a. gibelio.

Additionally C. auratus may be a tetraploid derivative of Carassius carassius. The native distribution of C. carassius is in Europe and western Asia, reaching northern drainages of the Caspian Sea in the southern limits of its distribution (Libosvárský, 1962). It differs from C. auratus in having a slightly convex margin to the dorsal fin (straight or slightly concave in C. auratus), caudal fin slightly emarginate (deeply emarginate), usually 6 branched anal rays (always 5), 23-33 gill rakers (37-53), 31-34 vertebrae, usually 32-33 (28-31, usually 29-30), 28-29 fin denticles posteriorly on the dorsal fin spine (10-11), peritoneum light (dark), black spot at the caudal fin base in young and some adults (absent), and a coppery gold body (silvery, pinkish gold, gold or red) (Szczerbowski in Bănărescu and Paepke, 2002). Berg (1948-1949) also cites the characters body rounded, back thick (body angular, back compressed) and scales weakly sculptured (rough), although his comparison is with C. a. gibelio.

Goldfish commonly hybridise with Cyprinus carpio to further confuse the identity of these fishes (L. Nico, http://nas.er.usgs.gov/fishes/accounts/cyprinid/ca_aurat.html, downloaded 24 May 2000). The identity of "goldfish" in Iran has not been thoroughly surveyed and, along with conflicting views on species and widespread introductions from many sources, make it simpler to refer to this taxon as C. auratus for now.

Al-Mukhtar and Al-Hassan (1999) describe a hybrid of this species and Barbus (= Mesopotamichthys) sharpeyi from Al-Hayei (= Al Ha'i), a seasonal lake between the Karkheh and Dez rivers in Khuzestan.

Key characters

The combination of spines in both the dorsal and anal fins and the absence of barbels is unique to this species. Szczerbowski in Bănărescu and Paepke (2002) distinguishes the subspecies auratus from gibelio by 21-36 lateral line scales (27-35 in gibelio) and a pink or gold colour (yellowish silver), not very diagnostic. Ilhan et al. (2005) give gill raker numbers of 34-40 for auratus, 42-56 for gibelio and 25-32 for C. carassius in Turkish waters (however note below that counts can increase with growth and see also under C. carassius for somewhat different counts and other distinguishing characters).

Morphology

Dorsal fin with 3-4 unbranched rays followed by 12-20 branched rays, anal fin with 2-4, usually 3, unbranched rays followed by 5-6, usually 5, branched rays, pectoral fin branched rays 11-18, and pelvic fin branched rays 6-9, usually 8. Dorsal and anal fin spine denticles coarse and few (about 10-15).

Lateral line scales 21-36. The anterior scale margin is wavy and there are very few anterior and posterior radii, as few as 3-4. The focus is slightly subcentral posterior. Circuli on the exposed part of the scale are more coarse and widely spaced than on the concealed part of the scale. Gill rakers long with serrated interior margins, reaching the fifth to eighth raker below when appressed with younger fish having longer rakers proportionately. Counts are size dependent in the range 34-54. Total vertebrae 25-34. Pharyngeal teeth 4-4, with very elongate, narrow, flattened and horizontal cusps arising from a much narrower stem. The gut is coiled with several loops. This species is variously reported as only diploid or as a tetraploid (2n=100-104); see above.

There are elongate specimens (morpha humilis, where fish density is high) and deep-bodied specimens (morpha vovki, where fish density is low) but these names have no taxonomic significance.

Meristic values for Iranian specimens are:- dorsal fin branched rays 16(4), 17(3), 18(3), 19(5) or 20(2); anal fin branched rays 5(17); pectoral fin branched rays 11(1), 14(1), 15(4), 16(10) or 17(1); pelvic fin branched rays 7(2) or 8(15); lateral line scales 28(6), 29(9) or 30(2); pharyngeal teeth 4-4(10); and total vertebrae 32(2).

Sexual dimorphism

Breeding males have small nuptial tubercles on the operculum, back and pectoral fin rays.

Colour

The golden or orange colour of artificially bred aquarium goldfish is distinctive. However populations in the wild, if they breed successfully, gradually revert to a wild-type of colour, without the appropriate diet supplement of aquarium fish and, as golden fish, are readily seen and eaten by birds and other fishes. Yanar and Tekelioğlu (1999) found that pigmentation increased with fish weight when specimens were fed the carotenoid zeaxanthin. Wild-type colour is an overall olive-green fading to a white belly. Flanks can be silvery to almost black. Fins are a dark olive-bronze, the membranes in particular being heavily pigmented. Young goldfish are usually green, brown or bronze to almost black and only after about 1 year do they take on the colour of adult auratus or gibelio. Peritoneum dusky to black.

Young fish at Ahvaz, Khuzestan, however, are a bright silvery overall (more so than Cyprinus carpio of similar size), the back is grey, the caudal fin is grey on the proximal half and hyaline distally, and the anal fin rays are white (and thus partly resemble gibelio).

Prussian carp (subspecies gibelio) is a dark steel colour with dark blue or greenish dorsally, silver-grey laterally and white ventrally, dorsal and caudal fins are dark grey and the paired fins and anal fin are light pinkish (Szczerbowski in Bănărescu and Paepke (2002).

Size

Attains 62.0 cm and about 5.0 kg, the subspecies gibelio being smaller, up to 45.0 cm and 1.24 kg.

Distribution

The native distribution is in northern Asia and China, reaching northern drainages of the Caspian Sea in the western limits of its distribution (Libosvárský, 1962; Plez, 1987). The goldfish has been widely introduced to garden ponds and released from aquaria in temperate to warm waters world-wide. In Iran it has been introduced throughout the Caspian Sea basin where it is reported from the Atrak, Gorgan, Gharasu, Tajan, Babol, Haraz, Sardab, Aras (including the middle Aras and lower reaches of its tributary the Qareh Chai), Tonekabon, Pol-e Rud, and Safid rivers, the Anzali Mordab where it is now the most abundant fish, Gorgan Bay and Alma- and Ala-Gol (Holčík and Oláh, 1992; Shamsi et al., 1997; Roshan Tabari, 1997; Abbasi et al., 1999; Kiabi et al., 1999; Abdoli, 2000; Gasmi and Mirzaei, 2004; Patimar, 2008; Abdoli and Naderi, 2009); the lower Talkheh and lower Zarrineh rivers in the Orumiyeh basin (Abdoli, 2000); the lower Shur, lower Qareh Chai and the Latian Reservoir in the Namak Lake basin (Armantrout, 1980; Hosseini, 1987; Abdoli, 2000); the Hamun Kushk and the Sistan Dam as well as throughout the hamuns in the Sistan basin (Ahmadi and Wossughi, 1988; Mansoori, 1994; J. Holčík, in litt., 1996; field work in the 1970s); Kerman-Na'in and Dasht-e Lut basins generally (Abdoli, 2000); lower Kashaf River in the Tedzhen basin (Abdoli, 2000); throughout Khuzestan where now common (N. Najafpour and M. Al-Mukhtar, pers. comm., 1995; field work 2000, absent in 1970s); middle and lower Hilleh and lower Mand rivers in the Gulf basin; middle Halil and middle to lower Bampur River (Abdoli, 2000); Dalaki and Shapour rivers (Pazira et al., 2005), Safid River, Zayandeh River, Zarivar Lake and the Hamun Lake (Shamsi et al., 2009), and found in garden and park ponds throughout Iran. Some introductions are probably discarded aquarium fish as goldfish are sold as pets and for the Now Ruz (= New Year) festivities. They may also have been introduced accidentally with the commercially important Chinese carps.

This species is also recorded from the Karakum Canal and Kopetdag Reservoir in Turkmenistan (Shakirova and Sukhanova, 1994; Sal'nikov, 1995) and may eventually reach Iranian waters from this source in the Tedzhen (= Hari) River basin.

The Prussian carp (subspecies gibelio) is less widely distributed and its presence and distribution in Iran are not known.

Zoogeography

This species has been introduced to Iran by man. Some are undoubtedly aquaculture pond escapees or aquarium releases. Goldfish are kept in aquaria as part of the Now Ruz (New Year) celebrations in March each year. Tehran television (and the Green Front of Iran, see below) urged people to release them into local waters rather than killing them after the New Year (J. Valiallahi, pers. comm., 2000).

Habitat

Goldfish are hardy and can live in winterkill water bodies with much aquatic vegetation, low oxygen, and high pollution (Gudkov, 1985). They can also survive several hours out of water (Pelz, 1987) and may bury themselves in mud, albeit temporarily when scared (Szczerbowski in Bănărescu and Paepke, 2002). Goldfish appear to favour ponds or pools in streams with aquatic vegetation but are often introduced into small bodies of water as ornamental fish. They are tolerant of turbidity, e.g. clay at 225,000 mg/l, pH from 4.5 to 10.5, very high temperatures (upper lethal limit 41.4°C), and high salinity (17‰). This species was killed under experimental conditions, when gradually acclimated to increasing salinity at 28,200 μmho and, by sudden exposure, at 19,200 μmho (Jassim, 1988). This is a greater tolerance than that shown by Cyprinus carpio, another exotic introduced to Iran. However, Carassius auratus appeared in the Basrah fish market when an increase in the Tigris River discharge reduced the salinity of the Shatt al Arab (N. A. Hussain, in litt., 1994).

In Iran it is one of two most abundant species in Caspian wetland areas along with Gambusia holbrooki (Iranian Fisheries Research and Training Organization Newsletter, 19:4, 1998).

Age and growth

Maturity is attained at 3-4 years in the Volga Delta for goldfish. Life span is 13 years with most growth in the first 2-4 years to a size of 15-20 cm (Gudkov, 1985; Kizina, 1986). Life span in captivity in China may exceed 50 years. Population numbers in confined areas are limited by a chemical released by the goldfish which represses more spawning. Prussian carp live up to 11 years.

In the Anzali Mordab, Holčík and Oláh (1992) found only 6 age groups (as did Bagirova et al. (1990) in reservoirs of Azerbaijan while Pipoyan and Rukhkyan (1998) found 9 age groups in Armenia) with the largest fish 32 cm standard length owing to intense fishing pressure. Growth in mm increments was successively 93, 47, 50, 42, 28, and 37. The population is entirely female (see below). Individual life span is greater in Armenia where males are scarce or absent than in bisexual populations (Pipoyan and Rukhkyan, 1998). Sayad Borani et al. (2001) studied this species (as C. auratus gibelio) in the Anzali Mordab at four localities and found a mean fork length of 19.5 cm (range 2.5-31.5 cm) and a mean weight of 196.8 g. The mean age was 2.6 years. The mean length, weight and age were higher in the Sia-Keshim area of the lagoon. The exploitation rate was 0.47, L was 36.0 cm and K was 0.23 per year. Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 41 Iranian fish measuring 5.65-8.17 cm standard length. The a-value was 0.0419 and the b-value 2.911 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases). Patimar (2009) examined fish from the Alma-Gola nd Ala-Gol wetlands in Golestan from 200 to 2002. Ages ranged from 0+ to 8+ with negative allometric growth in Alma-Gol, and positive allometric growth in Ala-Gol. The von Bertalanffy growth curves for mean total lengths were Ltmales = 183.33(1-e-0.31(t+1.05)) and Ltfemales = 245.66(1-e-0.19(t+1.21)) for Alma-Gol and  Ltmales = 224.79(1-e-0.24(t+0.83)) and Ltfemales = 242.80(1-e-0.23(t+0.80)) for Ala-Gol. The sex ratio was unbalanced for males:females at 1:10 and 1:12.7 for Alma-Gol and Ala-Gol respectively because of gynogenesis.

Fish in Buldan Dam Lake, Gediz River basin, Turkey referred to C. gibelio had a maximum age of 6 years and attained 25.5 cm and 269.1 g (Sarı et al., 2008). von Bertalanffy growth parameters were L = 31.66 cm, W = 635.91 g, k = 0.146 year-1 and t0 = -2.166 year. Ratios of total, natural and fishing mortality were calculated as 0.632 year-1, 0.456 year-1 and 0.176 year-1.

Food

Food is predominately zooplankton but also includes aquatic insects, crustaceans, molluscs, worms, detritus, filamentous algae, macrophytes and young fish, switching from one kind of food to another as circumstances warrant. Goldfish have a palatal organ on the roof of the mouth used to taste and touch food and their dense gill rakers aids in feeding on smaller food items. In the recovering Hawr al Hammar, Iraq, diet is 46.1% algae and 25.5% diatoms, with amounts of plants, crustaceans, insects, snails and fish being less than 10% each, in the Hawr al Hawizah 36.3% algae, 21.3% diatoms and 17.5% copepods, with amounts of plants, cladocerans, ostracods and insects being less than 10% each, in the Al Kaba'ish (= Chabaish) Marsh 45.5% algae, 25.2% diatoms, with plants, various crustaceans, insects and snails at less than 10% each (Hussain et al., 2006).

Reproduction

The fish in the Anzali Mordab are all female, reproducing through gynogenesis. Sayad Borani et al. (2001) found fish in Anzali Mordab to have a sex ratio of 99.3 females:0.7 males. Egg development is stimulated by sperm probably from Cyprinus carpio, Tinca tinca, Blicca bjoerkna or Scardinius erythrophthalmus. Here fish may mature at 1 year of age, and coupled with polycyclic ripening of eggs and intermittent spawning, this has led to the dominance of this species in the fresh waters of the lagoon (Holčík and Oláh, 1992). In Armenia, maturity appears to be linked with average annual temperature - at 12.0-13.1°C it occurs at the end of the first year of life while at 8.4-9.0°C it occurs at the end of third and fourth years (Pipoyan and Rukhkyan, 1998). Turkish populations in Topçam Dam Lake, Aydın (Şaşı, 2008) and Buldan Dam Lake, Gediz River basin (Sarı et al., 2008) referred to C. gibelio were 98.84% and 99.44% female. Spawning in the former locality was from March to August, suggesting multiple spawnings with mean fecundity ranging from 37,823 in August to 85,159 in March. Egg diameter reached 1.099 mm in June.

Patimar (2009) examined fish from the Alma-Gola nd Ala-Gol wetlands and found reproduction in February, March and April. Absolute fecundity reached 13,020 eggs.

Spawning begins in late April to mid-May in the Volga Delta and occurs in May-June in the Anzali Mordab (Sayad Borani et al., 2001). Eggs are laid in 2-5 batches over a spawning period extending into July. Up to 10 batches are laid elsewhere at 8-10 day intervals with up to 4000 greenish-yellow eggs in each batch. Fecundity reaches 253,200 eggs (elsewhere to 685,700 with absolute fecundity reaching 860,000 eggs). The largest eggs are 1.6 mm in diameter (Gudkov, 1985; Kizina, 1986; Szczerbowski in Bănărescu and Paepke, 2002). Each female is accompanied by 2 or more males and chases are reported with splashing and shooting through the water near the surface. The eggs are adhesive and attach to water plants and hatch in 5-8 days.

Parasites and predators

Mokhayer (1976b) records infectious dropsy and swimbladder inflammation in Iranian goldfish. Saprolegniosis has been reported from goldfish in Iran (Rahbari and Razavilar, 1982). Growths of the fungus Saprolegnia parasitica resembled tufts of cotton wool. Mokhayer (1989) reports metacercariae of the eye fluke, Diplostomum spathaceum from this species in Iran, which can cause complete blindness and death in commercially important species. Jalali and Molnár (1990a) record the monogeneans Dactylogyrus baueri, D. extensus, D. formosus and D. vastator from this species in the Safid Rud. Jalali and Molnár (1990b) report a variety of monogeneans from this species variously in fish farms throughout Iran, namely Dactylogyrus baueri, D. dulkeiti, D. formosus, D. vastator and D. vastator forma minor. Molnár and Jalali (1992) record the monogenean Dactylogyrus intermedius from this species in a petfish farm near Tehran. Gussev et al. (1993a) describe a new species of monogenean from goldfish on a fish farm near Tehran, Dactylogyrus intermedioides. Shamsi et al. (1997) report Clinostomum complanatum, a parasite causing laryngo-pharyngitis in humans, from this species. The helminth Anisakis sp. is recorded from the guts of this species in the Anzali wetland (Ataee and Eslami, 1999; www.mondialvet99.com, downloaded 31 May 2000). Mousavi (2003) records the monogeneans Gyrodactylus sp., G. kabayashi, D. extensus, D. baueri, Trichodina sp., the ciliates Ichthyophthirius multifilis and Ichthyoboda sp. and the copepods Lernaea cyprinacea and Argulus foliaceus from this species in ornamental fish in Iran. Aquarium specimens are often released in the wild at New Year (Now Ruz). Naem et al. (2002) found the following parasites on the gills of this species from the western branch of the Safid River, namely the protozoans Ichthyophthirius multifilis and a Trichodina species, monogenean trematodes Dactylogyrus anchoratum, and Gyrodactylus sp.. Jalali et al. (2002) and Jalali and Barzegar (2006) record Diplostomum spathaceum from this species in Lake Zarivar. Naem (2002) records the monogenean Dactylogyrus anchoratus from fish in Safid River. Mehdipoor et al. (2004) record the monogenean Dactylogyrus baueri in this fish in the Zayandeh River. Masoumian et al. (2005) recorded the protozoan parasite Ichthyophthirius multifilis from this species in the Aras Dam in West Azarbayjan (species identified as C. carassius, presumably goldfish). Jalali et al. (2005) summarise the occurrence of Gyrodactylus species in Iran and record G. kobayashii and G. sp. in fish from the Safid River. Khara et al. (2006a) record the eye fluke Diplostomum spathaceum for this fish in the Amirkalayeh Wetland in Gilan. Sattari et al. (2004, 2005) surveyed this species (as C. carassius) in the Anzali wetland, recording Raphidascaris acus (and larvae) Eustrongyloides excisus and Camallanus lacustris. Pazooki et al. (2007) recorded various parasites from localities in West Azarbayjan Province, and found Eustrongylides excisus. Sattari et al. (2007) record the nematode Raphidascaris acus, the digenean Diplostomum spathaceum and the monogeneans Dactylogyrus extensus,and Gyrodactylus sp. in this species in the Anzali wetland of the Caspian shore. Barzegar et al. (2008) record the digenean eye parasites Diplostomum spathaceum and Tylodelphys clavata from this fish. Khara et al. (2008) found the eye parasite Diplostomum spathaceum in this fish from Boojagh Kiashar Wetland in Gilan. Shamsi et al. (2009) found Dactylogyrus baueri, D. dulikeity, D. extensus, D. intermedius, D. intermedioides and D. wegeneri in this species from localities such as fish farms, the Safid River, Zayandeh River, Zarivar Lake and the Hamun Lake. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Lernaea sp. on this species.

Economic importance

This species is raised on Tehran fish farms for the pet trade (Molnár and Jalali, 1992). It forms part of the Now Ruz (= New Year, usually 21 March) celebrations in Iran where a bowl with goldfish forms part of the traditional Haft Sin table setting (so called for seven items that must be present, all beginning with the letter "S", each having a symbolic meaning, the goldfish is in addition to these). The goldfish in a bowl represents life within life, and the sign of Pisces which the sun is leaving.

Haft Sin Table, Iranian Embassy, Ottawa, 2009, Brian W. Coad.  aquarium goldfish at Iranian Embassy, Ottawa, 2009. Brian w. Coad.

Haft Sin table, Iranian Embassy, Ottawa, 2009

 

Ebrahimzadeh Mousavi and Khosravi (2004) report suspected epizootic ulcerative syndrome from ulcerated goldfish.

In the Anzali Mordab, 62% of the total catch is goldfish, an accidental introduction (Petr, 1987). The catch in the mordab in 1990 was 46,472 kg (Holčík and Oláh, 1992). As the salinity of this lagoon increases, the density of goldfish will decrease. Valeipour and Haghighy (2000) record the catch for 1992-1996 at 40% of the species taken. Safaei (2005) gives a goldfish catch figure of 45% of the 313 ton fishery there in 1992. The presence of goldfish in the Anzali Mordab led to a decline in the native fishery there.

This species is caught by anglers at Ahvaz in Khuzestan using bread or potato as bait.

It is known to control mosquito larvae in Bengal (Chandra et al., 2008).

The peculiar type of reproduction is very successful and affects the catches of other cyprinid species, being equivalent to a predatory effect (Holčík and Oláh, 1992).

There is some evidence that this fish disturbs the habitat of native species, muddying waters, and it may compete for food and space. Goldfish have destroyed some amphibian populations in other parts of the world by consuming frog eggs (Coad and Abdoli, 1993b). The Green Front of Iran recommended the release into pools of mosques, parks or natural lakes of the estimated 20 million goldfish kept in aquaria for the Iranian New Year celebrations in March each year. This would have a deleterious effect on habitats not yet colonised by this exotic species. A news report in 2005 cites the death of 5 million fish in transit from the store to the Iranian home at New Year, indicating perhaps that the numbers that do make it are much higher (www.politicalgateway.com, downloaded 5 August 2005). Newspaper articles suggested that goldfish should only be released into "pools" rather than rivers because of all the attendant dangers of this exotic. They are known to prevent reproduction of native species in Sistan (Iran Daily, 17 March 2005, p. 5).

This species is used in Iran as an experimental organism and for studies in reproductive biochemistry, e.g. in studying the effects of anionic detergents (shampoos, a common water pollutant) on blood parameters, on hepatic and renal pathology and serum biochemical parameters (Shahsavani et al., 2003; Shahsavani et al., 2004; Shahsavani et al., 2005; Shahsavani and Movassaghi, 2003); the use of phenytoin sodium on skin wounds (5mg/l showed best healing improvement while zinc oxide was not as effective) (Shahsavani et al., 2001, 2002, 2002); the formation of lesions and clinical changes in fish exposed to kerosene (Shahsavani et al., 2003); the effects of cortisol on testicular apoptosis (Bahmani et al., 2007); the adverse effects of phenytoin sodium, a drug used for healing skin lesions, on the gills, liver and kidney (Shahsavani et al., 2007); hormonal GnRHa and pituitary extract proved more effective on spermatological parameters than the hormone HCG (Zadmajid et al., 2008); the effects of seminal plasma indices on sperm motility (Zadmajid and Imanpour, 2009); the effects of hormones on seminal plasma biochemistry (Zadmajid et al., 2009).

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in textbooks, in aquaria and in aquaculture, as bait, as an experimental species and because it has been introduced outside its natural range. There are numerous, commercial aquarium forms with particular morphologies and colours that are assigned common names, e.g. common, veiltail, comet, fans, calicoe, black-moor, telescope-veiltail, lionhead, egg-fish, shubunkin. Balon (2006) reviews the origin of the species.

Conservation

This species is a successful exotic, in no need of conservation.

Further work

The Carassius species in Iran is generally regarded as C. auratus, the goldfish of aquaria, as it is used extensively in Now Ruz (New Year) celebrations and often released into natural waters. This needs confirmation for all major populations.

Sources

Iranian material: CMNFI 1979-0230, 41, 14.7-38.6 mm standard length, Sistan, Hamun-e Puzak (ca. 31º15'N, ca. 61º42'E); CMNFI 1991-0162, 1, 40.5 mm standard length, Mazandaran, Bagher Tangeh (36º42'N, 52º43'E); CMNFI 1993-0136, 64.0 mm standard length, uncatalogued material, 1, 93.5 mm standard length, Gilan, near Hendeh Khaleh (ca. 37º23'N, ca. 49º28'E); 1, 52.8 mm standard length, Gilan, near Hendeh Khaleh (ca. 37º23'N, ca. 49º28'E); 4, 16.4-50.3 mm standard length, Gilan, near Khoshk Bijar (ca. 37º22'N, ca. 49º47'E).

Carassius carassius
(Linnaeus, 1758)

Carassius carassius from Wikimedia Commons.

 

Carassius carassius from Wikimedia Commons.

Carassius carassius from Wikimedia Commons.

The crucian carp has been reported as introduced to Iran in the Karun River basin as aquarium releases by Armantrout (1980) without further details and there are other reports such as in the Gorgan River (Y. Keivany, in litt., 1992) and Mahabad Dam (Abdi, 1999; www.mondialvet99.com, downloaded 31 May 2000) but these may be confusion with Carassius auratus. Specimens are needed to confirm the presence of this species in Iran. The native distribution is in Europe and western Asia, reaching northern drainages of the Caspian Sea in the southern limits of its distribution (Libosvárský, 1962). It differs from C. auratus in having a slightly convex margin to the dorsal fin (straight or slightly concave in C. auratus), caudal fin slightly emarginate (deeply emarginate), usually 6 branched anal rays (always 5), 23-33 gill rakers (37-53), 31-34 vertebrae, usually 32-33 (28-31, usually 29-30), 28-29 fin denticles posteriorly on the dorsal fin spine (10-11), peritoneum light (dark), black spot at the caudal fin base in young and some adults (absent), and a coppery gold body (silvery, pinkish gold, gold or red) (Szczerbowski in Bănărescu and Paepke, 2002). Berg (1948-1949) also cites the characters body rounded, back thick (body angular, back compressed) and scales weakly sculptured (rough), although his comparison is with C. a. gibelio, itself recognised as a distinct Carassius gibelio (Bloch, 1782), also of uncertain occurrence in Iran.

Carassius gibelio from Wikimedia Commons.
Carassius gibelio from Wikimedia Commons.

Genus Chondrostoma
Agassiz, 1832

The nases are found from the Iberian Peninsula and France to the Caspian and Tigris-Euphrates basins. There are about 26 species of which 2 are known for Iran (Elvira, 1997). Chondrochylus Heckel, 1843 and Chondrochilus Heckel, 1843 are synonyms. Eschmeyer (1990) gives the year of publication for the genus as 1832 as opposed to other authors who give 1835 (e.g. Berg, 1948-1949; Reshetnikov et al., 1997). Doadrio and Carmona (2004) confirm the monophyly of the genus based on the cytochrome b gene with vicariant events accounting for distribution of taxa better than a dispersalist model. Middle East taxa belong to a single lineage with the more differentiated and basal species in the Caucasus and Mesopotamia, having been isolated in the Upper Miocene-Pliocene.

This genus is characterised by being of moderate size, with a somewhat compressed body, scales of moderate to small size (44-106 in the lateral line (Robalo et al. (2007) give a range of 52-78 for their more restricted genus)), scales squarish with radii in the anterior and posterior fields and a subcentral anterior focus, no barbels, an inferior and transverse or crescentic mouth with a cutting edge to the lower jaw, thin upper lip and no lower lip, pharyngeal teeth knife-like and in 1 row with a high count (5, 6 or 7, the same number on each arch or one more on the left), gill rakers short and moderately numerous (up to 40), short dorsal fin without a thickened ray opposite the pelvic fins, 7-10 dorsal fin branched rays, a moderately elongate anal fin with 8-12 branched rays, deeply forked caudal fin and usually concave dorsal and anal fins, a pelvic axillary process always present, 42-49 vertebrae, a black peritoneum, and a long, coiled gut. Elvira (1997) and Robalo et al. (2007) give osteological characters.

Bogutskaya (1997a) places the nominal Iranian species, C. regium and C. orientalis, in a group characterised by a straight or only slightly arched mouth cleft, high vertebral counts (total vertebrae modes 45-47 and abdominal modes 26-28) and often or commonly 4 unbranched rays in the dorsal fin.

Chondrostoma cyri
Kessler, 1877

Common names

shekamsiah-e Aras.

[Kur altagizi in Azerbaijan; chernobryushka or blackbelly, Kurinskii podust or Kura nase, uzkotelii Kurinskii podust, all in Russian; Kura undermouth, Kura nase].

Systematics

Earlier works by Elvira (1986; 1988; 1991) placed this species as a subspecies of C. oxyrhynchum but in Elvira (1997), using the phylogenetic species concept and following the studies of Smirnov (1992), this taxon is recognised as a species. C. oxyrhynchum is then found in more northerly rivers of the western Caspian Sea basin remote from Iranian waters. C. cyri orientalis (Bianco and Banarescu, 1982 is described from Fars (see below under C. regium).

Chondrostoma cyri Kessler, 1877 was described from the Kura River, Tiflis (= Tbilisi), Georgia and Chondrostoma oxyrhynchum from the Kuma River near Georgiyevsk, Russia in the Caspian Sea basin.

Alburnus alasanicus Kamenskii, 1901 described in part from the Alasan, Alazan' or Alazani River, a left bank Kura River tributary in Georgia, Chondrostoma schmidti, Berg, 1910 from the Alazan' River at Naporiri, and Chondrostoma leptosoma Berg, 1914 from the Kars-tchai, a tributary of the Aras River in Turkey, the Aras by Kopri-kei, near Erzurum, Turkey, and the lower Aras at Karadonly and Dzhulfa in the former U.S.S.R., are synonyms. Subspecies are not recognised (Elvira, 1991; 1997). C. leptosoma was founded on an elongate form from the Karasu in the Aras River basin.

Two syntypes of Chondrostoma cyri are also in ZISP (10919) from "Tiflis" collected by Kessler in September 1875. A syntype of Chondrostoma oxyrhynchum is in the Zoological Institute, St. Petersburg (ZISP 2881) from "Fl. Sunsha" collected in 1830 by Ménétries. According to Elvira (1988), the type locality is Kuma R. at Georgijewsk and 2 syntypes are under ZISP 10922. Another syntype of Chondrostoma oxyrhynchum is in the Natural History Museum, London (BM(NH) 1897.7.5:28 (184.8 mm standard length), formerly in ZISP, as is other syntype of Chondrostoma cyri (BM(NH) 1897.7.5:27 (correctly numbered 27, 78.4 mm standard length), formerly in ZISP)(Elvira, 1988; personal observations).

Five syntypes of Chondrostoma leptosoma are in the Zoological Institute, St. Petersburg (ZISP 9098) according to (Elvira, 1988) but there are 15 syntypes under this number from the "Reka Araks", 1888, Warpochowsky as well as additional material listed as syntypes with numbers ZISP 9107 ("Fl. Araxes", 1888, Warpochowsky, 12 fish), ZISP 5180 ("Kars-tschai", 1879, Dr. A. Brandt, 3 fish), ZISP 9099 ("Reka Araks", 1888, Warpochowsky, 4 fish), ZISP 15264 ("Reka Araks", 20.III.1911, 2 fish), and ZISP 15516 ("Reka Araks near settlement Djulfa", 17.VI.1911, 13 fish).

Key characters

This species is the only one in its genus in northern Iran and can be recognised by generic characters.

Morphology

Kuru (1981) gives the following meristic characters for 103 specimens from the Aras and Kura river basins in Turkey:- 10-12 dorsal fin rays, 10-11 anal fin rays, 9-10 pelvic fin rays, 9-15 pectoral fin rays, 52-62 lateral line scales, 13-18 scales around the caudal peduncle, 17-32 gill rakers, and 5-6 pharyngeal teeth on each arch (note that the statistical treatment in this paper is in error and the conclusion that species of Chondrostoma in Turkey are not distinct is therefore incorrect). There is clinal variation in scale numbers, the number increasing from south to north and Elvira (1988; 1991) gives the total range for characters of this species as dorsal fin branched rays 7-9, usually 8, anal fin branched rays 8-10, usually 9-10, pectoral fin branched rays 13-18, usually 14-16, pelvic fin branched rays 7-8, usually 8, lateral line scales 50-68 (to 73 in Kazancheev (1981) and from 48 in Chikova (1967)), scales above the lateral line 7-10, usually 8-10, scales below the lateral line 3-6, usually 4-6, pharyngeal teeth 6-5 or 5-5, more rarely 6-6 and mode 6-5, and gill rakers 21-29. Vertebrae number 43-45.

The mouth is arched with a thin horny layer on the lower jaw. Scales are rounded in overall shape with indentations above and below a central, rounded protuberance on the anterior margin. The anterior margin may be wavy. There are few anterior and posterior radii, few circuli and a subcentral anterior focus. There is a pelvic axillary scale. The gill rakers are short and reach the one below or just past it when appressed. Pharyngeal teeth are compressed and thin but deep with a long, thin and concave grinding surface. Teeth tips may be slightly hooked. The gut has numerous anterior loops.

Thirteen specimens from Djulfa (presumably in Azerbaijan opposite the Iranian town across the Aras River) have dorsal fin branched rays 8(12) or 9(1), anal fin branched rays 9(9) or 10(4), and pharyngeal teeth 6-5(5) or 6-6(1).

Sexual dimorphism

Unknown.

Colour

The flanks are silvery but may have dark pigment spots which may, or may not, form a stripe. Paired fins are orange to reddish and median fins grey. The dorsal and caudal fins have dark margins. The peritoneum is black.

Size

Reaches 80.0 cm and about 5.0 kg.

Distribution

Found in the rivers draining to the western coast of the Caspian Sea from the Kuma River in the north southward to the Kura and Aras river basins in the south. Recorded from the Aras River basin of Iran (Abdoli, 2000).

Zoogeography

This genus has a European and Middle Eastern distribution. Its relationships to other taxa are poorly known.

Habitat

Unknown. Found principally in streams and rivers.

Age and growth

Fish are mature at 2 years of age and life span is at least 5 years.

Food

Diet is assumed to consist of bottom organisms including aquatic insect larvae, detritus and vegetation scraped from the substrate.

Reproduction

Up to 16,217 eggs are produced and maximum diameter is 1.69 mm. The spawning season is in the spring, peaking in April in the Kura River basin (Abdurakhmanov, 1962).

Parasites and predators

None reported from Iran.

Economic importance

None.

Conservation

Kiabi et al. (1999) consider this species to be conservation dependent, in the south Caspian Sea basin according to IUCN criteria. Criteria include sport fishing, possibly few in numbers, limited range (less than 25% of water bodies), absent in other water bodies in Iran, absent outside the Caspian Sea basin.

Further work

Biology in Iranian waters needs study.

Sources

Morphology based on Bianco and Banarescu (1982), Elvira (1986; 1988), Nelva et al. (1988).

Type material: See above, Chondrostoma cyri (BM(NH) 1897.7.5:25, formerly in ZISP), Chondrostoma oxyrhynchum (BM(NH) 1897.7.5:28, formerly in ZISP), and Chondrostoma leptosoma (ZISP 15516).

Iranian material: None.

Comparative material: CMNFI 1980-0812, 2, 101.9-107.9 mm standard length, Turkey, Kars, Selim Çayi (40º28'N, 42º47'E).

Chondrostoma orientale
Bianco and Banarescu, 1982

Chondrostoma cyri orientalis Bianco and Banarescu, 1982 was originally described from the "Pulwar River near Persepolis".

The holotype (IZA 8170, 93.7 mm standard length, examined by me) and 19 paratypes (IZA 7833, 51 specimens under this number, 35.4-90.1 mm standard length) of Chondrostoma cyri orientalis are in the Istituto di Zoologia dell'Universitá di L'Aquila, Italy (Elvira, 1988). Two paratypes of Chondrostoma cyri orientalis are stored in the Field Museum of Natural History, Chicago (FMNH 94519)(Ibarra and Stewart, 1987), 1 paratype is in the Muséum national d'Histoire naturelle, Paris (1982-1014), 1 paratype is in the United States National Museum, Washington (USNM 227934), 2 paratypes are in the Academy of Natural Sciences, Philadelphia (ANSP 150985), and 6 paratypes are in the Canadian Museum of Nature, Ottawa (CMNFI 1982-0365, formerly IZA 7833, 37.8-88.7 mm standard length). The total number of paratypes is 75, originally under IZA 7833 but some dispersed as noted above, with 10 further fish in the Institutul de Stiinte Biologice, Bucurešti, Romania (ISBB) but uncatalogued (Bianco and Banarescu, 1982).

Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found  Lernaea cyprinacea on this species (as C. regium).

 

Barzegar and Jalali (2006) report parasites in this species from Kaftar Lake as Unio sp., Lernaea cyprinacea, Ichthyophthirius multifilis and Diplostomum spathaceum.

Chondrostoma regium
(Heckel, 1843)

Common names

jokhorak, nazok, nazi; heif-e nan (= waste of bread, i.e. valueless) in Khuzestan; سياه ديم (= siah deem in Behbehan); سياه دم (= siah dom, meaning blacktail); كپور پوزه دار (= kapur puzeh dar).

[baloot muluki, pangki; zurri (= the harmful one) at Mosul (also used for Alburnus mossulensis, Aphanius spp., Gambusia and any small fishes or large fishes when young); terris or terris achmar meleki (= royal red terris) at Aleppo (= Haleb, Syria), all in Arabic; based on Heckel (1843b) for zurri and terris; king nase].

Systematics

Chondrochilus regius Heckel, 1843 was described from the "Orontes" (= Asi) (but see below) and "Tigris". Elvira (1988; 1991; 1997) considers Chondrostoma orientale to be a valid species while Nelva et al. (1988) retain it as a subspecies of C. cyri. Bianco and Banarescu (1982) placed orientalis in C. cyri on the basis of similar dorsal and anal fin ray counts, scale counts and to a certain degree pharyngeal tooth formula.

Banarescu (1960) regarded C. regium as only a race of a widespread species, C. nasus (Linnaeus, 1758). C. nasus has larger scales on average and 6-6 pharyngeal teeth (Berg, 1949); Heckel (1846-1849c) found 47 C. nasus from the Danube River had 6-6 teeth, 2 had 6-7 and 2 had 5-6 while in 13 C. regium the count was 7-6 in 12 fish and 6-6 in 1 fish. Krupp (1985c) considers C. regium to be distinct while recognising the small degree of morphological variation between species in this genus. Data gathered for Iran show a wide range in scale and teeth counts (see below). Ladiges (1960) identified specimens from the same bodies of water in Turkey as members of both species. The earlier literature on the systematics of this genus remain confused (see Elvira (1988) for comments on Ladiges (1966) and Kuru (1981)) and the morphology summarised here for this species does not adequately resolve the problem. There may well be significant variation of a clinal nature, altitude and temperature may be important, and habitat types (lentic or lotic) may affect body form. Most samples examined previously are too small in numbers and differences due to size and sex could not be adequately assessed.

Twelve syntypes of Chondrostoma regium are in the Naturhistorisches Museum Wien (7 fish as NMW 52532-52535 from the Quwayq (= Kueik) River near Aleppo and 5 fish as NMW 52536-52538 from the Tigris River near Mosul)(Elvira, 1988). Krupp (1985c) gives further details. All material was collected by Th. Kotschy in 1842 from the Quwayq and in 1843 from Mosul and the range in standard length for the fish from the Quwayq is 102-166 mm and from Mosul 11.9-24.1 cm. The Vienna catalogue lists only 6 fish but the card catalogue in 1997 lists NMW 52532 (2 fish), 52533 (2), 52534 (2), 52535 (1), 52536 (2), 52537 (1) and 52538 (2) as syntypes. The type locality "Orontes" (= Asi) in Heckel (1843b) seems to be an error.

Key characters

This species is the only one in its genus in southern Iran and can be recognised by generic characters.

Morphology

?re-work

The following counts are from literature sources; my counts in the table below often show a wider range: lateral line scales 56-72 (47-55 for orientalis), scales above the lateral line 9-13 (8-9 in orientalis), and scales below the lateral line 5-6 (4-5 in orientalis). Lateral line scale counts for Iranian fish are as follows: Tigris - 50(1), 51(4), 52(3), 53(12), 54(5), 55(7), 56(4), 57(4), 58(4), 59(8), 60(7), 61(6), 62(5), 63(3), 64(3), 65(1), 66(2), 67(1) or 69(2); Kor (= orientalis) - 49(1), 50(2), 51(3), 52(8), 53(3), 54(7), 55(3) or 57(4). Despite a lower range, the counts for the Kor River basin are matched by a sample from Cheshmeh Javari near Ravansar, Kermanshahan (CMNFI 1979-0287) which have a range of 50-58, leading to a supposition of altitudinal or habitat variation :-

Dorsal fin branched rays 8-11, mode 9 (note Bogutskaya (1997a) gives a mode of 10) (7-9, mostly 8 for orientalis), anal fin branched rays 9-12, mode 11 (note that Bogutskaya (1997a) gives modes of 11 or 12) (9-10, mode 9 for orientalis), pectoral fin rays 14-18, mostly 15-17 (13-15, mostly 14 in orientalis) and pelvic fin rays 6-9, mostly 8 (7-8, mostly 8 in orientalis). Gill rakers 18-36 (probably some lower counts are of rakers on the lower arch only and ranges in single studies, presumably to a consistent technique, are 22-34, 24-31, 25-34 and 25-36) (22-28, 22-30 or 28-32 by different authors for orientalis). Counts for the whole arch on Iranian fish give a wide range of 19-34, highly correlated with size, larger fish having more (or more discernible) rakers than smaller fish (r = 0.5049, p<0.001, n = 90).

Scale radii are few and restricted to the posterior field. Total vertebrae 46-48. Pharyngeal teeth 6-5, 6-6, 6-7, 7-5, 7-6 and 7-7, mode 6-6 or 7-6 (6-5, 6-6, 5-6 and 7-5, mostly 6-6 for orientalis) but see above. The mouth is straight (= transverse) with a thick horny layer on the lower jaw. Esmaeili et al. (2010) give a diploid chromosome number of 2n=52 with 21 pairs of submetacentric and 5 pairs of subtelocentric chromosomes from the Fahlian River in Fars. The arm number was 58.Other Chondrostoma species have 2n=50.

Meristics for southern Iranian specimens of Chondrostoma:


Locality/Dorsal Fin Branched Rays

7

8

9

10

x

S.D.

Tigris River Basin

35

46

1

8.6

0.520

Kor River Basin

1

30

8.0

0.180

Locality/Anal Fin Branched Rays

8

9

10

11

12

x

S.D.

Tigris River Basin

4

45

21

9

3

9.5

0.892

Kor River Basin

23

8

9.3

0.445

Locality/Pelvic Fin Rays

7

8

9

x

S.D.

Tigris River Basin

3

77

2

8.0

0.248

Kor River Basin

1

29

1

8.0

0.258

Locality/Lateral Line Scales

Range

x

S.D.

Tigris River Basin

50-69

57.8

4.553

Kor River Basin

49-57

53.2

2.131

 

Locality/Total Vertebrae

42

43

44

45

46

47

48

49

x

S.D.

Tigris River Basin

1

20

17

9

14

10

9

2

45.1

1.833

Kor River Basin

5

20

5

1

43.1

0.680

Sexual dimorphism

Unknown.

Colour

The back is olive-brown with bluish tinges and the flanks and belly are silvery-white. The dorsal and caudal fins are greyish and the other fins hyaline. Some fish have bright orange fins, the pectorals paler, the pelvics and anal fins fringed by white. The dorsal and caudal fins have a black margin, wide on the caudal. These fin colours give them a flag-like effect (Heckel, 1843b). The caudal fin can be orange, distally black, with the extreme edge white in freshly dead fish.

Size

Attains 40 cm and 1 kg.

Distribution

Found in the Tigris-Euphrates basin and the Mediterranean basins of southeastern Turkey and the northern Levant. In Iran found in the Tigris River basin. Additional localities are springs (sarabs) near Kermanshah, the Marun River, the Hawr al Azim marsh (Wossughi, 1978; Abdoli, 2000). Ghorbani Chafi (2000) lists the Bazoft and Kuhrang rivers in the upper Karun River basin and also possibly the Zayandeh River of the Esfahan basin.

Zoogeography

This genus has a European and Middle Eastern distribution. Its relationships to other taxa are poorly known.

Habitat

Found in both rivers and lakes (and reservoirs) but habitat requirements have not been studied in Iran. Ünlü (2006) reports that this species prefers stone grounds and still waters in rivers and lakes in Turkey.

Age and growth

Khalaf et al. (1986) studied this species in the Diyala River, Iraq. Maximum age group is 7+ years, males and females show no difference in weight at the same length and samples from three adjacent areas show no major differences in growth rates. Length-weight relationship was W = 0.0480 L2.49 (n = 255, r = 0.88). Males mature at 15.0 cm and females at 19.0 cm in the Diyala River at Rustamiyah in Iraq (Allouse et al., 1986). A population at Al Kadhmia north of Baghdad in the Tigris River had four age classes dominated by the three year age class, with all fish being sexually mature during the second year. Fish smaller than 15 cm for males and 17 cm for females were immature. The disparity in age structure with the Diyala River population was attributed to pollution in the Diyala (Daoud and Qasim, 1999).

Polat and Gümüş (1995) aged a population of this species in the Bafra Altınkaya Dam lake in Turkey using vertebrae, otoliths, scales, opercle and subopercle. Age reached 5, perhaps 6, years and scales were found to be the best structure to use. Polat et al. (1999) found a similar age range in the Suat Uğurlu Dam, Turkey with annulus (hyaline ring) formation in October to February. Oymak (2000) examined growth characteristics of this species in the Atatürk Dam on the Turkish Euphrates River. Eight age groups were found and age-length and age-weight equations given for females and males were Lt = 38.67[1-e-0.136126(t+3.073799)], Wt = 527.52[1-e-0.136126(t+3.073799)]3.1986 and Lt = 35.01[1-e-0.168137(t+2.754214)], Wt= 724.73[1-e-0.168137(t+2.754214)]3.2779 respectively. The length-weight relationships were obtained as Log W = -5.4153 + 3.1986 Log FL in females and Log W = -5.6212 + 3.2779 Log FL in males. The condition factor was high in age group 7 and high in April and May, lowest in December and January. Gümüş et al. (2002) found deposition of hyaline rings was synchronous with decrease in food diversity in autumn in the Suat Uğurlu Dam, Turkey. Aydin et al. (2004) demonstrated a positive linear relationship between otolith length and fish length for this species in Keban Dam Lake, Turkey.

Food

This species is omnivorous taking insect larvae and eggs and fry of other fishes. Gut contents also include diatoms and algae as well as large quantities of sand. However, Gümüş et al. (2002) examined diet in the Suat Uğurlu Dam, Turkey and found Navicula, Cymbella and Synedra were the most frequently consumed organisms. This species feeds mostly on Bacillariophyta in this dam but also Chlorophyta, Cyanophyta, Xanthophyta, Euglenophyta and Rotifera. Diet varied with seasonable availability of food items.

Reproduction

Studies on the Diyala River population in Iraq found fish to be mature in December and by January females lacked eggs. Each female produces up to 6900 eggs and number of eggs increases linearly with length (Allouse et al. (1986). The breeding season at Al Kadhmia in the Tigris River near Baghdad was March-May (Daoud and Qasim, 1999). Al-Rudainy (2008) gives sexual maturity as 3 years, 25 cm total length and 250 g weight with spawning in February and March on gravel beds in shallow water with strong current. for Iraq. Ünlü (2006) reports up to 13,280 eggs for fish in the Tigris River of Turkey. Beckman (1962) states that this species probably spawns in May or June in Syria and Oymak (2000) found that condition factors were highest in April and May in the Atatürk Dam, Turkey.

Parasites and predators

Barzegar et al. (2004) examined this species for parasites in fish from the Beheshtabad river in Chahar Mahall va Bakhtiari Province and found Lernaea cyprinacea, Dactylogyrus ergensi, Ichthyophthirius multifilis and Myxobolus sp. Jalali et al. (2005) summarise the occurrence of Gyrodactylus species in Iran and record G. sp. from the Dez and Karun rivers in Chondrostoma nasus, presumably this species. Barzegar et al. (2008) record the digenean eye parasites Diplostomum spathaceum and Tylodelphys clavata from this fish. Riassy et al. (2009) found the digenean eye parasite Tylodelphys clavata in fish from Choghakhor Lagoon.

Economic importance

This species has been caught and used for food in Khuzestan.

Conservation

This species is relatively common and is not widely used as food; it may not need conservation. However it is listed as endangered in Turkey (Fricke et al., 2007).

Further work

Its biology in Iran needs study and this may reveal conservation needs.

Sources

?re-work

Type material: See above, Chondrostoma cyri orientalis (IZA 8170, IZA 7833, CMNFI 1982-0365, formerly IZA 7833).

Iranian material: CMNFI 1979-0025, 16, 22.1-119.0 mm standard length, Fars, Kor River at Marv Dasht (29º51'N, 52º46'30"E); CMNFI 1979-0028, 14, 32.2-139.1 mm standard length, Fars, Kor River drainage (no other locality data); CMNFI 1979-0059, 1, 72.2 mm standard length, Fars, Pulvar River (30º01'30"N, 52º57'E); CMNFI 1979-0061, 14, 9.5-56.5 mm standard length, Fars, stream tributary to Pulvar River (30º04'N, 53º01'E); CMNFI 1979-0245, 5, 35.3-47.1 mm standard length, Sharestan-e Bahktiari va Chahar Mahall, stream in Ab-e Shalamzar drainage (32º08'N, 50º51'E); CMNFI 1979-0247A, 4, 57.2-65.3 mm standard length, Sharestan-e Bakhtiari va Chahar Mahall (31º57'N, 51º01'E); CMNFI 1979-0248, 2, 39.2-65.2 mm standard length, Sharestan-e Bakhtiari va Chahar Mahall, stream 3 km east of Boldaji (31º55'N, 51º05'E); CMNFI 1979-0271, 11, 60.0-131.3 mm standard length, Lorestan, Kashkan River drainage (33º39'N, 48º32'30"E); CMNFI 1979-0272, 1, 58.5 mm standard length, Lorestan, river at Nokhor (ca. 33º40-47'N, ca. 48º28-45'E); CMNFI 1979-0279, 2, 61.8-134.0 mm standard length, Lorestan, Khorramabad River (33º37'N, 48º18'E); CMNFI 1979-0280, 1, 114.5 mm standard length, Lorestan, Kashkan River drainage (ca. 33º43-47'N, 48º12-15'E); CMNFI 1979-0283, 1, 137.0 mm standard length, Kermanshahan, river 15 km before Kermanshah (34º21'N, 47º07'E); CMNFI 1979-0287, 22, 56.6-112.5 mm standard length, Kermanshahan, Chashmeh Javari near Ravansar (ca. 34º42'N, ca. 46º40'E); CMNFI 1979-0286, 11, 77.4-100.4 mm standard length, Kermanshahan, Ravansar River at Ravansar (34º43'N, 46º40'E); CMNFI 1979-0289, 1, 131.5 mm standard length, Kermanshahan, Diyala River drainage (34º28'N, 45º52'E); CMNFI 1979-0368, 4, 54.0-84.5 mm standard length, Khuzestan, Karkheh River (32º24'30"N, 48º09'E); CMNFI 1979-0370, 6, 187.3-221.6 mm standard length, Khuzestan, Karkheh River (32º12'N, 48º14'30"E); CMNFI 1979-0382, 2, 37.7-62.5 mm standard length, Khuzestan, Karun River at Shushtar (32º03'N, 48º51'E); CMNFI 1979-0392, 1, 53.7 mm standard length, Khuzestan, Zard River (ca. 31º32'N, ca. 49º48'E); CMNFI 1979-0421, 5, 114.0-122.0 mm standard length, Boyer Ahmadi-ye Sardsir va Kohkiluyeh, stream in Khersan River drainage (30º24'N, 51º47'E); CMNFI 1979-0499, 1, 113.0 mm standard length, Fars, irrigation ditch 32 km from Kor River bridge (30º04'30"N, 52º36'E); CMNFI 1979-0500, 7, 94.8-110.5 mm standard length, Fars, Pulvar River at Naqsh-e Rostam (29º59'N, 52º54'E); CMNFI 2007-0100, 2, 165.4-165.7 mm standard length, Azarbayjan-e Bakhtari, Kalwi Chay near Piranshar (ca. 36º44'N, ca. 45º10'E); CMNFI 2007-0111, 2, 183.3-191.7 mm standard length, Kermanshahan, Alvand River near Sar-e Pol-e Zahab (ca. 34º36'N, ca. 45º56'E); CMNFI 2007-0113, 2, 106.7-145.0 mm standard length, Kermanshahan, Razavar River, Qareh Su tributary (ca. 34º25'N, ca. 47º01'E); CMNFI 2007-0115, 3, 72.5-96.5 mm standard length, Kermanshahan, Qareh Su basin north of Kermanshah (ca, 34º34'N, ca. 46º47'E).

Comparative material: BM(NH) 1931.8.12:1-3, 2, 136.0-172.2 mm standard length, Iraq, near Mosul (36º20'N, 43º08'E); BM(NH) 1971.4.2:6, 1, 147.7 mm standard length, Iraq, River Tigris near Mosul (36º20'N, 43º08'E); BM(NH) 1974.2.22:81-82, 1, 197.5 mm standard length, Iraq, Great Zab near Eski Kelek and near Bekhne Dam (no other locality data).

Genus Crossocheilus
Kuhl and van Hasselt, 1823

Kottelat (1987) retains the spelling Crossocheilus Kuhl and van Hasselt in van Hasselt, 1823 as first reviser. The name was spelt Crostocheilus early in the text but this has never been used again and Crossocheilus appeared with the description. Crossochilus Günther, 1868 is an incorrect emendation (Eschmeyer, 1990).

The genus is found chiefly in the Oriental Region but extends into Iran with one species. There are about 18 species.

The genus is characterised by an elongate body with a rounded belly; the mouth is inferior and transverse, the crenulated or fringed upper lip being continuous with the snout (not separated by a groove); the lower jaw has a horny covering and behind this are several rows of lobate papillae; 1-2 pairs of barbels; gill membranes attached to isthmus; dorsal and anal fins are short and spineless; the lateral line is complete; scales are large to moderate in size; the intestine is very long; and the peritoneum is black.

The lower surface of the head bears an "adhesive apparatus", the mechanism of which has been investigated by Singh (1993) for Crossocheilus latius latius, a subspecies not found in Iran. The fringed upper lip acts as a food strainer as well as part of the adhesive apparatus. Both this lip and the area behind the lower lip are heavily tuberculate with glandular openings and irregularly arranged hard ridges. Mucus from the glands in conjunction with the ridges holds the fish to the substrate.

Crossocheilus latius
(Hamilton, 1822)

Common names

None.

[ispigoar or dogra in Pakistan].

Systematics

Cyprinus latius was described from the Tista River in India/Bangladesh and types are unknown (Eschmeyer et al., 1996).

Bianco and Banarescu (1982) and Bănărescu (1986) consider Discognathus adiscus Annandale, 1919 described from Sistan (type locality given below) to be a synonym of this species which is represented in Sistan and Baluchestan by Crossocheilus latius diplocheilus (Heckel, 1838), originally described from Kashmir with syntypes in the Naturhistorisches Museum Wien under NMW 48820 (7 fish). Bănărescu (1986) cites 1 fish under NMW 48420 as possibly the holotype but this specimen is dated 1839 which is anachronistic. Berg (1949) considers Discognathus adiscus to be a distinct species. I concur with Bianco and Banarescu (1982) and Bănărescu (1986).

Characters advanced by Berg (1949) for separating the two species are number of barbels (4 in adiscus, 2 in latius diplocheilus where mouth angle barbels are absent or rudimentary, not the rostral ones as implied by Bianco and Banarescu (1982)), upper lip fringe (barely developed in adiscus, distinctly developed in latius diplocheilus), the posterior swimbladder (conical in adiscus, elongate cylinder in latius diplocheilus), and papillae on the lower lip and chin (rudimental in adiscus and latius diplocheilus but the latter has almost free lateral edges and an attached posterior end - this condition is not specified for adiscus). Bianco and Banarescu (1982) and Bănărescu (1986) found some latius diplocheilus specimens to have 4 barbels (and this is given too as a character of C. latius latius), and no difference in development of lip papillae in specimens from the Indus River basin (actually my reading of Berg (1949) cited above does not indicate that papillae development differs but that the sucker area has almost free lateral edges and an attached posterior end; this occurs in Sistan fish but not in 4 fish from the Hamun-e Mashkid and Makran basins of Iran which have a fold in the flesh behind the tuberculate area - these latter fish are very small however, 20.8-27.8 mm standard length, and I lack extensive comparative adult material from outside the Sistan basin in Iran and from neighbouring Pakistan to make an adequate analysis of nominal adiscus and latius diplocheilus in this and other characters). My observations of the posterior swimbladder development indicate a great individual variation in form for Sistan fish: the swimbladder may be conical, elongate and tapering, rounded posteriorly, expanded posteriorly, rounded posteriorly after a constriction, or even a narrow elongate cylinder supposedly characteristic of latius. Fringe development of the upper lip is also quite variable and seems to be relatively well-developed in larger Sistan fish.

Karaman (1971) described a new genus, Hemigarra, for Tylognathus elegans Günther, 1868 and Discognathus adiscus Annandale, 1919. He places Crossocheilus adiscus as the Sistan subspecies of his Hemigarra elegans (= Hemigrammocapoeta elegans here, q.v.) which is found in Mesopotamia. Karaman (1971) distinguishes the two subspecies by the former having densely arranged papillae on the chin as opposed to sparse papillae. Bianco and Banarescu (1982) and Bănărescu (1986) state that it is not related to Hemigrammocapoeta elegans but is a typical Crossocheilus species.

The type locality of Discognathus adiscus is Sistan by implication, as no locality is given for the holotype in Annandale (1919b). Menon and Yazdani (1968) concur. Distribution is given as "small watercourses and pools in the plains of Seistan" and "Nasratabad, irrigation channel in Consulate garden; pool in the desert 5 miles south of Nasratabad; pools in stream-bed 12 miles north of Nasratabad; channels in the reed-beds of the Hamun-i-Helmand near Lab-i-Baring, and channel leading out of the Hamun 12 miles east of Lab-i-Baring; small watercourse, Lutak, southern Seistan", and one of these is presumably the type locality.

Twenty syntypes of Discognathus adiscus are in the Zoological Survey of India, Calcutta (ZSI F9758/1) (Menon and Yazdani, 1968). Annandale (1919b) cites ZSI 9763/1 as the holotype catalogue number. Three syntypes are in the Zoological Institute, St. Petersburg (ZISP 25411) from "Nasratabad, Seistan, Indian Museum, Dr. Hora" and measure 38.0-43.4 mm standard length. Two syntypes (listed as cotypes) measuring 44.8-45.5 mm standard length from "Jellalabad" with the annotation "Ind. Mus. Ex. F 9762/1" are in the Natural History Museum, London (BM(NH) 1919.8.16:7-8; the outside has 1919.3.16:7-8, incorrectly).

Key characters

The characters of the genus, particularly in the mouth region, serve to identify the only species in Iran.

Morphology

Four short barbels are present, the rostral ones longer than those at the mouth corner. The upper lip covers the upper jaw, is granular or tuberculate and has a marginal fringe, variably developed and most apparent in larger fish. The lower lip is only apparent at the sides and the exposed lower jaw has a granular or tuberculate pad without a free posterior margin but with almost completely free edges.

Dorsal fin with 2-3 unbranched and 8-9 branched rays, anal fin with 2-3 unbranched and 5 branched rays, pectoral fin branched rays 14-17, and pelvic fin branched rays 7-9. Lateral line scales 33-39. Scales may have short dorsal and ventral projections from the margin at about one-third of the scale length from the posterior edge. There is a pelvic axillary scale. Scales have 9-10 radii on the posterior field and are elongate with a notably anterior focus. Radii in large fish are parallel rather than divergent. The anus is 4-5 scales in advance of the anal fin origin. Gill rakers 17-25, small reaching the adjacent or second raker when appressed. Pharyngeal teeth usually 3,3,5-5,3,3 or 2,4,5-5,4,2, depending on how the crowded teeth are counted; major row teeth are usually 5 but may be 4 or 6, middle row teeth are 3 or 4, and minor row teeth 2 or 3, more rarely 1 (this difficulty in assigning teeth to rows is the reason for omitting frequency distributions below). Supernumerary teeth may be present to further confuse counts. The crown of major row teeth are flattened, the anterior tooth may be rounded and some teeth may have a small hooked tip. The gut is very long and complexly coiled. The chromosome number is probably 2n=48 (Klinkhardt et al., 1995).

Iranian fish from Sistan and Baluchestan have the following meristic characters: dorsal fin branched rays 8(81) or 9(1), anal fin branched rays 5(81), pectoral fin branched rays 14(28), 15(37), 16(16), or 17(1), and pelvic fin branched rays 7(3), 8(76), or 9(3). Lateral line scales 33(1), 34(2), 35(12), 36(31), 37(32) or 38(4). Total gill rakers 20-25, but not countable with great accuracy since the smallest rakers are difficult to detect at the ends of the arch. Total vertebrae 34(6), 35(20), 36(9) or 37(1).

Sexual dimorphism

Unknown.

Colour

The back is bluish-grey in Sistan fish or brownish to greenish with irregular spots in other populations and the belly light pink to yellowish-white or silvery-white. Fins are pink and the dorsal and caudal fins have a grey tinge. The flank has a bluish, mid-lateral stripe in Sistan fish and in preserved ones scattered melanophores, or small blotches of less than scale size, or clumps of melanophores centred on upper flank scales and more dispersed on the lower flank. There is a broad stripe along the back mid-line. Fins in preserved fish from Sistan are mostly immaculate except in the larger fish with some melanophores lining rays basally. The caudal fin is distinctive in larger fish from Sistan in having the rays of the lower half of the fin heavily pigmented while the upper half rays are only lightly pigmented. Peritoneum is dark brown to black.

Size

Attains 14.6 cm although the largest fish recorded from Sistan was 93.2 mm standard length.

Distribution

Found in submontane areas of Afghanistan, Pakistan and India as well as eastern and southeastern Iran. The main areas of distribution are Sistan, the Hamun-e Mashkid basin including the Simish River and coastal streams of Makran from the Jagin to middle and upper Nikshahr rivers and the middle and upper Bahu Kalat River including its Sarbaz River reach. (Nikol'skii, 1899; Annandale, 1919b; Berg, 1949; Bianco and Banarescu, 1982; J. Holčík, in litt., 1996; Abdoli, 2000).

Zoogeography

This distribution in Iran marks the western limit for the genus and the relationships of the species lie to the east.

Habitat

Very abundant in small streams, including those with rocky or muddy beds, irrigation ditches, channels in reed beds and pools in Sistan, less common in Baluchestan streams. This species is found in large schools in Sistan in still or slow-flowing water, on the bottom during the day but it may swim at the surface in the evenings. It is common in the smallest permanent water channels but Annandale and Hora (1920) reported it to be in small numbers in the reed beds in winter and these were dead or dying, perhaps because of low oxygen conditions associated with vegetation decay. Large numbers die each year in drying stream beds as salt content increases and the water is fouled by sheep and goats. Tekrival and Rao (1999) report its aquarium preferences as 18-22°C, pH 6.5-7.2, algae as food, not too bright lighting, bottom dwelling with stones, roots and crevices preferred and cave brooding reproduction.

Age and growth

Unknown.

Food

Diet is algae on muddy bottoms. The type subspecies is a bottom feeding herbivore taking more than 90% plant food such as algae, diatoms and macrophytes as well as detritus (Sharma, 1984; Singh and Bahuguna, 1984). Iranian fish contain detritus and some insect remains, possibly as accidental inclusions.

Reproduction

Iranian adult specimens were caught in May in Sistan and show signs of developing reproductive organs suggestive of summer spawning.

Parasites and predators

Jalali et al. (2000) describe two new species of monogenean, Dactylogyrus faridpaki and D. eslamii, from this species in the Bahu Kalat River of Baluchestan.

Economic importance

This species is of no economic importance although Butt (1995) suggests it could be cultured as food and as a forage fish in Pakistan.

Conservation

This species does not appear to be under any major threat as it can survive drying of the Sistan lakes in small ditches and streams.

Further work

The biology of this species needs investigation as does the taxonomic status of Sistan populations.

Sources

Mirza (1972) for colour.

Type material: See above, Discognathus adiscus (ZISP 25411, BM(NH) 1919.8.16:7-8).

Iranian material: CMNFI 1979-0224, 8, 43.6-55.4 mm standard length, Sistan, effluent of Hirmand River (30º53'30"N, 61º27'E); CMNFI 1979-0226, 277, 29.7-78.8 mm standard length, Sistan, pool near Kuh-e Khajeh (30º57'N, 61º17'E); CMNFI 1979-0227, 4, 37.0-48.9 mm standard length, Sistan, naizar at Kuh-e Khajeh (30º57'N, 61º16'E); CMNFI 1979-0228, 1, 42.9 mm standard length, Sistan, ditch 1 km from Zabol (31º02'30"N, 61º31'E); CMNFI 1979-0229, 5, 52.3-93.2 mm standard length, Sistan, ditch 5 km from Zabol (31º03'N, 61º33'E); CMNFI 1979-0230, 1, 48.3 mm standard length, Sistan, Hamun-e Puzak (ca. 31º15'N, ca. 61º42'E); CMNFI 1979-0232, 9, 44.0-65.9 mm standard length, Sistan, ditch 11 km from Zabol (ca. 30º58'30"N, ca, 61º36'E); CMNFI 1979-0234, 17, 40.4-49.3 mm standard length, Sistan, effluent of Hirmand River (30º54'N, 61º40'E); CMNFI 1979-0318, 2, 24.0-27.8 mm standard length, Baluchestan, Sarbaz River at Huvar (26º09'N, 61º27'E); CMNFI 1979-0333, 2, 20.8-21.2 mm standard length, Baluchestan, Mashkid River west of Kuhak (ca. 27º05'N, ca. 63º12'E).

Genus Ctenopharyngodon
Steindachner, 1866

The grass carp genus contains only a single species found in East Asia but widely introduced for food and its ability to digest macrophytes.

This genus is characterised by a rounded body and broad head, the eyes are large and positioned at or above the body axis and often visible from the underside of the head, mouth wide and terminal, no barbels, moderate-sized scales, a complete lateral line, dorsal and anal fins short and lacking spines, branchial membranes attached to the isthmus, short unfused gill rakers, brown to black peritoneum, and pharyngeal teeth in 2 rows with the crowns strongly compressed and serrate and with a longitudinal groove on the grinding surface.

Ctenopharyngodon idella
(Valenciennes, 1844)

Common names

كپور علفخوار (= kapour-e alaf khaar or alaf khoar or kopur 'laf khoar, carp grass-eater or grass-eater), آمور (= amur), سفيد پرورشي (safid parvareshi or mahid safid parvareshi meaning cultured white fish, from a resemblance to mahi safid, i.e. Rutilus frisii kutum).

[grass carp, white amur].

Systematics

Leuciscus idella was originally described from China. A hybrid of this carp and Rutilus frisii has been bred at the Astaneh Ashrafie Fisheries Research Station and named "Samur" (Iranian Fisheries Research and Training Organization Newsletter, 11:6, 1996). See under Rutilus frisii for more information.

Key characters

This species is identified by the eyes being low on the side of the head, the anal fin is far back on the body close to the caudal fin, and pharyngeal teeth have large, parallel grooves on the grinding surface.

Morphology

Lateral line scales 34-47. Scales have a wavy anterior edge, central focus and moderate numbers of anterior and posterior radii. Dorsal fin branched rays 6-8, usually 7, after 3 unbranched rays, anal fin branched rays 7-9, usually 8, after 3 unbranched rays, pectoral fin branched rays 13-20 and pelvic fin branched rays 7-8. Gill rakers number 15-18 and touch the adjacent raker when appressed. Vertebrae 40-47. Pharyngeal teeth are 2,5-5,2, 2,4-5,2, 2,4-4,2, or 1,4-5,2 and are obviously serrated with a longitudinal grooves. The gut is long and complexly coiled. The diploid chromosome number is 48, the triploid 72 (Klinkhardt et al., 1995; Nowruzfashkhami et al., no date). Serum immunoglobulins have been characterised by Soltani et al. (2003).

Sexual dimorphism

Nuptial tubercles are evident on the male head, upper caudal peduncle, dorsal and caudal fins and in particular on the pectoral fins, the first ray of which is thickened, while the female has a distended belly and a swollen and pinkish vent.

Colour

The back is dark, olive to greenish-brown, the flanks are silvery but scales are marked with darker pigment on their posterior margin giving the appearance of a row of spots, and the belly is white to cream-yellow. Scale centres may reflect golden or yellowish tints. Upper scales are outlined with dark pigment to give a cross-hatching effect. The fins are grey-green, or grey to black, except the pelvics which resemble the belly colour. Peritoneum brownish black.

Size

Reputed to attain 1.6 m and about 50 kg in its native range; reports of weights up to 180 kg probably being exaggerations. Reaches 80 cm in the Tadjan River near Sari (A. Abdoli, pers. comm., 1995).

Distribution

The native distribution is in East Asia but it has been introduced to Iranian waters. Also introduced to Afghanistan, Pakistan, and Iraq (Shireman and Smith, 1983).

This species was first introduced in the 1950s according to Armantrout (1980) in the Anzali Mordab for vegetation control, adults surviving to the 1960s but no breeding population was established. Also introduced in 1966 from a hatchery in the Krasnodar region of the former U.S.S.R. and stocked in the Anzali Mordab (Anonymous, 1970b) and in October 1970 50,000 fingerlings from the U.S.S.R. were introduced to the Caspian Sea and Anzali Mordab (Griffiths et al., 1972). Three large fish (80 cm) were caught in January 1971 and believed to be from the October introduction and evidence of good growth although they may have been from an earlier stocking. It is reported from the Siah-Keshim Protected Region of the Anzali Mordab (Riazi, 1996), presumably recently stocked, and is stocked in a variety of reservoirs in the provinces of Gilan and Mazandaran but not as widely as silver carp (Hypophthalmichthys molitrix). It is pen cultured in Gomishan Reservoir, Mazandaran (Madbaygi, 1993b). Grass carp were introduced to Khuzestan in the 1970s to control vegetation in irrigation ditches. In April 1974, 1150 fish were released in the Dez Irrigation Project (Saadati, 1974). It is reported from Mahabad Dam (Abdi, 1999; www.mondialvet99.com, downloaded 31 May 2000), from the Safid River and Anzali Talab (Abbasi et al., 1999), from Lake Zaribar, Kordestan (Abzeeyan, 5(5):III, 1994), the Kor River in Fars (A. Alamdari, in litt., 1997), from the Sistan basin in Hamun Sabari, Hamun Kushk and the canal flowing into Chahnimeh (Ahmadi and Wossughi, 1988; Mansoori, 1994; J. Holčík, in litt., 1996), from the Haft Barm lakes near Shiraz in 1984 although these later disappeared, possibly eaten by introduced Sander lucioperca (Petr, 1987). As escapees from a fish farm, they have been found in Lake Famur. Also recorded from the Gorgan, Tajan and Safid rivers, and the Anzali Mordab (Kiabi et al., 1999), and it is mapped from the Kor, Kerman-Na'in, Hormuz, Dasht-e Lut and Sistan basins without exact localities; the Kashaf River in the Tedzhen River basin; middle reaches of the Atrak River, lower reaches of the Gorgan, Neka, Babol, Heraz and Safid rivers and in the Anzali Mordab, all in the Caspian Sea basin (Abdoli and Naderi, 2009), the middle to lower Talkheh and lower Zarrineh rivers in the Lake Orumiyeh basin, the middle to lower Abhar-Shur and Qom River in the Namak Lake basin, the middle to lower Zayandeh River in the Esfahan basin; the lower Karun and Jarrahi rivers in the Tigris River basin, and the lower Jovein and middle Kal Shur rivers in the Dasht-e Kavir basin (Abdoli, 2000).

It was introduced to the Soviet Caspian Sea in 1970-1974 where small populations became established in the Terek River and the Volga delta and to the Karakum Canal and Kopetdag Reservoir of Turkmenistan near the Iranian border (Baltz, 1991; Shakirova and Sukhanova, 1994; Sal'nikov, 1995; Opuszynski and Shireman, 1995).

Grass carp could establish breeding populations in the large rivers of southern Iran and Iraq if the environment proves favourable and there is enough uninterrupted river flow for eggs to hatch.

Zoogeography

This species is an exotic in Iran and has a native range from the Amur River basin of Siberia south to southern China. It has been widely introduced around the world for vegetation control.

Habitat

The natural habitat is large rivers but this species adapts easily to pond culture. Grass carp can live in the Caspian Sea at salinities of 5-8‰ although a few are found at 10-12‰. They enter rivers to spawn (Abdusamodov, 1986). Temperatures in the range 0-41°C and low oxygen concentrations (0.2 mg/l) are tolerated by this species as is high turbidity. Fry have an upper lethal temperature range of 33-41°C and temperatures greater than 38°C are lethal for adults. pH range is 5.0-9.0. Adults prefer densely vegetated inshore areas with depths of 1-3 m. Adults leave the river after spawning and feed in lakes, reservoirs and on floodplains, returning to the river in autumn to overwinter in deep holes separate from the juveniles. Young hide in vegetation of lakes, reservoirs and floodplains. Juveniles may migrate as much as 1000 km up- or downstream from the original spawning site in their native habitat. Young fish overwinter in deep holes in river beds.

Age and growth

Growth rate in Khuzestan canals was 1.8 g per day while in ponds growth was 6.6 g per day when fed alfalfa during a 5-month growing season from April to September (Saadati, 1974; Behnke, 1975a). Males begin to mature at 4 years and females at 5 years in the Terek River of Dagestan (Abdusamadov, 1986). Maturity is attained at 6-10 years in the Amur River, the native habitat, and as early as 10 months in Malaysia. Life span is over 33 years. Growth rate in this species is perhaps greater than in any other fish. Growth to 1 kg in the first year of life and 2-3 kg per year thereafter in temperate areas is very high; in tropical areas a 20 g fingerling can reach 8.5 kg in 1 year. Rates of 10-22 g per day have been reported in various areas of the world depending on local conditions.

Food

Grass carp are herbivores, except for quite small fish (20 mm total length or less) which consume zooplankton. In Khuzestan, the grass carp prefers to eat Potamogeton spp. and Alisma gramineum to Chara and Cladophora (Saadati, 1974). The grass carp can consume 100-150% of its body weight per day of aquatic vegetation. Peak feeding occurs at 25-30°C but food is taken in the range 15-35°C. Grass carp stocked in the Anzali Mordab and fish farms of the Caspian Sea basin consume fresh Azolla, an introduced fern. Grass carp stocked at 800/ha consume 400-500 kg of Azolla daily gaining 800-1200 g in 5 months. In China this species is known to eat grass, leaves, small fishes, insects and other items in addition to aquatic vegetation or when such vegetation is in low supply. About half the plant food passing through the digestive system is undigested and large quantities of plant material must be eaten to sustain life. This consumption rate is the reason for its success at aquatic vegetation control (Greenfield, 1973). Grass carp overwinter without feeding.

Reproduction

A spawning migration to a large river takes place at about 15-17°C water temperature. The female swims in the centre of the river at the surface accompanied by 2-3 males, they roll and rub their bodies together and often jump out of the water. A male prods the female's body to stimulate egg release and leans closely to one side. Eggs are semi-buoyant and require a slow and steady current to keep them off the bottom (minimum water velocity of 0.23 m/second or more to support them and allow hatching; this is found in large rivers where the eggs hatch as they drift downstream; at 20°C and a not unusual velocity of 1.2 m/second, hatching requires 180 km of river). Temperatures should be above 20°C and preferably 21-25°C, or 26-30°C in another source. At these temperatures hatching takes about 40 hours. Flow rates should be 0.7-1.8 m/sec. Spawning occurs after heavy rain in rising rivers, when turbidity may reduce predatory attacks on the semi-pelagic eggs (Greenfield, 1973). This regime is also required for newly hatched fry and such conditions are rare outside their native habitat. In the Terek River of the Caspian Sea basin, the spawning migration begins in mid-April at water temperatures of 15-17°C and continues until August although numbers begin to decrease from the end of May. Spawning takes place after a sharp rise in water level and current speed. Eggs are first found in the drift in the second week of June and hatch 34-70 hours later depending on temperature. Some larvae reach rice fields and live there until autumn when the fields dry up, some being lost, others migrating. Other larvae are carried into the Caspian Sea where they are sensitive to the prevailing salinity at 1-1.5 days old (Abdusamadov, 1986). Up to 100,000 eggs are laid at one time (Greenfield, 1973) and in the Terek River fecundity reaches 1,230,700 eggs (Abdusamadov, 1986). Absolute fecundity may reach 2 million eggs. Eggs are up to 2.5 mm in diameter before fertilisation and are greyish-blue to bright orange. In water they swell to over 5.3 mm in 2 hours, becoming buoyant in flowing water.

Parasites and predators

Mokhayer (1976b) reports the cestode Bothriocephalus gowkongensis and the acanthocephalan Pomphorhynchus perforator. Red-sore disease is reported from fish pond grass carp in Iran by Razavilar et al. (1981). It is caused by a bacterium Aeromonas hydrophila and treatment was unsuccessful. Mokhayer (1989) records metacercariae of the eye fluke, Diplostomum spathaceum from this species in Iran, which can cause complete blindness and death in commercially important species, as well as shedder scales (sic), Echinochasmus perfoliatus. Jalali and Molnár (1990b) record the monogenean Dactylogyrus lamellatus from this species at fish farms in Iran. Viral haemorrhagic disease has been reported from grass carp in Iran (Iranian Fisheries Research and Training Organization Newsletter, 6:6, 1994; 9:6, 1995). Pond-cultured grass carp were found to be infected by the tapeworm Bothriocephalus, with 70-80 parasites causing intestinal obstruction and lowered haemoglobin, haematocrit and erythrocyte values (Esmaeli and Abbasi, 1996). Esmaeli and Peighan (1997) record an Aeromonas-like bacteria from grass carp in Khuzestan Province. Ebrahimzadeh Mousavi and Khosravi (1999; www.mondialvet99.com, downloaded 31 May 2000) record the toxigenic fungi Aspergillus flavus, Alternaria, Penicillium and Fusarium from this species and the pond water at a fish farm in northern Iran. The crustacean parasite Lernaea elegans is reported from this species in the Mahabad Dam reservoir (Abdi, 1999; www.mondialvet99.com, downloaded 31 May 2000). The intestinal helminth Bothriocephalus gowkongensis was recorded from this species on fish farms in West Azarbayjan Province (Azarvandi et al., 1999). Naem et al. (2002) found the following parasites on the gills of this species from the western branch of the Safid River, namely the protozoan Ichthyophthirius multifilis, a copepod crustacean Lernaea sp., monogenean trematodes Dactylogyrus lamellatus, D. ctenopharyngodonis, and Gyrodactylus sp.. Jalali et al. (2002) and Jalali and Barzegar (2006) record Diplostomum spathaceum and Dactylogyrus lamellatus from this species in Lake Zarivar. Esmaeili et al. (2005) found a Flavobacterium columnaris-like bacterium on grass carp form Khuzestan fish ponds, suspected of either causing a 40% mortality or being a secondary factor in the fish kill. Pazooki et al. (2005) record Ergasilus peregrinus from this species in waterbodies of Zanjan Province. Araghi Soureh and Jalali Jafari (2005) recorded Dactylogyrus lamellatus from this species in the Mahabad River of the Lake Orumiyeh basin. Barzegar and Jalali (2006) report parasites in this species from Kaftar Lake as Trichodina sp., Dactylogyrus lamellatus, Lernaea cyprinacea and Diplostomum spathaceum. Barzegar et al. (2008) record the digenean eye parasites Diplostomum spathaceum and Tylodelphys clavata from this fish. Alishahi et al. (2009) examined moribund grass carp from 20 farms in Khuzestan for bacterial agents but found Aeromonas hydrophila, A. veroni and A. sohria in only 53 of 300 fish, secondary infections and not the cause of mortality. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Ergasilus sp., Ergasilus peregrinus, Lernaea sp. and Lernaea cyprinacea on this species.

Any piscivore will take this species.

Economic importance

This species has been introduced to Iran to control aquatic weeds in drainage and irrigation canals as an alternative to using polluting chemicals or mechanical removal. In some countries at is sought after by anglers. Grass carp may also help to control the snail-carried, human disease schistosomiasis, since the vegetation on which the snails live is severely reduced. They are also a food fish which relies on food sources not available to native fish (few fish consume whole plants). Grass carp consume vegetation at a rate of 100:1, i.e. for every 1 kg increase in grass carp biomass 100 kg of vegetation is consumed. Removal rates may exceed this figure since leaves are bitten off and branches clipped with not all of it being consumed. In the Dez Irrigation Project large amounts of this vegetation were removed daily from screens in test sections. Stocking in the Dez Irrigation Project in Khuzestan showed a removal rate approximately the same as mechanical control (Saadati, 1974; Behnke, 1975a). During a 5 month period the grass carp controlled 250 tons of aquatic vegetation per hectare. The fish preferred plant species which blocked the canals (Potamogeton spp. and Alisma gramineum) rather than those which grew close to the substrate (Chara and Cladophora) and did not interfere with water flow. Shireman and Smith (1983) give details on artificial propagation of this species.

Esmaeilzadeh et al. (2004) studied the nutrient composition and marinade qualities of this fish in Iran and compared them to those for safid mahi (Rutilus frisii) and found them to be preferable according to the organoleptic properties. The marinades could be stored for 6 months at 10ºC.

Fish farming of this species in Sistan was discontinued as its consumption of vegetation was reducing food for other species (www.netiran.com, downloaded 28 February 2005).

Holčík and Oláh (1992) report a catch of 315 kg in the Anzali Mordab in 1990. However Iran acounts for almost all the production of grass carp in the Near East and North Africa (4378 tonnes in 1994) (Food and Agriculture Organization, Fisheries Department, 1996). The aquaculture production in 1995 was 3942 tonnes (Bartley and Rana, 1998b). Grass carp sold for about U.S.$2.00/kg in 1995 (Rana and Bartley, 1998a). Marjan Iran Company was selling 1500-2000 g fish for U.S.$2.10/kg in August 2003 (http://groups.yahoo.com/groups/hilsa/message/25).

The inland waters of Turkmenistan had catches of 23 to 29.7 tonnes for the years 1971-1974 and a catch of 76 tonnes in 1970 when a ban on taking phytophagous fish was lifted.

Greenfield (1973) reviews the advantages and disadvantages of using this species as a weed control agent in the U.S.A. and Charyev (1984) in the Kara-kum Canal in Turkmenistan. Destruction of habitat for fishes and waterfowl, competition with native species and introduction of exotic diseases and parasites are all problems once this fish escapes into a main river suitable for reproduction. Their destruction of plants may interfere with waterfowl management, destroy breeding grounds for other species and facilitate the attacks of predators. Ideally triploids, produced by cold or warm shocks or by hydrostatic pressure on fertilised eggs, should be used initially as they cannot reproduce (Clugston and Shireman, 1987). However the chromosome number of each fish must be checked (by electronically measuring the volume of a red blood cell nucleus) as the process is not 100% effective. Grass carp are reproducing naturally in the Kara-kum Canal, vegetation is controlled, fish stocks have increased and some reduction of mosquitos has been obtained. However the ecosystem has been changed, spawning grounds of commercial species threatened, undesirable species have been introduced accidentally, and reduction in vegetation affects water quality. Grass carp are best used in restricted areas where improved flow and reduced mosquito populations are required but where there is no commercial fishery (Charyev, 1984).

The grass carp has a short gut and about half the plant material eaten is released to enrich the water and promote algal blooms. Oxygen levels and water clarity are reduced. The removal of plants can remove food sources for other fishes, shelter and spawning substrate. Additionally, as noted, the triploid treatment is not always effective and the species can become established.

Iran has had problems with disease outbreaks and poor survival of fingerlings which has led to production problems (Shehadeh, 1997).

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in aquaculture, as food and in textbooks. There are numerous studies on this species as an experimental fish and in relation to aquaculture. Some Iranian studies include Alboughobish and Khaksari Mahabadi (2005) on the histology of the liver and pancreas; Pahn et al. (2005) used electrocardiograms to determine that the anaesthetic ketamine had no marked effect on heart activities; Morovvati et al. (2006) on seasonal changes of pronephros lymphoid tissue; Nahavandi et al. (2006) on the chemiluminescent response to determine the effect of various concentrations of diazinon, an organophosphate, on phagocytosis in order to measure immunity after exposure to this toxin; Pourgholam et al. (2006) and Sharifpour et al. (2006) on the toxicity and histopathological effects of diazinon; Sharifpour et al. (2006) on the sub-lethal effects  of diazinon on various organs; Pourgholam et al. (2006) on the toxicity of diazinon and the effects of sub-lethal concentrations on haematological and biochemical indices; Rezaei et al. (2007) on sensory evaluation and lipid quality of fish stored in ice - good to excellent until the fourth day and good to acceptable to the tenth day; Khajeh et al. (2008) on haematological parameters in cultured fish and found some to be lower than in Mesopotamichthys sharpeyi; etc.

Conservation

No conservation is required for this exotic species. Krasznai (1987) and Petr (1987) give details of fish farms propagating this species in Iran. For example, 10 million were produced in the Safid Rud Fish Farm in 1986. 20 million carp, silver carp and grass carp fingerlings were produced in the Shahid Rajaae Hatchery in Sari for release across Iran in reservoirs and dams (Abzeeyan, Tehran, 4(7):VII, 1993). Feeding and growth studies on this species have also been carried out on this species in the Shaid Rajaee Hatchery (Ahmadi and Rezai, 1998). Experiments on induction of triploidy have been carried out in Iran using cold and heat shocks (M. Hassanzadehsaber, M. Pourkazemi, M. R. Nowruzfashkhami and A. Ghanaatparast (www.meeresschule.com/cgi-bin/abstracts/gastbuch.asp, downloaded 17 January2005).

Further work

Studies on the interactions of this species and native Iranian taxa should be carried out and introductions carefully controlled and monitored.

Sources

Shireman and Smith (1983) give a summary of the biology of this species. There is an extensive literature on herbivorous fishes, a recent book being Opuszynski and Shireman (1995), which has sections on grass carp. Gholipour (1996) has an account in Farsi.

Comparative material: BC65-381, 2, 95.4-98.8 mm standard length, Singapore, fish ponds (no other locality data).

Genus Cyprinion
Heckel, 1843

Scaphiodon Heckel, 1843 has been used for Cyprinion and Capoeta species in Southwest Asia.

Taki (1975) related members of this genus to a common ancestor with Onychostoma Günther, 1896, a Chinese and southeast Asian genus although Li et al. (2008) found this lineage to be unsupported on DNA evidence. Howes (1982) synonymises Semiplotus Bleeker, 1859, a genus found from Nepal to Viet Nam, and Scaphiodonichthys Vinciguerra, 1890, a genus from Indochina, with Cyprinion and refuted Taki's (1975) view using osteological characters, particularly of the jaws. Howes (1982) considers that Cyprinion cannot be defined on any uniquely derived characters. Krupp (1983) considers Howes' revision as unsatisfactory for the reasons that type specimens were not examined, relationships are based on jaw anatomy and other characters are largely excluded, variability of osteological characters within a species are largely unknown, and synapomorphies are not unequivocal. Bănărescu (1992b) and Banarescu and Herzig-Straschil (1995) regard Semiplotus as a distinct genus but probably related to Cyprinion. They comment that Semiplotus differs sharply from Cyprinion s.s. in the absence of barbels, a higher number of branched dorsal fin rays (20 or more), and in a lower number of branched anal fin rays (5 as in most related genera rather than the unusual 7 in Cyprinion). Scaphiodonichthys has 2 pairs of barbels (only 1 in Cyprinion), and 5 branched anal fin rays as well as differing from both Cyprinion and Semiplotus by having the lateral line closer to the ventral margin of the caudal peduncle and divergent rather than parallel striae on the scales. These latter 2 characters justify generic separation of Scaphiodonichthys. Bănărescu (1997) considers Scaphiodonichthys as valid and not a synonym of Cyprinion. Characters used by others to define Cyprinion such as expansion of the proximal part of the pelvic fin rays, interpelvic papillate flaps (Banister and Clarke, 1977) and a naked predorsal ridge (Mirza, 1969) do not occur in all species in this genus. If Semiplotus is included in Cyprinion then several osteological structures, particularly a synarthritic dentary joint, are uniquely derived or synapomorphic.

In the absence of a detailed revision, I have retained species within Cyprinion as the most familiar name in use in Southwest Asia for these fishes. Cyprinion s.s. is found from the Indus River basin west to the Arabian Peninsula and the Tigris-Euphrates basin but excluding northern drainages such as the Lake Orumiyeh, Caspian Sea and Hari River basins and excluding the westernmost edge of Southwest Asia such as the Jordan River basin and coastal drainages of Israel.

The genus Cyprinion is currently under revision by Florian Wicker at the Senckenberg Museum, Frankfurt and the status of the following species may undergo some changes.

Saadati (1977:45) refers to a new and undescribed Cyprinion species from Lar in southern Iran but the fish are Carasobarbus luteus.

A thorough study of the systematics of this genus in Iran depends to some degree on material from other areas which is not readily available, on large series of well-preserved adult specimens, and analyses which demonstrate consistency in characters used to define species. These conditions have not been achieved thus far in any studies undertaken and given the wide distribution and individual variation shown by Cyprinion species an adequate understanding of the species composition is not entirely possible.

This genus is characterised by a moderate sized, compressed body, a thick and blunt snout, an inferior mouth with a straight, crescentic or arched shape and a sharp horny edge to the lower jaw (which may fall off in preserved specimens), 1 pair of small barbels at the mouth corner, the last dorsal fin unbranched ray is thickened and bears weak to strong serrations (highly variable between individuals within a species and not a good character in species definitions), the dorsal fin is long (up to 16 branched rays) and the anal fin short (typically 7 branched rays), a ridge in front of the dorsal fin is formed internally from fused pterygiophores and lacks scales externally, pharyngeal teeth are in 3 rows and are compressed and spoon-shaped, scales large to moderate in size (lateral line counts (31-45), breast and belly scales may be absent (individually variable and not a good character), scale radii are restricted to the posterior field, peritoneum black, and gut very long and coiled (several times body length).

Cyprinion kais
Heckel, 1843


Dez River at Dez Wildlife Refuge, 24 April 2008, courtesy A. Mahjoor Azad

Common names

butak-e dehan kuchek, بوتك (= botak), butak dahan kuchek, butok, لوتك (= lotak), zanbour, زنبور دهان كوچك (= zanbour dahan kuchek).

[bunni saghir, bnaini; kais at Aleppo (= Haleb, Syria), hence the scientific name, all in Arabic; kais kingfish].

Systematics

Cyprinion Cypris Heckel, 1843 is a synonym, being a juvenile with keratinization of the lower jaw incomplete according to Howes (1982), although he did not examine the types. Krupp (1985c) and Banarescu and Herzig-Straschil (1995) agree with this synonymy. Berg (1949) placed C. kais (and C. cypris) in C. macrostomum, as the position of the dorsal fin in relation to the pelvic fins was variable in these fishes and not sufficient to warrant species status as Heckel (1843) stated in describing these species.

The type localities for Cyprinion Kais are "Aleppo" and "Mossul" and for Cyprinion Cypris the "Tigris bei Mossul" (Heckel, 1843b).

The syntypes of C. kais are in the Naturhistorisches Museum Wien comprising 3 fish in NMW 52801 (paralectotypes) and measuring 68.5-97.3 mm standard length, 2 fish in NMW 52802 measuring 120.6-164.3 mm standard length, and 2 fish in NMW 52803 (paralectotypes) measuring 153.4-154.2 mm standard length, the smaller of these being designated as the lectotype by F. Krupp in 1984. Eschmeyer et al. (1996) list possible syntypes in the Rijksmuseum van Natuurlijke Historie, Leiden under RMNH 2485 (2 fish, formerly NMW) and RMNH 2489 (1), and 1 syntype in the Senckenberg Museum Frankfurt (SMF 134, formerly NMW). The catalogue in Vienna lists 5 specimens.

A syntype of C. cypris is in the Senckenberg Museum Frankfurt (SMF 849, formerly NMW) (F. Krupp, pers. comm., 1985). Two syntypes, 63.5-106.2 mm standard length are under SMF 849, the larger one designated as a paralectotype (March 2007). Ten syntypes are in the Naturhistorisches Museum Wien (NMW 52804) measuring 51.2-115.1 mm standard length, the largest being designated as the lectotype (however Banarescu and Herzig-Straschil (1995) give 44.1-110.0 mm standard length for these 10 fish with one at 99.8 mm standard length as lectotype as selected by F. Krupp in 1984). Another specimen, 110.5 mm standard length, may also be a syntype (NMW 52800); and also NMW 59508, a dried specimen (Eschmeyer et al., 1996). The catalogue in Vienna lists 6 fish in alcohol and 1 fish stuffed.

Key characters

Mouth shape is distinctive. It is small and semicircular with a width about the size of the eye diameter and has large lateral lobes (= lower lips)(Kafuku, 1969). The cartilaginous sheath is thickened between the corners of the mouth and is rounded posteriorly with a distinct margin. The cartilage can form a tooth-like structure protruding anteriorly from the lower lip. The mouth in C. macrostomum is wider, arched and lacks the lateral lobes (see also illustrations in Kafuku (1969), Krupp (1985c) and Banarescu and Herzig-Straschil (1995)). These latter authors have the width of the mouth opening as only 13.5-22.0% of the head length (22.0-27.0% in C. macrostomum) for adult fish and the height of the arch or mouth opening (a line perpendicular from a line between the mouth corners to the tip of the lower jaw) 48-80% of the mouth width (29-47% in C. macrostomum), i.e. the mouth is narrower and more arched in C. kais. On this character, therefore, the two species can be distinguished as adults but there is potential for confusion in young fish. A single specimen identified as C. kais on the basis of mouth shape from the Dalaki River of Iran had values of 23.2% and 47.4% which are arguably C. macrostomum values. This specimen has a protruding tooth-like edge to the lower jaw in a u-shaped mouth with well-developed lips posterior to the "tooth".

The intestine is shorter and less complexly coiled in this species and the mean number of gill rakers is less in contrast to C. macrostomum (Kafuku, 1969). The back is higher and more curved, the eyes are larger and the anal fin is more posterior, in addition to the mouth shape (Heckel, 1843b). The dorsal fin origin arises over that of the pelvic fins (Heckel, 1846-1849a). The edge of the dorsal fin is more notched in C. kais than in C. macrostomum (the length of the fourth branched ray is 48-62% of the length of the first ray as opposed to 55-79% in C. macrostomum, with extreme values overlapping, according to Banarescu and Herzig-Straschil (1995)).

The form of the pharyngeal teeth is different from C. macrostomum (see Krupp (1985c) for illustrations where kais has hooked tips and macrostomum does not), there are fewer gill rakers (8-12 on the lower arch in kais, 12-16 in macrostomum), on average there are fewer dorsal fin rays, the last unbranched dorsal fin ray is longer, and interorbital width is smaller. However sample sizes in some studies are small (in Kafuku (1969) only 5 fish of each species were examined), morphometric characters are notoriously size-dependent, gill raker counts are also size dependent, and even pharyngeal tooth form varies with age (small macrostomum have hooked tips). C. kais may well be a good species but a wide-ranging comparison of adults and young and of localities is needed and material from Iran is scarce or equivocal. Further discussion is under C. macrostomum.

Morphology

Dorsal fin with 4 unbranched and 12-16 branched rays, anal fin with 3 unbranched and 7 branched rays. The dorsal fin has the last unbranched developed as a spine with strong teeth except at the extreme tip which is thin and flexible. Pectoral fin with 14-18 branched rays, and pelvic fin with 8-9 branched rays. Lateral line with 36-43 scales. The belly is scaled. There is a well-developed pelvic axillary scale. Scales have a subcentral anterior focus, fine circuli, few posterior radii and no or very few anterior radii. Total gill rakers 10-15, short and reaching the raker below when appressed. Rakers are absent on the anterior arch where there are only tubercles. Pharyngeal teeth 2,3,4-4,3,2, with variants 2,3,5-5,3,2 and 2,3,5-4,3,2, spoon-shaped with a small hook at the tip.

Meristics for Iranian material: dorsal fin branched rays 12(2), 13(2), 14(1) or 15(1); branched anal fin rays 7 (6); branched pectoral fin rays 14(1), 15(2), 16(2) or 17(1); branched pelvic rays 8(4) or 9(2); lateral line scales 38(5) or 39(1); and total gill rakers 12(2), 13(3) or 14(1).

Sexual dimorphism

Tuberculation in a 103.5 mm standard length specimen consisted of ca. 20 tubercles restricted to the area over the lachrymal bone. A specimen 147.5 mm standard length had small to minute tubercles in front of the eye, under the eye, on the mid-preoperculum and on the mid-operculum. Curiously the individual small tubercles on the operculum were connected by thin lines of horny tissue.

Colour

Overall colour is silvery to yellowish-white with the back grey-brown and the lower surfaces a lemon yellow. The lower jaw margin is a glossy yellow. The fish shown above may represent a spawning colouration, not seen in all specimens. The pelvic fins are a bright orange-red, the pectorals paler. Some fish have a less strong colour in the pelvic than in the anal fin. The anal fin is yellow, to orange or greenish, distally black and anteriorly most orange. The caudal fin has light orange to greenish tints. The dorsal fin is black with a yellow-tinged base becoming anteriorly reddish. In preserved fish, there is some concentration of pigment above and below each lateral line pore, scales on the back and upper flank are outlined with pigment, and there is some concentration of pigment into a few to moderate number of diffuse spots on the uppermost flank and back midline. The leading edge of the dorsal fin is very dark (but may be light), dorsal fin membranes are dark, anal fin membranes also dark but to a lesser extent, and the caudal, pectoral and pelvic fins have pigment lining the rays. Peritoneum black.

Size

Attains 21.5 cm total length, or to 25.0 cm total length in Iraq (Al-Rudainy, 2008).

Distribution

This species is found in the Tigris-Euphrates and Quwaiq basins. Abdoli (2000) maps the Jarrahi, Karun, middle to lower Dez, and Karkheh up to the Simarreh rivers of the Tigris River basin. It is also found in the Gulf basin, although rare, and specimens from the sugar cane fields of Khuzestan were seen in 2000 (personal observations, B. W. Coad).

Zoogeography

Zoogeographical comments are under the genus above.

Habitat

This species is recorded from a variety of habitats as listed above and is also known to inhabit canals but nothing is known of its environmental requirements.

Age and growth

Unknown.

Food

Gut contents are filamentous algae in the one specimen examined. Diet may be similar to Cyprinion macrostomus. Al-Rudainy (2008) gives aquatic insects and detritus for Iraq. Curiously, the mouth structure resembles that of the unrelated cutlips minnow, Exoglossum maxillingua (Le Sueur, 1817), from North America. This species feeds on insect larvae, with some molluscs and worms. Food is scraped from the bottom or poked out of crevices using the shovel-like lower jaw. Sand is also taken in and spat out, presumably after food items are extracted. The cutlips also picks out the eyes of other fishes in confined areas (Coad et al., 1995).

Reproduction

Generally unknown. Ünlü (2006) gives age at first maturity as 2 years in the Turkish Tigris River with spawning over sand, stones and gravel in May-June.

Parasites and predators

None reported from Iran.

Economic importance

None.

Conservation

This minnow appears to be rare, or at least is rarely collected, in Iran. Cyprinion macrostomum is much more common and is taken in most seine hauls in streams and rivers. The distribution and population numbers are unknown. Endangered in Turkey (Fricke et al., 2007).

Further work

The biology of this species needs to be investigated and the use of the peculiar jaw structure ascertained. Its great rarity, at least in Iran, leads to the suspicion that it may be a developmental anomaly of Cyprinion macrostomum - the few specimens at hand don't permit a detailed study of characters other than the strikingly different jaw (see comments under C. macrostomum and also above). Development of pharyngeal teeth, gill raker numbers, complexity of gut coils and morphometric characters are all size dependent and show individual and populational variations not analysable here.

Sources

Type material: See above, Cyprinion kais (NMW 52801, 52802 and 52803) and C. cypris (NMW 52804).

Iranian material: CMNFI 1993-0141, 1, 66.3 mm standard length, Bushehr, Dalaki River (29º28'N, 51º15'E); CMNFI 2008-0169, 5, 80.4-98.2 mm standard length, Khuzestan, irrigation ditch in sugar cane fields (31º58'42"N, 48º31'07"E); ZSM 25715, 2, 34.1-65.3 mm standard length, Khuzestan, Dez River at Harmaleh (31º57'N, 48º34'E).

Comparative material: BM(NH) 1920.3.3:50, 1, 83.6 mm standard length, Iraq, Basrah (30º30'N, 47º47'E); BM(NH) 1920.3.3:94-115, 40, 65.3-92.4 mm standard length, Iraq, Basrah (30º30'N, 47º47'E); BM(NH) 1931.12.21:3, 1, 129.8 mm standard length, Iraq, Mosul (36º20'N, 43º08'E); BM(NH) 1974.2.22:115-120, 5, 90.6-147.9 mm standard length, Iraq, Mosul (36º20'N, 43º08'E); BM(NH) 1974.2.22:1105, 1, 115.6 mm standard length, Iraq, Mosul (36º20'N, 43º08'E); BM(NH) 1974.2.22:1106, 1, 101.4 mm standard length, Iraq, Fao (29º58'N, 48º29'E); BM(NH) 1974.2.22:1214-1255 (in part), Iraq, Khalis (33º49'N, 44º32'E); BM(NH) 1984.4.18:30, 63.4 mm standard length, Iraq, Kut Hiwa (no other locality data); FMNH 51229, 1, 103.5 mm standard length, Iraq, Diyala River, 12 miles east of Baghdad (no other locality data); FMNH 51230, 6, 42.9-60.5 mm standard length, Iraq, Diyala River, 12 miles east of Baghdad (no other locality data); FMNH 51231, 2, 64.0-64.8 mm standard length, Iraq, Diyala River, 12 miles east of Baghdad (no other locality data); uncatalogued, 5, 49.1-66.7 mm standard length, Iraq, Shatt al Arab (no other locality data); uncatalogued, 1, 107.2 mm standard length, Turkey, Euphrates River 20 km west of Erzurum (ca. 41º03'N, ca. 39º55'E).

Cyprinion macrostomum
Heckel, 1843

Common names

بوتك (= botak); butok; لوتك (= lotak); butak-e dehan (or dahan) buzorg in Khuzestan; galuk (Mokhayer (1981c); kapour; zanbour (= bee) in Khuzestan and Boyer Ahmadi-ye Sardsir va Kohkiluyeh provinces; زنبور دهان بزرگ (= zanbour dahan bozorg); ?tumbuek (= hunting horn, possible name from Heckel (1843b)).

[hmarriya sefra or himriya sefra, surrah masih, bunni kaper, dunbuk kabir al-fam, benayne; kais at Aleppo (= Haleb, Syria) but see above species (Heckel, 1843b); dombok or dumbek at Mosul (= solid or compact flesh, a good source of food, according to Heckel (1843b)); all preceding in Arabic; large-mouthed barb, Tigris kingfish].

Systematics

Originally spelt macrostomus but correctly macrostomum (Berg, 1949). Cyprinion neglectus Heckel, 1849 from the "Tigris bei Mossul" is a synonym (Krupp, 1985c; Banarescu and Herzig-Straschil, 1995). Howes (1982) considered that Cyprinion tenuiradius (q.v.) was only a "variant" of this species but did not examine any material. Berg (1949) places C. kais (q.v.) in the synonymy of this species along with C. cypris (see C. kais).

The type locality of Cyprinion macrostomus is given by Heckel (1843b) as "Aleppo" and "Mossul". Krupp (1985c) lists 5 syntypes from Aleppo, 81-133 mm standard length in the Naturhistorisches Museum Wien (NMW 52805), the largest being selected as the lectotype (hence Aleppo is the type locality as designated by the publication of Banarescu and Herzig-Straschil (1995)). One syntype from Aleppo, 83 mm standard length, is in the Senckenberg Museum Frankfurt (SMF 70, formerly NMW; Eschmeyer et al. (1996) give SMF 870) and 4 syntypes from Mosul, 58-124 mm standard length are in the Naturhistorisches Museum Wien (NMW 52806). My measurements are 82.1-135.0 mm standard length for NMW 52805 and 59.1-126.2 mm standard length for NMW 52806. Another syntype is a dried specimen NMW 52503, and the Rijksmuseum van Natuurlijke Historie, Leiden has 1 syntype under RMNH 2487, formerly NMW) and 1 syntype under RMNH 2488, formerly NMW). The catalogue in Vienna lists 4 specimens.

Seven syntypes of Cyprinion neglectus from Mosul measure 54-131 mm standard length (NMW 52807), the largest being selected as the lectotype (Krupp, 1985c). My measurements are 53.3-131.9 mm standard length (Banarescu and Herzig-Straschil (1995) have 53.1-128.2 mm standard length). All material was collected by Th. Kotschy in 1842 for Aleppo and 1843 for Mosul. The catalogue in Vienna lists only 2 specimens under this name.

?Check lengths against data sheets

Key characters

Distinguished from C. kais by mouth and dorsal fin ray characters as described under that species, by having more gill rakers and a longer and more coiled intestine (Kafuku, 1969). The dorsal fin origin is in front of that of the pelvic fins (Heckel, 1846-1849a). See discussion under C. tenuiradius for distinction from that taxon.

Morphology

Dorsal fin with 4 unbranched and 12-17 branched rays (usually 14-15 according to Banarescu and Herzig-Straschil (1995) but 77% of fish in Iran are 13-14, see below). The last dorsal fin unbranched ray is strong and serrated to the tip. The anal fin has 3 unbranched and 6-7, usually 7, branched rays. In Iranian specimens, 96.1% of 127 fish have 7 rays, the remainder 6 rays. Pectoral fin branched rays are 12-17 and pelvic fin branched rays 7-9, usually 8. Lateral line scales 33-45 (usually 41-44 according to Banarescu and Herzig-Straschil, 1995) but a broader range in Iran, see below). The breast is covered with scales. The pelvic axillary scale is very elongate. Scales are squarish, being deeper than long, often with parallel dorsal and ventral margins (or rounded margins). The anterior margin has a marked central protuberance and the posterior margin is rounded. Radii are numerous on the posterior field and circuli are fine and numerous. The posterior field circuli break into "bubbles". The focus is subcentral anterior. Gill rakers 16-17, on the lower arm 12-16, in the literature but a much wider range in total rakers in Iran (see below). Rakers are short and only touch the raker below or a little further when appressed. Pharyngeal teeth 2,3,5-5,3,2, 2,3,4-4,3,2, and variations on 4 or 5 main row teeth. Teeth are spatulate with broad, flattened crowns. The tips of teeth are slightly hooked in small fish. The most anterior tooth in the main row may be very small or absent (or incompletely ossified and hard to distinguish). The gut is very elongate with complex coils. In small fish, the upper lip is not covered with a fold of the snout as in large fish. Also the gut is not as coiled in young fish as in adults. Chromosome number is 2n=48 (Ünlü et al., 1997).

Meristics for Iranian fish from the Tigris River basin: branched dorsal fin rays 12(4), 13(43), 14(52), 15(26) or 16(3)(mean = 13.9, S.D. = 0.861); branched pectoral fin rays 14(3), 15(44), 16(57) or 17(25)(mean = 15.8, S.D. = 0.771); branched pelvic fin rays 7(7), 8(121) or 9(1)( mean = 8.0, S.D. = 0.246); lateral line scales 33(3), 34(1), 35(12), 36(11), 37(3), 38(11), 39(29), 40(31), 41(25), 42(2) or 45(1)(mean = 38.8, S.D. = 2.211); total gill rakers 13(3), 14(8), 15(15), 16(23), 17(15), 18(24), 19(17), 20(14) or 21(6) (mean = 17.3, S.D. = 2.022); pharyngeal teeth 2,3,5-5,3,2(17), 2,3,4-5,3,2(8), 2,3,5-4,3,2(3), or 2,3,4-4,3,2(2); and total vertebrae ?.

The mouth is usually transverse or slightly arched and usually has a horny covering. Small fish have a crescentic mouth. A wide range of mouth arching is seen in fish of varying sizes and even in fish of the same size and locality of capture. Banarescu and Herzig-Straschil (1995) note that the syntypes of Cyprinion neglectus have a mouth arch which is more curved and not as wide, somewhat intermediate between C. macrostomum and C. kais, being closer to the former. This variation is attributed to the material possibly being from some tributary of the Tigris River, or from isolated ponds, where introgression with C. kais took place. It may well be that variation in mouth shape is more marked than limited sample sizes would indicate. Certainly in smaller fish, e.g. in 20 specimens of C. macrostomum (38.5-54.0 mm SL) examined by me from Iran, values for mouth width and depth as measured in Banarescu and Herzig-Straschil (1995) are not as clear cut and there is a variable developmental gradient in mouth shape. Mouth "height" as a % of width was 29.2-53.8 and width as % of head length was 22.1-36.6. Banarescu and Herzig-Straschil (1995) give "height" as 19-31% of width and width as 26-44% head length for macrostomum and 48-80% and 13.5-22.0% respectively for kais. Large macrostomum and kais (>100 mm SL) can be distinguished on mouth shape but not smaller specimens which bridge the gap between the two species. The possibility that kais is a developmental anomaly of macrostomum, retaining juvenile features, should be investigated.

Sexual dimorphism

Mature males have large tubercles on the snout in a broad band below the nostril level, extending back under the eye and breaking up into a few tubercles on the operculum. There is a large tubercle between the nostril and the eye. Fine tubercles are scattered over the top of the head. Three tubercles are found in rows on the first branched pectoral fin ray and very strong tubercles line each anal fin branched ray in single file. The anterior pelvic fin rays have the occasional 1-2 tubercles or a row of tubercles. Dorsal and caudal fin rays have fine tubercles, much smaller than those on the anal fin. Mid and posterior flank scales have 1-3 small tubercles, variably arranged on the exposed scale.

Colour

The back is bluish-grey to bluish-black or brown, flanks silvery or silvery-yellow and the belly whitish with silvery tints. The upper head is light brown. Scales are outlined with dark pigment and the anterior exposed scale base is darkened. The cleithrum area is pink or orange in some fish with pink or orange spots on up to 5 rows of flank scales but mostly along the anterior lateral line. Fish from a saline stream in Khuzestan had a pale-pink cleithrum and lateral line spots. There is a reddish-yellow spot at the base of the pectoral and pelvic fins. The pectoral, pelvic, anal and caudal fins are yellowish to pinkish or orange proximally and blackish distally. The dorsal fin has a narrow, yellow stripe at the base and the rest of the fin is black. The cartilaginous lower jaw is reddish-yellow to orange. The eye is slightly yellow. Small live fish are silvery overall with a white belly and olive back, the pectoral and pelvic fins slightly orange-yellow and other fins greyish although all fins may be hyaline. The peritoneum is black.

Small preserved fish have an indistinct blotch at the caudal fin base and a similar blotch on the back at the base of the spine in the dorsal fin. In very small fish these blotches are more distinct and there are 4-7 irregular blotches on the mid-flank above the lateral line and 3 blotches at the dorsal fin base. Development of blotches is individually variable, some fish being almost immaculate while in others the blotches extend vertically as bars as far as the back.

Size

Reaches 19.3 cm standard length (Krupp, 1985c).

Distribution

Found in the Orontes, (= Asi), Quwayq and Tigris-Euphrates basins. In Iran, it is found in the Tigris River basin including the Hawr Al Azim, Khersan, Jarrahi and Marun rivers (Berg, 1949; Abdoli, 2000) and the northern Gulf basin in the Shapur, Dalaki and Helleh rivers (Gh. Izadpanahi, pers. comm., 1995), the Zohreh River and possibly Lake Famur - some may be C. tenuiradius. Vossoughi (1998) reports this species from the western Hamun-e Jaz Murian basin based on a fishes with 13-15 branched dorsal fin rays, much higher than for C. watsoni, the taxon to be expected in this area.

Zoogeography

Zoogeographical comments are under the genus above.

Habitat

Known from a variety of habitats such as rivers, streams, reservoirs and ponds, as well as canals and gravel pits. Al-Habbib and Al-Habbib (1979) have demonstrated experimentally for a sample from "Nawaran Spring" north of Mosul, Iraq that this species can survive temperatures up to about 37°C. Akpinar and Aksoylar (1989) and Akpinar (1999) report this species from the Kangal Thermal Spring, Sivas, Turkey at a constant temperature of 35°C. This is the commonest species in catches in southwestern Iran, followed by Garra rufa. In areas under human influence in Lorestan, such as the lower reaches of rivers and near cities, it exceeds 80% in numbers in catches.

Age and growth

Maximum age reported for a population in the "Al-Nibaey" Lakes near Baghdad is 7+ years. Growth is slow and there is no difference in growth between males and females, although the habitat is not considered ideal for these fishes. Females tend to be slightly heavier than males of the same length especially in older fish. The length-weight relationship was W = 0.027 L2.67 (r = 0.78) for both sexes, W = 0.028 L2.65 (r = 0.90) for males and W = 0.020 L2.78 (r = 0.93) for females. Maturity is attained at 10.0-11.1 cm, corresponding to age group 2 (Allouse et al., 1989). The length-weight equation for commercially caught fish in the Tigris River was log W = 2.884 log L-4.623, condition factor was 1.15-1.47 (mean 1.28) and fish were immature up to age 2+ (Al-Nasiri, 1991). Haematology of this species from Sarao Subhana Agha near Sulaymaniyah was examined by Al-Mehdi and Khan (1984).

Food

Major food items in the Baghdad study are of plant origin with occasionally some chironomid larvae, copepods and cladocerans. Khan (1988) found for fish from near Sulaimaniyah, Iraq that diatoms and decayed organic matter are the main foods, with some green algae. Zooplankton are thought to be accidental food items. Guts contain mud and sand, evidence of a bottom feeding habit. Feeding increases at the start of the breeding season. The horny lower jaw covering is used to scrape algal food off hard bottom objects.

Reproduction

Near Baghdad, most fish are mature by April, the gonads occupying about one-third of the body cavity. Ovaries are orange to yellowish and testes milky white. Spawning occurs principally in May and June, with some in early July, but by July most fish are spent. Al-Rudainy (2008) gives a spawning season of May and June in Iraq on gravel beds in shallow water with fast current. Maturity is attained there at 2-3 years, 15 cm length and 50 g weight.

Iranian material shows minute but developing eggs in a 71.3 mm standard length fish caught on 31 January and specimens caught on 5 July have eggs 1.4 mm in diameter. The 31 January fish has tubercles on the snout and anal rays so tubercles develop quite early and in small fish. A fish caught on 20 September also shows tubercles around the snout. Small fish caught in January about 20 mm SL are presumably the young from the previous season and so show slow growth or are evidence of a prolonged or late spawning season.

Parasites and predators

Gussev et al. (1993a) describe a new species of monogenean from C. macrostomum in the Karun River, Dactylogyrus cyprinioni, and Jalali (1992) a new species of monogenean, Dogielius molnari, in the Dez River, both in Khuzestan. Jalali et al. (1995) describe a new species of monogenean, Dactylogyrus pallicirrus, from fish taken in the Dez River near Ahvaz.

Economic importance

Al-Mehdi and Khan (1984) report this species to be important in riverine and culture fisheries in northern Iraq. Ündar et al. (1990) identify this species and Garra rufa as the "doctor fish" of the Kangal hot spring in Turkey (Timur et al., 1983; Warwick and Warwick, 1989; Kürkçüoğlu and Öz, 1989; and various newspaper and television reports). High water temperatures reduce the amount of plankton available as fish food and the fish nibble away infected skin of humans who bathe in these waters. The fish is known as "striker" (and Garra rufa as "licker") from its behaviour in the spa pools. The healing properties are linked to the high level of selenium (1.3 p.p.m.) in the water, selenium being beneficial in some skin diseases, and possibly to UV light. The fish facilitate the action of the selenium and UV light by softening and clearing away psoriatic plaque and scale, exposing the lesions to the water and sunlight. However, some lesions are made worse and the fish can cause some new ones.

Conservation

This species is widely distributed in southern areas, particularly Khuzestan, and does not appear to be under threat other than that suffered by all species by pollution and water abstraction. Endangered in Turkey (Fricke et al., 2007).

Further work

See comments above on the need for further work to distinguish this species from C. kais, especially when young and below for distinction from C. tenuiradius.

Sources

Type material: See above, Cyprinion macrostomum (NMW 52805, 52806), C. neglectus (NMW 52807).

Iranian material: Tigris basin: and presumably macrostomum CMNFI 1979-0268, 13, 92.2-122.4 mm standard length, Lorestan, Dez or Karkheh drainage between Nowqan and Khorramabad (no other locality data); CMNFI 1979-0269, 4, 104.7-110.6 mm standard length, Lorestan, Dez or Karkheh drainage between Nowqan and Khorramabad (no other locality data); CMNFI 1979-0270, 10, 85.5-122.4 mm standard length, Lorestan, Kashkan River drainage (33º26'N, 48º19'E); CMNFI 1979-0271, 3, 100.7-144.8 mm standard length, Lorestan, Kashkan River drainage (33º39'N, 48º32'30"E); CMNFI 1979-0273, 9, ? mm standard length, Lorestan, Kashkan River drainage (33º26'N, 48º19'E); CMNFI 1979-0274, 14, ? mm standard length, Lorestan, Kashkan River drainage (33º27'N, 48º11'E); CMNFI 1979-0275, 2, 142.4-165.0 mm standard length, Lorestan, Kashkan River drainage (33º25'N, 47º58'E); CMNFI 1979-0278, 4, 93.5-114.1 mm standard length, Lorestan, Kashkan River drainage (33º34'N, 48º01'E); CMNFI 1979-0279, 9, 100.3-149.4 mm standard length, Lorestan, Khorramabad River (33º37'N, 48º18'E); CMNFI 1979-0283, 5, 93.0-144.0 mm standard length, Kermanshahan, Qareh Su drainage (34º21'N, 47º07'E); CMNFI 1979-0287, 1, 112.6 mm standard length, Kermanshahan, Chashmeh Javari 2 km from Ravansar (ca. 34º42'N, ca. 46º40'E); CMNFI 1979-0288, 1, 94.3 mm standard length, Ilam and Poshtkuh, Gangir River at Juy Zar (33º50'N, 46º18'E); CMNFI 1979-0289, 4, ? mm standard length, Kermanshahan, Diyala River drainage (34º28'N, 45º52'E); CMNFI 1979-0290, 11, 49.3-133.0 mm standard length, Kermanshahan, Diyala River drainage at Qasr-e Shirin (34º31'N, 45º35'E); CMNFI 1979-0291, 15, ? mm standard length, Kermanshahan, Diyala River drainage (34º24'N, 45º37'E); CMNFI 1979-0350, 18, ? mm standard length, Khuzestan, Marun River near Marun (30º39'30"N, 50º02'E); CMNFI 1979-0355, 1, ? mm standard length, Khuzestan, stream tributary to Karun River at Salmaneh (30º35'N, 48º22'E); CMNFI 1979-0356, 1, ? mm standard length, Khuzestan, stream at Hoveyzeh (31º27'N, 48º04'E); CMNFI 1979-0360, 2, ? mm standard length, Khuzestan, canal branch of Karkheh River (31º40'N, 48º35'E); CMNFI 1979-0361, 3, ? mm standard length, Khuzestan, jube in Karkheh River drainage (31º42'N, 48º33'E); CMNFI 1979-0363, 1, ? mm standard length, Khuzestan, Karkheh River (31º52'N, 48º20'E); CMNFI 1979-0364, 2, ? mm standard length, Khuzestan, river at Abdolkhan (31º52'30"N< 48º20'30"E); CMNFI 1979-0365, 24, ? mm standard length, Khuzestan, stream in Doveyrich River drainage (32º25'N, 47º36'30'E); CMNFI 1979-0366, 16, ? mm standard length, Khuzestan, stream west of Dehloran (32º45'30"N, 47º05'30"E); ID? CMNFI 1979-0367, 2, ? mm standard length, Khuzestan, Meymeh River 11 km north of Dehloran (32º44'30"N, 47º09'30"E) ID? CMNFI 1979-0368, 12, ? mm standard length, Khuzestan, Karkheh River (32º24'30"N, 48º09'E); CMNFI 1979-0371, 1, ? mm standard length, Khuzestan, stream in Karkheh River drainage (32º05'N, 48º19'E); CMNFI 1979-0373, 12, ? mm standard length, Khuzestan, Bala River north of Andimeshk (32º35'N, 48º17'E); CMNFI 1979-0374, 46, ? mm standard length, Khuzestan, stream tributary to Bala River (32º40'N, 48º15'E); CMNFI 1979-0376, 9, ? mm standard length, Khuzestan, river tributary to Karkheh River (32º48'30"N, 48º04'30"E); CMNFI 1979-0378, 10, ? mm standard length, Khuzestan, stream tributary to Karkheh River (ca. 32º48'N, ca. 48º04'E); CMNFI 1979-0379, 11, ? mm standard length, Khuzestan, Dez River (32º12'N, 48º27'E); CMNFI 1979-0380, 5, ? mm standard length, Khuzestan, stream tributary to Dez River (ca. 32º10'N, ca. 48º35'E); CMNFI 1979-0381, 28, ? mm standard length, Khuzestan, stream 40 km west of Shushtar (ca. 32º10'N, ca. 48º35'E); CMNFI 1979-0382, 67, ? mm standard length, Khuzestan, Karun River at Shushtar (32º03'N, 48º51'E); CMNFI 1979-0383, 1, ? mm standard length, Khuzestan, stream in Ab-e Shur drainage (31º59'30"N, 49º06'E); CMNFI 1979-0384, 7, 86.3-152.2 mm standard length, Khuzestan, Ab-e Shur drainage (32º00'N, 49º07'E); CMNFI 1979-0386, 4, ? mm standard length, Khuzestan, stream 21 km from Haft Gel (ca. 31º34'N, ca. 49º23'E); CMNFI 1979-0387, 6, ? mm standard length, Khuzestan, stream 12 km from Haft Gel, Jarrahi River drainage (31º25'N, 49º38'E); CMNFI 1979-0388, 2, ? mm standard length, Khuzestan, Zard River (31º19'N, 49º44'E); CMNFI 1979-0390B, 23, 36.2-156.2 mm standard length, Khuzestan, stream 3km south of Bagh-e Malek (31º29'N, 49º54'30"E); CMNFI 1979-0391, 1, 154.5 mm standard length, Khuzestan, stream in Marun River drainage (31º28'N, 49º51'E); CMNFI 1979-0392, 5, ? mm standard length, Khuzestan, Zard River (ca. 31º32'N, ca. 49º48'E); CMNFI 1979-0393, 2, 96.9-116.6 mm standard length, Khuzestan, Jarrahi River drainage (31º18'N, 49º37'E); CMNFI 1979-0394, 1, 130.2 mm standard length, Khuzestan, stream in Marun River drainage (31º01'N, 49º45'E); CMNFI 1979-0395, 4, ? mm standard length, Khuzestan, stream in Marun River drainage (ca. 30º57'N, ca. 49º51'E); CMNFI 1979-0396, 1, ? mm standard length, Khuzestan, Kheyrabad River (30º32'N, 50º23'30"E); ID? CMNFI 1979-0398, 23, ? mm standard length, Boyer Ahmadi-ye Sardsir va Kohkiluyeh, stream in Zohreh River drainage (30º24'30"N, 50º37'30"E); ID? CMNFI 1979-0399, 7, ? mm standard length, Fars, stream in Zohreh River drainage (30º19'30"N, 51º15'E); CMNFI 1991-0153, 1, 171.3 mm standard length, Khuzestan, Zohreh River (no other locality data); CMNFI 1991-0154, 1, 109.9 mm standard length, Khuzestan, Hawr al-Azim (ca. 31º45'N, ca. 47º55'E); CMNFI 1993-0128, 1, 110.7 mm standard length, Kermanshahan, Sarab-e Sabz 'Ali Khan (34º25'N, 46º32'E); CMNFI 1993-0149, 1, 121.7 mm standard length, Khuzestan, Karun River (no other locality data); CMNFI 2007-0111, 6, 24.7-173.8 mm standard length, Kermanshahan, Alvand River near Sar-e Pol-e Zahab (ca. 34º36'N, ca. 45º56'E); CMNFI 2007-0112, 6, 46.5-118.8 mm standard length, Kermanshahan, Kerend River basin near Shahabad-e Gharb (ca. 34º06'N, ca. 46º30'E; CMNFI 2007-0113, 1, 122.1 mm standard length, Kermanshahan, Razavar River, Qareh Su tributary (ca. 34º25'N, ca. 47º01'E); CMNFI 2007-0115, 6, 59.7-154.8 mm standard length, Kermanshahan, Qareh Su basin (ca. 34º34'N, ca. 46º47'E); CMNFI 2007-0116, 12, ?-93.0 mm standard length, Kermanshahan, Gav Masiab basin west of Sahneh (ca. 34º28'N, ca. 47º36'E); CMNFI 2007-0117, 1, ? mm standard length, Kermanshahan, Gav Masiab basin near Sahneh (ca. 34º24'N, ca. 47º40'E); BM(NH) 1980.8.28:1, 1, 90.3 mm standard length, Khuzestan, Dezful (32º23'N, 48º24'E); BWC95-20, 14, ? mm standard length, Khuzestan, Rud Zard at Rud Zard (31º22'N, 49º43'E); Gulf fish:- ? tenuiradius CMNFI 1979-0020, 56, ?, mm standard length, Fars, Mand River outside Kavar (29º11'N, 52º41'E); CMNFI 1979-0054, 14, 37.4-64.1 mm standard length, Fars, Shur River tributary (ca. 28º58-29º03'N, ca, 52º34-35'E); CMNFI 1979-0075, 123, 21.3-142.4 mm standard length, Fars, Mand River at Pol-e Kavar (29º11'N, 52º41'E); CMNFI 1979-0109, 5, 63.2-100.2 mm standard length, Fars, Mand River at Shahr-e Khafr (28º56'N, 53º14'E); CMNFI 1979-0128, 7, 19.2-103.8 mm standard length, Shur River (28º51'N, 52º31'E); CMNFI 1979-0131, 19, 16.4-41.7 mm standard length, Fars, Ab-Arak River (28º38'N, 52º49'E); CMNFI 1979-0132, 72?, 15.2-100.1 mm standard length, Fars, Ab-Arak River (28º35'N, 52º58'E); CMNFI 1979-0133, 50, 45.6-95.5 mm standard length, qanat stream near Qir (28º27'30"N, 53º03'E); CMNFI 1979-0135, 18, 21.8-49.2 mm standard length, Mand River tributary (28º08'N, 53º10'E); CMNFI 1979-0157, 4, 23.6-85.4 mm standard length, Fars, qanat stream at Hadiabad (28º52'N, 54º13'E); macrostomum? CMNFI 1979-0193, 1, 36.3 mm standard length, Fars, river 8 km from Darab (28º45'N, 54º27'30"E); macrostomum? CMNFI 1979-0195, 1, ? mm standard length, Fars, jube west of Darab (ca. 28º54'N, ca. 53º53'30"E); CMNFI 1979-0196, 1, 59.9 mm standard length, Fasrs, qanat and pool at Khanehnehrin (28º50'N, 53º31'30"E); not on data sheet check jar? CMNFI 1979-0197, 1, 51.3 mm standard length, Fars, spring nd stream 33 km from Fasa (28º45'N, 53º25'E); CMNFI 1979-0198, 23, 22.3-57.7 mm standard length, Fars, stream at Tadovan (28º47'N, 53º24'30"E); CMNFI 1979-0200, 8, 29.0-46.1 mm standard length, Fars, Mand River tributary (28º36'N, 53º36'30"E); CMNFI 1979-0202, 12, ? mm standard length, Fars, Mand River (29º01'N, 53º00'E); CMNFI 1979-0241, 18, 43.8-72.6 mm standard length, Fars, Shapur River at Shapur (29º47'N, 51º35'E); CMNFI 1979-0347, 2, 105.2-106.7 mm standard length, Fasr, Pol-e Berengie (29º27'30"N, 52º32'E); CMNFI 1979-0348, 4, 52.9-79.1 mm standard length, Fars, stream at Somduldul (ca. 29º28'N, ca. 52º32'E); CMNFI 1979-0404, 25, 20.2-127.9 mm standard length, Bushehr, stream 33 km south of Kaki (28º08'N, 51º47'E); CMNFI 1979-0405, 4, 33.5-36.7 mm standard length, Hormozgan, stream about 13 km north of Rostaq (28º29"N, 54º59'E); ID? CMNFI 1979-0497, 1, 85.6 mm standard length, Fars, Mand River at Band-e Bahman (29º11'N, 52º40'E); CMNFI 1979-0501, 17, 18.7-91.0 mm standard length, Fars, Mand River at Kavar (29º11'N, 52º41'E); CMNFI 1979-0504, 6, ?-93.0 mm standard length, Fars, stream at Pol-e Gaz in Lake Maharlu basin (no other locality data); CMNFI 1979-0789, 1, 164.6 mm standard length, Fars, Lake Parishan (29º45'N, 53º40'E); CMNFI 1993-0141, 1, 64.4 mm standard length, Bushehr, Dalaki River (29º28'N, 51º15'E); ID? CMNFI 2007-0061, 2, ? mm standard length, Fars, qanat pool at Ab-e Barik (ca. 27º52'N, ca. 54º09'E); CMNFI 2007-0063, 6, ? mm standard length, Fars, Mand River outside Jahrom (28º36'N, 53º37'E); USNM 205890, 2, 46.0-48.7 mm standard length, Fars, Lake Parishan (29º45'N, 53º40'E); ZSM 25705, 1, 107.0 mm standard length, Fars, Lake Parishan (29º45'N, 53º40'E).

Comparative material:- CMNFI 1980-0811, 2, 82.6-112.4 mm, Turkey, Akziyaret Deresi, Tigris River system (no other locality data);

BM(NH) 1931.12.21:1-2, 2, 69.5-78.5 mm standard length, Iraq, Mosul (36º20'N, 43º08'E); BM(NH) 1974.2.22:1184, 1, 130.2 mm standard length, Iraq, Sulaimaniyah (); BM(NH) 1974.2.22:1196, 1, 53.0 mm standard length, Hawiya Canal, Lesser Zab (); BM(NH) 1974.2.22:1214-1255 (in part), Khalis (33º49'N, 44º32'E).

Cyprinion milesi
(Day, 1880)

Common names

None.

[sabzug in Pakistan].

Systematics

Barbus milesi was described from "a spring at Tràl", Pakistan.

Berg (1949), Mirza (1969), Mirza et al. (1991) and Howes (1982) recognise this species as valid. If so, synonyms according to Berg (1949), would be Barbus bampurensis Nikol'skii, 1899 described from "Flum. Bampur", Scaphiodon daukesi Zugmayer, 1912 from "Irrigation channels and pools near Panjgur, Baluchistan, Pakistan", and Barbus baschakirdi Holly, 1929 from "Ein Bach bei Guadjik am Wege von Sarzeh in Biabun nach Darpahan in den Bergen von Baschakird, Südostpersien" (= a brook at Guadjik on the way from Sarzeh in Biabun to Darpahan in the Baschakird Mountains, southeast Persia).

Much of my material from southeastern Iran was assigned by me to C. watsoni. Specimens that resemble C. milesi (lacking a shallowly arched or sector mouth with a horny edge but having an oblique u-shaped mouth) are found at the same sample localities as typical C. watsoni. The mouth structure of the putative C. milesi resembles that of juvenile C. watsoni, possibly retained in the adult (paedomorphosis). A Principal Components Analysis does not separate these two forms when the mouth characters are not included in the analysis.

A specimen in the Naturhistorisches Museum Wien under NMW 52736, 34.4 mm standard length, is listed as a syntype under the name Cirrhina milesi but its locality is Gwadur, Hubb River and the type status may be an error.

Five syntypes of Barbus bampurensis, 32.0-64.8 mm standard length, are in the Zoological Institute, St. Petersburg (ZISP 11715) from "Flum. Bampur, 15-23.VII.1898, Zarudnyi". The jar label gives a date of 15-19.VII.1898.

The holotype of Barbus baschakirdi, 52.2 mm standard length, is in the Naturhistorisches Museum Wien under NMW 13798 and a cotype (syntype) of Scaphiodon daukesi, 102.8 mm standard length, is under NMW 19784.

Scaphiodon daukesi types in Munich were destroyed in World War II but one syntype is in the Naturhistorisches Museum Wien under NMW 19784, and two syntypes are in the Zoological Survey of India, Calcutta under ZSI F8028/1 and 8032/1 ((Menon and Yazdani, 1968; Eschmeyer et al., 1996; Neumann, 2006).

Key characters

The mouth is characteristically oblique, longer in lateral view than C. watsoni.

Morphology

The oblique mouth reaches back to the anterior eye margin in small fish and to the rear of the nostril in larger fish. Dorsal fin with 3 unbranched and 10-13 branched rays, anal fin with 2 unbranched and 7 branched rays, pectoral fin with 14-16 branched rays and pelvic fin with 7-8 branched rays. Total gill rakers 11-12. The following description is based mostly on Barbus bampurensis types. Dorsal fin spine strong and serrated, with large teeth in small fish. Lateral line scales 34-39. The scaleless groove before the dorsal fin is weakly expressed. Scales are present on the belly of large fish, almost absent on small fish. Upper flank scales may be regularly or irregularly arranged. Scales have few to no anterior radii, numerous posterior radii, numerous fine circuli, a subcentral anterior focus, and an anterior scale margin indented above and below the mid-line. A pelvic axillary scale is present. The head is more massive in relation to the body than for similar size C. watsoni/kirmanense specimens. The barbel is quite stubby at the base but tapers rapidly to the tip in larger fish. The type series of Barbus bampurensis (= C. milesi) has dorsal fin branched rays 10(4) or 11 (1), anal fin branched rays 7(5), pectoral fin branched rays 14(1) or 15(3) (one unclear), pelvic fin branched rays 7(1) or 8(4), lateral line scales 34(1), 36(1) and 37(3), and total gill rakers 11(3) or 12 (2). Two fish from Sib (see below) had dorsal fin branched rays 9(1) or 10(1), anal fin branched rays 7(2), pectoral fin branched rays 15(2), pelvic fin branched rays 6(1) or 7(1), lateral line scales 35(1) or 37(1), pharyngeal teeth 4,3,2 on the left side, total gill rakers 13(1) or 14 (1), and total vertebrae 38(1) or 39(1). Pharyngeal teeth have a slight hook on the anteriormost tooth with the rest in the main row with scooped-out crowns.

Sexual dimorphism

Tubercles line the anal fin rays and are apparent on the snout in males.

Colour

Copper-brown on the back and upper flank fading to a pinkish belly. Fins are pink and the lateral line has a bright orange streak along it. The preopercle also has orange-golden spots as does the base of the pectoral fins. There is a dark blotch at the base of the caudal fin. The caudal fin base bears a spot in small specimens and there are some much smaller, irregular spots on the caudal peduncle. Peritoneum brown to black.

Size

Attains about 19.0 cm.

Distribution

In Iran, it is recorded from the Sarbaz River of the Makran according to Saadati (1977), the Bampur River of the Hamun-e Jaz Murian basin according to Berg (1949) and the Dozdan River of the Hormuz basin (H. R. Esmaeili). Also in the Mashkid River basin in Pakistan and in rivers draining to the Indian Ocean.

Zoogeography

See under the genus.

Habitat

Unknown.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

None reported from Iran.

Economic importance

None.

Conservation

The distribution, abundance and biology of this species in Iran is poorly known and an assessment for conservation status cannot be given.

Further work

See above.

Sources

Type material: See above, Barbus bampurensis (ZISP 11715),Barbus baschakirdi (NMW 13798) and Scaphiodon daukesi (NMW 19784).

Iranian material: BM(NH)1883.8.2:2-3, 2, 72.2-130.9 mm standard length, Baluchestan, Sib near Dizak (27º15'N, 62º05'E). BWC97-4 no fish on cat sheet?

Comparative material: BM(NH) 1889.2.1:263-264, 2, 89.3-108.7 mm standard length, Afghanistan (no other locality data).

Cyprinion tenuiradius
Heckel, 1849

Common names

[Araxes kingfish (Fricke et al., 2007)]

Systematics

The type locality is the "Kara-Agatsch als aus dem Araxes" (= Qarah Aqaj River and the Kor River, Fars). Sometimes spelt tenuiradiatus (e.g. in Rainboth (1981) but this is incorrect). Syntypes of Cyprinion tenuiradius are in the Naturhistorisches Museum Wien according to Kähsbauer (1964) under NMW 52808 (1 specimen, 116.7 mm standard length), 52809 (2, 52.3-58.0 mm standard length), 52811 (4, 42.7-47.4 mm standard length), 52815 (1, 77.0 mm standard length) and 52816 (2, 75.5-80.8, although Kähsbauer lists only 1 while Banarescu and Herzig-Straschil (1995) list 2 as also found by me). Other material marked as syntypes from the "Kara-Agatsch. Th. Kotschy" includes NMW 52810 (2 , 103.7-110.0 mm standard length), NMW 52812 (2, 103.5-104.8 mm standard length), NMW 52813 (2, 97.7-103.1 mm standard length), NMW 52814 (1, 114.9 mm standard length), and 52817 (1, not examined). The catalogue in Vienna lists 8 specimens in one column and 26 in the adjacent column. Eschmeyer et al. (1996) add 2 fish from the Araxes River, formerly in NMW, now at the Rijksmuseum van Natuurlijke Historie, Leiden under RMNH 2486. The lectotype as selected by F. Krupp in 1984 is NMW 52814 and is published by Banarescu and Herzig-Straschil (1995) with NMW 52808, 52809, 52810, 52811, 52812, 52813, 52815 and 52816 as paralectotypes.

Karaman (1971) assigns this taxon as a subspecies of Cyprinion macrostomum and Bianco and Banarescu (1982) suggest it may be a subspecies in a polytypic species. Berg (1949) records it from the Tigris River where it may be sympatric with C. macrostomum. He considers it to be close to that species, perhaps its southeastern subspecies. Howes (1982) considers tenuiradius to be a variant of C. macrostomum.

Heckel (1846-1849b) distinguishes this species from C. macrostomum by a lower scale count (35-36 as opposed to 42; Berg (1949) gives 35-38 as opposed to 37-43); Krupp (1985c) gives 34-38 compared to 39-43 in macrostomum; Banarescu and Herzig-Straschil (1995) give 36-38, rarely 35 or 39 in C. tenuiradius compared to 41-44, rarely 40 or 45 in C. macrostomum), slenderer body, and a much thinner dorsal spine which is soft in its distal third. The mouth is arched and there is some lower lip development at the mouth corner as in C. kais (see illustrations in Krupp (1985c)). In addition, Berg (1949) gives a branched dorsal fin ray count of 12-13 in C. tenuiradius, 13-15 in C. macrostomum, although Banarescu and Herzig-Straschil (1995) give (12)13-15 for C. tenuiradius from the type locality of Kara-Agasch (sic). Krupp (1985c) states that tenuiradius has a smaller number of scale radii than macrostomum, radii are divergent and the posterior scale margin is curved. However, data for specimens examined by me show overlaps in meristic characters; although means differ, individual fish would be difficult to distinguish on counts alone.

The question then arises as to whether tenuiradius is distinct from macrostomum or merely a variant of a wide-ranging, variable species. The only absolute character is a weaker dorsal fin spine based on examination of type material; other, meristic characters overlap and minor variations in body form are difficult to quantify given a wide range of habitats (lowland rivers and marshes versus highland streams) which may affect shape. The species tenuiradius is retained here as distinct but would benefit from further analyses using new characters, if available, from molecular data.

Key characters

Distribution and a weak spine distinguish this taxon. The dorsal fin spine in macrostomum has teeth extending further along the spine, teeth are more well-developed even near the tip. Spine teeth in tenuiradius are more graded in size as they near the tip and are finer than in macrostomum.

Morphology

Dorsal fin with 4 unbranched and 11-15 branched rays (Berg (1949) has 12-13). The anal fin has 3 unbranched and 6-8 branched rays, usually 7. In 199 Iranian fish, 96.5% have 7 anal fin rays with the rest having 6 rays and 1, presumably anomalous fish, with 9 rays. Pectoral fin branched rays 13-18, pelvic fin branched rays 7-9. Lateral line scales 32-39. Gill rakers 10-21. Scales on the belly may be small and skin covered. There is a naked dorsal keel in front of the dorsal fin, although the area behind the occiput may be scaled and the groove begins nearer the dorsal fin. The mouth is transverse to more or less curved. The dorsal fin spine is weak and serrated only half way or two-thirds of its length. The chromosome number is 2n=50, comprising 13 metacentric, 5 submetacentric and 7 subtelocentric chromosomes pairs. Arm number is NF=86 (Esmaeili and Piravar, 2006).

Meristics for fish from Persian Gulf drainages of Fars, Bushehr and Hormozgan provinces including the Lake Maharlu endorheic basin:- dorsal fin branched rays 11(4), 12(51), 13(175), 14(74) or 15(9) (mean = 13.1, S.D. = 0.746); pectoral fin branched rays 13(3), 14(38), 15(117), 16(41), 17(2) or 18(1)(mean = 15.0, S.D. = 0.733); pelvic fin branched rays 7(23), 8(177) or 9(3)(mean = 7.9, S.D. = 0.345); total gill rakers 10(2), 11(16), 12(27), 13(24), 14(49), 15(35), 16(20), 17(14), 18(8), 19(3) or 21(1)(many counts are based on small specimens and may be low accordingly in comparison with Tigris River basin fishes; mean = 14.2, S.D. = 2.003); and lateral line scales 32(1), 33(15), 34(28), 35(41), 36(47), 37(56), 38(13) or 39(2)(mean = 35.7, S.D. = 1.431).

Sexual dimorphism

Unknown.

Colour

Overall colour is yellowish-white with a light grey back. Scale bases on the flank above the lateral line are brown. The pectoral and pelvic fins have an orange-yellow spot at their base.

Size

Reaches 16.3 cm (Berg, 1949).

Distribution

This species is found in the Gulf and Lake Maharlu basins in Iran (Bianco and Banarescu, 1982; M. Rabbaniha, pers. comm., 1995; Abdoli, 2000).

Heckel (1849) records this species as from the "Araxes", the modern Kor River in Fars. However, the catalogue sheets in Vienna for the types only list the "Kara Agatsch" (= Mand River) and no subsequent collections have been made of this species in the internal Kor River basin although Abdoli (2000) also maps it from the middle to lower Kor River, possibly based on Heckel's report. Berg (1949) records it from the Tigris River basin, perhaps in error, and Fricke et al. (2007) have it in Turkey from the Aras River system of eastern Turkey (presumably a confusion of the modern Aras or Araxes River with the classical Araxes or Kor River of Fars).

Zoogeography

See under the genus.

Habitat

Unknown in detail but found in springs, streams and rivers of varying descriptions.

Age and growth

Unknown. Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 40 fish measuring 5.04-13.49 cm fork length. The a-value was 0.0139 and the b-value 3.063 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases).

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

None reported from Iran.

Economic importance

None.

Conservation

The distribution, abundance and biology of this species in Iran is poorly known and an assessment for conservation status cannot be given. Endangered in Turkey (Fricke et al., 2007) but probably does not occur there.

Further work

See above.

Sources

Type material: See above, Cyprinion tenuiradius (NMW 52808, 52809, 52810, 52811, 52812, 52813, 52814, 52815, 52816).

Iranian material: ? see above and ID

Cyprinion watsoni
(Day, 1872)


ventral head

Common names

None.

[sehrgoar; sabzug = watsoni and microphthalmum - all in Pakistan].

Systematics

Scaphiodon irregularis Day, 1872 described from "rivers in the Sind hills", India, probably Scaphiodon microphthalmus Day, 1880 from "Quetta", Scaphiodon muscatensis Boulenger, 1887 from Muscat, Oman, Cirrhina afghana Günther, 1889 from "Nushki (N. Baluchistan)" and "small river at Kushk (N.W. Afghanistan), Badghis", Cyprinion kirmanense Nikol'skii, 1899 from "Schur-Ab in Kirmano orient.", Cirrhina afghana var. nikolskii Berg, 1905, Scaphiodon macmahoni Regan, 1906, Scaphiodon baluchiorum Jenkins, 1910 (see below for type locality), Scaphiodon watsoni var. belense Zugmayer, 1912 from the "Purali River, near Las Bela" (in Pakistani Baluchistan), Scaphiodon readingi Hora, 1923 from the "Salt Range, Punjab", India, and Cyprinion microphthalmum infraspecies nikolskii Berg, 1949 described originally in part as Cirrhina afghana var. nikolskii Berg, 1905, and Semiplotus dayi Fowler, 1958 are synonyms.

Semiplotus dayi was coined by Fowler to replace Scaphiodon aculeatus, a misidentification by Day (1880) for Chondrostoma aculeatum (= Capoeta aculeata). Fowler thought that Day's fish represented a new species which he named Semiplotus dayi. Howes (1982) considers Semiplotus dayi to be a synonym of Capoeta capoeta (since Karaman (1969a) synonymises Scaphiodon aculeata with C. capoeta. Day's Scaphiodon aculeatus is placed in the synonymy of Cyprinion microphthalmum infraspecies nikolskii by Berg (1949).

Syntypes (or at least specimens examined by Day) of Scaphiodon watsoni described from rivers on the Sind Hills and the Salt Range of the Punjab, India are in the Zoological Survey of India, Calcutta under ZSI 2596 (1), the Natural History Museum, London under BM(NH) 1889.2.1.370-9 (10, but 14 in jar September 2007, 35.6-93.4 mm standard length), the Australian Museum, Sydney under B.7751 (1), the Zoölogisch Museum, Universiteit van Amsterdam under ZMA 115.924 (2) and ZMA 119.255 (1), the Naturhistorisches Museum Wien under NMW 51671 (1), NMW 51672 (1) and NMW 51673 (1), the Museum für Naturkunde, Universität Humboldt, Berlin under ZMB 11042 (1)(132.6 mm standard length), the Rijksmuseum van Natuurlijke Historie, Leiden under RMNH 8704 (1) (or possibly 2552), the Zoological Institute, St. Petersburg under ZISP 8278 (4 but only 2 fish found by me, 63.6-79.6 mm standard length), and the Field Museum of Natural History, Chicago under FMNH 2303 ? 2302(4, 34.0-72.5 mm standard length as examined by me) (Whitehead and Talwar, 1976; Nijssen et al., 1993; Eschmeyer et al., 1996; Ferraris et al., 2000). The 3 fish in the Naturhistorisches Museum Wien measure 86.6, 80.8 and 93.3 mm standard length respectively and are listed there as syntypes.

ZISP 8279 comprising 3 fish, 51.5-52.1 mm standard length, has the same data as ZISP 8278 and may also be types. It is not clear if these are all types, those in ZISP not being marked as types and those in BM(NH) being marked as "possible types"; they may include material simply collected by Francis Day.

A cotype of Scaphiodon watsoni var. belense (NMW 19833) measures 136.9 mm standard length. Eschmeyer et al. (1996) report 2 fish under NMW 19833 although the Vienna card index in 1997 lists only one syntype under this number. In the Zoological Survey of India, Calcutta there are single syntypes under ZSI F827/1 (a misprint for 8027), ZSI F8029/1, ZSI F8030/1 and ZSI F8031/1 (see also Menon and Yazdani (1968)). The remainder of 42 syntypes were in the Munich Museum but were destroyed in World War II (Neumann, 2006).

Types of Scaphiodon microphthalmus are probably lost. The species was described from 2 specimens taken at Quetta in Pakistan. One specimen was sent to the Florence Museum but a recent search failed to locate it and the other specimen has not been located (Whitehead and Talwar, 1976; Banister and Clarke, 1977). A fish measuring 130.1 mm standard length in the Naturhistorisches Museum Wien is listed as a possible syntype (NMW 55897) and in the 1997 card index as "? holotype" (sic).

Note Howes (1982) and Mirza et al. (1991) consider Cyprinion microphthalmum to be a valid species with muscatensis, afghana, afghana var. nikolskii and baluchiorum as synonyms. Howes places macmahoni in watsoni rather than microphthalmum as Berg (1949) and Mirza (1969) do. Howes (1982) also includes irregularis, kirmanense, and readingi in watsoni.

A syntype of Scaphiodon irregularis is in the Australian Museum, Sydney under AMS B.7883 (Ferraris et al., 2000). Syntypes of Scaphiodon muscatensis are in the Natural History Museum, London under BM(NH) 1885.11.7:35-40 (6, 66.4-89.3 mm standard length) and BM(NH) 1887.11.11:289-291 (3, 72.1-79.3 mm standard length) (Eschmeyer et al., 1996; personal observations). Syntypes of Scaphiodon readingi are in the Zoological Survey of India, Calcutta under ZSI F10353/1 and ZSI 10354/1 (27) (sic, although the catalogue numbers seem to indicate only 2 fish) (Menon and Yazdani, 1968) and in the Zoological Museum of Moscow University (ZMMU) (P-1588) (Pavlinov and Borissenko, 2001).

Three syntypes of Scaphiodon baluchiorum (ZSI F9398 to F9400) and one syntype of Scaphiodon macmahoni (ZSI F1239/1) are in the Zoological Survey of India, Calcutta (Menon and Yazdani, 1968). A syntype of Scaphiodon macmahoni measuring 58.6 mm standard length from "Seistan" is in the Natural History Museum, London and was labelled as Cyprinion watsoni (BM(NH) 1905.11.29:27, 58.6 mm standard length). The type locality of Scaphiodon baluchiorum is "Gishtigan (Bampusht); Kalagan, 3,500 feet; Baluchistan". These localities are in Pakistani Baluchistan; Gishtigan being on the Kulushta River which drains into the Nihing River and then the Dashti River (Jenkins, 1910) (these are near the border of Iranian Baluchestan with the upper reaches of the Nihing being in Iran) and Kalagan possibly being the Kalugar River with headwaters in Iran and draining to the Hamun-i Mashkel in Pakistan. The type locality of Scaphiodon macmahoni is "affluents of the Helmand" (Regan, 1906), presumably an error for "effluents" or the delta of the Helmand.

The holotype of Cyprinion kirmanense, 61.6 mm standard length, is in the Zoological Institute, St. Petersburg under ZISP 11712 from "Schur-Ab in Kirmano orient. 27.VI." The 5 syntypes of Cirrhina afghana var. nikolskii are in the Zoological Institute, St. Petersburg (ZISP 11709) and are from the "Bampur River, 27 VII 1898, N. Zarudnyi" according to Berg (1949) but he mentions 2 additional fish with a somewhat deeper body, presumably also part of the type series. ZISP 11709 does have 7 specimens, 43.0-79.1 mm standard length, with a date 15-27.VII.1898. Four syntypes of Cirrhina afghana measuring 74.6-83.0 mm standard length from "Kushk" annotated Afghan. Boundary Comm. are in the Natural History Museum, London (BM(NH) 1886.9.21:150-154 - note that 150-154 indicates there should be 5 fish) with a further 6 syntypes measuring 44.9-99.5 mm standard length labelled "Nushki" and also annotated Afghan. Boundary Comm. (BM(NH) 1886.9.21:155-159 - note this indicates there should be 5 fish in this jar and probably one fish has been mixed up). Additional syntypes are in the Zoological Survey of India, Calcutta under ZSI 11474-11476 (3) and ZSI 11479-11485 (7) (Eschmeyer et al., 1996).

Berg (1949) places Cirrhina afghana var. nikolskii in his Cyprinion microphthalmum infraspecies nikolskii (see also Berg (1933a)). This infraspecies occurs together with Cyprinion microphthalmum but differs by a stronger osseous ray in the dorsal fin which is serrated almost to the summit (Berg (1949) states that transitions exist). The anterior belly region is scaleless also. ZISP 11709 fish mostly have their dorsal spines snapped off but one fish has osseous ray teeth between three-quarters and four-fifths along the spine and a second about three-quarters. ZISP 25406 from a qanat between Kerman and Bandar-e `Abbas comprises 12 fish, 31.0-53.6 mm standard length, belonging to infraspecies nikolskii according to Berg (1949). These fish, of all sizes, have the last quarter to a third of the osseous spine in the dorsal fin unserrated. The mouth form varies. One large fish has a terminal mouth, moderately oblique in lateral view, and no strong horny layer on the lower jaw. Others have a u-shaped or horny jaw positioned on the lower head surface so there is no real gape in lateral view. Some small fish are transitional between the two types. Fin serration, mouth form and development of scales on the anterior belly seem to be widely variable within samples of Cyprinion from a single locality and presumably a single species.

Berg (1949) recognises Cyprinion watsoni belense as a subspecies, rather than a variety as originally described, from Indian Ocean drainages of southeastern Iran and southwestern Pakistan (Baluchistan). It is distinguished by smaller scales (33-36) from the type form (31-34), hardly a sufficient criterion given the wide distribution range and individual variation shown by these fishes.

This species has not been adequately examined in southeastern Iran and most nominal species are referred to Cyprinion watsoni, the earliest available name for the taxon. C. watsoni is distinguished from other Iranian Cyprinion by having 9-11 dorsal fin branched rays (macrostomum and tenuiradius have 12-15; C. milesi also has a low dorsal ray count but has an oblique mouth, not transverse or arched (Berg, 1949)). Bianco and Banarescu (1982) consider that several subspecies may eventually be defined and that some of the names in synonymy here would then be used.

Berg (1949) also recognises C. irregulare as a distinct species with a low dorsal fin branched ray count as in C. watsoni but usually 37 or more scales in the lateral line, a scaleless groove on the back before the dorsal fin, and upper scale rows anteriorly arranged irregularly and not imbricate and C. microphthalmum with a low dorsal fin branched ray count as in C. watsoni but usually 37 or more scales in the lateral line, a scaleless groove on the back before the dorsal fin barely outlined, and upper scale rows anteriorly arranged regularly and imbricate. C. microphthalmum infraspecies nikolskii is described as having a strong dorsal fin spine with obvious teeth extending to the tip while typical C. microphthalmum has a weak ray with weak teeth only visible when the skin covering the fin is peeled away.

Berg (1949) later states that no great importance should be attached to the upper row scale arrangement and the groove development - if the groove is well-developed then the upper row scales are irregular and this phenomenon can be seen in some C. watsoni and C. microphthalmum specimens. Berg then suggests that irregulare could be regarded as an infraspecies of C. microphthalmum as this type of condition occurs in Capoeta fusca and in Garra rossica. Under the heading C. watsoni Berg also gives mouth shape, scale arrangement, dorsal fin spine serrations, and body form as characters which can vary greatly. These observations serve to confirm the great variability in characters for these fishes. Large series of adults and young would be needed to adequately define species and subspecies.

Mirza (1969) reports C. watsoni, C. microphthalmum and C. milesi from western Pakistan and Iran, the former in Makran drainages and the latter two in the Mashkel (= Mashkid) River basin. The characters used to separate these taxa are an oblique mouth and head length contained less than 4.5 times in total length (= C. milesi), an arched mouth, head length more than 4.5 times in total length, scaleless strip on back conspicuous, and 33-36 lateral line scales (= C. watsoni), and a transverse mouth, head length more than 4.5 times in total length, scaleless strip on back hardly visible, and 37-40 lateral line scales (= C. microphthalmum). Sample sizes in this study were small (22 fish) and these characters show considerable variation in larger samples and between fish of different sizes.

Key characters

The arched mouth and and 9-12, usually 10-11, branched dorsal fin rays serve to identify this species.

Morphology

Young fish have a more horseshoe-shaped mouth than larger and older fish where the mouth is a shallow arch, almost straight. The dorsal fin has 3-4 unbranched and 9-12 branched rays, the last unbranched ray of the dorsal fin being variably serrated and thickened. The extent of serrations appears to vary independently of size, from half to three-quarters or more of the spine length. The anal fin has 1-3, usually 3, unbranched and 6-8, usually 7 branched rays. In Iranian specimens, 89.7% of 419 fish had 7 branched anal fin rays, the remainder having 6 branched rays. Pectoral fin branched rays 11-18, usually 15-16, and pelvic fin branched rays 6-9, usually 8. Lateral line scales 31-43. Scales have well-developed anterior radii as well as posterior and some lateral radii. The scale focus is almost central on mid-flank scales. There is a naked median strip on the back in front of the dorsal fin, about one scale wide, in some fish. Some fish may show poor imbrication of scales on the belly and upper anterior flank. Gill rakers 8-18, reaching to or past the adjacent raker when appressed. Total vertebrae 38. Rarely with a tripartite swimbladder, usually bipartite (Mirza, 1971 - for his C. microphthalmum). Pharyngeal teeth 2,3,4-4,3,2 or 2,3,5-5,3,2, with spoon-shaped crowns. The back in front of the dorsal fin is naked in the mid-line. Mouth shape variable, from and arch to a transverse cutting edge. Scales on the belly can be embedded in the skin, obvious or even absent.

Meristics for Iranian specimens:-
 

Locality/Dorsal Fin Rays

9

10

11

12

x

S.D.

Hamun-e Mashkid

16

3

10.2

0.375

Hamun-e Jaz Murian

3

15

5

10.1

0.596

Dasht-e Lut

2

50

4

10.0

0.328

Makran

1

29

3

10.1

0.348

Hormuz

2

144

124

7

10.5

0.562

Sirjan

1

3

7

10.5

0.688

 

Locality/Pelvic Fin Rays

6

7

8

9

x

S.D.

Hamun-e Mashkid

6

13

7.7

0.478

Hamun-e Jaz Murian

1

21

1

8.0

0.302

Dasht-e Lut

1

54

8.0

0.135

Makran

33

8.0

0.000

Hormuz

2

31

237

8

7.9

0.400

Sirjan

2

9

7.8

0.405

 

Locality/Pectoral Fin Rays

11

13

14

15

16

17

18

x

S.D.

Hamun-e Mashkid

9

10

15.5

0.513

Hamun-e Jaz Murian

1

8

11

2

1

15.7

0.864

Dasht-e Lut

1

2

14

31

8

14.8

0.879

Makran

1

14

15

3

15.6

0.704

Hormuz

1

46

111

99

20

1

15.3

0.863

Sirjan

4

4

3

14.9

0.831

 

Locality/Total Gill Rakers

8

9

10

11

12

13

14

15

16

17

18

x

S.D.

Hamun-e Mashkid

1

4

3

4

3

4

13.8

1.718

Hamun-e Jaz Murian

3

4

10

4

1

1

12.0

1.492

Dasht-e Lut

1

7

8

16

12

10

2

12.2

1.427

Makran

1

6

9

4

8

3

1

1

13.0

1.794

Hormuz

3

7

33

45

64

60

41

10

2

1

12.2

1.569

Sirjan

2

2

1

5

1

12.1

1.375

 

 
Locality/Lateral Line Scales 33 34 35 36 37 38 39 40 41 42 43 x S.D.
Hamun-e Mashkid 1 3 9 1 1 1 2 1
Hamun-e Jaz Murian 1 9 11 1 1
Dasht-e Lut 5 12 26 10 2 1
Makran 2 1 17 8 4 1
Hormuz 1 2 17 57 99 72 20 7 3
Sirjan 3 4 3 1

? vertebrae

The holotype of Cyprinion kirmanense has 10 dorsal fin branched rays, 7 anal fin branched rays, 15 pectoral fin branched rays, 8 pelvic fin branched rays, 37 lateral line scales and 13 total gill rakers (not in above tables).

Sexual dimorphism

Males have snout tubercles and tubercles on the anal fin rays (Regan, 1906; Jenkins, 1910; Berg, 1949). Large tubercles are found on the snout in front of the nostrils, the top of the head, and in rows on the rays of the caudal and anal fin, following the ray branching, in a fish not yet fully mature (40.1 mm standard length). ZMB 11042 (132.6 mm SL) has tubercles thickly present on the snout extending back to the nostrils and then to the eyes, scattered all over the sides of the head, absent on top of the head (may be lost in this old specimen) and large tubercles on anal fin rays near the tip. There is a depression in front of the nostrils in adult males.

Colour

The back and upper flank are dark or copper brown, golden or dark olive, light green-brown or brown-grey, sometimes with bluish or orange tinges, fading to a light or yellowish pink on the lower flanks and belly. The flank scales are silvery and may be outlined in black. Black spots may be present along the flank as may be an orange stripe or a series of 7-9 orange spots above the lateral line anteriorly. The orange colour may be deep, almost red. Occasionally, there may be an orange spot below the lateral line. There may be a vertical orange line over the cleithrum or a spot at its postero-ventral corner. In some fish the whole cleithrum area is red-orange. The operculum, preoperculum and cheek can be iridescent blue. The bases of the pectoral and pelvic fins and part of the operculum may also be pink or orange-coloured. Young have a fine black streak above the lateral line. The dorsal and caudal fins are lead-coloured to black and other fins are pink to yellowish. All the fins except the paired fins may be hyaline. There can be a black caudal base spot, quite marked in some fish, particularly small ones. The peritoneum is black or dark brown.

Size

Attains 23 cm (Zugmayer, 1912).

Distribution

From southeastern Iran east to India. In Iran, it is recorded from the Dasht-e Lut, Hormuz, Hamun-e Jaz Murian, Sistan, Hamun-e Mashkid basins and in the Makran from the Jagin to the Bahu Kalat rivers (Berg, 1949; Spillman, 1972; Bianco and Banarescu, 1982; Abdoli, 2000; Esmaeili et al., 2011?).

Zoogeography

The occurrence of this species (as C. microphthalmum in Banister and Clarke (1977)) in Oman across the Straits of Hormuz is a result of the 120 m lowered sea level from 100,000 B.P. to 10,000 B.P. The Tigris-Euphrates then ran down the Gulf and presumably provided ready access across it. Banister and Clarke (1977) comment that it is surprising that only one species made the crossing but nothing is known of the climate during this 90,000 year period nor of the composition of the Iranian fish fauna. It may well have been quite impoverished. Hora (1956) describes fish paintings on pots from Nal in Pakistani Baluchistan dating from the third millenium B.C. One of the species not clearly represented is a Cyprinion, now found in that area (not extinct as Banister (1980) would have it). This potentially shows how the ichthyofauna in Southwest Asia can change over relatively short periods of time; the changes over 100,000 years must have been considerable and not readily traceable.

Dorsal fin branched rays have moderately strong modes of 10 for the Hamun-e Mashkid, Hamun-e Jaz Murian, Dasht-e Lut and Makran basins but modes of 10 and/or 11 for Hormuz and Sirjan, the westernmost basins. C. watsoni may be showing some introgression with Cyprinion species to the west which have higher counts. However, subsamples within the Hormuz basin do not show a clear pattern of higher modes or means towards the west.

Habitat

Kiabi and Abdoli (2000) found this species to be the commonest and to have the widest range in Hormozgan Province.

Age and growth

Unknown. Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 23 Iranian fish measuring 8.34-13.38 cm total length. The a-value was 0.0101 and the b-value 2.952 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases).

Food

Gut contents are primarily herbivorous items including filamentous algae such as Cladophora and Spirogyra, and a wide range of diatoms but some insect material is also found (Mirza, 1969; Farooq et al., 1996).

Reproduction

Spawning takes place in Pakistan at Islamabad from mid- to late March to mid-April (Shaikh and Jalali, 1989, 1991) and near Islamabad (33.3°N, 73.0°E) in April and May (Shaikh and Hafeez, 1993). Gonads begin to develop in December as photoperiod and temperature rise but a continuing warm temperature is the predominant factor for spawning to occur; a fall in temperature halts spawning. Eggs are dark yellow when mature, testes creamy when ripe. Spawning occurs once a year. Up to 150 eggs are recorded in fish from the Ab Garm-e Ganow with a diameter of 1.2 mm.

Parasites and predators

Males were reported as having snout tubercles and tubercles on the anal fin rays (Regan, 1906) but these were the encysted glochidia of a unionid mollusc (B. Prashad in Annandale and Hora (1920). Jalali et al. (1995) describe a new species of monogenean, Dactylogyrus pallicirrus, from fish taken in the Shur River, a Halil River tributary in the Hamun-e Jaz Murian basin. Jalali et al. (2005) summarise the occurrence of Gyrodactylus species in Iran and record Gyrodactylus sp. for fish from the Minab and Halil rivers.

Economic importance

This species is of no economic importance although Butt (1995) suggests that it could be a food source in Pakistan, occurring in shoals of considerable size in rivers that otherwise support little in the form of aquatic protein. It could be cultured as food and as a forage fish. This species has been used to study the effects of heavy metals in Pakistan (Shah, 2002). Higher concentrations of copper and zinc caused lethargy and loss of equilibrium.

Conservation

This species is widely distributed in various basins in southeastern Iran and neighbouring areas and does not appear to under any threat.

Further work

The biology of this species requires study and a thorough review of morphology in relation to named taxa should be carried out.

Sources

Type material: See above and note reservations on type status of some, Cirrhina afghana (BM(NH) 1886.9.21:150-154, BM(NH) 1886.9.21:155-159); Cirrhina afghana var. nikolskii (ZISP 11709); Cyprinion kirmanense (ZISP 11712); Scaphiodon macmahoni (BM(NH) 1905.11.29:27); Scaphiodon microphthalmus (NMW 55897); Scaphiodon muscatensis (BM(NH) 1885.11.7:35-40, BM(NH) 1887.11.11:289-291); Scaphiodon watsoni (FMNH 2303, NMW 51671, NMW 51672 and NMW 51673, ZMB 11042, ZISP 8278, ZISP 8279); Scaphiodon watsoni var. belense (NMW 19833).

Iranian material: CMNFI 1979-0138, 49, ?-66.8 mm standard length, Fars-Hormozgan border, stream in Rasul River drainage (ca. 27º32'N, ca. 54º58'30"E); CMNFI 1979-0143, 22, ? mm standard length, Hormozgan, marsh in Hasan Langi River drainage (27º21'N, 56º50'30"E); CMNFI 1979-0144, 76, ? mm standard length, Hormozgan, Minab River at Minab (27º09'30"N, 57º04'E); CMNFI 1979-0145, 139, ? mm standard length, Hormozgan, Geru River south of Minab (26º55'N, 57º01'30"E); CMNFI 1979-0149, 54, ?-93.8 mm standard length, Hormozgan, stream north of Bandar Abbas (27º36'N, 56º14'E); CMNFI 1979-0150, 34, 43.8-101.6 mm standard length, Hormozgan, stream at Gohreh (27º45'N, 56º05'E); CMNFI 1979-0152, 10, ? mm standard length, Homozgan, Shur River drainage (28º09'N, 55º43'E); CMNFI 1979-0153, 31 24.5-84.7 mm standard length, Fars, qanat stream and pool at Qaleh-ye Biabani (28º31'N, 54º53'E); CMNFI 1979-0154B, 49, ? mm standard length, Fars, stream channels at Koorsiah (28º45'30"N, 54º24'E); CMNFI 1979-0155, 26, ? mm standard length, Fars, springa t gavanoo (28º47'N, 54º22'E); CMNFI 1979-0156, 12, ? mm standard length, Fars, qanat at Rashidabad (28º47'N, 54º18'E); CMNFI 1979-0167, 25, ? mm standard length, Kerman, qanat at Bam (29º06'N, 58º20'E); CMNFI 1979-0168, 50, ?-93.8 mm standard length, Kerman, qanat at Shahabad (29º07'N, 58º16'E); CMNFI 1979-0173, 15, 26.5-84.0 mm standard length, Hormozgan, qanat at Hajjiabad (28º19'N, 55º54'E); CMNFI 1979-0176, 1, ? mm standard length, Hormozgan, Sarzeh River (27º30'30"N, 56º15'30"E); CMNFI 1979-0180, 5, ? mm standard length, Hormozgan, stream 3 km east of Essin (27º19'N, 56º17'30"E); CMNFI 1979-0181, 19, ? mm standard length, Hormozgan, Kul River (27º17'30"N, 56º03'30"E); CMNFI 1979-0183, 14, ? mm standard length, Hormozgan, stream in Rasul River drainage (27º11'30"N, 55º42'E); CMNFI 1979-0185, 4, ? mm standard length, Hormozgan, stream in Rasul River drainage (27º06'N, 55º45'E); CMNFI 1979-0186, 11, ? mm standard length, Hormozgan, strean and pools at Sar Khun oasis (ca. 27º24'30"N, ca. 56º25'E); CMNFI 1979-0187, 54, 18.9-73.6 mm standard length, Hormozgan, stream and pools at Sar Khun oasis (27º23'30"N, 56º26'E); CMNFI 1979-0188, 18, ? mm standard length, Hormozgan, jube at Gohreh (27º45'N, 56º05'E); CMNFI 1979-0189, 25, ? mm standard length, Hormozgan, jube and pool on road to Darab (27º08'30"N, 55º42'E); CMNFI 1979-0190, 44, ?-83.3 mm standard length, Fars-Hormozgan border, spring and pool at Galah Tuyeh (ca. 28º32'N, ca. 55º14'E); CMNFI 1979-0191, 35, 36.6-86.5 mm standard length, Fars, stream 10 km east of Furg (ca. 28º16'N, ca. 55º18'E); CMNFI 1979-0219, 19, 19.1-33.0 mm standard length, Kerman, jube 14 km west of Jiroft (28º37'N, 57º41'E); CMNFI 1979-0220, 4, 28.0-65.5 mm standard length, Kerman, jube 2 km south of Jiroft (28º39'N, 57º43'E); CMNFI 1979-0309, 2, 101.2-108.5 mm standard length, Kerman, Fahraj River at Azizabad (28º57'N, 58º42'E); CMNFI 1979-0310, 1, ? mm standard length, Baluchestan, qanat at Bazman (27º49'N, 60º12'E); CMNFI 1979-0311, 10, ? mm standard length, Baluchestan, Bampur River at Malakabad (27º11'N, 60º27'E); CMNFI 1979-0312, 39, ? mm standard length, Baluchestan, dam on Bampur River (27º11'N, 60º36'E); CMNFI 1979-0313, 68, ? mm standard length, Baluchestan, Bampur River at Bangharabad (27º20'N, 60º46'E); CMNFI 1979-0314, 10, 25.5-118.4 mm standard length, Baluchestan, qanat at Karavandar (27º50'N, 60º46'E); CMNFI 1979-0315, 71, ? mm standard length, Baluchestan, Bampur River 2 km north of Karavandar (27º51'N, 60º46'E); CMNFI 1979-0316, 22, 14.5-69.8 mm standard length, Baluchestan, stream 68 km south of Iranshahr (26º48'N, 61º02'E); CMNFI 1979-0317, 11, 16.5-118.6 mm standard length, Baluchestan, Sarbaz River at Bondan (26º35'N, 61º13'E); CMNFI 1979-0318, 11, ? mm standard length, Baluchestan, Sarbaz River at Huvar (26º09'N, 61º27'E); CMNFI 1979-0323, 6, ? mm standard length, Baluchestan, Sarbaz River (ca. 26º26'N, ca. 61º16'E); CMNFI 1984-0324, 4, ?39, ?51.9-117.8 mm standard length, Baluchestan, Bampur River at Sa'idabad (27º11'N, 60º22'E); CMNFI 1979-0325, 7, ? mm standard length, Baluchestan, qanat at Espakeh (26º51'N, 60º14'E); CMNFI 1979-0326, 10, ? mm standard length, Baluchestan, stream south of Pip (ca. 26º35'N, ca. 60º02'E); CMNFI 1979-0327, 10, 24.0-62.4 mm standard length, Baluchestan, stream in Geh River drainage (26º32'N, 59º57'E); CMNFI 1979-0329, 82, ? mm standard length, Baluchestan, stream at Zaminbandan (27º02'N, 61º20'E); CMNFI 1979-0331, 25, 13.1-50.3 mm standard length, Baluchestan, qanat in Saravan (27º22'N, 62º20'E); CMNFI 1979-0332, 9, 20.8-33.3 mm standard length, Baluchestan, qanat at Kalapurkan (27º14'N, 62º33'E); CMNFI 1979-0334, 4, 26.3-38.4 mm standard length, Baluchestan, Mashkid River (27º04'N, 62º54'E); CMNFI 1979-0335, 2, 66.8-72.2 mm standard length, Baluchestan, qanat at Esfandak (27º07'N, 62º50'E); CMNFI 1979-0338, 17, ? mm standard length, Baluchestan, Tahlab River drainage 8 km from Mirjaveh (28º58'N, 61º24'E); CMNFI 1979-0339, 24, 24.5-76.9 mm standard length, Baluchestan, Tahlab River drainage 16 km from Mirjaveh (28º56'30"N, 61º21'E); CMNFI 1979-0411, 8, ? mm standard length, Hormozgan, Minab River (27º24'N, 57º12'E); CMNFI 1979-0412, 22, 22.0-122.2 mm standard length, Hormozzgan, spring at Saras (27º30'N, 57º34'E); CMNFI 1979-0415, 5, ? mm standard length, Hormozgan, stream south of Ab Garm-e Ganow (27º17'30"N, 56º20'E); CMNFI 1979-0416, 2, 40.1-55.9 mm standard length, Hormozgan, Ab Garm-e Ganow (ca. 27º26'N, ca. 56º20'E); CMNFI 1979-0418, 5, 58.2-111.2 mm standard length, Hormozgan, river near Kahkom (28º09'N, 55º43'E); CMNFI 1991-0141, 3, ?, mm standard length, ; see original sheet? CMNFI 2007-0031, 12, ? mm standard length, Baluchestan, headwater of Bampur River (27º51'N, 60º46'E); CMNFI 2007-0033, 15, ? mm standard length, Baluchestan, Rusgay qanat in Iranshahr (27º13'N, 60º41'E); CMNFI 2007-0034, 3, ? mm standard length, Baluchestan, headwater stream on road to Zaboli (ca. 26º58'N, ca. 61º27'E); CMNFI 2007-0036, 8, ? mm standard length, Baluchestan, qanat at Bazman (27º49'N, 60º12'E); CMNFI 2007-0037, 7, 62.4-166.3 mm standard length, Kerman, Hosseinabad and Gamatabad qanats at Bam (29º06'N, 58º21'E); CMNFI 2007-0038, 9, 62.8-101.2 mm standard length, Kerman, Mehtiabad qanat at Bam (29º06'N, 58º21'E); CMNFI ?, 11, 46.5-69.1 mm standard length, ; 44 ? check, Sirjan on data sheet but no fish at this number in catalogue, check shelf CMNFI 2007-0049, 11, ? mm standard length, Hormozgan, ditches in upper Kol River basin at Hajjiabad (ca, 28º19'N, ca. 55º55'E); 45 CMNFI 2007-0050, 4, 61.2-92.4 mm standard length, Hormozgan, ditches in upper Kol River basin at Hajjiabad (ca. 28º19'N, ca. 55º55'E); 46 CMNFI 2007-0051, 7, 54.6-84.3 mm standard length, Hormozgan, upper kol River basin at Hajjiabad (28º19'N, 55º55'E); 47 CMNFI 2007-0052, 2, 70.7-92.3 mm standard length, Hormozgan, ditch at Qotbabad (27º46'N, 56º06'E); 48 CMNFI 2007-0055, 15, 24.5-75.3 mm standard length, Hormozgan, headwtaer stream in Minab River basin (ca. 27º47'N, ca. 57º12'E); 51 CMNFI 2007-0056, 14, 30.2-70.4 mm standard length, Kerman, qanat at Kahnuj (27º58'N, 57º45'E); 52 CMNFI 2007-0059, 9, ? mm standard length, Fars, Chashmeh Barashk (ca. 27º24'N, ca. 54º06'E); CMNFI 2007-0060, 3, 56.2-93.7 mm standard length, Fars, Chashmeh Ab-e Shirin near Lar (ca. 27º41'N, ca. 54º17'E); 57ID? BM(NH) 1883.8.2:4-9, 5, 42.4-114.9 mm standard length, Baluchestan, Jalq (27º36'N, 62º42'E); BM(NH) 1883.8.2:20-25, 6, 23.1-85.2 mm standard length, Baluchestan, Sib near Dizak (27º15'N, 62º05'E); NMW uncatalogued, 19, 18.1-52.9 mm standard length, Hormozgan, Ab Garm-e Ganow (ca. 27º26'N, ca. 56º20'E). BWC97-4, ?, ? mm standard length, Kerman, Halil River (no other locality data); no fish on cat sheet? BWC97-5, 1, ? mm standard length, Hormozgan, Hasan Langi River (no other locality data);

Comparative material: CAS 28722, 1, 117.4 mm standard length, India, Punjab, Salt Range, Katas Nallah (no other locality data);

Genus Cyprinus
Linnaeus, 1758

The carp genus is found in Europe and Asia and comprises several species of which one has been widely introduced as a food fish.

This genus is characterised by a compressed but heavy body, large size, rounded snout, 2 pairs of barbels, large molar pharyngeal teeth in 3 rows, a very long dorsal fin with the last unbranched ray spine-like and serrated, the anal fin short but with the last unbranched ray spine-like and serrated, the gut is moderately long, and the dorsal and lateral skull bones are sculptured.

Cyprinus carpio
Linnaeus, 1758

Common names

كپور (= kopur, kapur or kapoor (in Gilaki)), kapur-e ainehi (= mirror carp), كپور معمولي (kapur-e ma'mouli or mar'mulleh or ma'muli or maamoli (= common carp)), rashti or كپور رشتي (kapur-e Rashti) in Khuzestan (because their origin was Rasht), mahi-ye gul (= flower fish, meaning in this sense a good fish).

[car or carp shaeeh in Arabic; caki in Azerbaijan; geitan-tsatsan or dliter, both in Armenia; sazan for wild and karp for cultured carp in Russian; common carp, European carp, German carp, wild carp; mirror carp, leather carp, line carp, naked carp (last four referring to scalation), koi (aquarium variety)].

Systematics

Cyprinus Carpio was originally described from Europe. Cyprinus carpio var. caspicus Walbaum, 1792 is described from the mouth of the Volga and Don rivers but is infrasubspecific and the name has not been used in Iran nor has Cyprinus carpio fluviatilis Pravdin, 1945 described from floodplain lakes of the Volga River near Saratov. Mousavi Gel Sefid et al. (2007) give some morphometric and meristic charcters of fish from the Anzali Lagoon.

Key characters

This species is easily identified by the long dorsal fin, the spine in both the dorsal and anal fins, and the two pairs of barbels.

Morphology

Dorsal fin with 2-5 unbranched rays followed by 14-23, usually 18-20, branched rays, anal fin with 2-4, usually 3, unbranched rays followed by 3-7, usually 5, branched rays, pectoral fin branched rays 13-19, and pelvic fin branched rays 5-9, usually 8. The dorsal fin has the last unbranched ray developed as a toothed spine and the anal fin has a similar spine. Lateral line scales 26-41, mostly 36-39. Scales may be absent (leather carp), restricted to a few, enlarged scales (mirror carp), or only a mid-lateral row of scales (line carp), in cultivated varieties. Wild carp are fully scaled. Individual scales have a central focus, wavy anterior margin, few radii on the anterior and posterior fields in young fish, and medium numbers of radii on fish 12-14 cm standard length. There are numerous fine circuli and the posterior scale field breaks up into bubble-like structures. Gill rakers 17-29 (some literature counts may be lower arm of arch only and there may be size-related variation too) and vertebrae 32-39 (lower counts may not include Weberian vertebrae). Rakers touch the second raker below when appressed and have a row of knobs on their medial surface. Pharyngeal teeth 1,1,3-3,1,1 with variants 1,2,3-3,2,1, 1,2,3-3,1,1, 1,1,1,3-3,1,1, 1,1,3-2,1,1 and 1,3-3,1,1. Posterior major row teeth are large with flattened crowns bearing wavy ridges while more anterior teeth are a rounded knob, or even concave on top of the knob. The gut is elongate with several coils. This species is a tetraploid (2n=98-104)(Al-Sabti, 1986; Klinkhardt et al., 1995).

Meristic values for Iranian specimens are:- dorsal fin branched rays 16(1), 17(5), 18(18), 19(16), 20(10); anal fin branched rays 4(1) or 5(49); pectoral fin branched rays 14(1), 15(7), 16(25), 17(16) or 18(1); pelvic fin branched rays 6(1), 7(6) or 8(43); lateral line scales 34(2), 35(6), 36(10), 37(26) or 38(6); total gill rakers 18(1), 20(2), 21(11), 22(11), 23(15), 24(4), 25(4) or 26(2); total vertebrae 36(1), 37(19), 38(26) and 39(6); and pharyngeal teeth ?. Note that these samples may include individuals which are not native but introduced for fish farming from stocks outside Iran.

There are morphometric and meristic differences between carp from the southwestern and southeastern Caspian Sea but these are ecological not taxonomic. Also, carp from the Anzali Mordab differ from those in the sea by having a longer head, snout, eye and postorbital region (although of course some of these are redundant), greater interorbital width, shorter predorsal and preanal distances, shorter dorsal, anal, pectoral and caudal fins, and a lower anal fin (A. M. Shukolyukov in Berg (1948-1949)). Yousefian (2004) found carp from the Caspian Sea in Iran had a dominant genotype different from those in a fish farm. Meristics (scales and fin rays) and morphometrics (head length and body width) also differed.

Sexual dimorphism

Females are deeper bodied than males because of their eggs and the distance between the pectoral and pelvic fins and the pelvic and anal fins is more. Dorsal and anal fins in males are higher, the anal fin is longer at the base, the pectoral fin is longer and the lobes of the caudal fin are longer. This is accounted for by the greater swimming activity of males during spawning (Kuliyev and Agayarova, 1984). Breeding males have fine tubercles on the head, particularly on the anterior operculum and preoperculum and under the eye, above the lateral line and more frequently below it, and on the fin rays.

Colour

In semi-diadromous carp from the Kura region of the southwest Caspian Sea, the overall body colour is dark yellow, the flanks being golden-yellow with dark shading, and the back is black. The belly and fins are light yellow and the caudal fin is reddish. Lake populations are darker. Young fish from Iran are silvery on the flanks (but not as bright as Carassius auratus), greyish on the back, silver-pearl on the belly, the iris is silvery with grey above and below, the dorsal fin and upper caudal lobe are pale grey, the lower caudal lobe and anal fin are orange, the pelvic fin is pale orange, and the pectoral fin has only traces of orange. The caudal fin may be yellow-orange with lobe margins red. The freshwater resident form in the Anzali Mordab is yellowish, the semi-diadromous form dark. The peritoneum is grey to silvery and may be speckled.

Size

Carp resident in fresh waters are smaller than semi-diadromous carp. In the 1950s in Iran, carp catches were 20-41 cm long (Farid-Pak, no date). Fish up to 1.0 m long are caught in the Caspian basin (A. Abdoli, pers. comm., 1995). Maximum size exceeds 1.2 m and 44.9 kg, possibly to 1.5 m and 69.6 kg.

Distribution

Widely introduced in the Middle East for aquaculture. Found naturally in Iran in the whole Caspian Sea drainage, it is also widely stocked in the provinces of Gilan and Mazandaran (Petr, 1987; Abbasi et al., 1999). It is reported from the Atrak, Gorgan, Gharasu, Tajan, Babol, Aras, and Safid rivers, the Anzali Mordab and the Siah-Keshim Protected Region, Gorgan Bay, the southeast Caspian Sea, southwest Caspian Sea and south-central Caspian Sea as an introduced species, and in all these plus the Haraz and Pol-e Rud rivers as a native species (Riazi, 1996; Kiabi et al., 1999).

It is probably native to the Tedzhen River of Turkmenistan (the Hari River or Harirud in Iran) (Aliev et al., 1988). This species is also recorded from the Karakum Canal and Kopetdag Reservoir in Turkmenistan (Shakirova and Sukhanova, 1994; Sal'nikov, 1995) and may eventually reach Iranian waters in the Tedzhen (= Hari) River basin. These latter stocks may be introduced.

It has been introduced to the Sistan basin to a canal flowing into the Chahnimeh, the Sistan Dam, Hamun Lake and the Hamun Kushk (Ahmadi and Wossughi, 1988; J. Holčík, in litt., 1996; Shamsi et al., 2009)), to the Karaj River in the Namak Lake basin, to the Mahabad Dam (Abdi, 1999; www.mondialvet99.com, downloaded 31 May 2000), to the Haft Barm lakes near Shiraz in 1984 (Petr, 1987), to the Manjil Reservoir on the Safid Rud (Nümann, 1966), to Mehr Dasht Lake, Markazi, to the Zayandeh River Dam (Y. Keivany, in litt., 1992; Ghorbani Chafi, 2000), to Lake Zaribar, Kordestan (Abzeeyan, 5(5):III, 1994), to Lake Famur from nearby fish farms (Maafi, 1996b), to the Kor River basin in Fars (A. Alamdari, in litt., 1997; M. Hafezieh, in litt., 1997), throughout Khuzestan Province (not caught in the 1970s), to the Kol River, Hormozgan (Bagheri et al., 2010), and to numerous other water bodies and ponds throughout Iran where it reproduces in the wild. Mirror carp found in the Shadegan marshes of Khuzestan are escapees from fish farms. Carp in the Bahu Kalat and Sarbaz rivers of Baluchestan are also escapees from fish farms (A. Mobaraki, pers. comm., June 1999).

Abdoli (2000) records this species as present generally in the Dasht-e Lut, Sistan, Dasht-e Kavir and Kerman-Na'in basins; in lower reaches of rivers from the Gorgan to the Astara and along the Caspian coast, the middle Aras River; the Kashaf River in Khorasan; the middle and lower Talkheh River and Tatavi River in the Orumiyeh basin; the Qareh Chai and Qom rivers in the Namak Lake basin; the middle to lower Zayandeh River in the Esfahan basin; the Aravand, lower Karun, Jarrahi, lower Karkheh rivers in the Tigris River basin; the middle Halil River in the Hamun-e Jaz Murian basin; and the middle and upper Kul River including the Shur River tributary in the Hormuz basin. Jalali et al. (2005) records this species from the Dez, Karun and Zayandeh rivers, the Vahdat reservoir, and the Zarivar and Kaftar lakes. Abdoli and Naderi (2009) record it from the Atrak, Gorgan, Ghararasu, Tajan, Babol, Aras and Safid rivers, the Anzali Talab, Gorgan Bay, and the southwest, southeast and south-central Caspian Sea.

Zoogeography

The natural distribution of the carp is supposed to be Asia Minor and the Caspian Sea basin where its origins lie in the late Pliocene. From this area, modern wild carp spread east and west, perhaps as late as the last postglacial thermal optimum, and latterly aided by man (Balon, 1974; 1995; Van Damme et al., 2007). Iranian carp may be a mixture of native and introduced stocks (or species - see Kottelat (1997) - but this remains unresolved). It is probably not now possible to distinguish the native stocks morphologically because of admixtures of farmed stocks. All domestic forms probably originated from native Danube River stocks (Bănărescu, Barus and Peňáz in Bănărescu and Paepke, 2002). Khalili and Amirkolaie (2010) compared fish electrophoretically, meristically and morphologically from the Anzali lagoon, Qareh Su and Bandar Gaz. Morphometric differences between east and west were attributed to a relatively smaller body size in the east and the influence of domestic stocks. Electrophoretic differences, particularly obvious at Anzali, were attributed to the larger number of fish farms in this latter area.

Habitat

This species favours an abundance of soft vegetation in shallow water, necessary for successful reproduction. Still waters are preferred but they are found in the lower courses of lowland rivers with moderately flowing water, and occasionally in water exceeding 2 m/sec. They can often be seen basking at the surface or feeding on algae and their dorsal fins break the water surface. Large fish often move into shallows in the afternoon and evening. Carp also leap from the water but the reason is unknown. They rarely descend below 30 m in lakes and avoid fast water in streams. Carp overwinter in depressions in the lower reaches of rivers in the Caspian basin. There are both freshwater resident populations in the Caspian basin and diadromous (or semi-diadromous) ones, the latter living in brackish water near river mouths and only entering fresh water to spawn. The migration up the Ural River may reach 60 km from the river mouth. Some fish apparently spawn in the brackish waters of shallow coastal areas in the Caspian. Knipovich (1921) reports fish in the Iranian Caspian Sea down to 11.9-12.3 m. Riazi (1996) and Karimpour (1998) report that this species is both native (resident) to, and migrates to, the Siah-Keshim Protected Region of the Anzali Mordab.

Carp have a salinity tolerance under experimental conditions of up to 8‰, and for short periods 18.6‰ with acclimation, and this has significance for survival of carp in the Caspian Sea and in waters of southern Iran and Iraq where this species is farmed. Hafezamini and Oryan (2002) and Hafezamini et al. (2003) however found that, under experimental conditions, all fish in their study died at 18‰ in less than 12 hours. Carp eggs hatch in water up to 10‰, with the favourable level being up to 6.6‰ (Al-Hamed, 1971).

Low dissolved oxygen concentrations of 3 mg/l are tolerated and levels as low as 0.5 mg/l can be withstood for 2-3 hours. Normal growth has occurred in fish kept at 35°C. The immune response of carp following exposure to the organophosphate malathion has been studied by Soltani et al. (2003).

This species is considered to be a dominant species in the Karun River along with Barbus (= Tor) grypus (Iranian Fisheries Research and Training Organization Newsletter, 17:1, 1997).

Introduced to Iraqi waters in 1960 as juveniles, this species rapidly became established (Ahmed and Taher, 1988). Cyprinus carpio was caught in large numbers in the Shatt al Arab of Iraq down to the estuary after an increase in the discharge of the Tigris River reduced salinity (N. A. Hussain, in litt., 1994).

Age and growth

Fatemi et al. (2009) examined fish from beach seines along the Caspian shore of Iran for 2006-2007. Ten age groups were recorded and the catch was dominated by fish aged 4-5 years old. Growth parameters based on scale reading were FL = 71.52 cm and K = 0.16 per year for the total population, FL = 70.54 cm and K = 0.15 per year for males, and FL = 72.00 cm and K = 0.16 for females. Growth parameters based on length-frequency analysis gave values of FL = 72.0, 69.3 and 73.0 cm and K = 0.18, 0.15 and 0.18 per year respectively. The total (Z), natural (M) and fishing (F) mortalities were 0.71, 0.29 and 0.42 per year respectively for sexes combined. Exploitation (E) was 0.59 for sexes combined and no further fishing pressure was recommended.

Females are larger and mature a year later than males. Sexual maturity is attained in the second year of life, and in a few individuals even by the end of the first year, in the southeastern Caspian Sea; but in the southwestern Caspian Sea this occurs in the third and fourth years (Kuliyev and Agayarova, 1984). Resident carp in Dagestan mature in their third year at about 30 cm and have an average life span of 6 years whereas the semi-anadromous or semi-diadromous form matures in its fourth year at 35-36 cm and has an average life span of 8 years (Shikhshabekov, 1969). Growth is faster in the Kura River of Azerbaijan than in other populations in the Caspian but maturity is later at age 4 or more usually at 5 years. Freshwater residents in the Anzali Mordab are slow-growing compared to the semi-diadromous form and are less common. Fish taken in the commercial operations in Iran were 3-7 years old, 31.0-63.0 cm long and weighed 539-3375 g (Razivi et al., 1972). Mousavi Gel Sefid et al. (2007) give some growth equations for fish from the Anzali Lagoon.

This species was stocked in the Dukan and Derbendikhan dams of Iraq in the 1960s where fish up to 3 years of age were reported by Ciepielewski et al. (2001). Decreasing growth rates indicate conditions are not too favourable although growth in the first two years is comparable with that in lakes of central Iraq. The condition coefficient (K) was higher among smaller fish, e.g. fish from Dukan at 230 g had a K of 2.32, at 1 kg K was 1.75. Ahmed and Taher (1989) examined the growth of 0+ carp in Hawr al Hammar, Iraq found the length-weight relationship to be W = 0.00004627 L2.8022 and derived the growth equation Lt = 189.87 (1-e-0.0158 (t + 25.662)) with a calculated length at the end of the first year of life of 189.87 mm. This is relatively larger than for other parts of the world, indicating a successful introduction of this exotic. Al-Nasiri and Dawood (1991) found the smallest mature male was 182 mm and the smallest female was 184 mm in Hawr al Hammar, with sexual maturity achieved in the first year of life. Maximum life span was 8+ years. Epler et al. (2001) found the oldest age groups in Iraqi lakes to be 5+ in Lake Habbaniyah and 3+ in Lake Razzazah. The mean condition factor was 1.47 for Lake Habbaniyah and 1.50 for Lake Razzazah.

In Sariyar Dam Lake near Ankara in central Anatolia, ages range from 0 to 18 years (Ekmekci, 1996b). In their first year, fish have an average fork length of 103 mm and weigh 24 g, in 5 years they average 357 mm and 822 g, and in 10 years 580 mm and 3365 g. In Gölhisar Lake by contrast, a small water body in western Turkey, age composition was from 1 to 6 years and fish attained a maximum of 494 mm and 1922 g (Alp and Balik, 2000).

Maximum life span for this species is reported as 47 years for domestic fish.

Food

Food is derived from browsing on the substrate at all hours, if the temperature is favourable. Browsing muddies the water and can inhibit other species and uproot plants. Mouthfuls of bottom ooze are taken up, spat out and the food items selected. These include aquatic insects, crustaceans, worms and molluscs, and more rarely, fish. Plant material is ground up by the molar pharyngeal teeth and includes algae, seeds, wild rice, leaves and various aquatic plants. Organic sewage is also eaten. Some surface feeding on algal mats or insects will also occur. Feeding almost completely stops in winter and the fish go into a form of hibernation.

Feeding in the Hawr al Hammar, Iraq is related to temperature, the peak intensities being July and the minimum in January with peak activity in September and minimum in January. Feeding does occur year round and smaller fish (<200 mm) have highest feeding activity in spring while adults have this in summer (Hussein et al., 2000a). In a study of the recovering Hawr al Hammar, diet was 25.45% algae, 18.18% snails, 12.73% diatoms and 12.73% copepods, 10.91% insects with plants, cladocerans and shrimps at less than 10% each, in the Hawr al Hawizah diet was 27.3% snails, 18.2% insects and 12.1% for algae, plants and cladocerans with fish, diatoms and copepods at less than 10% each, and in the Al Kaba'ish (= Chabaish) Marsh 33.3% algae, 20.4% insects, 11.1% snails and diatoms and plants at 10.2, with various crustaceans at less than 10% each (Hussain et al., 2006). Hussein et al. (1991) examined diet in the Garma Marshes, Iraq and found crustaceans, molluscs, aquatic plants and seeds, aquatic insects, oligochaetes and fish to be dietary items, selection and numbers varying with carp size and season. Some fish were found to have fed exclusively on only a single, different mollusc species, presumably as opportunity presented. Al-Shamma'a et al. (1996) examined the food of this species in Al Qadisiyah Reservoir, Iraq and found plants, their seeds, molluscs and aquatic insects, all bottom foods. Mangalo and Akbar (1988a, 1988b) studied the food of carp in a farm pond at Al-Latifiyah, Baghdad where zooplankton was the principal diet. The gill rakers show an efficient structure for filtration, indicative of a phytoplanktivorous and omnivorous feeding (Salman et al., 1993). Salman et al. (1994) report a mixture of animal and plant foods, with zooplankton a dominant component of all length groups. The gut is coiled and 3.42 times standard length, indicating omnivory with plant food being important. Epler et al. (2001) gave the diet in Lake Habbaniyah, Iraq as 51.7% plants, 15.7% oligochaetes, 15.2% tendipedids, 7.2% molluscs, 5.2% detritus and 4.1% cladocerans. They also found that where there is significant competition between autochthonous species, as here, carp become another strong competitor for food. Dietary coincidence between carp and shabbout, gattan and himri was 58.5, 68.5 and 54.2 % respectively. Hussein et al. (2000b; 2000c) examined dietary overlap between this species and three native carps in the Hawr al Hammar. Overlap with Barbus (= Mesopotamichthys) sharpeyi was the weakest as this species is a herbivore but small B. luteus (<200 mm) and B. xanthopterus showed strong overlaps. This overlap may explain the decline in some native carps. Hussain and Ali (2006) also examined feeding relationships among fishes in the Hawr al Hammar and found this species to be a carnivore, 26.4% of the diet being crustaceans, 12.7% insects and 30.5% molluscs. Dietary overlap of 84% was found between this species and Barbus (= Luciobarbus) xanthopterus but the availability of food resources offset possible competition, contrasting with the conclusions above. Ciepielewski et al. (2001) found the diet of this species in the Dukan and Derbendikhan reservoirs in Iraq to be mainly algae, copepods and chironomids.

Reproduction

Under natural conditions, males spend more time on the spawning grounds than females and spawn several times. More than 7 million eggs up to 1.71 mm in diameter may be present in a female but only about 500 are laid at a time. Wild carp in the Atrek River had a fecundity range of 16,000 to 543,000 eggs (Bănărescu, Barus and Peňáz in Bănărescu and Paepke, 2002). The resident form is less fecund by about half than the semi-anadromous form (Shikhshabekov, 1969). In the Anzali Mordab a mass spawning run takes place in April with spawning in April-May. The first migratory fish are seen as early as January. Shallow weedy areas or the mouths of rivers are used as spawning sites and adhesive eggs are laid on plants.

Resident populations in Dagestan spawn earlier, by about a month, than the semi-anadromous population which spawn in early May (Shikhshabekov, 1969). Spawning time variations are governed by temperature and the most favourable temperature is 18-20°C.

Carp ascend the Kura River of Azerbaijan in spring and autumn. The spring run begins in mid-March and peaks in April while the weak autumn run lasts from August to mid-October.

Carp have feeding grounds in the coastal waters of the southeastern Caspian Sea and enter the Atrak River in winter to spawn between February and April. Young migrate downstream, this movement ending in July when the river flow is minimal or ceases. When there is no flood, spawning does not occur (Petr, 1987).

Fish in Iraqi ponds grew 25-30 cm in the first year of life and matured in 1-2 years. At 16-26ºC they spawned from late February to late April and again in the autumn (Al-Hamed, 1960). Palm tree fibres were used for egg deposition and eggs hatched in 4-8 days. Al-Nasiri and Dawood (1991) found a fecundity range of 14,150-1,492,500 eggs in Hawr al Hammar with a mean relative fecundity of 182 eggs/g of body weight, and a egg diameters of 0.90-1.02 mm. The gonadosomatic index indicated spawning in March and possibly October-November. Epler et al. (2001) studied reproduction in lakes Tharthar and Habbaniyah and found both sexes to achieve maturity in the first year of life at 13.5 cm for males and 12.6 cm for females. Spawning occurred in May and fecundity was 186-531 thousand eggs/kg body mass.

The spawning behaviour involves stimulation of a female while moving over vegetation and being accompanied by 2-3 males, active movement and spawning being induced by blows from the male(s). The eggs adhere to the vegetation or are lost. Most eggs are shed at night or in the early morning.

Parasites and predators

Eslami and Anwar (1971) record the cestode Caryophyllaeus fimbriceps from this species on the Caspian coast of Iran. Mokhayer (1976b) records the cestodes Bothriocephalus gowkongensis, the nematode larva Anisakis and the acanthocephalan Pomphorhynchus perforator. Mokhayer (1989) reports metacercariae of the eye fluke, Diplostomum spathaceum from this species in Iran, which can cause complete blindness and death in commercially important species. Jalali and Molnár (1990b) variously record the monogeneans Dactylogyrus anchoratus, D. extensus, D. sahuensis and D. vastator from carp on fish farms throughout Iran and Naem (2002) records D. anchoratus from fish in the Safid River.

Moghainemi and Abbasi (1992) record a wide range of parasites from this species in the Hawr al-Azim in Khuzestan. Mortazaei et al. (2000) record an infection rate of 66% (2 of 3 fish) with the worm Bothriocephalus opsariichthydis in Khuzestan marshes. Sattari and Faramarzi (1997) record Caryophyllaeus fimbriceps, C. laticeps and C. brachycollis from 38% of carp in the Anzali lagoon. Akhlagi (1999; 2000) reports that high temperatures (up to 32°C) stresses this species and leaves it open to infection with Aeromonas hydrophila. Safari and Khandagi (1999) record Clostridium botulinum from 3.8% of fresh and smoked samples of this species in Mazandaran Province. Mousavi and Khosravi (1999; www.mondialvet99.com, downloaded 31 May 2000) record the toxigenic fungi Aspergillus flavus, Alternaria, Penicillium and Fusarium from this species and the pond water at a fish farm in northern Iran. Farahnak (2000) records Anisakidae from this species in Khuzestan. Akhondzadeh et al. (2002) and Akhondzadeh Basti and Zahrae Salehi (2003) show that the psychotropic pathogen Listeria monocytogenes is found in market and fish farm samples of this species. Masoumian et al. (2002) investigated parasites from this fish in the Aras and Mahabad dams in northwest Iran and found the protozoan Goussia carpelli, also known from carp in the Safid River. Ebrahimzadeh Mousavi et al. (2005) isolated the fungus Branchiomyces spp. from gill lesions of farmed carp in northern Iran. Branchiomycosis or gill rot is a major problem in commercial fish production.

The intestinal helminth Bothriocephalus gowkongensis was recorded from this species on fish farms in West Azarbayjan Province (Azarvandi et al., 1999). Naem et al. (2002) found the following parasites on the gills of this species from the western branch of the Safid River, namely the protozoans Ichthyophthirius multifilis and a Trichodina species, a copepod crustacean Lernaea sp., monogenean trematodes Dactylogyrus anchoratum, and D. achmerowi. Jalali et al. (2002) and Jalali and Barzegar (2006) record parasites from this species in Lake Zarivar, namely Trichodina pediculus, Dactylogyrus extensus, Gyrodactylus stankovici, Diplostomum spathaceum two species of Argulus, Pseudocapillaria tomentosa and Lernaea cyprinacea. Jalali et al. (2005) summarise the occurrence of Gyrodactylus species in Iran and record G. cyprini, G. elegans, G. shulmani, G. sprostonae, G. stankovici and G. sp. from various localities for the carp. Farahnak et al. (2002) record Contracaecum sp. and Anisakis sp. from this fish in Khuzestan Province. Mehdipoor et al. (2004) record the monogenean Dactylogyrus extensus in Zayandeh River fish. Araghi Soureh and Jalali Jafari (2005) recorded Dactylogyrus extensus from this species in the Mahabad River of the Lake Orumiyeh basin. Jalali and Barzegar (2005c) record five species of monogeneans in the genus Dactylogyrus from both farmed and native Cyprinus carpio in Iran. These are D. extensus, D. anchoratus, D. achmerovi, D. vastator and D. sahuensis. Fry and fingerlings are more sensitive to these parasites and this sensitivity is increased with crowding in ponds. The paper also deals with gill histopathology and distribution of the parasites in Iran. Barzegar and Jalali (2006) report parasites in this species from Kaftar Lake as Lernaea cyprinacea,  Trichodina sp., Dermocystidium sp., Dactylogyrus extensus, D. anchoratus, Gyrodactylus sp. and Diplostomum spathaceum. Khara et al. (2006b) record the cestode Caryophyllaeus fimbriceps from this species in the Boojagh Wetland of the Caspian coast. Pazooki et al. (2007) recorded various parasites from localities in West Azarbayjan Province, namely Diplostomum spathaceum, Ligula intestinalis, Digrama sp. and Argulus foliaceus. Sattari et al. (2007) record the cestode Caryophyllaeus fimbriceps, the digenean Diplostomum spathaceum and the monogeneans Dactylogyrus extensus, Gyrodactylus sp. and Diplozoon sp. in this species in the Anzali wetland of the Caspian shore and also mention that the Caryophyllaeus laticeps is also known from this species in the Iranian Caspian Sea. Barzegar et al. (2008) record eye parasites from this fish including the monogenean Gyrodactylus stankovici, the digeneans Diplostomum spathaceum and Tylodelphys clavata, and the crustacean Lernaea cyprinacea. Khara et al. (2008) found the eye parasite Diplostomum spathaceum in this fish from Boojagh Kiashar Wetland in Gilan. Shamsi et al. (2009) found Dactylogyrus achmerovi, D. anchoratus, D. extensus, D. sahuensis and D. vastator in this species in fish farms, Safid River and Hamun Lake. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Argulus sp., Argulus foliaceus, Ergasilus sieboldi, Lernaea sp. and Lernaea cyprinacea on this species.

The Caspian seal, Pusa caspica, is a significant predator on this species (Krylov, 1984). Esox lucius, Sander lucioperca and Silurus glanis are all predators on small carp. A wide variety of other fishes and birds eat smaller carp as do predatory aquatic insects, frogs and toads. Carp eggs are eaten by many fishes. Adult carp are too large for most predators to take.

Economic importance

Balon (1974; 2006) details domestication of the carp which has been a cultivated fish for over 2000 years. The carp is an important table fish in Iraq where details of farming techniques are given by Ahmed and Taher (1988) and by Mangalo and Akbar (1988a; 1988b). Catches in former Soviet waters of the Caspian Sea were described by Kuliyev and Agayarova (1984). In the southwestern part, size range was 21-64 cm in 1960-1961 (92% of fish were 29-41 cm) but in 1972 95% of fish were 31-55 cm. The smallest carp are caught in the southeastern Caspian where maximum weight was 3.97 kg as opposed to 10.0 kg in the central Caspian. Spawning fish in the southeastern Caspian were only 0.58-1.0 kg while in the southwestern Caspian carp were 0.25-5.75 kg. About 17-30% of commercial catches in the Volga region are estimated to be comprised of hatchery production (Petr, 1987).

Nevraev (1929) gives catches for various fishing regions in Iran in the early twentieth century. For the Safid River region from 1899-1900 to 1917-1918 the catch was 30 to 30,500 individuals with no fish reported in some years, and in Astrabad (= Gorgan) region from 1900-1901 to 1912-1913 the catch was 14,200 to 851,800 individuals. The commercial catch in Iran from 1956/1957 to 1961/1962 varied from 3443 kg to 175,295 kg (Vladykov, 1964), from 1965/66 to 1968/69 varied from 184 to 333 tonnes (Andersskog, 1970) and from 1963 to 1967 from 7.0 to 108.8 t (on a yearly basis 41.1, 9.8, 7.0, 108.8, and 48.1 t respectively) (RaLonde and Walczak, 1970b). Catches from 1933/34 to 1961/62 varied from 34 kg to 1113 t in the Bandar Anzali region and the total catch of the Northern Shilot (Fisheries Company) varied from 9.8 t to 333 t from 1963/64 to 1968/69 (RaLonde and Walczak, 1972). The Food and Agriculture Organization, Rome gave the catches from Iran for the 6 years 1980 to 1985 as 1032, 2000, 1000, 52, 83, and 100 t respectively. The catch in the Anzali Mordab in 1990 was 6855 kg and from 1932 to 1964 the annual catch varied from 1.2-638.6 t (Holčík and Oláh, 1992). These wide fluctuations in reporting and in nominal catches are indicative of the poor fisheries statistics as well as the inter-year variability of catches in Iran.

March and April are the main fishing months in Iran (Farid-Pak, no date). It is the main fish in Iranian carp farms, when raised in polyculture with Chinese carps (Jalali and Molnár, 1990b). Salehi (2004) gives a figure of 25% as the share of common carp in Iranian polyculture (63% being silver carp) and gives a review of the economics of carp polyculture. Kohnehchahry and Heydarpur (1973) outline methods of raising carp using submerged cages which they believe would be suitable for Iranian waters. There were about 4000 ha of carp fish ponds (which presumably included Chinese carps such as Hypophthalmichthys spp.) with an annual production of 2-3 tonnes/ha (White, 1988). Plans were made in the 1980s to increase total pond area (including trout which was at 60 ha and produced 1000 t/year) to 35,000 ha over 10 years to yield an annual harvest of 100,000 t/year. However Bartley and Rana (1998b) report an aquaculture production of 6561 tonnes in 1995 and Salehi (2004) gives 28,060 t from warm-water fish farming, mostly Chinese carps. More than 700,000 carp fingerlings were released in the small and remote province of Chahar Mahall va Bakhtiari alone and 20 million carp, silver carp and grass carp fingerlings were produced in the Shahid Rajaae Hatchery in Sari for release across Iran in reservoirs and dams (Abzeeyan, Tehran, 4(7):VII, 1993). Salehi (1999) states that the marketable size of cultured carp is 1 kg with most harvested once annually and almost 90% supplied to market in October-March with a peak in March (the Iranian New Year when fish is a traditional food).

Marjan Iran Company was selling 1500-1800 g fish for U.S.$1.90/kg in August 2003 compared to $2.10 for Ctenopharyngodon idella (http://groups.yahoo.com/groups/hilsa/message/25). In Golestan Province, carp cost 1500-1700 tomans/kg in the early 2000s. Salehi (2006a) analysed the consumer market for this species and its products in Iran. Shabanpour et al. (2007) investigated preparation of surimi from this species.

An exotic species in some parts of the world, carp are a nuisance because they uproot vegetation used by native species for cover, food and spawning. This activity also increases water turbidity to levels which many native species cannot tolerate. Stirred up silt may smother eggs of native species. Carp also compete with other species for food and eat the eggs of other species.

Various studies have been carried out on its culture in Iran and neighbouring countries, e.g. see Freshwater Fishes of Iraq, and its use as an experimental organism, e.g. in Iran, see Mohagheghi and Hedayatifard (Farsi translation of Horváth et al. (1985a and 1985b); Khazraiinia et al. (2000) on acute ammonia toxicity; Peyghan et al. (2001) on use of ketamine and xylazine hydrochloride as anaesthetics; Rostami Bashman et al (2001) on histopathological lesions after exposure to copper, zinc, mercury and cadmium compounds; Soltani et al. (2001) on the anti-fungal effectiveness of formalin on hatch rate of eggs; Abtahi et al. (2002) on clove oil having no significant difference with MS222, another anaesthetic used in fish farms; Arabi and Heydarnejad (2002) on the deleterious effects of copper and mercury, used in combating algal blooms and weeds, on the gills of carp; Baradaran Noveyri et al. (2002) on the effects of cryopreservation on motility of spermatozoa; Peyghan et al. (2002) on differences between gonads of fishes from two culture seasons; Dorafshan et al. (2003) on induction of spawning using pituitary extract and GnRH analogue in combination with domperidone; Sharifpour et al. (2003) on the anaesthetic effects of clove oil under various pH and temperature regimes; Vahabzadeh Roodsari et al. (2003) comparing malachite green and hydrogen peroxide to control fungal infections (the latter is effective and less dangerous); Akhlaghi et al. (2004) on phagocytosis in relation to immunostimulants; Ebrahimi (2004) on the deleterious effects of copper, a pollutant, on sperm anatomy; Moini and Basimy (2004) on production of fish cake according to various recipes and its shelf life; Sharifpour (2004) on the histology of the response and the circumstances of wound healing; Soltani et al. (2004) on the effects of clove oil anaesthesia on haematological parameters, serum enzymes and some tissues, up to 200 p.p.m. being deemed safe; Oryan et al. (2005) on  the effects of baclofen on the pituitary system; Sharifpour et al. (2005) on the highly toxic effects of the pesticide endosulfan (with an LC50 of less than 0.1 mg/l); Yousefian (2005) on generating gynogenetic carp through irradiation; Baradaran Noveiri et al. (2006) on cryopreservation of spermatozoa using different extenders; Ghiasi and Mirzagar (2006) on lysozyme content in sub-lethal concentrations of cadmium; Kazemipour and Keivany (2005) on the use of garlic, mallow and motherwort in healing superficial wounds - garlic reduced recovery by one week; Kazemipour et al. (2006) on carp kept in aquaria at low concentrations of garlic (0.1 g/L) which healed superficial wounds more quickly than controls or aquaria having mallow or motherwort; Kohodabandeh and Abtahi (2006) on the use of sodium chloride, iodine and formalin to control Saprolegnia sp. on eggs (sodium chloride was recommended); Naji et al. (2007) on the 96h LC50 value of cobalt chloride at 327-328 mg/l; Naji et al. (2007) on the toxic effect of zinc sulphate on gill tissues which suffer hypertrophy and hyperplasia; Darafsh et al. (2008) on the use of scales as an indicator of heavy metal pollution; Golchinrad et al. (2008) on the effects of detergent on liver glycogen (decreased) and glucose (increased); Hasanabadizadeh et al. (2008) on the lack of improvement in wound healing after vitamin injections; Imnapour and Enayat Gholampour (2008) on the effect of broodstock migration time on various egg characters; Safari et al. (2008) on changes in muscle chemistry during maturation; Soleymani et al. (2008) on the effectiveness of vitamin C injections on survival of juveniles challenged by different doses of the theront of Ichthyophthirius multifilis; Darvish Bastami and Imanpour (2009) on sperm motility influenced by high concentration of ions; Baghfalki et al. (2009) on the effect of broodstock density on survival and growth of larvae and fingerlings in earthen ponds; Erfani Majd et al. (2009) on evaluating the response of incubated ovarian follicles to carp pituitary extract and cultivated pituitary cells secretion; Ghanbari et al. (2009) on the long term effects of changes in pH on haematological parameters in fingerlings; Imanpour and Safari (2009) on the effect of maturation stages on gonadal indices and chemical composition of the gonad; Imanpour et al. (2009) on stocking density and its effect on survival and growth in polyculture, up to 450 fish per hectare being optimal; Imanpour et al. (2009) on the effects of broodstock age on various egg dimensions, females age 3-6 years being found suitable for propagation; Kordjazi et al. (2009) on physicochemical parameters of water and their correlation with haematocrit indicators, growth and survival in farm ponds; Rostami Mina and Soltani (2009) on the histopathological effects of the aquaculture pollutant copper sulphate; Salamat et al. (2009) on pituitary primary cell culture and its secretion effect on endocrine activity of incubated ovarian follicles; Sheykhzadeh et al. (2009) on the effects of Eucalyptus essential oil on immunological variables; Yehganeh et al. (2009) on seasonal variation in the chemical composition and fatty acid profile of ovaries, necessary for embryogenesis; Ghiasi et al. (2010) on the effects of low concentrations of cadmium on the immune response in winter; Salamat et al. (2010) on ovarian follicular cells and their endocrine activity in cell culture; Soltani et al. (2010) on the immune responses to Zataria multiflora (Persian thyme) essential oil used as an anti-fungal for carp eggs; etc.

This species is actively angled for along the Caspian shore of Iran and in its rivers (e.g. see Noorbakhsh (1993a)), appears regularly in fish markets of Ahvaz, Khuzestan and is caught by anglers there using bread or potato as bait.

It is characterised as a fatty fish according to a lipid content 9-14% by wet weight of muscle in autumn (Hantoush et al., 1999). Al-Aswad et al. (1980) detailed the chemical composition of this species in Dukan Lake, Iraq including seasonal levels of moisture, fat, protein and ash, and the various types of fatty acids and amino acids. Hindi et al. (1996a) give the chemical composition of flesh of this species as 78.87% moisture, 2.46% fat, 17.06% protein and 1.35% ash, indicating a valuable food fish characterised as lean to medium fatty. Hindi et al. (1996b) give chemical indices for assessing fish freshness according to the month of capture and marketing (pH 6.28, total volatile nitrogen bases 11.07 mg/N/100g fish, thiobarbituric acid 0.47 mg, and free fatty acids 0.62%).

The roe or eggs of this species have been implicated in poisoning (Halstead, 1967-1970; Coad, 1979b) and should be avoided (see under the genus Schizothorax for more information on egg poisoning). Fish should be carefully cleaned in the spawning season to remove the eggs and ensure against contamination of flesh. Severe cases of egg poisoning in other species have resulted in death. Vaezzadeh et al. (2008) found the levels of the pesticide heptachlor in fish from Anzali and Ramsar could have a health risk to consumers.

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in aquaria and aquaculture, as food, in textbooks, for sport, as an experimental species and because it has been introduced outside its natural range. Brightly coloured varieties of carp are known as "koi" and are kept as ornamental fish. Colours include red, orange, white, black, blue and yellow in various combinations.

Conservation

Vladykov (1964) recommended that fishing for this species in Mazandaran and Gorgan be prohibited for 5 years because of reduced stocks. Krasznai (1987) and Petr (1987) give details of fish farms propagating this species in Iran. For example, 30 million fish were produced by the Safid Rud Fish Farm in 1986. In 1999-2000, 20 million juveniles were released into the Caspian Sea (Iranian Fisheries Research Organization Newsletter, 23:4, 2000). From October to March 2000, 3 million juveniles raised in the Shahid Ansari aquaculture and breeding centre in Gilan were released into the Caspian Sea and neighbouring water bodies (Iranian Fisheries Research Organization Newsletter, 26:2, 2001). Poaching was a problem in the Caspian basin of Iran (Razivi et al., 1972) and no doubt continues. The Atrak River stocks are an important fishery for both Iran and Turkmenistan but are susceptible to loss through the absence of flooding of the spawning grounds. Fish passes are needed to ensure access to the spawning grounds, timely release of water from a reservoir to flood the spawning areas in years of low water flow, enforcement of catch limits, and continued stocking (Petr, 1987). Yousefian et al. (2009) examined the normative reproduction values and genetic characters for native carp in the Caspian Sea. The average weight of sampled fish was 1441.6 g and standard length was 50.5 cm. Fertilisation rate was 62-79%, absolute fecundity was 114,347 and relative fecundity 82,007. There was no significant differences in haplotype distribution between areas in the southern Caspian Sea.

Lelek (1987) considered that the wild form, as opposed to domestic stocks, was vulnerable to endangered in Europe because of habitat modifications. Kiabi et al. (1999) consider this species to be of least concern in the south Caspian Sea basin according to IUCN criteria. Criteria include commercial fishing, sport fishing, abundant in numbers, habitat destruction, widespread range (75% of water bodies), present in other water bodies in Iran, and present outside the Caspian Sea basin. Karami et al. (2005) found river fish had greater exposure to organophosphorus compounds than Caspian Sea and well-fed fish.

Further work

Where this species is introduced, its interactions with native species should be carefully studied and monitored in an effort to protect the native fauna.

Sources

A recent summary on this species is Bănărescu, Barus and Peňáz in Bănărescu and Paepke (2002).

Iranian material: CMNFI 1970-0510, 1, 47.8 mm standard length, Gilan, Golshan River (37º26'N, 49º40'E); CMNFI 1970-0522, 1, 108.5 mm standard length, Gilan, Safid River at Astaneh Bridge (37º16'30"N, 49º56'E); CMNFI 1970-0563, 3, 28.6-58.6 mm standard length, Gilan, Caspian Sea at Kazian Beach (ca. 37º29'N, ca. 49º29'E); CMNFI 1970-0568, 1, 125.8 mm standard length, Gilan, Caspian Sea at Kazian Beach (ca. 37º29'N, ca. 49º29'E); CMNFI 1970-0582, 3, 63.3- 134.0 mm standard length, Mazandaran, Aliabad Reservoir (36º56'N, 54º50'E); CMNFI 1970-0587, 2, 47.5-76.7 mm standard length, Mazandaran, Babol River at Babol Sar (36º43'N, 52º39'E); CMNFI 1979-0455, 6, 46.9-67.1 mm standard length, Markazi, Manjil Dam (36º45'N, 49º17'E); CMNFI 1979-0476, 3, 44.1-85.3 mm standard length, Mazandaran, Qareh Su near Kord Kuy (36º51'N, 54º05'E); CMNFI 1979-0479, 19, 28.0-143.8 mm standard length, Mazandaran, dam on Gorgan River (37º09'30"N, 54º41'30"E); CMNFI 1979-0685, 2, 31.5-45.7 mm standard length, Gilan, Safid River (ca. 37º22'N, ca. 49º57'E); CMNFI 1979-0788, 2, 104.2-123.5 mm standard length, Mazandaran, Gorgan River at Khadje Nafas (37º00'N, 54º07'E); CMNFI 1980-0128, 3, 60.2-75.2 mm standard length, Mazandaran, Qareh Su (36º49'30"N, 54º03'30"E); CMNFI 1980-0132, 4, 56.0-64.6 mm standard length, Gilan, Safid River at Kisom (37º12'N, 49º54'E); CMNFI 1980-0157, 1, 126.3 mm standard length, Mazandaran, Gorgan River estuary (36º59'N, 53º59'30"E); CMNFI 1980-0905, 2, 78.1-92.9 mm standard length, Mazandaran, Gorgan River at Khadje Nafas (37º00'N, 54º07'E); CMNFI 1980-0908, 1, 54.4 mm standard length, Gilan, Safid River estuary (ca. 37º28'N, ca. 49º54'E).

Genus Garra
Hamilton, 1822

The genus Garra is found throughout Southwest Asia and from Africa to southeast Asia. There are about 73 species and 4 are recognised from Iran. The genera Discognathus Heckel, 1843 and Discognathichthys Bleeker, 1859 are synonyms. Hora (1921) presents anatomical arguments for including Discognathus in Garra.

This genus is characterised by a small to moderate-sized body, elongate and almost cylindrical, a rounded snout with the mouth inferior and crescent-shaped, the lower jaw has a horny edge, the upper lip is usually fringed and continuous with the snout, the lower lip and chin area modified into a suctorial disc with free posterior margin (in Iran, elsewhere a some have a callous pad (Stiassny and Getahun, 2007) and the smallest specimens lack full disc development), the anterior disc margin free or adherent (species with the latter condition were placed in a separate genus, Discognathus or Discognathichthys), 1 or 2 pairs of short barbels (species with the former condition were placed in a separate genus, Discognathus or Discognathichthys), vomero-palatine organ vestigial or regressed, eyes small, usually large scales, lateral line complete, small dorsal and anal fins without thickened rays, pectoral and pelvic fins placed horizontally on the body, first two or more pectoral fin rays prominent and often unbranched, pharyngeal teeth in 3 rows (typically 2,4,5-5,4,2) with hook-shaped tips and spoon-shaped crowns, vent may be midway between pelvic and anal fin bases or nearer the latter, elongate and coiled gut, a black peritoneum, and 2n=50.

The Farsi common name used generally for these fishes is gel cheragh (= mud-eater, mud-grazer), سنگ ليس (sang lis) and ماهي سنگي (mahi sangi), not repeated in each Species Account.

These fishes are found in mountains streams and other flowing waters, maintaining position with their suctorial disc, reduced swimbladder, flattened belly and large, splayed and horizontal paired fins. Also known to occur in slow-moving or stagnant waters. They scrape algae from rocks. These are oily fishes which are eaten in India (Hora, 1956).

Menon (1954) considers that the members of this genus spread westwards along the Himalayas as late as the early Pleistocene. Kosswig (1952) indicates their presence in the Araxes (= Aras) of Turkey but this seems to an error.

Garra persica
Berg, 1913

Common names

Persian stone lapper.

Systematics

This species is recognised only as a subspecies of Garra rufa by Bianco and Banarescu (1982) while Menon (1964) and Karaman (1971) synonymise it with Garra rufa. Karyotype analysis separates this species from Garra rufa (Esmaeili et al., 2009).

The syntype specimens are in the Zoological Institute, St. Petersburg under catalogue numbers ZISP 11707 (6 specimens from the "River Bampur in Eastern Persia. N. Zarudnyi 1898, 15-27.VII") and 11706 (1 specimen from "Kiabad in Zirkuh (Eastern Khorassan). N. Zarudnyi 1898, 3.V") according to Berg (1913) where the original description is founded on these fish, implying all are types. The latter is also given as "settlement Kiabad between Zirkuh province and Sistan" in the catalogue (this locality may be at or near Kuh-e Ziri at 32°48'N, 59°50'E according to Coad (1981d)). These dates are old style and corrected to new in Berg (1949) (27.VII-8.VIII and 15.V respectively). In St. Petersburg under ZISP 11707 there are 10 fish 24.0-46.5 mm standard length and ZISP 11706 is not listed as a type in the catalogue nor in Berg (1949). Berg (1949) lists 10 fish in 11707 too. These specimens were formerly identified as Discognathus lamta by Nikol'skii (1899) who lists 1 fish in 11706 and 6 in 11707. Three syntypes are in the Zoological Survey of India, Calcutta (ZSI F11101/1) listed under Garra rufa obtusa and received from the Zoological Institute, St. Petersburg, Russia on exchange (Menon and Yazdani, 1968). There are more apparent types available than those listed by Berg (1913).

Key characters

Two pairs of barbels are present, the adhesive disc is well developed with a free posterior margin, the dorsal fin has 7 branched rays, and the caudal fin modally 16 branched rays. The caudal fin ray count is unique in cyprinids from Iran and very rare elsewhere. Almost all cyprinids show a strong mode of 17 branched caudal fin rays.

Morphology

Dorsal fin with 4 unbranched and 6-8 branched rays, anal fin with 2-3 unbranched and 4-6 branched rays, branched pectoral fin rays 12-16, pelvic fin branched rays 6-8, caudal fin branched rays number 15-17 with a strong mode at 16, lateral line scales 28-38, total gill rakers 15-22 with lower counts in smaller fish, pharyngeal teeth usually 2,4,5-5,4,2, and total vertebrae 34-35.

Scales are regularly arranged and only lacking on the anterior isthmus in some fish. A pelvic axillary scale is present but is not always well-developed. Scales have numerous radii on all fields with the focus broken up into a network of lines. There are 4 short barbels. Upper lip well-developed, the rostral fold weakly fringed. The adhesive disc and rostral fold are greatly papillose although the disc centre is not as papillose as the margins. The gut is very elongate and greatly coiled. Chromosome number 2n=48 with 15m, 8Sm and 1St (Esmaeili et al., 2009).

Counts for 12 Iranian topotypic specimens from the Bampur River are as follows:- dorsal fin with 4(12) unbranched and 7(12) branched rays; anal fin with 3(12) unbranched and 4(3) or 5(9) branched rays; branched pectoral fin rays 12(1), 13(6) or 14(4); caudal fin branched rays number 15(1), 16(8) or 17(1) in the type series and 15(1), 16(10) or 17(1) in topotypes; lateral line scales 29(2), 30(1), 31(1), 32(2), 33(2), 34(2) or 35(2); total gill rakers 15(6); pharyngeal teeth 2,4,5-5,4,2(5); and total vertebrae 34(9) or 35(3).

Meristic values for topotypes and other material: dorsal fin branched rays 6(3), 7(115) or 8(4); anal fin branched rays 4(5) or 5(118); branched pectoral fin rays 12(8), 13(40), 14(49), 15(20) or 16(5); branched pelvic fin rays 6(9), 7(100) or 8(13); caudal fin branched rays number 15(10), 16(113) or 17(9); lateral line scales 28(1), 29(5), 30(5), 31(77), 32(18), 33(28), 34(20), 35(22), 36(11), 37(4) or 38(2); total gill rakers 15(16), 16(13), 17(18), 18(17), 19(11), 20(4) or 21(2); pharyngeal teeth 2,4,5-5,4,2(5); and total vertebrae 32(4), 33(44), 34(34) or 35(4).

Sexual dimorphism

Males develop numerous breeding tubercles around the snout, between the nostril and the eye and between the nostrils. There is a transverse depression anterior to the nostrils on the snout. A postspawning individual from the Hormuz basin measuring 148.7 mm SL has small tubercles under the eye running forward onto the snout as a band, the most evident tuberculation. The swollen snout tip bears no tubercles. The top of the head has tubercles but these are smaller and sparser than the band under the eye. Scattered large tubercles are present on the gill cover. Evident tubercles line the dorsal, anal, caudal and pectoral fin rays (pelvic fins not present on specimen), the largest being those on the anal fin.

Colour

The back and flanks are an orange-brown to golden-brown. There is a blue spot on the flank near the postero-dorsal edge of the operculum (dark black in preserved fish). The dorsal fin bears elongate blotches on the posterior half of each fin membrane, usually fading distally, but in some fish occupying the whole membrane. Proximally there is a gap between these blotches and 3-5 bars which originate at the posterior edge of the base of branched ray three and succeeding rays, and extend distally across the ray and then along the ray and the membrane to the gap. These bars are much more heavily pigmented than the dorsal blotches. There is a bluish tinge or spot around the pectoral fin base, sometimes developed as a bar along the edge of the gill cleft, becoming dark blue dorsally. There is a dark bar or a roundish, poorly-delimited spot on the caudal peduncle at the base of the caudal fin. Fins are generally pink to light orange. The caudal fin pigmentation is individually variable. Some are blotched irregularly on both rays and membranes, in others there is a trace of a band in mid-fin extending from the dorsal to the ventral margin following the posterior outline of the fin, while others have pigment heavily concentrated only in the mid-fin clear of the margins. The pelvic fin has little or no pigment and the anal fin has a very few irregular light blotches on both rays and membranes. The pectoral fin is pigmented near the dorsal base with some pigment on anterior rays and membranes. In live fish the paired fins are a light orange and other fins show less marked orange tinges. The peritoneum is black.

Size

Attains 7.5 cm total length (Berg, 1913).

Distribution

This species is found in the Hormuz, Makran and Hamun-e Jaz Murian basins and possibly the Sistan and Kerman-Na'in basins (Bianco and Banarescu, 1982; Abdoli, 2000). Localities include springs in the upper Kul River basin near Darab, small springs in the upper Sarbaz River, and at Jaghin.

Zoogeography

The species of the genus Garra are thought by Menon (1964) to have colonised Iran from a centre of origin in southern China by a series of "waves". The earliest wave arrived in the Miocene and is represented in Iran today by the species rossica and variabilis, characterised by the primitive condition of a weakly-developed adhesive disc without a free posterior border, the posterior chamber of the swimbladder is cylindrical and well-developed, there is no proboscis on the snout and the vent is close to the anal fin. A second wave is represented in Iran by rufa (and by implication persica) characterised by a well-developed disc and a tuberculated snout marked off by a transverse groove. There were 6 "waves" all told but only the first two are relevant to Iran. Karaman (1971) criticises this complex interpretation on two grounds. He maintains that it is unlikely that fishes would immigrate from southern China to Iran but leave no extant forms between these two remote places and that the species assigned to the various waves show no characteristics which would make them more adaptive and capable of replacing earlier wave members. The characters of rossica and variabilis (one pair of barbels, weak disc, reduced squamation) could equally be loss characters and a more recent specialisation rather than the primitive condition.

Habitat

Kiabi and Abdoli (2000) found this species to have the widest altitudinal range in Hormozgan Province.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

None reported from Iran.

Economic importance

None.

Conservation

This species is widely distributed in various smaller water bodies in eastern Iran and does not appear to be under threat.

Further work

The biology of this species remains unknown and would repay study. It is a significant component of a variety of water bodies.

Sources

Type material: See above, ZISP 11707 and 11706.

Iranian material: CMNFI 1979-0138, 1, 25.7 mm standard length, Fars-Hormozgan, Rasul River drainage (ca. 27º32'N, ca. 54º58'30"E); CMNFI 1979-0139, 1, 30.6 mm standard length, Fars-Hormozgan, Rasul River drainage (ca. 27º25'30"N, ca. 54º59'E); CMNFI 1979-0144, 1, 27.3 mm standard length, Hormozgan, Minab River at Minab (27º09'30"N, 57º04'E); CMNFI 1979-0145, 4, 14.8-25.4 mm standard length, Hormozgan, Geru River drainage (26º55'N, 57º01'30"E); CMNFI 1979-0149, 7, 29.0-49.4 mm standard length, Hormozgan, stream north of Bandar Abbas (27º36'N, 56º14'E); CMNFI 1979-0152, 1, 62.2 mm standard length, Hormozgan, Shur River drainage (28º09'N, 55º43'E); CMNFI 1979-0178, 23, 25.1-66.9 mm standard length, Hormozgan, Sarzeh River draiange (27º36'N, 56º15'E); CMNFI 1979-0180, 1, 42.7 mm standard length, Hormozgan, stream 3 km east of Essin (27º19'N, 56º17'30"E); CMNFI 1979-0181, 1, 44.0 mm standard length, Hormozgan, Kul River drainage (27º17'30"N, 56º03'30"E); CMNFI 1979-0186, 8, 30.2-64.6 mm standard length, Hormozgan, stream and pools at Sar Khun (ca. 27º24'30"N, ca. 56º25'E); CMNFI 1979-0187, 9, 32.1-57.9 mm standard length, Hormozgan, stream and pools at Sar Khun (ca. 27º23'30"N, ca. 56º26'E); CMNFI 1979-0312, 10, 26.6-35.6 mm standard length, Baluchestan, dam on Bampur River (27º11'N, 60º46'E); CMNFI 1979-0315, 1, 23.8 mm standard length, Baluchestan, Bampur River 2 km north of Karevandar (27º51'N, 60º46'E); CMNFI 1979-0324, 1, 29.6 mm standard length, Baluchestan, Bampur River at Sa'idabad (27º11'N, 60º22'E); CMNFI 1979-0329, 2, 25.4-30.8 mm standard length, Baluchestan, stream at Zaminbandan (27º02'N, 61º20'E); CMNFI 1979-0411, 1, 60.4 mm standard length, Hormozgan, Minab River (27º24'N, 57º12'E); CMNFI 1979-0412, 9, 22.9-39.3 mm standard length, Hormozgan, spring at Saras (27º30'N, 57º34'E); CMNFI 1979-0416, 39, 15.1-46.8 mm standard length, Hormozgan, Ab Garm-e Ganow (ca. 27º26'N, ca. 56º20'E); CMNFI 2007-0051, 10, 29.5-43.7 mm standard length, Hormozgan, upper Kol River basin (28º19'N, 55º55'E); CMNFI 2007-0055, 5, 30.9-44.6 mm standard length, Hormozgan, Minab River basin (27º47'N, 57º12'E); CMNFI 2007-0056, 2, 32.1-54.2 mm standard length, Kerman, qanat at Kahnuj (27º58'N, 57º45'E); CMNFI 2007-0058, 7, 36.7-51.7 mm standard length, Fars, headwaters of Gowdar River (ca. 27º24'N, ca. 54º16'E).

Garra rossica
(Nikol'skii, 1900)

Common names

mahi-e sangi, mahi-e sang lis, anjarak.

[diskognat in Russian; pathar chat or patherchatta in Pakistan].

Systematics

Discognathus phryne Annandale, 1919 described "from Seistan" is a synonym. Menon and Yazdani (1968) give Nasratabad, Seistan as the type locality. G. rossica has been placed in the genera Discognathus Heckel, 1843 and Discognathichthys Bleeker, 1859, here considered to be synonyms.

Discognathus rossicus var. nudiventris Berg, 1905 was described from "Schiwar" in Iran for specimens with a naked abdomen, thoracic region and groove on the back anteriorly. The distribution of these specimens overlaps with that of the type form and they are not given independent taxonomic recognition here. Berg (1949) later placed them as an infraspecies.

The syntypes of Discognathus rossicus are in the Zoological Institute, St. Petersburg (ZISP 10365), the type locality in Latin on page 239 being "Flum. Tedschent in prov. Transcasp. Zarudnyi. 1892 (4)" while on p. 240 are the localities "Habitat in flumine Tedshent in provincia Transcaspiensi, nec noc in Persia orientale ad Kirmanum orientale" (Nikol'skii, 1900), and confirmed by me. However there were 3 fish in the jar (45.0-54.5 mm standard length) although 4 are listed in the catalogue and in the type description. Berg (1905) lists 3 fish but in Berg (1949) lists only 2. Other materials listed by Nikol'skii (1900) from eastern Iran and Kerman (ZISP 11113, 11703, 11704, 11705, 11708) are apparently not types of rossicus although Berg (1949) indicates that 11704 ("Neizar in Seistano") and 11705 ("Ljabeab in Seistano") are part of the type series from Iran. Eschmeyer et al. (1996) also list ZIL (= ZISP) 10665 (4) as part of the type series, perhaps a misprint for ZISP 10365.

The types for the var. nudiventris are in ZISP 11113, listed by Berg (1905) as 2 fish from Schivar, by Berg (1949) as being 4 fish, not numbered in the ZISP catalogue and with 5 fish in the jar (45.6-66.2 mm standard length). ZISP 11113 appears to have been renumbered in part as 11703 and 11708 according to Berg (1949). ZISP 11708 is listed in Berg (1949) as 13 specimens and 11703 seems to be also 13 specimens. The type localities for var. nudiventris are for ZISP 11113 "Shivar, north of Nikh (Nekh), north-east Kerman, basin of L. Hamun, 23 VI 1896, N. Zarudnyi", for ZISP 11708 "Podaghi, 28°08'N, north-north-east of Bazman, eastern Kerman, 6 VII 1898, N. Zarudnyi" and 11703 is probably "Neizar in Seistan, N. Zarudnyi" according to Berg (1949) (the catalogue number 11703 does not appear under the description of materials in Berg (1949), possibly omitted in error, and it is deduced here that it should have preceded the locality cited). This is not critical as the variety or infraspecies has no taxonomic status.

The holotype of Discognathus phryne is in the Zoological Survey of India, Calcutta (ZSI F9787/1) (Annandale, 1919b; Menon and Yazdani, 1968), a syntype (listed as a cotype) measuring 42.2 mm standard length from "Baluchistan" with the annotation "Ind. Mus. Ex. F 9789/1" is in the Natural History Museum, London (BM(NH) 1919.8.16:1) with another syntype (cotype) from "Quetta" measuring 32.2 mm standard length with the annotation "Ind. Mus. Ex. F 9790/1" (BM(NH) 1919.8.16:2).

Key characters

The single pair of small maxillary barbels (sometimes an anterior pair), absence of a free anterior margin to a weakly developed adhesive disc on the lower head surface, gill raker count, and distribution distinguishes this species. It is separated from the closely related, but geographically separated G. variabilis by smaller size, head length longer than caudal peduncle length, head length equal to or longer than pectoral fin length, distinctly emarginate caudal fin, and dorsal fin origin mid-way between snout tip and caudal fin base or closer to caudal fin base (Berg, 1949).

Morphology

Dorsal fin with 2-3 unbranched and 6-7, usually 7, branched rays, anal fin with 2-3 unbranched and 5 branched rays, pectoral fin with 11-16 branched rays and pelvic fin with 7-8 branched rays. Lateral line scales 33-46. The mid-line of the back, and the chest and belly are naked in some populations. Some fish have only 3-4 rows of scales below the lateral line and this character is extremely variable (Berg, 1949). Scales are a vertical ovoid with an anterior or subcentral anterior focus. The posterior scale margin is rounded and elongate, the dorsal and ventral margins are rounded and merge into the posterior margin and the anterior margin has slight indentations above and below a shallow, rounded central protuberance. However some scales may be squarish with rounded corners and scale shape can be very variable. Circuli are fine. Radii are found on all fields, moderate to very numerous although this is individually variable. A pelvic axillary scale may be present or absent. Gill rakers on the lower arm 9-11, 10-12 total (but see below), almost reaching the adjacent raker when appressed. Pharyngeal teeth 2,4,5-5,4,2, 3,4,5-5,4,3, 2,4,5-5,4,3, 3,4,5-5,4,2, 2,4,5-5,3,2, 2,4,5-5,4,1, 2,4,4-4,4,2, 2,3,5-5,3,2 or 1,3,5-5,3,1. Teeth are conical to flattened with oblique but flattened crown which is slightly concave. Rarely crowns are blade-like and lack the flattened crown. Specimens from the Hari River basin have scales on the thorax and ridge of the back in front of the dorsal fin while those from Sistan lack scales in these areas (Annandale and Hora, 1920; Saadati, 1977). The anterior pair of barbels are usually absent but may be present and minute or even moderately well-developed. Some fish may lack barbels entirely. The gut is very elongate and coiled.

Meristics for Iranian specimens:- dorsal fin branched rays 6(5) or 7(54); anal fin branched rays 5(59); pectoral fin branched rays 11(2), 12(14), 13(14), 14(15), 15(8) or 16(4); pelvic fin branched rays 7(39) or 8(14); lateral line scales 34(3), 35(6), 36(17), 37(10), 38(4), 39(7), 40(1), 43(1), 44(1), 45(1) or 46(1); total gill rakers 11(7), 12(18), 13(18), 14(9), 15(3) or 16(2); pharyngeal teeth 2,4,5-5,4,2 (15), 2,4,5-5,3,2(2), 2,4,5-5,4,1(1), 3,4,5-5,4,2(1) or 2,4,5-5,4,3(1); and total vertebrae 34(11), 35(37) or 36(11).

Sexual dimorphism

Males in spawning condition bear small but evident tubercles on the operculum and the head above the operculum, fine tubercles on top of and anteriorly on the head, small tubercle son the pectoral fin rays following the branching of the rays and on the first unbranched ray, and there are a few minute tubercles on anterior flank scales.

Colour

The upper flanks and back are dark to greyish-brown, greenish-brown or golden-brown and there may be large spots on the upper flanks. The body is silvery overall. There is often a dark bar at the base of the tail. Fins are colourless and the belly and lower head surface are white to yellowish-white. The belly and lower head may be bright yellow as are the neighbouring fin bases. The bases of dorsal fin branched rays 3-5, and sometimes 6, have small dark spots. Young have a bluish mid-lateral stripe along the flank. The peritoneum is black. Colour is darker in clear than in muddy water.

Size

Reaches 9.5 cm.

Distribution

This species is found in the Mashkel (= Mashkid) River basin of Pakistan (Mirza, 1992), the Tedzhen and Murgab River drainages of the former U.S.S.R., and the Tedzhen River, Bejestan, Sistan, Dasht-e Lut, Hamun-e Jaz Murian and Makran basins of Iran (Nikol'skii, 1900; Berg, 1905; Berg, 1949; Menon, 1964; Spillman, 1972; Abdoli, 2000). Localities from these authors include the following and indicate the wide and common distribution of the species: Nekh, Shivar north of Nekh basin of Lake Hamun, Lab-i-ab, Hamun-e Farah in the Sistan basin, Deh Salm in the Lut basin, Laadas, Sarhad Region near Kuh-i Taftan, Podaghi north-northeast of Bazman, Chanf between the Makran and the Bampur River, headwaters of the Sarbaz River, Salabad 130 km south of Birjand, Shusef qanat, Khunik Pa'in and Khaneh Sharef on Birjand to Zahedan highway, and Hormak south of Zabol.

Records of Discognathus variabilis Heckel, 1843 from Sistan by Nikolskii (1899) and Regan (1906) are G. rossica (Menon, 1964).

Zoogeography

See under Garra persica. G. rossica is related to Garra variabilis of the Tigris-Euphrates basin.

Habitat

Found in pools and slow-flowing ditches and channels and in reed beds in Sistan.

Age and growth

Sexual maturity is attained at 2-3 years. Females grow more rapidly than males.

Food

Gut contents include green filamentous algae, higher plant fragments and sand grains.

Reproduction

Spawning occurs in the summer and up to 984 eggs are produced. Some fish still contain undeposited eggs in late July. Egg diameter is up to 1.06 mm (Nikol'skii, 1945). In southern Iran, fish with 1.1 mm eggs have been collected on 14 November and on 2 December eggs were 1.3 mm. On 8 May eggs were 1.2 mm suggesting either early spawning or prolonged retention of eggs. The most tuberculate males were found in both November and May.

Parasites and predators

None reported from Iran.

Economic importance

None.

Conservation

This species is widely distributed in eastern Iran and does not appear to be under significant threat.

Further work

Further studies on its biology would clarify details of life history. Its wide distribution in isolated basins may have led to some divergence and speciation that is not readily detected with morphological studies and molecular work on this species might be rewarding.

Sources

Type material: See above, Discognathus rossicus (ZISP 10365), var. nudiventris (ZISP 11113), and Discognathus phryne (BM(NH) 1919.8.16:1 and BM(NH) 1919.8.16:2).

Iranian material: CMNFI 1979-0091, 5, 24.6-58.3 mm standard length, Khorasan, qanat at Nehbandan (31º32'N, 60º02'E); CMNFI 1979-0226, 5, 37.2-49.0 mm standard length, Sistan, pool near Kuh-e Khajeh (30º57'N, 61º17'E); CMNFI 1979-0227, 6, 48.1-61.1 mm standard length, Sistan, naizar at Kuh-E Khajeh (30º57'N, 61º16'E); CMNFI 1979-0230, 2, 29.2-43.4 mm standard length, Sistan, Jehil-e Puzak at Gohoor-ghoori (ca. 31º15'N, ca. 61º42'E); CMNFI 1979-0236, 7, 18.3-47.3 mm standard length, Sistan, irrigation ditch 27 km from Zabol (ca. 30º52'N, ca. 61º22'E); CMNFI 1979-0238, 11, 15.3-29.2 mm standard length, Sistan, irrigation ditch 11 km south of Zabol (30º57'N, 61º27'30"E); CMNFI 1979-0315, 2, 24.9-50.3 mm standard length, Baluchestan, Bampur River 2 km north of Karevandar (27º51'N, 60º46'E); CMNFI 1979-0316, 9, 35.7-53.4 mm standard length, Baluchestan, stream 68 km south of Iranshahr (26º48'N, 61º02'E); CMNFI 1979-0326, 1, 39.0 mm standard length, Baluchestan, Ughin River south of Pip (ca. 26º35'N, ca. 60º02'E); CMNFI 1979-0327, 5, 37.3-46.7 mm standard length, Baluchestan, stream 26 km south of Pip (26º32'N, 59º57'E); CMNFI 1979-0330, 68, 14.7-65.1 mm standard length, Baluchestan, stream 22 km west of Qaleh-ye Zaboli (27º02'30"N, 61º26'E); CMNFI 1979-0336, 30, 22.4-31.6 mm standard length, Baluchestan, qanat 7 km from Khash (28º10'N, 61º15'E); CMNFI 1979-0339, 3, 40.6-51.0 mm standard length, Baluchestan, Tahlab River drainage 16 km from Mirjaveh (28º56'30"N, 61º21'E); CMNFI 2007-0025, 8, 36.6-47.1 mm standard length, Khorasan, qanat south of Birjand (ca. 32º24'N, ca. 59º49'E); CMNFI 2007-0026, 19, 36.3-62.9 mm standard length, Khorasan, qanat at Shusf (31º48'N, 60º01'E); CMNFI 2007-0027, 13, 31.4-60.7 mm standard length, Khorasan, qanat at Khvansharaf (31º34'N, 60º06'E); CMNFI 2007-0028, 13, 36.3-58.9 mm standard length, Khorasan, qanat at Khunik-e Pa'in (31º28'N, 60º06'E); CMNFI 2007-0029, 7, 35.2-60.7 mm standard length, Baluchestan, qanat at Hormak (29º58'N, 60º51'E); CMNFI 2007-0031, 2, 36.0-45.5 mm standard length, Baluchestan, headwater of Bampur River (27º51'N, 60º46'E); CMNFI 2007-0035, 9, 28.9-50.1 mm standard length, Baluchestan, stream west of Zaboli (ca. 26º58'N, ca. 61º27'E); USNM 205905, 6, 30.2-36.9 mm standard length, Baluchestan, small springs in upper Sarbaz River basin (no other locality data).

Garra rufa
(Heckel, 1843)

Garra rufa courtesy of F. Krupp, Senckenberg Museum, Frankfurt.

Garra rufa from Wikimedia Commons.

Common names

mahi-e sangi, mahi-e sang lis, shirbot, gel khorok, gel ra (and even gararufa or gara in Farsi).

[djulake; kokur ahmar or karkoor ahmar; garagoor; gassur achmar (= red gassur) or gassur hadjari (gassur of the pilgrims) at Aleppo (= Haleb, Syria) according to Heckel (1843b), all in Arabic; red garra].

Systematics

Discognathus obtusus Heckel, 1843 described from "Aleppo" and "Mossul", Discognathus crenulatus Heckel, 1849 described from the "Confluenten des Araxes, als aus den Quellen des Saadi und dem Kara-Agatsch" (= probably includes the Pulvar (= Sivan) River near Persepolis; Sa`di at 29°37'N, 52°35'E, now within the city of Shiraz; and the Qarah Aqaj River; all in Fars), Garra rufa gymnothorax Berg, 1949 from "Kulihan, Karun R. basin, 6 IV 1904, N. Zarudnyi" and Garra rufa turcica Karaman (1971) from the Ceyhan River basin in Turkey are synonyms (Krupp, 1985c). Records of Garra lamta (Hamilton, 1822) from Iran are in error (Menon, 1964).

The types of Discognathus rufus are from "Aleppo" according to Heckel (1843b). The syntypes of Discognathus rufus according to Krupp (1985c) are in the Naturhistorisches Museum Wien under NMW 53240, 8 specimens, 59-108 mm standard length from Aleppo and 1 syntype is in the Senckenberg Museum Frankfurt under SMF 553 (formerly NMW), 103 mm standard length and also from Aleppo. The catalogue in Vienna lists 6 specimens. One specimen from NMW 53240, 112.3 mm standard length (as measured by me), was designated as the lectotype and 7 fish, 60.2-97.5 mm standard length as paralectotypes by F. Krupp, 29 October 1984, and published in Krupp and Schneider (1989).

Four syntypes of Discognathus obtusus, 46-134 mm standard length, are under NMW 53238 and 2 syntypes, 65-92 mm standard length, are under SMF 5408 (formerly NMW and also numbered SMF 447)(65.1-93.1 mm standard length). A further 10 fish under NMW 53257 and measuring 31.5-106.2 mm standard length are also indicated on the jar in Vienna as syntypes but this is probably in error as the catalogue lists 6 fish. Eschmeyer et al. (1996) give only 1 syntype under NMW 53257, 4 syntypes under NMW 53238, 1 dried syntype under NMW 79372, 1 syntype in the Senckenberg Museum Frankfurt under SMF 447 and 2 syntypes under SMF 5408 (formerly NMW).

The syntypes of Discognathus crenulatus are in the Naturhistorisches Museum Wien under NMW 53236 (14 specimens) from the Qarah Aqaj River and 53237 (6) from Sa`di measuring 33-79 mm standard length (Kähsbauer, 1964). The 14 specimens under NMW 53236 measure 24.0-75.9 mm standard length and 7 (not 6) specimens under NMW 53237 measure 35.4-56.6 mm standard length according to my observations. Neither the record of Kähsbauer (1964) nor my own data from jars on the shelves accord with the catalogue in Vienna which gives 10 or 8 and 6 or 5 specimens respectively for these two syntype localities.

The syntypes of Garra rufa gymnothorax are in the Zoological Institute, St. Petersburg (ZISP 13214), there being 6 fish in the catalogue and 6 in the jar although Berg (1949) lists 7 in his description. They measure 30.5-44.9 mm standard length. The date in Berg (1949) is 6.VI.1904 while in the catalogue it is 4.III.1904 and in the jar 24.III.1904, variations not accountable by old and new styles of dating (13 days apart). A further collection listed by Berg (1949), (ZISP 24435), is not listed as type material in the text nor in the jar but the catalogue suggest that they are (Eschmeyer et al. (1996) list these 10 fish as syntypes). ZISP 24354 is from "Ziaret-Seid-Hasan, Mesopotamiya". The type series may be only ZISP 13214 as in the text. Eschmeyer et al. (1996) list ZIL (= ZISP) 13215, 17 fish, and ZIL 24436, 3 fish, as syntypes also.

Bianco and Banarescu (1982) referred their material from the Hablehrud and Mand River to Garra rufa crenulata as these fish had fewer scales (29-32 for Mand and 33-34 for Hablehrud versus 35-38 from the Tigris-Euphrates)  and fewer gill rakers (15-21 versus 25-27 in Tigris-Euphrates specimens). Possible syntypes of crenulata had intermediate scale counts (31-34) between Mand and Hablehrud fish. These authors suggest that there may be distinct subspecies in these two rivers. Their sample sizes are too small in my opinion to warrant subspecies recognition. Berg (1949) was uncertain of the status of this taxon.

A Principal Components Analysis on 448 fish from the Hormuz, Lake Maharlu, Gulf and Tigris River basins and Sa`di's Tomb using 20 morphometric and 5 meristic characters did not separate any of these groups. Note that fish form the Hormuz basin, rivers draining to the Straits of Hormuz, modally had 7 branched dorsal fin rays (as in G. persica, see table below) but branched caudal fin rays were modally 17 (not 16 as in G. persica). There may be some introgression in this region of Iran. Ghalenoei et al. (2010) examined fish from 13 stations in the Tigris and Persian Gulf basins and found the Mand River population was separated from the rest, which overlapped each other.

Menon (1964) and Karaman (1971) consider Garra persica to be a synonym but this species is regarded here as distinct. Both these authors refer specimens from the Tigris River basin of Iran (and therefore all Iranian specimens) to a subspecies, Garra rufa obtusa, distinguished from the type subspecies in Syria, Lebanon, Israel and Jordan by having a variable number of dorsal fin branched rays (7-8), the anal aperture further forward, and the anal fin origin half way between the pelvic fin base and the caudal fin base as opposed to closer to the pelvic base. Krupp (1985c) synonymises Garra rufa obtusa with the type subspecies.

Fish lacking scales on the breast were named by Berg (1949) as Garra rufa gymnothorax but this is a variable character widespread among cyprinid fishes in Iran and is unlikely to be of systematic significance.

Key characters

Two pairs of barbels are present, the adhesive disc is well developed with a free anterior margin, the dorsal fin has 8 branched rays modally, and the caudal fin 17 branched rays modally.

Morphology

Dorsal fin with 2-3, usually 3, unbranched and 7-9 branched rays with a very strong mode at 8, anal fin with 2-3 unbranched and 4-6 with a very strong mode at 5 branched rays (99.6% of 534 specimens from Iran). Pectoral fin branched rays 12-14 (in literature, but see table), pelvic fin branched rays 7-8 (but see table). Lateral line scales 31-38 (but see table), scales from the dorsal fin origin to the lateral line 3-6, from the lateral line to the pelvic fin origin 2-5, predorsal mid-line scales 9-13, and scales around the caudal peduncle 12-17 with a strong mode at 16. Pharyngeal tooth formula 2,4,5-5,4,2 or 2,4,4-4,4,2 (3,3,5-5,3,3 in Heckel (1843b)). Teeth are hooked at the tip. The short gill rakers number 16-21, 12-17 on the lower arm. In Iranian specimens the range is 14-26 (a range only is given since rakers are difficult to count on the arch ends with accuracy and may be related to age). The upper lip is delicately fimbriated. Pharyngeal teeth in Iranian specimens? Total vertebrae in Iranian specimens 32-37 (see table). The chromosome number is probably 2n=52 (Klinkhardt et al., 1995) although Ünlü et al. (1997) give 2n=38 for Turkish specimens with 26 meta- to submetacentric chromosomes and 12 telo- to subtelocentric chromsomes (NF=64) and Gözükara and Çavaş (2004) gave 2n = 44 for Turkish specimens with 22 metacentric and 20 submetacentric chromosomes and 2 acrocentric ones (NF=85). Esmaeili and Piravar (2007) examined fish from the Rodbal River in Fars and found 2n=50 with arm number NF=84 and the karyotype formula of 10 metacentric, 24 sub-metacentric and 16 sub-telocentric chromosomes.

Some fish are very rounded in cross section while others are more terete, possibly related to habitat as observed in other cyprinid fishes..

Meristics for Iranian specimens:-

 

Basin/Dorsal fin branched rays

7

8

9

x

S.D.

Hormozgan

27

6

7.2

0.39

Lake Maharlu

90

2

8.0

0.15

Sa`di's Tomb

5

20

7.8

0.41

Gulf

16

173

1

7.9

0.29

Tigris River

15

176

3

7.9

0.30

 
Basin/Pectoral fin branched rays 10 11 12 13 14 15 16 18 x S.D.
Hormozgan 1 3 18 10 1 14.2 0.78
Lake Maharlu 1 2 15 50 23 1 14.1 0.78
Sa`di's Tomb 5 16 2 2 1 13.2 1.24
Gulf 1 4 57 84 42 1 13.9 0.81
Tigris River 1 1 10 54 93 33 2 13.8 0.88

 

Basin/Pelvic fin branched rays 6 7 8 9 x S.D.
Hormozgan 20 12 7.4 0.49
Lake Maharlu 8 84 7.9 0.28
Sa`di's Tomb 9 16 7.9 0.27
Gulf 1 110 79 7.4 0.50
Tigris River 2 51 138 3 7.7 0.50

 
Basin/Lateral line scales 25 26 27 28 29 30 31 32 33 34 35 36 37 38 40 x SD
Hormuz 2 7 19 10 1 33.0 0.87
Lake Maharlu 1 4 13 21 32 30 9 31.2 1.54
Sa`di's Tomb 2 6 6 7 4 33.1 1.39
Gulf 1 1 5 16 25 24 25 42 36 19 5 31.2 2.06
Tigris River 2 2 24 44 59 45 9 2 1 1 33.9 1.34

 

Basin/Total vertebrae 32 33 34 35 36 37 x S.D.
Hormozgan 7 24 7 3   34.2 0.78
Lake Maharlu   6 38 21 3   34.3 0.70
Sa`di's Tomb     14 9     34.4 0.50
Gulf 2 15 36 34 9   34.3 0.93
Tigris River   23 52 76 17 2 34.5 0.92

Sexual dimorphism

Large males become heavily tuberculate on the front and sides of the snout and in a band from the eye to the nostril and across to the other nostril and eye as illustrated in Fowler and Steinitz (1956) for a specimen from `Ain Umm Keishik in Iraq. A deep, tubercle-free groove is apparent between the upper band of tubercles through the nostrils and the tubercles on the snout above the mouth.

Colour

Overall colour is brownish-olive to dark green with darkly mottled flanks and a yellowish to whitish belly. The head and flanks may be a rusty-red, bronze or golden. A dark or bluish-green band runs along the whole flank ending in a spot on the caudal fin base. Much of the body may be blackish with only the belly creamy. Others are a light olive-green with lime-green highlights giving an iridescent effect especially on upper anterior flank scales. There is a black, greenish-blue, lime-green or dusky-blue spot behind the upper corner of the gill opening, sometimes extending as a bar to the pectoral fin base where the skin is also blue. Fins can be yellowish with darker margins. The pectoral fins can be orange-pink dorsally, grey-white or slightly orange-pink ventrally. The pelvic and anal fins may be orange with the fin rays yellow posteriorly in the anal fin but yellow mesially in the pelvic fin. The bases of the pectoral, pelvic and anal fins are orange-red in breeding males and the caudal fin is orange. The caudal fin can be orange to red ventrally and yellow dorsally. There is a black spot at the caudal fin base and the upper caudal lobe may have a few dark grey spots. The dorsal fin is dark green with reddish pigment at its middle. There is usually a dark spot at the bases of each of the middle 4-5 dorsal fin rays. In some specimens the dorsal fin is orange with yellow posterior rays. The pectoral, dorsal and caudal fin rays may be olive to black rather than yellow or orange. The iris is bright yellow, orange or red.

There is variation in colouration. Some fish are pale while others are very dark; the spots on the dorsal fin may extend two-thirds of the way up the fin rather than being restricted to the base; and the flanks may not be mottled. Fish from muddy water are a sickly grey with the body mottled and the lower caudal lobe dark. Their colour darkens and becomes brighter after immersion in ice water. Fish from deep in qanats are very pale.

Size

Attains 24 cm total length in the Tigris River in Iraq (Rahemo, 1995). Reaches 15.9 cm (Krupp, 1985c), over 17 cm according to Heckel (1843b). Fish up to 18.5 cm total length are known from Khuzestan.

Distribution

Found in the Tigris-Euphrates basin and the Ceyhan, Orontes (= Asi), Quwayq and Jordan river basins and coastal drainages of the eastern Mediterranean as well as much of southern Iran.

In Iran it is found in the Tigris River, Gulf, Lake Maharlu, and Kor River basins and the Hormuz basin (Berg, 1949; Menon, 1964; Bianco and Banarescu, 1982; Gh. Izadpanahi, pers. comm., 1995; M. Rabbaniha, pers. comm., 1995; Abdoli, 2000).

Zoogeography

The wide distribution in Southwest Asia and inadequate examination of variation may mask distinct taxa, although this is not apparent on morphometric and meristic grounds. If such variation is valid, then taxa may reflect vicariant events.

Habitat

This is the commonest species in catches in southwestern Iran, followed by Cyprinion macrostomum. In areas under human influence in Lorestan, such as the lower reaches of rivers and near cities, it is more common than in higher, pristine waters. As well as grazing on exposed rock surfaces in streams, it can be found under pebbles on the stream bed.

Age and growth

Rahemo (1995) reported fish up to age 7 from the Tigris River, Iraq in a parasitological study. Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 291 Iranian fish measuring 2.28-11.82 cm standard length. The a-value was 0.0265 and the b-value 2.919 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases)

Abedi et al. (2010) examined the age in a population in the Armand Stream in Chahar Mahall va Bakhtiari Province. Length and weight were significantly correlated (W = -5.076 + 3.112 log TL for sexes combined, -5.092 + 3.134 log TL for males and -5.036 + 3.089 log TL for females), growth was isometric, numbers of males and females were not significantly different, life span was up to 4 years and most fish were 2-3 years old. Patimar et al. (2010) examined fish from the Kangir River in Ilam and found a maximum age of 5+ years, the most frequent size class was 65-70 mm for both sexes, negative allometric growth in both sexes, a balanced overall sex ratio but with males predominant at smaller sizes and females at larger sizes, and von Bertalanffy growth parameters were L = 108.453 mm, K = 0.449 year-1, t0 = 0.192 years for males and L = 115.516 mm, K = 0.420 year-1, t0 = 0.088 years for females. Males grew faster than females. The growth index ф' was 8.572 for males and 8.631 for females. Fish were smaller and of lower weight than fish from the Zanjiran spring stream system in Fars, southern Iran studied by Yazdanpanah (2005), and this was attributed to severe ecological conditions. The Zanjiran fish has a similar most frequent size class of 60-70 mm, but growth was positively allometric and the sex ratio was more balanced.

Food

Gut contents include diatoms, algae and large quantities of sand in fish examined from Iran. Younis et al. (2001b) found Shatt al Arab fish feeding mainly on organic detritus, followed by diatoms and algae, with arthropods ranking third. A study by Yalçin-Özdilek and Ekmekçi (2006) in the Asi (= Orontes) River in Turkey demonstrated that this species is a grazer on aquatic plants, mostly consisting of benthic cyanobacteria, chrysophytes and phytoplankton with included rotifers and protozoans. Both season and location in a stream affected the composition of the diet with season the most important factor.

Reproduction

Al-Rudainy (2008) gives sexual maturity at 2-3 years, 10 cm in length and a weight of 50 g for Iraqi fish. Spawning took place in May and June with eggs deposited on vegetation and rocks with a relative fecundity up to 542 eggs/g. Bardakci, Ozansoy and Koptagel (at www.epress.com/w3jbio/vol5/bardakci/paper.htm, downloaded 29 March 2001) note depression of vitellogenesis in a hot spring population in Turkey, perhaps due to temperature and starvation. A nearby stream population has a higher gonadosomatic index. Ovaries increased in size and weight from May to July in both localities although the hot spring had fewer mature oocytes and more atretic oocytes at various development stages. High temperatures and poor food conditions in some Iranian habitats may be limiting factors in reproduction for this species.

The Armand Stream population (Abedi et al., 2010) had a prolonged and active reproductive season from March to September, an adaptation to unstable environmental conditions. All mature oocytes are spawned at once but some may be retained for later spawning. In addition, different individuals release eggs and sperm at different times. Average egg diameter was 0..67 mm, maximum 1.98 mm, with highest diameter in May and the lowest in November. Absolute and relative fecundity were 1179.65 and 109.4 respectively on average. Maximum absolute fecundity reached 3794 eggs. The Kangir River fish (Patimar et al., 2010) had a maximum fecundity of 13,927 eggs and a maximum relative fecundity of 2345.72 eggs/g. Egg diameters reached 1.7 mm. Reproduction occurred in April-May with the highest average gonadosomatic index for males of 4.21 in April and for females of 7.85 in May. The Zanjiran population (Yazdanpanah, 2005) had the highest gonadosomatic index for males in March and for females in April with maximum averages at 6.49 for males and 16.98 for females. The range and mean of absolute fecundity for this population were much lower than that in the Kangir River (184-2396, mean 760 versus 1680-13,927, mean 5806), as was relative fecundity.

Parasites and predators

Jalali and Molnár (1990a) record two monogenean species, Dactylogyrus spp., from this species in the Dez River. Gussev et al. (1993b) describe two new species, Dactylogyrus rectotrabus and D. acinacus, from this species in the Dez River, Khuzestan. Jalali et al. (2005) summarise the occurrence of Gyrodactylus species in Iran and record G. sp. from fish in the Helleh River.

Eaten by Silurus triostegus at Baghdad (notes on a specimen in the Field Museum of Natural History, Chicago (FMNH 51251)).

Economic importance

Ündar et al. (1990) identify this species and Cyprinion macrostomum as the "doctor fish" of the Kangal hot spring in Turkey (Warwick and Warwick, 1989; Kürkçüoğlu and Öz, 1989; Bardakci, Ozansoy and Koptagel at www.epress.com/w3jbio/vol5/bardakci/paper.htm, downloaded 29 March 2001, Bilke, 2004; Anonymous, 2007; and various newspaper and television reports). High water temperatures around 35°C reduce the amount of plankton available as fish food and the fish nibble away infected skin of humans who bathe in these waters. This fish is known as "licker" (and Cyprinion macostomum as "striker") from its behaviour in the spa pools. The healing properties are linked to the high level of selenium (1.3 p.p.m.) in the water, selenium being beneficial in some skin diseases and possibly UV light. The fish facilitate the action of the selenium and UV light by softening and clearing away psoriatic plaque and scale, exposing the lesions to the water and sunlight. However some lesions are made worse and the fish can cause some new ones. Garra rufa is now widely used in commercial facilities around the world for treating skin diseases and removing dead skin. Jarvis (2011) gives a biological synopsis of this species, published in Canada, since it is increasingly being used in private spa facilities and may escape into the wild.

Garra rufa as "doctor fish" from Wikimedia Commons.

Conservation

A common species with a wide distribution and not under any specific threat. Vulnerable in Turkey (Fricke et al., 2007).

Further work

Meristic counts summarised above show overlapping but significant variation between drainage basins and may reflect recognisable taxa. Molecular studies would help clarify this situation but is should be noting that variation of this nature is to be expected in a wide ranging species and does not always warrant taxonomic distinction.

Sources

Type material: See above, Discognathus rufus (NMW 53240 and SMF 553), Discognathus obtusus (NMW 53238 and SMF 5408), Discognathus crenulatus (NMW 53236 and 53237), and Garra rufa gymnothorax (ZISP 13214), and see comments on other possible types.

Iranian material: CMNFI 1970-0540, 7, 22.2-41.4 mm standard length, Fars, qanat south of Kazerun (no other locality data); CMNFI 1979-0018, 48, 21.5-64.9 mm standard length, Fars, Barm-e Shur (29º28'N, 52º41'30"E); CMNFI 1979-0019, 4, 28.9-35.4 mm standard length, Fars, Barm-e Baba Haji (29º23'N, 52º40'E); CMNFI 1979-0026, 2, 21.5-22.3 mm standard length, Fars, Shapur River at Shapur (29º47'N, 51º35'E); CMNFI 1979-0027, 1, 50.5 mm standard length, Fars, Chehel Cheshmeh (ca. 29º43'N, ca. 52º04'E); CMNFI 1979-0033, 34, 23.7-72.0 mm standard length, Fars, Barm-e Shur (29º28'N, 52º41'30"E); CMNFI 1979-0036, 1, 36.4 mm standard length, Fars, Shapur River at Shapur (29º47'N, 51º35'E); CMNFI 1979-0045, 16, 20.0-59.2 mm standard length, Fars, spring at Sa'adi's Tomb, Shiraz (29º37'N, 52º35'E); CMNFI 1979-0046, 1, 43.7 mm standard length, Fars, qanat at Barm-e Dalak (ca. 29º35'N, ca. 52º38'E); CMNFI 1979-0047, 2, 36.2-67.6 mm standard length, Fars, spring source of Ab-e Paravan marshes (ca. 29º34'N, ca. 52º42'E); CMNFI 1979-0048, 1, 40.1 mm standard length, Fars, spring and marsh northeast side of Lake Maharlu (ca. 29º32'N, ca. 52º48'E); CMNFI 1979-0075, 31, 12.5-57.1 mm standard length, Fars, Mand River at Pol-e Kavar (29º11'N, 52º41'E); CMNFI 1979-0109, 1, 74.3 mm standard length, Fars, Mand River at Shahr-e Khafr (28º56'N, 53º14'E); CMNFI 1979-0111, 8, 30.7-62.0 mm standard length, Fars, stream on Shiraz-Bushehr road (29º37'30"N, 52º21'E); CMNFI 1979-0112, 5, 55.0-77.1 mm standard length, Fars, stream draining Soltanabad Marshes (29º29'N, 52º38'30"E); CMNFI 1979-0113, 4, 40.8-63.7 mm standard length, Fars, spring at Sa'adi's Tomb, Shiraz (29º37'N, 52º35'E); CMNFI 1979-0115, 5, 57.7-66.5 mm standard length, Fars, spring at Sa'adi's Tomb, Shiraz (29º37'N, 52º35'E); CMNFI 1979-0120, 5, 27.6-66.3 mm standard length, Bushehr, Dalaki River near Konar Takhteh (29º28'N, 51º21'E); CMNFI 1979-0125, 2, 99.0-121.4 mm standard length, Bushehr, Dalaki River near Dalaki (ca. 29º28'N, ca. 51º21'E); CMNFI 1979-0128, 6, 26.6-32.1 mm standard length, Shur River between Atashkadeh and Firuzabad (28º51'N, 52º31'E); CMNFI 1979-0129, 43, 24.8-46.9 mm standard length, Fars, spring 2 km north of Farrashband (28º54'N, 52º04'E); CMNFI 1979-0131, 10, 16.9-46.6 mm standard length, Fars, Ab-Arak River 24 km from Qir (28º38'N, 52º49'E); CMNFI 1979-0132, 11, 23.1-49.5 mm standard length, Fars, Shur River 54 km from Firuzabad (28º35'N, 52º58'E); CMNFI 1979-0155, 11, 27.1-42.9 mm standard length, Fars, spring at Gavanoo village (28º47'N, 54º22'E); CMNFI 1979-0156, 20, 33.8-56.7 mm standard length, Fars, qanat at Rashidabad (28º47'N, 54º18'E); CMNFI 1979-0157, 10, 40.6-88.6 mm standard length, Fars, qanat at Hadiabad (28º52'N, 54º13'E); CMNFI 1979-0158, 13, 35.3-54.2 mm standard length, qanat over Qasook River (28º54'N, 53º53'30"E); CMNFI 1979-0161, 11, 43.1-92.2 mm standard length, Fars, qanat on Shiraz-Neyriz road (29º10'30"N, 53º41'E); CMNFI 1979-0193, 3, 22.8-27.1 mm standard length, Fars, river 8 km from Darab (28º45'N, 54º27'30"E); CMNFI 1979-0195, 3, 44.3-59.4 mm standard length, Fars, jube stream of road to Fasa (ca. 28º54'N, ca. 53º53'30"E); CMNFI 1979-0199, 4, 37.3-45.0 mm standard length, Fars, qanat 18 km from Jahrom (ca. 28º23-25'N, ca. 53º31-40'E); CMNFI 1979-0200, 5, 27.6-43.6 mm standard length, Fars, Mand River tributary 13 km from Jahrom (28º36'N, 53º36'30"E); CMNFI 1979-0202, 9, 19.1-25.1 mm standard length, Fars, Mand River (29º01'N, 53º00'E); CMNFI 1979-0206, 2, 50.9-51.9 mm standard length, Fars, qanat near Runiz-e Pa'in (29º12'N, 53º40'E); CMNFI 1979-0241, 6, 35.7-49.2 mm standard length, Fars, Shapur River at Shapur (29º47'N, 51º35'E); CMNFI 1979-0271, 8, 34.7-56.7 mm standard length, Lorestan, Kashkan River drainage (33º39'N, 48º32'30"E); CMNFI 1979-0273, 15, 40.9-58.2 mm standard length, Lorestan, Kashkan River drainage 5 km from Khorramabad (33º26'N, 48º19'E); CMNFI 1979-0274, 3, 24.4-32.8 mm standard length, Lorestan, Kashkan River drainage 20 km from Khorramabad (33º27'N, 48º11'E); CMNFI 1979-0275, 7, 38.6-60.5 mm standard length, Lorestan, Kashkan River near Ma'mulan (33º25'N, 47º58'E); CMNFI 1979-0276, 10, 42.1-70.0 mm standard length, Lorestan, Chamesk River (ca. 33º19'N, ca. 47º53'30"E); CMNFI 1979-0277, 1, 128.4 mm standard length, Lorestan, Kashkan River drainage (33º30'N, 47º59'30"E); CMNFI 1979-0278, 12, 53.9-89.3 mm standard length, Lorestan, Kashkan River draiange, Sarab Dowrah (33º34'N, 48º01'E); CMNFI 1979-0279, 9, 33.5-117.3 mm standard length, Lorestan, Khorramabad River (33º37'N, 48º18'E); CMNFI 1979-0288, 43, 22.9-102.3 mm standard length, Ilam and Poshtkuh, Gangir River at Sarab Ewan (33º50'N, 46º18'E); CMNFI 1979-0289, 5, 57.2-101.9 mm standard length, Kermanshahan, Diyala River drainage (34º28'N, 45º52'E); CMNFI 1979-0290, 6, 24.1-54.9 mm standard length, Kermanshahan, Diyala River drainage in Qasr-e Shirin (34º31'N, 45º35'E); CMNFI 1979-0291, 7, 24.9-66.8 mm standard length, Kermanshahan, Diyala River draiange 15 km from Qasr-e Shirin (34º24'N, 45º37'E); CMNFI 1979-0293, 8, 92.1-101.3 mm standard length, Fars, Mand River at Kavar (29º11'N, 52º41'E); CMNFI 1979-0304, 2, 41.6-43.0 mm standard length, Fars, Lake Famur (ca. 29º31'N, ca. 51º50'E); CMNFI 1979-0350, 4, 28.1-33.5 mm standard length, Khuzestan, Marun River near Marun (30º39'30"N, 50º02'E); CMNFI 1979-0364, 4, 36.0-50.6 mm standard length, Khuzestan, Karkheh River branch at Abdolkhan (31º52'30"N, 48º20'30"E); CMNFI 1979-0365, 2, 32.1-32.8 mm standard length, Khuzestan, stream in Doveyrich River drainage (32º25'N, 47º36'30"E); CMNFI 1979-0366, 2, 22.7-29.2 mm standard length, Khuzestan, stream 17 km west of Dehloran (32º45'30"N, 47º05'30"E); CMNFI 1979-0367, 1, 38.0 mm standard length, Khuzestan, Meymeh River 11 km north of Dehloran (32º44'30"N, 47º09'30"E); CMNFI 1979-0368, 4, 25.3-47.7 mm standard length, Khuzestan, Karkheh River (32º24'30"N, 48º09'E); CMNFI 1979-0369, 4, 24.9-37.7 mm standard length, Khuzestan, Shush River at Shush (32º12'N, 48º14'30"E); CMNFI 1979-0371, 1, 37.3 mm standard length, Khuzestan, stream in Karkheh River drainage (32º05'N, 48º19'E); CMNFI 1979-0374, 1, 62.2 mm standard length, Khuzestan, stream tributary to Bala River (32º40'N, 48º15'E); CMNFI 1979-0375, 10, 37.9-71.1 mm standard length, Khuzestan, stream tributary to Bala River (ca. 32º45'N, ca. 48º14'30"E); CMNFI 1979-0378, 1, 49.3 mm standard length, Khuzestan, stream tributary to Karkheh River (ca. 32º48'N, ca. 48º04'E); CMNFI 1979-0379, 1, 63.1 mm standard length, Khuzestan, Dez River (32º12'N, 48º27'E); CMNFI 1979-0382, 3, 34.7-38.0 mm standard length, Khuzestan, Karun River at Shushtar (32º03'N, 48º51'E); CMNFI 1979-0383, 5, 31.9-53.0 mm standard length, Khuzestan, stream in Ab-e Shur drainage (31º59'30"N, 49º06'E); CMNFI 1979-0384, 1, 63.3 mm standard length, Khuzestan, river in Ab-e Shur drainage (32º00'N, 49º07'E); CMNFI 1979-0385, 2, 44.9-46.8 mm standard length, Khuzestan, stream in Ab-e Shur drainage (ca. 32º01'N, ca. 49º07'30"E); CMNFI 1979-0387, 2, 48.2-64.5 mm standard length, Khuzestan, stream in Jarrahi River drainage (31º25'N, 49º38'E); CMNFI 1979-0388, 1, 55.2 mm standard length, Khuzestan, Zard River 21 km north of Ramhormoz (31º19'N, 49º44'E); CMNFI 1979-0389, 2, 43.8-57.7 mm standard length, Khuzestan, Zard River at Bagh-e Malek (31º31'N, 49º53'30"E); CMNFI 1979-0390B, 10, 33.8-69.7 mm standard length, Khuzestan, stream tributary to Zard River (31º29'N, 49º54'30"E); CMNFI 1979-0391, 1, 39.7 mm standard length, Khuzestan, stream in Marun River drainage (31º28'N, 49º51'E); CMNFI 1979-0392, 1, 42.6 mm standard length, Khuzestan, Zard River 25 km north of Ramhormoz (ca. 31º32'N, ca. 49º48'E); CMNFI 1979-0396, 3, 22.7-35.1 mm standard length, Kheyrabad River 20 km from Behbehan (30º32'N, 50º23'30"E); CMNFI 1979-0397, 2, 52.4-59.5 mm standard length, Khuzestan, stream tributary to Kheyrabad River (30º30'N, 50º28'E); CMNFI 1979-0398, 1, 37.6 mm standard length, Boyer Ahmadi-ye Sardsir va Kohkiluyeh, stream in Zohreh River drainage (30º24'30"N, 50º37'30"E); CMNFI 1979-0399, 1, 41.0 mm standard length, Fars, stream near Basht in Zohreh River drainage (30º19'30"N, 51º15'E); CMNFI 1979-0497, 2, 52.4-60.3 mm standard length, Fars, Mand River at Band-e Bahman (29º11'N, 52º40'E); CMNFI 1979-0501, 8, 24.7-48.9 mm standard length, Fars, Mand River at Kavar (29º11'N, 52º41'E); CMNFI 1987-0217, 5, 35.8-55.6 mm standard length, Khuzestan, Karun River at Kut Abdollah (31º13'N, 48º39'E); CMNFI 2007-0063, 5, 40.1-65.9 mm standard length, Fars, Mand River tributary near Jahrom (28º36'N, 53º37'E); CMNFI 2007-0065, 1, 81.2 mm standard length, Fars, Barm-e Dalak (ca. 29º35'N, ca. 52º38'E); CMNFI 2007-0066, 2, 47.1-55.3 mm standard length, Fars, qanat under Sa'di's Tomb, Shiraz (29º37'N, 52º35'E); CMNFI 2007-0100, 2, 48.4-54.8 mm standard length, Azarbaijan-e Gharbi, Kalwi Chay near Piranshahr (ca. 36º44'N, ca. 45º10'E); CMNFI 2007-0109, 10, 54.7-78.3 mm standard length, Kordestan, Qeshlaq River basin (ca. 35º16'N, ca. 47º01'E); CMNFI 2007-0110, 1, 84.9 mm standard length, Kordestan, Yuzidar River basin (ca. 35º05'N, ca. 46º56'E); CMNFI 2007-0111, 11, 26.5-64.5 mm standard length, Kermanshahan, Alvand River near Sar-e Pol-e Zahab (ca. 34º36'N, ca. 45º56'E); CMNFI 2007-0112, 19, 43.1-54.3 mm standard length, Kermanshahan, Kerend River basin near Shahabad-e Gharb (ca. 34º06'N, ca. 46º30'E); CMNFI 2007-0116, 4, 25.8-31.7 mm standard length, Kermanshahan, Gav Masiab River basin west of Sahneh (ca. 34º28'N, ca. 47º36'E).

Comparative material: BM(NH) 1931.12.21:9-10, 2, 94.9-95.5 mm standard length, Iraq, Mosul (36º20'N, 43º08'E); BM(NH) 1973.5.21:187-188, 2, 67.6-73.1 mm standard length, Iraq, Tigris River at Jadriyah (no other locality data); BM(NH) 1974.2.22:1418, 1, 67.4 mm standard length, Iraq, Khalis (33º49'N, 44º32'E); BM(NH) 1974.2.22:1441-1444, 4, 48.9-86.3 mm standard length, Iraq, Baghdad (33º21'N, 44º25'E); BM(NH) 1986.2.14:2-3, 2, 64.8-91.8 mm standard length, Iraq, Baghdad (33º21'N, 44º25'E); NMW 91123, 5, ?, Turkey, Cheilani bei Cizre, Tigris basin (ca. 37º20'N, ca. 42º10'E).

Garra variabilis
(Heckel, 1843)

Common names

See under genus account.

[karkoor mit-la'oon, gassur diseileki or gassur isivid (gassur = colour of strawberries, isivid referring to the spotted, almost black fish according to Heckel (1843b), all in Arabic; variable garra].

Systematics

Menon (1964) considers this species to be the most primitive in the genus. It has been placed in the genus Discognathichthys Bleeker, 1860, e.g. by Berg (1949).

Heckel (1843b) gave the type localities as "Mossul" and "Aleppo". The syntypes of Discognathus variabilis are in the Naturhistorisches Museum Wien under NMW 53239, 8 specimens, 38-112 mm standard length and in the Senckenberg Museum Frankfurt under SMF 403 (formerly NMW), 4, 72-86 mm standard length, all from Aleppo (Krupp, 1985c). In Vienna I made counts on types as listed below under Sources. Material under NMW 53238 (3) from Aleppo may also be types. Eschmeyer et al. (1996) list NMW 532339 (= an error for 53239) (8), NMW 53260-69 (1, 2, 2, 2, 3, 2, 2, 2, 2, 2), NMW 53272 (4), SMF 403 (4) (formerly NMW) and in the Museum für Naturkunde, Universität Humboldt, Berlin, ZMB 3301 (3) (formerly NMW; 82.6-99.2 mm standard length measured in February 2006).

Key characters

The single pair of maxillary barbels, absence of a free anterior margin to a weakly developed adhesive disc on the lower head surface without papillae on the rear part, scaled back, chest and belly, gill raker count and distribution distinguishes this species. It is separated from the closely related but geographically separated G. rossica by larger size, head length shorter than caudal peduncle length and pectoral fin length, slightly emarginate caudal fin, and dorsal fin origin closer to snout tip than the caudal fin base (Berg, 1949).

Morphology

Dorsal fin with 2-3, usually 3, unbranched and 6-8, modally 7, branched rays, anal fin with 2-3 unbranched and 5 branched rays. Pectoral fin branched rays 11-14, pelvic fin branched rays 7-8. Lateral line scales 32-40, scale from the dorsal fin origin to the lateral line 4-6, scales below the lateral line to the pelvic fin origin 3-4, scales around the caudal peduncle usually 16, and predorsal scales in mid-line 12-14. The chest and belly are scaled. The upper lip is not fimbriate. There may be 2 pairs of barbels in some larger fish. Pharyngeal tooth formula 2,4,5-5,4,2 (2,3,5-5,3,2 in Berg (1949), 3,3,5-5,3,3 in Heckel (1843b)) and the short gill rakers number 13-20, on the lower arm of the gill arch only. Gut very elongate and coiled. The diploid chromosome number was 2n=102 with karyotype formulae being 42m + 18sm + 24st + 18a (FN=186) for females and 41m + 18sm + 24t + 19a (FN=185) for males in fish from Savur Stream, Turkey (Karahan and Ergene, 2010).

Sexual dimorphism

Specimens from NMW 91121 had the top and sides of the head finely tuberculate and scales on the back before the dorsal fin with fine tubercles lining the scale margins. The upper lip, lip sides and sucker (except for a naked central area) have keratinised tubercles. Tubercles line the dorsal surface of pectoral fin rays, fading medially and following the ray branching in single rows.

Colour

Overall colour is olivaceous brown or greyish with darker mottlings. The flanks may have large, irregularly-arranged dark spots. The upper corner of the operculum may have a black spot. The belly is reddish-yellow. The middle 3-4 rays of the dorsal fin each have a small, black spot at their bases. There is a black spot at the caudal fin base. The lateral line may occasionally have a double row of black spots as in certain Alburnoides spp. Young fish may have a dark lateral stripe. Peritoneum black.

Size

Reaches 15.5 cm or according to Heckel (1843b) 5 Zoll (= about 21 cm).

Distribution

Found in the Quwayq (= Kueik), Orontes (= Asi) and Nahr al-Kabir rivers of the Levant and the Tigris-Euphrates basin (Menon, 1964; Krupp, 1985c). In Iran, this species is found in the Tigris River basin. Abdoli (2000) maps the Jarrahi, middle Karun, lower Dez, Karkheh, lower Simarreh, and lower Kashkan rivers.

Keyserling (1861) recorded this species from Sistan but this is a misidentification. Records of this species (as Discognathus variabilis) from Sistan by Nikolskii (1899) and Regan (1906) are G. rossica (Menon, 1964).

Zoogeography

This species is related to Garra rossica of eastern Iran. See also under Garra persica. Krupp (1985c) considers this species to belong to the indo-asiatic line of Garra.

Habitat

Garra rufa and this species seem to exclude each other, variabilis being more common in faster water (F. Krupp).

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

None reported from Iran.

Economic importance

None.

Conservation

This species appears to be very rare in Iran and if specific localities are found they should be protected.

Further work

The presence of this species in Iranian waters and its biology there need study.

Sources

Type material: See above, Discognathus variabilis (NMW 53239, SMF 403 and ZMB 3301) and see comments also on other possible types.

Iranian material:- None.

Type material used for counts: NMW 53260, 1, 40.6 mm standard length, NMW 53261, 2, 87.6-97.8 mm standard length, NMW 53262, 2, 51.4-52.1 mm standard length, NMW 53263, 2, 101.8-103.1 mm standard length, NMW 53264, 3, 52.3-54.2 mm standard length, NMW 53266, 2, 87.6-92.7 mm standard length, NMW 53267, 2, 61.5-68.3 mm standard length, NMW 53268, 7, 43.9- 71.4 mm standard length (dried at some point), NMW 53269, 2, 100.3-109.1 mm standard length, NMW 53272, 4, 84.9-92.1 mm standard length, all previous Tigris at Mosul; NMW 53239, 8 36.9-111.0 mm standard length, Aleppo.

Comparative material: BM(NH) 1935.9.12:27-40, 5, 60.8-69.1 mm standard length, Iraq, Karasu (no other locality data); BM(NH) 1935.9.12:53, 1, 76.0 mm standard length, Iraq, Tchaiy Su (no other locality data); BM(NH) 1986.2.14:2-5, 2, Iraq, Baghdad (33º21'N, 44º25'E); BM(NH) 1968.12.13:290-297, 8, 30.7-71.6 mm standard length, Syria, Tigris River at `Ayn Diwar (37º17'N, 42º11'E); BM(NH) 1968.12.13:298-304, 7, 44.4-66.7 mm standard length, Syria, Quweiq River at Masslemiyeh (no other locality data); NMW 91121, 10, 71.9-115.4 mm standard length, Turkey, Wadi Mahmedian Tschai (ca. 38°20'N, ca. 40°45'E).

Genus Gobio
Cuvier, 1816

The gudgeon genus includes about 24 species found from the British Isles through Europe and northern Asia to Korea. There is one species in Iran and two other species are now in the genus Romanogobio.

The body shape is distinctive, being elongate and fusiform with moderately large scales (36-51), the throat is naked or scaled, the mouth is inferior or terminal, horseshoe-shaped and has a barbel at each corner, the lower lip is thin like the upper lip but is interrupted medially, gill rakers are short and widely spaced, pharyngeal teeth are in 2 rows (usually 5 in the main row and 2-3 in the second) and are obviously hooked at the tips, both dorsal and anal fins are short and spineless, the gut is short and the peritoneum silvery, and the vent is remote from the anal and pelvic fin origins.

Gobio gobio
(Linnaeus, 1758)

Common names

mahi kopur kafzi (= bottom-dwelling carp fish), گاو ماهي (= gav mahi, probably in error for Neogobius and related gobies).

[Turkestanskii peskar' or Turkestan gudgeon in Russian].

Systematics

Cyprinus Gobio was originally described from England.

Bungia nigrescens Keyserling, 1861 described from "Fluss Heri-Rud bei Herat" (the Harirud at Herat in Afghanistan, formerly part of Persia) is a synonym. Bungia Keyserling, 1861 is a synonym of Gobio Cuvier, 1816 (Eschmeyer, 1990).

This species is represented in Iran by the subspecies Gobio gobio lepidolaemus Kessler, 1872, originally described as Gobio fluviatilis var. lepidolaemus from Ak-darja and Chodshaduk in the Zeravshan River basin, Uzbekistan and the Syr Darya at Khodzhent, Tajikistan. This subspecies is distinguished from the typical gudgeon by having a scale-covered throat, deep body, deep and short caudal peduncle, slightly notched or emarginate caudal fin and small size (Amanov, 1972). Berg (1948-1949) considers that these characters would be sufficient to distinguish this taxon as a full species but there are intermediate forms. Interestingly, Reshetnikov and Shakirova (1993) list Gobio lepidolaemus as a full species.

Also in the western Caspian Sea basin as Gobio gobio lepidolaemus natio holurus Berg, 1914 (the Terek gudgeon or Terskii peskar') but not recorded from Iranian waters and considered to represent intergrades between G. gobio gobio and G. gobio lepidolaemus by Bănărescu in Bănărescu (1999).

A syntype from Khodzhent of Gobio gobio lepidolaemus measuring 49.7 mm standard length is in the Natural History Museum, London under BM(NH) 1897.7.5:26, formerly in St. Petersburg University, a syntype is in the Zoological Institute, St. Petersburg under ZISP 2078 or 2076, and a possible syntype is in the Zoological Museum of Moscow State University under MMSU P.1052. Svetovidova (1978) refers to ZISP 2078 as the holotype on page 257 and ZISP 2076 as the holotype on page 262 (Eschmeyer et al., 1996).

Key characters

See above. This is the only gudgeon in eastern Iran and is separated from other cyprinids by the meristic characters, presence of barbels, absence of fin spines, mouth not transverse or crescentic but horseshoe-shaped, and colour pattern. It is separated from the other related species in Iran (Romanogobio macropterus and R. persus) by having the body and caudal peduncle compressed (caudal peduncle depth at anal fin insertion greater than caudal peduncle width) and by well-defined spots on the dorsal and caudal fins. This subspecies is also characterised by all members of the population having scales on the breast, absent or variably developed in other subspecies and populations to the west.

Morphology

Usmanova (1975) found differences between populations of this species in Uzbekistan associated with habitat. Fish from more stable habitats have deeper and wider bodies and reduced fin sizes.

Dorsal fin branched rays 6-8, usually 7, after 2-4 unbranched rays, anal fin branched rays 5-8, usually 5 (Amanov, 1972 - an error, see below) or 6 (Usmanova, 1975; Banarescu and Nalbant, 1973) after 2-3 unbranched rays, pectoral fin branched rays 13-17 and pelvic fin branched rays 7-8. Lateral line scales 33-46, usually 37-42 and averaging less than 40 (Bănărescu in Bănărescu (1999)). Scales have posterior radii only (or, if present, very few anterior radii) and the scale focus is subcentral anterior but not very eccentric in small fish, very eccentric in large fish. The anterior scale margin is rounded to wavy. There is a pelvic axillary scale. The anus is separated from the anal fin origin by 5-6 closely overlapping scales and is near the end of the pelvic fins but underneath them. Pharyngeal teeth 3,5-5,3 usually but Pipoyan (1998) found 21 variant counts for 141 Gobio gobio in Armenia with 3,5-5,3 (34.0%), 2,5-5,2(22.7%), 2,5-5,3(10.0%) and other combinations at about 2% or less. The anterior teeth are blunt with small hooks, conical and short and are followed by long, thin, strongly hooked teeth (the description of Bungia nigrescens may be in error in stating that there is only one row of teeth but Pipoyan (1998) notes that Gobio gobio in Armenia are exceptionally uniserial). Total gill rakers 1-7, only developed rakers being counted and anterior rakers reduced to bumps not included. Developed rakers are stubby and may or may not touch the adjacent raker when appressed. Total vertebrae 33-42 (this wide literature range may reflect the wide species range but could also include specimens counted in varying ways). The gut is an elongate s-shape with a slight anterior loop. The chromosome number is 2n=50 (Klinkhardt et al., 1995; Bănărescu in Bănărescu, 1999).

Meristics in Iranian fish are as follows: branched dorsal fin rays 7(4), branched anal fin rays 6(4), branched pectoral fin rays 15(2), 16(1) or 17(1), branched pelvic fin rays 7(4); lateral line scales 37(1), 38(2) or 39(1), scales around caudal peduncle 14(2) or 16(2); total gill rakers 5(1) or 6(2); pharyngeal teeth 3,5-5,3(1), 3,5-4,3(1) or 3,4-5,2 or 3(1); and total vertebrae 38(2) or 39(1).

Sexual dimorphism

Snout length, depth of the head at the occiput, eye diameter, greatest body depth and thickness, predorsal distance, pectoral-pelvic fin distance, and barbel length are greater in females than males while caudal, pectoral and pelvic fin lengths and height of dorsal fin are greater in males. Females reach larger sizes than males (Usmanova, 1975). Males darken in the spawning season. Fine tubercles develop on the side and upper surface of male heads, on the upper flank and back anteriorly, and the 8 outer rays of the pectoral fin.

Colour

The top of the head, the back and the flanks above the lateral line are dark brown and may have a greenish tinge. The lower flanks are paler and may have a silvery tinge or be a light yellow. The mid-flank bears a row of 6-13 dark spots which may merge into a line, merge in pairs or form a lattice. The back may have 4-5 longitudinal dark bands with a variegated pattern. The dorsal and caudal fins have 3-5 rows of spots, the pectoral fins have several rows of small spots and the pelvic and anal fins may also have 2-6 rows of spots but are often colourless. The peritoneum is silvery.

Colour may vary with habitat, being more uniform and darker on a monotonous background and more spotted and lighter on a gravelly background.

Size

Attains 20.0 cm total length for the species, 11.1 cm for lepidolaemus.

Distribution

Found from the British Isles and north of the Pyrenees through much of Europe north of the Alps to Siberia. The subspecies considered here is found only in the Tedzhen or Hari River basin in Iran (Abdoli, 2000). It is also found in north flowing rivers of the Kopetdag in Turkmenistan and eastwards in Central Asia to the Chu River in Kyrgyzstan. The natio holurus is found in the Kuma, Terek and Sulak rivers of the western Caspian shore but not as far south as waters neighbouring Iran.

This species is also recorded from the Karakum Canal and Kopetdag Reservoir in Turkmenistan (Shakirova and Sukhanova, 1994; Sal'nikov, 1995) and may eventually reach Iranian waters in the Caspian Sea basin.

Zoogeography

This is a widely distributed and very variable species with many taxa listed as subspecies, potentially species if further investigated. As part of a gobionine fauna found across Eurasia, this wide distribution may be suggestive of further work that could be done to clarify relationships of these fishes.

Habitat

This gudgeon is found in rivers near the bank where there is a sand or fine gravel bottom and in inlets, backwaters and weed beds of irrigation canals in the Kashkadar'ya of Uzbekistan (Usmanova, 1975). It prefers stagnant water in the Surkhandar'ya of Uzbekistan (Amanov, 1972). Gudgeons are generally resistant to pollution and varying environmental conditions although there is little information on the eastern Iranian subspecies.

Age and growth

Sexual maturity is reached at the age of 1 year and a length of 3.9-4.3 cm in females from the Kashkadar'ya in Uzbekistan, at 4.6-5.3 cm in the Surkhandar'ya, or at 2-3 years and 3-7 cm in reservoirs in Uzbekistan; also reported as 2-3 years in the Issyk Kul' and the Chu River (Usmanova, 1975). Life span is over 4 years (Amanov, 1972).

Food

Food is mainly benthic invertebrates, chiefly insect larvae such as chironomids in Iranian fish, but varied items may be taken depending on the water body and food availability. Abdoli (2000) lists Chironomidae, Ephemeroptera, Plecoptera and Trichoptera. Some insects falling on the water surface are taken. Remains of terrestrial plants, green algae and detritus have also been recorded (Usmanova, 1975). Detritus and vegetation (mostly diatoms) dominate in some gut samples, followed by insect larvae and benthopelagic crustaceans, and occasionally fish eggs. Other samples show chironomid larvae to be the main diet item or cladocerans. Vegetation fragments are apparently seized accidentally with such foods as chironomids (Amanov, 1972).

Reproduction

Spawning is intermittent and takes place from May to August in Uzbekistan, May-July in the Issyk Kul' and May-June in the Chu River (Usmanova, 1975). Up to 12,900 eggs are laid on a clay-sand bottom at 18-20°C (Amanov, 1972). Well-developed eggs are present in Iranian fish caught on 10 November. Elsewhere gudgeons slap the water surface with the rear part of the body, and males and females rub their bodies together while releasing eggs and sperm (Bănărescu in Bănărescu, 1999).

Parasites and predators

None reported from Iran.

Economic importance

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in aquaria and textbooks. Gudgeons in Europe have been used as bait fish and as food.

Conservation

Lelek (1987) classifies populations of this species in Europe as intermediate to rare.

Further work

The biology of this species in Iran requires study.

Sources

Type material: See above, a syntype of Gobio gobio lepidolaemus (BM(NH) 1897.7.5:26).

Iranian material: CMNFI 2007-0014, 3, 81.5-85.0 mm standard length, Khorasan, Kuh-e Sang Park, Mashhad (ca. 36º18'N, ca. 59º36'E).

Comparative material: SMF 17137, 5, 61.3-84.1 mm standard length, Afghanistan, tributary of the Harirud near Herat (34°21'N, 62°14'E).

Genus Hemiculter
Bleeker, 1859

This genus contains several species with a native distribution in the Amur River basin of the Russian Far East and in China, Taiwan and Viet Nam. A single species has been accidentally introduced to Iran.

The sawbellies are characterised by an elongate body; moderate-sized scales with a deeply-decurved lateral line only 1-3 scales above the mid-ventral line; a scaleless keel from the pectoral fins to the vent; a short dorsal fin with a spine and an elongate anal fin; pharyngeal teeth in 3 rows; gill rakers short and numerous; and pelagic eggs.

Hemiculter leucisculus
(Basilewsky, 1855)

Common names

tizeh kuli or teez-e-kooli (= sharp or spiny fish, kuli being any small fish), shakam tiz (= ?sharp belly), kuli-e mordab (= lagoon fish).

[vostrobryushka or sharpbelly in Russian; common sawbelly, knifefish].

Systematics

Culter leucisculus was originally described from Peking, China. Hemiculter eigenmanni (Jordan and Metz, 1913) is a synonym.

Key characters

The sharp keel, lacking scales, extending from the vent or anus to the throat below the pectoral fin on the mid-ventral surface is distinctive, especially when combined with the dorsal fin spine and the three rows of pharyngeal teeth.

Morphology

The last dorsal fin unbranched ray is a sharp spine with a flexible tip. The lateral line curves downward from its origin on the head to the level of the end of the pectoral fin and then parallels the lower body margin eventually to curve upward (sometimes sharply) at the end of the anal fin level and run along the middle part of the caudal peduncle. There is a tubercle on the lower jaw which fits into a notch on the upper jaw. Dorsal fin branched rays 6-8, usually 7, after 2-3 unbranched rays and anal fin branched rays 10-18, mostly 13-14 (but see below), after 3 unbranched rays. Lateral line scales 43-55, scales above the lateral line 8-11, and scales below the lateral line to the pelvic fin origin 1-3. Scales bear numerous fine circuli and a few posterior radii. Total gill rakers 17-29, reaching the second raker below when appressed. Pharyngeal teeth, 2,4,5-5,4,2, 2,4,5-4,4,2, 2,4,4-5,4,2, 2,4,4-4,3,2, 3,4,5-4,3,2 or 2,3,5-4,3,2. Teeth are hooked at the tip with an elongate and narrow grinding surface. Total vertebrae 42. The gut is an elongate s-shape. The chromosome number is 2n=48.

Meristic values for Iranian specimens are:- dorsal fin branched rays 6(1) or 7(6), anal fin branched rays 11(1), 12(5) or 13(1), lateral line scales 47(1), 48(1), 50(1), 52(1), 53(1) or 54(2), total gill rakers 17(1), 18(4) or 19(2), and pharyngeal teeth 2,4,5-5,4,2(2), 2,4,4-5,4,2(2), 2,4,5-4,4,2(2) or 2,4,4-4,3,2(1).

Sexual dimorphism

Unknown.

Colour

Overall colour dark above, silvery on the flanks and whitish on the belly. There is a dark stripe along the upper flank. The lips are dark. The dorsal, caudal and anterior pectoral and anal fin ray edges and their fin membranes are lightly pigmented with melanophores. The peritoneum is silvery with some melanophores giving it a brownish pigmentation in preserved fish.

Size

Reaches 25 cm.

Distribution

The native range of this species is from Maritime Russia south through China to Korea and Viet Nam. First reported from the Anzali Mordab by Holčík and Razavi (1992) and apparently not uncommon there. Abbasi et al., (1999), Kiabi et al. (1999), Abdoli (2000) and Gasmi and Mirzaei (2004) record this species from the lower Safid River and the Anzali Talab, and in the middle Aras River. Patimar et al. (2002a; 2002b; 2008) and Patimar (2008) report it from the International Wetlands of Alma-Gol, Adji-Gol and Ala-Gol and Esmaeili et al. (2010) from Zarivar Lake. Found in ab-bandans along the Caspian shore of Iran (Jolodar and Abdoli, 2004). Also reported in the Tedzhen River, Karakum Canal and Kopetdag Reservoir of Turkmenistan (as H. eigenmanni)(Aliev et al., 1988; Shakirova and Sukhanova, 1994; Sal'nikov, 1995) and so may eventually be found in the Tedzhen (= Hari) River basin of Iran. Now recorded from the Hawizah Marsh in southern Iraq (Coad and Hussain, 2007).

Zoogeography

This species is introduced to Iran, probably by accident along with commercial shipments of Chinese major carps from Central Asia in the former U.S.S.R. and/or Rumania in 1967. The Chinese major carps in Central Asia came from the Amur River basin in the Far East and sawbellies were accidentally transferred with them in the 1950s-1960s (Holčík and Razavi, 1992).

Habitat

Found in rivers, lakes, small ponds and swamps but little appears to be known about its habitat requirements.

Age and growth

Patimar et al. (2002a; 2002b; 2008) report six age groups in the International Wetlands of Alma-Gol, Adji-Gol and Ala-Gol. The smallest mature specimens there were 2 years old. It is an 'r' strategist, forming dense stunted populations in any new environment.

Food

Macrophytes, fish, crustaceans and insects are eaten by this species and young fish feed on zooplankton. Iranian fish contain large plant fragments and filamentous algae.

Reproduction

Up to 1,180 eggs are produced. Fish from a swamp near Hendeh Khaleh in Gilan taken on 9 August contained well-developed eggs but Patimar et al. (2002a; 2002b; 2008) report a peak spawning in March in the International Wetlands of Alma-Gol, Adji-Gol and Ala-Gol.

Parasites and predators

Sattari et al. (2007) record the digenean Diplostomum spathaceum and the monogenean Diplozoon sp. in this species in the Anzali wetland of the Caspian shore. It is food for Sander lucioperca, Silurus glanis and Aspius aspius in Turkmenistan (Aliev et al., 1988).

Economic importance

This species has potential as a food fish and is canned in China but this is probably outweighed by its competition with native species for food and the possibility of predation on fish eggs and young. The sawbelly is easily able to switch from one food to another as conditions warrant (Holčík and Razavi, 1992) and is known to show more rapid growth and higher fecundity than under native conditions. Welcomme in Courtenay and Stauffer (1984) regards this species as a pest when introduced.

It is found on the fish market at Bandar Anzali and a catch of 41 kg is reported from the Anzali Mordab in 1990 (Holčík and Oláh, 1992).

Conservation

None required for an introduced species.

Further work

The spread of this species and its effects on native species and habitats should be monitored.

Sources

Based on data in Holčík and Razavi (1992) for 5 Iranian specimens from the Anzali Mordab (134.5-143.4 mm standard length).

Iranian material: Uncatalogued material, 2, 99.6-113.1 mm standard length, Gilan, swamp near Hendeh Khaleh (37°23'N, 49°28'E).

Comparative material: CMNFI 2006-0028, 2, 112.5-123.6 mm standard length, Iraq, Hawizah Marsh (31º38'30"N, 47º35'21"E and 31º36'02"N, 47º33'09"E).

Genus Hemigrammocapoeta
Pellegrin, 1927

This genus comprises 5 species of which 1 is found in Iran. The genera Tylognathoides Tortonese, 1938, Neotylognathus Kosswig, 1950 and Hemigarra Karaman, 1971 are synonyms (Krupp, 1985c; Krupp and Schneider, 1989).

The genus is characterised by having a distinct rostral flap without separate lateral lobes, underslung mouth, horny covering to the upper and lower jaws, no disc on the chin behind the lower lip but this area papillose, no or up to 2 pairs of barbels (mouth and lip structures can be quite variable), short dorsal and anal fins, complete or incomplete lateral line, and pharyngeal teeth in 3 rows with somewhat hooked tips and spoon-shaped crowns, count uniquely 2,4,5-5,4,2 (but see below). Most species are found in the Levant. The genus is closely related to Garra according to Krupp (1985c).

Hemigrammocapoeta elegans
(Günther, 1868)

Common names

None.

Systematics

Originally described in the genus Tylognathus Heckel, 1843 and also placed in the genus Hemigarra Karaman, 1971. Tylognathus elegans was described from "Mesopotamia?" (sic). Six syntypes are in the Natural History Museum, London (BM(NH) 1850.10.21:31-36), one specimen (number 36) being an alizarin preparation. One of the specimens had been designated as a lectotype but was loose in the jar, and the jar contains a total of 17 fish measuring 36.0-44.5 mm standard length.

Key characters

Characters of the genus serve to identify this species in Iran, particularly around the mouth region.

Morphology

Dorsal fin with 3 unbranched and 6-8, usually 8 branched rays, anal fin with 3 unbranched and 5 branched rays, pectoral fin with 11-17 branched rays and pelvic fin with 8 branched rays. Last unbranched dorsal fin ray thickened although concave posteriorly, tapering distally to a thin ray. Unbranched ray anterior to the last one also thickened in large fish. Lateral line scales 33-38. Flank scales have a wavy anterior margin, very fine and numerous circuli, few anterior and more numerous posterior radii, posterior scale field with tubercles and a subcentral anterior focus. Pelvic axillary scale present. There is a scale between the anus and the anal fin origin. Gill rakers short, just reaching the base of the second raker below when appressed. Barbels 4, thin but moderately long. Snout projecting over inferior mouth but not strongly folded over the upper lip. Upper lip free. Lower lip thicker in the middle but no sucker, posterior edge wavy. Horny edge to lower jaw. Weakly papillose behind the lower lip. Large nasal flap and large first suborbital bone. Pharyngeal teeth in the major row quite massive and obviously hooked with a concave space below. The most anterior tooth is the largest and posterior teeth are slender. Minor row teeth are similar but smaller. Gut complexly coiled. Total vertebrae 46-48.

Meristic values for Iranian specimens are:- dorsal fin branched rays 6(1, ? deformed) or 8(7), anal fin branched rays 5(8), pectoral fin branched rays 16(2) or 17(6), pelvic fin branched rays 8(8); lateral line scales 34(1), 36(5), 37(1) or 38(1) and caudal peduncle scales 12(8); total gill rakers 17(1), 18(5), 19(1) or 20(1); pharyngeal teeth 2,3,5-5,3,2 (5) (sic); and total vertebrae 46(1), 47(5) or 48(2).

Sexual dimorphism

Unknown.

Colour

There is an indistinct silvery stripe along the flank. Dark pigment ends about 1 scale above the lateral line along the flank. Peritoneum black with some silvery patches.

Size

Reaches 10.9 cm total length.

Distribution

Found in the Tigris-Euphrates basin. Initially recorded from Harmaleh on the Dez River of Khuzestan (ZSM 25716, 25717), Abdoli (2000) maps distributions from the upper Karun, upper Dez, and upper Karkheh including the Simarreh rivers.

Zoogeography

The relatives of this species are confined to the Levant, a distribution matched by various other unrelated taxa indicating former connections between these basins (see Krupp (1985c; 1987)).

Habitat

Unknown.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

No parasites reported from Iran. One specimen examined was from the gut of a Silurus triostegus taken at Harmaleh on the Dez River.

Economic importance

None.

Conservation

This species is rare in Iran and is poorly known elsewhere. Assuming this is not the result of inappropriate collecting techniques, then the status of this species should be assessed by field surveys.

Further work

The distribution and biology of this species is very poorly documented and needs attention.

Sources

Type material: See above (BM(NH) 1850.10.21:31-36).

Iranian material: ZSM 21862, 5, 60.5-86.9 mm standard length, Khuzestan, Dez River at Harmaleh (31º57'N, 48º34'E); ZSM 25716, 2, 76.4-79.3 mm standard length, same locality as preceding; ZSM 25717, 1, 78.5 mm standard length, "from the stomach of a wels", same locality as preceding.

Comparative material: BM(NH) 1973.5.21:185, 1, 57.5 mm standard length, Iraq, Tigris River at Jadriyah, Baghdad (no other locality data); FMNH 51235, 1, 43.9 mm standard length, Iraq, Tigris River at Rustam Farm near Baghdad (no other locality data); FMNH 51236, 12, 38.7-59.5 mm standard length, same locality as preceding.

Genus Hypophthalmichthys
Bleeker, 1859

The silver carp genus contains 3 species with a native distribution in eastern Asia. Two species have been widely introduced for food in aquaculture and for phytoplankton control.

The genus is characterised by an elongate and compressed body, very small scales (usually over 100 in the lateral line), eyes low on the head with their lower margin below the mouth corner level, a terminal mouth, no barbels, gill rakers long and thin, a very long gut, branchiostegal membranes joined and free of the isthmus, a short dorsal and elongate anal fin, both spineless, pharyngeal teeth in 1 row, and a ventral keel from the throat or pelvic fins to the anus.

Hypophthalmichthys molitrix
(Valenciennes, 1844)

Hypophthalmichthys molitrix from Wikimedia Commons.

Common names

فيتوفاگ (= fitofag or phytophag, i.e. phytophage or plant eater), كپور نقره اي (= kopur-e noqrehi or kapoor-e-noghreie, meaning silver carp), آزاد پرورشي (= azad-e parvareshi meaning free farmed).

[silver carp, thickforehead, white thickforehead; belyi tolstolobik, tolpyga and maksun in Russian].

Systematics

Leuciscus molitrix was originally described from China. Populations in the Anzali Mordab are hybrids with Hypophthalmichthys nobilis (J. Holčík, in. litt., 1989).

Key characters

The abdomen has a compressed keel extending from the breast to the vent, the eyes are positioned low such that they are visible from the underside of the head, and scales are minute. The similar bighead (H. nobilis) can be distinguished by the long pectoral fins which extend past the origin of the pelvic fins, a shorter keel (pelvic fins to anus), and gill raker structure (free, no spongy root mass).

Morphology

Dorsal fin with 2-3 unbranched rays followed by 6-7 branched rays, anal fin with 2-3, usually 3, unbranched rays and 11-15 branched rays, pectoral fin branched rays 17, and pelvic fin branched rays 7. Lateral line scales 85-125 Scales are rounded to oval with a posterior focus and very few posterior radii. Total vertebrae 36-40. Pharyngeal teeth 4-4, well-developed, compressed and with striated grinding surfaces. Gill rakers exceed 650 and are longer than the gill filaments. The left and right sides of the gill arches are united by a mucous membrane to form a continuous band; the gill rakers are distinguishable distally but the roots form a spongy mass. The gut is long and complexly coiled. The diploid chromosome number is 48 (Klinkhardt et al., 1995; Varasteh et al., no date). There were 6 pairs of metacentric, 14 pairs of submetacentric and 4 pairs of acrocentric chromosomes in the latter study.

Sexual dimorphism

Unknown.

Colour

Overall colour is silvery, the back bluish to grey-black with upper flanks olivaceous but silver laterally and ventrally, and the fins red or immaculate.

Size

Reaches 1.50 m and 56 kg.

Distribution

The natural distribution is from the Amur River in the former U.S.S.R. southward to southern China. In Iran, it has been introduced to the Sistan basin including the Hamun Sabari and the canal flowing into Chahnimeh (Ahmadi and Wossughi, 1988; Mansoori, 1994; J. Holčík, in litt., 1996), the Voshmgir Reservoir on the Gorgan River (Petr, 1987) as well as widely stocked in Mazandaran and Gorgan provinces in reservoirs and lakes by both governmental agencies and private companies. It is also introduced to the Anzali Mordab and its outlets (Holčík and Oláh, 1992), the Gorgan, Tajan, Aras, and Safid rivers, and the Anzali Mordab (Kiabi et al., 1999; Abbasi et al., 1999), Lake Zaribar, Kordestan (Abzeeyan, 5(5):III, 1994), reservoirs and fish farms in Khuzestan, Mahabad Dam (Abdi, 1999; www.mondialvet99.com, downloaded 31 May 2000; Shamsi et al., 2009).

Abdoli (2000) and Abdoli and Naderi (2009) record this species generally from the Dasht-e Kavir, Kerman-Na'in, Sistan and Hamun-e Jaz Murian basins; from the middle and lower Kor River and the Pulvar River in the Kor River basin; from the middle and lower Zayandeh River in the Esfahan basin; from the Khersan River, lower Karun River, the lower Jarrahi river, the lower Dez River the upper Karkheh River and the lower Kashkan River in the Tigris River basin; the middle and lower Qareh Su and Qom rivers, the lower Karaj and Shur rivers in the Namak Lake basin; the lower Zarrineh river and the middle and lower Talkheh River in the Orumiyeh basin; the middle Kashaf River in the Tedzhen basin; the middle and lower Atrak and Gorgan rivers, the Aras River, the lower Babol, Heraz, Chalus, Tajan, Tonekabon and Safid rivers, the Anzali Talab and the Caspian coast generally in the Caspian Sea basin.

This species is also recorded from the Karakum Canal and Kopetdag Reservoir in Turkmenistan (Shakirova and Sukhanova, 1994; Sal'nikov, 1995) and may eventually reach Iranian waters in the Tedzhen (= Hari) River and Caspian Sea basins. It is pond-cultured in Iraq and is known from open waters such as the Shatt al Arab and Tigris River (Al-Hassan, 1994).

Zoogeography

This species is an exotic, introduced to Iran from a variety of sources. It may become established in the large river systems of southern Iran and Iraq from escapees (Al-Hassan, 1994).

Habitat

This species is a riverine fish in its native habitat, or is found in water bodies connected to rivers, but is extensively cultivated in ponds for food throughout Asia. Silver carp can live in the Caspian Sea at salinities of 5-8‰ although a few are found at 10-12‰. It enters rivers to spawn (Abdusamodov, 1986). Temperatures in the range 0-40ºC are tolerated although 26-30ºC or 30-34ºC is preferred in different studies. It is more cold-tolerant than bighead carp. This species can be difficult to catch as it will jump over nets, to a height of about 2 m, and when frightened by noise has been known to jump into boats. Malek Nedjad and Parivar (1993) consider that the level of lead pollution in the Anzali Mordab (average 0.124 p.p.m. in surface waters, 0.1956 p.p.m. in deeper waters) caused 8% mortality in fertilised eggs and nearly 18% of eggs are useless for fisheries work.

Age and growth

Terek River silver carp first mature at 4 years for males and 5 years for females. About 15% of females mature at 4 years but 87% of the females and 85% of the males are in the 5-7 age groups (Abdusamadov, 1986). Maturity varies with locality, at 2-8 years, with males maturing a year earlier than females. Silver carp can reach 18-23 kg in 4-5 years. Life span is at least 20 years. Abdolmalaki (2004) examined the fishery for this carp in the Mahabad Reservoir, Iran and found mean fork length was 51.15 cm and mean weight was 2272.1 g. The length-weight relationship was W = 0.013L3.04. The von Bertalanffy growth parameters were L = 150 cm, K = 0.128, instantaneous rate of total (Z), natural (M) and fishing (F) mortality were 1.68/year, 0.22/year and 1.46/year. The exploitation rate (E) was calculated to be 0.82. The biomass was 158.5 t with a maximum sustainable yield estimated at 68 t. A decreased fishing effort was recommended as the annual catch in 1998-1999 was 101,123.5 kg.

Food

The gill rakers form a very fine, sponge-like mesh used to filter small planktonic food, aided by the epibranchial organ that produces mucus to trap very small particles. This species is a pump filter feeder, taking smaller particles than bighead carp. Food in Lake Kinneret, Israel is phytoplankton from February to August and predominately zooplankton from September to January, a response to a decrease in phytoplankton biomass in summer-fall (Spataru and Gophen, 1985). Cladocerans and cyclopoid copepods dominate the biomass of zooplankton taken. The ability to take cyclopoids is due to the large mouth, strong sucking power and the high filtration rate when feeding. Food is taken passively rather than selectively. Mohammadi et al. (2003) found detritus and protozoans to be the main items in gut contents, with also other algae, diatoms and green algae respectively.

Reproduction

Silver carp require cool, flowing water to breed. The spawning migration begins at the end of April in the Terek River of Dagestan at 16-17°C, with a peak between the middle of May and the beginning of June. Generally spawning occurs between 18 and 26°C. Fecundity reaches 1,340,500 large, greyish eggs (and in the Kara-kum Canal 1,525,000 eggs; elsewhere to 5.4 million eggs). Water hardened eggs are 4.9-5.6 mm, smaller than those of bighead carp. Spawning takes place after a sharp rise in water level and current speed. Spawners chase each other near the water surface where eggs and sperm are shed. Eggs are first found in the drift in the second week of June and hatch 34-70 hours later depending on temperature. Some larvae reach rice fields and live there until autumn when the fields dry up where some are lost, others migrating. Other larvae are carried into the Caspian Sea where they are sensitive to the prevailing salinity at 1-1.5 days old (Abdusamadov, 1986). It does not breed in the wild in Iran.

Parasites and predators

Mokhayer (1989) reports metacercariae of the eye fluke, Diplostomum spathaceum from this species in Iran, which can cause complete blindness and death in commercially important species, as well as black-spot, Posthodiplostomum cuticola. Jalali and Molnár (1990b) record the monogenean Dactylogyrus hypophthalmichthys from this species at fish farms in Iran. Masoumian and Pazooki (1998) surveyed myxosporeans in this species in Gilan and Mazandaran provinces, finding Myxobolus pavlovskii. Akhlagi (1999) reports that high temperatures (up to 32°C) stresses this species and leaves it open to infection with Aeromonas hydrophila. Safari and Khandagi (1999) record Clostridium botulinum from 1.1% of fresh and smoked samples of this species in Mazandaran Province. Ebrahimzadeh Mousavi and Khosravi (1999; www.mondialvet99.com, downloaded 31 May 2000) record the toxigenic fungi Aspergillus flavus, Alternaria, Penicillium and Fusarium from this species and the pond water at a fish farm in northern Iran. Akhondzadeh et al. (2002) and Akhondzadeh Basti and Zahrae Salehi (2003) show that the psychotropic pathogen Listeria monocytogenes is found in market and fish farm samples of this species. Naem et al. (2002) found the following parasites on the gills of this species from the western branch of the Safid River, namely the protozoans Ichthyophthirius multifilis and a Trichodina species and the  monogenean trematode Dactylogyrus hypophthalmichthys. Jalali et al. (2002) and Jalali and Barzegar (2006) record several parasites from this species in Lake Zarivar, namely Dactylogyrus hypophthalmichthys, D. suchengiaii, Diplostomum spathaceum, two species of Argulus, and myxosporean plasmodia. Jalali et al. (2005) summarise the occurrence of Gyrodactylus species in Iran and record G. sprostonae from Safid River fish. Araghi Soureh and Jalali Jafari (2005) recorded Dactylogyrus hypophthalmichthys and D. suchengtaii from this species in the Mahabad River of the Lake Orumiyeh basin, the latter species being a new record for Iran. Barzegar et al. (2008) record eye parasites from this fish including the digeneans Diplostomum spathaceum and Tylodelphys clavata, and the crustacean Lernaea cyprinacea. Shamsi et al. (2009) found Dactylogyrus hypophthalmichthys and D. suchengtaii in this species from fish farms, and Zarivar Lake, and Mahabad reservoir, respectively. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Argulus sp., Argulus foliaceus, Lernaea sp. and Lernaea foliaceus on this species.

Economic importance

This species is the most productive freshwater fish in the world, with 3.1 million metric tons produced in 1997 (versus 2.2 million mt for Cyprinus carpio).

Holčík and Oláh (1992) report a total catch of 6585 kg in the Anzali Mordab in 1990, at 8.8% of the total fish catch being the third largest catch. This species is used in polyculture with common and grass carps and comprises 50-63% of the fishes. They are fed through pond fertilization without supplementary feeding (Emadi, 1993b). Silver carp in oil are packaged in northern Iran (http://www.netiran.com/business.html, downloaded 31 October 2003). They are also found in fish stores in Ahvaz, Khuzestan (personal observations, September 1995). Iran shares with Uzbekistan the most production of this species among North African and Near Eastern aquaculture; the Iranian catch increasing from nothing in 1989 to 24,720 tonnes in 1994 (Food and Agriculture Organization, Fisheries Department, 1996). Bartley and Rana (1998b) however give a production of 15,228 t for 1995. Market price in 1995 was about U.S.$1.00/kg in 1995, lower than for grass carp at about $2, but silver carp have the higher stocking ratio (Rana and Bartley, 1998a). Kals and Bartels (2004) give some recommendations for improving silver carp farming in Iran.

Abdolmalaki (2004) reported that 67.3% of the 150,261 kg of fishes caught in Mahabad Reservoir in the1998-1999 fishing season were this species. The fishing effort was 69 beach seine hauls and 2530.7 fishing effort units with 7-15 cm gill nets, each unit being 100 m of gill net for 24 hours.

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in aquaculture and food. The food of this species being phytoplankton it has been introduced to areas lacking filter feeders such as Europe and Israel in efforts to control algal blooms associated with eutrophication. The results in Israel were controversial and difficult to interpret because of chemical treatments for nuisance dinoflagellates and introductions of other fish species (Opuszynski and Shireman, 1995). In some cases the effects of the introduction were negative because the silver carp fed from September to January on zooplankton which themselves reduce algal blooms. There was also competition with a more valuable native species of cichlid for the same zooplankton resource and the native cichlid was a more efficient consumer of the nuisance dinoflagellate. Elsewhere, the consumption of plankton may remove food for native, larval fishes.

This species has been studied extensively as a commercial species in various parts of the world. Studies specific to Iran include Azari Takami et al. (2000) on female broodstock selection; Nazifi et al. (2000) on the effects of the organophosphate trichorofon on serum biochemistry; Nazifi et al. (2001) on haematological parameters after experimental intoxication with trichlorofen; Javadian et al. (2003) on the effect of ice storage on lipid and other chemical changes, moisture and texture and appearance; Moradian et al. (2003) on the effects of the hormone thyroxine on the survival of early growth stages (the ratio of hatched eggs in 0.5 p.p.m. thyroxine was highest); Kashani Sabet et al. (2004) on inducing ovulation with hormones combined with dopamine antagonists; Peyghan et al. (2002, 2006) on deleterious effects of malachite green, used to prevent fungal and parasite infections, on eggs and larvae; Alavi Talab et al. (2007) on use of skins and fins for gelatin extraction; Moeini et al. (2008) on producing surimi from this fish; Roshan and Moini (2009) on washing minced flesh with cold water and brine to increase quality of surimi (1.5% brine for 10 minutes was best); Shabanpour et al. (2008) on lipid quality changes of surimi during frozen storage; Alavi Talab et al. (2010) on optimising encapsulation of oil from this species; Choobkar et al. (2010) on the use of an essential oil from Persian thyme (Zataria multiflora) to combat Staphylococcus aureus in salted fillets; etc.

This fish can be dangerous to boaters as it leaps out of the water (Kolar and Lodge, 2002). Akhondzadeh Basteh et al. (2006) found the bacterial pathogens Listeria monocytogenes in fresh and smoked H. molitrix, Staphylococcus aureus in smoked H. molitrix, Escherichia coli and Salmonella dublin in fresh H. molitrix and Vibrio haemolyticus in smoked H. molitrix.

The Iranian Fishery Research Institute has made ice cream using this fish. The product is deoderised so there is no fishy flavour and a protein in the fish blood is reported to lower the temperature at which ice crystals form meaning less cream or fat is needed in the product (Iran Daily, 11 February 2009).

Conservation

Krasznai (1987) and Petr (1987) give details of fish farms propagating this species in Iran. For example, 30 million fish were produced by the Safid Rud Fish Farm in 1986. 20 million carp, silver carp and grass carp fingerlings were produced in the Shahid Rajaae Hatchery in Sari for release across Iran in reservoirs and dams (Abzeeyan, Tehran, 4(7):VII, 1993). Piri and Ordog (1999) describe the effects of herbicides and insecticides on this species. These chemicals are used extensively on rice fields in Gilan where aquaculture is widely developed.

Hybridisation (presumably with H. nobilis) has led to poor growth and a decline in the fishery (Shehadeh, 1997).

Further work

The biology of this species in Iran needs investigation in relation to its effects on native fishes and its distribution monitored.

Sources

Opuszynski and Shireman (1995) summarise the biology and culture of this species.

Comparative material: CMNFI 1977-0590, 3, 120.3-135.5 mm standard length, Israel, Kibbutz `En Hamifraz (32º59'N, 35º05'E).

Hypophthalmichthys nobilis
(Richardson, 1844)

Hypophthalmichthys nobilis, Wikimedia Commons.

Common names

sar gondeh or sargundeh (= bighead) in the Caspian basin, كپور سرگنده (= kopur-e sargondeh, meaning bighead carp), mahi kopur sar gondeh, fitofag.

[pestryi tolstolobik in Russian; bighead, painted thickforehead].

Systematics

Leuciscus nobilis was originally described from Canton, China. Eschmeyer et al. (1996) give the date of publication as 1845, Reshetnikov et al. (1997) as 1846.

Howes (1981) reaffirms the placement of this species in Hypophthalmichthys Bleeker, 1860, considering that the characters of abdominal keel length, pharyngeal dentition and gill raker form are insufficient to place this species in the distinct genus Aristichthys Oshima, 1919. Other authors disagree (see Eschmeyer, 1990; Reshetnikov et al., 1997).

Populations in the Anzali Mordab are hybrids with Hypophthalmichthys molitrix (J. Holčík, in litt., 1989). Artificial hybrids with Ctenopharyngodon molitrix have been developed in Iran and their karyology studied (Dorafshan and Kalbasi, 2007). The F1 hybrids were triploids with 3n=72 and NF=132.

Key characters

The similar silver carp (H. molitrix) can be distinguished by the short pectoral fins which do not extend past the origin of the pelvic fins, a longer keel (throat to anus), and gill raker structure (continuous band uniting both sides, roots fused into a spongy mass).

Morphology

Dorsal fin unbranched rays 3 followed by 7 unbranched rays, anal fin unbranched rays 3 with 11-14 branched rays, pectoral fin branched rays 16-19, and pelvic fin branched rays 7-9. Lateral line scales 92-115. Scales are a rounded oval with a slightly posterior focus, very few posterior radii and numerous fine circuli. Total vertebrae number 36-41. Pharyngeal teeth 4-4, with smooth grinding surfaces. Gill rakers are very numerous. The gut is elongate and convoluted. The diploid chromosome number is 48 (Klinkhardt et al., 1995).

Sexual dimorphism

Males have a sharp edge along the dorsal surface of several anterior pectoral fin rays.

Colour

Overall colour is silvery and the body has numerous scattered small black spots which makes some fish speckled or darker in overall colour. A few larger blotches may be present. The head is often darker and contrasts with the silvery body. Fins are greyish and similar to the adjacent body colour with the caudal fin darkest, sometimes reddish.

Size

Attains 1.57 m in length and 77.5 kg, possibly to 1.95 m.

Distribution

This species is a native of China and was first introduced to Iran in 1966 from a hatchery in the Krasnodar region of the former U.S.S.R. (Anonymous, 1970b). It is reported from fish farms in the Caspian Sea basin and is stocked in reservoirs throughout Iran (Coad and Abdoli, 1993b). Introduced to the Kor River in Fars (A. Alamdari, in litt., 1997) and reported from the Gorgan, Tajan, Aras, and Safid rivers, and the Anzali Mordab (Abbasi et al., 1999; Kiabi et al., 1999; Jalali et al., 2005). Abdoli (2000) and Abdoli and Naderi (2009) record it generally from the Sistan, Hormuz, Kor, Kerman-Na'in, Dasht-e Kavir, Esfahan, Namak, Tigris, and Orumiyeh basins and from the Aras and Safid rivers, lower Gorgan, Neka, Babol, Haraz, Tajan, Chalus and Tonekabon rivers, the Gorgan and Anzali mordabs and along the Caspian coast. Also in Lake Zarivar (Shamsi et al., 2009).

Also reproducing naturally in the Karakum Canal and recorded from the Kopetdag Reservoir in Turkmenistan (Shakirova and Sukhanova, 1994; Sal'nikov, 1995) and may eventually reach Iranian waters in the Tedzhen (= Hari) River basin. Introduced to Iraq for fish farming.

Zoogeography

An exotic introduced to Iran.

Habitat

In their natural habitat, bigheads are found in large rivers and associated floodplain lakes. They migrate upstream to spawning grounds when water levels rise, moving to flooded land afterwards, and returning to the river channel as water levels fall. Bigheads can live in the Caspian Sea at salinities of 5-8 p.p.t. although a few are found at 10-12 p.p.t.. They can adjust gradually to salinities of 15-20 p.p.t. They enter rivers to spawn (Abdusamodov, 1986) but are known to spawn in the Karakum Canal of Turkmenistan. Preferred temperatures for feeding and reproduction are within the general range 20-30°C. Activity almost ceases at 10°C and the critical thermal maximum is 38.8°C.

Age and growth

Males achieve first maturity at age 5 and males of 5-7 years and 81-90 cm make up 90% of the run in the Terek River. Most females mature at 6 years and 81% of the females on the spawning run are 6-7 years old and 75-100 cm (Abdusamodov, 1986). Males mature at 2-4 years and females at 3-5 years (and 10 kg) in Turkmenistan. Sexual maturity varies widely with environmental conditions, 2-6 years for males and a year later for females. Life span is up to 16 years. Growth is rapid, attaining 18-23 kg in 4-5 years.

Food

Zooplankton is almost exclusively the food of this species. Phytoplankton and detritus may be taken when zooplankton biomass is low. Most feeding occurs during summer and peaks daily in the range 1200 to 2000 hours. This species is both a pump feeder, using the buccal pump to push food-laden water through the gill rakers, and a ram feeder, swimming with the mouth open to force water through the gills, with intermittent gulps. Feeding often occurs at the water surface, in contrast to silver carp, as well as in the water column and on the bottom.

Reproduction

A spawning migration of this species enters the Terek River in the second week of May at water temperatures of 18-19°C, numbers increasing until the end of June. Spawning takes place after a sharp rise in water level and current speed. Males actively chase females near the water surface, occasionally butting the female's belly, and sometimes leaping out of the water. Eggs and sperm may be cast into the air. Fecundity attains 1,860,800 eggs. Unswollen eggs are 1.4-1.5 mm in diameter and water hardened eggs are 5.7-6.2 mm. Eggs are first found in the drift in the second week of June and hatch 34-70 hours later depending on temperature. Some larvae reach rice fields and live there until autumn when the fields dry up where some are lost, others migrating. Other larvae are carried into the Caspian Sea where they are sensitive to the prevailing salinity at 1-1.5 days old (Abdusamadov, 1986).

Parasites and predators

Jalali and Molnár (1990b) record the monogeneans Dactylogyrus aristichthys and D. nobilis from this species in Iranian fish farms. Naem et al. (2002) found the following parasites on the gills of this species from the western branch of the Safid River, namely the protozoan Trichodina sp., monogenean trematodes Dactylogyrus nobilis, D. aristichthys, and Gyrodactylus sp.. Jalali et al. (2005) summarise the occurrence of Gyrodactylus species in Iran and record G. sprostonae and G. sp. in fish from the Safid River. Masoumian and Pazooki (1998) surveyed myxosporeans in this species in Gilan and Mazandaran provinces, finding Myxobolus pavlovskii. Alishahi and Peyghan (2008) found a heavy infestation with Lernaea cyprinacea from a fish pond in Tehran. Barzegar et al. (2008) record the digenean eye parasite Diplostomum spathaceum from this fish. Shamsi et al. (2009) found Dactylogyrus aristichthys, D. nobilis and D. taihuensis in this species from fish farms and Zarivar Lake. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Lernaea sp. and Lernaea cyprinacea on this species.

Economic importance

Holčík and Oláh (1992) report a catch of 466 kg in the Anzali Mordab in 1990. Aquaculture production in 1995 was 1269 tonnes (Bartley and Rana, 1998b). Marjan Iran Company was selling 1500-1800 g fish for U.S.$1.90/kg in August 2003 (http://groups.yahoo.com/groups/hilsa/message/25).

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in aquaculture and food. It ranks fourth on world aquaculture production. This species has been used in Israel to reduce zooplankton populations in reservoirs in an effort to improve water quality (Opuszynski and Shireman, 1995). It has a higher net production in culture systems than H. molitrix and Ctenopharyngodon idella. The consumption of plankton may remove food for native, larval fishes, and affect the diet of piscivorous fishes and birds. This species can also cause habitat alteration, increasing turbidity, and introduce diseases and parasites. In Iran, it has been used in aquaria for investigations on the effect of lead nitrate on blood serum electrolytes (Jamili et al., 2006).

Conservation

Krasznai (1987) and Petr (1987) give some details of propagation of this species in Iran. Some populations are hybrids (see above) and there is a danger of loss of genetic purity in fish farm stocks should breeding adults be captured in the wild. As an exotic, there is no need for conservation.

Further work

The biology of this species in relation to native species should be investigated for Iran.

Sources

Jennings (1988), Opuszynski and Shireman (1995) and Kolar et al. (2005) summarise the biology and culture of this species.

Comparative material: CMNFI 1980-0530, 2, 230.6-255.8 mm standard length, Japan, pond cultured (no other locality data).

Genus Iranocypris
Bruun and Kaiser, 1948

This genus contains a single species found only in Iran and the characters of the species are therefore the characters of the genus. This blind cave species is placed in a world-wide context by Proudlove (1997a; 1997b).

Iranocypris typhlops
Bruun and Kaiser, 1948

    
Dorsal view of head                                             Ventral view of head

Courtesy of R. Mehrani

Courtesy of Amir Hosin Zalaghi, 11-19 May 2010 (all fish returned to cave alive)

Courtesy of Kiavash Golzarianpour

Iranocypris typhlops movie.3gp, courtesy of Kiavash Golzarianpour

Common names

mahi-ye kureghar or mahi-ye kur-e qar (= blind cave fish), kopur mahi kureghar, ماهي كور (= mahi kur).

[Iran cave barb].

Systematics

The holotype is in the Zoological Museum of Copenhagen (ZMUC P 26475) and measures 46.5 mm total length and 38.5 mm standard length (Nielsen, 1974; personal observations). The paratypes number 5 (in Nielsen (1974)), or 6 (in Bruun and Kaiser (1948)) but only 4 were found in ZMUC in December 1999. Paratypes (ZMUC P 26476, 26477, 26478, 26480) measure 19.5-42.0 mm total length and 16.5-34.5 mm standard length according to Bruun and Kaiser (1948). Two fish (P 26476 and P 26480) were used in histological studies and one consists of the body only. The type locality is given below and the fish were collected by E. Kaiser on 6-5-1937 from "lok 80" (= locality 80; but no field notes by E. Kaiser are available in ZMUC).

The date of authorship for this species is variously listed as 1943 on an official reprint, as 1944-49 in one set of Contents and "ready from the press 1944" in another set of contents. Proudlove (2006) states that is did not appear until 1948 because of World War II.

Bruun and Kaiser (1948) believe this species to be related to the genus "Barbus", members of which also have two pairs of barbels, although Saadati (1977) considers this unlikely since most "Barbus" from the Tigris River basin are large fishes.

Key characters

The only eyeless, depigmented cyprinid species in Iran, it is very distinctive.

Morphology

The body is compressed and the head somewhat flattened. There are two pairs of barbels, one pair at the mouth corners and one about half way along the upper lip. The upper lip has a feebly crenulated edge. The mouth is subterminal and horseshoe-shaped. A mental disc is developed in some fish, absent in others. There are significant differences in some morphometric and meristic characters in fish with, and without, a disc (Sargeran et al., 2008). There is no visible trace of eyes in most fish. The skin is naked except for a few rows of scales behind the pectoral fin base, although some individuals may have more flank scales. There are about 32 myomeres  along the flank. A lateral line is present. The dorsal fin has 3 unbranched and 7-8 branched rays, the anal fin 3 unbranched and 4-5 branched rays, the pectoral fin 14-17 branched rays and the pelvic fin 5-7 branched rays. Pharyngeal teeth in 3 rows, 1 to 3 in the outer row, 3 to 4 in the middle row and 3-5 in the inner row. Anterior teeth are very enlarged and conical, appearing as rounded knobs while the posterior teeth in the main row are flattened and slightly hooked. Smaller fish have less conical anterior teeth with a tiny hook at the tip and posterior teeth have a short, flat to slightly concave surface below the tip. Tooth counts are difficult to make with accuracy as it is not always clear to which row a tooth belongs. Smaller fish can be interpreted as 2,3,5-5,3,2 while larger fish may possibly lose a tooth and have a 2,3,4-4,3,2 count. Abbasi and Gharezi (2003) give a 3,5,5-5,3,3 count. Gill rakers very short, not reaching the adjacent raker when appressed and numbering 10-13 total. Total vertebrae 34-36. Gut s-shaped. The morphology and histology of the digestive tract was examined in detail by Abbasi and Gharezi (2003).

Meristics in specimens examined, including the holotype and 2 paratypes: dorsal fin branched rays 7(30) or 8(29); anal fin branched rays 4(1) or 5(49); pectoral fin branched 12(1), 13(9), 14(8), 15(23) or 16(8); pelvic fin branched rays 6(18), 7(31) or 8(1); total gill rakers 10(2), 12(4) or 13(1); pharyngeal teeth 2,3,5-5,3,2(1), 3,4,5-5,4,2(1), 2,3,4-5,3,2(1) or 2,3,4-4,3,1(1); total vertebrae 34(3), 35(8) or 36(1).

Sexual dimorphism

None reported.

Colour

This species is almost entirely unpigmented although live fish are pinkish to red from the blood showing through the skin. The gill filament area is bright red and some fish give an overall impression of red like a goldfish. Small, black pigment cells were visible in two small fish over the brain and just behind it and in these two fish and three others a very small, black pigment spot deep in the tissues on the side of the head may indicate a rudimentary but non-functional eye. Gut contents are visible through a semi-transparent body wall. Preserved fish are yellowish-white.

Size

Reaches 55.0 mm total length (Kiavash Golzarian, pers comm., 6 April 2008).

Distribution

Found only at "Kaaje-ru" above the garden "Bagh-e Loveh", "Lowa" or "Levan" (probably Loven at 33°04'N, 48°37'E) which is about 4 km from kilometre 382 on the railway from Bandar Shapur to Tehran and approximately 12 km north of the railway station Tang-e Haft. The stream below the cave locality is the "Ab-e Serum" which runs into the "Ab-e Zezar" which is a tributary of the Dez River, in Lorestan Province. Further locality details are given in Bruun and Kaiser (1948). The locality is at 744 m and 33°04'38.6"N, 48°35'33.1"E according to the Iranian Fisheries Research and Training Organization Newsletter, 21:3, 1998 and Kiavash Golzarian, pers comm., 6 April 2008).

Zoogeography

The relationships of cave species, with their reduced characters, are problematical but the three rows of pharyngeal teeth and mouth structures indicate a possible relationship with Garra.

Habitat

Known only from a well-like but natural outlet of a subterranean system. The outlet overflows to form a small stream from January to May (Smith, 1979) during the snow-melt period in the Zagros Mountains but in April to June this flow ceases (the precise timing of flow and its cessation is estimated from villager's comments and scientific visits and also varies with precipitation). Pictures of show flowing water in May 2010 are shown in the account of Paracobitis smithi Nemacheilidae). The well area is about 5 by 3 m and gradually decreases as the year progresses. Divers descended to a depth of 60 feet (= 18.3 m) in 1977 in the "well" until the resurgence narrowed (Farr, 1977). A rope was let down by R. Mehrani (pers. comm., 2000) and reached 23 m before the rope ran out and yet it was not at the bottom. Smith (1979) reports divers descending to 60-70 feet (18.3- 21.3 m). The pool shelves deeply under the cliff rearwards but the whole pool surface is exposed to light. There is no vegetation in the pool except for some encrusting algae on the rocky sides. The shale fragments forming the outermost floor of the pool have a thin layer of mud on them which may contain algae.

It seems probable that a complex of flooded but narrow and inaccessible passages is the habitat of this species and the well is merely the surface manifestation of this complex (Bruun and Kaiser, 1948; Smith, 1978; Banister, 1992). There is a smaller pool (about 2 m across narrowing rapidly inside) and flowing exit stream lower down the gorge, about 50 m away from the main locality, where a blind fish was seen but not caught in December 2000 (Smith (1979) also tentatively reports sighting a fish here). This is assumed to be evidence of the interconnectivity of subterranean passages. The main pool was not flowing at this time. The stream from the smaller pool increases in flow downstream, possibly tapping more groundwater, and eventually has a moderate flow. No fish were seen in it. The stream falls over a high waterfall (estimated at 10-15 m high by Smith (1979) which seems about right) so the well localities are isolated from the local fishes in the main river. The main river houses Garra rufa and nemacheilid species. The stream shows evidence of recent higher flow which tends to confirm overflow from the main well. More photographs can be seen in the description of Paracobitis smithi (Nemachelidae).

The fish may be seen swimming freely in the well, up to a 20 at a time may be counted. They can be caught with a dip-net.

Sampling in December 2000 recorded a water temperature of 18.5°C, pH 7.5 and a conductivity of 334 µS. Aquarium specimens have been maintained at 5-28°C and were very resistant to changes in oxygen levels (R. Mehrani, (pers. comm., 2000). Amir Hosin Zalaghi recorded the following parameters on 19 May 2010:-

pH 7.3, 18.0ºC, conductivity 506.0 µS/cm, TDS 255.0 mg/l, CO3 0.0 mg/l, HCO3 152.0 mg/l, Cl 35.0 mg/l, SO4 65.0 mg/l, Ca 59.1 mg/l, Mg 27.8 mg/l, K 0.0 mg/l, Th 250.0 mg/l, turbidity 0.65 NTU, COD 3.0 mg/l. BOD 0.0 mg/l, total alkalinity 101.0 , alkalinity-f 0.0, DO 7.9 mg/l, TSS 0.5 mg/l, NO3 0.5 mg/l, NO2 0.0 mg/l and Na 12.0 mg/l.  

 

 

Cave locality with R. Mehrani, 4 December 2000

 

Cave locality showing friable rock surrounds

Age and growth

Unknown although R. Mehrani (per. comm., December 2000) kept fish in aquaria for 18-24 months.

Food

Unknown but the aquarium specimens referred to above were fed Artemia, dried and fine-ground Gammarus, zooplankton and phytoplankton. Faecal contents were phytoplankton and one fish was observed to scrape the aquarium wall. Occasionally aquarium fish will swim upside down with the snout at the water surface and may be feeding on an algal film. Fish with a disc can attach to and graze on the substrate; a significantly longer intestine in such fish may be indicative of a detrital feeding habit (Sargeran et al., 2008).

Reproduction

Unknown.

Parasites and predators

None reported from Iran.

Economic importance

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in textbooks and its status as a cave fish.

Conservation

A fine of 10,000 rials (U.S.$139.94, 15 March 1978) was imposed specifically for illegal fishing of this species (Anonymous, 1977-1978), now 100,000 rials (U.S.$11.04, 7 April 2008). It is on the 1994 IUCN Red List of Threatened Animals as one of two rare fish species from Iran (see also Paracobitis smithi) and is on the 2000 IUCN Red List and subsequent ones as VU D2 (Vulnerable, acute restriction in its area of occupancy; see also Proudlove (2001)). The habitat is of unusual importance for studies on evolution in unique environments. Coad (2000a), using 18 criteria, found this species to be one of the top 4 threatened species of freshwater fishes in Iran.

B. Sandford (in litt., 1979) considered this fish to be endangered. The cave appeared to be a recently collapsed system and the network of fissures could be quite small. The main pool is at the end of a narrow cleft, overhung by a cliff of friable shale. Shale fragments fall spontaneously and the nearer end of the pool has a floor of shale fragments. Coupled with recent collecting the number of extant specimens may be quite low but this is impossible to confirm.

Local informants in December 2000 estimated that 5-6 parties visit the site each year. The number of specimens taken is unknown but an estimated 66+, possibly more than 100, have been collected in recent years (from 2000 to 2008). Eight specimens are referred to in the literature, 4 specimens were caught in 1998 (R. Mehrani, pers. comm., 2000), in the two years 1999-2000 13 specimens were collected by one party, 18 by another in December 2000 (R. Mehrani and IFRO staff, N. Najafpour, IFRO, F. Razi, Darabad Museum, Tehran and B. W. Coad), 10 specimens by Ali Ebrahimi (pers. comm, 25 January 2006), 11 by Kiavash Golzarian (pers comm., 6 April 2008), and more than 10 by others.

Four fish collected in 1998 survived 2 years in an aquarium (R. Mehrani, pers. comm., December 2000). They were fed on Artemia, zooplankton, phytoplankton and fine-ground Gammarus. Water temperature ranged from 5 to 28°C and resistance to changes in oxygen levels was high. Fish were sometimes observed to swim upside down at the water surface.

The establishment of a small park or reserve around the site and education of the local people to maintain a watch on the cave would be most useful to protect this species, and the other cave species at this site, from unauthorised collectors. A survey of the local people and the Department of the Environment files should be made to determine the numbers of visitors to this remote site.

Further work This is a small species of fish of unusual appearance and provenance and could be bred and sold as an aquarium and experimental species, providing that numbers at the site warrant removal of breeding stock. If successful, this would ensure survival of the species. Captures at present appear to be fortuitous and give no real picture of the population size; removal of more specimens would have to be carefully planned and monitored. Surveys of groundwater recharge in the area and a more thorough investigation of the cave system should be undertaken to assess the status of the habitat.

Sources

Movaghar (1973) is an additional reference, in Farsi, on this species.

Type material: See above (ZMUC P 26475, P 26476, 26477, 26478, 26480).

Iranian material: CMNFI 2007-0124, 8, 27.3-42.2 mm standard length, Lorestan, type locality as above.

Genus Kosswigobarbus
Karaman 1971

?

Much of the past literature on this genus appeared under Barbus (q.v.).

 

Kosswigobarbus kosswigi
(Ladiges, 1960)

Common names

ابوحنج (abu henej or abu hanaj= father of the hook or spine; possibly abu hanash or abu henesh, father of the snake), shebeh shirbot.

[Kosswig's barbel].

Systematics

This species was described as Cyclocheilichthys kosswigi Ladiges, 1960 from the "Batman suyu" (the holotype is possibly a female, 162.7 mm, Turkey, Siirt Province, Batman suyu (the Batman stream enters the Tigris River at 37°47.30"N, 41°00'E near Batman), April 1939, C. Kosswig). The holotype is in the Zoologischen Instituts und Zoologischen Museums der Universität Hamburg (ZMH H1148). The genus Cyclocheilichthys Bleeker, 1860 is found only in Southeast Asia.

A new and monotypic genus, Kosswigobarbus, was erected for this species by Karaman (1971) but this was synonymised with Barbus by Coad (1982f). However Bănărescu (1997) and Ekmekçi and Banarescu (1998) consider Kosswigobarbus to be valid. Borkenhagen (2005) considers that kosswigi and sublimus should be placed in Barbus (Carasobarbus) or Carasobarbus, favouring treating Carasobarbus as a subgenus until Barbus is revised thoroughly. Borkenhagen et al. (2011) include the two species in the genus Carasobarbus. Karaman (1971) distinguished the genus on the basis of the fin ray characters, a well-developed rostral flap, numerous fine pores on the head, and a large lachrymal bone. These characters are found in other "Barbus" species in Southwest Asia and the whole complex of large-scaled Barbus requires a detailed revision (see also under Carasobarbus luteus and K. sublimus).

The relationships of this species appear to lie with other Southwest Asian species formerly in the catchall genus "Barbus" which possess a compressed body, large scales with counts of 38 or less in the lateral line, a smooth dorsal fin spine, 9 or more branched dorsal fin rays and 6 anal fin branched rays such as Barbus apoensis Banister and Clarke, 1977, B. exulatus Banister and Clarke, 1977 (both in Southwest Arabia), B. chantrei (Sauvage, 1882) and B. canis Valenciennes, 1842 (both from the Levant), B. luteus (Heckel, 1843) from the Tigris-Euphrates and neighbouring basins, and B. sublimus Coad and Najafpour, 1997 from Khuzestan in the Tigris-Euphrates basin of Iran. Borkenhagen (2005) regards these species as a monophyletic group characterised by 6 branched anal fin rays, smooth and ossified last unbranched dorsal fin ray, modally 10 branched dorsal fin rays, less than 40 scales in the lateral line, medium body size, large, round to shield-shaped scales with numerous parallel radii, pharyngeal teeth usually 2,3,5-5,3,2, gill rakers short, stout and slightly curved, and barbels short.

Borkenhagen et al. (2011) found the number of nucleotide differences between individuals of this species to be surprisingly high, attributing this to rarity and small populations which give rise to genetic drift and founder effects. Both this species and K. sublimus live in small mountain streams and were considered to be less likely to migrate through lowland rivers than, for example, Carasobarbus luteus, a more widespread and generalist species.

Key characters

This species is characterised by having two pairs of thin barbels, 6 branched anal fin rays, the last unbranched dorsal fin ray strong and sharp-edged but smooth, 9-11 branched dorsal fin rays, large scales (38 or less in the lateral line), and a deep and compressed body.

Morphology

The rostral flap shows variable development, sometimes overlapping the upper lip to become visible in ventral view and other times not so well developed. Lips are thick, continuous and fleshy and there is a large median lobe to the lower lip. The mouth is small, ventral and u-shaped. The dorsal fin origin lies over or slightly in advance of the pelvic fin origin. The dorsal fin margin is strongly concave and the last unbranched dorsal fin ray is a strong spine without teeth. The caudal fin is deeply forked. The anal fin is long and may overlap the caudal fin base.

Scales are regularly arranged over the body. A low sheath of scales is found at the base of the anal and dorsal fins, being most evident anteriorly, and enclosing the anal papilla. There is a pelvic axillary scale. Anterior scale radii are few (5-11 in five scales from one specimen 126.6 mm SL) while posterior radii are numerous (35-40). There is a scaled keel or ridge before the dorsal fin as the back narrows dorsally. Pharyngeal tooth formulae 2,3,5-5,3,2, 2,3,5-4,3,2, 2,3,4-5,3,2 and 2,3,4-4,3,2. The teeth are quite small even in the largest specimens. Teeth are hooked at the tip and strongly recurved there, teeth are conical and have a small, concave to irregular or even rounded grinding surface below the tip. The fifth and most anterior tooth in the main row is small to minute in most fish and may be absent but this is not size related as both large and small specimens have or lack this tooth. The gut is elongate and coiled.

Meristic data from Iranian and other Tigris-Euphrates specimens: dorsal fin branched rays 9(5), 10(34), 11(1), mean 9.9 after 4 unbranched rays; anal fin branched rays 6(40) (not 7 as in the original description) after 3 unbranched rays; pectoral fin branched rays 15(2), 16(6), 17(4), 16.1; and pelvic fin branched rays 7(1), 8(11). Lateral line scales 29(1), 31(2), 32(1), 33(2), 34(6), 35(3), 36(1), 37(1), 38(1), 41(1) mean 34.2 (Kuru's (1975) range is 32-36); scales above the lateral line 6(7), 7(10), 8(1), mean 6.7; scales below the lateral line 5(5), 6(13), mean 5.7; scales between lateral line and pelvic fin 4(11), 5(1), mean 4.1; predorsal scale rows 11(1), 12(2), 13(5), 14(3), 15(1), mean 13.1; and caudal peduncle scales 13(2), 14(2), 15(6), 16(2), mean 14.7. Total gill rakers 10(2), 11(2), 12(3), 13(3), 14(1), mean 11.9. Pharyngeal teeth 2,3,5-5,3,2(4), 2,3,4-4,3,2(3) or 2,3,4-5,3,2(1). Total vertebrae 39(5), 40(4), mean 39.4.

Caudal peduncle length in head length 1.2-1.3, mean 1.3; caudal peduncle depth in caudal peduncle length 1.5-1.7, mean 1.6; pelvic fin length in standard length 4.6-5.4, mean 5.0; pelvic fin length in pelvic fin origin to anal fin origin distance 1.2-1.6, mean 1.3; dorsal fin spine length in head length 0.8-0.9, mean 0.9; and longest dorsal fin ray in head length 0.9-1.0, mean 0.9.

Sexual dimorphism

Sample sizes are too small to investigate accurately.

Colour

Upper flank scales are outlined by pigment, most evidently anteriorly on each scale. Fins are lightly pigmented with scattered melanophores on both rays and membranes with some concentration on dorsal fin membranes although the extent varies individually. The peritoneum is black.

Size

Reaches 19.4 cm total length (the holotype).

Distribution

Found in the Tigris-Euphrates basin of Turkey and Iran (Coad, 1982f; Coad and Najafpoiur, 1997; Abdoli, 2000). It may also occur in the Zohreh River (Gh. Izadi, pers. comm., 2001).

Zoogeography

Karaman (1971) considers that the closest relatives of this species are to be found in the Indo-Malayan region.

Habitat

This species is found in large rivers in Iran which, however, in mid-summer are more stream-like in water flow. Collections are from the plains of Khuzestan and from altitudes in excess of 1600 m in the Zagros Mountains. Temperatures in early July range from 21 to 23°C. One locality was polluted and others were cloudy or muddy. The river beds are composed of stones.

Age and growth

Unknown.

Food

The elongate gut and black peritoneum suggest a plant component to the diet but examination of two gut contents reveal insect remains including chironomid larvae.

Reproduction

Unknown.

Parasites and predators

Sohrabi and Jalali (2002) report the nematode Schulmanella petruchewskii from the liver of this species caught in the Dez River.

Economic importance

This species is too rare in Iran to be of any economic importance.

Conservation

Recommendations are difficult to make since the ecological requirements of this species are unknown. It appears to be rare but this may only be inadequate sampling techniques. Further collections in addition to the holotype have been made in southern Anatolian Turkey (Kuru, 1978-1979) but it does not seem to be common. Endangered in Turkey (Fricke et al., 2007).

Further work

Intensive field work utilising a wide variety of techniques should be directed to determining the abundance and distribution of this species. An adequate material base would then enable ecological studies to be carried out and conservation measures determined.

Sources

Some counts from Kuru (1975) on Turkish material.

Type material: See above, Cyclocheilichthys kosswigi (ZMH H1148).

Iranian material: CMNFI 1979-0275, 1, 126.6 mm standard length, Lorestan, Kashkan River drainage (33º25'N, 47º58'E); CMNFI 1979-0277, 1, 116.5 mm standard length, Lorestan, Kashkan River drainage (33º30'N, 47º59'E); CMNFI 1979-0289, 1, 103.5 mm standard length, Kermanshahan, Diyala River drainage (34º28'N, 45º52'E); CMNFI 1979-0290, 2, 120.1-122.1 mm standard length, Kermanshahan, Diyala River drainage (34º31'N, 45º35'E); CMNFI 1979-0368, ?, ? mm standard length, Khuzestan, Karkheh River (32º24'30"N, 48º09'E); uncatalogued, 1, 173.3 mm standard length, Khuzestan, Karkheh River near Shush (no other locality data); uncatalogued, 3, 140.1-179.0 mm standard length, Khuzestan (no other locality data); uncatalogued, 1, 160.0 mm standard length, Khuzestan, Karun River basin near Izeh (no other locality data).

Comparative material: BM(NH) 1974.2.22:1281, 1, 31.2 mm standard length, Iraq, Al Hadithah (34º07'N, 42º23'E); BM(NH) 1974.2.22:1292-1296, 4, 35.5-98.5 mm standard length, Iraq, Al Hadithah (34º07'N, 42º23'E).

Kosswigobarbus sublimus
(Coad and Najafpour, 1997)

Common names

None.

Systematics

The holotype is CMNFI 1995-0009, female, 113.5 mm, Iran, Khuzestan, A'la River at Pol-e Tighen, 31°23.5'N 49°53'E, 20 September 1995, B. W. Coad, N. Najafpour and party. Paratypes are CMNFI 1995-0009A, 41.9 mm, same locality as the holotype (lost in the mail while on, loan September 2005), CMNFI 1995-0010, female, 115.3 mm, A'la River, 2 km above Pol-e Tighen, 31°23.5'N 49°54'E, 20 September 1995, B. W. Coad, N. Najafpour and party, and CMNFI 1995-0011, 3 females, 90.5-98.6 mm, same locality as holotype, early December 1994, Gh. Eskanderi (one specimen lost in the mail while on loan, September 2005).

The species was named after its river of capture, the only known locality for this species. A`la means "most high" or "exalted".

Borkenhagen et al. (2011) used limited molecular data (a small number of base pairs in the cytochrome b gene) and found evidence of paraphyly of kosswigi with sublimus, indicating a recent speciation event.

Key characters

A member of the genus Barbus sensu lato characterised by the unique combination of the following characters: large scales (24-27 in the lateral line), 37-38 total vertebrae, 10-11 branched dorsal fin rays, 6 branched anal fin rays, a relatively short and smooth dorsal fin spine (spine length in head length 1.0-1.1), lower lip with a rounded median lobe and a posterior free flap, a compressed body (depth 3.3-3.5 in standard length), a short caudal peduncle (length in head length 1.5), long pelvic fins (length in standard length 4.1-4.5), and a short dorsal fin (longest dorsal fin ray in head length 1.1-1.2).

Morphology

Dorsal fin branched rays 10(5), 11(1), mean 10.2; anal fin branched rays 6(6); pectoral fin branched rays 14(1), 15(5), mean 14.8; and pelvic fin branched rays 8(6). Lateral line scales 24(1), 25(2), 26(2), 27(1), mean 25.5; scales above the lateral line 4(1), 5(5), mean 4.8; scales below the lateral line 4(3), 5(3), mean 4.5; scales between lateral line and pelvic fin 3(6); predorsal scale rows 9(5), 10(1), mean 9.2; and caudal peduncle scales 12(5). Total gill rakers 10(1), 11(1), 12(2), 15(1), mean 12.0. Total vertebrae 37(2), 38(4), mean 37.7. A specimen from the Khersan River had 39 total vertebrae and one from the Ardal River had 40 total vertebrae; both these fish being unusual in other counts too. Esmaeili et al. (2006) give the following characters for their 6 specimens from Fars: 11 branched dorsal fin rays, 6-8 anal fin branched rays, 16-18 branched pectoral fin rays, 24-28 lateral line scales, and 10-12 total gill rakers.

The body is relatively deep (depth 3.3-3.5 times in standard length) and compressed. The snout is rounded and overhangs the upper part of the thick upper lip. The extent of overlap varies individually. The lower lip is also thick but has a rounded protuberance at its centre, visible in lateral view. The protuberance is variably developed as a flap which is free posteriorly and at the rearmost sides. The posterior barbel is longer and thicker than the anterior barbel. The anus lies just anterior to the anal fin origin.

Scales are regularly arranged over the whole body, there is a pelvic axillary scale, and scales at the anterior base of the anal fin form a small sheath around the bases of the anal rays. Radii are found on the anterior and posterior fields of each scale, being most numerous posteriorly, about three times as many. Some radii extend into the lateral fields. Circuli are numerous and on the posterior field break up into bubble-like shapes.

The dorsal fin is slightly to strongly concave on its margin. The spine tapers and is thin and flexible at the tip. The dorsal fin origin lies over the pelvic fin origin. The caudal fin is deeply forked with the lower lobe more developed and with longer rays than upper lobe. The anal fin reaches or obviously passes the base of the caudal fin rays. This variation in length does not appear to be size or sex related. The posterior margin of the anal fin is straight to concave. The pelvic fin has a straight to rounded posterior margin. The pectoral fin margin is concave and in some fishes is falcate.

The gut is elongate with anterior and posterior loops. Gill rakers are short and reach to the adjacent raker when appressed. Pharyngeal teeth are rounded with a hooked tip and a flattened area below the tip. On three specimens counts were 2,3,5-4,3,2, 1,3,5-4,3,2 and 3,3,4-4,3,2.

Sexual dimorphism

Sample size of the type series is too small to document sexual dimorphism.

Colour

The overall live colour of the species is silvery with the back olive-green. Scales are outlined with dark pigment. The pectoral, pelvic, anal and caudal fins are a faintly pigmented with orange to yellow hues, most apparent when the fin is collapsed. Much of these fins is grey to hyaline. The dorsal fin is grey to hyaline. The eye is silvery with grey-brown pigment at the upper margin. The peritoneum is silvery with numerous melanophores merging to give an overall dark appearance.

In 70% ethanol the pigmentation pattern is as follows. Upper to mid-flank scales have the margins and bases pigmented with melanophores, outlining the scales. Most pigment is concentrated at the scale base giving a slight appearance of rows of spots. Larger fish are more fully pigmented so the back and upper flank then appear dark. The dorsal surface of the head is finely speckled black. The dorsal fin has dark pigment on the membranes, on the distal half or the whole fin, with less pigment on the rays. The caudal fin is mostly hyaline with dark pigment lining the rays. The pectoral and anal fins have some dark pigment lining or on the anterior rays and, in larger fish, on the membranes. The pelvic fin is hyaline. The smallest specimen has a distinct mid-caudal base spot and another spot on the back at the anterior dorsal fin base. Fins are more hyaline than in larger fish.

Size

Reaches 115.3 mm standard length. The maximum size is 15.5 cm (Iranian Fisheries Research and Training Organization Newsletter, Tehran, 18:5, 1997).

Distribution

Known from the A'la River in Khuzestan Province in the Tigris River basin and the Fahlian River in Fars (Esmaeili et al., 2006).

Zoogeography

This species is known from the A`la River, which joins with the Rud Zard (rud = river), and emerges from the foothills of the Zagros Mountains onto the Khuzestan plains where it is tributary to the Jarrahi River. The Jarrahi feeds the Shadegan Marshes and is mostly lost there. In flood times, there may be a connection through the marshes to the Karun River and thence to other large river systems in the Tigris-Euphrates basin. However, it is suspected that the ecological requirements of this species limit it to fast flowing rivers over hard substrates and the marsh system isolates it from other river systems. Collections in the Rud Zard at Rud Zard village and Bagh-e Malek on several occasions have not included the new species although the Rud Zard would appear to be a suitable habitat.

The range extension of 380 km southwest of the A`la River to the Fahlian River near Noorabad in Fars places this species in the headwaters of the Zohreh River which drains to the northern Persian Gulf. This may indicate headwater captures or possibly former interdigitating drainages on the Khuzestan plain.

Habitat

The type habitat is a cloudy river in a wide flood plain at about 800 m. The river bed is stones and pebbles. Water is led off from the river at intervals to irrigate the rice fields of the villages of Meydavud-e `Olya (31°24'N. 49°52'E) and Meydavud Pa'in (31°23'N. 49°49'E) which extend along the bank of the A`la River. This water abstraction is a potential threat to the well-being of fishes in this river system. The water demands of rice growing are large and there is little or no rain through the summer months in this area. Air temperatures in September can exceed 40°C and evaporation from the fields and the river is commensurate.

The fish were caught at the type locality in relatively fast water (0.9 m.s-1) over a one hour fishing period. In September 1995, the river was at the seasonal low water and the type locality was 10 m wide, 40 cm deep and had a discharge of ca. 2.9 m3.s-1. The water was also cloudy for the collection in December 1994 at the type locality but the river was wider and had more flow after rain. The second locality had more flow and was deeper and wider than the type locality, to about 30 m and 80 cm. Fishes were caught by electroshocker and cast-net and were difficult to catch and few in number. Other species captured were the cyprinids Luciobarbus barbulus, Tor grypus, Barilius mesopotamicus, Capoeta trutta, Cyprinion macrostomus and Garra rufa, and the sisorid catfish Glyptothorax silviae.

The Fahlian River capture site was shallow, had relatively clear water, a heterogenous bed morphology (sand, gravel, stone, pebbles, rock, etc.), and an absence of aquatic and riparian vegetation (Esmaeili et al., 2006).

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

None reported.

Economic importance

None reported.

Conservation

Known only from the type series, its conservation status is unknown. It appears to be rare and possibly restricted to areas with running water year round.

Further work

The distribution of this species should be investigated within the A'la River and neighbouring drainages and accurate estimates of its numbers and biology procured as part of a process to determine its conservation status.

Sources

Type material: See above, CMNFI 1995-0009, CMNFI 1995-0009A, CMNFI 1995-0010, CMNFI 1995-0011.

Genus Labeo
Cuvier, 1816

Labeo dero
(Hamilton, 1822)

Mashkel (= Mashkid) River basin in Pakistan on the southeastern border of Iran (Zugmayer, 1913; Mirza, 1971; 1972; 1974; 1975; 1992). No Iranian record.

Labeo gedrosicus
Zugmayer, 1912

Mashkel (= Mashkid) River basin in Pakistan on the southeastern border of Iran (Zugmayer, 1912; Mirza, 1971; 1972; 1974; 1975; 1992; Neumann, 2006). No Iranian record.

Labeo macmahoni
Zugmayer, 1912

Makran basin (Dasht River) in Pakistan on the southeastern border of Iran (Zugmayer, 1912; Mirza, 1971; 1972; 1974; 1975; Mirza and Saboohi, 1990; Neumann, 2006). Placed in a new subgenus, Tariqilabeo by Mirza and Saboohi (1990). Kullander et al. (1999) and Mirza and Arshad (2008) consider this taxon to be, questionably, a synonym of Crossocheilus diplocheilus (Heckel 1838). A syntype is in the Naturhistorisches Museum Vienna (NMW 81256). No Iranian record.

Genus Leucaspius
Heckel and Kner, 1858

The genus Leucaspius has not been recently revised in detail and its composition remains uncertain. There may be several species in Europe but only one has a wide distribution and this is found in Iran.

The genus is characterised by a moderately compressed and elongate body; an incomplete lateral line on up to about 13 scales; moderately large, easily detached scales; short dorsal and somewhat longer anal fin; belly without a keel but somewhat compressed; terminal mouth with lower jaw entering the depression of the upper; pharyngeal teeth usually in 2 rows; and gill rakers of moderate size and density.

Leucaspius delineatus
(Heckel, 1843)

Leucaspius delineatus from Wikimedia Commons.

Common names

mahi-ye riz-e noqrei or mahi-e-rize-noghreie (= small silvery fish).

[gafgaz ustuzani in Azerbaijan; ovsyanka, verkhovka, Kavkazskaya verkhovka or Caucasian verkhovka in Russian; sunbleak, white aspe, rain bleak; belica; Moderlieschen in German].

Systematics

Squalius delineatus was originally described from Wien and Mähren, Austria. The Caspian Sea basin taxon is given by Berg (1948-1949) as Leucaspius delineatus delineatus natio caucasicus Berg, 1949, described from Transcaucasia, which is distinguished by a lower average dorsal fin branched ray count (7-8 rather than 8 or rarely 9 for the typical form of Europe). This natio has no taxonomic standing but has been applied as a subspecies by some authors (Arnold and Längert, 1995).

Key characters

The large, rounded papillae around the genital opening are distinctive in females, and for both sexes the combination of an incomplete lateral line with moderately large scales is distinctive.

Morphology

Dorsal fin with 2-3, usually 3, unbranched rays followed by 7-10 branched rays (usually 8 in Europe but counts of 7 and 8 are about equally frequent in the Caucasian populations according to Berg (1948-1949) but Abdurakhmanov (1962) gives a frequency of 94% for 8 rays and only 6% for 7 rays in fish from Azerbaijan), anal fin with 3-4, usually 3, unbranched rays followed by 9-17 branched rays (10-12 in the Caucasian subspecies), pectoral fin branched rays 11-16 and pelvic fin branched rays 7-8. Lateral series scales 36-53; lateral line incomplete with 0-13 pored scales anteriorly. Scales bear few anterior and posterior radii, have few circuli, a subcentral anterior focus and are a vertical oval in shape. Gill rakers 10-17 (rarely 20, usually 13-16), reaching the second raker below when appressed. Vertebrae 36-40. Pharyngeal teeth very variable 5-5, 5-4, 4-4, 4-5, 1,5-5, 5-5,1, 1,5-4, 5-4,1, 1,4-4, 4-4,1, 1,5-5,1, 1,5-5,2, 1,4-4,1, 1,4-5,1, 1,5-4,1, 1,5-4,2, 1,4-5,2, 2,5-4,1, 2,5-4,2, 2,5-5,2 and even 2,5-6,1. The frequency of various counts varies with locality, and even whether single row counts dominate over two-rowed counts (Arnold and Längert, 1995). Teeth are hooked at the tip and slightly to strongly serrated. The belly is compressed in the mid-line between the pelvic fins and the vent but does not form a strong keel. The gut is an elongate s-shape. The chromosome number is 2n=50 (Klinkhardt et al., 1995).

Meristic values for Iranian specimens are:- branched dorsal fin rays 8(10) or 9(1); branched anal fin rays 10(4) or 11(7); branched pectoral fin rays 11(7), 12(3) or 13(1); branched pelvic fin rays 7(10) or 8(1); scales in lateral series 39(3), 40(5), 41(1), 43(1) or 45(1); total gill rakers 13 (4) or 14(7); and total vertebrae 36(1) or 37(9). One specimen showed fusions of abdominal vertebrae.

Sexual dimorphism

Females have a unique fold of skin in the shape of two, large, rounded papillae around the genital opening. The male is a little smaller than the female. Males develop prominent nuptial tubercles on the dorsal head surface, snout, on the lower jaw in three pairs and on the upper jaw in two pairs for a total of about 60 tubercles. The male genital opening is depressed.

Colour

The back is olive-green to brown and the flanks and belly silvery-white. A steel blue or bluish-green stripe begins at the rear third of the body and extends back, broadening, to the tail base. Fins are hyaline or slightly yellowish. The peritoneum is light.

Size

Attains 12 cm total length although only up to 5.6 cm total length in the Caucasian form.

Distribution

Found in western and central Europe from the Rhine and north of the Alps east to northern drainages of the Black Sea and the western and northern drainages of the Caspian Sea. Leucaspius delineatus caucasicus is found in the north Caucasus including the Black Sea parts and in Transcaucasia. In the southern Caspian Sea basin, it is found in the lower reaches of the Kura River, Imeni Kirova Bay and the Lenkoran region of Azerbaijan (Kuliev, 1989). A single specimen from Iran was collected by Mr. Akbar Nasrollazadeh near Siah Darvishan (which is at 37°22'N, 49°26'E) in Gilan on 27 May 1993. In June 1996 over 50 specimens were caught in the Anzali Mordab (= Talab) by K. Abbasi and A. Sarpanah of the Gilan Fisheries Research Centre (Iranian Fisheries Research and Training Organization Newsletter, 15:4, 1997). Also reported from the Anzali Talab by Abbasi et al. (1999) and present in the Safid River and Amirkelayeh Wetland (Nasrollazadeh, 1999; K. Abbasi, pers. comm., 2001).

Zoogeography

The Caspian shore of Iran has been surveyed in some detail during the 20th century and it is curious that this species was only discovered towards its end. It may simply have been confused with other small, silvery minnows although it should be noted that some of the surveys were carried out by Russian workers familiar with this species. It may be a recent introduction with other, commercial exotics, and therefore may not be from a Caucasian population.

Habitat

Found in still or slowly flowing water with vegetated shores in large schools. It can be found in fish ponds, ditches, gravel pits and quarries as well as natural habitats. Still water is required for reproduction. It is tolerant of a wide range of temperatures, pH and salinity depending on adaptation, e.g. temperature range of 3-32.8°C (Arnold and Längert, 1995). This small fish is found in large schools near the water surface. It may appear in small ponds without any apparent connection to other water bodies, hence the German name that has been interpreted as "Moderlieschen" or motherless. However, the German name may more correctly mean mud lover (G. H. Copp, in litt., 16 June 2004).

Age and growth

Life span is about 4-6 years with growth fairly continuous over this period.

Food

Diet comprises plankton such as cladocerans, copepods and rotifers, benthic chironomids, flying insects which land on the water surface, and also some algae and detritus.

Reproduction

There is often a spawning migration against the water flow (up to 2-3 m/sec) to new waters. Eggs are laid in strings which are wound spirally around plants by the female, aided by the fold of skin around the genital opening. They may also be laid in a disc-shaped patch on any flat surface. Several spawnings occur over a few weeks in March to September in Europe. The eggs are guarded and fanned by the male who covers them with a bacteriostatic dermal mucus. Up to 485 eggs are found in females and have diameters up to 0.5 mm in Azerbaijan, up to 3500 eggs and 1.5 mm in Europe. Maximum egg production over two seasons is about 500-600 (Abdurakhmanov, 1962). Clutch sizes are about 50-350 eggs (Arnold and Längert, 1995). A minimum temperature of 18°C is required for reproduction.

Parasites and predators

None reported from Iran but eaten by a wide variety of other fishes in Europe and numerous parasites reported (Arnold and Längert, 1995).

Economic importance

The scales have been used in the production of artificial pearls as with Alburnus alburnus (a relative of A. hohenackeri). It has also been used in aquaria and garden ponds and as bait by anglers.

Conservation

Lelek (1987) classifies this species as rare to vulnerable in Europe. Kiabi et al. (1999) consider this species to be conservation dependent in the south Caspian Sea basin according to IUCN criteria. Criteria include few in numbers, habitat destruction, limited range (less than 25% of water bodies), absent in other water bodies in Iran, present outside the Caspian Sea basin.

Further work

More specimens from Iranian waters need to be examined to determine if this species belongs to a distinct subspecies or is an exotic population of the European type subspecies. Biology of the Caucasian subspecies is unknown.

Sources

Arnold and Längert (1995) summarise biology of European populations in detail.

Iranian material: Uncatalogued material, 11, 31.2-37.5 mm standard length, Gilan, swamp near Hendeh Khaleh (37º23'N, 49º28'E).

Genus Luciobarbus
Heckel, 1843

?

Much of the past literature on this genus appeared under Barbus (q.v.). Valiallahi (2000) describes "Barbus persicus", a new species, in his thesis but this has not been formally described. The name may be preoccupied by Bertinius longiceps persicus Karaman, 1971.

Seyed Mortezaei et al. (2008) give details of 11 myxozoan and protozoan parasites of barboid fishes from rivers, reservoirs and marshes in Khuzestan.

Luciobarbus barbulus
(Heckel, 1849)

Common names

lab pahn (= broad lip), برزم (berzem or barzam); berzem lab pahn in Khuzestan and Iraq to distinguish it from B. pectoralis; boz mahi (= goat fish) or سس ماهي (= sos or sas mahi in the Dalaki and Shapur river basins); dolenj.

[abu-barattum (= owner or father of lips), abu baratem, abu bratum or nabbash in Arabic; Orontes barbel].

Systematics

Howes (1987) places this species in Barbus sensu stricto. Karaman (1971) places this species in the synonymy of Barbus rajanorum but other authorities consider it to be Luciobarbus pectoralis (q.v.). Almaça (1983) placed this species as a subspecies of Barbus mystaceus but later (1984a, 1984b, 1986, 1991) retained barbulus as a full species, known only from the Levant, despite Heckel's record from both the Qarah Aqaj (= Mand) of Fars, Iran and the Quwayq (= Kueik) River of the Levant. I retain it as a species under Heckel's name for this taxon until the systematics of this and related species can be worked out as indicated above. It is separated from mystaceus according to Almaça (1983) by having thinner lips, shorter barbels, the last unbranched dorsal ray weaker and shorter, more dense denticles spread over a shorter length of ray, higher anal fin, gill rakers less numerous and the upper dorsal profile is rectilinear and oblique to the back.

The type locality of Barbus Barbulus is the "Fluss Kara-Agatsch....bei dem Dorfe Geré" (= Qarah Aqaj or Mand River, Fars; possibly near Kereft, 29°01'N, 52°52'E) and presumably the "Kueik bei Aleppo" (Heckel, 1846-1849b). J. Valiallahi, pers. comm., 2001 and Edmondson and Lack (2006) suggest Jereh at 29°15'N, 51°58'E but this is in the Hilleh River drainage, a Dalaki River tributary. In addition, "Geré" takes a hard G in German, not a J. There may be some confusion of names and rivers here.

A possible syntype of barbulus from the Qarah Aqaj was located by Almaça (1983, 1986) in the Naturhistorisches Museum Wien (NMW 53957) and seen by me but is in too poor condition to be of much value, being mostly bones. Another syntype is listed as NMW 6596 and measures 119.3 mm standard length. In 1997, this was the only syntype recognised and is possibly the same as NMW 53957 re-numbered as the latter was not located in 2002. The catalogue in Vienna lists only 1 fish, while Heckel's description refers to several fish. NMW 6596 is mostly bones and is dried. The fleshy lip fold of the original description could not be discerned, teeth are missing and the dorsal fin is broken off short.

"Syntypes" of mystaceus are in the Naturhistorisches Museum Wien from Mosul on the Tigris River (NMW 16472 (1 specimen), NMW 50394 (2), NMW 54384 (2)) and NMW 54385 (2) but note that authors such as Karaman (1971) and Almaça (1983, 1991) refer the species description to Heckel (1843) in error. These were not marked as being syntypes as observed on a 1997 visit to Vienna.

Key characters

This species is characterised by having two pairs of barbels, a serrated and very strong dorsal fin spine similar to that in Capoeta trutta in its proportions relative to the body, usually 8 dorsal fin rays (never 10), fleshy lips, and 47 or more lateral line scales.

Morphology

The inferior mouth is moderate in size, with moderate to thick lips and with or without a median lower lip lobe. Some fish have very thick lips so a central lobe is apparent. Some show such a degree of lip development as to appear almost abnormal while fish of similar size or larger lack this hypertrophy. In the latter case, the anterior head may be bluntly rounded and foreshortened rather than having an almost straight upper margin tapering to a pointed end. Barbels are relatively thin, occasionally quite thick. The anterior barbel does not extend past the anterior eye margin level and the posterior one not past the posterior eye margin in all sizes of fish. Rarely the anterior barbel extends to mid-eye level and the posterior one almost to the anterior operculum margin.

Dorsal fin with 4 unbranched and 8-9, usually 8, branched rays, anal fin with 3 unbranched and 5 branched rays. The last unbranched dorsal fin ray is usually very strong with a moderate density of denticles extending along much of the ray but its strength is variably developed. Pectoral fin branched rays 17-19, pelvic fin branched rays 8-9, usually 8. Lateral line scales 47-59. Scale focus subcentral anterior, many fine circuli, and numerous radii on all fields, curved in the lateral fields. A pelvic axillary scale is present but not strongly developed or apparent. Gill rakers 14-24, reaching the second raker when appressed. The interior raker surface may be covered with spinules, the internal base is heavily tubercular and the tips may become club-shaped. Pharyngeal teeth 1 or 2,3,4 or 5-5 or 4,3,2 or 1, hooked at the tip but spoon-like below with the fourth tooth of the inner row molariform, with or without a blunt projection (hooked in small fish) and much larger than the third, and the fifth tooth very small and rounded and sometimes absent apparently independent of size. The gut is elongate and complexly coiled with one anterior and 3 posterior loops. Total vertebrae 44 (Howes, 1987).

Meristics in Iranian fish are as follows: dorsal fin branched rays 8(26); anal fin branched rays 5(26); pectoral fin branched rays 17(6), 18(15), or 19(5); pelvic fin branched rays 8(25) or 9(1); lateral line scales 47(1), 48(3), 49(4), 50(4), 51(2), 52(5), 53(4), 54(1), 56(1) or 57(1); total gill rakers 15(1), 17(1), 18(4), 19(3), 20(6), 21(5), 22(2), 23(3) or 24(1); pharyngeal teeth 2,3,5-5,3,2(18), 2,3,5-4,3,2(3), 2,3,4-5,3,2(3), 2,3,5-5,3,1(1) or 2,3,4-4,3,2(1); total vertebrae ?.

Sexual dimorphism

Unknown.

Colour

The back and upper flank are brownish, the lower flank yellowish and the belly whitish. Upper flank scales are outlined with pigment, and the anterior edge of the dorsal fin and the caudal fin margin are black in preserved fish. Small fish have a few spots on the upper to mid-flank or may be profusely speckled in preservative.

Small live fish are silvery overall and have anal and caudal fins orange to bright red, especially the lower caudal fin lobe. The dorsal fin is grey and the pectoral and pelvic fins yellowish. The operculum is greenish. The lower flank is greenish-golden and the upper flank brown to grey. Large specimens are silvery with clear fins. The belly in small and large fish is white and the back grey or green to brown. The iris is silvery. The peritoneum is black.

Size

Reaches 62 cm total length (Atabak Mahjoor Azad, pers. comm., 16 June 2008); J. Valiallahi (pers. comm., 2001) believes this species reaches 1.5 m and 90 kg in the Zagros rivers of western Iran.

Distribution

Found in the Tigris-Euphrates basin, the Orontes River and the Quwayq River. In Iran it is found in the Tigris River basin (Abdoli, 2000; Ghorbani Chafi, 2000), in the Gulf basin from the Zohreh River and from the Shapur and Dalaki rivers (Gh. Izadpanahi, pers. comm., 1995), the lower Mand River (M. Rabbaniha, pers. comm., 1995), the Helleh, Dozgah, Dasht-e Palang (and its tributary the Shur) (Abdoli, 2000), in the Kor River basin (Abdoli, 2000), although not confirmed by specimens seen by me, and possibly in the Hormuz basin.

Zoogeography

Almaça (1991) believes that this species originated from a colonisation wave from South Europe.

Habitat

van den Eelaart (1954) records this species from rivers in Iraq, moving into lakes and marshes on the floods but never far from rivers.

Age and growth

Al-Rudainy (2008) gives sexual maturity at 2.8 years, 31.5 cm length and 305 g in Iraq. Hashemi et al. (2010) examined fish from the southern Karun River in Iran and found a size range of 20-94 cm and 52-4675 g, growth was isometric, and growth and mortality parameters were L = 132.9, K = 0.17, t0 = -0.66, M = 0.33, F = 1.04, Z = 2.72 and E = 0.76. Relative yield per recruitment (Y'/R) was 0.021, relative biomass per recruitment (B'/R) was 0.25, exploitation ratio maximum sustainable yield (Emax) was 0.42, precautionary average target (Fopt) was 0.16 year-1, and limit (Flimit) as 0.21 year-1. The stock was overfished and fishing regulations are required.

Food

Diet is benthic organisms including insects. Large plant remains and detritus are also present in gut contents of Iranian fish.

Reproduction

Al-Habbib et al. (1986) report spawning during July and August in fish from the Tigris River at Mosul, Iraq. Al-Rudainy (2008) cites a major spawning in April and a lesser one in October, with eggs deposited on gravel beds in fast water. Absolute fecundity is about 100,000 eggs. Reproduction in Iran has not been studied.

Parasites and predators

Peyghan et al. (2001) record the cestode Bothriocephalus sp. and the nematode Rhabdocona sp. from fish from Khorramabad rivers. Masoumian et al. (2008) recorded the myxosporeans Myxobolus karuni and M. persicus from gills of fish captured in the Karun and Karkeheh rivers and Shadegan Marsh.

Economic importance

This species is a preferred catch of anglers at Ahvaz in Khuzestan, second only to shirbot (Tor grypus). Peyghan et al. (2001) report that is is an economically important species with a good market value in the Khorramabad region.

Conservation

The population numbers of this species have not been well-studied nor has its distribution been well-documented. Since it does appear on fish markets in Khuzestan, is a large species and its habitats are under threat, it may require protection. Endangered in Turkey (Fricke et al., 2007).

Further work

The biology, distribution and population numbers of this species need investigation in Iranian waters.

Sources

Type material: See above, Barbus barbulus (NMW 6596), and note comments.

Iranian material: CMNFI 1979-0024, 1, 128.7 mm standard length, Fars, neighbourhood of Shiraz (no other locality data); CMNFI 1979-0109, 2, 91.1-91.6 mm standard length, Fars, Mand River at Shahr-e Khafr (28º56'N, 53º14'E); CMNFI 1979-0135, 1, 215.4 mm standard length, Fars, tributary to Mand River (28º08'N, 53º10'E); CMNFI 1979-0271, 1, 61.8 mm standard length, Lorestan, Kashkan River drainage (33º39'N, 48º32'30"E); CMNFI 1979-0290, 1, 139.1 mm standard length, Kermanshahan, Diyala River drainage in Qasr-e Shirin (34º31'N, 45º35'E); CMNFI 1979-0293, 1, 210.8 mm standard length, Fars, Mand River at Kavar (29º11'N, 52º41'E); CMNFI 1979-0349, 1, 126.0 mm standard length, Fars, Mand River at Kavar (29º11'N, 52º41'E); CMNFI 1979-0393, 1, 112.1 mm standard length, Khuzestan, Jarrahi River drainage (31º18'N, 49º37'E); CMNFI 1979-0497, 2, 117.4-134.4 mm standard length, Fars, Mand River at Band-e Bahman (29º11'N, 52º40'E); CMNFI 1980-0907, 1, ? mm standard length, Iran (no other locality data); CMNFI 1991-0153, 1, 230.0 mm standard length, Khuzestan, Zohreh River (no other locality data); CMNFI 2007-0109, 3, 85.1-138.7 mm standard length, Kordestan, Qeshlaq River basin south of Sanandaj (ca. 35º16'N, ca. 47º01'E); CMNFI 2007-0110, 1, 191.1 mm standard length, Kordestan, Yuzidar River basin (ca. 35º05'N, ca. 46º56'E); CMNFI 2007-0111, 1, 153.0 mm standard length, Kermanshahan, Alvand River near Sar-e Pol-e Zahab (ca. 34º36'N, ca. 45º56'E); CMNFI 2007-0113, 2, 123.9-139.6 mm standard length, Kermanshahan, Qareh Su tributary northwest of Kermanshah (ca. 34º25'N, ca. 47º01'E); CMNFI 2007-0117, 4, 43.4-155.5 mm standard length, Kermanshahan, Gav Masiab near Sahneh (ca. 34º24'N, ca. 47º40'E); uncatalogued, 1, 60.2 mm standard length, Khuzestan, Rud Zard at Rud Zard (31º22'N, 49º43'E).

Comparative material: BM(NH) 1920.3.3:23-30, 9, 80.2-98.9 mm standard length, (); BM(NH) 1931.12.21:4, 172.5 mm standard length, (); BM(NH) 1971.4.2:5, 1, 140.3 mm standard length, Iraq, Tigris near Mosul (36º20'N, 43º08'E); BM(NH) 1972.3.16:2, 69.4 mm standard length, Iraq, 10 km northwest Qala Dize (); BM(NH) 1974.2.22:1270, 174.6 mm standard length, (); BM(NH) 1974.2.22:1271-1272, 2, 91.9-210.2 mm standard length, (); BM(NH) 1974.2.22:1273-1274, 58.4-62.0 mm standard length, (); BM(NH) 1974.2.22:1275-1277, 3, 182.4-201.0 mm standard length, (); BM(NH) 1974.2.22:1278, 81.9 mm standard length, (); BM(NH) 1974.2.22:1289, 173.3 mm standard length, ();

Luciobarbus brachycephalus
(Kessler, 1872)

Common names

zardek, زرده پر (= zardehpar), سس ماهي (= sos or sas mahi), سس ماهي خزري (sas mahi khazari), sassmahi-ye Daryaye-Khazar.

[xazar sirbiti or shirbit in Azerbaijan; Kaspiiskii usach or Caspian barbel and korotkogolovyi ustach or short-headed barbel in Russian; Aral barbel; short-headed barbel].

Systematics

Barbus brachycephalus was originally described from the Syr Darya in Uzbekistan.

Howes (1987) considers the generic placement of this species to be problematical. It has slender barbels, 7 branched dorsal fin rays and the cranium is broad and flat, all characters at odds with Barbus sensu stricto. Doadrio (1990) places it in the subgenus Luciobarbus Heckel, 1843 based on a series of 4 synapomorphic osteological characters, namely the exoccipital contacts the pterotic "largely" (sic, probably broadly), high medial process of the urohyal, narrow exoccipital apophysis of the pterotic, and wide 4th and 5th infraorbitals.

Barbus obtusirostris (non Valenciennes, 1842) Jakovlev, 1870 (nomen praeoccupatum), described from the Volga River delta, Russia, is a synonym.

A possible syntype of B. brachycephalus from the Aral Sea is in the Naturhistorisches Museum Wien (NMW 53971) (Almaça, 1986). The NMW card index lists this fish plus 2 fish in NMW 53972 and 1 fish in NMW 53973 as syntypes. Syntypes in St. Petersburg, Russia are lost (Bogutskaya in Bănărescu and Bogutskaya, 2003). Syntypes of Barbus brachycephalus caspius are in the Zoological Institute, Russian Academy of Sciences, St. Petersburg under 2892 (8 fish), Transcaucasia, 3895 (8), Lenkoran, 9076 (22), 9085 (10), 9109(2), 9117(11), 9118(1), 9124(8), 9128(9), all from the lower Aras River and Lenkoran, 17042(2), 17043(1), 17044(1), all from the Bank Fishery along the lower Kura River. Syntypes under 10619 are apparently lost and a fish under 9108 is actually a Luciobarbus capito (Bănărescu and Bogutskaya, 2003).

The Caspian Sea basin subspecies is Luciobarbus brachycephalus caspius (Berg, 1914), described originally from the Caspian Sea basin (Eschmeyer et al., 1996). Karaman (1971), however, considers differences with the type subspecies of the Aral Sea basin to be minor and not worthy of subspecific recognition. Differences are in body proportions and the Caspian barbel has a smaller eye, lower dorsal fin, less deep body and head, longer pectoral-pelvic distance, shorter pelvic-anal distance, and dorsal fin further back than in the Aral barbel (Berg, 1948-1949). Fricke et al. (2007) list this taxon as a full species but also have brachycephalus in the same system in Turkey (Kura-Aras).

Key characters

The 7 branched dorsal fin rays and the predorsal distance shorter than the postdorsal distance distinguishes this species from B. capito, and colour pattern distinguishes it from B. lacerta, the other Caspian Sea barbels. ?and mursa

Morphology

The mouth is moderate in size and subterminal. Lips are thin to moderate, without a median lobe on the lower lip, and barbels are of moderate thickness. The anterior barbels can reach the level of the posterior eye margin and the posterior barbels reach or pass the preopercle level but barbel lengths show marked individual variation.

Dorsal fin with 3-5, usually 4, unbranched and 6-8, usually 7, branched rays, anal fin with 2-3, usually 3, unbranched and 5-6, usually 5, branched rays. Pectoral fin branched rays 14-17 and pelvic fin branched rays 7-8, usually 8. The dorsal fin denticles on the last unbranched ray are usually moderate in number, but may be lost in very large adults, are usually well-developed and extend along four-fifths of the ray (Karaman, 1971). This ray is very strong. Lateral line scales 62-90, commonly 65-77. Scales are elongate with a central focus and few anterior and posterior radii in young fish. There is a pelvic axillary scale. Gill rakers 16-25, short and reaching the one below when appressed. Pharyngeal teeth 2,3,5-5,3,2, hooked at the tip with the fourth tooth of the inner row large and blunt and the first three spatulate, rarely 2,3,4-5,3,2 or 2,3,4-4,3,2. Total vertebrae 45-50, usually 46-49, mode 48. The gut is coiled anteriorly. The chromosome number is 2n=100 (Klinkhardt et al., 1995).

Iranian specimens have the following meristics: branched dorsal fin rays 7(3), branched anal rays 5(3), branched pectoral fin rays 16(1) or 17(2), and branched pelvic fin rays 8(3). Lateral line scales 69(2) or 71(1). Total gill rakers 18(1) or 19(2). Pharyngeal teeth 2,3,5-5,3,2(2) or 2,3,4-5,3,2(1). Total vertebrae ?.

Sexual dimorphism

Abdurakhmanov (1962) reports on fish from the Kura River basin where males have a longer dorsal fin base and females have a greater maximum body depth, width and girth. Bogutskaya in Bănărescu and Bogutskaya (2003) report that males have a shorter head and longer unpaired fins; nuptial tubercles and colouration are absent.

Colour

The back is dark green, flanks and belly lighter, and the two areas may contrast as in Luciobarbus capito. No dark spots on the body. Fins greyish. Peritoneum brown.

Size

Reaches 22.5 kg (Robins et al., 1991) and 1.2 m.

Distribution

Found in the Caspian and Aral seas and their tributaries. In Iran, it was formerly known from the Anzali Mordab but is probably no longer present (Holčík and Oláh, 1992; but see below) and it was listed as rare in the Safid Rud (Derzhavin, 1934). Nedoshivin and Iljin (1929) and Nevraev (1929) recorded it from the Gorgan, Astrabad and Enzeli (= Anzali) regions. Recent works place it in rivers from the Astara to the Neka and Gorgan Bay peninsula, in the Anzali Mordab, and along the whole Caspian Sea coast but these are summaries of past and present distributions (Riazi, 1996; Abbasi et al., 1999; Kiabi et al., 1999; Abdoli, 2000; Abdoli and Naderi, 2009). This species is now very rare in the Caspian Sea basin of Iran, with only a couple of specimens found in a recent survey (M. Ramin, pers. comm., 2000).

This species is also recorded from the Karakum Canal and Kopetdag Reservoir in Turkmenistan (Shakirova and Sukhanova, 1994; Sal'nikov, 1995) and may eventually reach Iranian waters in the Tedzhen (= Hari) River basin.

Zoogeography

Almaça (1984b) considers this species to be a Sarmatian Sea remnant, a Neogene brackish-water basin.

Habitat

Enters rivers to spawn but does not ascend as high as Luciobarbus capito. It prefers deep sections of rivers with stony and gravel bottoms. In the Caspian Sea it may be found at 13-25 m depth. On the Kura River in Azerbaijan there is a spring run and one in August-September. The spring run begins in March and lasts about 50 days; the summer run starts after a short interruption and lasts about 190 days. The water temperature at the start of the spring run is 6.7-11.0°C but the most intensive migration is in summer at 25.2-27.2°C (Bogutskaya in Bănărescu and Bogutskaya, 2003). Spring run fish spawn in the same year. This species has been recorded at depths of 11.0-11.9 m in the Iranian Caspian Sea (Knipovich, 1921). Riazi (1996) reports that this species migrates into the Siah-Keshim Protected Region of the Anzali Mordab. Jolodar and Abdoli (2004) state that it is more abundant in Gilan than in Mazandaran coastal waters.

Young females usually enter the sea immediately but males may remain in fresh water for 3-5 years. Spawners return to the sea.

Age and growth

Most fish examined by Razivi et al. (1972) from commercial catches in Iran were 2-7 years old, 38.0-69.0 cm long and weighed 698-4658 g. Low recruitment is attributed to poor spawning success, a result of water abstraction during its spawning season. Sexual maturity is attained at 6-8 years. Holčík and Oláh (1992) note that the Anzali region catches are dominated by 3-5 year old fish, 38-71 cm fork length, with rapid growth and a weight of 2 kg attained during the fifth year of life. Abdurakhmanov (1962) gives a maximum life span of 13 years in Azerbaijan. Females live longer than males which only reach 10 years (Bogutskaya in Bănărescu and Bogutskaya, 2003).

Food

No detailed literature reports but gut contents of small specimens from Iran contain crustaceans, and insects such as, curiously, ants, thrips and mosquitos. This fish evidently feeds on insects taken at the surface and is reported as leaping out of the water to take flying insects (Bogutskaya in Bănărescu and Bogutskaya, 2003). Mayflies and caddisflies are also taken and gut contents includes detritus. Crustaceans are the main food taken in the Caspian Sea (Abdurakhmanov, 1962) but molluscs are also recorded as well as small fish.

Reproduction

This barbel spawns in swift streams over pebbles or sand during July and August in Iran and the eggs attach to rocks (Razivi et al., 1972). Holčík and Oláh (1992) and Makeeva and Pavlov (2000) state that eggs are semipelagic, hatching as they drift downstream over 2 days at 25°C. Fry are carried downstream. Up to 1,259,000 bright-yellow eggs are produced of 1.4 mm diameter and the spawning season on the Kura River begins at the end of April, peaks in June and ends at the end of August. Favoured temperatures are 20-23°C (Abdurakhmanov, 1962). First spawning is at 5-7 years of age with females taking a year longer to mature than males (Bogutskaya in Bănărescu and Bogutskaya, 2003).

Parasites and predators

Molnár and Jalali (1992) record the monogenean Dactylogyrus affinis from this species in the Safid River. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Lamproglena polchella on this species.

Economic importance

This species is caught as a food fish in Iran. Nevraev (1929) records catches of 37 to 962 individuals from the Anzali region for the years 1914-1915 to 1917-1918. It was abundant in the Anzali Mordab with total catches for Iran of 54.6 t and 32.9 t in 1969/70 and 1970/71 (28.7 t and 14.4 t for the Anzali region alone) but few fish are captured now (Holčík and Oláh, 1992) (note that these figures were taken from Appendix 11, on page 10 they are reversed). They are caught in rogas (outflowing rivers from the Anzali Mordab) and inflowing rivers of the mordab (lagoon) in late winter and early spring. On the Kura River of Azerbaijan average weight in catches was 5.6 kg for females and 3.5 kg for males and the catch from 1920-1944 varied from 0.2 to 3.6 thousand centners.

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use as food and in aquaria.

Conservation

Vulnerable in Turkey (Fricke et al., 2007). Stocks of this species have declined because of poor habitat for spawning and the construction of dams and weirs which restricted access to spawning grounds. Water abstraction for irrigation during the summer spawning season would have to be balanced against the requirements of the fish. Larvae of spring spawners are lost when they enter irrigation channels and become stranded in fields (Razivi et al., 1972).

Once known from the Anzali Mordab, it is now absent to rare there and apparently replaced by Luciobarbus capito (Holčík and Oláh, 1992). Kiabi et al. (1999) consider this species to be critically endangered in the south Caspian Sea basin according to IUCN criteria. Criteria include commercial fishing, sport fishing, few in number, habitat destruction, limited range (less than 25% of water bodies), absent in other water bodies in Iran, and absent outside the Caspian sea basin.

This species is regarded as critically endangered through illegal overfishing, pollutants and the destruction of breeding and nursery grounds. Only 2 specimens were caught in the 3 years prior to 2000 during a study of "Barbus" species in Iran. Additionally, during the 6 month beach seine fishing season (October to April) for the years 1998 and 1999 along the Caspian shore, no specimens were caught in 138 beach seines used 51,000 times (M. Ramin, pers comm., 2000).

Further work

Detailed surveys, perhaps returning captures alive, need to be carried out to monitor the status of this species in Iran.

Sources

Type material: ?

Iranian material: CMNFI 1970-0553, 2, ? mm standard length, Gilan, Sowsar Roga (37º27'N, 49º30'E); CMNFI 1980-0120, 1, 115.3 mm standard length, Mazandaran, Babol River at Babol Sar (36º43'N, 52º39'E);

Luciobarbus capito
(Güldenstaedt, 1773)

L. capito (above) and L. mursa (below) Aras River, 2 October 1994
Courtesy of Asghar Abdoli

Common names

usach bulatmai, usach chanari; zardi, zardek, zardak, زرده پر (= zardehpar), zard pareh, اورنج (oranj, orenj, orenge or ourange, possibly from the yellowish fin colour)سس ماهي (= sos, sas or sass mahi), pulad mahi (= steel fish from body colour), ses mahi bozorg (= big ? fish, ses being a word of unknown meaning).

[zardapar, shirbit, yastibas zardapar for natio platycephalus, all in Azerbaijan; tchanari in Georgian; bulatmai in Turkish; usach (or usatch) bulatmai and usach chanari in Russian]. Bulatmai is derived from Farsi, bulat = pulad or steel, mai = mahi or fish in reference to the colour on the upper flank].

Systematics

Cyprinus capito was originally described from the Kura River, Transcaucasia. No types are extant.

Howes (1987) places this species in Barbus sensu stricto. Doadrio (1990) places it in the subgenus Luciobarbus Heckel, 1843 based on a series of osteological characters (listed under Luciobarbus brachycephalus).

Cyprinus bulatmai Hablizl, 1783 (after Berg (1948-1949; Rainboth (1981) has Gmelin, 1774 as the author while Eschmeyer et al. (1996) have Gmelin, 1784 (originally described from Anzali, Iran), Cyprinus chalybatus Pallas, 1814 (originally described from Anzali, Iran), Cyprinus mystaceus Pallas, 1814 (partim, from Tiflis), Barbus conocephalus Kessler, 1872 described from the Zeravshan River, Uzbekistan, Barbus lacertoides Kessler, 1872 described from the Syr-Darya in the neighbourhood of Khodzhent (= Leninabad), Tajikistan, Barbus capito var. tiflissica Kamenskii, 1899 described from the Kura River at Tiflis (= Tbilisi), Georgia, and Barbus bilkewitschi Bulgakov, 1923 (originally described from the "Atrek", i.e. the Atrak River in Turkmenistan on the northeastern border of Iran; also spelt bilkewitchi on page 236 in Bulgakov but bilkewitschi on the plate), are synonyms. Barbus capito serratus Sokolinskii, 1927 is a subspecies from the southern Caspian Sea and Barbus capito platycephalus Abdurakhmanov, 1960 is a subspecies or a natio in the lower Kura River basin (see Abdurakhmanov (1962) for further details). Berg (1948-1949) and Karaman (1971) consider Barbus capito serratus to be a synonym of B. c. capito.

Bianco and Banarescu (1982) record this species from the Hablehrud and the Kul River basin at Darab in Persian Gulf drainages. The 2 specimens have 52 lateral line scales, 8 branched dorsal fin rays and 18-19 gill rakers. They acknowledge that these 2 fish have fewer scales than L. capito from the Caspian Sea basin but believe they may represent a new subspecies. These fish are presumed to be misidentifications as L. capito is restricted to the Caspian Sea basin.

Laloei et al. (2003) using the mitochondrial cytochrome-b gene found no separable populations of this species in 60 samples from the Iranian Caspian Sea coast and rivers.

Key characters

The 8 branched dorsal fin rays and the predorsal distance considerably longer than the postdorsal distance distinguishes this species from L. brachycephalus, and colour pattern distinguishes it from Barbus lacerta, the other Caspian Sea barbels.  ?mursa

Morphology

There is a rounded keel on the back in front of the dorsal fin. The mouth is moderate in size, inferior and horseshoe-shaped. Lips are fleshy and well-developed with tubercles but there is no free median lobe on the lower lip. Barbels can be the most developed in thickness in this species among the Luciobarbus considered here but this can vary. The anterior barbel extends back between the anterior eye margin level and its middle and the posterior barbel extends to the posterior eye margin level or almost to the preopercle in young and some adults.

Dorsal fin with 3-5, usually 4-5, unbranched rays followed by 7-9, usually 8, branched rays and anal fin with 2-4, usually 3, unbranched and 5-6, usually 5, branched rays. Pectoral fin branched rays 15-19 and pelvic fin branched rays 7-9. The dorsal fin denticles on the last unbranched ray may be lost in very large adults but are evident for two-thirds or more of the spine length in most fish (Karaman, 1971; Almaça, 1981). The last unbranched ray is moderately strong and the denticles are of moderate density along it. Lateral line scales 51-72, usually 60-66 (Karaman (1971) gives 36-70 but he includes 8 subspecies over a wide range within his definition of the species). There is no obvious pelvic axillary scale although scales in this region are elongate. The scale focus is slightly subcentral anterior, there are numerous fine circuli, and there are radii on all fields with those on the lateral fields few and often curved. Gill rakers 12-19, rarely to 22, increasing in number with the size of the fish, reaching the one below or slightly further when appressed, rounded and knobbed tip, and a large internal rounded extension. Pharyngeal teeth usually 2,3,5-5,3,2 with minor variants, hooked and spoon-like below with the depression below the crown filled in, the fourth one in the inner row the largest and pointed or blunt and rounded, the fifth smaller and blunt. The gut is long and complexly coiled with several anterior and posterior loops. Total vertebrae 42-45 (Howes, 1987), 45-47 (Elanidze, 1983), 43-49 (Bogutskaya, Bănărescu and Almaça in Bănărescu and Bogutskaya, 2003). Chromosome number 2n=100, NF=172 (Pourali Darestani et al., 2006).

Iranian fish have the following meristics: branched dorsal fin rays 7(1) or 8(49), anal fin branched rays 5(50), pectoral fin branched rays 16(3), 17(27), 18(18) or 19(2), pelvic fin rays 7(2) or 8(48); lateral line scales 53(3), 54(4), 55(7), 56(10), 57(6), 58(7), 59(7), 60(5) or 61(1); total gill rakers 13(3), 14(15), 15(18), 16(9), 17(4) or 18(1); pharyngeal teeth 2,3,5-5,3,2(33), 2,3,5-5,3,1(1), 2,3,4-5,3,2(2), 2,3,5-4,3,2(1), 2,3,5-5,2,1(1), 2,2,5-5,3,2(1); and total vertebrae ?

Sexual dimorphism

Unknown, apparently no spawning colouration or breeding tubercles.

Colour

The upper flank and head are steel-grey (hence bulat mahi) and the lower flank and belly are a strongly contrasting pale yellow or pearly-white. Occasionally fish with a uniform coloration are found and preserved material may be uniform. The steel-grey upper flank may be comprised of dark scale margins surrounding a silvery-grey scale centre. The lateral line may be darkly pigmented. Spots may occur individually on the body. The iris is silvery with a grey exterior ring and a very narrow interior golden ring. Barbels are white with grey on the inner surface. The dorsal fin is greyish and may have some dark grey spots. The caudal fin has a greyish or yellowish or slightly orange upper lobe, sometimes with faint dark grey spots, a more strongly coloured and larger yellow-orange to canary-yellow lower lobe and pink margins. The pectoral fin is whitish with a little or considerable amount of pink or yellow. The pelvic and anal fins are canary-yellow to orange with a white margin. Young fish may be darkly speckled and mottled on the mid and upper flank rather like Barbus lacerta. Peritoneum dark brown.

Size

Reaches 1.05 m and 15 kg in literature reports. A specimen from the Sardabrud was 85 cm and 5.5 kg (A. Abdoli, pers. comm., 1995).

Distribution

Found in the basins of the Black, Caspian and Aral seas. Karaman (1971) gives a distribution from the Iberian Peninsula and North Africa to Southwest Asia but he includes 8 subspecies within his definition of "Barbus" capito.

In Iran, this species is found in the Caspian Sea basin, in rivers from the Aras to the Atrak includng the Gorgan, Tajan, Babol, Haraz, Sardab, Pol-e Rud and Safid rivers,the Anzali Mordab, the Qezel Owzan and Shahrud in the upper Safid River basin, and the along the sea coast (Derzhavin, 1934; Bianco and Banarescu, 1982; Almaca 1984a; Aliev et al., 1988; Holčík and Oláh, 1992; Kiabi et al., 1994; Abbasi et al., 1999; Kiabi et al., 1999; Abdoli, 2000; Abdoli and Naderi, 2009). Valiallahi (2000) considers this species to be present in western Iran, in the Tigris River basin.

Luciobarbus capito conocephalus (Kessler, 1872) is reported from the Karakum Canal, Kopetdag Reservoir and Uzboi lakes in Turkmenistan (Shakirova and Sukhanova, 1994; Sal'nikov, 1995) and may eventually be recorded from the Tedzhen River and Caspian Sea basins in Iran.

Zoogeography

Almaça (1984b) considers this species to be a Sarmatian Sea remnant, a Neogene brackish-water basin, and related to Euro-Mediterranean "Barbus".

Habitat

This species avoids muddy bottoms (Solak, 1977) although Bogutskaya, Bănărescu and Almaça in Bănărescu and Bogutskaya (2003) report that it prefers warm, deep, slowly-flowing water above gravel, sand or mud and can be found in lacustrine habitats. Spawning migrations in the Kura River of Azerbaijan go as far up as Aragva and generally it ascends to the uppermost tributaries of rivers it enters. The spawning run in the Kura lasts almost the whole year except for the two coldest months. However the main spawning runs are in September-October and April (Bogutskaya, Bănărescu and Almaça in Bănărescu and Bogutskaya, 2003). The Caspian Sea form is anadromous but there are also resident forms in the rivers there. Knipovich (1921) reports this species at depths of 9.15-14.2 m, possibly deeper, in the Iranian Caspian Sea. There are both resident and andromous populations in the Anzali lagoon (Karimpour, 1998).

Age and growth

Solak (1989c) examined a population of this species in the Aras River in Turkey and found a life span of over 4 years, but over 6 years in the Çoruh River of the Black Sea basin of Turkey. In the Caspian Sea basin fish may live up to 8 years (Abdurakhmanov, 1962). Anadromous fish are heavier than fish of the same length that are river residents. Maturity is attained at 3-5 years with females mature one year later than males. Spring migrants spawn that summer while summer or autumn migrants overwinter to spawn the following spring or summer (Bogutskaya, Bănărescu and Almaça in Bănărescu and Bogutskaya, 2003). Shajiee et al. (2002) found a sex ratio of 3:1 for male:female fish in the Caspian Sea off Gilan and a life span of 8 years. Gonadosomatic and hepatosomatic indices, length-weight relationships and other growth and fecundity indices were given.

Food

Stomach contents consist of insects, crustaceans and worms, and filamentous algae and other plant material with associated invertebrates. Terrestrial insects, small fishes and frogs are also taken. Abdoli (2000) reports Ephemeroptera, Trichoptera and Chironomidae. One specimen from Iran had fish remains, possibly a small Luciobarbus capito. Abdurakhmanov (1962) reports grasshoppers and ants, presumably taken at the surface.

Reproduction

Eggs number up to 193,600 and diameters up to 1.8 mm in Azerbaijan (Abdurakhmanov, 1962). A fish with well-developed testes was caught in the Gorgan River on 7 July, suggesting a spawning season of late spring and summer, agreeing with egg diameters of fish from Azerbaijan which are largest in June. Eagderi et al. (2006) studied the reproductive cycle in male fish migrating to the Safid and Polrud rivers. Spermatogenesis developed rapidly from late March with the process continuing up to July.

Parasites and predators

Molnár and Jalali (1992) record the monogenean Dactylogyrus linstowi from this species in the Safid Rud. Masoumian and Pazooki (1998) surveyed myxosporeans in this species in Gilan and Mazandaran provinces, finding Myxobolus musculi. Masoumian et al. (2005) recorded the protozoan parasites Ichthyophthirius multifilis and Trichodina perforata from this species in water bodies in West Azarbayjan. Masoumian et al. (2003) record Myxobolus musculi while Pazooki et al. (2003) and Pazooki (2006) record Rhabdochona hellichi, Paradiplozoon homoion and Pseudocapillaria tomentosa, all reports from fishes captured in the Tajan and Zarem rivers of Mazandaran. Sattari et al. (2002) and Sattari (2004) records the presence of the nematode, Eustrongylides excisus, in the body cavity. This parasite can damage muscles in commercial species and render them unsuitable for sale. Sattari et al. (2004, 2005) surveyed this species in the inshore area of the Caspian Sea, recording Eustrongyloides excisus and Anisakis sp. Pazooki et al. (2007) recorded various parasites from localities in West Azarbayjan Province, including Neoechinorhynchus rutili from this species. Miar et al. (2008) examined fish in Valasht Lake and the Chalus River, Mazandaran and found the metazoan Bothriocephalus gowkongensis.

Economic importance

Holčík and Oláh (1992) report a catch of only 9 kg in the Anzali Mordab for 1990. This species had a catch of 17 tonnes in 1997, 28 t in 1998 and 7 t in 1999 during the 6 month beach seine fishing season (October to April). For the years 1998 and 1999, 138 beach seines were used 51,000 times (M. Ramin, pers. comm., 2000). This species was of minor importance commercially in the former U.S.S.R. and is a sport fish in Georgia (Bogutskaya, Bănărescu and Almaça in Bănărescu and Bogutskaya, 2003). In East Azarbayjan it reaches sizes large enough for sport fishing and as a commercial species (Ghasemi, 2002).

Conservation

Kiabi et al. (1999) consider this species to be conservation dependent in the south Caspian Sea basin according to IUCN criteria. Criteria include commercial fishing, sport fishing, medium in numbers, habitat destruction, widespread range (75% of water bodies), absent in other water bodies in Iran, and present outside the Caspian Sea basin. Mostafavi (2007) lists it as conservation dependent in the Talar River, Mazandaran. Vulnerable in Turkey (Fricke et al., 2007).

Further work

Biology and numbers of this species needs investigation.

Sources

Type material: ?

Iranian material: CMNFI 1970-0521, 7, ?-102.5 mm standard length, Gilan, Safid River near Lulaman (no other locality data); CMNFI 1970-0525, 5, 111.9-133.4 mm standard length, Gilan, Safid River near Mohsenabad (no other locality data); CMNFI 1970-0526, 19, ? mm standard length, Gilan, Safid River 6 km below Astaneh Bridge (37º19'N, 49º57'30"E); CMNFI 1970-0531, 1, 157.3 mm standard length, Mazandaran, Larim River (36º46'N, 52º58'E); CMNFI 1970-0536, 1, 194.4 mm standard length, Gilan, Siah River estuary near Rudbar (36º53'N, 49º32'E); CMNFI 1970-0538, 10, 36.7-188.5 mm standard length, Gilan, Qezel Owzan River near Manjil (36º44'N, 49º24'E); CMNFI 1970-0543A, 1, 170.4 mm standard length, Gilan, Caspian Sea at Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1970-0546, 10, 39.3-61.8 mm standard length, Gilan, Safid River canal (no other locality data); CMNFI 1970-0553, 1, 58.1 mm standard length, Gilan, Sowsar Roga (37º27'N, 49º30'E); CMNFI 1970-0563, 1, 70.1 mm standard length, Gilan, Caspian Sea at Kazian Beach (ca. 37º29'N, ca. 49º29'E); CMNFI 1970-0568, 8, 62.5-132.0 mm standard length, Gilan, Caspian Sea at Kazian Beach (ca. 37º29'N, ca. 49º29'E); CMNFI 1970-0581, 6, 41.3-65.5 mm standard length, Gilan, Caspian Sea near Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1970-0587, 3, 69.0-91.4 mm standard length, Mazandaran, Babol River at Babol Sar (36º43'N, 52º39'E); CMNFI 1979-0431, 2, 240.9-265.5 mm standard length, Mazandaran, bazaar at Now Shahr (no other locality data); CMNFI 1979-0437, 1, ? mm standard length, Gilan, Safid River 2 km west of Astaneh (37º16'30"N, 49º56'E): CMNFI 1979-0452, 2, 53.5-56.5 mm standard length, Azarbaijan-e Khavari, Qezel Owzan River 6 km from Mianeh (37º23'N, 47º45'E); CMNFI 1979-0486, 2, 69.2-78.8 mm standard length, Mazandaran, Atrak River draiange (37º44'N, 56º18'E); CMNFI 1979-0488, 1, 95.8 mm standard length, Mazandaran, Atrak River at Maraveh Tappeh (37º55'N, 55º57'30'E); CMNFI 1979-0491, 1, 191.5 mm standard length, Mazandaran, Gorgan River 15 km northeast of Kalaleh (ca. 37º33'N, ca. 55º44'E); CMNFI 1979-0686, 19, ? mm standard length, Gilan, Safid River (37º24'N, 49º58'E): CMNFI 1979-0695, 4, ? mm standard length, Gilan, Safid River at Manjil Bridge (36º46'N, 49º24'E): CMNFI 1979-0788, 2, 152.0-202.4 mm standard length, Mazandaran, Gorgan River at Khadje Nafas (37º00'N, 54º07'E); CMNFI 1980-0116, 8, ?-62.4 mm standard length, Gilan, Safid River at Astaneh Bridge (37º16'30"N, 49º56'E); CMNFI 1980-0123, 8, ? mm standard length, Gilan, Safid River (ca. 37º22'N, ca. 49º57'E): CMNFI 1980-0127, 1, ? mm standard length, Gilan, Caspian Sea near Hasan Kiadeh (37º24'N, 49º58'E): CMNFI 1980-0132, 8, ? mm standard length, Gilan, Safid River at Kisom (37º12'N, 49º54'E); CMNFI 1980-0138, 2, 132.5-137.6 mm standard length, Gilan, Safid River estuary (ca. 37º28'N, ca. 49º54'E); CMNFI 1980-0905, 1, 188.9 mm standard length, Mazandaran, Gorgan River at Khadje Nafas (37º00'N, 54º07'E); CMNFI 1980-0908, 3, 67.2-91.3 mm standard length, Gilan, Safid River estuary (ca. 37º28'N, ca. 49º54'E); uncatalogued, 2, 245.7-272.8 mm standard length, Markazi, Shah River (no other locality data);

Luciobarbus esocinus
Heckel, 1843



Darreshahr, Simarreh River, April 1987,
photo by N. Atarody; courtesy of B. Kiabi

Kermanshah, Simarreh River,
courtesy of B. Kiabi

more photos

Common names

سونگ (= soong) or بچ ( = bach) in northern Khuzestan and Lorestan, anzeh, anzah, narbach, and anzeh-bach at Ahvaz and in southern Khuzestan (meanings unknown); بل زرد (= balzard).

[bizz (in Iraq), farkh-el-biz (= cheerful one (Heckel, 1843b) or baby of bizz), farch or mangar in Arabic; "Tigris salmon", "Euphrates salmon", pike barb, pike barbel].

Systematics

Howes (1987) places this species in Barbus sensu stricto. Labeobarbus Euphrati Sauvage, 1882 described from "Biredjik (Euphrates)", Turkey (not "Irak" as in Bertin and Estève (1948)) is a synonym.

Karaman (1971) places this species in the synonymy of "Barbus" xanthopterus as he considered the only difference to be scale count and the range of variation for these species is unknown. Almaça (1983, 1986) agrees that several meristic characters are similar while the main differences are a shorter head and barbels in esocinus and dotted coloration in esocinus as opposed to uniform in xanthopterus (isn't the reverse true?). He maintains them as separate species because information on variability in characters is lacking.

Examination of the types of L. esocinus (NMW 54088, 2, 58.5-61.5 mm standard length, 54091, 372.4 mm, 54092, 321.3 mm) and L. xanthopterus (NMW 54841a (a syntype), 216.5 mm, 54786 (not a type), 292.8 mm) in Vienna showed the following differences. Head size differs in the two taxa in that esocinus postorbital length is very elongate and the head tapers anteriorly in a distinctive fashion. Head length in standard length is 3.2-3.6, mean 3.4 for esocinus and 4.0-4.2, mean 4.1 for xanthopterus and postorbital length in standard length is 5.9-7.2, mean 6.5 for esocinus and 7.7-7.8, mean 7.8 for xanthopterus with the higher values for esocinus based on smaller fish which tend to have proportionately larger heads. Total gill raker counts are 8-10, mean 9.3 for esocinus and 12-13, mean 12.5 for xanthopterus. Larger esocinus appear to lose anterior rakers with age but still have fewer than xanthopterus of similar size. Lateral line scale counts are 63-70, mean 67.3 in esocinus and 57-60, mean 58.5 in xanthopterus. On this limited basis I am maintaining the two species as distinct. An Iranian specimen, 284.3 mm standard length (ZSM 21830 from the Dez River), falls within the ranges given above. see xanthopterus?

Almaça (1986) records syntypes of Luciobarbus esocinus in the Naturhistorisches Museum Wien from the type locality as given by (Heckel, 1843b) "bei Mossul in Tigris", Iraq (NMW 54088 (2 specimens), NMW 54091 (1), and NMW 54092 (1) but Heckel (1843b) does not specify the number of types). A syntype is in the Senckenberg Museum Frankfurt (SMF 454, formerly NMW; 281.2 mm standard length) and another syntype is also there but dried (SMF 6785, formerly NMW) (F. Krupp, pers. comm., 1985). The catalogue in Vienna lists 2 fish in spirits and 2 fish stuffed.

The mounted holotype of Labeobarbus euphrati is in the Muséum national d'Histoire naturelle, Paris (MNHN A.6961) and measures 1650 mm total length (Bertin and Estève, 1948). Eschmeyer et al. (1996) indicate that the catalogue number may be A.6971.

Key characters

This species is characterised by large size, a long, tapering and depressed head (rather pike-like in shape), two pairs of barbels, a serrated dorsal fin spine, lateral line scale count high (63-78), moderately developed lips, and no large flank spots. Head length in standard length 3.1-3.7, mean ? and postorbital length in standard length 5.9-7.2, mean ? for ? specimens ?-? mm standard length including the types listed above. (includes SMF454 281.2 mm SL, HL 75.6, postorb 44.1, snout 21.5

Morphology

Dorsal fin with 4 unbranched and 8 branched rays, anal fin with 3 unbranched and 5 branched rays, pectoral fin with 16-18 branched rays and pelvic fin with 8 branched rays. Lateral line scales 62-78. Scales are regularly arranged, the smallest being on the isthmus anterior to the pectoral fin bases. There is a pelvic axillary scale. Scales have a central focus, numerous fine circuli, a wavy or rounded anterior margin, and radii on the anterior and posterior fields with a few widely spaced ones on the lateral fields. Gill rakers 8-12, well spaced and just touching the one below when appressed. Pharyngeal teeth 2,3,5-5,3,2, hooked with the third tooth of the inner row slightly larger than the fourth and the fifth smaller. Heckel (1843b) gives 2,3,4-4,3,2, and teeth from large specimens seen at Ahvaz in 1995 by me had 2,3,4-4,3,2 and 2,3,5-4,3,2, the anteriormost tooth being small or absent. Even small specimens (85.7 mm standard length) may have the anteriormost tooth absent. Total vertebrae 48 (Howes, 1987) or 48-50 based on comparative materials listed below. The last unbranched dorsal fin ray is very strong, with a low density of denticles but with fine denticles extending over much of the ray. The mouth is large, terminal and almost horizontal and extends back to the anterior eye margin. Lips are thin to moderate without a median lobe to the interrupted lower lip, and barbels are thin to very thin. The anterior barbel does not reach past the nostril level and the posterior barbel does not pass the mid-eye to rear eye level. The nostril is elongate and closer to the eye than the snout tip. The cephalic canals on the suborbital series have numerous branches. The gut is an elongate s-shape with several anterior loops.

Meristics for an Iranian specimen:- dorsal fin branched rays 8; anal fin branched rays 5; pectoral fin branched rays 17; pelvic fin branched rays 8; lateral line scales 69; and total gill rakers 9.

Sexual dimorphism

Unknown.

Colour

The back has numerous scattered, black spots on an olivaceous background, the spots extending onto the base of the dorsal fin. Spots may be weak or absent but this is comparatively rare. Overall colour is silvery with the anal and caudal fins dark red. The flanks and belly are lighter. The eye is yellowish in colour. Young fish have a yellow tinge or sulphur yellow colour to the fins.

Size

Frequently up to 3 hundredweights (= 152.4 kg) in the Zab River of Iraq southeast of Mosul (Heckel, 1846-1849a); a fish 6'4" (1.93 m) long with a girth of 3'10" (1.17 m) and a weight of 215 lbs (97.6 kg) from the Euphrates River at Hakika (Light, 1917; wrongly identified as "Barbus" scheich according to the editors in an article by Gudger (1945a)); 69 inches (1.75 m) measured over the curve of a back with a 38 inch (0.97 m) girth and a weight of 123 lbs (55.8 kg) caught in the Diyala River, Iraq on a light 14-foot rod taking 1½ hours to land (Bagnall, 1919); 96 lb (43.6 kg) fish caught near Kizil Robat (= As Sa`diyah) in the Diyala River on a lump of atta (a ball of dough)(MacKay, 1919)(Bagnall, a Major, out-doing MacKay, a Brigadier-General); 140 lbs (63.6 kg) Tigris salmon caught on a 2" spoon at Samarra (Lane, 1920); hundreds of good weight up to 112 lbs (50.8 kg), one caught on a hand-line at 170 lbs (77.2 kg), one netted at 252 lbs (114.4 kg), and reputedly over 300 lbs (136 kg)(Radcliffe, 1926); up to two yards (1.83 m) as evidenced by a photograph of a specimen draped over a donkey in Iraqi Kurdistan (Hamilton, 1937); 2 m and 150 kg in Iraq (van den Eelaart, 1954; Herzog, 1967); a 167 lb (75.8 kg) Tigris specimen and a 213 lb (96.7 kg) specimen at Nassiriyah on the Euphrates, called both gattan and "Euphrates salmon" but it was presumably the latter (Vesey-Fitzgerald and Lamonte (no date)); weights up to 300 lbs (136 kg) and the largest taken on rod-and-line as 220 lb (100 kg) and 7 feet (2.1 m), baits used included atta and dates, and chicken or sheep liver (Mahdi, 1962). Beck (pers. comm., 2000) reports the largest fish seen in the 1990s along the Syrian Euphrates and its tributaries weighed 198 kg. A fish caught in 2001 on the Euphrates River near Birecik in Turkey with a net weighed 111 kg and was 2.4 m long (www.fishing-worldrecords.com, downloaded 16 February 2007).

Iranian records of large specimens include one by Mr. Chabok-Savar, a Game Warden or biologist of the Department of the Environment who caught a specimen about 80 kg in the Simareh River in 1973 and N. Atarody, also a Game Warden or biologist, caught two large specimens in April 1987 from "Tang-e Gheer" on the Simareh near Darreh Shahr (Abzeeyan, Tehran, 3 (August-September):19, 1992). A 1.65 m and 75 kg specimen is reported from the Dez River and a 2.1 m specimen is reported from the market at Ahvaz in 1993 (this last fish may have weighed 150 kg, original report not seen; J. Valiallahi, pers. comm., 2001). The Gav Masiab, a river in Kordestan, is reputedly named for these large fishes ("river with fishes as large as a cow")(J. Valiallahi, www.modares.ac.ir, downloaded 4 July 2000). Floor (2003) gives a photograph of a large specimen from the Karun River.

Distribution

This species is found in the Tigris-Euphrates basin including its Iranian portion and the adjacent northern Gulf basin (Marammzai, 1995; Abdoli, 2000). It is reported as common in the Dez Dam (Gh. Eskandary, pers comm., 2000).

Zoogeography

Almaça (1984b, 1991) considers that the origin of this species lies with a group that migrated southwards in the late Pliocene from the Dacian Lake of the Sarmatian Sea and speciated in Mesopotamia.

Habitat

Found in large rivers and dams but also the more limited environment of palace ponds at Baghdad. Details of environmental requirements unknown.

Age and growth

Life span is at least 10 years (Ahmed, 1982). Fish in Khuzestan were found to have a sex ratio of 4.2:1.0 male:female (Gh. Eskandary, pers. comm., 2000; Eskandari et al., 2004). In the Dez Dam of northern Khuzestan females had a length range of 156-1350 mm and a weight range of 31.7-26,500 g while for males figures were 183-1065 mm and 48-12,208 g. Males matured faster than females, annual growth is slow and asymptotic length is more than 2 m. It appears to have a longer reproductive life compared to pre-maturation life (Eskandari et al., 2004).

Food

This species is a predator on other fishes. In the Dez Dam, all samples had fish in their stomachs although the gut to body length ratio indicates omnivory (Eskandari et al., 2004). Al-Rudainy (2008) cites also crustaceans, aquatic insects and zooplankton in Iraq.

Reproduction

van den Eelaart (1954) reports spawning in Iraq in March and Al-Rudainy (2008) gives April to May. Eggs are laid between large stones in the deep part of rivers, with absolute fecundity in Iraq at 600,000 eggs (Al-Rudainy, 2008). Some fingerlings drift down into lakes and marshes. Eskandari et al. (2004) report a very short spawning season in the Dez Dam in spring after reservoir water levels rise through spring flooding. The fish is a total spawner with eggs released in upstream areas and shallows of the reservoir over gravel at 24ºC. Ünlü (2006) gives age at first maturity as 4 years in the Turkish Tigris River while Al-Rudainy (2008) gives sexual maturity as 10 years in Iraq.

Parasites and predators

Masoumian et al. (2008) recorded the myxosporeans Myxobolus karuni and M. persicus from gills of fish captured in the Karun and Karkeheh rivers and Shadegan Marsh.

Economic importance

In Iraqi Kurdistan these fish were caught and tethered by a cord passed through the lips until eaten by the villagers (Elliot, 1977). At Altan Keupri on the Lesser Zab River in Iraq a drugged bait was used to stupefy the fish so it could be netted and dragged to shore (Hamilton, 1937).

This species was being considered for aquaculture during the year 2000 in Khuzestan although fish larger than 1 m are needed to be ripe adults. Anglers and commercial fishermen seek this fish in the Iranian Zagros Mountains using ducklings (!) as bait (J. Valiallahi, pers. comm., 2001).

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in textbooks.

Conservation

This species is under severe threat in the Syria Euphrates and its tributaries. A survey in 1997-1998 caught only a single juvenile and the commercial fisheries had not more than two dozen fish. Blast fishing and poisoning had led to a decline in age of catches since 1993. Large scale water abstraction, dam building and pollution had destroyed habitats (R. Beck, pers. comm., 2000). It is listed by Stone (2007) as one of the world's largest freshwater fishes, presumed to be threatened.

A report of fish kill, presumably of this species, in the "Cham Ghorah" River near Mahabad in July 1999 numbering about half a million fish was owing to desiccation of the habitat (J. Valiallahi, www.modares.ac.ir, downloaded 4 July 2000).

Further work

Distribution and numbers are needed for a conservation assessment.

Sources

Type material: See above, NMW 54088.

Iranian material: ZSM 21830, 1, 284.3 mm standard length, Khuzestan, Dez River at Harmaleh (31º57'N, 48º34'E). Some specimens observed in the IFRTO laboratory at Ahvaz (pharyngeal arches).

Comparative material: BM(NH) 1892.9.1:30, 1, 197.3 mm standard length, Iraq, Al Faw (29º58'N, 48º29'E); BM(NH) 1920.3.3:80-82, 3, 85.7-147.0 mm standard length, Iraq, Basrah (30º30'N, 47º47'E); BM(NH) 1931.8.12:5, 1, 111.8 mm standard length, Iraq, near Mosul (no other locality data); BM(NH) 1974.2.22:1297, 1, 166.5 mm standard length, Iraq, Diyala River (no other locality data); BM(NH) 1974.2.22:1810, 1, 220.1 mm standard length, Iraq (no other locality data).

Luciobarbus kersin
(Heckel, 1843)

Common names

برزم (= berzem).

[shissan, jassan, gassan, djissan, barsam or bunni, kersin at Aleppo, all in Arabic; kersin barbel].

Systematics

Karaman (1971) places this species as a synonym of his Barbus capito pectoralis. Almaça (1983) suggests that kersin may be only subspecifically distinct from Barbus (= Luciobarbus) pectoralis (q.v.) but later (Almaça, 1984b) retains it as a full species until further information becomes available. Krupp (1985c) also synonymises this species with Barbus (= Luciobarbus) pectoralis.

Syntypes of Barbus Kersin from "Aleppo", the type locality given by Heckel (1843b) or "Gewässern von Aleppo" (Heckel, 1846-1849a), are in the Naturhistorisches Museum Wien (NMW 54212 and 54215) (Almaça, 1986). Krupp (1985c) lists the following syntypes of B. kersin all from Aleppo and collected by Th. Kotschy: 1 specimen, 141.2 mm standard length as measured by me (NMW 54212), 4, 89.1-135.1 mm standard length as measured by me (NMW 54213), 1, 166.0 mm standard length as measured by me (NMW 54215) and 1, 152 mm standard length (formerly NMW, now in the Senckenberg Museum Frankfurt as SMF 610). The card catalogue in 1997 listed NMW 54215 as "? lectotype" and NMW 54213 as "? paralectoptypes" (sic). Eschmeyer et al. (1996) list 1 syntype in the Museum für Naturkunde, Universität Humboldt, Berlin (ZMB 3237, formerly NMW). The catalogue in Vienna lists 6 specimens.

Key characters

This species differs from the similar L. pectoralis by the smaller scales and body depth being greater than head length (equal in pectoralis) (Berg, 1949). ?see BWC95-32 for a kersin after Valiallahi, its a dark Barbus without the very large D spine and a different colour pattern

Morphology

Dorsal fin unbranched rays 3-4, usually 4, branched rays 7-8, anal fin with 3 branched and 5-6, usually 5, unbranched rays. Pectoral fin branched rays 17, pelvic fin rays 8. Lateral line scales 49-58. Gill rakers 19. Pharyngeal teeth 2,3,5-5,3,2. The last unbranched dorsal fin ray is strong (as in pectoralis)with a low density of coarse denticles extending over much of the ray. The mouth is moderate in size and subterminal. The highly rounded snout projects a little. Lips are thin to moderate but not fleshy and lack a median lobe. The upper lip is covered partly by the snout. Barbels are thin, the anterior barbel not extending back beyond the anterior eye margin and the posterior barbel not beyond the middle of the eye. Body depth is equal to or greater than head length in the types examined by me.

Sexual dimorphism

Unknown.

Colour

The body lacks distinctive markings and is olive to reddish-brown above, silvery on the flanks and white below. The dorsal and caudal fins have a blackish margin.

Size

Attains 70.1 cm total length (Menon, 1956). Reaches 2 m and over 100 kg (Khalaf, 1961).

Distribution

Found in the Tigris-Euphrates, Quwayq and Orontes River basins. In Iran, it is found in the Tigris River basin in the Karun and Karkheh rivers, and the northern Gulf basin in the Zohreh and Helleh rivers and questionably the southern Gulf basin (Abdoli, 2000).

Zoogeography

Almaça (1984b, 1991) considers that the origin of this species lies with a group that migrated southwards in the late Pliocene from the Dacian Lake of the Sarmatian Sea and speciated in Mesopotamia.

Habitat

The main habitat of Iraqi fish is rivers, entering marshes and lakes during floods but returning to rivers in June (van den Eelaart, 1954).

Age and growth

Unknown.

Food

This species is said to eat a wide range of food items (Beckman, 1962), including aquatic insects and plants (Al-Rudainy, 2008).

Reproduction

Eggs are deposited on clay or gravel bottoms during mid-February to early March in Iraq (van den Eelaart, 1954). Al-Rudainy (2008) gives the spawning season as March to April in Iraq.

Parasites and predators

Gussev et al. (1993a) describe new species of monogeneans from this species in the Dez River, Khuzestan, namely Dactylogyrus deziensis, D. deziensioides and D. kersini. Ebrahimzadeh and Kailani (1976) record parasite species in the cestode genera Caryophyllaeus and Isoglaridacris and the protozoan Myxosoma from Barbus (= Luciobarbus) kersin taken in the Karun River.

Economic importance

None.

Conservation

Endangered in Turkey (Fricke et al., 2007) but status in Iran unknown.

Further work

The biology of this species in Iran needs study and a molecular comparison with putative synonyms would be of value in clarifying distinctiveness.

Sources

Type material: See above, NMW 54212, NMW 54213 and NMW 54215.

Iranian material: None.

Comparative material: BM(NH)1974.2.22:1324, 186.2 mm standard length (); BM(NH)1920.3.3:41-50, 12(5 examined), 110.9-165.3 mm standard length (); BM(NH)1920.3.3:31-40, 10, 141.7-310.9 mm standard length ();

Luciobarbus mursa
(Güldenstaedt, 1773)

L. capito (above) and L. mursa (below) Aras River, 2 October 1994
Courtesy of Asghar Abdoli

Common names

سس ماهي (= sos, sas or sass mahi), mahi siah (= black fish), zardek-e qalami (= slender or straight yellow one), ses mahi koloft safid rud (= Safid River thick fish, the meaning of ses, sos or sas being unknown but referring to "Barbus"), sas mahi-ye lab koloft (= thick lip "Barbus" fish), زرده پر (= zardehpar).

[mursa or shchirbit in Azerbaijan; murtsa or mursa in Georgian; murtsa or Araksinskaya murtsa, both in Russian].

Systematics

Cyprinus mursa was originally described from the Kura River at Tbilisi, Georgia. Syntypes are presumed lost (Bogutskaya, Bănărescu and Almaça in Bănărescu and Bogutskaya, 2003)

Barbus miliaris De Filippi, 1863 described from a "fiumicelli presso Teheran" (= a stream near Tehran), Barbus mursoides Kessler, 1877 described from Transcaucasia (presumably the Kura-Araks basin), Barbus microphthalmus Sauvage, 1882 from "Tiflis" (presumably the Kura River at Tbilisi, Georgia) (and Barbus macrophthalmus and Barbus mycrophtalmus in Chantre (1882) which are presumably misspellings of this name; the former is in any case preoccupied by B. macrophthalmus Bleeker, 1855 described from Indonesia), Barbus kessleri Derzhavin, 1929 described in Latin from the "Keredsh flumen" (= Karaj River near Tehran), and Barbus dageti Fowler, 1958 are synonyms. Barbus dageti was coined because Fowler believed Barbus kessleri was preoccupied by Puntius kessleri Steindachner, 1866; Puntius Hamilton, 1822 is not now considered a synonym of Barbus sensu lato (Eschmeyer, 1990) although Eschmeyer et al. (1996) have Barbus kessleri listed as preoccupied by Puntius kessleri. Eschmeyer et al. (1996) record 3 syntypes "whereabouts unknown" for Barbus dageti, i.e. Derzhavin's Barbus kessleri types. Dadikyan (1986) refers Aras River fish from Armenia to Barbus mursa mursoides.

Three specimens (presumably syntypes)(MZUT N.676) of Barbus miliaris are stored in the Istituto e Museo di Zoologia della R. Università di Torino (Tortonese, 1940).

The holotype of Barbus mursoides is in the Zoological Institute, St. Petersburg (ZISP 2863) from the Caucasus collected by Hohenacker in 1838.

Two syntypes of Barbus microphthalmus, measuring 340 mm total length, are in the Muséum national d'Histoire naturelle, Paris (MNHN A.3923, formerly MNHN 1881-1007 and MNHN 1881-1008) (Bertin and Estève, 1948).

Howes (1987) considers the generic placement of Barbus miliaris as problematical. It has a series of preanal scales and a prominent genital papilla similar to schizothoracines, and a lachrymal bone similar to Barbus (= Tor) grypus and B. sharpeyi. Karaman (1971) considers Barbus miliaris from the Namak Lake basin of Iran to be a subspecies of the Caspian Sea basin type subspecies, differentiated by larger scales (78-92 compared to 85-103), less fleshy lips, an undeveloped lower lip lobe, feebly ossified last dorsal fin spine, and shorter pectoral fins. Derzhavin (1929b) in describing his Barbus kessleri on fish 121-154 mm total length with well-developed gonads states that the lower lip is clearly trilobate. Berg (1949) recognises miliaris as distinct from mursa on the basis of a shorter snout, somewhat larger scales, fewer scale rows above the lateral line, smaller dimensions and different colour. Bianco and Banarescu (1982) and Almaça (1984b) retain it as a full species although Bianco and Banarescu (1982) also suggest that this species may be a subspecies of their wide-ranging taxon Barbus cyclolepis Heckel, 1837. Almaça (1984a) points out that his conclusion is based in part on small specimens in poor condition and that there is not enough data to take a sound decision (Almaça notes that gill raker counts are low and the lower lip lobe undeveloped in accordance with Karaman (1971) but these are characters which I believe may be size and age related). Almaça (1992) also distinguishes the two taxa on the shorter barbel in miliaris (not exceeding the middle of the eye as opposed to not exceeding the rear border of the eye), slope of the dorsal fin oblique in miliaris as opposed to oblique to nearly perpendicular, and pharyngeal teeth in miliaris 5,3 (or 4),2 as opposed to 4-5,3,2. These characters too may be size dependent or individually variable, as are those of Berg (1949). I consider that miliaris is at most a subspecies of mursa.

Key characters

The high scale counts are an important character as is the presence, usually, of a fleshy three-lobed lower lip.

Morphology

Dorsal fin unbranched rays 3-5, usually 4, followed by 7-8, usually 8, branched rays, anal fin unbranched rays 3 followed by 5 branched rays. Pectoral fin branched rays 13-17 and pelvic fin branched rays 6-8, usually 7. Lateral line scales 74-103, often 85 or more. Scales are small, horizontally elongated and almost rectangular, with a anterior margin variably indented, a very anterior focus, relatively few and well-spaced circuli, and few radii on all fields. A single pelvic axillary scale is not developed but a series of enlarged scales may be separated from other scales by a fold of skin. Gill rakers 7-18 (10-18 in Bogutskaya, Bănărescu and Almaça in Bănărescu and Bogutskaya (2003), lower counts from the literature perhaps not including rakers on the upper arm of the arch. There may also be differences due to size and, independent of size, the rakers on the lower arch anteriorly are variably developed, sometimes being reduced to bumps which were counted and sometimes not even bumps are present. The larger rakers reach the second adjacent raker when appressed. Vertebrae 41-45. Pharyngeal teeth 2,3,5-5,3,2, rarely 1,2,3,5-5,3,2,1; or with only 4 teeth in the main row (e.g. see Heckel (1843b)). Teeth are hooked and the fourth inner row tooth is slightly larger or smaller than the third. The fifth tooth is smaller (sometimes minute) than teeth 3 and 4 and may be pointed or blunt. The grinding surface below the tip is short, uneven and concave to rounded. The mouth is moderate in size, inferior, horseshoe-shaped with moderate to thick fleshy lips and an undeveloped to strongly developed median lower lip lobe (see above). Barbels are thick, the anterior one not extending back beyond the nostril level and the posterior one not exceeding the middle or posterior eye margin. The last dorsal fin spine is moderate to strong and has many, closely-packed denticles from one-half to four-fifths of the spine length, although denticles are lost in adults. The gut is elongate with 2-3 anterior loops. Chromosome number 2n = 100, NF 140 (Pourali Darestani et al., 2006).

Meristic variation in Iranian specimens:- dorsal fin branched rays 7(1) or 8(18); anal fin branched rays 5(19); pectoral fin branched rays 14(1), 15(2), 16(13) or 17(3); pelvic fin branched rays 7(13) or 8(6); lateral line scales 75(1), 80(2), 82(1), 84(1), 85(1), 87(1), 88(3), 89(3), 90(3), 91(1), 92(1) or 95 (1); total gill rakers 9(2), 10(1), 12(1), 13(5), 14(4), 15(2) or 16(3); pharyngeal teeth 2,3,5-5,3,2(7), 2,3,4-5,3,2(5), 2,3,5-4,3,2(2) or 3,4,5-5,3,2; and total vertebrae 43(1), 44(6) or 45(2).

Sexual dimorphism

Etessami (1982) reports an hermaphrodite in this species in the Namak Lake basin. A female specimen, 112.5 mm standard length, caught on 15 July had tubercles on the top and upper sides of the head. Male tuberculation in large adults has not been reported on.

Colour

Overall colour is a pale grey to olive-grey to brownish, slightly darker over the back, and the belly is white to yellowish-brown. The sides of the head and flanks can have golden tints. The iris is grey with a narrow rim of silver immediately around the pupil or may be yellow-gold. The dorsal and caudal fins are pale grey to dark reddish-brown. The caudal fin bears several series of small dark spots. The pectoral and pelvic fins have pale brown rays and transparent membranes but may be pink. The anal fin may be colourless except for a little grey pigment over the last unbranched and first branched rays to an overall reddish-brown. The margins of the pelvic and anal fins are well-developed and white, while the pectoral fin has a very narrow white margin. Young may have numerous dark spots on the back and upper flank, lost in adults.

Size

Attains 39.5 cm or 43 cm total length (Jolodar and Abdoli, 2004).

Distribution

This species occurs in the Kura River basin of the southwestern Caspian Sea and in southern tributaries of the Caspian from Iran. In Iran, it is reported in the Caspian Sea basin from the Aras to the Gorgan rivers including the Tajan, Babol, Haraz, Sardab, Aras, Tonekabon, Pol-e Rud and Safid rivers (Abbasi et al., 1999; Kiabi et al., 1994; 1999; Abdoli, 2000; Abdoli and Naderi, 2009), the Namak Lake basin (Wossughi, 1978; Rainboth, 1981; Almaca, 1984a; Bianco and Banarescu, 1982; Abdoli, 2000), and the Lake Orumiyeh basin in the Arnar Chay, Nowruzlu Chay, Tatavi and Zarrineh rivers (Günther, 1899; Abdoli, 2000).

Zoogeography

This species is possibly a Caspian Sea endemic, depending on the status of populations in the Namak and Orumiyeh Lake basins.

Habitat

Avoids muddy bottoms, preferring streams with rapid water, gravel and sand bottoms and a rich benthos. It may also be found in lacustrine habitats (Solak, 1977; Bogutskaya, Bănărescu and Almaça in Bănărescu and Bogutskaya, 2003).

Age and growth

Solak (1989b) reports a life span of 6 years in the Aras basin of Turkey. Maturity is attained at 2-3 years (Bogutskaya, Bănărescu and Almaça in Bănărescu and Bogutskaya, 2003).

Food

Food items include chironomids, as much as 70-100% of the diet at times, crustaceans such as copepods and ostracods, insects, worms, plankton, vegetation and detritus (Abdurakhmanov, 1962; Bogutskaya, Bănărescu and Almaça in Bănărescu and Bogutskaya, 2003). Iranian fish guts contain plant fragments, aquatic insects such as chironomids and Ephemeroptera (mayflies), and crustaceans such as amphipods.

Reproduction

Fecundity is up to 25,000 eggs. The spawning season is probably in May and June as noted for Georgian fish in Abdurakhmanov (1962) but may extend from April to August, the peak depending on locale (Bogutskaya, Bănărescu and Almaça in Bănărescu and Bogutskaya, 2003). Fish caught on 6 July in Mazandaran, 64 km west of Dasht had large, possibly atretic, eggs measuring about 1.5 mm although Bogutskaya, Bănărescu and Almaça in Bănărescu and Bogutskaya (2003) report a maximum egg diameter of 2.5 mm.

Parasites and predators

Masoumian et al. (2003) record Myxobolus azerbajdzanicus, M. kovali, M. squamae, M. tauricus, M. rutili and M. osmaniae while Pazooki et al. (2003) and Pazooki (2006) record Rhabdochona hellichi, Bothriocephalus gowkongensis and Paradiplozoon homoion, all reports from fishes captured in the Tajan and Zarem rivers of Mazandaran.

Economic importance

Said to taste even better than trout (Abdurakhmanov, 1962), it is caught by some anglers but is not commercially important (Bogutskaya, Bănărescu and Almaça in Bănărescu and Bogutskaya, 2003).

Conservation

Kiabi et al. (1999) consider this species to be near threatened in the south Caspian Sea basin according to IUCN criteria. Criteria include sport fishing, medium in numbers, habitat destruction, widespread range (75% of water bodies), present in other water bodies in Iran, and absent outside the Caspian Sea basin. Mostafavi (2007) lists it as near threatened in the Talar River, Mazandaran. Bogutskaya, Bănărescu and Almaça in Bănărescu and Bogutskaya (2003) report that is is extremely rare in Azerbaijan. Endangered in Turkey (Fricke et al., 2007).

Further work

The presence of this species in the Lake Orumiyeh basin and the taxonomic status of Namak Lake basin populations need careful examination.

Sources

Type material: ?

Iranian material: CMNFI 1970-0525, 1, 49.3 mm standard length, Gilan, Safid River near Mohsenabad (ca. 37º22'N, ca. 49º57'E); CMNFI 1970-0538, 5, 42.5-82.7 mm standard length, Gilan, Qezel Owzan River near Manjil (36º44'N, 49º24'E); CMNFI 1970-0545, ?, ? mm standard length, (); CMNFI 1970-0589, 1, 110.0 mm standard length, Gilan, Safid River opposite Kisom (37º12'N, 49º54'E); CMNFI 1979-0084, 2, 92.5-96.8 mm standard length, Mazandaran, Chalus River (no other locality data); CMNFI 1979-0253, ?, ? mm standard length, (); CMNFI 1979-0456, 2, 44.1-50.2 mm standard length, Markazi, Shah River at Lowshan (36º37'30"N, 49º31'E); CMNFI 1979-0481, 1, 142.7 mm standard length, Mazandaran, stream 3 km west of Ghalahleekesh (37º18'30"N, 55º31'E); CMNFI 1980-0132, 1, 112.5 mm standard length, Gilan, Safid River at Kisom (37º12'N, 49º54'E); CMNFI 1991-0158, ?, ? mm standard length, (); CMNFI 1993-0136, 1, 109.9 mm standard length, Mazandaran, Sardabrud (36º39'42'N, 51º22'36'E); CMNFI 2007-0086, 1, 182.2 mm standard length, Azarbaijan-e Khavari, Qareh Su basin near Nir (ca. 38º02'N, ca. 48º00'E); Behnke 8 Behnke 79 CMNFI 2007-00, ?, ? mm standard length, (); FMNH 51245, 2, 108.4-128.5 mm standard length, Markazi, Rayy (35º35'N, 51º25'E); ZMH 2429, 98.1 mm standard length, Markazi, Tehran (no other locality data).

Luciobarbus pectoralis
(Heckel, 1843)

Common names

basan, برزم  (berzem or barzam), ? tu'ini.

[nebbash, sheikh san or shabbout pectoralis in Arabic; Heckel's Orontes barbel (Fricke et al., 2007)].

Systematics

Howes (1987) places this species in Barbus sensu stricto. Barbus perniciosus Heckel, 1843 described from "Gewässern bei Damascus", Luciobarbus Schejch Heckel, 1843 described from "Mossul" (also spelt schech on p. 1019 and p. 1098 in Heckel, presumably in error, and sometimes emended to scheich), Labeobarbus Orontis Sauvage, 1882 from the "Canal de l'Oronte à Antioche", Turkey and possibly Barbus (= Luciobarbus) kersin Heckel, 1843 (q.v.) and possibly Barbus (= Luciobarbus) barbulus Heckel, 1849 (q.v.) are synonyms (see Krupp (1985c)). Barbus pectoralis was described from the "Orontes" (Heckel, 1843b) but the catalogue in Vienna reads "Damascus" (possibly in confusion as this part of the catalogue has been overwritten).

The holotype of Labeobarbus orontis is in the Muséum national d'Histoire naturelle, Paris (MNHN A.3868), with a length of 600 mm (Bertin and Estève, 1948). The catalogue in the Naturhistorisches Museum Wien appears (the catalogue is overwritten here) to list a single specimen opposite each of the names Barbus perniciosus and Barbus pectoralis, probably the holotypes. Krupp (1985c) records the holotype of L. pectoralis as being 116 mm standard length (NMW 54474 and the holotype of B. perniciosus as being 105 mm standard length (NMW 54472). My examination of NMW 54474 showed a length of 117.8 mm standard length.

Karaman (1971) places pectoralis as a subspecies of Barbus (= Luciobarbus) capito but Almaça (1986) disagrees on several grounds, especially on the number of pharyngeal teeth (5 in capito and 4 in pectoralis in the main row (yet Heckel (1843b) gives 5 main row teeth for pectoralis, 4 for the synonym schejch), see also below). Luciobarbus kersin has 5 main row pharyngeal teeth, an indication that it may be distinct. Almaça (1986) points out that the specimen in the Naturhistorisches Museum Wien (NMW 54475) referred to as the holotype of pectoralis by Karaman (1971) is from the wrong locality and was collected at a later date.

Barbus Rajanorum Heckel, 1843 described from "Aleppo" and later in Heckel (1846-1849a) from "Gewässern von Aleppo" is a hybrid of this species and Capoeta damascina (F. Krupp, in litt., 1986) and Almaça (1991) also believes it to be founded on a hybrid; see also Almaça (1983; 1991), Berg (1949) and Karaman (1971) for conflicting views). Valiallahi (2000) also considers this species to be a hybrid, with pectoralis or barbulus. Almaça (1983) could not find any specimens attributable to Barbus rajanorum and the holotype housed in the Naturhistorisches Museum Wien is lost. However, the type locality for this taxon is "Aleppo" (Heckel (1843b) and Krupp (1985c) states that the holotype is NMW 54494, 190 mm standard length, Aleppo, 1842, Th. Kotschy. The catalogue in Vienna lists a single specimen and the card catalogue in 1997 lists this fish as the holotype.

Karaman (1971) places Barbus (= Luciobarbus) barbulus, Luciobarbus schejch and fish Heckel (1843) referred to Luciobarbus mystaceus (Pallas, 1814) as synonyms of B. rajanorum. Berg (1949) also places L. mystaceus of Heckel in B. rajanorum. However mystaceus of Pallas would have priority (authors such as Karaman (1971) and Almaça (1983, 1991) refer the species description to Heckel (1843) in error) but, as Berg (1949) points out, Pallas's Cyprinus mystaceus is partly Barbus (= Luciobarbus) mursa and Barbus (= Luciobarbus) capito. Almaça (1983) recognises Barbus mystaceus with two subspecies, mystaceus from Aleppo, Tigris at Mosul and the Euphrates and barbulus (see above under this latter species). Krupp (1985c) places Barbus (= Luciobarbus) barbulus and Heckel's Luciobarbus mystaceus in Barbus pectoralis.

I am uncertain as to the identity of Barbus mystaceus (Pallas, 1814) reported by Heckel (1843b) from the "Tigris bei Mossul", Iraq, in regard to Iranian Luciobarbus species and do not assign any Iranian specimens collected by me to it. F. Krupp (in litt., 1987) considers Heckel's mystaceus to be identical with B. barbulus but that Heckel's mystaceus differs from that of Pallas, as previously noted by Berg (1949). Heckel's B. mystaceus is most probably either B. barbulus or B. pectoralis.

Barbus schejch is recognised as a distinct species by Almaça (1983, 1991) but only one specimen, a syntype from the Tigris (in the Naturhistorisches Museum Wien, NMW 50399), was available to him. It measures 136.5 mm standard length. Two other specimens identified as syntypes of this this species are under NMW 54520 with standard lengths 175.4 and 270.7 mm. The barbels in the 50399 are very short, not reaching the eye and about equal in length while in the other two syntypes the posterior barbel reaches the mid-eye and the barbels are subequal. The lips are fleshy, like Luciobarbus barbulus, but there is no central lobe in 50399, present in the smaller of the two other syntypes and poorly developed in the larger. The complete dorsal fin spine bears 29 teeth in the 50399 and 29 or 35 in broken spines of the other two fish. Gill rakers number 22 in 50399 and 16 or 18 in the other two fish. Lateral line scales number 52 (or 54 to end of scale row on caudal fin) in the syntype and 57(58) or 58(60). Main row pharyngeal teeth are 4-4 in 50399, missing in the other two fish. These data are somewhat contradictory and further data are required to resolve the status of this nominal species. The catalogue in Vienna lists 4 fish in spirits and 4 fish stuffed.

The synonymy of Luciobarbus barbulus with L. pectoralis remains uncertain. The putative holotype of B. pectoralis (NMW 54474) was compared with a specimen of similar size from Iran referred to L. barbulus (CMNFI 1973-0393). The L. pectoralis specimen is partly dried so direct measurement comparisons are not possible. The L. pectoralis specimen has more teeth in the dorsal fin spine (27 teeth even though it is broken off, much more than 30 presumably in the intact spine), barbels in pectoralis are shorter, the posterior one reaching the anterior half of the eye, the anterior one short of the mouth angle, mouths similar in shape but lips appear to be less fleshy, gill rakers number 16, lateral line scales number 44, and 4 main row pharyngeal teeth but there is a trace of a fifth tooth not fully ossified. ? check counts on NMc fish?

Key characters

The dorsal spine is much stronger than in Luciobarbus barbulus and arises from an elevated base that supports the dorsal fin base. The body is deeper than in L. barbulus and the lips usually less fleshy.

Morphology

Dorsal fin with 4 unbranched and 7-9 branched rays (7 in the holotype, usually 8), anal fin with 3 unbranched and 5, rarely 6, branched rays. Pectoral fin branched rays 16, pelvic fin rays 8. Lateral line scales 42-60 (44 in the holotype; 42 in Barbus perniciosus). Gill rakers 14-17 (to 21 if Barbus schejch is included (Almaça, 1986)). Pharyngeal teeth 2,3,4-4,3,2 in 16 fish, 2,3,5-5,3,2 in 9 fish examined by Krupp (1985c), rarely 2,3,5-4,3,2 or 2,3,4-5,3,2 (1 fish each), spoon-shaped or pointed with the fourth tooth of the inner row large and globose. Larger fish usually have 4 teeth in the main row and the fourth tooth is globose. Smaller fish with 5 teeth in the main row have cylindrical teeth. All intermediates stages exist (Krupp, 1985c). The mouth is moderate in size and subterminal. Lips are thin to moderate and the median lobe of the lower lip may be present or absent. Barbels are thin to moderate, the anterior one not extending back beyond the nostril to anterior eye margin level and the posterior one not beyond the middle to the posterior margin of the eye. The last unbranched dorsal fin ray is moderate to very strong with a low density of denticles extending along much of the ray. Larger specimens have a lesser extent of denticles along the ray. The body form is extremely variable.

Sexual dimorphism

Nuptial tubercles develop on 88.9% of mature males (Ghafari and Jamili, 2010).

Colour

The back is brown to bluish-green and the flanks yellowish to silvery-white.

Size

46.9 cm standard length (Krupp, 1985c).

Distribution

Tigris-Euphrates basin and the Orontes and Quwayq rivers. In Iran, it is found in the Tigris River basin in the Hawr Al Azim and the lower Karkheh, Karun and Jarrahi rivers, in the Kor River basin, and in the Gulf basin in the middle and lower Helleh, middle and lower Mand and Dasht-e Palang rivers (Abdoli, 2000).

Zoogeography

Almaça (1984b) considers that the origin of this species (as Barbus schejch) lies with a group that migrated southwards in the late Pliocene from the Dacian Lake of the Sarmatian Sea and speciated in Mesopotamia and later (Almaça, 1991) that this species (as Barbus (= Luciobarbus) pectoralis) originated from a colonisation wave from South Europe.

Habitat

Unknown.

Age and growth

Al-Rudainy (2008) states that in Iraq it reaches sexual maturity in 4 years at 30 cm length and 350 g weight.

Food

Unknown.

Reproduction

Ghafari and Jamili (2010) sampled fish from the Karun River and found the breeding season was from January to February at 14.3ºC and pH 7.75. Eggs were released in shallow water over gravel or sand in a single batch. Males matured at 3 years with LM50 35-40 cm and females at 4 years with LM50 50-55 cm. Absolute fecundity range was 7144-332,196 eggs and relative fecundity was 3845 to 164,753 eggs/kg. Maximum egg diameter was 2.0 mm in February. Sex ratio was 1:2 in favour of females.

Al-Rudainy (2008) gives the spawning season in Iraq as March to April on gravel in shallow water with a strong current.

Parasites and predators

Masoumian et al. (2008) recorded the myxosporeans Myxobolus karuni and M. persicus from gills of fish captured in the Karun and Karkeheh rivers and Shadegan Marsh.

Economic importance

Occasionally caught and used for food. Ghafari and Jamili (2010) consider it suitable for aquaculture because of its good taste and size.

Conservation

Endangered in Turkey (Fricke et al., 2007). Confusion over its identity has made a conservation assessment difficult for Iranian waters.

Further work

The taxonomic status of this species and its presence in Iranian waters should be resolved.

Sources

Type material: See above, Barbus pectoralis (NMW 54474).

Luciobarbus subquincunciatus
(
Günther, 1868)

Common names

سليماني (= solimani or soleimani).

[abou khazzama, a'djzan, agzan or adzan, all in Arabic; black spot barb, leopard barbel, Mesopotamian barbel].

Systematics

The type locality of this species is unknown. Günther (1868) gives the following account:- "From the Collection of the East-India Company.- Although no record of the history of this specimen has been preserved, it is probable that it came from Mesopotamia, as other examples from this country are preserved in precisely the same manner". The type specimen is a "Skin, 15 inches long" (= 5.9 cm).

Krupp (1985a) removes this species from Bertinius Fang, 1943 since the enlarged molariform pharyngeal teeth on which this genus was erected are due to convergence and are not evidence of monophyly. Howes (1987) places this species in his Barbus sensu stricto.

Key characters

The numerous, large, dark spots arranged in an almost quincunx pattern are distinctive.

Morphology

Dorsal fin with 3-4 unbranched and 8 branched rays, anal fin with 3 unbranched and 5 branched rays. The last unbranched dorsal fin ray is very strong and bears denticles along almost its whole length or three-quarters of the length. Pectoral fin branched rays 14-18, pelvic fin rays 7. Lateral line scales 75-88. Scales have few radii on all fields, fine circuli and a focus slightly subcentral anterior. Total gill rakers about 10-13, broad based and triangular in shape with highly tubercular distal or foliose margin. The longest raker reaches the one below when appressed. Total vertebrae 45 (Howes, 1987). Pharyngeal teeth 2,3,3,-3,3,2, occasionally 2,3,4-4,3,2, the usual number of teeth in the inner row in large specimens being 3 (Krupp, 1985a). The third inner row tooth is the biggest by far and is molariform. Juveniles have 5 inner row teeth (Krupp, 1985c). Total vertebrae 46. The mouth is horseshoe-shaped, small and inferior. Lips are well-developed and fleshy. The median lobe of the lower lip is undeveloped. The barbels are thick. The gut has many anterior loops, the number increasing with size.

Meristics for Iranian material:- dorsal fin branched rays 8(1), anal fin branched rays 5(1), pectoral fin branched rays 14(1), and pelvic fin branched rays 5(1). Lateral line scales 83(1). Total gill rakers ?. Pharyngeal teeth 2,3,4-4,3,2 (1).

?others

Sexual dimorphism

Unknown.

Colour

The whole body, head, fins, barbels, lips and even eyeball are covered with dark spots about the same size as or larger than the eye. Some larger flank spots are 2-3 times the eye diameter. Spots on fins are elongated along the fin length. These spots are arranged in patterns similar to a quincunx, hence the species name. A quincunx comprises four spots, one at each corner of a square with the fifth spot in the middle of the square. Sometimes a spot runs into an adjacent one. Some spots below the lateral line may be elongate, three times longer than wide, and arranged vertically. Occasional fish lack spots on the mid-flank but are still distinctively spotted elsewhere. The overall colour is greenish to brownish-yellow with the belly white. Peritoneum dark brown to black.

Size

Reaches 33.8 cm total length (Menon, 1956), 45.7 cm (Khalaf, 1961) or 60 cm (Sauvage, 1884).

Distribution

Found in the Tigris-Euphrates basin including its Iranian portion in such rivers as the Jarrahi (Wossughi, 1978; Rainboth, 1981; Abdoli, 2000).

Zoogeography

Almaça (1991) believes that this species originated in Mesopotamia.

Habitat

Unknown but recorded from rivers and artificial reservoirs.

Age and growth

Şen et al. (1992) examined 9 fish in Keban Dam Lake, Turkey and found age groups 3-7, growth rings being best expressed in sectioned dorsal fin rays. The length-weight relationship was logw = - 5.78723 + 3.27533 logl and the mean K(TL) was 0.8234. The b value indicates the habitat is suitable for the species.

Food

The molariform pharyngeal teeth and evidence from gut contents showed this species is an obligate molluscivore (Krupp, 1985a). However, Al-Rudainy (2008) gives diet in Iraq as insects, as well as detritus and aquatic plants, but this may apply to younger fish.

Reproduction

Sexual maturity is attained at 3-4 years, 35 cm length and 500 g weight in Iraq. Spawning takes place in April and May with eggs deposited on rocks (Al-Rudainy, 2008).

Parasites and predators

None reported from Iran.

Economic importance

This species occasionally occurs in commercial catches in Khuzestan but is not a common food fish compared to other Barbus sensu lato species. It has been investigated for aquaculture in Khuzestan but fish are rare and so adults are caught and released.

Conservation

This species is now very rare in Iran and "critically endangered". Reports of 1 fish taken in the Gav Masiab River in 1991, 4 fish from the Karun River in 1995 and 1 fish from the Karun River at Ahvaz in 1997 were the only records for the 1990s (M. Ramin, pers. comm., 2000). The stock of this species in the Gav Masiab River is severely reduced and during 4 years of collecting in western Iran only one fish was caught (J. Valiallahi, www.modares.ac.ir, downloaded 4 July 2000; pers. comm., 2001; Valeolahy, 2000).

Syrian populations in the Euphrates River and parts of its tributaries are also in a parlous state (R. Beck, pers. comm., 2000).

Further work

The biology of this distinctive species should be investigated.

Sources

Type material: ?

Iranian material:- CMNFI 1993-0133, ?, ? mm standard length, (): ZMH 2506, 1, 308.0 mm standard length, Kermanshahan, Karasu-Gamasiab-Seymarreh, Kermanshah (); FMNH70794,1, ?, Javanarud near Kermanshah (); and market specimens from Khuzestan.

Comparative material:- CMNFI 1980-1036, 1, 177.5 mm standard length, Turkey, Keban Dam on Murat Nehri near Elazig (38º41'N, 39º14'E); CMNFI 1986-0676, 1, 283.0 mm standard length, Turkey, Keban Dam on Murat Nehri (no other locality data); BM(NH) 1874.4.28:15, 1, 415.3 mm standard length, Iraq, Tigris River near Baghdad (33º21'N, 44º25'E); BM(NH) 1875.1.14:3-5, 3, 377.6-468.2 mm standard length, Iraq, Tigris River (no other locality data); BM(NH) 1974.2.22:1353, 1, 253.4 mm standard length, Iraq, Sirwan River, Diyala (no other locality data);

Luciobarbus xanthopterus
Heckel, 1843

Common names

گطان (gatan or gattan).

[gattan or ghattan, nobbash, or thekar, all in Arabic; yellowfin barbel].

Systematics

Howes (1987) places this species in Barbus sensu stricto. Almaça (1983) briefly reviews the placement of this species in synonymy; most ichthyologists now regard it as a distinct species. Luciobarbus xanthopterus has been considered as a variant of L. schejch but differs in gill raker count (10-13 in xanthopterus, 21 in schejch) and main row pharyngeal tooth count (5 in xanthopterus and 4 in schejch) (Almaça, 1983; see also discussion under Luciobarbus pectoralis; Heckel (1843b), however, gives the main row count for xanthopterus as 4). It has also been considered as a synonym of esocinus (q.v.) (Almaça, 1986).

Fayazi et al. (2006) used mtDNA to study differentiation between populations of this species in the Karun, Karkheh and Jarrahi rivers in Iran. Diversity was low although the Karun and Karkheh fish grouped together, leading to the recommendation that fish from the Jarrahi should not be used to stock other river basins.

Almaça (1986) records syntypes of Luciobarbus xanthopterus from the type locality given by Heckel (1843b) "Tigris bei Mossul", Iraq in the Naturhistorisches Museum Wien under NMW 54841 (10 specimens, one large fish at 216.5 mm standard length and 9 smaller fish at 48.6-63.4 mm; one of these was noted as being listed as the lectotype in 1997 (? the largest), and the catalogue number was 54841a) and NMW 54786 (1 specimen, 292.8 mm, not listed as a type in 1997). Material listed under NMW 1843 (? catalogue number or collection date) may also be syntypes. Eschmeyer et al. (1996) list 1 dried syntype under NMW 91215. The catalogue in Vienna lists 4 fish in spirits and 2 fish stuffed.

Key characters

This species is characterised by two pairs of barbels, a serrated dorsal fin spine, small scales (57-68 in lateral line), and a subterminal to terminal and oblique mouth. The elongate postorbital length is also seen in Luciobarbus esocinus but is more marked in the latter (check this?) and scale counts are different (check?).

Morphology

Dorsal fin with 4 unbranched and 7-9, usually 8, branched rays, anal fin with 3 unbranched and 5 branched rays. Pelvic fin branched rays 8. Pectoral fin branched rays 14-18 (Jawad, 1975). Lateral line scales 57-68. Scales have rounded dorsal, ventral and posterior margins and an anterior margin with a central protuberance and indentations above and below. Circuli are fine and radii are found on the anterior and posterior fields and sometimes the lateral fields. The focus is subcentral anterior. There is no distinct pelvic axillary scale. Gill rakers 7-13, short and reaching the adjacent raker when appressed. Pharyngeal teeth 2,3,5-5,3,2, strongly hooked the fourth tooth of the inner row being the largest and anterior teeth being rounded with a small flat or concave grinding surface below the tip. Qasim and Niazi (1975) gave a tooth formula of 4,3,2-2,3,4, i.e. 2,3,4-4,3,2 as does Heckel (1843b) and teeth were molariform. Total vertebrae 44 (Howes, 1987), 40-42 (Qasim and Niazi, 1975), 42 (Wossughi, 1978) or 46 (BM(NH) 1973.5.21:198). The last unbranched dorsal fin ray is moderately to very strong, has a low denticle density and is serrate along much of its length. Barbels are thin, the maxillary barbels are longer than rostral barbels but both are short, the rostral ones not extending beyond the level of the nostrils and the maxillary ones not extending back beyond mid-eye to rear eye level. The gut has one anterior and two posterior loops in an elongate s-shape. The mouth is moderate in size, inferior and an elongate u-shape in young fish and as development progresses becomes terminal in adults (Karaman, 1971; Almaça, 1984b). Lips are thin to moderate and the lower lip has no median lobe.

Iranian specimens had the following meristics:- dorsal fin branched rays 8(2); anal fin branchd rays 5(2); pectoral fin branched rays 18(2); pelvic fin branched rays 8(2); lateral line scales 57(1) or 68(1); total gill rakers 7(1) or 10(1); pharyngeal teeth 2,3,5-5,3,2(2); and total vertebrae ?.

Sexual dimorphism

Unknown.

Colour

The body is without distinctive marks. The back is brownish to bluish-grey, the flanks silvery to silvery-yellow, and the belly white. The scales are outlined by melanophores. The overall colour from a marsh habitat is darker than from a riverine habitat, the pigment outlining scales being thicker for example especially at the scale base. The eye is red in marsh specimens, white to yellowish elsewhere (M. Al-Mukhtar, pers. comm., 1995). All fins are lemon-yellow to orange with some darker melanophores. The unbranched dorsal fin rays and the uppermost caudal fin rays are black.

Two small specimens from Iran have irregular spots and blotches on the flank. The peritoneum is silvery with melanophores developed dorsally.

Size

Al-Hassan et al. (1986) report a specimen 1.5 m total length and 8.6 kg from the Abu Al Khasib area in the Shatt al Arab, Iraq.

Distribution

This species is found in the Tigris-Euphrates basin including its Iranian portion such as the lower Karun River and adjacent lower reaches of the Jarrahi River (Abdoli, 2000), the Qareh Su at Kermanshah, the Karkheh River and Hawr al Azim.

Zoogeography

Almaça (1984b, 1991) considers that the origin of this species lies with a group that migrated southwards in the late Pliocene from the Dacian Lake of the Sarmatian Sea and speciated in Mesopotamia.

Habitat

van den Eelaart (1954) and Al-Hamed (1966b; 1972) describe the habitat for this species in the Tigris River as distributed in the deep, open waters of lakes and vegetated marshes and to a lesser extent in the river and its tributaries. Mature fish move upstream to the spawning grounds in February-March and spent fish descend to their original habitat in lakes and marshes. In summer, beginning in June, under low water level conditions and high temperatures, the smaller fish remain in the deepest depressions of lakes but the large fish (3 kg or more) migrate up rivers and tributaries in search of cooler water, returning in September and October when temperatures fall to fatten over winter.

In Khuzestan, this species is most abundant in the Karkheh River in March and in the Hawr al-Azim in December, migrating from the wetland to the river in spring. Younger fish are more abundant in the wetland and older fish in the river (Iranian Fisheries Research Organization Newsletter, 22:3, 2000; Eskandary et al., 2000). Another study showed this species to be most abundant in the Karkheh River in December, with a migration from wetlands in spring to the main river (Tehran Times, 1 October 2000).

Age and growth

Life span is at least 11 years (Al-Ahmed, 1966a). Al-Hamed (1966b; 1972) working on Tigris River fishes found males to mature at about 43 cm and females at about 48 cm, maturity being attained in the fourth year of life and spawning occurring at the beginning of the fifth. Some fish mature at age group 3 and some as late as age group 5. Males outnumber females on the spawning grounds, comprising 62% of the population. Tigris River and Al-Tharthar reservoir fish in Iraq had 7 age groups with growth good in the first three years and slower thereafter (Ali, 1979). In Keban Dam Lake, Turkey, age determination was best made on sectioned dorsal fin rays (of scales, otoliths, vertebrae and opercula) and up to 9 age groups were detected (Duman and Şen, 1995).

In the Karkheh River, male fish are mature at 151-200 mm (one year old) and females at 501-550 mm (3 years old). The sex ratio is 1:1.31 for males:females but this is not significantly different from 1:1 (Eskandary et al., 2000).

Food

Al-Hassan et al. (1986) report isopods and molluscs. Al-Hamed (1965) considers this species to be an omnivore, consuming filamentous algae, detritus, frogs, molluscs and fishes and even planktonic organisms. Organic matter is obtained in periods of food shortage by engulfing mud from the pond bottom. van den Eelaart (1954) reports food to be plants, epiphytes and plankton. In cold winters they take no food. Ali (1979) for Iraqi waters gives insects and plankton as the principle foods. Al-Shamma'a et al. (2009) found fish from Lake Habbaniyah, Iraq to feed mostly on animal materials (76.25) including molluscs and insects and their larvae. Feeding was most active in spring but high feeding intensities were also observed in autumn.

In Khuzestan it is omnivorous, feeding mainly on insects and vegetation, but also taking secondarily shrimps, snails and ostracods (Iranian Fisheries Research Organization Newsletter, 22:3, 2000; Eskandari et al., 2003). In the Karkheh River food is insects and vegetation mainly, with shrimps, gastropods and ostracods secondary food choices (Tehran Times, 1 October 2000). The intestine fullness is greater in fish in the Hawr al Azim, less in the Karkheh River which is used mainly for spawning (Eskandari et al., 2003).

Reproduction

van den Eelaart (1954) and Al-Hamed (1966b; 1972) studied the reproduction of this species in Iraq. Eggs are deposited on fine gravels overlying a layer of coarse sand in shallow, wide holes excavated by the fish. Water depth varies from 30 to 150 cm. Egg diameter is 1.0 mm and fecundity up to 340,000 grey eggs. Al-Hassan et al. (1986) record up to 350,000 eggs for their large fish from the Shatt-al-Arab. The spawning season on the Tigris River between Beled and Tigrit is April and May. Fish appear on the spawning grounds in schools just before dark and remain there until shortly before midnight, making loud noises by splashing, jumping and chasing.

In Khuzestan, spawning fish are 63.7-80.0 cm total length with a relative fecundity of 18.9-142.5 eggs/g body weight and a minimum and maximum absolute fecundity of 136,924 and 549,211 eggs (Iranian Fisheries Research Organization Newsletter, 22:3, 2000; Eskandari et al., 2003). In the Karkheh River, spawning took place at surface water temperatures of 25.5-28.65°C in turbid water after a spring migration from wetlands into the river (Tehran Times, 1 October 2000; Eskandary et al., 2000). Spawning occurs annually in May and June in the Karkheh River and maximum egg diameter is 2.25 mm (Eskandary et al., 2000).

Parasites and predators

Bykhovski (1949) reports a new species of monogenetic trematode, Dactylogyrus inutilis, from this species in the Karkheh River, Iran. Ebrahimzadeh and Nabawi (1975) list Anisakidae from this species in the Karun River. Moghainemi and Abbasi (1992) record a wide range of parasites from this species in the Hawr al-Azim in Khuzestan. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Argulus sp., Ergasilus sp. and Lamproglena compacta on this species.

Economic importance

This species appears regularly in the markets of Ahvaz and Wossughi (1978) states it is of great economic importance. Sharma (1980) reports that gattan is the third most important fish species at Basrah fish market, accounting for 510,503 kg for the period from October 1975 to June 1977. Petr (1987) reports the annual catch for 1976 in Iraq was 2543 t. This species has been studied for pond culture in Khuzestan where over 95% of young survived, using hormones to stimulate reproduction (Iranian Fisheries Research Organization Newsletter, 28:3, 2001). Zadeh et al. (2009) investigated  the optimal dietary carbohydrate to lipid ratio for fingerlings of this species. Mortezavizadeh et al. (2009) induced reproduction in this species with carp pituitary extract. Sperm production was increased, 86% of females responded positively, 480 eggs/gram were produced and mean survival rate was 77.95%. Propagation was best at the beginning of March at 19.0-24.5°C.

englers in Iraq catch this fish which will reject any bait showing resistance, requiring a fast strike at the first indication that the fish has taken the bait.

Conservation

Several hundred thousand gattan juveniles have been introduced into the Hawr al Azim in Khuzestan in order to restock and protect this resource. The fish were artificially bred from breeders using hormone treatment (Iranian Fisheries Research Organization Newsletter, 39:3, 2004; Network of Aquaculture Centres in Asia, downloaded 11 January 2007). Endangered in Turkey (Fricke et al., 2007).

Further work

?

Sources

Type material: ?

Iranian material: ZMH 4071, 1, 151.6 mm standard length, Kermanshahan, Qareh Su at Kermanshah (?). uncatalogued, 2, 93.7-112.8 mm standard length, Kermanshahan, Sarab-e Yavari (34º28'N, 46º56'E);

Comparative material: BM(NH) 1893.6.23:25, 1, 198.0 mm standard length, Iraq, Al Faw (29º58'N, 48º29'E); HL51.5, Postorb 26.6, for body proportions; g.r 13; BM(NH) 1973.5.21:197, 1, 227.8 mm standard length, Iraq, Shatt al Arab (no other locality data) HL 60.3, postorb 28.8, g.r. 21;  BM(NH) 1973.5.21:198, 1, 143.0 mm standard length, Iraq, Shatt al Arab (no other locality data). HL 34.6, postorb 15.1, g.r. 18.

Genus Mesopotamichthys
Karaman, 1971

?

Much of the past literature on this genus appeared under Barbus (q.v.)

 

Mesopotamichthys sharpeyi
(Günther, 1874)

Common names

بني (benni, benny, beni, binni, bini, binny (meaning possibly nose)), سليماني (= solimani or soleimani, meaning unknown).

[binni, bunni, bunnei, bunia or aradah, all in Arabic].

Systematics

Howes (1987) places this species outside the genus Barbus sensu stricto as defined by him because it has the non-elongate lachrymal bone with a sensory canal running along the antero-dorsal border, a derived condition. Karaman (1971) erected a new genus for this species, Mesopotamichthys, which is not accepted here nor by Krupp (1985c). However Bănărescu (1997) and Ekmekçi and Banarescu (1998) recognise this genus as valid. The large-scaled "Barbus" of Southwest Asia require a detailed revision probably with additional, molecular characters (see also under Carasobarbus luteus and Kosswigobarbus kosswigi).

Barbus faoensis Günther, 1896 described from "Fao (Persian Gulf)", Iraq is a synonym; Karaman (1971) retains it as the subspecies of the lower part of the Tigris-Euphrates basin with the type subspecies in the upper part of the Tigris River basin.

Barbus sharpeyi was described from "Baghdad". The three syntypes are in the Natural History Museum, London (BM(NH) 1874.4.28:20 labelled "R. Tigris nr. Baghdad. Sharpey", well sealed in its jar and not measured accurately, and BM(NH) 1874.4.28:27 and BM(NH) 1875.1.14:16 labelled "R. Tigris. Sharpey" and measuring 147.6-178.0 mm standard length). The large holotype of Barbus faoensis is also there, strongly sealed in its jar, labelled "Persian Gulf. Kurrachee Museum" (BM(NH) 1888.5.17:4).

Al-Mukhtar and Al-Hassan (1999) describe a hybrid of this species and Carassius auratus from Al-Hayei (= Al Ha'i), a seasonal lake between the Karkheh and Dez rivers in Khuzestan.

Key characters

The absence of barbels, the last dorsal fin unbranched ray moderately ossified but lacking teeth, and the low scale count are characteristic.

Morphology

Dorsal fin with 4 unbranched and 7-9, usually 8, branched rays, anal fin with 2-3 unbranched and 4-5, usually 5, branched rays, pectoral rays 13-19, usually 16-17, and pelvic fin branched rays usually 8. The last third or quarter of the last unbranched dorsal fin ray is thin, flexible and tapering. The gut has several loops, two anteriorly and three posteriorly. Pharyngeal tooth formula is 2,3,5-5,3,2, sometimes with only 4 teeth in the main row but the anterior tooth is missing in both small and large fish and so does not appear to be age related, with teeth hooked at the tip but not strongly on the posterior teeth which are spoon-shaped with the hollow of the spoon filled in with bone. Total gill rakers number 13-19, reaching the raker below or just beyond when appressed. Total vertebrae 38-42 (lower values, 38-39, may not include hypural plate). Lateral line scales 29-37. A small pelvic axillary scale may be present or scales in this area may be so weakly developed as not to be an apparent axillary scale. Scales have a slightly anterior focus, fine concentric circuli, many radii on all fields and the posterior, exposed field bears numerous small tubercles. The mouth is slightly subterminal. Lips are well-developed but not fleshy and the lower lip is interrupted in the middle. Microscopic studies of the pharynx and oesophagus have been carried out by Alboghobeish and Moosavi (1998) who confirm that it is adapted for herbivory. Chromosome number 2n=98 (Balasem et al., 1994). Alboghobeish and Hamidian (2006) studied the distribution of alarm cells in the skin of this species.

Iranian fish have the following meristics: branched dorsal fin rays 8(2), branched anal fin rays 5(2), branched pectoral fin rays 16(2) and branched pelvic fin rays 5(2). Lateral line scales 30(1) or 31(1). Total gill rakers 16(1) or 18(1). Pharyngeal teeth 2,3,5-5,3,2(2). Total vertebrae 41(1) or 42(2) based on CMNFI 79-0087 (42) and 87-0017 (41 and 42). 41(3) or 42(2) based on BM(NH) 1920.3.3:71-75, 42 (1973.5.21:195), 41 (1973.5.21:196), 42 (1874.4.28:27), 41 (1875.1.14:16)

Sexual dimorphism

Unknown.

Colour

Overall colour is greenish to light brown or golden brown with the belly white to silvery or yellowish-brown. Scales on the back and uppermost flank have solid dark brown pigment on the exposed part of the scale. The scale edge is thinner and so appears lighter. The eye is brownish orange, golden or silvery. Fins are darker than the adjacent body, a deep reddish-brown, with melanophores on rays and membranes in preserved fish. The peritoneum is black.

Size

Attains 55 cm and 4 kg (van den Eelaart, 1954; Al-Hamed, 1966b; 1972). Reaches at least 3.5 kg in Khuzestan (J. Gh. Marammazi, pers. comm., 1995).

Distribution

This species is found in the Tigris-Euphrates River basin including its Iranian portion in such marshes as the Hawr Al Azim and in rivers and in the northern Gulf basin in the Zohreh River (Marammazi, 1995; Abdoli, 2000).

Zoogeography

Karaman (1971) considers that this species originated from the Indian line of the Torini, a tribe of Cyprinidae, in which Karaman includes such genera as Carasobarbus and Kosswigobarbus, Garra, Hemigarra (recognised as Hemigrammocapoeta here) which have Iranian members.

Habitat

van den Eelaart (1954) and Al-Hamed (1966b; 1972) report some movement from lakes and marshes, from the end of February to the beginning of March, to rivers in the Tigris-Euphrates basin of Iraq during floods for about 3 weeks. There is a return to lakes and marshes for spawning in mid-March to mid-April. However, most fish remain in marshes and lakes for most of the year, in overgrown areas avoiding open water. Low water levels and high temperatures in the lakes and marshes may cause a migration to their deepest parts or into the lower reaches of the main and more permanent rivers. This species is less tolerant of low oxygen than Luciobarbus xanthopterus which probably accounts for them not being caught together in any number. Arzi et al. (2009) compare organochlorine residues in this species in three cities in Khuzestan.

Marammazi (1994) considers this species to be stenohaline and so restricted in its distribution in the Zohreh River which drains to the northern Persian Gulf. The influence of salinity on growth rate is examined by Orian et al. (1993).

Age and growth

Al-Hakim et al. (1976) studied some aspects of the biology of this species in Razzaza Lake, Iraq. Females are longer and heavier than males at advanced ages. Life span of females is 9 years and for males 8 years. Maturity starts in the third year at 32-35 cm total length. Males mature earlier than females. Al-Hamed (1966a; 1966b; 1972) found Tigris River fish in Iraq to mature at 25 cm for males and 28 cm for females in the second year of life and spawning took place early in the third year. A few matured in age group 1 and some as late as age group 3. Males are somewhat more abundant than females on the spawning grounds, averaging 57.4% of the fish caught. Maximum age is 6 years. Ali (1982b) found this species to mature in the fourth year of life in Iraq, with growth better in the marshes than in Tharthar Reservoir. Epler et al. (1996) found fish up to age 6+ years in fresh and salty Iraqi lakes. Nasir et al. (1989) reports on the biology of this species in the Al-Hammar Marsh, Iraq and found a sex ratio of 1 female:3 males for all months and length groups caught. No explanation for this skewed ratio was found.

Food

Al-Hamed (1965) found this species to be strictly herbivorous, feeding on unicellular Chlorophyceae, diatoms and filamentous algae when young and on higher plants and detritus when older. Nasir et al. (1989) and Epler et al. (1996) confirm that this species in Iraq is completely herbivorous although some copepods and molluscs are taken, most probably incidental to filamentous algae, diatoms and detritus. van den Eelaart (1954) reports feeding even in cold winters. In the Karun River, diet includes such plants as Potamogeton, Salvinia, Nuphar and Phragmites (Annual Bulletin 1993-94, Iranian Fisheries Research and Training Organization, Tehran, p. 91-92, 1995).

Reproduction

van den Eelaart (1954) and Al-Hamed (1966b; 1972) studied reproduction in this species on the Tigris River in Iraq and Al-Nasih (1992) in fish ponds. Spawning occurs chiefly in lakes and marshes, with some spawning in the lower reaches of rivers. Eggs are deposited on submerged, or partially submerged, vegetation, from the surface down to about 1 m depth. Eggs are large, yellow and measure up to 1.7 mm in diameter and number up to 158,000. Epler et al. (1996) give a relative fecundity of 10,021 to 28,471 eggs for fish 4+ to 6+ in age from Iraqi lakes with fish spawning in April in a freshwater lake and February/March in a saline lake. Al-Nasih (1992) gives details of larval development. The spawning season in Lake Saniyah just north of Amara is March and April, with some ripe fish caught in May. Fish appear on the spawning grounds about sunset and left before darkness is complete. They return in the early morning and leave again at about 0800 hours. These fish chase each other, dart about singly or in pairs and sometimes come to the surface and splash. Al Mukhtar et al. (2006) investigated this species in the Hawizah Marsh as a source of spawners for aquaculture. Ripe eggs appeared in January and 25% were running in February and 30% in March. Half of the fish were spent in April. The spawning migration was lead by males in October and December with females increasing rapidly in February. Males disappeared in April. Absolute fecundity reached 236,160 eggs. Al-Rudainy (2008) gives an absolute fecundity of up to 358,343 eggs in Iraq or up to 145 eggs/g body weight and a diameter of 2.0 mm

Petr (1987) reports spawning in Iran at 15-16°C in February in clean water of rivers with sandy bottoms. In the Karun River this species spawned in March-April in river estuaries (Annual Bulletin 1993-94, Iranian Fisheries Research and Training Organization, Tehran, p. 91-92, 1995). A specimen caught in March had well-developed testes. Spawning in Shadegan Marsh, Khuzestan is in March and in branches of the Karkheh River in March to April (Al Mukhtar et al., 2006). Shadegan Marsh is one of the most important spawning areas in Iran (Mohammadi and Marammazi, 2001).

Parasites and predators

Bykhovski (1949) reports a new species of monogenetic trematode, Dactylogyrus pavlovskyi, from this cyprinid in the Karkheh River, Iran. Ebrahimzadeh and Nabawi (1975) list species of the protozoans Trichodina and Myxosoma and the trematode Dactylogyrus as well as the nematode Camallanus lacustris, from this species in the Karun River. Jalali and Molnár (1990a) record two monogenean species, Dactylogyrus spp., in the Dez River and Molnár and Jalali (1992) a new species of monogenean, Dogielius persicus, from this species in the Dez and Karun rivers of Khuzestan. Gussev et al. (1993b) record Dactylogyrus pavlovskyi in the Dez River.

Masoumian et al. (1994) describe a new species of Myxosporea from the gills of this species in the Karun River, namely Myxobolus persicus, and later (Masoumian et al., 1996) another new species of Myxosporea, Myxobolus nodulointestinalis, in the gut lining of this species and also from rivers of southwestern Iran. Masoumian et al. (1996) describe a new species of Myxosporea, Myxobolus bulbocordis, from the heart of fish caught at various localities in Khuzestan. Molnár et al. (1996) report additional new species from this fish in Khuzestan, namely Myxobolus iranicus in the spleen and Myxobolus sharpeyi in the gill cartilage. Myxosporeans are potentially dangerous to fishes such as Mesopotamichthys sharpeyi which may be used in fish culture in Khuzestan. Masoumian and Pazooki (1999) list Myxobolus persicus, M. karuni, M. sharpeyi, M. nodulointestinalis, M. bulbocordis and M. iranicus from this species in various localities in Khuzestan. Shamsi et al. (2009) found Dactylogyrus nchoratus in this species from fish farms and the Karun River.

Peyghan (1994) reports ichthyophthiriasis in cultured Barbus (= Mesopotamichthys) sharpeyi in Khuzestan. This parasite causes severe skin and gill damage and mortality reaches 80%. A combination of formalin and malachite green with transfer of fish to another pond having a better environment cured the condition.

Molnár and Pazooki (1995) record philometrid nematodes from this species in the Karun River, and these are presumed to be a new species. Pazooki and Molnár (1998) later describe Philometra karunensis as the new species from the swimbladder and adominal cavity of this fish.

Moghainemi and Abbasi (1992) record a wide range of parasites from this species in the Hawr al-Azim in Khuzestan. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Argulus sp. and Ergasilus sieboldi on this species.

The monogeneans Dactylogyrus povlovskyi, D. barbioides, D. anchoratus, D. carassobarbi and Dogielius persicus are recorded from this species in the Karun River with heavier infestations in spring and summer than in autumn and winter. These gill parasites caused no serious injuries but were thought to be important in respect of monitoring infestation levels on fish farms in Khuzestan (www.avz1.8m.com/fulltext.htm, downloaded 28 October 2002). Papahn et al. (2004) record the monogeneans Dactylogyrus pavlovskyi, D. barbioides, D. anchoratus, D. carassobarbi and Dogielius persicus from this species in the Karun River at Ahvaz. Mokhayer et al. (2006) collected this species from four sites in Shadegan Marsh and found Dactylogyrus anchoratus, D. carassobarbi and Dogelius persicus. The first species had more parasites in the right gill compared to the left gill, the second had more in the upper holobranch and the  third more in the lower holobranch. Barzegar et al. (2008) record the digenean eye parasite Diplostomum spathaceum from this fish.

Economic importance

This species is second in importance after sobour (Tenualosa ilisha) at the Basrah fish market in Iraq with a weight from October 1975 to June 1977 of 772,775 kg. Nasir et al. (1989) record a total catch for Iraq of 5000 tonnes per year and Petr (1987) for Iraq in 1976 a catch of 4243 t. Young (1976) noted that this species was regarded as the tastiest fish available from the marshes of Iraq.

Petr (1987) has suggested investigating fish farming of this species in Khuzestan and Al-Nasih (1992) carried out such an investigation for Iraq (see below). The Khuzestan Fisheries Research Centre at Ahvaz has successfully bred this species in pond and pen culture using hormone stimulation of broodstocks (Emadi, 1993a; Iranian Fisheries Research and Training Organization pamphlet; Iranian Fisheries Research and Training Organization Newsletter, 5:2, 1994; Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 49, 1996; Mohammadian et al., 2009) and in polyculture with Chinese carps such as Ctenopharyngodon idella (Annual Bulletin 1993-94, Iranian Fisheries Research and Training Organization, Tehran, p. 93-94, 1995; Annual Report, 1995-1996, Iranian Fisheries Research and Training Organization, Tehran, p. 36-37, 1997). Private companies also culture this species in Khuzestan. In Khuzestan, over 95% of young survived, using hormones to stimulate reproduction. Khajeh et al. (2008) have examined haematological parameters in cultured fish and found some to be higher than in Ctenopharyngodon idella. Mortezazadeh et al. (2009) studied the effects of propofol as an anaesthetic on cultured fish.

Yazdipour et al. (1991) give a report on propagation of this species in Iran.

Sharifian (2000) gives details of whole body analysis. The highest protein content was in the 30-95 mm and 100-140 mm length groups.

Conservation

Local fishermen in Khuzestan believe numbers of this species declined in the Shadegan marshes after young Hypophthalmichthys molitrix from the Caspian were released. As a food fish, its population biology should be monitored in Khuzestan. Several hundred thousand juveniles have been introduced into the Hawr al Azim in Khuzestan in order to restock and protect this resource (Network of Aquaculture Centres in Asia, downloaded 11 January 2007) and Mohammadian et al. (2009) list production of 1-5 million 1-2 g fry used to restock Hawr al Azim and Shadegan March annually.

Further work

Al-Nasih (1992) investigated the use of this popular food fish for aquaculture in Iraq. Although its growth rate is slower than in Cyprinus carpio, a popular fish for aquaculture, its plankton feeding makes it adaptable to pond life without competition with Cyprinus carpio, it has tasty flesh, reaches 2 kg, and has a relatively high fecundity. Hormonal injections with hypophysial extract from the more readily available Cyprinus carpio induced breeding in this species. Natural production can be increased to 450-600 kg/ha with the use of mineral fertilizers in ponds to stimulate plankton growth. The biology of this species has been investigated in Khuzestan with a view to aquaculture (Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 6, 1996).

Sources

Type material: See above, Barbus sharpeyi (BM(NH) 1874.4.28:20, BM(NH) 1874.4.28:27, BM(NH) 1875.1.14:16), Barbus faoensis (BM(NH) 1888.5.17:4).

Iranian material:- CMNFI 1979-0087, 1, 228.0 mm standard length, ? (); CMNFI 1991-0154, 277.8 mm standard length, ?; material observed on market stalls in Ahvaz, Khuzestan.

Comparative material:- CMNFI 1987-0017, 146.0-175.4 mm standard length, Iraq, vicinity of Basrah (no other locality data); BM(NH) 1920.3.3:71-75, 17, 58.7-115.0 mm standard length, Iraq, Basrah (30º30'N, 47º47'E); BM(NH) 1920.3.3:76-77, 1, 261.4 mm standard length, Iraq, Basrah (30º30'N, 47º47'E); BM(NH) 1922.5.24:1, 1, 113.5 mm standard length, Iraq, Basrah (30º30'N, 47º47'E); BM(NH) 1973.5.21:195, 1, 185.5 mm standard length, Iraq, Shatt-al-Arab (no other locality data); BM(NH) 1973.5.21:196, 1, 186.1 mm standard length, Iraq, Shatt-al-Arab (no other locality data).  

Genus Mylopharyngodon
Peters, 1881

This genus comprises a single species so the characters of of the genus are the same as for the species.

Mylopharyngodon piceus
(Richardson, 1846)

Common names

None.

[black carp, snail carp, black amur, Chinese black carp, Chinese roach, black Chinese roach]

Systematics

This species was originally described from Canton, China.

Key characters

The pharyngeal teeth are strong, grooveless molars in 1-2 series and the overall colour is black. distinguishing it from the similar grass carp, Ctenopharyngodon idella.

Morphology

The mouth is terminal to slightly subterminal with thin lips. Dorsal fin origin slightly in advance of level of pelvic fin origin. Dorsal fin with 3 unbranched and 7-9 branched rays, anal fin with 3 unbranched and 7-10 branched rays, pectoral fin branched rays 16, pelvic fin branched rays 8, and lateral line scales 39-46. Gill rakers short and stout, 14-23. Total vertebrae 36-41. The molar pharyngeal teeth are very strong and in  one or two rows, commonly 1,4-4,1 or 4-5. Intestine short, 1-2 times total length. The chromosome number is 2n = 48. Artificial hybrids with Ctenopharyngodon idella, Cyprinus carpio, Hypophthalmichthys molitrix and H. nobilis have been bred.

Meristics on Iranian material from Abbasi (2003): Dorsal fin branched rays 7(2), anal fin branched rays 8(2), lateral line scales 41(1) and 42(1), gill rakers 17(1) and 19(1), and pharyngeal teeth 4-5 (1) and 5-5(1).

Sexual dimorphism

Nuptial tubercles are present on the operculum, interorbitally, pectoral fin rays and on the scales of males.

Colour

The overall colour of the body and fins is black to blackish-brown or blue-grey. Fins are characteristically dark but may be somewhat lighter than the body, a blackish-grey, but darker than fins usually are in cyprinids. Scales have dark edges conferring a cross-hatched appearance. The lower head surface may be whitish and the belly greyish-white.

Size

Reaches 2.0 m and 106.0 kg.

Distribution

The native distribution is in East Asia from Amur River basin to south China and perhaps Vietnam but it has been introduced worldwide in suitable waters as a farm fish. The Food and Agriculture Organization web site "Database on Introductions of Aquatic Species" lists this species as introduced to Iran for aquaculture and research from China in 1992. It is reported from the Karakum Canal and the Kopetdag Resevoir in Turkmenistan (Shakirova and Sukhanova (1994) and Sal'nikov (1995) and may eventually reach the Tedzhen River basin in Iran. It is recorded from the Aras River in Armenia (Gabrielyan, 2001) and may eventually be recorded from the Iranian reach of this river. A large specimen from Gilan in the Caspian Sea basin shown me in Tehran in December 2000 appeared to be this species but was frozen solid so certain key characters could not be examined. The Iranian Fisheries Research and Training Organization Newsletter (27:2, 2001) has an illustration of this specimen which weighed 7.5 kg and was 96 cm long. Abbasi (2003) gives the  length of this fish as 97 cm total length and a capture date of October 2000 in a beach seine near Bandar Anzali coastal waters. In March 2001 a fish 80 cm total length and weighing 4.8 kg was caught in the same general area. Shamsi et al. (2009) record it from fish farms in Gilan.

Zoogeography

An exotic species introduced to Iran.

Habitat

The main habitat is floodplain lakes and the lower reaches of rivers, down to about 10 m. It may also be found in canals and reservoirs. Fast flowing water is required for reproduction. Temperature tolerances are similar to the grass carp, Ctenopharyngodon idella. Oxygen levels as low as 2 mg/l are tolerated.

Age and growth

Life span is possibly 20 years or more, with maturity attained at 3-11 years in various habitats. Fish introduced to the Karakum Canal showed a relatively rapid growth attaining 48 cm and 1.47 kg in their first year. At six years old fish were 125 cm and 29 kg.

Food

The principal food item of this species is molluscs, snails and clams, although apparently it does not eat large groups of zebra mussels. A 4-year-old fish can eat 1-2 kg of molluscs per day. The molar pharyngeal teeth crush the mollusc shells, the soft tissues are ingested and the broken shells spat out. Young fish eat zooplankton and aquatic insects and adults may take some crayfishes and other benthic invertebrates.

Reproduction

A stretch of river is required for drifting eggs to mature. before settling in quieter areas of floodplain lakes or the river itself. The length of river varies with temperature and velocity of the particular river. Adults migrate up rivers to spawn. Spawning starts at 26-36ºC, and perhaps as low as 18ºC, when rivers begin to rise in flood in spring and summer. Up to 1,180,000 eggs are produced annually in a single batch, more in hatchery fish where an average of 1.5-2.1 million eggs is cited. Swollen eggs have a diameter of 5.6 mm, swelling 4-5 times during hydration. Eggs hatch in 1-2 days.

Parasites and predators

Shamsi et al. (2009) found Dactylogyrus magnihamotus in this species from fish farms. Various Caspian Sea basin species are reported as predators including Esox lucius, Silurus glanis, Sander lucioperca, Aspius aspius, Perca fluviatilis and Lota lota.

Economic importance

None in Iran as not extensively used. This species is widely used on fish farms elsewhere to control parasite problems occasioned by molluscs acting as hosts for trematodes. Both diploid and triploid forms are used in farming. It is also important as a food fish.

Conservation

This species is an exotic and requires no conservation.

Further work

Additional field work is required to determine distribution and abundance in the wild as it can seriously affect native mollusc populations.

Sources

Iranian material: The specimen noted above.

Nico et al. (2005) summarise the biology of this species on which much of the above is based.

Genus Parabramis
Bleeker, 1865

Parabramis pekinensis
(Basilewsky, 1855)

Reported from the Karakum Canal and Kopetdag Reservoir in Turkmenistan (Shakirova and Sukhanova, 1994; Sal'nikov, 1995) as an exotic from China. May eventually be found in the Tedzhen (= Hari ) River and Caspian Sea basins of Iran. No Iranian record.

Genus Pelecus
Agassiz, 1835

The sabre carp genus contains only a single species found from the Baltic to the Black, Caspian and Aral Sea basins including Iran. The characters of the genus are the same as under the species.

Pelecus cultratus
(Linnaeus, 1758)

Pelecus cultratus from Bloch (1795-97) and Wikimedia Commons.

Common names

shamshir mahi (= scimitar or swordfish), shamshir mahi ab shirin (= freshwater swordfish), kuli.

[gilincbalig in Azerbaijan; chekhon' in Russian; sichel, ziege, razorfish, sabre carp, sabrefish].

Systematics

Pelecus cultratus was originally described from the Baltic Sea.

Pelecus cultratus kurensis Smirnov, 1943 is the Kura River basin subspecies but Berg (1948-1949) considered that other populations over the range of this species had been insufficiently studied to validate this subspecies.

Key characters

This species is easily recognised by the scaleless keel extending from the throat to the anal fin and the decurved and wavy lateral line.

Morphology

The body is elongate and strongly compressed. The mouth is almost vertical. The pectoral fins are long and curved, used for rapid manoeuvring when swimming normally and folded against the body when swimming rapidly. The lower lobe of the caudal fin is larger than the upper, with more rays and a stiffer ventralmost ray. The lower jaw is hooked in older specimens and has a tubercle which fits into an upper jaw notch. Gill openings are very wide with the branchiostegal membranes attached far forward, under the eye level. Muscles on the back extend forward to reach the anterior eye margin. Dorsal fin unbranched rays 2-3, usually 3, followed by 6-10, usually 7, branched rays, anal fin unbranched rays 2-3, usually 3, and branched rays 23-31, pectoral fin branched rays 13-17, and pelvic fin branched rays 6-8. Lateral line scales 88-120. Scales have extremely fine circuli, a central to subcentral posterior focus and very few posterior radii. Gill rakers number 15-26 (reaching the second or third raker below when appressed) and vertebrae 46-52. Pharyngeal teeth number 2,5-5,2, are narrow and are very strongly hooked at the tip with obviously serrate edges. Variants include 2,5-4,2, 2,4-5,2 and 2,5-5,3. The gut is an elongate s-shape. The chromosome number is 2n=50 (Klinkhardt et al., 1995).

Sexual dimorphism

Unknown.

Colour

The back is greenish and the flanks silvery. Fins are hyaline to grey, although the paired fins and the anal fin can be a bright yellow.

Size

Reaches 60 cm and 3.5 kg but most fish in the Volga-Caspian region are 80-180 g.

Distribution

Found in the drainages of the Baltic, Black, Caspian and Aral seas. This species is reported as rare in the lower Safid River (Derzhavin, 1934) and is found in the Anzali Mordab (Abbasi et al., 1999; Kiabi et al., 1999). Recorded by Abdoli (2000) from the middle to lower Safid River, Anzali Talab and adjacent Caspian coast. Recorded from the Atrak River but not yet from its Iranian reach (Reshetnikov et al. (1997). Also reported from the Karakum Canal and Kopetdag Reservoir of Turkmenistan (Shakirova and Sukhanova, 1994; Sal'nikov, 1995) and so may eventually be found in the Tedzhen (= Hari) River basin of Iran.

Zoogeography

A European and western Asian species with its origins in a Danubian or Sarmatian fauna.

Habitat

This species lives primarily in the sea but is also anadromous and may live permanently in larger tributaries. Jolodar and Abdoli (2004) note two forms in Iran, one resident in fresh water and a migratory form. It inhabits surface waters, aided by fin and mouth modifications for this mode of life (Adamicka, 1984). It migrates to the fresh water of large rivers to spawn. In the sea and larger rivers it occurs in schools.

Age and growth

Sexual maturity in the Volga River is attained at 3-4 years and a minimum length of 20 cm in males, 4-5 years and 22 cm for females. The largest immature males are 24 cm long, females 25 cm and the maximum age of non-spawners of both sexes does not exceed 6 years. The spawning stock is mostly fish 4-10 years old. Males have a slightly smaller maximum size than females and life span is at least 16 years (Sil'chenko, 1976). Spawning takes place first at age 2 in the Kura River of Azerbaijan. Growth is faster than in other populations, the Kura fish being the same size at age 2 as age 3 fish from the Don River of the Black Sea and age 4 fish of the Aral Sea.

Food

Food is taken by the vertical mouth from surface waters and includes insects and spiders. Young fish feed on zooplankton and even adults will do this if crustaceans are abundant. In the sea, various crustaceans are taken and these may be pursued near the bottom. Full grown Pelecus (50-60 cm) will capture fish such as Clupeonella, gobies (Gobiidae), Cyprinidae and even sticklebacks (Gasterosteidae) (Adamicka, 1984).

Reproduction

Spawning takes place in the latter half of May in the Kyubyshev Reservoir of the Volga River at 13.5-14.1°C or as high as 18-22°C. Water depths are 2.0-3.5 m. The main spawning takes place around sunset over a period of 24 days with a peak period of 10-12 days. Fecundity reaches 71,400 eggs. Eggs sink to the bottom but swell to an average diameter of 4.7 mm within an hour of fertilisation. Once swollen, any slight movement of the water will suspend the eggs in the water column. Eggs develop pelagically in floodplains, main rivers, side channels, bays and lakes but all these diverse habitats have a high oxygen content through flowing water or wind mixing. In the Volga flow rates are 0.28-1.1 m/sec on a sand-gravel bottom. Spawning may also take place in brackish water where the eggs float. Spawning in the Kura River of Azerbaijan takes place at the end of April and in May. The larvae are phototropic and active swimmers.

Parasites and predators

The Caspian seal, Pusa caspica, is a predator on this species (Krylov, 1984).

Economic importance

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in textbooks and for food. The scales contain silvery crystals of guanine which are extracted and used to make essence d'orient for artificial pearls. In the period 1909-1913, the catch in the Volga-Caspian region was more than 14 million fish annually. The flesh is fatty and bony and so this species is best smoked.

Conservation

Lelek (1987) classifies this species as intermediate to rare in Europe but fairly common in the Caspian Sea basin. Kiabi et al. (1999) consider this species to be critically endangered in the south Caspian Sea basin according to IUCN criteria. Criteria include sport fishing, few in numbers, habitat destruction, limited range (less than 25% of water bodies), absent in other water bodies in Iran, and present outside the Caspian Sea basin.

Further work

The distribution and biology of this species in Iranian waters needs study although it is probably too rare to be easily captured in numbers.

Sources

Meristics based partly on Wais (1995).

Iranian material: None available.

Comparative material: CMNFI 1971-0825, 1, 232.0 mm standard length, Czechoslovakia, Lake Ozirna (no other locality data); CMNFI 1987-0220, 2, 96.9-105.7 mm standard length, Rumania, Lake Călăraşi (44º20'N, 27º20'E).

Genus Petroleuciscus
Bogutskaya, 2002

This genus currently comprises 7 species of which 3 are found in Iran. Some species in this genus were formerly placed in the subgenus Squalius of the genus Leuciscus; Squalius is now recognised as a distinct genus too. The genus is distinguished by the small size of adults, the reduced number of vertebrae (modally 34-38 in total, rarely 39 or 40 ?check esfahani), few sensory cephalic pores (7-10 in the supraorbital canal, 12-19 in the infraorbital canal and 12-17 in the preoperculo-mandibular canal), a relatively small supraethmoid-mesethmoid block, narrow infraorbitals, and a deep neurocranium with a normally developed interorbital septum (Bogutskaya, 2002). Perea et al. (2010) using mitochondrial and nuclear DNA concluded that Petroleuciscus is not monophyletic. P. persidis was included with Abramis, Acanthobrama, Acanthalburnus, Ballerus, Blicca, Mirogrex and Vimba. The genus is found in the basins of the Aegean, Black, Caspian and Aral seas, the upper Tigris River , Lake Orumiyeh, and central and south-central Iran.

Petroleuciscus esfahani
Coad and Bogutskaya, 2010

Holotype

Common names

None.

Systematics

The type locality is Esfahan, stream at Dizaj in the southern Zayandeh River basin, 31°55’N, 51°30’E (holotype, CMNFI 1979-0249, female, 106.7 mm SL). Paratypes are from Esfahan, stream 1 km east of Daran, Pelasegan River tributary in the northern Zayandeh River basin, 32°59’N, 50°26’E (CMNFI 1979-0251, 134, 22.1–83.7 mm SL). The species is named after the central Iranian drainage basin in which it was found, itself named for the principal city of Esfahan, the third largest city in Iran.

Key characters

This species is characterised by having a pharyngeal tooth count of 2,5-4,2, modally 8 branched dorsal fin rays, modally 10-11 anal fin branched rays, small lateral line scales numbering 44-54, and in particular a high total vertebrae count of 40-42 (the abdominal + caudal vertebral formulae being usually 22+19, 22+20 and 21+20).

Morphology

The snout is short, its length equals to or slightly smaller than the orbit width. The tip of the mouth cleft is on a level with the upper margin of the pupil in large-sized individuals such that the mouth is slightly superior or somewhat below, at about the level of the middle of the eye, in small-sized ones. The lower jaw projects slightly compared to the upper jaw, often forming a distinct chin especially visible in larger specimens.

Dorsal-fin with 3 unbranched rays and 7(22), 8(108) or 9(5) branched rays; anal fin with 3 unbranched rays and 9(14), 10(91), 11(25) or 12(5) branched rays; pectoral fin with 14(5), 15(17) or 16(9) branched rays; pelvic fin with 7(1), 8(28) or 9(2) branched rays; lateral-line scales to hypural fold 45(3), 46(1), 47(3), 48(8), 49(3), 50(1), 51(4), 52(4), 53(2), 54(1), or 56(1); total lateral line scales 45(1), 46(2), 47(4), 48(2), 49(6), 50(14), 51(16), 52(7), 53(5), 54(2), 55(1) or 56(1); among 134 paratypes there are also 8 specimens with the lateral line widely interrupted (28–40 pored lateral-line scales from 50–53 scales in lateral series); scales around caudal peduncle 14(3), 15(12), 16(13), 17(2) or 18(1); predorsal scales 23(12), 24(3), 25(5), 26(10) or 27(1); scales between lateral line and dorsal-fin origin 9(9), 10(17) or 11(5); scales between lateral line and anal-fin origin 4(6), 5(22) or 6(3); scales between lateral line and pelvic-fin origin 3(2), 4(17) or 5(12); total gill rakers 12(2), 13(4), 14(8), 15(10), 16(6) or 17(1); total number of vertebrae 39(2), 40(22), 41(71), 42(35) or 43(5); abdominal (precaudal) vertebrae including the Weberian and intermediate vertebrae (those vertebrae that lost articulation with ribs and differ in the degree of transformation of the parapophyses into the haemal arch with the haemal spine) 21(39), 22(87) or 23(9); caudal vertebrae 18(12), 19(66), 20(53) or 21(4); and the vertebral formula is 22+19(47), 22+20(28), 21+20(23), 21+19(13), 22+18(9), 23+19(6), 22+21(3), 23+20(2), 21+18(2), 23+18(1) or 21+21(1). The number of predorsal vertebrae (in front of the first dorsal pterygiophore) is 14(52), 15(76) or 16(7) and the number of intermediate vertebrae is 3(17), 4(69), 5(46) or 6(2) Pharyngeal teeth 2.5–4.2 (8) or 2.5–5.2(2), hooked at the tip and strongly serrated below it. The gut is an elongate s-shape.

The sensory canal system was examined in the holotype and 19 paratypes, so the numbers of examined paired canals is 40. The supraorbital canal is not lengthened in its posterior section and has 8–12, commonly 9 or 10 pores (mean 9.6, standard deviation 1.33), with 3 or 4 (3 in 66%) and 5–8 (counts 6 and 7 found each in 30%) canal openings on the nasal and frontal bones, respectively. The infraorbital canal has (14, 15)16–18 pores (a mode of 16 found in 38% of canals; mean 16.6, standard deviation 1.01) with 4 or 5 (in 75%) canal openings on the first infraorbital. The preoperculo-mandibular canal is complete, with 15–17 pores (15 in 58%; 15.5, 0.74) with 4 or 5 (in 83%) and (8)9–10 (9 in 50%) canal openings on the dentary and preoperculum, respectively. It always communicates with the infraorbital canal in the pterotic, passing through the antero-dorsal process of the operculum. The supratemporal canal is complete (60% of specimens) or incomplete, with 4–6 pores. Other osteological characters are given in Coad and Bogutskaya (2010).

Sexual dimorphism

Males have a longer dorsal fin base and females a longer anal fin base, possibly associated with pair spawning. Tubercles on two male fish, 82.5–83.7 mm SL, are fine and scattered on the upper head extending down onto the upper half of the operculum but weak to absent on the snout (Fig. 2). Fine tubercles lining scales are prominent behind the head dorsally but become less evident posteriorly and ventrally. The pectoral fin bears fine tubercles on the anterior rays but numbers decrease to none on more posterior rays.

Colour

There is no distinctive colour pattern in preserved fish. The flank above the lateral line bears a fine dark speckling of melanophores and is mostly unpigmented below the lateral line. Melanophores are present on the upper part of the operculum and extend down along the posterior edge. Melanophores on the cheek ring the lower orbit. Melanophores on the lateral sides of the head are larger and more evident particularly in smaller fish. The back bears a predorsal and postdorsal stripe. The dorsal and caudal fins have very fine melanophores on the rays only. The pectoral fin has similar fine melanophores on its anterior rays only. The anal and pelvic fins are almost unpigmented. The peritoneum is silvery to cream with very few, widely scattered, melanophores.

Size

Reaches 106.7 mm standard length.

Distribution

This species is found in two streams in the Esfahan basin, one a right bank tributary of the Zayandeh River and the other a tributary to the Pelasagan Riveritself a left bank tributary of the Zayandeh River.

Solid square = holotype of Petroleuciscus esfahani, open square = paratypes;
solid circles = P. persidis holotype and paratypes (southern circle) and paratypes (northern circle), 
open circles = other material of
P. persidis;
solid triangles = P. ulanus syntypes (northern triangle);
solid triangles = P. gaderanus syntypes (other triangles),
open triangles = other material of
P. gaderanus;
star = P. kurui holotype and paratypes).

Zoogeography

This species is morphologically closest to P. gaderanus of the Lake Orumiyeh basin.

Habitat

Habitat data is confined to notes made when the type specimens were collected. The type locality was a fresh, clear stream at an altitude of 2300 m. Water temperature at 1705 hours on 9 June was 22°C, pH was 6.2, conductivity was 0.455 mS, stream width was 2–8 m, maximum depth was 1 m, current was slow to moderate, aquatic plant material was of the submergent type, the shore was grassy, and the stream bottom was a mix of pebbles and mud. A Capoeta species was also caught. The second locality was a fresh stream with clear water at an altitude of 2410 m. Water temperature at 1225 hours on 10 June was 17°C, pH was 6.2, conductivity was 0.4 mS, stream width was 4 m, maximum depth was 70 cm, current was slow to moderate, aquatic plant material was of the encrusting type, the shore was grassy, and the stream bottom was a mix of pebbles, sand and mud.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

None.

Conservation

This species is known only from two localities from collections made in 1977. Dam construction on the Zayandeh River may have altered the ecology of the river system, exacerbated by population growth and its demands on limited water resources in a desert environment. However, the species may be relatively secure in higher tributary streams.

Further work

The ecology of this species should be studied, detailed distributional information compiled, and its conservation status investigated.

Sources

Based on Coad and Bogutskaya (2010).

Type material: See above.

Petroleuciscus persidis
(Coad, 1981)

Common names

'rus mahi persidis.

[Persian chub].

Systematics

Trewavas (1972) has discussed the generic nomenclature of Pseudophoxinus Bleeker, 1859 in which this species was originally described. It is more closely related to the genus Leuciscus according to N. Bogutskaya (pers.comm., 1994; 1996). Bogutskaya (1996) placed this species in her Leuciscus borysthenicus (Kessler, 1859) group which also includes L. ulanus and deserves subgeneric rank, but later erected a new genus Petroleuciscus. Perea et al. (2010) using mitochondrial and nuclear DNA propose placing this species in Acanthobrama.

Saadati (1977) described, but did not name, a new species in the related genus Phoxinellus Heckel, 1843 from the Bid Sorkh River in the Simareh River basin between Sahneh and Kangavar in Kermanshahan (the Bid Sorkh Pass is at 34°26'N, 47°49'E). I was unable to find this species in his collections and no further material has come to light. It had 8-9 dorsal fin branched rays, 8-9 branched anal fin rays, 8 branched pelvic fin rays, 47-56 scales in a complete lateral line, and 20 long and crowded gill rakers.

The type locality of Pseudophoxinus persidis is the "upper Shur River drainage at "Koorsiah" village, near Darab on Darab-Fasa road, 28°45.5'N, 54°24'E, Fars". The holotype is a 54.7 mm standard length male held at the Canadian Museum of Nature, Ottawa under CMNFI 1979-0154A (see figure above). Paratypes comprise 95 fish, 34.7-58.8 mm standard length, from the same locality as the holotype under CMNFI 1979-0154B, and 5 fish, 74.7-92.4 mm standard length, under CMNFI 1979-0499 from an "irrigation ditch at village 32 km west of Kor River bridge on road to Dariush Dam, 30°04.5'N, 52°36'E, Fars". Paratypes were distributed to the following institutions from CMNFI 1979-0154B: BM(NH) (2), ROM (2), UAIC (1), UBC (2) and UMMZ (2).

Key characters

This species is characterised by having a pharyngeal tooth count of 1,5-4,1, modally 7 branched dorsal fin rays, anal fin branched rays 7-9, pelvic fin rays 7-8, pored lateral line scales 35-43 in a complete lateral line, short gill rakers numbering 10-14 on the whole first arch, total vertebrae 34-37, a light coloured peritoneum but with numerous melanophores, and flanks with a lateral stripe evident posteriorly but fading anteriorly and not reaching the head.

Morphology

The mouth is terminal and oblique and the rictus reaches back to a level just anterior to the anterior eye margin. A pelvic axillary scale is present. The subcircular to oval scales bear numerous fine circuli and radii on the anterior and posterior fields (total radii number 23-44). Pharyngeal teeth are strongly hooked, concave below the hook and serrated along the margins of the concave surface, particularly on the anterior edge. In the largest fish, the hook may be much reduced to absent, serrations may be absent, and anterior teeth in the main row are rounded. Gill rakers reach the adjacent raker when appressed. The gut is a short and simple s-shape.

Meristic values are as follows: dorsal fin branched rays 6(2) or 7(189) after 3 unbranched rays, anal fin branched rays 7(3), 8(37) or 9(10) after 3 unbranched rays, pectoral fin branched rays 13(10), 14(20) or 15(20), pelvic fin branched rays 7(31) or 8(19), lateral line scales 35(2), 36(19), 37(4), 38(10), 39(3), 40(6), 41(5) or 43(1), total gill rakers 10(1), 11(9), 12(32), 13(6) or 14(2), pharyngeal teeth 1,5-4,2(1), 1,5-4,1(13) or 1,5-4,0(1), and total vertebrae 34(1), 35(20), 36(27) or 37(2). Karyotype 2n=50 (Esmaeili and Piravar, 2006a).

Sexual dimorphism

Number of anal fin branched rays is significantly higher in females (mean 8.3 versus 8.0 in males). Pectoral and pelvic fin lengths, longest dorsal and anal fin rays and caudal peduncle length are shorter in females than in males while head length, head width and predorsal length are longer in females than in males.

Colour

The back and upper flanks are dark and the belly cream. The straight lateral stripe extends from a diffuse area on the tail base to a level at or in front of the dorsal fin origin but does not reach the head. The stripe overlaps the lateral line on the caudal peduncle but lies above it on the flank, paralleling the back. Dorsal fin membranes are lightly speckled with melanophores which tend to be concentrated along the fin ray margins. The caudal and anal fin membranes are mostly clear with pigment restricted to fin ray margins. The pectoral fin may be pigmented on the membranes and there is often strong pigment along the posterior edge of the first unbranched ray. The pelvic fin has little or no pigmentation. Large fish are much darker overall than small fish, obscuring the stripe and with more pigment on fin rays and membranes.

Size

Reaches 9.2 cm standard length.

Distribution

This species is found in the Kor River and Hormuz basins (Coad, 1981e; M. Hafezieh, pers. comm.). Abdoli (2000) reports this species from the Kor and Pulvar rivers, the Hilleh, middle Mand, Shur (tributary of the Dasht-e Palang) and the Kul rivers.

Zoogeography

Durand et al. (2000) place this species in a Leuciscus (= Squalius) cephalus "complex", i.e. descendents of peripheral isolates of a widespread ancestral species, later re-invaded by Danubian S. cephalus.

Habitat

Habitat knowledge is restricted to field data of collections. This species was collected in large rivers, streams, springs, irrigation ditches and qanats, but all these have in common a stream-like environment for much of the year. They are relatively shallow (20 cm to 2 m), variable width (0.5-75 m), medium to slow current, some submergent and emergent aquatic vegetation and a bottom varying from pebbles and gravel to mud. Water temperatures varied from 15 to 23°C from October to January and presumably would be over 30°C in the summer. Conductivity ranged from 0.3 to 1.0 mS. Altitude ranged from 980 to 1940 m. The lower reaches of some of the capture rivers are salty and may not support this species.

Age and growth

Unknown. Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 10 fish measuring 4.21-8.33 cm standard length. The a-value was 0.0178 and the b-value 3.229 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases).

Food

Gut contents include insect remains and fragments of large plants. Diet presumably consists mostly of aquatic invertebrates.

Reproduction

Reproduction in this species is unknown but egg development in adult fish collected in winter and young of the year collected in October suggest spring and early summer as the spawning season.

Parasites and predators

Jalalai et al. (2000) record Dactylogyrus sphyrna, a monogenean, from the gills of this species in the Kor River basin. Barzegar and Jalali (2006) report parasites in this species from Kaftar Lake as Lernaea cyprinacea, Trichodina sp., Ichthyophthirius multifilis and Dactylogyrus sphyrna. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Lernaea foliaceus on this species.

Economic importance

None.

Conservation

The distribution of this species in various small habitats probably means that it is not in immediate danger. However water is in short supply in this part of Iran and abstraction may threaten its survival.

Further work

The ecology of this species should be studied and detailed distributional information compiled.

Sources

Based on Coad (1981e).

Type material: See above, Pseudophoxinus persidis (CMNFI 1979-0154A, 1979-0154B, and 1979-0499).

Iranian material: CMNFI 1979-0025, 11, 19.4-79.6 mm standard length, Fars, Kor River at Marv Dasht (29º51'N, 52º46'30"E); CMNFI 1979-0114, 1, 47.1 mm standard length, Fars, Mand River (29º41'N, 52º06'E); CMNFI 1979-0156, 6, 43.8-64.3 mm standard length, Fars, qanat in Rashidabad (28º47'N, 54º18'E); CMNFI 1979-0163, 8, 31.1-48.3 mm standard length, Fars (no other locality data); CMNFI 1979-0164, 15, 28.8-52.6 mm standard length, Fars (no other locality data); CMNFI 1979-0194, 3, 48.9-58.2 mm standard length, Fars, upper Shur river drainage (28º45'30"N, 54º24'E); CMNFI 1979-0292, 7, 37.0-54.0 mm standard length, Fars, Lapu'i spring near Zarqan (29º48'N, 52º39'E); CMNFI 1979-0305, 2, 23.8-25.5 mm standard length, Fars, Pulvar River at Pasargad (30º12'N, 53º12'E); CMNFI 1979-0342, 33, 23.6-43.2 mm standard length, Fars, Kor River at Band-e Amir (29º46'N, 52º51'E): CMNFI 1979-0503, 4, 32.2-113.5 mm standard length, Fars (no other locality data); USNM 258445, 18, 15.5-44.5 mm standard length, Fars, Chasht Khvar (ca. 29º44'N, ca. 53º12'E).

Petroleuciscus ulanus
(
Günther, 1899)

Common names

None.

Systematics

This species was originally described in the genus Leuciscus. Saadati (1977) considers this species to be in the genus Alburnus but did not examine the types. Leuciscus gaderanus Günther, 1899 is a synonym, even Günther (1899) the describer of the two species indicating that this may be the case.

The 2 syntypes of Leuciscus ulanus, 68.0-84.5 mm standard length, are in the Natural History Museum, London and are "from Ula on the Zola Chai", Günther (BM(NH) 1984.10.10:1-2). The 3 syntypes of Leuciscus gaderanus, 44.6-73.0 mm standard length, are "from the Gader Chai" which is near Ocksa (BM(NH) 1899.9.30:113-115). Additionally there are 5 specimens of Leuciscus gaderanus, 11.0-54.5 mm standard length "from near the mouth of the Nazlu Chai at Superghan" (BM(NH) 1899.9.30:108-112). Günther (1899) refers to 3 young specimens from the latter locality as syntypes but of these 5 fish under this catalogue number and locality given as "Superghan" on the label only 2 fish are small (11.0-12.5 mm SL) and the other 3 fish are larger (45.8-54.5 mm SL). Another collection comprising 1 specimen, 27.0 mm standard length, is from the Urmi River according to the label (BM(NH) 1899.9.30:107). Its type status is unclear since the locality is wrong but the size is small and it could be the third of Günther's "three young specimens" since its catalogue number is in sequence.

Key characters

Distribution and meristic characters identify this species.

Morphology

Dorsal fin unbranched rays 3, branched rays 7-9, anal fin unbranched rays 3, branched rays 7-11, pectoral fin branched rays 12-14, and pelvic fin branched rays 7-8. Lateral line scales 36-45. There is a pelvic axillary scale. Scales with a slightly anterior focus and numerous radii on the anterior and posterior fields. Total gill rakers 12-16, reaching the first or second raker below when appressed. Pharyngeal teeth 2,5-4,2, more rarely 2,4-5,2, 2,5-5,2 or 2,4-4,2, hooked at the tip and strongly serrated below on the larger teeth. Total vertebrae 37-38. The mouth is oblique and extends back to just behind the front margin of the eye and the lower jaw protrudes slightly or hardly at all (Günther (1899) has the upper jaw slightly overlapping the lower). The gut is an elongate s-shape and may have an anterior loop to the left.

Meristic values for Iranian specimens are:- dorsal fin branched rays 7(5) or 8(15); anal fin branched rays 7(1), 8(6), 9(11) or 10(2); pectoral fin branched rays 12(5), 13(10), or 14(5); pelvic fin branched rays 7(19) or 8(1); lateral line scales 38(3), 39(6), 40(6), 41(1), 42(1) or 45(2); total gill rakers 12(2), 13(7), 14(6), 15(3) or 16(2); pharyngeal teeth 2,5-4,2(9), 2,4-5,2(2), 2,5-5,2(1) or 2,4-4,2(1); and total vertebrae 37(4) or 38(1).

Sexual dimorphism

Males bear tubercles on the pectoral fins mostly in a single file, occasionally two together, branching with the rays. Small tubercles are also found thickly on the top and sides of the head but no pattern was discernible in the specimens examined (types of L. ulanus).

Colour

There is a narrow, straight black stripe running from the upper half of the eye to the end of the lateral line separating the bluish back from the silvery flanks (Günther, 1899), best developed in preserved specimens posteriorly. Live fish have a grey-silver flank with some spots, a brown-green back and a light silver abdomen. Lower fins are pale to light yellow and the dorsal and caudal fins are light grey (Abbasi and Sabkara, 2004a; 2004b). The lower half of the operculum below the mid-eye level has few or no spots while the upper half is heavily pigmented. There is also no pigment below the eye or it is restricted to a thin line around this lower margin. The flanks are dotted with minute pigment spots. The back mid-line has a black stripe, most obvious predorsally. The rays and membranes of the dorsal fin, caudal fin and anterior pectoral fin bear melanophores; melanophores weak to absent on the anal and pelvic fins. The peritoneum is silvery brown with scattered melanophores.

Size

Reaches 14.2 cm for length (Abbasi and Sabkara, 2004b).

Distribution

This species is endemic to the Lake Orumiyeh basin (Günther, 1899). This species was found only in the Gadarchai and Mahabad River and was not caught in the Zolachai, Nazluchai, Baranduzchai and sampled areas of the Siminerud and Zarrinerud in recent collections by Abbasi and Sabkara (2004a; 2004b).

Zoogeography

Petroleuciscus kurui (Bogutskaya, 1995) from the upper Tigris River system of Turkey is the closest relative of this species (Bogutskaya, 1995) although other members of the Orumiyeh ichthyofauna are related to fishes from the Caspian Sea basin.

Habitat

Essentially unknown, found in stream or river habitats.

Age and growth

Fish caught by Abbasi and Sabkara (2004b) in the Gadarchai and Mahabad River were 34-142 cm in fork length and 1.4 years old. Males comprised 28.8% and females 71.2% of the total population. Maturity was attained at 2 years.

Food

Generally unknown but traces of insects and large quantities of filamentous algae were found in the gut contents of a few specimens. The algae may be an accidental inclusion as the gut is short and probably cannot digest plant material. Fish caught by Abbasi and Sabkara (2004a; 2004b) however contained a wide range of phytoplankton (19 genera) and zooplankton (7 groups), as well as benthic organisms (4 groups). Daphnia, Chydrus and chironomids were dominant gut contents. It was considered to be an omnivorous fish preferring to feed on zooplankton and other mid-water animals.

Reproduction

Spawning of fish in caught by Abbasi and Sabkara (2004b) took place from 1 April until 1 July with absolute fecundity estimated at 1810-16,115, average 6437 eggs.

Parasites and predators

Jalali et al. (2005) summarise the occurrence of Gyrodactylus species in Iran and record G. sp. in fish from the Baranduz and Halaj rivers.

Economic importance

None.

Conservation

This species is known only from the type series and a few other collections. This suggests that it is quite rare or has a restricted habitat preference.

Further work

The biology, distribution and abundance of this poorly known species should be investigated to see if it requires protection.

Sources

Type material: See above for L. ulanus (BM(NH) 1984.10.10:1-2) and L. gaderanus (BM(NH) 1899.9.30:113-115 and BM(NH) 1899.9.30:108-112).

Iranian material: CMNFI 1970-0560, 35,11.4-48.8 mm standard length, Azarbayjan-e Bakhtari, Mamiyand Chay (ca. 36º59'N, ca. 45º39'E); CMNFI 2007-0096, 6, 38.8-68.5 mm standard length, Azarbayjan-e Bakhtari, Baranduz Chay basin (ca. 37º25'N, ca. 45º10'E); BM(NH) 1899.9.30:107, 1, 27.0 mm standard length, Azarbayjan-e Bakhtari, Urmi River (no other locality data).

Genus Pimephales
Rafinesque, 1820

This genus is endemic to North America where there are 4 species in Arctic, Atlantic and Gulf of Mexico drainages. It occurs in Iran as an exotic.

These fishes are small, less than 12 cm standard length, with a stout to slender body. They are uniquely characterised by the last, unbranched dorsal fin ray in males being short, blunt and separated from the succeeding rays by a membrane. There are usually 7 branched dorsal and 6 branched anal rays. The lateral line is complete to incomplete. The head and back before the dorsal fin is flattened and the predorsal scales are small, irregularly arranged and crowded. The intestine varies from long to short, the peritoneum from black to silvery. Pharyngeal teeth are in one row. The flank has a dark band often terminating in a spot at the caudal fin base.

The species of this genus are of some commercial importance as bait or food for sport fishes.

Pimephales promelas
Rafinesque, 1820

Common names

None.

[fathead minnow].

Systematics

This exotic is unrelated to native Iranian fishes and was described from Lexington, Kentucky, U.S.A..

Key characters

This species is unique in Iran in that the last, unbranched dorsal fin ray is short and separated by a membrane from the next, longer ray in males. Other characters include the absence of a barbel, scales in front of the dorsal fin are much smaller and more crowded than flank scales, the back is flattened, the lateral line is incomplete to nearly complete, the mouth is small and terminal, and by the colour and tubercle patterns described below.

Morphology

Dorsal fin branched rays 7, rarely 8, anal branched rays 6, sometimes 7, pectoral rays 14-18 and pelvic rays 8-9. Scales in lateral series 39-56, lateral line pores usually ending before the dorsal fin origin but rarely may be almost complete. Scales are oval with a markedly anterior focus, few circuli and moderate numbers of radii restricted to the posterior field. Vertebrae number 35-38.

Pharyngeal teeth 4-4, slightly hooked at the tip, compressed and with elongate and concave cutting surfaces. The gut is elongate with several loops. Gill rakers 12-16, short and touching the adjacent raker when appressed.

Sexual dimorphism

Large tubercles in males are found on the snout in 3 (rarely 4) rows with 4-15 in the lower row, and with up to 11 tubercles on the lower jaws. There are also smaller tubercles on top of the head and pectoral rays. Tubercles first develop in early April in Canada and males begin to lose them in late July. By early September only scars remain. Males also develop a blue-black to grey, spongy, wrinkled pad on the back between the head and dorsal fin. Fin membranes swell. Breeding males darken, particularly the head and dorsal fin, and the stripe is not apparent. The body can be completely black with a white band at the head-body region and under the dorsal fin. This occurs only during aggression or sexual activity. The lateral banding enhances the robust appearance of fish which must maintain a territory over several weeks without much opportunity to feed. Weight loss is replaced by water to help maintain the image of a fat and vigorous male. Very frightened fishes blanch.

Females have a protruding ovipositor.

Colour

The back and upper flank are dark olive-green to brown, silvery to golden flanks and a whitish belly. There is a mid-flank stripe and a predorsal stripe. Scales on the upper flank are outlined by pigment. A spot at the tail base is faint to absent. Peritoneum black.

Size

Reaches 10.2 cm total length but usually much less.

Distribution

This species was reported from a reservoir, probably the Yengi Kand where bass (Micropterus salmoides) and bluegills (Lepomis macrochirus) were introduced, south of Tehran at 75 km from Asia-beg (Andersskog, 1970) (presumably Asia Bak at 35°19'N, 50°30'E on the Tehran-Esfahan highway). The MMTT catalogue also has a record for the "Dusadj Reservoir", 90 km west of Saveh, presumably in Markazi Province and the Namak Lake basin too. These reservoirs may be the same locality as "Dusadj" or Duzaj, Yang-e Kand and Yanguikand are all villages in the same general area west of Saveh. The reservoir is deduced from maps to be at about 35°19'N, 49°55'E.

A report for the Golpayegan Reservoir (Golpayegan is at 33°27'N, 50°18'E) in the Namak basin by Saadati (1977) was an error (R. J. Behnke, in litt., 1979).

? Map

Zoogeography

The fathead minnow is native to North America and was introduced to Iran accidentally in shipments of largemouth bass, Micropterus salmoides, and bluegills, Lepomis macrochirus. Fathead minnows are pond-raised in North America as food for bass.

Habitat

Fathead minnows are found in ponds, small lakes and slow-flowing brooks, often associated with vegetation. They tolerate high turbidity, high temperatures, low oxygen and high alkalinity and so are able to survive in desiccating or other conditions unfavourable to most fishes.

Age and growth

Males grow faster and larger than females, typical of nest defending species. Life span is about 3 years with maturity attained as early as 1 year, rarely in the year of birth.

Food

Food is bottom sediment for its organic content including plant material, aquatic insects and zooplankton.

Reproduction

Spawning runs from April to August in North America, once water temperatures reach 14°C and light-dark hours are 16-8. Males choose a spawning site under a log, rock, plant stems or even a lily pad or any solid artificial structure in shallow water. Cavity spawning in mud-bottomed habitats prevents the eggs from being smothered as well as offering protection from predators for the relatively few eggs spawned from a small fish. The male will clean out the cavity, spending up to 10 hours on the task, using his tubercles to scrape, pulling debris with his mouth and sweeping with his tail fin. The spongy pad on the back may serve to test spawning sites and eggs chemically. The pad secretes a mucus which is smeared on the spawning site perhaps to improve it for egg survival since mucus protects against disease and parasites. Diseased eggs are eaten by the male. The mucus may also serve to indicate ownership of a nest site. Spawning male fatheads lose their ability to produce alarm or fright chemicals on skin injury, otherwise the continual pad rubbing would disturb spawning activities by releasing alarm substance and scaring away females.

Females may enter the spawning site casually, be chased there by a male or enticed by a face to face encounter and leading to the nest. The male lifts and presses the female on her side between himself and the roof of the spawning site and the female rapidly undulates through an s-shape. Egg deposition is thought to use the same unusual mechanism as in the bluntnose minnow (Pimephales notatus (Rafinesque, 1820)). As each egg is extruded, it is transferred to the upper side of the female and the undulation rolls it along between her side and the nest cavity roof. When the adhesive egg reaches the tail she presses it against the roof where it sticks. How the female transfers the egg from her papilla to her side is unknown and the question remains why this sideways process is used when other roof spawners simply turn upside down and deposit eggs directly. The process may serve to roll eggs into a vacant roof space, occupying the roof most efficiently. Eggs laid on top of others may not attach well or prevent proper development of the underlying eggs.

Males court females by swimming rapidly up to them and then freezing at 3-5 cm away, and by leading females with a zig-zag or straight-line motion from the female to the nest site. Males will also display to females by erecting their fins for 2-3 seconds and by jump-swims in which a male swims upwards to a female then rolls on his side and swims back down. Butting and lateral quivering also occur, perhaps attempts to assess spawning condition of the female. Males defend the eggs against other fishes, including female fatheads, by using the snout tubercles to butt and tail swipes to intimidate by sending a pressure wave sensed by the lateral line system. Chasing and biting are common and 2 males may carousel (or circle head-to-tail) trying to contact each other. Leeches and turtles are also driven away. Some eggs are lost while the male is distracted chasing away predators. Repeat spawning may be necessary to replace lost eggs. Males also aerate and clean the eggs with fin movements. A nest will contain eggs in various stages of development as the male will spawn with several females. Females will deposit eggs in several nests. Orange, mature eggs are up to 1.6 mm in diameter with 12,000 or more per nest. A female will release up to 10,164 eggs in a season but from 9 to 1136 at a time. Spawning intervals are 2 to 16 days.

Parasites and predators

It is an important food for many other fishes and aquatic birds.

Economic importance

In the U.S.A. fatheads are raised as bait fish and as forage fish for introduction into bass fishing lakes. They are also used extensively as a laboratory animal for tests of toxic compounds. They have even been used to evaluate the biological effects of materials from the moon.

Conservation

This exotic species may have deleterious effects on the native fishes and its conservation is of no utility. It is extremely fecund and could compete with native species for food and habitat. A variety of parasites have been reported from this species including some which can have devastating effects on fish populations such as Ligula intestinalis. The introduction of this fish into Europe led to an outbreak of enteric redmouth disease (Yersinia ruckeri) which spread to native and farmed stocks of commercially important fishes such as trout (Michel et al., 1986; Welcomme, 1988). Native fishes may be less well adapted to withstand the depredations of exotic diseases and parasites. Ideally it should be extirpated but, if this is not possible, its interactions with native fishes should be studied and attempts made to prevent its spread to other water bodies.

Further work

The status and numbers of this exotic should be checked by field work. Any data on this exotic species held in files of the Department of the Environment should be published so that this "grey" literature is not lost to future students of Iranian fishes.

Sources

Based on North American literature summaries such as Scott and Crossman (1973) and Becker (1983) as well as personal observations. The biology of Iranian populations is unknown.

Genus Pseudogobio
Bleeker, 1859

Pseudogobio rivularis
(Basilewsky, 1855)

Reported from the Tedzhen River in Turkmenistan on the northeastern border of Iran by Aliev et al. (1988) in their text but not in their distributional table. This exotic species is from China. Also recorded from the Karakum Canal and Kopetdag Reservoir (Shakirova and Sukhanova, 1994; Sal'nikov, 1995) and may eventually reach the Caspian Sea basin. No Iranian record.

Genus Pseudorasbora
Bleeker, 1859

This genus contains 3 species with a native distribution in eastern Asia including the Amur River basin shared between Russia and China, in Japan, other parts of China, and in Korea. One species is an exotic now found in Europe and accidentally introduced in Iran.

The genus is characterised by a small and transverse mouth positioned at the top of the snout rather than the anterior tip, the lower jaw has a trenchant edge and projects slightly beyond the upper jaw, no barbels, pharyngeal teeth are in a single row, the gut is short, scales are large, gill rakers are rudimentary, dorsal and anal fins are short and spineless, and there is no keel on the abdomen.

Pseudorasbora parva
(Temminck and Schlegel in Siebold, 1842)

Pseudorasbora parva from Wikimedia Commons.

Common names

آمورچه (amurcheh or amorcheh), amoornama, parva.

[stone moroko, topmouth gudgeon, topmouth minnow, false rasbora; chebachek or Amur chebachok in Russian].

Systematics

Leuciscus parvus was originally described from Nagasaki, Japan.

Subspecies have been described in China but exotic introductions are usually referred to the type subspecies. Reshetnikov et al. (1997) and Bănărescu in Bănărescu (1999) give the date for this species as 1846. The phenotype of this species is highly influenced by environmental conditions and this is suggested to be one of the attributes that make this fish a successful invasive species (Záhorská et al., 2009).

Key characters

The mouth structure is unique, being very small and lying entirely before the nostril level, almost vertical, opening antero-dorsally with the gape entirely visible in dorsal view. The lower jaw protrudes to form the most anterior part of the head.

Morphology

Dorsal fin unbranched rays 3 followed by 7-8, usually 7, branched rays, anal fin with 2-3, usually 3, unbranched rays and 5-7 branched rays, usually 6, pectoral fin branched rays 11-14, and pelvic fin branched rays 6-8, usually 7. Lateral line scales 30-40, with the lateral line rarely incomplete. A pelvic axillary scale is present. The scale radii are restricted to the posterior field. Gill rakers are rudimentary and are only well-developed at the junction of the upper and lower arms of the gill arch. These rakers are stubby and rounded, reaching the adjacent raker when appressed, and bearing fine, fleshy fimbriae which extend onto the adjacent parts of the gill arch. Anterior rakers are absent and patches of fimbriae are found. Rakers number 6-16, usually 9-13. Total vertebrae 31-38 presumably the result of different counting methods; Naseka (1996) gives 36-38 and fish from Turkey have 34-37, cf. below. Pharyngeal teeth usually 5-5, rarely 6-5, with the tips strongly hooked and the area below the hook flattened and without ridges or only very weakly ridged on some teeth. The gut is an elongate s-shape. The chromosome number is 2n=50 (Klinkhardt et al., 1995).

Meristic values for Iranian specimens are:- dorsal fin branched rays 7(11) or 8(1), anal fin branched rays 6(12), pectoral fin branched rays 13(7) or 14(5), pelvic fin branched rays 7(4) or 8(8), lateral line scales 34(4), 35(5) or 36(3), pharyngeal teeth 5-5(9), 6-5(2) or 4-4(1), and total vertebrae 34(2) or 35(10).

Sexual dimorphism

A horny pad develops on the jaws in males and females during spawning and strong, sharp tubercles in males. One tubercle is found between the eye and the nostril, one below the nostril (this may be absent), one next to the upper lip on a line across from the one below the nostril, 5-8 in a row from the extreme corner of the mouth along the side of the head over the flesh of the cheek, and 2-3 below the lower lip from the tip of the lower jaw to the end of the jaw on the lower head surface. Lower head surface tubercles may coalesce at the base but each tubercle bears a single rounded cusp. Rarely a tubercle may have a single base but two cusps.

Males are larger than females and have larger fins. Spawning males are darker than females and the flank has a metallic violet sheen.

Colour

The head and body has a mid-lateral stripe but this is obscured in adults by crescentic speckles situated posteriorly on each scale. The back is light grey, the flanks silvery and the belly whitish. Dorsal and anal fins are speckled and turn almost black in spawning fish.

Preserved fish have a cream coloured belly with the back much darker. The head is black dorsally and fades to cream ventrally. The scales on the back and flanks, but not the belly, carry a broad band of pigment which follows the scale margin distally. The extreme edge of the scale is hyaline but the arc of pigment effectively defines the posterior scale margin and outlines the scale pattern of the back and flanks. Pigmentation on fins is mostly restricted to the rays and their margins but is found also on fin membranes to varying extents. Pigmentation is strongest distally on all fins. The dorsal fin, particularly in smaller fish, bears patches on the membranes posterior to branched rays 1 or 2 through 5 or 6, starting on ray 1 or 2 below the mid-point of the ray length and descending gradually behind successive rays to lie near the base behind the last ray. These patches are vertically short and do not touch the succeeding ray. The leading edge of the dorsal, anal, pectoral and pelvic fins and the upper and lower edges of the caudal fin are black in large, and some small, fish. Pigment may be concentrated along the mid-line forming a thin stripe, only apparent posteriorly in some fish. There is a dark line along the mid-line of the back. The peritoneum is silvery with some scattered melanophores.

Size

Attains 12.0 cm (Movchan and Kozlov, 1978).

Distribution

The natural range of this species is in eastern Asia as given above under the genus. It has been introduced to Iran by accident (Abdoli, 1992; Coad and Abdoli, 1993b). It is now found in ab-bandans at Avaness, Hasan Tabeeb and Shaeed Ziaee (all about 40-45 km east of Gorgan), Teer Tash and Lemrask (about 20-25 km east of Behshahr), Lapoo (about 4 km east of Babol Sar) on the Caspian Sea coast, and at Gorgan-Aliabad, Mazandaran, from the Safid River estuary and neighbouring waters and the Anzali Talab (Iranian Fisheries Research and Training Organization Newsletter, 6:8, 1994; Anonymous, 1994; Abbasi et al., 1999), from the Atrak, Gorgan, Gharasu, Tajan, Babol, Haraz, Sardab, Pol-e Rud, and Safid rivers (Kiabi et al., 1999), from fish ponds at Arak probably inadvertently carried there with carp fingerlings imported from Gilan on the Caspian shore; at Mashhad in northeastern Iran, in springs near Kermanshah and in Lake Zeribar, Kordestan (Coad, 1996g); in Sistan at Hamun Kushk, and the canal flowing into the Chahnimeh (J. Holčík, in litt., 1996), and from the International Wetlands of Alma-Gol, Adji-Gol and Ala-Gol (Patimar et al., 2002a; 2002b; Patimar, 2008).

Abdoli (2000) records it generally from the Dasht-e Kavir, Dasht-e Lut, Kerman-Na'in and Sistan basins; the lower Kashaf River in the Tedzhen River basin; the middle Atrek, lower Neka, Babol, Heraz, Chalus, Tonekabon, and Safid rivers and the Anzali Talab in the Caspian Sea basin; the middle to lower Talkheh and lower Zarrineh rivers in the Orumiyeh basin; the lower Shur and middle and lower Qareh Chai in the Namak Lake basin; the middle and lower Zayandeh River in the Esfahan basin; and the Simarreh and lower Gav Masiab rivers in the Tigris River basin. Jolodar and Abdoli (2004) record it from most water bodies on the Iranian Caspian coast and Abdoli and Naderi (2009) from the Atrak, Gorgan, Gharasu, Tajan, Babol, Heraz, Sardab, Pol-e Rud and Safid rivers there. Esmaeili et al. (2010) and Esmaeili et al. (2011) add Lake Zarivar and the Gulf and Lake Maharlu basins and it is now probably distributed throughout Iran.

It is also recorded in the Karakum Canal, Kopetdag Reservoir and Tedzhen River of Turkmenistan (Shakirova and Sukhanova, 1994; Sal'nikov, 1995) so may well reach the Tedzhen (= Hari) River basin of Iran eventually. Pipoyan (1996a) reports it from the Araks River in Armenia.

Zoogeography

This species was first recorded in western Eurasia in Romania in 1960 as an accidental introduction with Chinese carps from the lower Yangtze River of China. The species is now widespread in Europe and is becoming common in western Asia including Kazakhstan, Uzbekistan, Kyrgyzstan and southern Anatolian Turkey as well as Iran (Wildekamp et al., 1997).

Habitat

This species prefers well-vegetated areas as protection from predators. It may be found in streams, rivers and ponds, and more rarely in the shallows of large lakes. It is apparently quite resistant to pollution (Bănărescu in Bănărescu, 1999) and is found in waters that freeze over and that attain 30ºC in summer (Boltachev et al., 2006).

Age and growth

Life span is about 5 years with maturity attained at 1-2 years, usually at 1 year in Europe or the second year of life in the Crimea (Boltachev et al., 2006). Most fish in a population are 2-3 years old or 1-2 years in the Crimea. Patimar et al. (2002a; 2002b) report 4 age groups from the International Wetlands of Alma-Gol, Adji-Gol and Ala-Gol, with the smallest mature specimens found at 2 years. Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 33 Iranian fish measuring 3.29-5.99 cm standard length. The a-value was 0.0286 and the b-value 2.763 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases).

Food

This species feeds on benthos but also some zooplankton. Food items include various aquatic insects such as stone flies, caddis flies, chironomids, water sawbugs and midge larvae but guts also contain sponges, bryozoans, Spirogyra, detritus, and fragments of higher plants (Movchan and Kozlov, 1978). It may also feed on the eggs and juveniles of native fishes. Young fish take zooplankton (Movchan and Kozlov, 1978). Bănărescu in Bănărescu (1999) reports also isopods and aquatic worms and, in fish ponds, artificial food.

Reproduction

Reproduction begins at 16-18°C and lasts two months in its native Amur River basin. Fecundity is about 5000 elliptical eggs with a diameter of 2.0-2.5 mm, and this species has intermittent spawning with up to 85 eggs per batch in introduced populations in Central Asia (Makeyeva and Mokhamed, 1982). Up to 60 batches may be laid in a spawning season. The spawning site is cleaned of ooze and plant material. Adhesive eggs are deposited on the lower surface of stones, and occasionally sticks or empty mollusc shells, and are protected by the male using the head tubercles to drive away other fishes. The ellipsoidal eggs are laid in strips, usually of 5 eggs but as many as 10. Males clean the eggs and remove dead ones. The spawning season in Central Asia is April to August and spawning takes place in warm, shallow and calm waters in the morning. Spawning in the Crimea is in second half of May or in June, late May to July in the Ukraine and from the end of June to the beginning of August in the Amur (Boltachev et al., 2006). Female specimens from Iran collected in March, April and May are ripe and males have well-developed breeding tubercles and Patimar et al. (2002a; 2002b) report a spawning peak in April in the International Wetlands of Alma-Gol, Adji-Gol and Ala-Gol of Iran.

Parasites and predators

Malek and Mobedi (2001) report Clinostomum complanatum from this species in Mazandaran, in the Shiroud. Sander lucioperca and Silurus glanis are predators in Turkmenistan (Aliev et al. (1988). Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Lernaea foliaceus on this species.

Economic importance

Makeyeva and Mokhamed (1982) and Movchan and Kozlov (1978) report competition with commercial species and predatory behaviour on carp larvae if there is insufficient food. Male reproductive aggression may inhibit breeding of native fishes. Boltachev et al. (2006) report that it is a facultative parasite of other fishes in enclosed areas including commercial species such as the silver carp (Hypopthalmichthys molitrix). Areas of the body are attacked such as behind the dorsal fin and above the anal fin out of sight of the affected fish. Skin and muscles are eaten away. Welcomme in Courtenay and Stauffer (1984) regards this species as a pest when introduced. Bănărescu in Bănărescu (1999) reports this species as a competitor for food with native species in Europe.

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in textbooks.

Conservation

None required for an introduced species.

Further work

The distribution of his species as an exotic in Iran should be thoroughly documented and its biology and effects on native species studied.

Sources

Iranian material: CMNFI 1991-0160, 3, 47.8-55.8 mm standard length, Mazandaran, Abgeere Avanes (37º03'N, 54º47'E); CMNFI 1991-0161, 1, 52.5 mm standard length, Mazandaran, Madarso River (37º23'N, 55º47'E); CMNFI 1993-0134, 7, 45.0-59.1 mm standard length, Mazandaran, Gorgan-Aliabad (37º01'30"N, 54º47'36"E); uncatalogued material, 1, 86.9 mm standard length, Kermanshahan, sarabs near Kermanshah (no other locality data).

Comparative material: CMNFI 1983-0204, 7, 51.7-76.6 mm standard length, Turkey, Edirne, Meriç River at Ipsala (40º55'N, 26º23'E); CMNFI 1983-0343, 5, 49.9-83.4 mm standard length, same locality as preceding.

Genus Rhodeus
Agassiz, 1832

The bitterlings comprise about 18 species in Europe, Asia Minor, the Caspian Sea basin, China, Japan and Korea with 1 species in Iran.

They are small fishes with deep, compressed bodies, an incomplete lateral line (about 10 pored scales or less), large to moderate-sized scales, females with an ovipositor, males larger than females (unusual in cyprinid fishes), brightly coloured and tuberculate when spawning, pharyngeal teeth in 1 row and not or only slightly serrated, mouth small, oblique and subterminal, no barbels, dorsal fin short to moderately long and spineless, anal fin of similar length, gill rakers short, intestine long and spirally coiled, and peritoneum black.

Rhodeus amarus
(Bloch, 1782)

Common names

ماهي مخرج لوله اي (= mahi-ye makhraj lulehi or mahi-e-makhraj looleei, meaning tube-like vent fish), rodeus.

[karka in Azerbaijan; gorchak in Russian; bitterling, European bitterling].

Systematics

Holčík and Duyvené de Wit (1964) reviewed the systematic status of Rhodeus sericeus (Pallas, 1776) in Europe and western Asia (but not Iran) in comparison to the Chinese populations of this species. They refer European and western Asian populations to Rhodeus sericeus amarus (Bloch, 1782) and Iranian populations were long regarded as this subspecies. Cyprinus sericeus was described from Dauriya; Cyprinus amarus was described from the Elbe basin, Germany and no types are known (Eschmeyer et al., 1996). Later, Holčík and Jedlička (1994) consider subspecies not to be warranted as the key characters used in distinguishing them (pored lateral line scales, scales in lateral series, gill rakers) showed clinal variation with longitude and the number of segments was also related to latitude, elevation, mean annual air temperature, and fish size. Bohlen et al. (2006) consider Rhodeus from the Vistula to the Volga to belong to the R. amarus eastern clade based on cytochrome b sequences.

Kottelat (1997) considers Rhodeus amarus to be a distinct species since it is diagnosable (although differences with Rhodeus sericeus are slight and largely overlap), and it is separated by 2-4 million years in time and 4000 km in space. However he does admit that the immense distributional gap may be a source of bias. Holčík in Bănărescu (1999) considers amarus to be synonymous with sericeus. Bogutskaya and Komlev (2001) found no characters to clearly confirm the specific status of R. amarus. The name amarus is retained here as an indication that the taxa are geographically isolated. Van Damme et al. (2007) refer the Caspian bitterling to a species as yet undescribed.

This species was described from Lake Müggelsee near Köpenik, Berlin, Germany and 3 syntypes are in the Museum für Naturkunde, Humboldt-Universität zu Berlin (or Zoologisches Museum Berlin, ZMB 3393).

Key characters

The presence of an ovipositor in females, the flank stripe and few pored scales are distinctive.

Morphology

Dorsal fin with 2-4, usually 3, unbranched and 7-11, usually 9, branched rays, anal fin with 2-4, usually 3, unbranched and 6-12, usually 9, branched rays, pectoral fin branched rays 10-13, and pelvic fin branched rays 4-8, usually 6-7. Pored lateral line scales 0-11, usually 4-6, along the flank 28-45, usually about 30-32 in some reports but 32-38 in Abdurakhmanov (1962), 37-40 in Holčík and Jedlička (1994), 30-38 in Pipoyan (1996b) and 35-40 in Holčík in Bănărescu (1999). Gill rakers 9-16, usually 10-14, reaching the raker below when appressed. Vertebrae 33-39. Pharyngeal teeth 5-5, rarely 5-4, 4-5 or 4-4. Teeth are elongate and narrowly compressed with a slight to strong hook at the tip and a very long, narrow and concave grinding surface below the tip. The gut has numerous coils. The chromosome number is 2n=48 (Klinkhardt et al., 1995).

Meristic values for Iranian specimens are:- dorsal fin branched rays 8(4), 9(52), 10(1) or 11(1); anal fin branched rays 8(18), 9(39) or 10(1); pectoral fin branched rays 10(3), 11(24), 12(29) or 13(2); pelvic fin branched rays 4(1), 6(3), 7(53) or 8(1); scales in lateral series 32(1), 33(2), 34(22), 35(29) or 36(4); pored lateral line scales 3(1), 4(7), 5(37), 6(11) or 7(2); total gill rakers 10(3), 11(14), 12(27), 13(10) or 14(4); pharyngeal teeth 5-5(20); and total vertebrae 34(9), 35(36), 36(12) or 38(1). Holčík and Jedlička (1994) give ranges in their Iranian sample of 33-41 for lateral series scales, 0-6 pored scales and 9-14 for gill rakers, in general agreement with data here.

Sexual dimorphism

Males develop a triangular or crescent-shaped patch of 7-20 tubercles on each side of the snout and small tubercles are found above the eyes. The female develops an ovipositor near the genital opening and it may be longer than the body. The flank stripe is longer and about one half wider in males compared to females, and male colouration is distinctive. Males are generally larger than females.

Colour

Males are particularly colourful in the spawning season: the top of the head and back are olive to bright green, reddish or dark violet, the iris is bright red, flanks are iridescent with violet and steel-blue colours most evident, the throat and belly are orange to blood-red, dorsal and anal fins are bright red and margined with black, the caudal fin is green at the base and yellow distally, and pectoral and pelvic fins are yellowish. Females are more yellowish and less iridescent than males in the spawning season. Outside the breeding season both sexes are similar in colour with a grey-green back, silvery flanks and yellowish belly. A grey-green to greenish-blue stripe originates under the dorsal fin and extends back to the tail base, broadening posteriorly. The dorsal fin is blackish and other fins reddish to yellowish. The dorsal, and sometimes the anal fin, has a dark interrupted stripe. The iris is silvery or yellowish. The peritoneum is dark.

Size

Reaches a reputed 18.0 cm but usually not more than 7.0-9.0 cm.

Distribution

Found from western Europe north of the Pyrenees and Alps to the Caspian Sea basin. In Iran, it is recorded from Astara to the Gorgan River including the Anzali Talab, Gorgan, Tajan, Babol, Haraz, Sardab, and Safid rivers (Derzhavin, 1934; Holčík and Oláh, 1992; Abbasi et al., 1999; Kiabi et al., 1999; Abdoli, 2000; Abdoli and Naderi, 2009). Also probably in the Araks River of Armenia (Pipoyan, 1996b). They may have been introduced into the Zarrineh River of Azarbaijan-e Gharbi based on photographs by Saber Shiri (pers. comm., 14 June 2008).

Zoogeography

Bohlen et al. (2006) assumed a continuous distribution of Rhodeus from Europe through Siberia to east Asia during the Pliocene, loss of the Siberian population in the late Pliocene or early Pleistocene, subsequent isolation, and later post-glacial expansion from several refugia in the Euro-mediterranean zoogeographic subregion. The Caspian Sea populations were not examined but seem to belong to R. amarus although the authors suggest a revisionary study is required for European bitterlings. Van Damme et al. (2007) consider this species was restricted to Ponto-Caspian and Aegean regions and its presence in western and central Europe is associated with the spread of Cyprinus carpio cultivation, and more recently, anthropogenic alterations to habitats and temperature changes.

Habitat

This species favours heavily vegetated areas of small lakes, ponds and slow-moving rivers, rarely in faster water. It is found in the lower reaches of rivers on the Caspian coast of Iran (Jolodar and Abdoli, 2004). The bottom is usually a fine sand or a thin layer of mud. Swan mussels (Anodonta) and freshwater clams (Unio) share this kind of habitat and are necessary for reproduction. Other genera of clams include Pseudanodonta, Cristaria, Margaritifera and Dahurinaia (Smith et al., 2004). Spawning occurs at water temperatures of 12-24°C, although 15-21°C is optimal (Holčík in Bănărescu, 1999), Van Damme et al. (2007) giving 23°C as optimal. Its distribution is limited by the 16°C July isotherm (Van Damme et al., 2007).

Age and growth

Maximum life span is 5-8 years, the higher values being uncertain (Holčík in Bănărescu, 1999). Maturity is attained during the second or third year, the life span of most individuals. Rarely some fish may mature as early as before the age of 1 year or as late as the fourth year. Growth is faster in ponds than in rivers. Males outnumber females at a ratio of 1.2-1.5:1, especially in spawning areas, although females apparently outlive males (Holčík in Bănărescu, 1999).

Food

Food consists of diatoms and detritus with aquatic insects and crustaceans being mostly accidental inclusions. They also take eggs of Rutilus rutilus, Cyprinus carpio and Rutilus frisii, and also their own eggs which fail to be deposited in the clam or mussel. Most gut contents in Iranian fish examined were plant fragments, filamentous algae, detritus and sand grains.

Reproduction

Reproduction takes place mainly in April and May, but can run from the end of February (in Azerbaijan) to August. The female develops an ovipositor from the genital opening, up to 6 cm long. In Iranian fish the best developed ovipositor was seen in a fish collected on 12 May when it extended back two-thirds along the caudal fin length. Eggs in this fish were 2.2 mm. Fish taken in September, October, November and January had short ovipositors progressively increasing in length with time. A fish taken on 4 July had an ovipositor extending only half way along the anal fin but no large eggs. Spring spawning is indicated but an ovipositor can be seen in varying degrees of development year round in adult females. Holčík in Bănărescu (1999) reports ovipositor length up to 126.5% of standard length in the Anzali Mordab in April.

The ovipositor lays eggs inside freshwater clams and mussels, using the excurrent siphon as the entry route. Apparently the flow of water and a high oxygen content out of this siphon encourage egg laying. Before egg laying, the female nudges the clam repeatedly to accustom the mollusc to stimulus so that it does not close up its shell. In the absence of clams, the bitterling does not become sexually mature. Clams and mussels with high numbers of bitterling larvae or filled with glochidia are avoided, as is Anodonta cygnaea, a mussel which is able to eject eggs and larvae and has low oxygen levels in its excurrent siphon (Kottelat and Freyhof, 2007). Males select and defend a particular clam against other males. They may headbutt each other or strike flanks, dislodging scales. Sneaking occurs despite this. The female deposits eggs 1-2 at a time and the male sheds sperm which are sucked into the clam on its feeding current. This process may be repeated with the same or different females. The female deposits about 40-100 eggs at each spawning and can spawn at least 5 times a day. The movement of the ovoid eggs through the ovipositor can be clearly seen as they are somewhat larger than the distensible ovipositor. Spawning bouts last 1-3 days with intervals of 5-7 days, variable with feeding conditions and temperature (Smith et al., 2004). Fecundity is up to 22,136 eggs and maximum egg diameter to 3.1 mm with width up to 1.52 mm. Eggs hatch 2-5 weeks later and the young leave the clam after 2 days when the yolk sac has been absorbed. Young fish leave the clam singly or in pairs. The fish gain the advantage that the eggs and young are protected inside the clam, even should the shore area dry out since the clam will move to deeper water. The clam is unharmed and is able to disperse its own young or glochidia by their attachment to the fins of the adult fish. Bitterlings seem however to have an immunological response to glochidia, having far fewer than other fishes that are less intimately associated with clams. Circumstantial evidence laid out by Smith et al. (2004) involving inhibition of free water circulation by the fish embryos, damage to clam gills, and increased consumption of oxygen by clams, indicate the relationships is a parasitic rather than a commensal one.

Smith et al. (2004) give a detailed account of reproduction, behaviour and development in this species.

Parasites and predators

None reported from Iran.

Economic importance

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in textbooks and aquaria and because it has been introduced outside its natural range. The species is used as a model in behavioural studies. Van Damme et al. (2007) regard it as a parasite on freshwater mussel populations in western and central Europe outside its native range, escaping glochidia infection and having low egg ejection rates.

Conservation

Lelek (1987) classifies this species as rare to vulnerable in Europe. Vulnerable in Turkey (Fricke et al., 2007).Kiabi et al. (1999) consider this species to be of least concern in the south Caspian Sea basin according to IUCN criteria. Criteria include medium in numbers, widespread range (75% of water bodies), absent in other water bodies in Iran, and present outside the Caspian Sea basin.

Further work

Although well-studied elsewhere, the biology of this species in Iran has not been investigated in detail.

Sources

Iranian material:  CMNFI 1970-0510, 21, 34.5-47.5 mm standard length, Gilan, Golshan River (37º26'N, 49º40'E); CMNFI 1970-0512, 25, 26.6-43.9 mm standard length, Gilan, Shalman River (37º08'N, 50º15'E); CMNFI 1970-0514, 2, 36.6-38.3 mm standard length, Gilan, Shafa River estuary (37º35'N, 49º09'E); CMNFI 1970-0520, 7, 28.9-45.0 mm standard length, Gilan, Astara River (ca. 38º25'N, ca. 48º52'E); CMNFI 1970-0526, 4, 30.7-43.2 mm standard length, Gilan, Safid River below Astaneh Bridge (37º19'N, 49º57'30"E); CMNFI 1970-0579, 8, 32.6-49.0 mm standard length, Gilan, Old Safid River estuary (37º23'N, 50º11'E); CMNFI 1970-0580, 5, 35.8-53.8 mm standard length, Mazandaran, river near Iz Deh (36º36'N, 52º07'E); CMNFI 1979-0265, 9, 16.1-47.4 mm standard length, Gilan, Anzali Mordab at Abkenar (37º28'N, 49º20'E); CMNFI 1979-0472, 6, 37.7-47.7 mm standard length, Mazandaran, stream west of Mahmudabad (36º37'N, 52º12'E); CMNFI 1980-0117, 3, 42.2-47.8 mm standard length, Gilan, Golshan River (37º26'N, 49º40'E); CMNFI 1980-0127,9, 32.6-43.0 mm standard length, Gilan, Caspian Sea near Hasan Kiadeh (37º24'N, 49º58'E).

Rhodeus ocellatus
(Kner, 1866)

Reported from the Karakum Canal and Kopetdag Reservoir in Turkmenistan (Shakirova and Sukhanova, 1994; Sal'nikov, 1995) as an exotic from China. May eventually be found in the Tedzhen (= Hari) River and Caspian Sea basins of Iran. No Iranian record.

Romanogobio
Bănărescu, 1961

There are about 17 species in the genus found from Europe to China with two species recorded from Iran. Members of the genus have a shallower body than the related Gobio, an elongated, usually cylindrical, caudal peduncle, epithelial keels on dorsal scales, an anus placed closer to ventral fins, absolute or average prevalence of caudal vertebrae over abdominal ones, and a higher number of preanal vertebrae (Naseka, 1996). Bănărescu in Bănărescu and Paepke (2002) notes that he considers Romanogobio as a subgenus and further work is needed to resolve this difference of opinion. Most literature is under the genus Gobio.

Romanogobio macropterus
(Kamensky, 1901)

Common names

 گاو ماهي (= gav mahi, probably in error for Neogobius and related gobies).

[Kur gumlagcisi in Azerbaijan; Kurinskii peskar' or Kura gudgeon in Russian].

Systematics

Gobio macropterus Kamenskii, 1901 was originally described from the Caucasus. The 14 specimens in the type series were deposited in the Georgian State Museum, Zoological Section, Tbilisi (ZMT) and in Kharkov University, Ukraine. Naseka et al. in Bănărescu (1999) cites types of Gobio macropterus in the Museum of the Caucasus (Tbilisi, presumably the Georgian State Museum) under numbers 128a (1 fish from Alazan) and 129 (4 fish from Kars-tschai) and 1-2 fish from the Kura (catalogue number unknown) in Khar'kov University. See below under R. persus for its changing status.

Key characters

The 7 branched dorsal fin rays are characteristic and this is the only Romanogobio species in the Caspian Sea basin of Iran (see also below under R. persus). It is separated from the related species in Iran (Gobio gobio) by having the body only slightly compressed and the caudal peduncle being cylindrical (caudal peduncle depth at anal fin insertion less than or about equal to caudal peduncle width) and by faint spots on the dorsal and caudal fins.

Morphology

Sexual dimorphism

Colour

Size

Distribution

Found in the Kura and Aras river basins of Armenia, Azerbaijan and Iran. Abdoli (2000) maps the middle Aras River and its lower Qara Chai tributary.

Zoogeography

Habitat

Age and growth

Food

The chitinous remains of aquatic insects comprised 30% of the gut contents from samples in the Kura River basin, caddisflies 21%, mayflies 14%, chironomids 12%, sevryuga eggs 6%, fish scales 3%, and much of the remainder was detritus at 20% (Abdurakhmanov, 1962).

Reproduction

Spawning takes place in May in Azerbaijan (Abdurakhmanov, 1962) and each female may spawn several times in a season. Eggs number up to 15,840 and are up to 1.62 mm in diameter. Water temperatures of 12-18°C are recorded for Georgia in April to June (Naseka et al. in Bănărescu, 1999).

Parasites and predators

Economic importance

Conservation

Vulnerable in Turkey (Fricke et al., 2007).

Further work

Sources ?check

Iranian material: CMNFI 1980-0155, 3, 45.9-69.3 mm standard length, Azarbayjan-e Khavari, Qareh Su near Ardebil (ca. 38º15'N, ca. 48º18'E).

Comparative material: CMNFI 1980-0806, 1, 93.4 mm standard length, Turkey, Kars Çayi, Kars (40º37'N, 43º05'E); CMNFI 1986-0007, 4, 49.4-84.0 mm standard length, Turkey, Kars, Kars Çayi north of Kars (41º00'N, 43º00'E).

Romanogobio persus
(Günther, 1899)

Common names

 گاو ماهي (= gav mahi, probably in error for Neogobius and related gobies).

[Persian gudgeon].

Systematics

This species was once thought to belong to the genus Rheogobio Bănărescu, 1961 variously regarded as a synonym of Gobio, a subgenus or a distinct genus. P. M. Bănărescu (in litt., 1984) and A. Naseka (pers. comm., 1994; in litt., 1995) place Gobio persus in the subgenus Romanogobius sensu Bănărescu (1961; 1992a) and Naseka (1996) elevates Romanogobio to a genus.

A. Naseka (pers. comm., 1994; in litt., 1995; Naseka et al. in Bănărescu, 1999) has studied R. persus from the Orumiyeh basin and from the Kars, Kura and Aras rivers of Turkey, Georgia and Azerbaijan. He distinguished two subspecies, R. p. persus from the Orumiyeh basin and R. p. macropterus from the Kura River basin which includes the Kars River in Turkey and the Aras River on the border of Iran and Azerbaijan. Gobio macropterus Kamensky, 1901 is now regarded as a distinct species (Naseka and Freyhof, 2004). 

The type series of Gobio persa consists of 7 specimens, 53.1-65.9 mm standard length, in the Natural History Museum, London from "Ocksa in the Gader Chai" in the description and "Ockra. NW Persia. Günther" in the jar, the former being more accurate (BM(NH) 1899:30:90-96).

Key characters

The 7 branched dorsal fin rays are characteristic and this is the only Romanogobio species in the Orumiyeh basin. The number of lateral line scales is 40 to 42 with modes of 40 or 41 (41 to 45 with modes of 42 and 43 in R. macropterus), total vertebrae are 37 to 40 with modes of 38 and 39 (38 to 42 with modes of 40 and 41), and the connection between the supraorbital and infraorbital head canals is usually absent (present). The Orumiyeh fish also have a shorter caudal peduncle, a shorter snout, a shorter barbel, and a longer predorsal distance.

It is separated from the related species in Iran (Gobio gobio) by having the body only slightly compressed and the caudal peduncle being cylindrical (caudal peduncle depth at anal fin insertion less than or about equal to caudal peduncle width) and by faint spots on the dorsal and caudal fins.

Morphology

The barbel is broad and fleshy, the mouth subterminal and horseshoe-shaped with thick, papillose lips. The lower head surface between the jaws and the sides of the head are papillose in a mature female.

Dorsal fin with 3, rarely 4, unbranched and 7, rarely 8, branched rays, anal fin with 3, rarely 2, unbranched and 5-7, usually 6, branched rays, pectoral fin branched rays 11-16, and pelvic fin branched rays 6-8. Lateral line scales 39-45. A pelvic axillary scale is present. The throat, breast and anterior belly are naked. Dorsal scales bear epithelial keels, usually one central keel and one to several lateral ones. Scales have a very anterior focus and few posterior radii. Gill rakers 0-6, small and irregularly spaced. Vertebrae 36-42. Pharyngeal teeth 2,5-5,2 or 3,5-5,3, or more rarely 2,5-5,1, 2,5-5,3, 2,4-5,3, 2,4-6,3, and 3,5-5,2. Teeth are strongly hooked at the tip, broadly concave or flattened below the tip, the surface sloping medially. The gut is s-shaped with a slight anterior loop to the left. The anus lies between the pelvic fins, remote from the anal fin origin. The karyotype is 2n=50.

Meristic counts in Iranian fish are: dorsal fin branched rays 7(22); anal fin branched rays 6(21) or 7(1); pectoral fin branched rays 11(1), 12(1), 13(6), 14(7), 15(6) or 16(1); pelvic fin branched rays 7(22); lateral line scales 39(3), 40(7), 41(3), 42(2) or 43(2); pharyngeal teeth 3,5-5,3(5); and total vertebrae 37(1), 38(7), 39(10), 40(4), 42(1).

Sexual dimorphism

Males and females bear irregular-shaped, elongate tubercles on the head. In a female specimen (and presumably males too), the pectoral and pelvic fin rays and adjacent membranes bear tubercles both dorsally and ventrally although the latter are less well developed. Males have longer pectoral and pelvic fins than females, reaching the pelvic fin and anal fin origin respectively in males. The snout to anus distance is more than half body length in females and about equal in males, head width is less than head depth at nape in females and equal in males, and snout length is longer than postorbital distance in females and about equal in males.

Colour

The upper flank is a light yellow-grey to brown with each scale outlined with, or partially filled in with, dark pigment, fading to a yellowish colour below. The mid-flank has 6-12 elongate to rectangular black spots. The lateral line pores may have small dark spots above and below reminiscent of Alburnoides species. The back also bears vague dark spots. Dorsal, caudal and pectoral fins bear up to 5 thin to broad discontinuous bars, the pigment being on the rays. Scales above the lateral line in mature males have longitudinal streaks. Peritoneum silvery with scattered melanophores.

Size

Reaches ?

Distribution

Found in the Lake Orumiyeh basin (Günther, 1899). Abdoli (2000) maps the middle and lower Talkheh, lower Tatavi and Zarrineh rivers in the Lake Orumiyeh basin.

Zoogeography

This species is part of a gobionine fauna found across Eurasia and this wide distribution may be suggestive of further work that could be done to clarify relationships of these fishes.

Habitat

Presumed to be similar to its relative.

Age and growth

Presumed to be similar to its relative.

Food

The principal food is aquatic insects and crustaceans but detritus and vegetation are also taken and presumably, though rarely, the eggs and fry of other fishes. Abdoli (2000) lists Trichoptera, Ephemeroptera and Chironomidae as food items.

Reproduction

Presumed to be similar to its related species. An Iranian specimen had relatively large eggs (1.2 mm) when captured on 23 June.

Parasites and predators

None reported from Iran.

Economic importance

None, although the recorded habit of feeding on fish eggs by its relative may be true of this species and  make it important in conservation of other species.

Conservation

Endemic to the Lake Orumiyeh basin where few specimens have been caught and deposited in museums. The numbers of this species in the wild are unknown.

Further work

The biology of this species, and its conservation status, have not been thoroughly investigated.

Sources check?

Type material: See above, BM(NH) 1899:30:90-96.

Iranian material: CMNFI 2007-0101, 12, 28.8-64.3 mm standard length, Azarbayjan-e Bakhtari, Tat'u River (ca. 36º54'N, ca. 46º07'E).

Genus Rutilus
Rafinesque, 1820

The roaches are found in Europe and western Asia where there are about 15 species (Bogutskaya and Iliadou, 2006). Three species are found in Iran.

The genus is characterised by having pharyngeal teeth in one row, usually 6-5, more rarely 6-6 or 5-5, with conical crowns on the anterior teeth and posterior teeth slightly hooked and truncated, scales are large to moderate in size, numbering 33-68 and with numerous fine circuli and radii on all fields, few and short gill rakers (17 or less), short gut, usually a light peritoneum, few to moderate numbers of dorsal and anal fin rays (7-13), the dorsal fin commonly having 4-5 unbranched rays, abdomen behind the pelvic fins rounded or with a slight but scaled keel, and various osteological characters (Bogutskaya and Iliadou, 2006).

Rutilus caspicus
(Yakovlev, 1870)

?

Caspian Sea species, R. rutilus freshwater residenth

Barzegar et al. (2008) record the digenean eye parasite Diplostomum spathaceum from this fish.

Rutilus kutum
Kamenskii, 1901

Common names

ماهي سفيد (= mahi safid or safid mahi, meaning white fish), sifid mahyi and asbalan mahi (in Gilaki), mahi safid daryacheh khazar or mahisephid-e-daryaye khazar (= Caspian Sea white fish), talaji (in Mazanderani).

[kutum, ak-balyk (= white fish) or ziyad in Azerbaijanian; akbalyk or kutum in Turlmenian; kutum in Russian; Southern Caspian roach; pearl roach; Caspian kutum].

Systematics

Leuciscus Frisii was originally described from the market in Odessa and the Danube, Dniester, South Bug, Dnieper and Don rivers, draining to the Black Sea.

Rework?

Rutilus frisii kutum Kamenskii, 1901 is the generally accepted Caspian Sea basin subspecies although J. Holčík (pers. comm., 1994) considers that this is not a good taxon (see also Holčík and Jedlička, 1994). It was originally described as Leuciscus Frisii var. kutum from the Caspian Sea, essentially in the southern part, spawning in the rivers and streams of Transcaucasia and Persia (Kura, Araxes streams of the Lenkoran district) and in lesser numbers elsewhere. This subspecies is distinguished from the type subspecies from the Black Sea by having fewer lateral line scales (about 55-58 versus about 60-66), shallower body (depth equal to or less than head length versus exceeding head length), anal fin longer than high versus shorter than high, lower lobe of caudal fin usually shorter than head versus longer (young R. f. kutum have this lobe as long as head), and dorsal fin as high as long versus higher than long (young R. f. kutum have dorsal fin higher than long). Kotlík et al. (2008) discuss divergence and gene flow between Black and Caspian Sea populations, the majority of the migrations occurring during the Pleistocene (ca. 10,000-1,800,000 years ago). The refugial populations in the Black and Caspian seas have diverged despite periods of migrations between them. Kavan et al. (2009) investigated the population genetic structure of Iranian and Azerbaijani fish using microsatellite markers. Most of the variation was within rather than between locations. Abdolhay et al. (2010) found independent populations existed in the Lemir, Safid, Shirud and Tajan rivers based on morphometric data. This has implications for the extensive stocking programmes for this species.

Leuciscus frisii caspius Lönnberg, 1900 described "from the Volga delta" has priority over Rutilus frisii kutum which may be a nomen nudum as it is listed in Radde (1899) as "Leuciscus Frisii Nordm. var. Kutum Kam." without a description, the description only appearing in Kamenskii (1901). However, the name caspius has not been used while kutum appears widely in the literature as well as being the common Russian name of the fish. Additionally, as mentioned above, there may be no need of a subspecific name. Kottelat (1997) considers kutum to be a distinct species as do Bogutskaya and Iliadou (2006) and Fricke et al. (2007).

Leuciscus friesii Kessler, 1870 from the Volga delta is presumably a misspelling.

Ghasemi et al. (2009) found the spring and autumn races of this species to be independent populations using microsatellite markers.

A hybrid of this species and Ctenopharyngodon idella has been bred at the Astaneh Ashrafie Fisheries Research Station (Sefidrud Research Station) and named "Samur" (Iranian Fisheries Research and Training Organization Newsletter, Tehran, 11:6, 1996; 18:6, 1997; Khara et al., 2002; Nouruz Fashkhami et al., 2002). Gynogenesis may have occurred. Artificial hybrids with Rutilus rutilus (possibly including R. caspicus) and Abramis brama have been bred in Iran (Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 39-40, 1996).

Key characters

This species is distinguished from the related Rutilus rutilus and R. caspicus by the higher scale count (usually >50) and the posterior part of the swimbladder being elongate and conical or pointed rather than rounded.

Morphology

Dorsal fin with 3 unbranched and 8-10, usually 9, branched rays, anal fin with 3 unbranched and 9-12, usually 10, branched rays, pectoral fin branched rays 16-19 and pelvic branched rays 8-9. Lateral line scales 47-68, mostly 55-58. Scales are regularly arranged over the body. A pelvic axillary scale is present. Scales have numerous circuli, numerous posterior radii, few but distinctively crowded anterior radii (more than in Rutilus rutilus and R. caspicus) and an almost central focus which is broken up into a network of lines. The posterior scale margin is crenulate and the anterior margin is wavy to crenulate but scale margins vary greatly between individual scales. Total gill rakers number 7-12 and are very short, hardly reaching the one below when appressed. Pharyngeal teeth usually 6-5, crowns rounded above a slender stalk, posterior teeth with a weakly hooked tip, posteriormost tooth margin may be serrated. The gut is an elongate s-shape. Chromosome number 2n=50 (Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 43, 1996; Noruz Fashkhani and Kosroshahi, 1995; Klinkhardt et al., 1995). Adibmoradi and Sheibani (2002) carried out a histological study of the brain of this species and concluded, in part, that vision was sharp and the fish were good chasers of food.

Meristic values for Iranian specimens are:- dorsal fin branched rays 8(1), 9(58) or 10(1); anal fin branched rays 9(24), 10(35) or 12(1); pectoral fin branched rays 16(15), 17(30), 18(14) or 19(1); pelvic fin branched rays 8(49) or 9(11); lateral line scales 47(1), 49(1), 50(1), 51(1), 52(7), 53(8), 54(6), 55(5), 56(11), 57(7), 58(7) or 59(5); total gill rakers 8(1), 9(21), 10(25) or 11(13); pharyngeal teeth 6-5(23), 5-5(1) or 5-4(1); and total vertebrae 40(1), 41(7), 42(34), 43(3) or 44(2).

Sexual dimorphism

Females are larger than males. The breeding tubercles are evident in males and may develop as early as the end of summer before the spring spawning season in the following year. Large tubercles are found on the top of the head above the eye level, behind the eye, between the eye and the mouth and on the snout as well as on upper flank scales. Tubercles are white in contrast to the dark head. Imanpour and Shirmohammadli (2008) found that fish from the Gorgan River showed correlations between increase in number of tubercles and total length, sperm volume, and gonad weight but not with sperm motility, spermatocrit, sperm density and gonadosomatic index.

Colour

Small specimens up to a year old are silvery on the flanks and belly and the back is steel-grey to pale brown or pale olive. In adults, back scales are circled with black and there is a strong contrast between the back and flank. The anterior part of each flank scale, particularly those of the lateral line, is darkly pigmented. Lateral line scales may have two dots, one above and one below the opening as in some Alburnoides spp. but not as pronounced. The sides of the head are silvery with some yellow and darker pigment, the latter particularly in front of the eye. Adults are a bright silvery on the flank. The belly is pearly-white. The iris is silvery, with some spots, and with a marked dark spot above. The dorsal and caudal fins have some grey and a faint orange tint while the pectoral, pelvic and anal fins are colourless to lightly pigmented with black. The pectoral fin may be orange.

Size

Attains 66 cm body length and 4.065 kg in Iran (Farid-Pak, 1968a). Ouseley (1819-1823) ate one almost 3 feet long (ca. 0.9 m). In Iran during the 1950s, catches were 36-67 cm long (Farid-Pak, no date). Reaches 7.0 kg in weight.

Distribution

Found in drainages of the Caspian Sea. In Iran, it is found along the whole Caspian coast, entering almost all the rivers to spawn including Atrak, Gorgan, Gharasu, Tajan, Babol, Haraz, Sardab, Aras, Tonekabon, Pol-e Rud and Safid rivers, the Anzali Talab, Gorgan Bay, and in the southeast, southwest and south-central Caspian Sea (Nedoshivin and Il'in, 1929; Kozhin, 1957; Holčík and Oláh, 1992; Riazi, 1996; Oryan et al., 1998; Abbasi et al., 1999; Kiabi et al., 1999; Abdoli, 2000; Lasheidani et al., 2008; Abdoli and Naderi, 2009).

It is also found in Valasht Lake, Mazandaran (landlocked and introduced about 1850 to provide food for the royal family which had a summer residence there) where it possibly hybridises with native Alburnus chalcoides (Armantrout, 1980).

von den Driesch and Dockner (2002) record this species as faunal remains in an excavation at the Great Mosque in medieval Siraf on the Persian Gulf coast. This may well be a misidentification but, if not, indicates that a fondness for this species has a long history considering that a preserved fish or its remains had to be transported all the way from the Caspian Sea.

Zoogeography

A European and western Asian species with its origins in a Danubian or Sarmatian fauna.

Habitat

This species is migratory, spawning in rivers in March-April and returning to the Caspian Sea. Rezavi (1997) found three populations in Iran, one autumn and two spring populations. Il'in (1927a) records schools of fish entering the Anzali Bay as numbering in the several tens of thousands. On one fishing ground next to the Djifrud in February 1914, 12,000 fish were taken simultaneously. However, while the majority of fish migrate into lowland rivers not far from the sea to spawn among bulrushes and cattails, some migrate far upstream into the mountains of Tavalesh (= Talish) and Gilan where spawning conditions are very different. These fish may well overwinter in the river (Derzhavin, 1934). Such fish reached altitudes of about 1000 m before environmental changes inhibited the migration. Young fish migrate downstream, the migration ending in August, and in Azerbaijan enter the sea within 20-50 days. Holčík (1994; 1995) stated that Iranian young from the Anzali Mordab never entered the sea but remained in fresh or brackish water for 1-2 years. Riazi (1996) reports that this species migrates into the Siah-Keshim Protected Region of the Anzali Mordab. Young fish descend the Atrak River and feed in the sea at depths of 4.0-8.5 m. They winter in Iranian waters (Savenkova, 1985). Some fish are reported at depths of 36.6-53.0 m in the Iranian Caspian Sea (Knipovich, 1921).

Age and growth

Azari Takami et al. (1990) described the biology of this species in Iran. Males normally mature between their third and fourth year, sometimes earlier, females during their fourth year. Life span is at least 9 years in Dagestan (Shikhshabekov, 1979) and 8 years in Iran (Holčík and Oláh, 1992) and maximally about 12 years (Afraei Bandpei et al., 2010). Spawners are 3-8 years old and the principal age groups are 4-5 years for males and 5-6 years for females in the Anzali Mordab (Holčík and Oláh, 1992; Holčík, 1995), older than the 3-4 years of fish reported in 1970-1971. However, recently males are maturing at age 2 and females at age 4 with most spawners at age 3 and 4 years respectively (Holčík and Oláh, 1992). The average size of mature females (700 g) has been decreasing (Bartley and Rana, 1998b). Males grow faster than females until about the third year of life and then more slowly. Males are smaller and have a shorter life span than females (Holčík and Oláh, 1992). An annual increase in length of 17% over the first four years was observed in tagged fish, decreasing to 4.7% annually in subsequent years (for weight the figures are 61.2% and 25.7%). Sex ratios in spring were 3.08 males:1 female and in autumn 0.78 males:1 female (Annual Report, 1995-1996, Iranian Fisheries Research and Training Organization, Tehran, p. 55, 1997).

A series of studies on this species, apparently sampling in somewhat different areas or from different commercial fish catches give several growth parameters as follows. Abdolmaleki and Ghaninejad (2007) examined the commercial catch in 2003 -2004 and found mean fork length was 36.7 cm (range 21-69 cm) and mean age was 3.82 years (range 1-8 years). Age groups 3-5 years comprised 87% of the total catch age composition. von Bertalanffy growth parameters were L = 70.1 cm, K = 0.138/year, t0 = -1.557 years, total mortality = 1.1/year, natural mortality 0.28/year and fishing mortality 0.83/year. Abdolmaleki et al. (2007) sampling the commercial catch found the total catch of kutum, including poaching, was 6612.5 t of 43.3% of all commercial bony fishes in the 2004-2005 season. Mean fork length was 38.2 cm, age range was 1-8 years and mean age was 4.2 years. Age groups 3-5 years made up 85.5% of the age composition. von Bertanffy growth parameters were  L = 60.7, K = 0.15 year-1, t0 = -1.75 years. Instantaneous total mortality coefficient (Z) was estimated as 0.88 year-1, instantaneous natural mortality coefficient (M) was 0.31 year-1, and annual fishing mortality coefficient (F) was 0.52 year-1. Growth rate has decreased in comparison to previous years probably due to non-selective artificial breeding over more than 20 years. Afraei Bandpei et al. (2008) surveyed biological characteristics in samples from Iranian coastal waters. Average fish size was 11.73 cm fork length, range 8.9-29.0 cm, average weight was 22.74 g and range 8.8-321.2 g. Females were larger than males (12.27 cm and 32.26 g on average compared to 11.03 cm and17.07 g). Sex ratio was 1.3:1 male:female. Overall condition factor was 1.23. Afraei Bandpei et al. (2010) also recorded features of fish from Iranian coastal waters but found a range in size of 21 to 58 cm and 148 to 2450 g, average fish size  was 39.0 cm fork length and 830.3 g for females and 38.2 cm and 719.1 g for males, sex ratio was 0.65:1 in favour of females, and condition factor varied from 1.2 to 1.3. The von Bertalanffy growth parameters were L = 63.00, K = 0.21 year-1, t0 = -0.88 years for both sexes, L = 62.03, K = 0.21 year-1, t0 = -0.80 years  for females, and L = 54.52, K = 0.27 year-1, t0 = -0.75 years. the growth performance index was 2.89 for both sexes. Life span was 9 years in females ands 8 years in males with a mean age of 3.99 years overall and four-year-old fish dominating in the sample. The length-weight relationship value b was 3.02 indicating growth is isometric. Growth was seen to be rapid in this southern Caspian Sea population, attributed to warm temperatures, available food resources and the brackish water environment. Historically there has been a decrease in size of fish over three decades as length and weight used to be 67 cm and 7 kg. Afraei Bandpei et al. (2010), in another study, investigated the population dynamics in Iranian fish during the fishing season October to April, and found the von Bertalanffy growth parameters to be L = 59.85 cm fork length (FL), K = 0.27 year-1, C = 0.25, WP (winter point) = 0.40, instantaneous total mortality coefficient (Z) was estimated as 1.28 year-1, instantaneous natural mortality coefficient (M) was 0.46 year-1, instantaneous fishing mortality coefficient (F) was 0.82 year-1, current exploitation rate (E) was 0.64, mean length at first capture (Lc) was 36.8 cm FL, maximum exploitation rate (Emax) was 0.76, and the current exploitation rate (E) was less than Emax. The highest growth oscillation occurred in December and was attributed to active feeding before wintering and migration to deeper waters. These figures indicate that the fish population is moderately exploited and agree in general with other studies in the Caspian Sea. Variations in growth performance could be due to food availability, ecological conditions, geographical changes and genetic variability.

Fish may spend 1 or 2 years in fresh water after hatching (Holčík and Oláh, 1992) and this affects growth, 1 year fish growing more quickly than those spending 2 years in fresh water, maturing earlier and having a shorter lifespan. Growth in the Anzali Mordab is slower than it was 20 years prior to the report of Holčík and Oláh (1992), probably owing to a higher population density and more competition for food resources. The commercial catch in Iran was 3-7 years old, 39.0-57.1 cm long and weighed 613-2525 g (Razivi et al., 1972).

Growth in Iranian fish farms is up to 1.5 g in 8 weeks after a hatching rate of 75% (Aslaanparviz, 1994).

Food

Zarbalieva (1987) provides data on feeding of this species in the Caspian Sea off Azerbaijan. Fish concentrate on a sandy-shellrock bottom and remain there for most of the year to feed. The crab Rhithropanopeus harrisii dominates in the diet, 67.9-93.7% by weight. Molluscs, mainly Cerastoderma lamarckii, comprise 30% by weight of the food of fish 30-40 cm long. Fish larger than 40 cm seldom take molluscs but occasionally Clupeonella spp. Molluscs used to be the main diet item of this fish. Juveniles in the Anzali Mordab of Iran feed mostly on phytoplankton in contrast to the zooplankton reported for Azerbaijan fish (Holčík, 1995). This is a consequence of the poor productivity of this lagoon. Small Iranian specimens from the Caspian Sea had bivalve shell remains, plant remains and in one case a worm. Oryan et al. (1998) state that Cardium is the main food of this species on the eastern and western coasts of Bandar Anzali. Crabs and Balanus are also important. The hepatosomatic index is highest in February and March, the prespawning period. Afraei Bandpei et al. (2008) found diet in the southern Caspian Sea was Mytilaster (48.46%), Gastropoda (31.84%), Cerastoderma (19.65%) and Balanus (0.05%).Afraei Bandpei et al. (2009) sampled fish from the commercial catch caught from October to April and found the diet was dominated by bivalves (59%) with Cerastoderma lamarcki the dominant prey (57%).Other foods were cirripeds (21%), gastropods (13%), malacostracans (3%), fish eggs (2%), amphipods (1%) and filamentous algae (1%). Fish fed on a wider variety of food groups in November compared to other months and the lowest feeding activity was in January and April, the latter being the peak of the spawning season.

The hybrid with Ctenopharyngodon idella fed principally on macrophytes with phytoplankton as a secondary food in pond sin Gilan (Khara et al., 2002).

Reproduction

Fish spawn annually according to an Iranian study by Azari Takami et al. (1990) and probably return to their river of birth. Temperature, and probably river flow, are the factors determining the entrance of fish into the rivers on the spawning migration. Male fish run before females. Adeli Mosabbab and Piri (2005) report spawning in Iranian rivers from the end of March to the middle of April, beginning at 10°C. A large female, 60 cm long, 3.24 kg and 7 years old contained 124,712 eggs but some large females may have over 250,000 eggs (see also Farid-Pak (1968a) where fish 38-40 cm long have a mean value of 53,900 eggs, ranging up to 174,400 for fish 62-64 cm long; Yousefian and Mosavi (2008) give March to May as the spawning season. Aminian Fatideh et al. (2008, 2009) determined stages of sexual maturity of migrating fish, 87% achieving stage V in April and 94% stage V in June. Abdurakhmanov (1962) gives a fecundity of 290,000 eggs for fish in Azerbaijan). Maximum egg diameter is 2.0 mm (Farid-Pak, 1968a). There are two spawning migrations, winter and spring, and so two forms or stocks of this species (Azari Takami et al., 1990). There may be three stocks based on electrophoretic studies of blood proteins associated with the two spawning migrations (Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 41, 1996). The winter form enters rivers with emergent or submerged aquatic plants at the end of autumn or the beginning of winter. Eggs are deposited on these plants. The principal rivers were the Nahang Roga, Pir Bazar Roga, Sowsar Roga, and Anzali Roga (all draining the Anzali Mordab) but the stock has declined with overfishing and destruction of the habitat. The Safid River is listed as the main spawning ground by the Caspian Sea Biodiversity Database (www.caspianenvironment.org). The spring form enters rivers at the end of winter or the beginning of spring and spawns on gravel or sand. Adults return downstream after spawning, in the early morning. The run occurs both by day and night but increasing water clarity near the end of the spawning season limits runs to the night.

The migration into the "Dinachal" (? = Denya Chal) River occurred on 1 November in 1975, when sexually immature fish were caught. By 11 February, 5 spawners were caught and the peak migration occurred on 26 March when 3,845 fish were caught at a water temperature of 10.5°C. The migration was continuous until 11 May. Water temperatures between 28 March and 23 April in 1976 and 1977 varied between 9.1°C and 13.5°C when peak migrations occurred in the "Havyg" (= ? Haviq) River. Here the spring form of this species started to migrate at 6°C and the optimum water temperature appears to be between 11 and 13°C. Yousefian and Mosavi (2008) found a spawning temperature range of 5 to 21°C centred on 11°C. Fertilisation rate in 4 rivers was 75-92% and hatchability was high. Rivers with high turbidity had low hatchability because of egg damage. Further details of natural and artificial reproduction are given by these two authors.

The main migration (99%) in the Anzali Mordab began in February and lasted 3 months at 8-10°C with a much smaller run (only about 1%) in November-December. The Mordab is now overgrown with reeds or has a silty, vegetated bottom and is no longer a spawning site (Holčík and Oláh, 1992), an economic and poetic loss. Holčík and Oláh (1992) translate Il'in (1927a) on Anzali Mordab mahi safid "During this time the reeds rustle as if of wind as the fish rubs pushing their flanks to the reed stalks or pushing forward among reeds standing in rows".

Khaval (1998) reports a spawning migration into the Safid River despite construction, sand removal and pollution. The migration peaks in the second half of March at a water temperature of 11°C.

Each female is flanked by two males at spawning in shallow water. Sometimes the backs of the fish protrude from the water. Females sharply rub their lower abdomen and pectoral area on the gravel bottom, males and females convulse, and eggs are shed and fertilised. Eggs are adhesive and hatch in 20 days at 10-14°C. Embryonic development in glass incubators at 14-16°C is described by Parivar et al. (1993). Cleavage and gastrulation begin within 24-30 hours, hatching at 216 hours (10 days after fertilisation) and resorption of yolk is complete at day 16. Some males and females are injured through contact with the gravel stream bed. Embryonic development takes 10-15 days at 8-16°C and 5-6 days at 20°C (Aslaanparviz, 1994). Young fish go down to the sea in summer according to some works although Holčík and Oláh (1992) believe that they stay in fresh water or brackish river mouths for 1-2 years before entering the sea.

Kousha et al. (2007) examined some aspects of reproductive endocrinology in this species, specifically on sex steroid-binding protein in plasma. Najafipour et al. (2008) examined 17-αhydroxy progesterone hormone levels, egg diameters and liver weight in fish caught at sea and in the Chalvand River near Astara. These factors were used to indicate physiological indices of reproduction, hormone levels and egg diameters increasing in fish caught in the river, while liver weight decreased. Egg diameters increased from 1.55 to 2.14 mm. Sabet et al. (2009) studied gonad steroid levels in fish from the Safid River and found estradiol and testosterone reached their highest levels in April and were closely correlated to ovarian development and the gonadosomatic index.

Parasites and predators

Mokhayer (1976b) records the digenetic trematodes Aspidogaster limacoides and Asymphylodora macracetabulum. Eslami and Kohneshahri (1978) report various helminths from this species in Iranian waters. These are the monogenean Diplozoon paradoxum, the aspidogastrean Aspidogaster limacoides, the digenean Asymphylodora kubanicum and larvae of the nematode Anisakis sp. This fish therefore is potentially a source for human infection with Anisakis. Mokhayer (1989) reports metacercariae of the eye fluke, Diplostomum spathaceum from this species in Iran, which can cause complete blindness and death in commercially important species. Jalali and Molnár (1990a) record the monogeneans Dactylogyrus frisii from this species in the Khosk (= ? Khoshk) River (Caspian basin) and D. rarissimus and two monogenean species, Dactylogyrus spp., from the Safid Rud. Jalali and Molnár (1990b) record the monogeneans Dactylogyrus frisii and D. rarissimus at fish farms in Iran. Molnár and Jalali (1992) report the monogeneans Dactylogyrus suecicus, D. haplogonus and D. turaliensis from this species in the Safid Rud. Gussev et al. (1993b) record the monogenean Dactylogyrus haplogonus from this species in the Safid Rud. Masoumian and Pazooki (1998) surveyed myxosporeans in this species in Gilan and Mazandaran provinces, finding Myxobolus bramae. Safari and Khandagi (1999) record Clostridium botulinum from 2.2% of fresh and smoked samples of this species in Mazandaran Province. Shamsi and Jalali (2001b; 2001c) detail monogenean parasites for Safid Rud and Caspian Sea samples, including 6 species of Dactylogyrus. Jalali et al. (2005) summarise the occurrence of Gyrodactylus species in Iran and record G. prostae and G. sp. in fish from the Safid River. Khara et al. (2008) found the eye parasite Diplostomum spathaceum in this fish from Boojagh Kiashar Wetland in Gilan. Barzegar et al. (2008) also record the digenean eye parasite Diplostomum spathaceum from this fish.

Amini (2006) records Gyrodactylus sp., nematodes, Diplostomum sp., Trichodina sp., Aeromonas sp. and Hydrophila sp. in fingerlings from hatcheries in Iran. The latter two parasites caused a heavy mortality in the Shahid Rajaii hatchery ponds.

Tavassoli and Moghir (20020 describe a squamous cell carcinoma in the oral cavity of this species, the first record for the Caspian Sea.

Economic importance

Safid mahi is the most popular fish in Iran with the highest economic value (Azari Takami et al., 1990) and Afraei Bandpei et al. (2008) record 10,773 fishermen making a living from its capture. More than 70% of bony fish caught in the coastal Caspian Sea of Iran are this fish (Yousefian and Mosavi, 2008) or 78% and 76.6% of the income The average annual catch from 1991 to 2001 was about 9600 t (Afraei Bandpei et al., 2010).

It may be available out of season as mahi qachaq or bootleg fish, an indication of its popularity. Batmanglij (1999) notes that this fish is baked and served with herbed rice. A stuffing consists of garlic, parsley, tarragon, scallions, coriander, mint, ground walnuts, barberries, raisins, lime juice, salt and pepper, sautéed in butter (he also incorrectly refers to mahi-e safid as striped bass). In 1996 a news report stated that "In the port city of Anzali, one small white fish sells for $10" (http://www.iran-e-azad.org/english/boi/03400201.96). In Rasht, 60% of the inhabitants consume no other fish and "86% of ladies in Rasht can only cook one kind of seafood" (Khairkhah, 1994). Ouseley (1819-1823) reported that this species "seemed most abundant, and was found in all the great rivers of this country near the sea; for several days it had furnished the principal dish of my dinners and often of my breakfasts". Holmes (1845) recorded catches with cast-nets in the Anzali Mordab over 150 years ago which were even then worth £1400-1500. O'Donovan (1882) reported that a small stream in the Atrak River drainage was so crowded with this species that individuals could only move by floundering and jumping over one another, and the horses in crossing the stream, trod them to death by scores. Lönnberg (1900) reported an annual catch of some millions in Persian waters.

It is caught in rivers and lagoons and by large, mechanically hauled beach nets in the Caspian Sea. Sea nets can be 1500 m long and 18 m deep. The fishing season begins in October and reaches a maximum between 20 February and 10 March, ready for the "Now Ruz" or New Year celebrations when many Iranians eat this fish with rice (Emadi, 1979).

The roe of this species is also eaten, salted or unsalted (the fish are called asbalan mahi and the roe shur-e asbal in Gilaki, the local dialect of Gilan). Whole female fish are soaked for one year in a mixture of salt and madder in special clay jars. The jars are traditionally buried in the ground and hermetically sealed but a modern technique has the roe removed from the fish after 20 days. Esmaeilzadeh et al. (2004) studied the nutrient composition and marinade qualities of this fish and compared them to those for grass carp (Ctenopharyngodon idella), the latter being preferable according to the organoleptic properties. The marinades could be stored for 6 months at 10ºC. Alipour et al. (2009) studied teh effects of different brine concentrations and temperatures in traditional smoking of this fish, finding a brine concentration of 26% gave the better quality and taste.

The average weight of fish caught in the Anzali Mordab in the early years of this century was 2 kg, the catch in the 1914-1915 season was 47,000 fish and in 1913-1915 123,000 fish. On 26 February 1914 a single haul in the sea near Anzali took 41,045 fish (Il'in, 1927a; Berg, 1948-1949). Il'in (1927a) estimated a yearly catch in Anzali Bay to be 3-4 million fish, a figure conflicting with these other reports.

Nevraev (1929) gives catches for various fishing regions in Iran in the early twentieth century. For the period 1901-1902 to 1913-1914 the catch in the Astara region was 0 to 29,053 individuals, for 1901-1902 to 1917-1918 the catch in the Anzali region was 2565 to 124,195 individuals, in the Safid River region from 1904-1905 to 1917-1918 the catch was 100 to 31,799 individuals, and in Astrabad (= Gorgan) region from 1900-1901 to 1912-1913 the catch was 4000 to 323,500 individuals. The total catch for Iran in the 1914-1915 season was 443,000 fish. The catch in Iran from 1956/1957 to 1961/1962 varied from 197,884 kg to 2,066,580 kg (Vladykov, 1964), from 1965/66 to 1968/69 it varied between 159 and 1252 tonnes (Andersskog, 1970), from 1963/64 to 1968/69 it varied between 121.3 and 1252 t (RaLonde and Walczak, 1970b; 1972), from 1987 to 1991 it varied between 3500 t and 8855 t (Holčík and Oláh, 1992), and between 1989 and 1998 it varied between 11,792 kg and 14,336 kg (Caspian Environmental Programme, 2001a) - catch figures are at variance with each other. Holčík and Oláh (1992) report a catch of 3107 kg in the Anzali Mordab for 1990 and for 1932-1964 a range of 95.1-3488.9 t. They are also caught in rogas and inflowing rivers of the mordab in late winter and early spring. Moghim et al. (1994) estimate that coastal areas of the southern Caspian Sea have a total biomass of 24,000 t with a maximum sustainable yield of 7000 t. In 1993-1994 the total catch of this species, including the illegal catch, was 11,175 t with the total stock estimated at 25,400 t and a maximum sustainable yield of 9300 t. More than 25% of the catch was young fish indicative of non-standard methods being used (Annual Report, 1995-1996, Iranian Fisheries Research and Training Organization, Tehran, p. 19-20, 1997). In 1994-1995, the biomass of this species in Iran was 241,000 t (sic, probably 24,000 t) and the maximum sustainable yield was 9000 t (Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 37, 1996). About 62% of the bony fish catch in the Caspian Sea of Iran in 1993-1994 was this species with the mullet Liza aurata second at 22% (Annual Bulletin 1993-94, Iranian Fisheries Research and Training Organization, Tehran, p. 83, 1995). The catch from beach seine cooperatives along the Iranian coast was 8477 t in in 2004-2005, a 2500 t decrease over the previous year, and 45.5% of the total bony fish catch. Catch per unit effort  was 93.3kg/set. The biomass of this species in Iranian coastal waters for 2003-2004 was estimated at about 25,000 t (Abdolmalaki, 2006a; Abdolmaleki and Ghaninezhad, 2007)).

Bartley and Rana (1998a; 1998b) comment that the fishery collapsed in 1980 but has risen from 500 tonnes in 1981 to around 10,000 t in 1996 after restocking from around 400,000 fingerlings/year in 1981 to around 142 million/year in 1997. Rana and Bartley (1998a) note that 7 million 1 g fingerlings were released into the Caspian Sea in 1997 which contradicts their earlier report. There is only a state supported stocking programme but ERM-Lahmeyer International GmbH, DHI Water & Environment and GOPA Consultants (2001a) note that this fish is being successfully managed in Iran while it has been fished almost to extinction in waters off Daghestan and Azerbaijan. Market price is high at about U.S.$5.00/kg, five times the price for silver carp. Export prices are higher, 700 g of smoked "mahi-sefid" cost £21.00 in 2004 (www.superhormuz.com, downloaded 19 January 2004). However this species is difficult to culture beyond the 40-50 g stage and growth is slower than carp.

The conflicting ranges seen in Andersskog and RaLonde and Walczak are typical of the wide variations in reports; figures can only be taken as general guides for many Iranian fisheries. Abdolmalaki (2006b) states that fluctuations are due to destruction of spawning grounds, overfishing and release of fingerlings. The mean catch sizes were 3110 tonnes in 1937-1947, 990 t in 1967-1977 and 8505 t in 1987-1997. A minimum catch of 121 t was taken in 1964 and a maximum of 11,175 t in 1994. Catch-per-unit-effort also showed high variation,. The calculated stock biomass was 1300 t in 1071 and between 18,489 and 25,400 t between 1990 and 2000. The mean biomass in the past 10 years was 22,750 t, a 17-fold increase over the year 1971. The catch in the previous 10 years was 35-46% of the annual stock. A decrease in stock size during 1998-1999 was attributed to an exploitation rate being more than maximum sustainable yield, a decreased mean weight of released fingerlings and a lowered return rate.

Catches in the Bandar-e Anzali area have been as high as 5,480 t in 1939-1940 but fell to 85 t in 1961-1962 (Vladykov, 1964; RaLonde and Walczak, 1972) but have risen again as indicated above. This is attributed to a massive stocking effort with 170 million fingerlings released in Iran in 1991 (cf. Emadi (1993a) below), about a quarter in the Anzali Mordab (Holčík and Oláh, 1992). The Sari hatchery produced 400 million "white fish" over the previous 10 years (Tehran Times, 30 May 1998). Fingerling production in 1996 was 142.1 million (Bartley and Sana, 1998a). Sea ranching increased the yearly catch to 8500 t in 1991, the highest recorded catch in the past being 5850 t in 1940 (Emadi, 1993a). Natural stocks in the past were very high. Migrating fish were so dense at the Mordab mouth that they were caught in buckets and jumping fish literally fell into boats.

Savenkova (1990) comments on the marketing of this species in southwestern Turkmenistan. Salehi (2003) analysed the economics of production of finglerings in Iran. The cost of labour was 50%, feed and fertiliser 20%, maintenance 10%, and harvesting, handling and releasing 6%. The average cost per fingerlings was 37 rials in 2001 (rising to 123 rials in 2003 (Salehi, 2008)). As the average rate of fingerling return was 8.3% and the average weight and age of commercially caught fish was 815 g and 3.7 years, it was expected that 19,257,494 individuals weighing 16,000 t would be harvested over the Iranian year 2004-2005. The value would be an estimated 345 billion rials at the 2001 price (8050 rials to a U.S. dollar in 2001; $42.86 million). Salehi (2008) estimated the wholesale price of the catch as 505 billion rials in 2004 and reviewed the comparative economics of fingerling production for 2001-2003.

Experimental culture of triploid mahi safid has been carried out at the Gilan Fisheries Research Centre (Iranian Fisheries Research and Training Organization Newsletter, 2:2, 1993) to increase fish weight for exploitation but apparently was unsuccessful although the techniques worked with grass carp, Ctenopharyngodon idella (Annual Bulletin 1993-94, Iranian Fisheries Research and Training Organization, Tehran, p. 70-71, 1995). In addition, monoculture and polyculture of this species has been investigated, the latter with grass carp (Ctenopharyngodon idella) and silver carp (Hypophthalmichthys molitrix) (Iranian Fisheries Research and Training Organization Annual Report, 1992-93; Danesh Khoshashi, 1997). From an average initial weight of 7 g, monocultured fish weighed 158 or 177 g on average at the end of each of two one-year periods, polycultured fish weighed 158 or 168 g, and maximum weight attained was 250 or 300 g (Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 38, 1996). Danesh Khoshasi (1997) gives figures of 177 g for mono-cultured fish after 6 months in ponds, poly-cultured fish weighed 185 g (with a maximum of 300 g) while stocking with 7 g fish gave 168 g and 192 g for mono- and poly-culture fish respectively (maximum weight 250 g)(these two reports on the same experiment have confusing figures). The fish were fed on pellets especially made for this species, nitrate and phosphate fertiliser and cow and chicken manure were added to the ponds, and water temperature was 15.0-27.64°C.

Experimental studies have been carried out on different lipid levels in food fed to cultured fry, a level of 12% increasing growth levels (Novirian et al., 2007). Ghanbari et al. (2009) investigated  production of a bacteriocin or anti-bacterial from intestinal bacteria of this fish species. This bacteriocin has potential for use as a biopreservative for food.

This species has been used in Iran for experimental studies and analysed for pollutants, e.g. on the acute LC50 and bioconcentration of mercuric chloride (Gharaei et al., 2006; Gharaei and Esmaili-Sari, 2008); on the toxicity and LC50 of phenol and 1-naphthol (Shariati et al., 2004); the effects of anionic detergents and diazinon on the LC50, both separately and in combination (Tehranifard et al., 2002; Tehranifard et al., 2007); Razavilar and Tavakoli (2006) on the prevalence of human toxigenic Clostridium botulinum and the need for food safety control measures; Foroughi et al. (2007) on mercury content which varied between tissues, was not significantly different between sexes and was lower than permissible limits; Vaezzadeh et al. (2008) found the levels of the pesticide heptachlor in fish from Hashtpar and dieldrin in fish from Kiashahr could have a health risk to consumers; etc.

The hybrid with grass carp, Ctenopharyngodon idella, reached an average weight of 100 g and a length of 22 cm after 5 months in the first report cited above and 6.7 g and 9.17 cm after 4 months in the second report. Larvae were fed live food twice a day and hybrids larger than fingerling size ate grass. The hybrid phenotype resembled the safid mahi but was a herbivore. It was expected that this hybrid would be used as a new culturable "species".

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in aquaculture and as food.

Conservation

Azari Takami et al. (1990) list the following reasons for a decline in the commercial catch of this species:- a) regression of the Caspian Sea which decreased the surface area of the Anzali Lagoon and increased the growth rate of aquatic vegetation, b) mechanisation of farming and a consequent increased demand for irrigation water leading to reduced river flows during the spawning migration, c) use of fertilisers and pesticides in rivers draining into the Anzali Mordab (and pesticides and herbicides such as Diazinon, Malathion, Machete and Saturn have a highly toxic effect on fingerlings of this species (Piri et al., 1999); and the herbicide butachlor reduces sperm volume and increases abnormal sperm (Lasheidani et al., 2008)), d) pumping of river water for irrigation causing mass mortalities of fry, e) industrial development increasing the pollution load, and f) excessive catches of adults to the extent that all spawners in a river were taken. Emadi (1979) added such factors as road construction and the removal of sand from banks and river beds, erosion caused by felling trees and shrubs along river banks and in the mountains, construction of bridges and raising of their substructures which formed barriers to migration, and climatic changes. Illegal fishing and non-standard nets threaten the stocks while fingerling release (120-140 million) and improvement of natural spawning areas through rises in water level have contributed to stock increases (Annual Report, 1995-1996, Iranian Fisheries Research and Training Organization, Tehran, p. 19-20, 1997). Three million fingerlings weighing 3-5 g were released into the Anzali Lagoon (Iranian Fisheries Research Organization Newsletter, 49:4, 2006).). Poaching is rampant, even on the spawning grounds, and is also a significant factor in decline of this species (RaLonde and Walczak, 1972). Savenkova (1990) lists several reasons for reductions in catches in the Atrak River stock of Turkmenistan on the border with Iran. These include ineffective fish passes, poor marketing strategies, and the low quality of fish reaching the spawning grounds.

In the 1970s, rivers were rented to fishermen to exploit to an unlimited degree such that all spawning fish were taken and there was no recruitment for 4 years (Carl Bond Archives, Oregon State University, Corvallis).

The catch declined from 5854 t in 1918 to 172 t in 1937. Actual catches are about 20-30% larger because of local sale and consumption (Emadi, 1979). RaLonde and Walczak (1970b) note a decline from 1556 t in 1957 to 162.1 t in 1967 so the catch does not decline evenly. Emadi (1979) also pointed out that intensive sea fishing and fishing in rivers hindered spawning of the winter form, and led to a situation where only the spring form spawned. Leasing of rivers to fishermen was discontinued in an attempt to alleviate the decline in this species. The migration distance up rivers has decreased from about 25 km to about 8 km in recent years because of water abstraction and dams for agriculture (Bartley and Rana, 1998b). However, Yousefian and Mosavi (2008) report that artificial stocking has increased catches from 1000 t in 1981 to 10,000 t in 2002. Ghasemi et al. (2009) report a catch of 17,000t in 2008.

A further problem is the restocking programme only takes spring-run fish; the fall-run stock may no longer exist (but see below). In addition, stocks from various rivers are mixed and may result in outbreeding depression, the loss of adaptations to specific rivers in terms of migration patterns, spawning time, behaviour and other factors (Bartley and Rana, 1998b). Ghasemi et al. (2009) note that the spring run comprised over 98% of the catch and that the autumn run stock has declined from deterioration of spawning grounds, overfishing and other factors.

Measures to combat loss of this valued fish included a ban on fishing in the Anzali Mordab and its tributary rivers (catches here were up to 1000 t per year (Emadi, 1979)), effective control of illegal fishing and artificial spawning experiments (although the latter was insufficient to replace stocks). For several years starting in 1925, artificial breeding raised larvae for release in the ten most important rivers. In 1976-1977, fingerlings were raised to increase the stock in the Caspian Sea. Over 5 million fry were produced by the "Havyg" (? = Haviq) Hatchery alone in 1977. Mesh size of nets used to catch this species are apparently not harmful to younger fish as these have a chance to escape from the wings, with a wider mesh, before the bag net, with a smaller mesh, comes into play (Afraei Bandpei et al., 2010).

Emadi (1979) recorded releases of 28-44 million fingerlings and 250 million larvae "in recent years". A farm in the Siah Kal region of Gilan Province near Rasht was expected to produce 40 million white fish "roe", presumably fry, in 1985 (Kayhan International, 20 May 1984). The Dr. Beheshti Hatchery near Rasht expected to release more than 60 million fingerlings in the Iranian year 1993-1994 (Abzeeyan, Tehran, 4(5):VI, 1993). This hatchery had a peak production for the period 1973-1993 of over 140 million fingerlings in 1989, rising from the low period of 1973-1982 with less than 10 million fingerlings, and with subsequent decreases to about 50 million fingerlings in 1993 (Abzeeyan, Tehran, 5(3 & 4):IX-X, 1994). The number of fingerlings released from 1986 to 1991 climbed from 38 million to 170 million (Holčík and Oláh, 1992). Krasznai (1987) also refers to propagation of this species at "Sad-e Sangar" (Dr. Beheshti) and Siah Kal Fish Farms near Rasht in Gilan. Emadi (1993a) gives fingerling production at government hatcheries as follows: 25.3 million in 1983, 28.3 million in 1984, 38.0 million 1985, 51.7 million in 1986, 72.0 million in 1987, 84.3 million in 1988, 140.2 million in 1989, 156.3 million in 1990, 110.0 million in 1991, and 145.0 million in 1992. The Shahid Rajaee Hatchery in Sari released 358 million fingerlings, possibly in a single year (Abzeeyan, Tehran, 4(7):VII, 1993), although 70 million are reported as released annually to Mazandaran rivers in 1995 (Abzeeyan, Tehran, 6(8):III, 1995) and production of 70 million kutum fingerlings annually is reported in 2001 (Iranian Fisheries Research Organization Newsletter, 28:3, 2001). In 1997, 142 million fingerlings were produced for restocking (Bartley and Rana, 1998b). In 1999-2000, 150 million juveniles were released into the Caspian Sea (Iranian Fisheries Research Organization Newsletter, 23:4, 2000). From October to March 2000, 80 million juveniles raised in the Shahid Ansari aquaculture and breeding centre in Gilan were released into the Caspian Sea and neighbouring water bodies (Iranian Fisheries Research Organization Newsletter, 26:2, 2001). Billard and Cosson (2002) cite a mean of 100 million alevins released per year, reaching 140 million in some years, and give a brief overview of production facilities. Mosabbab and Piri (2005) record the release of fry into the Gorgan River of the southeast Caspian Sea as 16,663 million in 2000, 19,119 in 2001, 12,263 million in 2002, 11,931 in 2003 and 10,413 in 2004, the decrease being due to a fall in capture of brood fish. Amini (2006) reports release of fingerlings into the Larim, Goharbaran, Shirood, Tonekabon, Sardabrud, Mirud, Babol, Asbuchin and Sorkhrud rivers in 2000-2001 numbering 53.7 million and 62.5 million. Fork length was 36.7 mm and 43.3 mm and condition factor 1.13 and 1.2 0 respectively by year. In 2006, a report has more than 150 million juveniles being released into the Caspian Sea every year (www.iranfisheries.net, downloaded 28 July 2006). Sources obviously conflict on exact numbers, nevertheless marked variations in production are evident over short periods. The South Caspian Fisheries of Azerbaijan also released larvae in Soviet waters, more than 150 million in 1983 for example (Zarbalieva, 1987).

The Inland Water Aquaculture Research centre in Anzali has artificially propagated the autumn or fall-run stock (www.iranfisheries.net, downloaded 28 July 2006). Three million fish were released in July 2006, probably at 3-4 g with more to be released at a heavier weight. Paykhan Heyrati and Dorafshan (2007) compared the effectiveness of two kinds of dopamine antagonist combined with a gonadotropin releasing hormone analogue on ovulation and fertilisation success under hatchery conditions. Spawning induction techniques using hormones provide high quality gametes in quantity for aquaculture programmes.

Azari Takami et al. (1990), Woynarovich (1985) and Bartley and Rana (1998b) detail the technique for artificial spawning, incubation and raising of fingerling mahi safid. Adults are caught as they enter rivers on the spawning migration by blocking the river with wooden tripods interlaced with twigs as screens and using a meshed net. The Shaeed Ansari Fish Farm (of Shilat, the Iranian Fisheries Company) takes fish from 5 main rivers, forming a brood stock of 10,000 females and 20,000 males (Bartley and Rana, 1998b). Almost 100% of migrating fish are caught. Ripe fish are stripped and the eggs fertilised with sperm. Eggs from 2 females are pooled and mixed with milt from 2 males. The adhesive layer is washed away with water and continuous stirring. Once the eggs are completely separated from each other they are placed in incubator jars. Eggs may be placed in net-bottomed trays in the river for 2 days before being sent to a hatchery. Egg development takes a minimum of 7 days but larval development only takes 1-2 days. Larvae are fed with a mixture of milk and eggs until they are 5 days old and then put in the earthern rearing ponds. In the Shaeed Ansari Fish Farm, fingerlings are released in a river mouth at 1 g and most enter the Caspian Sea within 3 days (Bartley and Rana, 1998b). Fingerlings are grown in ponds to a weight of 2-3 g and then released in the Safid River which carries them down to the sea (Petr, 1987). Production of 1-2 g fingerlings attains 3 tonnes per hectare in fish farms (Emadi, 1993a). Farabi et al. (2007) studied brood stocks and fingerlings in the Shirod, Tonekabon, Tajan and Goharbaran rivers from March 2004 to March 2005. The mean length, weight and condition factor for broodstock females and males were 43.75 and 36.5 cm, 1189.5 and 678.13 g and 1.42 and 1.38 respectively. Eggs collected over a 62 day period weighed 4931 kg, mostly at the end of March and beginning of April. Percentage of survival of eggs in the four rivers listed above was 94.5, 95.1, 87.7 and 96.9 respectively. Newly hatched larvae averaged a total length of 6±2 mm and a weight of 2±0.2 g. The number of fish produced at less than 1g was 16,942,454 representing 19.9% of the total released in Mazandaran in 2004. Fish in the 1.0-1.5 g weight class, suitable for release, numbered 62,905,247. Parasites of fry were Diplostomum, Dactylogyrus, Butriocephalus, nematodes, Trichodina and Epistelis.

Studies on the best methods of rearing fingerlings and fry have been carried out in Iranian hatcheries and in rivers. The use of rotifers (Brachionis plicatilis) as food is more effective in larval growth than concentrated food preparations; after 40 days the rotifer-fed fish reached 440 mg and 31.36 mm on average and those fed concentrated food reached 17.2 mg and 14.75 mm (Fallahi et al., 2004; Kapourchali, 2006). Neverian et al. (2005) studied variation in protein levels fed to advanced fry at an isocaloric digestable energy of 3400 kcal/kg. The highest treatment at protein level 35% showed increased body protein, decreased fat and improved flesh quality in the carcass. Trichlorfon, an organophosphate pesticide is used to protect zooplankton such as rotiferans, food for larval safid mahi, against predators. However it does slow growth of the fish and the larvae must only be added to treated water 72 hours after the last application of the chemical (N. Chookbar, www.netiran.com, downloaded 18 April 2005; Chobkar et al., 2005; 2008). The sex steroid binding protein has been investigated as a biomarker for sexual behaviour in this species (Kousha et al., 2006). Spawning induction studies have been carried out on broodfish in Iran using carp pituitary extract and GnRH analogue, the latter alone or combined with metoclopramide. The latter treatment gave 90% ovulated females, 71.3% ovulation index and 68.4% fertilisation success, significantly higher than values for carp pituitary extract. However, the latency period was longer. Paykhan Heyrati et al. (2006) and Paykhan Heyrati and Dorafshan (2007) induced spawning with a gonadotropin releasing hormone analogue, both alone and in combination with the dopamine antagonists domperidone and metoclopramide, the combination with both antagonists being equally successful. Bahmani et al. (2007) found greater spermatozoan activity and quantity in fish that were captured further upriver on the spawning migration thus suggesting better sampling areas for broodstock; Ghara et al. (2007) found growth and survival of larvae in polyethylene tanks was better than fibreglass and glass aquaria; Nikou et al. (2007) studied levels of various hormones during transport of spawners and found clear indications of stress; Amiri et al. (2008) found that the optimum salinity for growth and survival of 1 g fingerlings was 8-10 p.p.t.; Keyvan et al. (2008) examined the effect of frozen storage time on the deterioration of  the product. Neverian et al. (2008) determined the optimum level of vitamin premix for the diet of advanced fry. Noverian et al. (2008) studied the effect of dietary digestible energy level on growth indices of advanced fry. Body protein and fat improved when the dietary energy was raised from 2500 to 2800 kcal kg-1. Haghighi et al. (2009) investigated the effects of dietary lipid levels on fry, growth, survival, protein and energy retention decreasing with an increase of dietary lipid level; a lipid level of 8% and 4314 kcal/kg bringing about the best growth performance. Akbarian et al. (2009) found that mid to high light intensities and long photoperiods could be useful in hatcheries where no particular light regime had been used. Kapoorchali et al. (2009) found increased growth of larvae when cow manure was used to increase abundance of zooplankton on a fish farm. Kousha et al. (2009) examined the fluctuation in sex steroid hormones during the spawning migration and annually, levels of some varying with variations in sea and river temperatures. Gholami (2010) found that Daphnia magna enriched with cod liver oil optimised nutrition of fry. Nikoo et al. (2010) studied physiological stress responses in fish caught on their upstream migration. Yasemi and Nikoo (2010) studied the impact of captivity on fertilisation and stress levels in broodstock and found short-term stress did not affect gamete quality levels.

A fine of 1,500 rials was imposed specifically for illegal angling of this species (Anonymous, 1977-1978).

Kiabi et al. (1999) consider this species to be of least concern in the south Caspian Sea basin according to IUCN criteria. Criteria include commercial fishing, sport fishing, abundant in numbers, habitat destruction, widespread range (75% of water bodies), absent in other water bodies in Iran, and present outside the Caspian Sea basin. The 2000 IUCN Red List lists this species as DD (Data Deficient). Coad (2000a), using 18 criteria, found this species to be one of the top 4 threatened species of freshwater fishes in Iran. Extinct in Turkey (Fricke et al., 2007).

Further work

The heavy demand for this fish as a traditional food warrants extensive management and conservation of stocks.

Sources

Iranian material: CMNFI 1970-0513, 9, 51.1-66.1 mm standard length, Gilan, Shafa River estuary (37º35'N, 49º09'E); CMNFI 1970-0516, 27, 42.3-61.0 mm standard length, Gilan, Lomir River (38º14'N, 48º52'30"); CMNFI 1970-0518, 9, 43.4-60.7 mm standard length, Gilan, Haviq River estuary (38º10'N, 48º54'E); CMNFI 1970-0520, 19, 46.7-73.9 mm standard length, Gilan, Astara River (ca. 38º25'N, ca. 48º52'E); CMNFI 1970-0563, 24, 40.8-58.1 mm standard length, Gilan Caspian Sea, Kazian Beach (ca. 37º29'N, ca. 49º29'E): CMNFI 1979-0088, 1, 104.3 mm standard length, Gilan, Safid River (no other locality data); CMNFI 1979-0471, 1, 106.0 mm standard length, Mazandaran, Caspian Sea west of Alamdeh (36º35'N, 51º48'E); CMNFI 1979-0686, 30, 42.8-52.2 mm standard length, Gilan, Safid River above ferry (37º24'N, 49º58'E); CMNFI 1980-0159, 1, 117.0 mm standard length, Gilan, Caspian Sea at Kazian Bridge (37º28'30"N, 49º28'E); CMNFI 1980-0160, 1, 88.5 mm standard length, Iran, Caspian Sea basin (no other locality data); CMNFI 1993-0140, 2, 64.5-71.5 mm standard length, Mazandaran, Tirom River, Ramsar (36º51'48"N, 50º48'E).

Comparative material: BM(NH) 1879.11.14:31-32, 2, 410.0-430.0 mm standard length, Russia, Astrakhan (ca. 46º24'N, ca. 48º05'E).

 Rutilus rutilus
(Linnaeus, 1758)

need to separate out caspicus information?

Common names

كلمه (= kolme, kollme, kolmeh or koolmeh, meaning unknown), mahi kolme, kolme Gorgan, telagi or talaji (in Gilaki and Mazandarani, meaning unknown, latter three used for R. r. caspius natio knipowitschi), mahi cheshm qermez (= redeye fish), kolme Kura or kolme Anzali for R. r. caspius natio kurensis, kolme Gorgan or kolme Turmenistan for R. r. caspius natio knipowitschi

[kulma, kyumen, xazar kulmasi and Kur kulmasi in Azerbaijan; kasli akcapagy in Turkmenian; Armyanskaya plotva or Armenian roach, karmraki and last for schelkovnikovi, all in Armenia; vobla, severo-kaspiiskaya vobla or North Caspian vobla, Astrakhanskaya vobla or Astrakhan vobla, Kurinskaya vobla or Kura vobla, Astrabadskaya vobla or Astrabad vobla, Turkmenskaya vobla or Turkmenian vobla, all in Russian; roach being the English equivalent of vobla].

Systematics

Cyprinus Rutilus was originally described from lakes in Europe, the type being from Sweden.

Holčík and Skořepa (1971) revised the roach and found no reason to maintain subspecies, of which they list 12. Rutilus rutilus caspicus (Yakovlev, 1870) (North Caspian or Astrakhan vobla) was the subspecies found in the Caspian Sea basin with natio knipowitschi Pravdin, 1927 (Astrabad or Turkmenian vobla) in Gorgan (= Astrabad) and Gasan-kuli bays and the Gorgan, Atrak and Qareh Su rivers and natio kurensis Berg, 1932 (Kura vobla) in Kyzylagach Bay, the Kura River, Astara and rarely the Anzali Mordab. The Kura vobla was said to differ from the Astrabad vobla by a greater body depth, smaller eye and more rapid growth. R. r. caspicus was distinguished from other subspecies by having modally 9 branched dorsal fin rays, darker fins, more inferior mouth, higher dorsal and anal fins, longer pectoral and pelvic fins, a deeper head, and larger eyes (see Berg (1948-1949) for more details). R. r. caspicus was originally described as Leuciscus rutilus var. caspicus from the Volga River delta, Russia. The types are unknown (Eschmeyer et al., 1996). Eschmeyer et al. (1996) also list Leuciscus rutilus var. wobla Grimm, 1896 described from the North Caspian Sea and mouths of rivers (Volga, Ural, Emba, Terek, Kura, Astara) entering the Caspian Sea. This is presumably a synonym of Rutilus rutilus judging from the subspecific name which is the Russian word for this species (wobla = vobla or roach). Kottelat (1997) lists it as a nomen nudum.

Holčík and Skořepa (1971) recognize the Caspian Sea populations as the morph or morpha migratorius, perhaps a first step towards differentiation and speciation. Mironovskii and Kas'yanov (1986; 1987) however retained Rutilus rutilus caspicus as a distinct subspecies in the Caspian Sea based on a multivariate analysis of 12 meristic and 7 morphometric characters. Mironovskii (1991; 1992) also demonstrated differences between Turkmen and Azerbaijan samples. Kuliyev (1984) considered these variations to be responses to different and changing conditions of the environment. Naddafi et al. (2002) also demonstrated differences in meristic (anal fin rays, predorsal scale number and total body vertebrae of 12 characters studied) and morphometric characters (7 measurements of 28 studied) for fish from the Anzali Mordab and the Gorgan River estuary of Iran. Keyvanshokooh and Kalbassi (2006) however found the value of Nei's genetic distance (d = 0.04) to be small between these two populations using DNA. The populations had similar levels of polymorphism. Keyvanshokooh et al. (2007) compared populations in the Anzali Wetland and Gorgan Bay using microsatellite markers and found differences between both populations were not significantly different for average number of alleles per locus nor for observed heterozygosities. Parafkandeh Haghighi and Rezvani (2005) and Parafkandeh Haghighi (2006) used the trace element content in otoliths to demonstrate the presence of two different populations in the southern Caspian Sea, the Anzali-Kura and the Gorgan-Turkmen, as noted above on meristic and morphometric values. Patimar et al. (2005) found morphological differences between fish from the Gomishan Wetland and those from the Adji-gol and Alma-gol wetlands in southeast Iran, and between the latter two wetlands. Inter- and intra-population variation was higher in the Gomishan fish. Rezvani et al. (2006) used mitochondrial DNA to compare Anzali and Turkmen roach and found the former to be more genetically variable, attributing this to there being more rivers, and therefore more spawning populations, in that area. Reyhani et al. (2010) used microsatellite markers and found higher heterozygosity in Anzali Wetland fish compared to those from Gorgan Bay. However, gene flow was high and population differentiation was non-significant.

Various body forms of this species have been given names as morpha, sometimes taken from species names regarded as synonyms, although as morpha these have no taxonomic significance, e.g. prasinus from Leuciscus prasinus Agassiz, 1835 (= elongata from Leuciscus rutilus var. elongata Fatio, 1882) for an elongate body and rutiloides from Leuciscus rutiloides Selys-Lonchamps, 1842 (= elata from Leuciscus rutilus var. elata Fatio, 1882) for a wide-backed body. A golden-orange or red aberration was known as aurata from Leuciscus rutilus var. aurata Fatio, 1882.

Bogutskaya and Naseka (2004) and Kottelat and Freyhof (2007) recognise Rutilus caspicus (Yakovlev, 1870) as the semi-anadromous species in the Caspian Sea with R. rutilus in rivers and lakes. Curiously, R. caspicus and R. frisii showed low genetic divergence in Ketmaier et al. (2008), a study using DNA but sample sizes were low.

Three "types" of Rutilus rutilus caspicus have been reported from Iran (Annual Report, 1995-1996, Iranian Fisheries Research and Training Organization, Tehran, p. 54-55, 1997). One type lives in the Anzali Mordab and is relatively small, the second is larger and migrates between Anzali and the Kura River of Azerbaijan and the largest migrates between the northern and southern coasts of the Caspian Sea.

Rutilus rutilus schelkovnikovi Derzhavin, 1926 was described from the Karasu which falls into the Aras River 5 km above the mouth of the Zanga River (Razdan River), Echmiadzin District, Armenia, and Rutilus rutilus uzboicus Berg, 1932 from lakes in the Uzboi River Valley (with the type from Lake Yaskhan) in Turkmenistan north of the Iranian border. Rutilus rutilus var. fluviatilis (Jakovlev, 1870) described from the delta of the Volga River is another synonym.

Hybrids with Blicca bjoerkna are reported from the Aras River basin in Armenia, which is shared with Iran. Artificial hybrids with Rutilus frisii kutum and Abramis brama have been bred in Iran (Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 39-40, 1996).

Key characters

This species is distinguished from the related Rutilus frisii by the lower scale count (< 50) and the posterior part of the swimbladder being rounded rather than pointed.

Morphology

Dwarf and large forms are recognised in Dagestan by Shikhshabekov (1969) with different life histories and distinct spawning stocks are known from the north Caspian Sea.

Lateral line scales 39-48, mostly 42-47. Scales have fine circuli around a central focus. Radii may be found on all fields but fish often lack radii on lateral fields. Usually there are very few anterior and posterior radii (e.g. in a fish 150.7 mm standard length there are 2 anterior and 4 posterior radii; but counts are quite variable, at least 7-14 radii total). Scales have a wavy anterior margin and a finely crenulated posterior margin. There is a pelvic axillary scale and a scaled keel behind the pelvic fins. Dorsal fin branched rays 8-12, usually 9 in the Caspian Sea (but see below), after 2-5 unbranched rays, anal rays 8-12, mostly 9-10, after 2-4 unbranched rays, pectoral fin with 14-18 branched rays, and pelvic fin with 7-9 branched rays. Gill rakers 9-17, usually 10-14, short and stubby and reaching the adjacent raker when appressed. Vertebrae 38-43. Lake populations generally have more vertebrae than river populations and the number of vertebrae is genetically fixed, generally following the maternal genotype (Izyumov and Kas'yanov, 1995). Pharyngeal teeth 6-5 or 5-5, rarely 6-6, 4-5, 5-4, or 5-6 with the grinding surface slightly folded to form a long, hollow crown for the largest teeth. There is a rounded and hooked tip to teeth 1-4. The tooth margin in the posteriormost 2 teeth is serrated. The anteriormost 2 teeth are rounded. Teeth are more serrated and hooked in smaller fish. The gut is s-shaped with a small anterior loop. The chromosome number is 2n=50 (Klinkhardt et al., 1995).

Meristics for Iranian specimens:- dorsal fin branched rays 9(22) or 10(18); anal fin branched rays 9(16), 10(23) or 11(1); pectoral fin branched rays 14(1), 15(4), 16(22), 17(12) or 18(1); pelvic fin branched rays 7(4), 8(33) or 9(3); lateral line scales 40(1), 41(4), 42(16), 43(11), 44(5) or 45(3); total gill rakers 11(4), 12(15), 13(18) or 14(3); pharyngeal teeth 6-5(22), 6-6(1), 6-4(1) or 5-5(1); and total vertebrae 37(1), 39(2), 40(9), 41(64) or 42(19).

Sexual dimorphism

In a male caught on 26 April, moderate-sized tubercles are evenly distributed over the top and sides of the head, with fewer tubercles on the undersurface. Scales bear one strong tubercle in the exposed mid-scale, anterior and dorsal scales have smaller tubercles on the scale margin and belly scales have minute tubercles on the margin and exposed scale base. Fine tubercles follow the branching fin rays in single file on all fins, both dorsally and ventrally on the pectoral and pelvic fins, and in 1-2 rows on the last unbranched ray of each fin. Some fish have some scales with 1-3 tubercles, usually 2, one above the other. When there are 3 tubercles these are variably arranged, e.g. in an L-shape or a reversed L. Females also bear tubercles on the head and scales but these are less well-developed and are not present on fins.

Colour

Overall colour is silvery. The back is grey and the sides of the head golden. The belly is pearly-white. The iris is silvery with a black spot above the pupil, some gold on the upper part, and may be orange to red. Pectoral, pelvic and anal fins light grey with a darker band on the edge. The middle of the pelvic and anal fins may be transparent, orange to pink or red. The peritoneum is silvery with moderately dispersed but evident melanophores.

Size

Attains 56.0 cm and 3.13 kg. Larger fish may be hybrids with Rutilus frisii. In Iran during the 1950s, catches were 16-35 cm long (Farid-Pak, no date).

Distribution

Found from the British Isles and France eastwards to Siberia, north of the Alps; in the Black Sea basin but not in Turkey, except the northwestern corner and the Yesilirmak River basin, and on all shores of the Caspian Sea basin including that of Iran, its rivers and bays (Nedoshivin and Il'in, 1929; Berg, 1948-1949; Kozhin, 1957; Riazi, 1996; Abbasi et al., 1999; Kiabi et al., 1999).

Abdoli (2000) and Abdoli and Naderi (2009) have natio knipowitschi in Gorgan Bay and the neighbouring Caspian coast and in the Gorgan, Gharasu and Atrak rivers, and natio kurensis in the lower Safid River, Anzali Talab and neighbouring Caspian coast to Astara and the middle Aras River.

Rutilus rutilus aralensis Berg, 1916 (but see above on subspecies status) is reported from the Karakum Canal and Kopetdag Reservoir in Turkmenistan (Shakirova nd Sukhanova (1994; Sal'nikov, 1995) and may eventually be found in the Tedzhen (= Hari) River basin of Iran.

Zoogeography

A European and western Asian species with its origins in a Danubian or Sarmatian fauna.

Habitat

This species is eurytopic, living in rivers, streams, lakes, reservoirs, and fresher parts of seas. It occurs in schools close to vegetation in fresh waters.

This species gathers in front of the Volga River delta in the north of the Caspian Sea in large shoals in autumn when water levels decrease. At water temperatures of 5°C they descend to the sea floor and "lie in winter sleep" (Holčík and Skořepa, 1971). It is not known if this occurs off the warmer Iranian shore. Generally the vobla lives in the sea itself and migrates to rivers for spawning. Although most spawning takes place in fresh water, some occurs at a salinity of 6.5‰. This species is the most tolerant of the semi-anadromous fishes, the lethal salinity level being 15-16‰. In the north Caspian Sea, this fish are found mostly shallower than 7.5 m but Knipovich (1921) reports this species at 36.6-53.0 m in the Iranian Caspian Sea.

A mass migration of the Kura vobla (natio kurensis) into the Anzali Mordab occurred between 21 and 28 February 1915, only to disappear after 4 April. Riazi (1996) reports that this species migrates into the Siah-Keshim Protected Region of the Anzali Mordab. Males migrate into the Anzali and Gomishan wetlands of Iran earlier than females and the larger fish enter first (Naddafi et al., 2005).

Natio knipowitschi lives in the sea in Gorgan and Gasan-kuli bays and enters rivers there. Savenkova (1994) reported as many as 2000 million juveniles descending the Atrak River in May and June in high productivity years and as few as 83 million in low productivity years. The sea in July and August is as warm as 28-30°C off the Iran-Turkmenistan border and the juvenile fish from the Atrak descend to 11-12 m. Savenkova (1994) gives further details of movements in relation to temperature, e.g. when waters cool to 8-12°C off Turkmenistan, young fish move to warmer Iranian waters to overwinter. ERM-Lahmeyer International GmbH, DHI Water & Environment and GOPA Consultants (2001a) report that the Gomishan Lagoon at 37º11N, 53º57'E is an important staging area for this species.

Age and growth

Generally growth is a function of type of feeding (a diet favouring molluscs gives higher growth than herbivory) and temperature. High growth rate is shown by semi-migratory fish but is also shared by some populations resident in lakes and rivers. Kas'yanov et al. ( 1995) review the complex relationships of environmental factors affecting growth in this species.

A life span up to 19 years has been reported although most fish are 10 years or less. Growth is fastest in the second and third years of life when most fish mature sexually (Strubalina and Chernyavskiy, 1992).

Some fish may mature at one year of age and a length of 9-11 cm in the Atrak River population (Petr, 1987). Growth in the Atrak population juveniles is heavily dependent on temperature regimes in the sea, the timing of descent to it, and stock abundance. Fish which descend earlier because of low flow and have a longer feeding season, find a favourable, warm regime and few competitors will have faster growth (Savenkova, 1994).

Young-of-the-year Turkmen fish reach 4.9-5.8 cm in July while in the northern Caspian they only attain 3.3-4.1 cm, although the latter have a higher condition factor, possibly because of higher benthic biomass and heterogeneity (Savenkova, 1994). The dwarf form in Dagestan is 11 cm and 31 g at 3 years of age while the large form is 20.3 cm and 130 g. Both are sexually mature (Shikhshabekov, 1969).

The commercial catch in Iran during 1971/72 was 3-6 years old, 20.4-26.0 cm long and weighed 146-300 g (Razivi et al., 1972). The average weight in 1995 was 200-300 g (Abzeeyan, Tehran, 5(9):V, 1995).

In Gorgan Bay, the age composition is from 1+ to 6+ with age group 3+ and 4+ each accounting for about 30%. Growth rate for all south Caspian populations is higher than the north Caspian populations, two-year-olds being the same length as three-year-olds. Average weight is 247 g with larger fish at 30 cm weighing 583 g. Numerous fish entered Gorgan Bay in mid-January in the early years of this century and by the end of that month ascended all the tributary rivers, in particular the Qareh Su (= Karasu).

Growth rate in the Anzali Mordab was higher than in the Gomishan Wetland of the southeastern Caspian Sea, limited by higher salinity in the latter (Naddafi et al., 2002a, 2002b, 2005; Paghe et al., 2005). The population density was higher in Gomishan as Anzali suffers more from pollution, overfishing, illegal fishing and other human impacts. Mean condition factors for the Gomishan population were 1.84 for males and 2.09 for females while in the Anzali population these were 2.03 and 2.2 respectively. The most abundant age groups in the commercial fishery of both areas were 3+, 4+ and 5+. Maximum age at Gomishan was 8+ years. Growth was rapid in the first year with a sharp decline in the second year and a steady, less rapid decline in subsequent years. Females were longer than males at in each age class. Sex ratios were did not differ significantly from 1:1. The Anzali population had longer body lengths than fish from other studies, perhaps due to a longer growing season and higher water temperatures in the south Caspian Sea, better food supply and genetic factors.

Food

The young feed first on phytoplankton and then switch to zooplankton and benthic insect larvae with growth. Adults eat benthic animals and plants with molluscs an important diet item where present as in the southern Caspian Sea. A mollusc diet occasions significant wear on the pharyngeal teeth (Holčík and Skořepa, 1971). There has been a long-term variation in the diet of roach in the Caspian Sea, molluscs declining and being replaced by crustaceans and plants (Strubalina and Chernyavskiy, 1992). Plant foods in Europe include water thyme, milfoil, duckweed, stonewart and algae scraped from reeds and rocks (Muus and Dahlstrøm, 1999). There is little feeding during spawning in rivers. Young-of-the-year in the Atrak delta feed on the polychaete worm Nereis (up to 89.5%) and with growth the mollusc Cerastoderma lamarckii (up to 70.8%), and to a lesser extent the exotic mollusc Abra ovata (5.2%) since this abundant species is partially buried and is not readily available. Polychaetes and molluscs have high biomasses in the silty sediments of around 30-50 g/sq m. Sometimes protozoa and crustaceans figure prominently in the summer diet but diet varies considerably between months and years (Savenkova, 1994).

Reproduction

A spawning migration runs up rivers in the Caspian Sea basin in spring, the larger fish first. Spawning grounds are mostly flooded meadows that warm up easily (Caspian Sea Biodiversity Database, www.caspianenvironment.org). Immature fish only migrate in autumn after their gonads have matured, spending the summer in deeper water. The fish return to the sea after spawning in the same sequence as the upriver migration. Young remain in the river until early summer when they too go down to the sea. However some populations of this species are exclusively freshwater and do not migrate.

Spawning is continuous and takes about 5-6 hours. Fecundity reaches about 202,000 eggs and egg diameters 1.6 mm. In the Volga delta this species spawns in the same areas as Abramis brama. However, it spawns earlier at temperatures of 7-16°C on last years dead vegetation while the bream spawns later at 15-23°C on new growth. Spawning can occur on stony bottoms in the absence of vegetation. Spawning is a noisy process which can be heard from some distance. Each female is accompanied by 2-3 males who push on her belly, sometimes lifting her part way out of the water, as they swim in circles. With many fish engaged in this process, the water foams. The fish gradually approach the shore where the eggs are shed in water only 5-20 cm deep as the female is squeezed by two males. Other males take part in the process, as many as 10 at a time. Incubation takes 4 days at 17-20°C. Young actively move downstream once they reach 20-25 mm.

The breeding migration in the Atrak River starts in January-February and continues until April, the fish traveling 70-80 km upriver (Petr, 1987). Nümann (1969) states that the spawning migration takes place from November to December in Iranian waters. The migration in the Atrak peaks at 10-12°C and lasts 20-25 days. Spawning peaks at 15-18°C in March on the lower Atrak floodplains (in 1908 spawning in the lower Atrak was observed in mid-April). In the absence of flooding, no spawning occurs. After spawning, females migrate back to the sea first. In the Qareh Su of Gorgan Bay, spawning took place 20-25 km upriver from the mouth in 1907 and 1908. It started in mid-February, peaked in the second half of February and the first half of March, and ended between 20 and 30 March. Males with running milt first appeared between 14 and 20 February in 1915 and ripe females a week later. Mass spawning took place at the beginning of March (Nedoshivin and Il'in, 1929). The main spawning rivers in Iran are the Qareh Su, Goharbaran, Larim, Siah-Darvishan and Shafa River (Iranian Fisheries Research and Training Organization Newsletter, 9:5, 1995).

Fecundity in the Gomishan Wetland (4262-98,804 eggs) was significantly higher than in the Anzali Mordab (6035-32,141) for a given body size. Egg diameters ranged from 0.9 to 1.45 mm and were not significantly different between the two wetlands. The peaks for the gonadosomatic index (GSI) curves were early March for males and mid-March to early April for females. The GSI was higher in Gomishan (Naddafi et al., 2005).

Parasites and predators

The main predator of this species in the north Caspian Sea is Sander lucioperca, accounting for 65% of its food. Silurus glanis and Esox lucius and various birds such as the pelican are also important predators (Kushnarenko, 1978). The Caspian seal, Pusa caspica, is a significant predator on this species (Krylov, 1984). On the spawning grounds, this fish can be picked up by hand and falls easy prey to birds and other predators.

Mokhayer (1976b) records the cestode larva Diagramma. Jalali and Molnár (1990a) record the monogenean Dactylogyrus turaliensis from this species in the Safid River. Masoumian et al. (2002) recorded parasites from fish captured form the coast of the southeast Caspian Sea as Anisakis larvae, Aspidogaster limacoides, Bothriocephalus gowkongensis, Dactylogyrus turaliensis and Diplostomum spathaceum. Youssefi et al. (2005) record the pleroceroid larvae of Ligula intestinalis in fish from the Aras Dam. A toxin produced by this parasite can cause infertility and weight loss in the fish and can be harmful to humans. Khara et al. (2006a) record the eye fluke Diplostomum spathaceum for this fish in the Amirkalayeh Wetland in Gilan, having the highest infection rate in 7 species examined. Khara et al. (2006b) record the cestode Caryophyllaeus fimbriceps from this species in the Boojagh Wetland of the Caspian coast. Khara et al. (2008) found the eye parasite Diplostomum spathaceum in this fish from Boojagh Kiashar Wetland in Gilan. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Lernaea sp. on this species.

Economic importance

This species is one of the most important commercial species in the Caspian Sea with a catch in the north Caspian and lower reaches of rivers in 1930 being 2,591,000 centners (15,000,000,000 fish) or 45.5% of the total catch of fishes in the Caspian Sea or nearly 20% of the whole Soviet fish catch. Archaeological studies on the eastern Caspian shore in the former Soviet Union has shown this species to have been fished for 5000 years ago (Tsepkin, 1986). It is a significant food for the sturgeon Huso huso (Keyvanshokooh and Kalbassi, 2006).

The main fishing season in Iran is February and March (Farid-Pak, no date). Nevraev (1929) gives catches for various fishing regions in Iran in the early twentieth century. For the Anzali region from 1914-1915 to 1917-1918 the catch was 6000 to 244,800 individuals (the catch declining steadily over this period), and in Astrabad (= Gorgan) region from 1909-1910 to 1912-1913 the catch was 8,348,800 to 21,790,000 individuals. The catch in Iran from 1956/1957 to 1961/1962 varied from 2989 kg to 1,092,719 kg (Vladykov, 1964), from 1965/66 to 1968/69 it varied from 32 to 74 tonnes (36, 74, 32 and 35 t respectively) (Andersskog, 1970) and from 1963 to 1967 from 2.4 to 47.0 t (23.1, 2.4, 3.7, 30.7, 47.0 t respectively) (RaLonde and Walczak, 1970b). Vladykov (1964) reports catches from the Anzali region for the period 1933/1934 to 1961/1962 to vary from 57 to 716,974 kg, with none reported in some years. Holčík and Oláh (1992) report an annual catch in the Anzali Mordab from 1932-1964 of 0.8-449.7 t. The total catch in 1989 (or 1990) was only 26 t but increased to 120 t in 1995 because of the rising Caspian Sea level which provided more spawning grounds (Abzeeyan, Tehran, 5(9):V, 1995; Iranian Fisheries Research and Training Organization Newsletter, 9:5, 1995). Conflicting values from reports where they overlap are typical of fisheries statistics from Iran which can only be taken as general trends rather than absolute values (see below also). The Food and Agriculture Organization, Rome gave the catch for Rutilus spp., presumably this species, for the years 1980-1985 as 0, 0, 0, 121, 347, and 350 t respectively. The catch in Gorgan Bay 20-30 years ago according to Petr (1987) was about 4000 tonnes per year but is now negligible. This figure conflicts with the ones cited above. In the 1914-1915 fishing season in Gorgan (= Astrabad) Bay, 24.2 million fish were taken, of which 21.9 million were taken in the Qareh Su. Petr (1987) cites catches of 830 t in 1978 for the southeastern Caspian Sea and for 1979 only 120 t, probably including Soviet catches. Recruitment falls when the Atrak River, a major spawning ground, fails to flood adequately. The dwarf form in Dagestan is of no commercial value and may have arisen with changing environmental conditions (Shikhshabekov, 1969).

Moeini and Daneshnuran (20010 and Moini et al. (2005) have investigated the production of marinades using this fish using various recipes, examining their chemical composition over time and their comparative tastes.

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in aquaculture and as food, in sport and in textbooks and because it has been introduced outside its natural range. Golpour and Imanpour (2010) consider this species to be threatened in Iran from overfishing and deterioration of spawning grounds. Everard (2006) is one book of many describing the biology and importance of the roach as Rutilus rutilus.

Conservation

Holčík and Oláh (1992) report a catch of only 5 kg in the Anzali Mordab for 1990. This species has been cultured at "Sad-e Sangar" Fish Farm near Rasht (3 million larvae a year), at Astara and at Anzali on the Caspian among other localities (Mokhayer, 1972b). In 1999-2000, 30 million juveniles were released into the Caspian Sea (Iranian Fisheries Research Organization Newsletter, 23:4, 2000). Noroozi et al. (2006) detail capture of brood stock from the Gorgan River estuary for release into 2 ha earthen ponds enriched with manure and fertiliser. The optimum temperature for spawning brood stocks is 12-17ºC, found in mid-March to late April. Pine branches were placed in the ponds as spawning sites. Ponds were stocked with 700 females averaging 150 g and 350 males averaging 100 g. Eggs hatched on the sixth day. Larvae were fed on natural zooplankton (Rotatoria and Daphnia) and artificial food.

The Atrak River population can be conserved by regulating the discharge so that the spawning grounds flood at the appropriate time (Petr, 1987). Beach seines with mesh sizes of 28 mm, 33 mm and 36 mm caught fish of ages 2.8 years, 2.9 years and 3.3 years respectively. This data indicates that 58.3%, 50.5% and 3.6% of the total catch is non-standard respectively. Seines with a cod end of at least 36 mm are recommended for stock protection (Iranian Fisheries Research and Training Organization Newsletter, 6:8, 1994). Catches are increasing with the rise of the Caspian Sea water level as noted above, indicating how natural events beyond human control can have a significant effect on stocks.

Imanpour et al. (2009) studied the relationship between fish size and various spermatological parameters. Sperm volume did not increase with body length although gonadal weight did. Spermatocrit, sperm motility, sperm density, gonadosomatic index and hematocrit were not influenced by fish body size but sperm density decreased significantly with increasing gonadal weight. Golpour and Imanpour (2010) studied relationships between seminal and blood plasma composition during the reproductive season, as part of improving artificial fertilisation.

There have been few studies on the effects of environmental factors on this species in Iran. Gerve'ei et al. (2008) determined the LC50 and gill tissue lesions caused by aluminium sulphate under laboratory conditions and Keramati et al. (2010) studied the effect of the agricultural organophosphate diazinon on enzyme activity, also under laboratory conditions.

Further work

As an important food fish, this species should be effectively managed and conserved.

Sources

Wossugh-Zamani (1991d) and Akbari-Pasand (1996) give accounts of this species in Farsi.

Iranian material: CMNFI 1970-0509, 3, 73.4-82.9 mm standard length, Gilan, Safid River at Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1970-0510, 4, 58.6-83.8 mm standard length, Gilan, Golshan River (37º26'N, 49º40'E); CMNFI 1970-0531, 14, 65.5-115.8 mm standard length, Mazandaran, Larim River (36º46'N, 52º56'E); CMNFI 1970-0532, 1, 74.9 mm standard length, Gilan, Caspian Sea near Bandar-e Anzali (37º28'N, 49º27'E); CMNFI 1970-0543A, 1, 119.4 mm standard length, Gilan, Caspian Sea at Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1970-0549, 1, 109.7 mm standard length, Mazandaran, Qareh Su near Alm Imamzadeh (no other locality data); CMNFI 1970-0563, 5, 40.3-112.2 mm standard length, Gilan Caspian Sea, Kazian Beach (ca. 37º29'N, ca. 49º29'E): CMNFI 1970-0583, 6, 35.7-84.9 mm standard length, Gilan, Nahang Roga River (37º28'N, 49º28'E); CMNFI 1979-0431, 1, 157.0 mm standard length, Mazandaran, fish bazaar at Now Shahr (no other locality data); CMNFI 1979-0788, 2, 64.9-66.6 mm standard length, Mazandaran, Gorgan River at Khadje Nafas (37º00'N, 54º07'E); CMNFI 1980-0126, 4, 162.2-178.3 mm standard length, Gilan, Caspian Sea near Bandar-e Anzali (37º28'N, 49º27'E); CMNFI 1980-0136, 4, 72.1-109.7 mm standard length, Mazandaran, Fereydun Kenar River estuary (36º41'N, 52º29'E); CMNFI 1980-0148, 1, 85.4 mm standard length, Gilan, Pir Bazar Roga River (37º21'N, 49º33'E); CMNFI 1980-0157, 2, 73.1-132.0 mm standard length, Mazandaran, Gorgan River estuary (36º59'N, 53º59'30"E); CMNFI 1980-0905, 9, 98.5-203.0 mm standard length, Mazandaran, Gorgan River at Khadje Nafas (37º00'N, 54º07'E).

Genus Scardinius
Bonaparte, 1837

Howes (1981) placed this genus in Rutilus Rafinesque, 1820 on osteological grounds. Bogutskaya (1988) disagrees. Scardinius is usually separated from Rutilus by having 2, as opposed to 1, rows of pharyngeal teeth and a ventral keel on the body. Howes (1981) considers pharyngeal teeth to be of only species importance and the keel is variously developed in Rutilus.

The genus contains perhaps 6 species and is found from the British Isles and the Iberian Peninsula throughout Europe to the Caspian and Aral Sea basins, with one species in Iran.

The genus is characterised by a pharyngeal tooth count of 3,5-5,3 with major row crowns laterally compressed and bearing 5-8 serrations, scales moderate in size in a complete lateral line, few, short gill rakers, a keel on the belly behind the pelvic fins covered with scales, a short gut and light peritoneum, dorsal and anal fins of moderate length, dorsal fin origin well behind the origin of the pelvic fins, terminal and oblique mouth,

Scardinius erythrophthalmus
(Linnaeus, 1758)

Scardinius erythrophthalmus from Wikimedia Commons.

Scardinius erythrophthalmus from Wikimedia Commons.

Scardinius erythrophthalmus from Wikimedia Commons.

Common names

sorkh pareh or sorkh par (= red fin), mahi sorkh baleh, ماهي چشم قرمز (= mahi chesm ghermez).

[giziluzkac in Azerbaijan; krasnoperka in Russian; rudd, redeye, redfin, pearl roach].

Systematics

See above under the genus. Cyprinus Erythrophthalmus was originally described from northern Europe. This species is widely known to spawn with other cyprinid fishes making hybrids a common occurrence. Some Iranian material appears to be hybrids of this species and another, unknown parental species but this has not been investigated.

Key characters

This species is often confused with Rutilus rutilus and R. caspicus but can be distinguished by the posterior position of the dorsal fin (in relation to the pelvic fins), the belly keel, the upturned mouth, and the serrated pharyngeal teeth in 2 rows.

Morphology

Dorsal fin with 2-4 unbranched and 7-10, usually 8 branched rays, anal fin with 3-4 unbranched followed by 9-13, usually 11, branched rays (Abdurakhmanov (1962) initially gives 8-9 anal fin branched rays for Azerbaijan fish but this may be a misprint as a subsequent table lists 9-11 rays), pectoral fin with 13-16 branched rays, and pelvic fin with 7-9 branched rays. Lateral line scales 36-45. There is a pelvic axillary scale. Scales are squarish in shape, with sharp dorsal and ventral anterior corners, a wavy anterior margin, central focus, fine circuli which are coarser on the posterior field, and very few anterior and posterior radii (e.g. 2 anterior and 3 posterior primary radii reaching the focus from the margin). The scale margin is indented where radii terminate and the thick posterior radii are visible on the flank. Gill rakers short and widely spaced, touching the adjacent one when appressed, and numbering 8-13. Vertebrae 37-42. Pharyngeal teeth mostly 3,5-5,3 with variants 3,5-5,2, 3,5-5,1, 3,5-4,3, 2,5-5,3 and 2,5-5,2, narrow and elongate, slightly hooked and with about 5-8 strong serrations on each tooth. There is a strongly-developed, scaled keel between the vent and the pelvic fin base. The gut is s-shaped with an anterior loop. The chromosome number is 2n=48-50 (Klinkhardt et al., 1995).

A single Iranian specimen had the following meristics:- dorsal fin branched rays 8; anal fin branched rays 10; pectoral fin branched rays 16; pelvic fin branched rays 8; lateral line scales 37; total gill rakers 10; pharyngeal teeth 3,5-5,3, and total vertebrae 39..

Sexual dimorphism

Males develop breeding tubercles on the head and body.

Colour

The back is blue-black to greenish- or olive-brown, the flanks are brassy and the belly silvery-white. Upper flank scales have dark bases. The tips of the caudal, anal and pelvic fins are a bright, blood red in the spawning season and the dorsal fin is black proximally and red distally. The iris is yellow to orange, or gold, with a red spot at the top. Peritoneum silvery with scattered melanophores. Young are much less brightly coloured than adults.

Size

Attains 62.0 cm and 3.01 kg.

Distribution

Found from the British Isles and north of the Pyrenees east to the Caspian and Aral sea basins. It is recorded from the Lenkoran in Azerbaijan and, in Iran, from the Aras River, Anzali Mordab, Bojaagh wetland, Safid River, Haraz River and Babol River (Derzhavin, 1934; Holčík and Oláh, 1992; Abbasi et al., 1999; Abdoli, 2000; Jolodar and Abdoli, 2004; K. Abbasi, pers. comm., 21 February 2005; Patimar et al., 2010).

Zoogeography

This species is part of a European and West Asian fauna whose origins may lie in a Danubian or Sarmatian fauna.

Habitat

Rudd can favour heavily overgrown areas (Shikhshabekov, 1979) and are generally found in shallow warm lakes or slow moving rivers. They are usually inhabitants of midwater or near the surface but they overwinter in deep water. They are regarded as fairly hardy and are adapted to eutrophic, and presumably therefore, polluted waters. In Iranian waters its density is highest in the Anzali Mordab (K. Abbasi, pers. comm., 21 February 2005) and the population there is stunted through poor habitat quality (Patimar et al., 2010).

Age and growth

Sexual maturity is attained at 3-4 years in Dagestan at lengths of 17-29 cm and 80-530 g. A stunted form is found in rice paddies at an age of 2 years, 7.5-11.0 cm and 10-23 g (Shikhshabekov, 1979). Elsewhere life span is at least 17 years.

Patimar et al. (2010) studied a stunted population in the Anzali Lagoon and found maximum total length was 146 mm and maximum weight was 46.93 g. Males reached 4+ years and females 5+ years with 92.5 % of fish age 2+ to 3+ years. Females were longer and heavier than males. Growth was isometric. There was no significant difference in condition coefficients between sexes and no temporal differences when considered separately for each date of sampling. The condition coefficient was highest in early May for males and mid-May for females and was lowest in early June for both sexes. Females dominated in older age classes and males in the younger.

Food

Food is aquatic macrophytes as well as insect larvae, crustaceans, molluscs and more rarely fish eggs and fry. The young feed on zooplankton.

Reproduction

Spawning takes place at water temperatures of at least 18-20°C in June-July in Dagestan. Each female can be accompanied by two males, one on each side. Two batches of eggs may be spawned in this period (Shikhshabekov, 1979). In the Volga Delta, spawning takes place from April until the end of June. Eggs attach to water plants. The young remain attached to vegetation until the yolk-sac is absorbed. Fecundity is up to 232,000 eggs with a diameter of 1.5 mm. Hatching takes 3 days at 20-22°C.

The Iranian study of Patimar et al. (2010) found a reproductive period of mid-April to late May and showed that the gonadosomatric index peaked in mid-May for both sexes and then decreased sharply. Egg diameters reached 1.23 mm, maximum fecundity reached 59,620 eggs and relative fecundity 1737.69 eggs/g.

Parasites and predators

Masoumian and Pazooki (1998) surveyed myxosporeans in this species in Gilan and Mazandaran provinces, finding Myxobolus pfeifferi. Sattari et al. (2004) record the nematode Raphidascaroides acus larvae from this species in Gilan. Khara et al. (2006b) record the nematode Raphidascaris acus from this species in the Boojagh Wetland of the Caspian coast. The Caspian seal, Pusa caspica, is a predator on this species (Krylov, 1984).

Economic importance

Holčík and Oláh (1992) report a catch of 98 kg in the Anzali Mordab in 1990.

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in aquaria and aquaculture, as food, in sport and in textbooks and because it has been introduced outside its natural range.

Conservation

Lelek (1987) classifies this species as vulnerable in Europe. Kiabi et al. (1999) consider this species to be conservation dependent in the south Caspian Sea basin according to IUCN criteria. Criteria include sport fishing, medium numbers, habitat destruction, limited range (less than 25% of water bodies), absent in other water bodies in Iran, present outside the Caspian Sea basin. Near threatened in Turkey (Fricke et al., 2007).

Further work

The distribution and biology of this species and its potential for hybridisation with other cyprinids requires study in Iran.

Sources

Iranian material: Uncatalogued, 1, 103.7 mm standard length, Gilan, swamp near Hendeh Khaleh (37°23'N, 49°28'E).

Genus Schizocypris
Regan, 1914

This genus of medium-sized snow trouts contains only 2 species found in Pakistan, Afghanistan and Iran. Coad and Keyzer-de Ville (2005) revised the genus.

It is characterised by a rounded and moderately elongate body, scales small but larger near the shoulder region, belly scaleless, a wide and transverse mouth with the snout projecting, no barbels or barbels vestigial, pharyngeal teeth with a flat tip unlike Schizothorax and a formula of 2,3,4-4,3,2 rather than 2,3,5-5,3,2 as in Schizothorax, dorsal and anal fins short but 6 branched anal fin rays not 5 as in related genera in the same area, dorsal fin with a strongly serrated spine, scales in the vent region are split and enlarged to flank the urogenital region, and radii are on all scale fields.

Rainboth (1981) includes Capoeta trutta, Capoeta fusca and Capoeta nudiventris (= C. fusca) in this genus but this is incorrect. These species show some enlargement of scales around the anus and anal fin region but it is not as marked and definitive as in true Schizocypris and other characters of the genus are absent.

Schizocypris altidorsalis
Bianco and Banarescu, 1982

Common names

gorgak (= small wolf), anjak or khaju (A. A. Pasand, pers. comm., 5 November 2000 but see under Schizothorax zarudnyi and S. pelzami).

Systematics

Schizocypris brucei, non Regan, 1914 (Annandale and Hora, 1920) is a synonym. Schizocypris brucei Regan, 1914 is thus restricted to the Gomal River drainage in the Indus River basin of Pakistan (originally described from the Wana Toi, a Gomal River tributary). A third species in the genus Schizocypris is S. ladigesi Karaman, 1969 from the Kankai River, also in the Indus River basin.

Kähsbauer (1964) reports a hybrid between Schizothorax schumacheri and Schizopygopsis stoliczkae from Sistan which may in fact be this species.

The holotype of Schizocypris altidorsalis is in the Istituto di Zoologia dell'Universitá di L'Aquila, Italy under IZA 8169 and is 73.7 mm standard length (my measurment) (Bianco and Banarescu, 1982). The type locality is "Nahr-Taheri near Zabol, Seistan". Paratypes include 5 fish from the type locality under IZA 7841, 35-65 mm standard length (4 fish seen by me, 35.4-62.0 mm standard length) with further specimens in the Institutul de Stiinte Biologice, Bucuresti, Romania (ISBB 3136). Three paratypes, 68-73 mm standard length, from "Rud-Sistan, 8 km from Zabol, Seistan" are under IZA 7844 (69.9-74.3 mm standard length measured by me) with further specimens under ISSB 3137. Two paratypes from the Nahr Taheri are in the Zoologischen Instituts und Zoologischen Museums der Universität Hamburg (ZMH 6091, 77.2-81.9 mm standard length) (Wilkens and Dohse, 1993; examined by me), 2 paratypes are in the American Museum of Natural History, New York (AMNH 40952), 1 paratype is in the Muséum national d'Histoire naturelle, Paris (1982-1018), 1 paratype is in the United States National Museum, Washington (USNM 227928), 2 paratypes are in the Academy of Natural Sciences, Philadelphia (ANSP 150977), and 2 paratypes are in CMNFI 1982-0368 (formerly IZA 7841).

Syntypes of Schizocypris brucei are in the Natural History Museum, London under BM(NH) 1913.4.15:100-109 (10 fish) and in the Zoological Survey of India, Calcutta under ZSI F9832/1 (1) (Menon and Yazdani, 1968; Eschmeyer et al., 1996).

Berg (1949) advanced the possibility that this species is the juvenile of Schizothorax zarudnyi since barbels and scale cover develop with age. However the tooth formula is very distinctive as is the anal fin branched ray count and lack of barbel development at all sizes in S. altidorsalis (Coad and Keyzer-de Ville, 2005).

Key characters

This species is characterised by a very high dorsal fin with a strongly denticulated spine. The spine is longer than the head and the denticles easily catch the skin when the fish are handled. This is particularly true of small fish, larger ones are not so snaggly. Bianco and Banarescu (1982) give values for spine length of 24.4-29.8% of standard length in altidorsalis, 19.4-20.4% in brucei, and 14.7-19.5% in ladigesi. For 35 altidorsalis 66.1-175.1 mm standard length examined here, spine length is 23.3-31.3% of standard length and 14.0-18.5% for 20 brucei 102.3-170.8 mm standard length. Dorsal spine length in head length is 0.7-0.9, mean 0.9 for altidorsalis, 1.2-1.6, mean 1.4 for brucei (see also below). Scales in the lateral line are 87-96 (brucei has 74-81 and ladigesi 78-88) according to Bianco and Banarescu (1982). Specimens examined by me have lateral line counts of 81-95, mean 87.4 for 60 altidorsalis and 73-91, mean 79.6 for 56 brucei, showing some overlap but scales are definitely smaller on average in altidorsalis.

Bianco and Banarescu (1982) describe the body as mostly scaled except on the anterior part of the breast (scaled on the mid-line of the back in front of the dorsal fin as in ladigesi, naked in brucei), and scales embedded on most of the body except the caudal peduncle and the lateral line; but see below.

Morphology

Dorsal fin unbranched rays 4, branched rays 7-8, anal fin unbranched rays 3, branched rays 5-6, pectoral fin rays 14-20, pelvic fin rays 7-9, lateral line scales 82-96, total gill rakers 24-30, reaching the third raker below when appressed in large fish but only one raker below in small fish, pharyngeal teeth usually 2,3,4-4,3,2(16) with variants 2,3,4-4,3,3(1), 2,3,5-4,3,2(2) and 2,3,5-4,4,2(1), and total vertebrae 43-45. Since spines are often broken off, the height of the dorsal fin can be measured as the longest branched ray. For this species it is 16.5-28.8% standard length (mean 24.3, 60 fish) while in S. brucei, the taxon fish in Sistan were formerly assigned to, it is 11.6-20.0% (mean 16.8, 56 fish), clearly distinguishing these species.

Scales are regularly arranged over most of the body. In some fish, scales near the tail are difficult to distinguish. Shoulder scales are moderately large anteriorly above the lateral line and decrease in size posteriorly. The back is naked in a narrow band for a short distance anterior to the dorsal fin (not so according to Bianco and Banarescu (1982) but visible in fish examined by me). Flank scales are small but those at the dorsal fin base are a little larger. Lateral line scales are larger than those on the flank but only on the anterior lateral line. The breast is scaleless according to Bianco and Banarescu (1982) but is scaled on the breast in large specimens and some small ones too. There is a pelvic axillary scale. The scale focus is anterior with radii on all fields and in overall shape is oval to rounded. Radii are found on all fields but are few in number (15).

The mouth is inferior, transverse, and slightly arched. The lower lip is developed only laterally. The lower jaw is covered by a horny sheath in some specimens, lost in others. There are usually no barbels (n = 105) although one fish, 70.1 mm standard length, had a minute pair of barbels hidden in the lip grooves and two other fish of similar size had respectively a single right and a single left minute barbel. S. brucei has small but protruding barbels on both sides in 15 fish, a left barbel only in 3 fish, a right barbel only in 5 fish and no barbels in 8 fish.

The gut is very elongate and complexly coiled. The anterior main row pharyngeal tooth is peg-like while the rest are spatulate with a deep central groove and crowns flared on each side of the groove.

Meristics for Iranian specimens are:- dorsal fin branched rays 7(2) or 8(38); branched anal fin rays 5(1) or 6(39); pectoral fin branched rays 14(1), 16(11), 17(23), 18(4) or 20(1); pelvic fin branched rays 7(1), 8(35) or 9(4); lateral line scales 81(1), 82(2), 83(3), 84(4), 85(6), 86(8), 87(10), 88(7), 89(5), 90(4), 91(3), 92(3), 93(3) or 95(1); total gill rakers 24(1), 25(4), 26(5), 27(10), 28(12), 29(6) or 30(2); pharyngeal teeth 2,3,4-4,3,2(16), 2,3,4-4,3,3(1), 2,3,5-4,3,2(2), or 2,3,5-4,4,2(1); and total vertebrae 43(16), 44(33) or 45(11).

Sexual dimorphism

Unknown.

Colour

The back is bluish and the flanks and belly are silvery. The flanks may have a few small to numerous black spots.

Size

Reaches 17.5 cm standard length.

Distribution

This species is endemic to the Sistan basin of Iran, and presumably adjacent Afghanistan (Bianco and Banarescu, 1982; J. Holčík, in litt., 1996). The distribution of this species (as S. brucei) in Khorasan and Gorgan reported by Wossughi (1978) is incorrect.

Zoogeography

A relative of other schizothroacine species found along the mountain chain from Iran to China. See also the genus Schizothorax.

Habitat

Occurs in a wide range of habitats from ditches to hamuns. Reported from pools in dry river beds and still, reedy channels in Sistan. E. Penning (pers. comm., 28 July 2005) states that this was the dominant fish species in Hamun-e Puzak and Hamun-e Saberi in April 2005 after a dry period when the hamuns flooded at the end of February. The fish enter the flooding hamuns from the upstream parts of rivers. In July, water levels fell from1-2 m to less than 1 m and this species was absent, presumably having returned to the more permanent rivers. They were also observed swimming up a fish staircase at the Sistan Dam.

Age and growth

Unknown.

Food

The principal food is aufwuchs and detritus as evidenced by the sectorial mouth and elongate gut. Gut contents are a fine mush. E. Penning (pers. comm., 28 July 2005) observed filamentous algae in the gut.

Reproduction

Most fish examined by me were young and no data on reproduction is available. E. Penning (pers. comm., 28 July 2005) noted that fish at 20-30 cm caught in April 2005 had eggs 1 mm in diameter and were approaching or ready for spawning.

Parasites and predators

Jalali et al. (1995) describe a new species of monogenean, Dactylogyrus schizocypris, from fish taken in the Chahnimeh water reservoir near the Hamun Lake in Sistan. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Lamproglena compacta and Lernaea sp. on this species.

Economic importance

Individual fishermen caught 5-10 kg per day of this species in 2005 although they considered individuals as small and catches very low. Fish were 10-20 cm long with some larger ones at 20-30 cm (E. Penning, pers. comm., 28 July 2005). Rahimabadi et al. (2009) assessed the lipid quality of this species and found it to be less than that of Schizothorax zarudnyi.

Conservation

Further knowledge of biology is needed to assess conservation status.

Further work

The biology of this species needs study.

Sources

Type material: See above, Schizocypris altidorsalis (IZA 8169, IZA 7841, IZA 7844, ZMH 6091 and CMNFI 1982-0368).

Iranian material: CMNFI 1979-0072, 2, 120.8-162.9 mm standard length, Sistan, river near Zabol (ca. 30º58'N, ca. 61º28'E); CMNFI 1979-0223, 1, 19.9 mm standard length, Sistan, ditch south of Lutak (30º45'N, 61º24'E); CMNFI 1979-0224, 13, 48.2-77.5 mm standard length, Sistan, Hirmand River effluent (30º53'N, 61º27'E); CMNFI 1979-0225, 1, 147.7 mm standard length, Sistan, Hirmand River effluent (30º58'N, 61º28'E); CMNFI 1979-0226, 107, 60.0-82.5 mm standard length, Sistan, pool east of Kuh-e Khajeh (30º57'N, 61º17'E); CMNFI 1979-0228, 16, 18.1-73.8 mm standard length, Sistan, ditch 1 km from Zabol (31º02'N, 61º31'E); CMNFI 1979-0229, 11, 61.0-84.9 mm standard length, Sistan, ditch 5 km from Zabol (31º03'N, 61º33'E); CMNFI 1979-0231, 2, 17.5-19.9 mm standard length, Sistan, irrigation jube 3 km from Zabol (31º01'N, 61º32'E); CMNFI 1979-0232, 23, 41.8-78.8 mm standard length, Sistan, irrigation ditch 11 km from Zabol (30º58'N, 61º36'E); CMNFI 1979-0233, 2, 66.0-71.5 mm standard length, Sistan, irrigation ditch 15 km from Zabol (ca. 30º57'N, ca. 61º38'E); CMNFI 1979-0234, 15, 13.9-82.4 mm standard length, Sistan effluent of the Hirmand River near Zahak (30º54'N, 61º40'E); CMNFI 1979-0236, 1, 14.4 mm standard length, Sistan, irrigation ditch 27 km from Zabol (ca. 30º52'N, ca. 61º22'E); CMNFI 1979-0237, 5, 21.4-72.1 mm standard length, Sistan, irrigation ditch 18 km south of Zabol (30º53'N, 61º27'E).

Comparative material: USNM 182276, 5, 95.7-153.1 mm standard length, Afghanistan, Arghandab Reservoir (ca. 31º51'N, 65º55'E); USNM 182277, 1, 159.7 mm standard length, Afghanistan, Arghandab River at Kandahar (ca. 31º35'N, 65º45'E); USNM 182281, 3, 71.2-79.9 mm standard length, Afghanistan, Lashka-dah, Helmand River (ca. 31º35'N, ca. 64º21'E); USNM 182282, 5, 142.1-159.4 mm standard length, Afghanistan, Laskha-dah area (ca. 31º35'N, ca. 64º21'E); ZMUC 261629-34, 6, 131.4-203.7 mm standard length, Afghanistan, Sistan, Feyzabad (31º28'N, 61º31'E).

Material of Schizocypris brucei and S. ladigesi, nominal species not found in Iran, are listed in Coad and Keyzer-de Ville (2005)

Genus Schizopygopsis
Steindachner, 1866

This genus contains about 8 species distributed from Iran along the Himalayas to China. There is a single species in Iran.

The osmans are characterised by an elongate and almost cylindrical body, almost scaleless, scales being restricted to the complete lateral line, the flank above the pectoral fin and as enlarged scales around the vent and anal fin base, the mouth is terminal to inferior, the lower jaw is sometimes covered by a horny sheath, lips are present but may only be developed at the mouth corners, no barbels, pharyngeal teeth in 2 rows, spatulate or cochleariform, short dorsal and anal fins, dorsal fin last unbranched ray thickened (but not strongly) and serrated or denticulate (denticles lost with age), peritoneum black, and gut very elongate.

Schizopygopsis stoliczkai
Steindachner, 1866

Common names

kopur-e barfi (= snow trout).

[lozhnyi osman or false osman in Russian; Pamir snowcarp].

Systematics

The type locality of this species is a stream near Hanle Monastery, Ladakh, India. Syntypes are in the Naturhistorisches Museum Wien under NMW 9255 (1), NMW 9256 (1), NMW 51472 (9) and NMW 51473 (2) (Eschmeyer et al., 1996; 1997 Vienna card catalogue).

The species name is spelt Stoličkai by Steindachner (1866) which becomes correctly stoliczkai since accents are not used in Latin nor capitals for the scientific species name. It is often spelt stoliczkae in the general literature.

Schizopygopsis stoliczkae infraspecies sewerzowi Herzenstein, 1890, originally described from the Bulun Kul and Karasu, Amu Darya basin in the Pamir Mountains has no taxonomic validity (it occurs together with the typical form in the upper Amu Darya) but it was used to characterise the form in the Sistan basin of Iran (Berg, 1948-1949). However, Annandale and Hora (1920) report both this and the stoliczkae form, without intermediates, in Sistan. The sewerzowi form differs from the typical form by having a larger eye (1.2-1.3 times in interorbital width as opposed to 1.5-1.7 times and 3.8-4.4 times in head length as opposed to 4.8-5.7 times), a spotted rather than monotone body, smaller size (much less than half that of the typical form), somewhat deeper body, and more oblique mouth with apex at the lower eye level. Data presented below under Food indicates that this species can be very plastic in its characters so subspecies designations would require extensive study of both characters and ecology. The Iranian fish are poorly represented by specimens and are referred here simply to the species.

Syntypes of sewerzowi (dated 1891 in Eschmeyer et al. (1996)) are in the Zoological Institute, St. Petersburg under ZISP 8747 (2), ZISP 8748 (2), ZISP 8749 (2), ZISP 8780 (1), ZISP 8901 (2) (Eschmeyer et al., 1996).

Kähsbauer (1964) reports a hybrid between Schizothorax schumacheri and Schizopygopsis stoliczkae from Sistan which may in fact be Schizocypris altidorsalis.

Key characters

The anal fin base has a sheath of enlarged scales and there are a few scales in the shoulder region and along the lateral line but the body is nearly naked, the anal fin has 5 branched rays, barbels are absent, and pharyngeal teeth are in 2 rows.

Morphology

Dorsal fin branched rays 7-9, usually 8, after 3-4 unbranched rays, anal fin branched rays 5-6, usually 5, after 2-3 unbranched rays, pectoral fin branched rays 12-20, and pelvic fin branched rays 8-10. Total vertebrae 48 (Howes, 1987). The lateral line is complete, a pelvic axillary scale is present and the anus and anal fin base are sheathed in enlarged scales. Scales are also present on the anterior lateral line and between there and the anterior pectoral fin base. A double row of scales extends forward from the anal sheath to the pelvic fin bases. Lateral line scales 96-120.

The dorsal fin spine is weakly developed and lacks denticles in large fish while young have many well-developed denticles. The gut is very elongate and coiled. The lower jaw is strong, with a dark brown, horny plate. The pharyngeal teeth number 3,4-4,3 and have a rounded base becoming spatulate distally with a rounded, hooked tip. Some teeth have a weak, flat cusp with a bump posteriorly below a rounded tip.

Meristic values for Iranian specimens are:- dorsal fin branched rays 8(3); anal fin branched rays 5(3); pectoral fin branched rays 17(1), 18(1) or 19(1); pelvic fin branched rays 9(3); lateral line scales 91(1) or 92(1); total gill rakers 11(3), reaching the raker below when appressed; and total vertebrae 48(1) or 50(2).

Sexual dimorphism

Tubercles are present on the anal fin of two male specimens from Sistan in a single file on the last unbranched and first 3 branched rays. They number up to 8 and are often widely spaced. Tubercles are found low on the flank above the anal fin and anus and on the upper flank behind the dorsal fin level. They also line the posterior lateral line. There are also a very few tubercles on the top of the head, side of the snout, below the eye and on the operculum, widely scattered and small.

Colour

Dark or yellowish with small dark brown or blackish spots extending onto the fins, or olive with large grey spots, or bluish-grey. Spots can be so numerous as to give the appearance of an overall blackish colour. Tilak (1987) however states that blackish spots are absent in both old and freshly preserved material. Both spotted and non-spotted forms occur in Sistan. The belly is whitish. Fins are pink. Peritoneum dark brown to black.

Size

Reaches 75.4 cm but the Sistan form is only up to 22.0 cm and is regarded as a dwarf form by Annandale and Hora (1920).

Distribution

Found in the mountainous areas of Afghanistan, Pakistan, India and western China. In Iran, restricted to the lowland Sistan basin where reported from the Hirmand River delta and 8 miles east of Lab-e Baring (Annandale, 1921; Vijayalakshmanan, 1950).

Zoogeography

The presence of this species in Sistan is an example of a riverine highway enabling species usually found at higher altitudes to penetrate into lowlands. See also the genus Schizothorax.

Habitat

Outside Iran the adults favour the main river while young are found in shallow streams and pools.

Age and growth

Savvaitova et al. (1989) cite a life span up to 22 years. Maturity in the Sistan form is attained at 18.0 cm. Berg (1949) reports a male fish 14 cm long with fairly well-developed testes and tubercles on the anal fin which bears out the dwarf nature of the Sistan fish.

Food

Savvaitova et al. (1989) examined feeding in certain lakes of the Pamirs at altitudes over 3220 m. This is much higher than the Sistan populations and the data may not be relevant. However the false osman is a poorly studied species so such information gives a basis for comparison and future research. The false osman is a very plastic species and can adapt to a variety of conditions. The osmans of the Pamirs were divisible into four groups: herbivores, detritivores, molluscivores and predators reflected in the structure of the gut and its length, the number of gill rakers, eye diameter, length and position of fins, and the shape of the horny plate on the lower jaw. Food includes higher aquatic plants, aquatic insects, diatoms and blue-green algae, detritus, molluscs and fish. In herbivores consuming periphyton, the horny plate on the lower jaw is rasp-shaped with the pointed end aimed anteriorly. In detritivores, the plate is sharper, and in predators, it is larger with the pointed edge directed upwards to grasp prey. Predatory behaviour only develops in fish over 30 cm, until which they eat plants.

Chaudhary et al. (1991) indicate that in a Pakistani population gut length increases with age and diet changes gradually from an omnivorous to a herbivorous one.

Reproduction

Spawning in Asia occurs generally in June and July. Eggs are large at 2.0 mm diameter.

Parasites and predators

None reported from Iran.

Economic importance

None.

Conservation

This species is rarely collected in Sistan and may require conservation measures.

Further work

Detailed surveys are needed to document the occurrence of this species in Sistan and to study its biology in a lowland habitat. Comparative studies with samples of the species from other, remote localities may demonstrate some level of taxonomic distinction as is seen in other schizothoracines of Sistan.

Sources

Iranian material: BM(NH) 1905.4.7:1-2, 2, 113.0-116.5 mm standard length, Sistan (no other locality data); ZISP 25854, 1, 111.5 mm SL, Sistan (no other locality data).

Comparative material: BM(NH) 1931.10.26:2-4, 2, 85.5-138.6 mm standard length, India, Ladkah, stream into Pangong Lake (no other locality data).

Genus Schizothorax
Heckel, 1838

The snow trouts, mountain barbels or Indian trouts are found from Iran to China, favouring mountainous areas but occasionally in lowlands. There are about 56 species (more if some other genera are included), with 3 in Iran.

The genera Racoma McClelland and Griffith in McClelland, 1842 and Aspiostoma Nikol'skii, 1897 are synonyms of Schizothorax Heckel, 1838 (Eschmeyer, 1990).

Schizothorax intermedius, and other species, have been placed in the genus Schizothoraichthys Misra, 1962 (e.g. in Tilak (1987). However Schizothoraichthys is regarded as a synonym of Schizopyge Heckel, 1847 by Jayaram (1981) or of Schizothorax (see Talwar, 1978; Eschmeyer, 1990). Schizopyge is itself regarded as a synonym of Schizothorax by some authors, e.g. Talwar (1978). Talwar (1978) separates the genus Oreinus McClelland, 1839 from Schizothorax by the margin of the lower jaw having a firm and hard horny covering which is thickest internally and a thick lower lip with a free posterior edge forming a sucker. Schizothorax has a non-suctorial lower lip and a lower labial fold interrupted or entire in the middle. However Talwar and Jhingran (1991) contradict this view and use Schizothoraichthys for Schizothorax and Schizothorax for Oreinus. Tilak (1987) recognises the name Schizothorax for fishes with strip of papillated tissue on the chin and Oreinus as a synonym; Schizothoraichthys is used then for fishes without the papillated chin. Oreinus is regarded as a synonym of Schizothorax by Jayaram (1981) and Eschmeyer (1990). Mirza (1991a; 1991b) recognises a tribe Schizothoracini, with the genera Schizothorax, Schizopyge, Racoma McClelland (and Schizocypris). I have retained Schizothorax here as the oldest name in view of these conflicting opinions.

There are various records of nominal Schizothorax species from the Helmand River basin in Afghanistan summarised in Coad (1981d); they have not been reported from the Sistan lowlands of Iran.

This genus is characterised by an elongate and almost cylindrical body, very small scales, over 100 in the series next to the lateral line, scales in complete lateral line somewhat larger, the vent and anal fin base are sheathed in enlarged scales and there may be enlarged scales near the pectoral fin and edge of the gill opening, dorsal and anal fins are short, dorsal fin with a thickened last unbranched ray bearing denticles (denticles lost with age), pharyngeal teeth in 3 rows and hooked at the tip, 4 barbels (rostral and maxillary), mouth inferior or subterminal, lower jaw may have a horny sheath, a papillated area on the chin may be present or absent, the lower labial fold may be interrupted or not in the middle, elongate gut and black peritoneum, and poisonous eggs. Members of the Schizothoracinae tribe are of polyploid origin with 2n=98 and 3n=148.

The ancestors of the schizothoracines in general were barbinines in the eastern part of the Qinghai-Xizang Plateau as it rose and water temperatures decreased in the late Miocene to early Pliocene (Sizhong, 1995). Primitive genera like Schizothorax migrated westwards earlier and further than more specialised genera such as Schizopygopsis (although both reach their westernmost distribution in Iran).

These fishes generally prefer rapids and pools of the larger streams at temperatures of 8-22°C although some occur in lakes with inflowing streams (Sharma, 1988). They are found in streams above 3000 m. Food varies from detritus to insects, plankton and fish depending on the species. The spawning season may be in late summer and early fall or in spring. Egg counts vary from a few hundred to over 50,000 and egg diameters may attain 3.6 mm. Some species show a spawning migration from warm lakes to cold streams.

Deaths have occurred from eating poisonous eggs of members of this genus but none are reported from Iran (see under S. zarudnyi). Symptoms include abdominal pain, nausea, vomiting, diarrhoea, dizziness, headache, fever, bitter taste, dryness of the mouth, intense thirst, sensation of chest constriction, cold sweats, rapid irregular weak pulse, low blood pressure, cyanosis, pupillary dilatation, syncope, chills, dysphagia and tinnitus. Severe cases show muscular cramps, paralysis, convulsions, coma, and death. Victims generally recover within 3-5 days with supportive treatment but it may take longer. Treatment is symptomatic and there is no known antidote or therapeutic data available. The patients' stomach should be evacuated as soon as possible after ingestion of eggs (Halstead, 1967-1970; Coad 1979b). Fish eaten during the breeding season should be cleaned with care to remove all traces of the eggs to avoid contamination of the flesh as cooking does not destroy the toxin.

Schizothorax esocinus
(Heckel, 1838)

Reported from the Helmand River drainage of Afghanistan; records summarised in Coad (1981d). No Iranian record.

Schizothorax intermedius
McClelland, 1842

Variation in ventral head structure, above and below

Common names

mawda (in Sistan), marinka.

[marinka obyknovennaya or common marinka in Russian.

Systematics

Schizothorax intermedius was described from the "Cabul river at Jullalabad. Tarnuck River" in the Indus River basin.

Oreinus plagiostomus McClelland, 1842 described from the "Helmund river at Girdun Dewar" in Afghanistan, Racoma brevis McClelland, 1842 described from the "Helmund River", Racoma labiatus McClelland, 1842 described from "Pushut, Koonar river near Jullalabad" in the Indus River basin but also reported from the Helmand River basin by Annandale and Hora (1920) and Schizothorax ritchieana McClelland, 1842 described from "Affghanistan. In the Helmund there is a variety of this species ..... which will probably prove to be distinct." are probably synonyms (Berg, 1949).

Berg (1948-1949) reviews three morphae or forms of this species which indicate the great variation in this taxon. He also notes how barbel length varies independently of morpha and how the lips may be very strongly developed. The forms are as follows and the latter two were originally described as distinct species and are recognised as such by some authors: typica, eurystomus Kessler, 1872 and fedtschenkoi Kessler, 1872. Eurystomus has a transverse jaw covered by horny padding and a strong or weak dorsal fin spine. Fedtschenkoi has a lower jaw without the horny pad and the lower lip continuous or interrupted (the form with an interrupted lip is called morpha irregularis Day, 1896; the date is incorrect and should be 1877) and the dorsal fin spine is weak with 10-22 denticles extending to the mid-point or two-thirds along the ray. The forma typica has a crescent-shaped lower jaw without horny padding, the dorsal fin spine is well-developed with 12-32 denticles extending distally beyond the mid-point, and the lower lip is interrupted.

Schizothorax schumacheri Fowler and Steinitz, 1956 described from "Zabol, Eastern Iran" is also a synonym (Saadati, 1977). The holotype of Schizothorax schumacheri (ANSP 71950) at 244 mm total length and a paratype (ANSP 71951) at 130 mm to the end of the broken caudal fin, are stored in the Academy of Natural Sciences of Philadelphia (Böhlke, 1984).

Kähsbauer (1964) reports a hybrid between Schizothorax schumacheri and Schizopygopsis stoliczkae from Sistan which may in fact be Schizocypris altidorsalis.

Tilak (1987) reports Schizothorax richardsonii (Gray, 1832) from Sistan based on 2 fish in the Zoological Survey of India, Calcutta (F. 1226-1227/1), which I have not seen, and this may be the correct name for this fish.

Key characters

Gill raker counts and distribution separate this Schizothorax from others in Iran.

Morphology This is a very variable species, depending on habitat (Mirzaev, 1998). Dorsal fin with 2-4 unbranched and 5-9, usually 8 (Kullander et al. (1999) give 6-7 for their Kashmir specimens of S. curvifrons), branched rays, anal fin with 1-3 unbranched and 4-7, usually 5, branched rays, pectoral fin branched rays 14-19 and pelvic fin branched rays 7-10, usually 8. Lateral line scales 85-121, scale series next to the lateral line 115-165. Pharyngeal teeth 2,3,5-5,3,2. Gill rakers 10-17 (Kullander et al. (1999) give 21-28 for their Kashmir specimens of S. curvifrons). Total vertebrae 48 (Howes, 1987) or 40-43  (2000a) – presumably excluding 4 Weberian vertebrae). There is considerable variation in lower jaw form in specimens attributed to this species. The lower jaw may be crescent-shaped with or without a sharp horny sheath, or covered with a deciduous horny layer, or transverse and covered by a horny sheath. Lips may be interrupted medially or continuous, and can be very strongly developed. The dorsal fin spine may be well-developed with numerous denticles or weakly-developed with denticles not beyond the middle of the spine. Various morpha or infraspecies have been described to refer to these forms (see Berg, 1948-1949). Barbel length is highly variable. Young about 30 mm long have a naked body and no barbels. The karyotype is 2n=98-100. Usually silvery and occasionally with minute black spots on the upper half of the body but usually without spots. The head is olive-green. In preservative a pale greyish brown above, flanks and lower surfaces brilliant silvery or light yellow. Fins greyish to pale olive with lower ones whitish. Iris bright silvery white.

SizeReaches 60.0 cm (Solijonov, 2007).

Sexual dimorphism

Unknown.

Colour

Usually silvery and occasionally with minute black spots on the upper half of the body. In preservative a pale greyish brown above, flanks and lower surfaces brilliant silvery. Fins greyish to pale olive with lower ones whitish. Iris bright silvery white.

Size

Reaches 50 cm.

Distribution

Found in the basins of the Indus, the Amu Darya and Syr Darya rivers, the Tarim basin and the Helmand (= Hirmand) of Afghanistan and Iran.

Zoogeography

See under genus description.

Habitat

Reported from both lotic and lentic environments but little is known of its environmental requirements.

Age and growth

Life span is at least 8 years. Solijonov (2007), in a study of fishes in the Pamir-Alai Transboundary Conservation Area, found this species as being most active in the evening and so is seldom caught in the daytime when it hides in refuges among rocks. It maintains station in fast water of Pamir-Alai mountain streams behind rocks and in whirlpools. Males mature here at 2-3 years and females at 3-4 years and spawning takes place in mid-May.

Food

Food is small aquatic fauna, vegetation and detritus. Akhrorov and Kondur (1981) found macrophytes, detritus and molluscs to be important foods in a Pamir lake, varying with the year of sampling such that molluscs dominated in one year and macrophtyes in another.

Reproduction

Spawning takes place between May and September, depending on locality, and up to 8678-59,895 eggs are produced in fish 21.5-37.1 cm and 211-913 g (Mitrofanov et al., 1988). Spawning is probably non-annual in some areas (Maksunov, 1971).

Parasites and predators

None reported from Iran.

Economic importance

None.

Conservation

This species is not well-documented in Iran and no assessment of conversation needs can be made. It may be a stray from higher latitudes in Afghanistan.

Further work

Surveys should be done to confirm its presence in Iran.

Sources

Iranian material: None.

Schizothorax labiatus
(McClelland, 1842)

Reported from the Helmand River drainage of Afghanistan; records summarised in Coad (1981d). May be a synonym of Schizothorax intermedius (McClelland, 1842). No Iranian record.

Schizothorax pelzami
Kessler, 1870

Common names

shir mahi.

[Zakaspiiskaya marinka or Transcaspian marinka, and forel, in Russian].

Systematics

Schizothorax raulinsii Günther, 1889 described from a skin from the "Hari-rud River, near Khusan" and Schizothorax pelzami iranicus Karaman, 1969 are synonyms. Wossughi (1978) considers Schizothorax pelzami iranicus to be only a large specimen of S. p. pelzami and Coad and Keyzer de Ville (2004) concur.

The holotype of Schizothorax Poelzami is in the Zoological Institute, St. Petersburg (ZISP 8036, 265 mm total length) and is from "Fl. Schach-rud. accursus fl. Sefid-rud in Persia. 1889. Univ. Petropol.". The Shah Rud or Shah River is a tributary of the Safid Rud of the Caspian Sea basin but this species does not occur there. Berg (1948-1949) cites A. N. Derzhavin who suggests that this Shah-rud is south of Astrabad (= Gorgan). There is a Shahr Now River in the Tedzhen or Hari Rud basin where this species is found (shahr is the Farsi for city and may have been a general term for a major river of northeastern Iran as it flows through a city). However, the type probably came from the environs of the city of Shahrud (= Emamshahr) in the Damghan basin, a sub-basin of the Dasht-e Kavir basin (see below).

Another specimen listed as a type from the "Schah-Roude. Persia. Pelzame. St. Petersburg University" measuring 78.6 mm standard length is in the Natural History Museum, London (BM(NH) 1897.7.5:24). Kessler (1870) and Eschmeyer (1998) list 4 syntypes so two appear to be lost but Coad and Keyzer de Ville (2004) point out a disparity in size range (120-180 mm for the syntypes according to Kessler (1870)) while the London fish is too small and may not be a type despite its label.

The type of Schizothorax raulinsii is from the Hari-rud River, near Khusan, Afghanistan. BM(NH) 1886.9.21:181, is a skin marked as a syntype, 312.8 mm standard length, Hari-rud River, near Khusan, presumably the skin referred to by Eschmeyer et al. (1996). The skin is also marked Schizothoraxaitchisonii” (= S. raulinsii, a synonym of S. pelzami); Albert Günther of the British Museum apparently confused the names Aitchison and Raulins, collectors of specimens on the Afghan Delimitation Commission; aitchisonii was never used and the skin is a syntype of raulinsii. Eschmeyer lists a skin and 2 other syntypes in the Natural History Museum, London and 2 syntypes in the Zoological Survey of India, Calcutta (ZSI F11477-78). These 4 additional specimens are presumably the 4 smaller fishes mentioned by Günther (1889) in his original description as being collected at Bezd on the Jam River in Iran. BM(NH) 1886.9.21:171-172, 2, 87.8-100.9 mm standard length, Bezd, are not currently listed as syntypes in the British Museum however.

Schizothorax pelzami iranicus Karaman, 1969 is described from "Teheran in Quelle" (= Tehran in a spring) based on a single specimen. Schizothorax pelzami does not occur in the Namak Lake basin in which Tehran lies and the subspecies may have come from the Damghan part of the Dasht-e Kavir basin. The subspecies differs from the type subspecies by having a weakly ossified spiny ray in the dorsal fin (only the first half with small teeth), smaller eyes, longer snout and an overall brown to blackish-grey body colour with all fins, lips and barbels dark-coloured as opposed to the sharp boundary between the dark brown dorsal side and the light ventral side (Karaman, 1969). The Damghan part of the Dasht-e Kavir basin is the type locality of the species and S. p. iranicus is a synonym (Coad and Keyzer de Ville, 2004).

The holotype of S. p. iranicus is in the Zoologischen Instituts und Zoologischen Museums der Universität Hamburg (ZMH 4116, 327.5 mm standard length).

Starostin (1936) reports a hybrid of this species and Capoeta heratensis (= Capoeta capoeta) from Turkmenistan.

Key characters

This is the only schizothoracine species in northeast Iran and is easily recognised by its high lateral line scale count and the enlarged scales around the anus and anal fin.

Morphology

Dorsal fin branched rays 7-8 after 3-4 unbranched rays, anal fin branched rays 5-6, usually 5, after 3 unbranched rays. Pectoral fin branched rays 16-21, pelvic fin branched rays 7-9. The dorsal fin spine is very strong and thick with well-developed and widely-spaced teeth. Lateral line scales 84-108, lateral series scales 155-170, about 32 between the dorsal fin spine and the lateral line and about 27 between the lateral line and pelvic fin. The belly is scaled up to the isthmus. There is a scaled pelvic axillary process. The anal papilla and anal fin lie in a groove formed by enlarged scales, the groove extending about one third to half way between the anal fin origin and pelvic base. Scales are oval and obliquely inserted into scale pockets on mid-flank, sloping backwards postero-dorsally. The focus is subcentral anterior and radii are present on all fields. Circuli are few in these small scales. Pharyngeal teeth usually 2,3,5-5,3,2. Teeth are rounded with an evident hooked tip and posterior teeth have a short to medium flat grinding surface below the tip. Teeth may also be spatulate or have a spatulate shape with the hollow filled in. Gill rakers 9-15, relatively short and reaching the adjacent raker or slightly beyond when appressed. Occasional rakers are forked. The mouth is inferior. The lower lip is interrupted medially. The lower jaw may have a sharp horny sheath but this is mostly lacking. Mouth shape varies from a u-shape to a sector mouth (a gentle arch), the latter with a horny edge. The anterior barbels extend back to the anterior eye margin or the mid-eye while the posterior barbels extend to the rear eye margin or beyond. Barbel size is variable and not obviously related to size. The gut is very elongate and coiled.

Meristic values for Iranian specimens are:- dorsal fin branched rays 7(11) or 8(22); anal fin branched rays 5(33); pectoral fin branched rays 16(2), 17(4), 18(12), 19(6), 20(8) or 21(1); pelvic fin branched rays 7(2), 8(30) or 9(1); lateral line scales 85(2), 86(2), 88(2), 89(1), 90(2), 91(2), 92(2), 93(2), 95(2), 98(2), 99(7), 100(3), 102(1), 104(1), 105(1) or 108(1); total gill rakers 9(1), 10(5), 11(6), 12(7), 13(8), 14(1), 15(2), 16(1), 17(1) or 18(1); pharyngeal teeth 2,3,5-5,3,2(15), 2,3,4-5,3,2(2), 2,3,5-4,3,2(1), 2,2,5-5,3,2(1), 1,3,5-5,3,2(1); and total vertebrae 43(6), 44(10), 45(1), 46(2), 47(2), or 49(1).

Sexual dimorphism

A male specimen, 123.9 mm standard length, caught on 6 April had small to moderate sized tubercles on the top and sides of the head but these were not fully developed. The largest tubercles are found between the nostrils and the upper lip on the snout. No tubercles were noted on the fins. A fish taken on 5 November also had small but distinctive tubercles.

Colour

The overall coloration is silvery without any pattern but the back and upper flank are blackish to olive or brassy and the belly is whitish in small fish to a strong yellow in large fish. The back may be iridescent blue-green. The lateral line may be lighter than the surrounding flank, appearing as a thin, whitish line. The lips, pectoral, pelvic and anal fins are yellow. Fins bases are bright orange, the gill slit has a bright orange streak and the isthmus is bright orange. All fins may be flushed with red in freshly caught material and the lower flank and belly can have pinkish tinges. The iris is red dorsally. The peritoneum is a dark brown.

Preserved fish have a uniform brown colour with faint to dark speckles arranged irregularly on the flank. There are no obvious patterns on the fins although they are darkened by melanophores on both rays and membranes.

Günther (1889) reports the caudal fin as black, but this is possibly a dried or otherwise abnormal specimen. The colouration of the iranicus nominal subspecies in the original description (cited above) is not borne out by specimens from Damghan, the presumed locality of the type specimen, and again may be an artefact of preservation or simply a variation.

Size

Attains 54 cm (Muhomedieva, 1967) and reputedly 3 kg in qanat specimens (R. J. Behnke, in litt., 1981).

Distribution

Found in the Tedzhen and Murgab rivers of Afghanistan and Turkmenistan including Iranian drainages of the former known as the Hari River in its Iranian reach (Aliev et al., 1988). It is recorded from the Jam River, the Sharak River, the Akhland River near Mashhad, the Kashaf Riverand various smaller water bodies in Khorasan, the upper Kal Shur, Jajarm and Jovein rivers in the Dasht-e Kavir basin, as well as Cheshmeh Ali at Damghan and Cheshmah Badash near Shahrud further west, the westernmost distribution of the schizothoracine fishes (Günther, 1889; Nikol'skii, 1897; 1899; Abdoli, 2000; Coad and Abdoli, 2000b).

Wossughi (1978) records this species from the Hamun-See (= Sistan) but this is an error.

Zoogeography

Saadati (1977) found slight differences between fish from the Dasht-e Kavir basin and from the Hari River basin, in raker counts and caudal peduncle depth. He concludes that isolation in the Dasht-e Kavir basin is relatively recent and migration has occurred westwards in the past 15-25,000 years. The occurrence of this species in the western Dasht-e Kavir basin (the Damghan basin) is the westernmost distribution of the schizothoracine fishes. See also the genus Schizothorax.

Habitat

Found in springs, streams, rivers and qanats, the principal habitats of northeast Iran but environmental requirements are unknown.

Age and growth

Life span exceeds 7 years (Muhomedieva, 1967). Abdoli et al. (2007) found males reached 23.3 cm and 145.6 g and females 34.0 cm and 428 g in the Laiinsoo River. The sex ratio was male:female 2.5:1.

Food

The diet is 93-99% fishes including Cyprinus carpio (Muhomedieva, 1967). Other foods include small Capoeta capoeta, Cyprinus carpio, chironomids, caddis flies, dragonflies, other aquatic insects, and plant material (Aliev et al., 1988). Crustaceans, plant fragments and filamentous algae, and possibly fish eggs, may also be found in gut contents. Abdoli (2000) lists Plecoptera, Ceratopogonidae, Trichoptera, Ephemeroptera, Chironomidae and Simuliidae. Abdoli et al. (2007) found fish in the Laiinsoo River had a diet dominated by chrironomid pupae and simuliid larvae, with 9 other benthic invertebrate food items taken.

Reproduction

Egg diameter reaches 2.0 mm and numbers reach 36,300 eggs (Aliev et al., 1988). Iranian fish caught on 6 April had developing eggs suggesting a spring or early summer spawning period.

Parasites and predators

Barzegar et al. (2008) record the digenean eye parasite Diplostomum spathaceum from this fish from the Hamun Lake, Sistan (sic - misidentification of the fish species or locality confusion).

Economic importance

None.

Conservation

The numbers and details of distribution need to be examined before an assessment can be made.

Further work

The biology of this species in Iran is unknown in detail.

Sources

Type material: See above, S. pelzami (ZISP 8036) (and possibly (BM(NH) 1897.7.5:24)) and S. p. iranicus (ZMH 4116).

Iranian material: CMNFI 1993-0124A, 1, 118.7 mm standard length, Semnan, Cheshmeh Ali-Damghan (36º17'N, 54º05'E); CMNFI 2007-0003, 8, 94.3-188.1 mm standard length, Semnan, Cheshmeh Ali (ca. 36º17'N, ca. 54º46'E); CMNFI 2007-0004, 6, 75.6-109.5 mm standard length, Semnan, Cheshmeh Bedasht (ca. 36º35'N, ca. 55º03'E); CMNFI 2007-0012, 2, 77.8-127.6 mm standard length, Khorasan, qanat at Bagh-e Jan (ca. 36º00'N, 58º38'E); CMNFI 2007-0013, 2, 55.9-80.4 mm standard length, Khorasan, qanat 5 km north of Boghai (ca. 36º02'N, ca. 59º31'E); CMNFI 2007-0014, 3, 88.7-98.1 mm standard length, Khorasan, Kuh-e Sang Park, Mashhad  (ca. 36º18'N, ca. 59º36'E); BM(NH) 1914.1.1:16-17, 2, 98.0-163.9 mm standard length, Khorasan, Kashaf River, Mashhad (no other locality data); BM(NH) 1914.1.1:18-20, 3, 138.5-242.0 mm standard length, Khorasan, small stream near Mashhad (no other locality data); BM(NH) 1914.1.1:21-23, 2, 151.2-165.7 mm standard length, Khorasan, Cheshmeh-e Saby (no other locality data); BM(NH) 1914.1.1:24-29, 5, 147.6-188.8 mm standard length, Khorasan, Langar, Jam River (35º23'N. ca. 60º25'E).

Schizothorax plagiostomus
Heckel, 1838

Described from Kashmir; records summarised in Coad (1981d) for the Helmand River basin in Afghanistan but these may refer to Oreinus plagiostomus, a probable synonym of Schizothorax intermedius (q.v.). No Iranian record.

Schizothorax zarudnyi
(Nikol'skii, 1897

Common names

hamun mahi (= hamun or lake fish), ماهي خواجو (= mahi khvaju or khaju, after the historic island in Lake Hamun), sefidak (=white fish) or vatani (A. A. Pasand, pers. comm., 5 November 2000), shir mahi (= milk fish), anjak (Fowler and Steinitz (1956) report three kinds of fish being caught by fishermen in Sistan named anjaq (hence Oreinus anjac, see below), mawda (? unknown) and mahrmahé (= mar mahi, snake fish, probably a nemacheilid)).

Systematics

The holotype of Aspiostoma zarudnyi is in poor condition with the tail detached and the body impaled on a wooden spike when examined by me. Nikol'skii (1897) in his original description states in Latin "Specimen valde destructum". It is in the Zoological Institute, St. Petersburg (ZISP 11195a) and measures about 265 mm standard length. Nikol'skii (1897) gives the catalogue number as 11115 (sic, incorrectly according to Berg (1949)) and the type locality as "Palus Neizar in Seistan. 3.VI.96.". Berg (1949) gives the collection locality as "Neizar near the southern tip of Lake Hamun-i-Farah, western edge of the Helmand delta in northwestern Seistan" based on Zarudnyi (1901).

This species was originally described in the genus Aspiostoma Nikol'skii, 1897, a synonym of Schizothorax Heckel, 1838 (Eschmeyer, 1990). Bianco and Banarescu (1982) place this species as Schizopyge zarudny, the species name being a mis-spelling. Schizopyge Heckel, 1847 is regarded by these authors as the correct name for snow trouts without a suctorial disc on the chin (synonym Schizothoraichthys Misra, 1962 (see discussion above under the genus Schizothorax).

Barbus microlepis Keyserling, 1861 described from "Flüsschen bei Anardareh, zwischen Herat und Lasch" is a synonym and is pre-occupied by Barbus microlepis Bleeker, 1851. Oreinus anjac Fowler and Steinitz, 1956 from "Zabol, Eastern Iran" is also a synonym as suggested by Saadati (1977) although it may be a hybrid.

The holotype of Oreinus anjac (ANSP 71949) at 281 mm total length is stored in the Academy of Natural Sciences of Philadelphia (Böhlke, 1984).

This species is closely related to Schizothorax intermedius but is distinguished by much smaller paired fins, longer and narrower branchial isthmus, and the scales slightly enlarged at the base of all fins, especially the dorsal and anal (Annandale and Hora, 1920).

Key characters

The only common Schizothorax species in Sistan, is is recognised by the large barbels and enlarged scales around the anal fin.

Morphology

Dorsal fin with 4 unbranched and 7-8 branched rays, anal fin with 3 unbranched and 5 branched rays. The last unbranched dorsal fin ray is moderately strong and serrated, the serrations being proportionately longer and spinier in young. Pectoral fin branched rays 18 and pelvic fin branched rays 9. Lateral line scales 97-114, at least 190 in the scale series immediately above the lateral line. The breast is naked or sparsely scaled. There is a pelvic axillary process. The scale sheath around the anal papilla and anal fin extends about one third to half way between the anal fin origin and the pelvic fin base. Scales are very small, horizontally ovoid and have an almost central focus. Scales are obliquely inserted in the scale pockets on the mid-flank above the lateral line and below the dorsal fin. Scales on the nape are none to minimally imnbricate. Radii are found on all fields and are numerous. Gill rakers on the lower arm 30-39 (Nikol'skii's (1897) count of 25 is incorrect (Berg, 1949); but see below for wider variation). Gill rakers are long, reaching the third to the sixth adjacent raker when appressed. The interior margin of each raker is serrated. Pharyngeal teeth usually 2,3,5-5,3,2, spoon-shaped with a slightly hooked tip. Anterior teeth are more rounded and thicker. There are 2 pairs of barbels, the anterior ones long to rudimentary in literature sources. The barbels are subequal in length, the anterior ones not reaching the eye and the poterior ones not reaching beyond the eye. The mouth is usually slightly subterminal but can be terminal or have the lower jaw projecting slightly. The gut is elongate and coiled. The chromosome number is 2n=96, NF=142, comprising 9 pairs of metacentric, 14 pairs of submetacentric and 25 pairs of acrotelocentric chromosomes, and the fish is a tetraploid (Hosseini and Kalbasi, 2003; Kalbassi et al., 2008).

Meristics for Iranian specimens: dorsal fin branched rays 7(2) or 8(34); anal fin branched rays 5(35); pectoral fin branched rays 16(1), 17(7), 18(10) or 19(16); pelvic fin branched rays 9(30) or 10(5); lateral line scales 93(1), 97(1), 98(2), 99(3), 100(4), 101(5), 103(4), 105(2), 106(5), 107(2), 108(1), 109(2), 113(1) or 114(1); total gill rakers 24(1), 25(1), 26(1), 27(1), 28(1), 29(1), 31(5), 34(6), 35(2), 36(8), 37(1), 38(2), 40(1) or 41(1); pharyngeal teeth 2,3,5-5,3,2(14), 2,3,5-5,3,3 (1), 2,3,5-5,2,1(1), 2,3,5-5,3,1(1), 2,2,5-5,3,2 (1), 2,2,5-4,2,2(1), 2,3,4-5,3,2(2), 2,3,4-4,3,2(1) or 1,2,5-5,3,2(1); and total vertebrae 47(11), 48(11) or 49(4).

Sexual dimorphism

Males develop prominent nuptial tubercles on the snout and on the scales. Females have a soft and distended belly during the breeding season (CIRSPE, 2006b).

Colour

Overall colour is silvery, the back and head darker with indistinct fine dots. The flanks may be spotted with black and some small areas may be more lightly pigmented and appear as indistinct spots or blotches. There are melanophores on the fin rays and membranes. Adult males may have reddish fins and dull red specks on the dorsal surface. Young are more silvery than adults. Generally there is no distinctive pattern on the body and fins.

Colour varies with the environment. In muddy water, the back and fins are pale olive-green, the flanks tinged with green or pale yellow and the belly pure white while in the yellow water of the reed beds the back and flanks are much darker, almost black, and even the belly is darkish. The peritoneum is brown to black.

Size

Reaches 62.1 cm total length and and over 2.2 kg. Ahmadi and Wossughi (1988) give average weights of 300 to 2000 g in commercial catches while fish more than 12 kg are reported recently (Iranian Fisheries Research Organization Newsletter, 30-31:5, 2002).

Distribution

This species is restricted to the Sistan basin including the Chahnimeh (Nikol'skii, 1897; Bianco and Banarescu, 1982; J. Holčík, in litt., 1996. Ghanbari and Jami (2011a) report it to be present only in the Chahnimeh Reservoir as the hamun dried out; presumably it recolonises the hamuns from rivers and reservoirs in wetter seasons.

Zoogeography

See under the genus description.

Habitat

Found in the open lake, in reed beds and in pools in Sistan (Annandale, 1921). It is the only species in Sistan common in the open lake in winter. Young probably make their way up upstream in the flood season as only adults are found in the lake in winter. The species is extremely abundant in pools left in stream beds when the floods recede. Spawning may occur in rivers as fry have not been found in the lakes (Iranian Fisheries Research Organization Newsletter, 30-31:5, 2002). Zabihi (2006) characterises it as a potamodromous species and notes that in March and April, if there is no flow in the rivers and thus no migration from the lake is possible, female gonads are reabsorbed. Ghanbari and Jami (2011a) note that mature fish migrate during April-May from rivers and lakes to cold and well-oxygenated streams for spawning, and migrate back leaving the young behind .

Age and growth

Sexual maturity may only be attained after 4 years. Zabihi (2006) examined 697 specimens with a length and weight range of 24.5-62.1 cm and 137-2204 g. Half the male fishes were mature at 29-31 cm and for females at 38-40 cm. Males mature a month earlier than females (Ghanbari and Jami, 2011a)

Food

The diet comprises almost exclusively small fishes.

Reproduction

Eggs in fish caught in spring by me were developing but very small. Specimens with mature, yellow eggs have been caught in December. However this species is mature in Esfand (20 February-20 March), incubation is 6-7 days at 16.20-17.75ºC, maximum egg diameter is 3.8 mm when washed and the yolk sac is absorbed at 6-7 days (M. Abedi, Islamic Azad University of Savad Kooh, abstract). Zabihy et al. (2004) and Zabihi (2006) found maximum oocyte size in March and April at 14-18ºC when the gonadosomatic index was highest at 7.9-9.6. The mean absolute and relative fecundities for fish 460-1380 g was 26,256 and 34,418 eggs respectively. The species was a total spawner showing a synchronous ovary. Eggs are adhesive to prevent them being washed away by strong currents (CIRSPE, 2006b).

Parasites and predators

Datta (1937) describes the male of the acanthocephalan Eosentis rigidus from the intestine of this species. Barzegar et al. (2008) record the digenean eye parasite Diplostomum spathaceum from this fish. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Lernaea sp. on this species.

Economic importance

Annandale in Annandale and Hora (1920) described the fishery for this species in the early years of the 20th century. Annandale commented that the flesh tasted like trout but was bony. The bones can be softened by cooking in vinegar. This fish is still caught and eaten and appears in local markets. Ahmadi and Wossughi (1988) cite an annual catch of 300-500 t while Iran Daily (24 August 2006) gives 700-1000 t before the drought. Rahimabadi et al. (2009) assessed the lipid quality of this species and found it to have a higher nutritional value than Schizocypris altidorsalis.

Annandale and his assistants became sick from eating the eggs of this species but he maintained that the local fishermen ate it without any deleterious effects. Fish were caught in a net about 4 feet (1.22 m) deep and 100 feet (30.5 m) or more long anchored at each end by a tamarisk stick stuck into the lake bottom and with the bottom of the net on the lake bed and the top of the net slightly above the surface. The net was positioned in relation to the wind, and therefore the prevailing currents, so it formed a semi-circle. The net was arranged in a gap in the reed beds or just outside the reeds in the open lake if the weather was exceptionally calm. Pools in the reeds were kept open to facilitate the fishing. The fishermen riding their tutins (reed boats) would drive the fish into the net by beating the water with poles and ululating. The two ends of the net were then lifted out of the water by the men in the two end boats such that the net formed a bag. The net was drawn into the two boats as rapidly as possible. A similar but shorter net was used to drag small channels while the men using it waded. A small-meshed bag net attached to one horizontal and two upright poles was also used in pools of dry stream-beds. The net was dragged by ropes, the men wading through the water. Some large fish were killed in the flooded swamps by striking at them with swords. Another net consisted of a bag about 7 feet (2.14 m) long and 6 feet (1.83 m) by 2 feet (0.61 m) at the mouth. The mouth was held open by poles tied together at one end to make a fork. The fork pivoted on a post on the bank. The mouth of the net had fine lines across it, the lead string of which was held by the fishermen to warn him that a fish had entered the net so he could pull the net out of the water to retrieve the fish. The net was placed along a bank where the current swirled forming a backwater, at the mouth of a small canal, or as the focus of a line of stakes blocking a channel. The season for this type of net began as early as August or not until October. It lasted several weeks. March and April could also be a favourable time if the river was not very full but the fish did not move as actively. The large fish caught were attracted to feed on the numerous small fish which migrated up river along the shore and were checked where backwaters met the main flow.

A more recent description of fishing in Sistan is found in Fowler and Steinitz (1956). Fishing takes place in fall on rivers and in the lake, preferably the early evening or morning when it is cool. River fishing is preferred to lake fishing and some fishermen never go out onto the lake. Lake fishing using boats may last up to 3-5 days at a time. River fishing is carried out with a cotton-thread seine weighted by stones at the bottom and with gourds as floats. The seine is tied to sticks at each end and the sticks to ropes leading to shore. Four to ten men can own a seine and the catch is divided among those fishing that day. Fishing is done in teams of 4-7 men or often in two teams with two nets. Half the men are on one side of the river and half on the other, pulling each other's nets in and out of the water. Only large, fat fish are kept, the others being thrown back. One net catches 30-40 fish in a day which are then sold in Zabol. Women never fish nor are present during fishing as they bring bad luck. Fish are always cooked before eating as eating raw fish is reputedly fatal.

Conservation

Ahmadi and Wossughi (1988) state that introductions of various fishes such as Cyprinus carpio, Carassius auratus, Ctenopharyngodon idella and Hypophthalmichtys molitrix are a possible source of competition for native species such as this schizothoracine since they are voracious, take spawning sites and carry diseases and parasites. Native catches have decreased in favour of introduced species. M. Abedi, Islamic Azad University of Savad Kooh, has studied artificial reproduction in this species including larval development. CIRSPE (2006b) and Iran Daily (24 August 2006) give details on artificial reproduction of this species. Males over 600 g and females over 900 g were found to be suitable for breeding, ideal water temperature range was 18-22ºC, fecundity range was 24,300-37,640 eggs for fish 0.8-1.1 kg, egg size was 1.54 mm (presumably not water hardened; see above), and survival from egg to 11 mm fingerling was about 10%.

Ghanbari and Jami (2011a) class this species as endangered.

Further work

The molecular relationships of this species to other Schizothorax species and related genera could be revealing.

Sources

Type material: See above, Aspiostoma zarudnyi (ZISP 11195a).

Iranian material:- CMNFI 1979-0072, 1, 199.1 mm standard length, Sistan, Hirmand River near Zabol (30º58'N, 61º28'E); CMNFI 1979-0223, 1, 34.4 mm standard length, Sistan, irrigation jube 1 km south of Lutak (30º45'N, 61º24'E); CMNFI 1979-0225, 11, 182.2-225.3 mm standard length, Sistan, effluent of Hirmand River (30º58'N, 61º28'E); CMNFI 1979-0226, 2, 146.8-167.2 mm standard length, Sistan, pool near Kuh-e Khajeh (30º57'N, 61º17'E); CMNFI 1979-0231, 2, 23.1-25.8 mm standard length, Sistan, irrigation jube 3 km from Zabol (31º01'N, 61º32'E); CMNFI 1979-0232, 2, 26.3-26.5 mm standard length, Sistan, irrigation jube 11 km from Zabol (ca. 30º58'30"N, ca. 61º36'E); CMNFI 1979-0235, 13, 148.4-193.7 mm standard length, Sistan, effluent of Hirmand (30º54'30"N, 61º41'E); CMNFI 1979-0237, 8, 24.4-45.3 mm standard length, Sistan, irrigation jube 18 km south of Zabol (30º53'N, 61º27'30"E); BM(NH) 1920.1.20:35, 1, 232.5 mm standard length, Sistan, near Lab-e Baring (ca. 31º07'N, ca. 61º12'E); ZMH 5902, 2, 372.3-369.7 mm standard length, Sistan, Hamun See (no other locality data); ZMH 5903, 3, 274.2-298.7 mm standard length, Sistan, Hamun See (no other locality data); ZMH 6088, 2, 48.3-136.1 mm standard length, Sistan, Rud Sistan (no other locality data).

Genus Squalius
Bonaparte, 1837

The members of this genus were formerly placed in the dace genus Leuciscus Cuvier, 1816. Some subgenera have been elevated to generic rank and vice versa. The genus Leuciscus is not monophyletic based on allozyme data for a limited number of European taxa (Hänfling and Brandl, 2000). There may be about 22 species in the genus Squalius with 3 species reported from Iran. It has been suggested that species formerly considered to belong to the subgenus Squalius should be simply regarded as part of a Leuciscus cephalus complex characterised by serrated pharyngeal teeth in 2 rows (2,5-5,2) and an almost straight or convex anal fin margin. This complex would include L. cephalus, L. gaderanus (= L. ulanus) and L. lepidus among Iranian species (Bogutskaya, 1994). Bogutskaya (2002), however, places L. persidis and L. ulanus in a new genus Petroleuciscus, q.v. and places L. cephalus and L. lepidus in the genus Squalius, and this is followed here.

Bogutskaya (2002) gives the characters of Squalius as numerous total vertebrae (commonly more than 40, up to 48); increased number of sensory cephalic pores (up to 12-20 in the supraorbital canal) in most species; often fused and very expanded fourth and fifth infraorbitals; and depressed neurocranium with a reduced interorbital septum. Other characters are a somewhat compressed body, moderate to large scales, a complete lateral line, no barbels, mouth terminal or subterminal, no notch in the upper jaw accommodating a tubercle on the lower jaw, thin lips with the lower one interrupted medially, a short dorsal fin without a thickened ray, a moderately long anal fin, long and hooked pharyngeal teeth in 2 rows (2,5-4,2, 2,5-5,2 or 3,5-5,3 modally) usually with hooked tips and spoon-shaped crowns, short gut, no keel on the belly, and short and relatively few gill rakers.

Rückert-Ülkümen (2000) and Rückert-Ülkümen and Matzke-Karasz (2000) document Lower Miocene fossils of Leuciscus, presumably Squalius, from western Turkey.

Squalius cephalus
(Linnaeus, 1758)

Common names

mahi safid (= white fish) in the Caspian basin and in central Iran; mahi-ye safid rudkhanehi or mahi-e-sephid-e-roodkhaneie (= literally white river, presumably in the sense of white river fish) in Khuzestan; kuli; 'rus mahi or aroos mahi.

[enlibas or gafgaz enlibasi, nour enlibasi for natio kaznakovi, all in Azerbaijan; tepug in Armenia; bir-aan siphaloos, baeaan, barayan, or berak (= breast perhaps in allusion to the broad and fleshy chin (Heckel, 1843b)) at Aleppo, all in Arabic; Kavkazskii golavl' or Caucasian chub in Russian; European chub].

Systematics

Cyprinus Cephalus was originally described from southern Europe.

Leuciscus orientalis Nordmann, 1840 described from Abkhazia, Georgia, Squalius Berak Heckel, 1843 described from "Aleppo" (earlier in the same work - p. 1041 - spelled Berag without a description, presumably an error), Squalius cephalopsis Heckel, 1843 described from "Aleppo", Squalius orientalis Heckel, 1849 described from "Flusse Kueik bei Aleppo", Squalius turcicus De Filippi, 1865 described from "Dell' Arasse presso Erzerum" (Aras River at Erzerum in Turkey), and Squalius agdamicus Kamenskii, 1901 from Agdam in the Kura River basin, are synonyms.

Several natio and varieties within what is now Squalius cephalus have been described from Iran or contiguous drainages and are listed as follows. They have no taxonomic validity but the names may reoccur. Leuciscus cephalus orientalis natio kaznakovi Berg, 1912 was described from Lake Nour near Vandam, Nukha District, Azerbaijan (in the Tur'yan-chai basin of the Kura River basin but not connected to it). Squalius turcicus var. platycephala Kamenskii, 1897 was described from Lake Taparavani (= Lake Paravani at 41°26'N, 43°48'E) and the Kyrchbulach River in the upper Kura River basin, Georgia. Leuciscus cephalus orientalis natio aralychensis Barach, 1934 was described in Latin from "Turcia, Aralych. fl. Kara-su", and in Russian "Reka Kara-su u vblizi Aralykha podnoshchiya Ararata" (= Kara-su River at the foot of Ararat in the vicinity of Aralykha). Leuciscus cephalus orientalis natio zangicus Barach, 1934 was described in Russian from "R. Zanga v Armenii" (i.e. at Erivan). Leuciscus cephalus orientalis natio ardebilicus Barach, 1934 was described in Latin from "Ardebil, fl. Balyk-tchai in systemate fl. Arax. inf." and in Russian "R. Balyk-chaya, vblizi Ardebilya v Persii" (i.e. Balyk River or Balyk-chai) in the upper Aras River basin of Iran.

Iranian populations were usually recognised as Leusciscus cephalus orientalis (Nordmann, 1840). Bianco and Banarescu (1982) correctly point out that there has been no critical revision of the subspecies and that differences are slight. Recognition of subspecies is disputable according to Reshetnikov et al. (1997).

Turan et al. (2007) examined morphological variation in Turkish populations from the Black, Caspian, Aegean, Mediterranean and Tigris-Euphrates basins and found three groups, but no discrete taxa.

The nominal Iranian subspecies, L. c. orientalis, differs from the European subspecies, L. c. cephalus, by having a more elongate body, a dark stripe behind the operculum and on average fewer scales and anal fin branched rays (Berg, 1948-1949; 1949).

The syntypes of Cyprinus cephalus are in the Naturhistoriska Riksmuseet, Stockholm under LP 81 (1 fish) and in the Zoologiska Museet, Uppsala Universitet, Uppsala under ZMUU Linnaeus Collection 213 (1) (Eschmeyer et al., 1996).

Three syntypes of Squalius orientalis, 111-146 mm standard length from Aleppo are in the Naturhistorisches Museum Wien under NMW 49438 (Krupp, 1985c). Two more fish from Aleppo are under NMW 49440 and measure 130.0-168.3 mm standard length.

Six syntypes of Squalius berak are in the Naturhistorisches Museum Wien under NMW 48915 and 3 syntypes are in the Senckenberg Museum Frankfurt under SMF 469, formerly in NMW, and 3 syntypes of Squalius cephalopsis are under NMW 49438 and 2 other syntypes are under NMW 49440 (Eschmeyer et al., 1996). The Vienna catalogue lists 6 specimens as Squalius berak and 2 specimens as Squalius cephalopsis but the Vienna card index in 1997 agrees with Eschmeyer et al. (1996) for the NMW specimens.

Natio ardebilicus syntypes are in the Georgian State Museum, Zoological Section, Tbilisi under ZMT 136h, natio aralychensis syntypes are under ZMT 22-11 (10), and natio kaznakovi syntypes are in the Zoological Institute, St. Petersburg under ZISP 12089 (5) (Eschmeyer et al., 1996).

A hybrid between Squalius cephalus and Chalcalburnus (= Alburnus) chalcoides is reported from Turkey (Ünver and Erk'akan, 2005).

Key characters

The relatively large scales outlined by pigment give this fish a distinctive appearance and, combined with large body size, fin ray counts, rounded belly and the absence of barbels, separates it from other Iranian cyprinids.

Morphology

Dorsal fin branched rays 7-9, usually 8, after 2-3, usually 3, unbranched rays, anal fin branched rays 7-10, usually 8 after 3 unbranched rays, pectoral fin branched rays 14-19, and pelvic fin branched rays 6-9, usually 8. Lateral line scales 38-48. Scales have few to moderate numbers of radii on the anterior and posterior fields. The focus is central to subcentral anterior and circuli are fine. On the posterior field, circuli break up into "bubbles" and are coarser than on other fields. The anterior scale margin is wavy, sometimes irregular and others with a small central protuberance and indentations above and below. Gill rakers short, 7-12, touching the adjacent raker when appressed. Vertebrae 40-46. Pharyngeal teeth 2,5-5,2, with variants 2,5-4,2, 2,4-4,2, 2,5-5,1, 1,5-5,2, 1,5-5,1, 2,5-5,3, 2,6-5,2, or even 1,5-5,1,2 and 1,2,5-5,2,2. Teeth are very narrow, strongly hooked at the tip and strongly serrated so that there is no obvious flat surface. The serrations on the teeth of orientalis are stronger than in the typical subspecies. The gut is an elongate s-shape. Chromosome number is 2n=50 (Al-Sabti, 1986; Klinkhardt et al., 1995).

Meristics for Iranian specimens: dorsal fin branched rays 7(3), 8(103) or 9(1); anal fin branched rays 7(3), 8(60) or 9(44); pectoral fin branched rays 14(3), 15(13), 16(39), 17(46), 18(5) or 19(1); pelvic fin branched rays 6(1), 7(1), 6(100) or 9(5); lateral line scales 40(2), 41(20), 42(45), 43(30), 44(9) or 46(1); total gill rakers 7(2), 8(14), 9(58), 10(31) or 11(2); pharyngeal teeth ?; and total vertebrae 40(6), 41(29), 42(45) or 43(6).

Sexual dimorphism

Abdurakhmanov (1962) reports that head length, eye diameter, caudal peduncle length, dorsal and anal fin heights, pectoral and pelvic fin lengths and lower caudal fin lobe length are all greater in males while postorbital length, interorbital width, head depth and pectoral-pelvic fin distance are greater in females.

Colour

Overall colour is silvery to grey. Scales of the lateral line and upper back have a strong dark pigmentation along the posterior margin and a distinct dark spot anteriorly. Scales may have some gold in their centre. The back is dark brown or reddish-brown to blue-grey and the belly and lower head are pearly-white to a silvery yellow. The operculum is a strong copper-yellow colour and the opercular opening is dark, nearly black. The iris is silvery and has very little gold, or is golden with a lot of dark grey pigment. There may be an upper spot of dark grey on the iris. The dorsal and anal fins are grey and may have some pink. The dorsal fin is blackish distally. The pectoral fin has a little pale grey pigment and there may be a yellow or yellow-pink spot at its upper base with this colour extending onto the first 2-3 rays. The pelvic and anal fins are pink to red-pink with somewhat colourless posterior margins, especially in spawning males. The caudal fin is a pale pink to dirty reddish with some dark pigmentation to the posterior margin.

Size

Possibly as long as 80 cm although 45 cm is a more likely maximum (Berg, 1948-1949) and a weight of 6-8 kg. Reaches 43 cm total length in the Madar Su of Golestan National Park (A. Abdoli, pers. comm., 1995). Reaches 39.2 cm and 870 g in Taham Dam in northern Iran (Iranian Fisheries Research Organization Newsletter, 54 & 55:4, 2008). Attains 39.0 cm fork length in Turkey (Yerli et al., 1999). Maximum size is possibly 85.0 cm and 10.0 kg.

Distribution

Found from the British Isles and the Iberian Peninsula eastwards in its southern distribution to Turkey and Iraq, the northern half of Iran, and the whole Caspian and Aral seas drainages. In Iran, it is recorded from rivers along the Caspian Sea coast including the Anzali Mordab (Holčík and Oláh, 1992; Roshan Tabari, 1997; Shamsi et al., 1997; Abbasi et al., 1999; Kiabi et al., 1999) but also the Aras River, the Namak Lake basin (Wossughi, 1978; Bianco and Banarescu, 1982), the Lake Orumiyeh basin (Günther, 1899); Lake Zaribar, Kordestan (Abzeeyan, 5(5):III, 1994), and the Esfahan basin and the Tigris River basin (Berg, 1949). Abdoli (2000) and Abdoli and Naderi (2009) map this species from the middle Atrak and Gorgan, middle and upper Neka, middle and lower Babol and Heraz, Chalus, Tajan, Tonekabon, Sardab, Pole-e Rud, Safid, Shahrud and Qezel Owzan, middle Aras and its tributary Qareh Chai, all in the the Caspian Sea basin, the middle and lower Talkheh, Zarrineh and Tatavi in the Lake Orumiyeh basin, the Karaj, Shur, Abhar, Qareh Chai and Qom rivers in the Namak Lake basin, and the Marun, upper Karun and middle Khersan, Dez, middle and upper Karkheh, Simarreh, Kashkan and lower Gav Masiab rivers in the Tigris River basin.

Zoogeography

This widely distributed species has been split into a number of subspecies but the principal one from the Middle East or Southwest Asia is recognised as S. c. orientalis. This has not been examined in detail recently using classical techniques and the relationships between isolated populations are unknown. Durand et al. (1999), working on the cytochome b of European populations, found that a lineage from a Ponto-Caspian refugium recolonised the Baltic area in the Holocene after glaciation. Durand et al. (2000), again examining cytochrome b, consider that S. cephalus may have originated from Mesopotamia and, in the late Pliocene, used the large inland lake of Anatolia existing at that time for dispersion. Uplift of the Anatolian Plateau, climatic changes and river isolation was probably the main vicariant event leading to a quick radiation in these chubs.

Habitat

In the Caspian Sea basin, it is found mainly in upstream waters (Jolodar and Abdoli, 2004).

Age and growth

Öztaş (1988; 1989), Öztaş and Solak (1988) and Türkmen et al. (1999) studied the chub in the Aras River of Turkey and found its condition factor to be higher in summer and autumn and to vary between age groups. Life span there is over 8 years, compared with 7 years in Lake Aksehir, Anatolia (Altindag, 1996) and in the Savur stream in the Tigris River basin of southeast Anatolia (Ünlü and Balcı, 1993a; 1993b), over 8 years in the Euphrates River in Turkey (Özdemir and Şen, 1986), and 10 years in Sariyar Dam Lake near Ankara (Ekmekci, 1996a). In the Aras, females grow faster than males and there are more sexually mature females in the older age groups. Sexual maturity is attained at age 2-3 in males and ages 3-4 for females. Mean condition factor for Aras males was 1.326 and for females 1.333. In the Savur stream in the Tigris River basin, 80% of females and 75% of males reaching sexual maturity at their third (14.6 cm fork length) and second years (13.0 cm) respectively. Females grow larger than males and live about 2 years longer as they do in Lake Aksehir. Sexual maturity is attained at ages 3-4 in Sariyar Dam Lake and these fish at age 1 have a mean fork length of 90 mm, mean weight of 9.4 g and a condition factor of 1.263 which by age 10 can reach 370 mm, 910 g and 1.805. Aksehir Lake fish reached 44 cm forked length and 1766 g for females and 31 cm and 557 g for males. In Lake Tödürge, Turkey, Ünver (1998) found that males reached maturity in their second or third year of life while females were in their third or fourth year. Over 68% of the catch was female and life span attained 7 years. Yerli et al. (1999) invetsigated growth in Lake Çıldır, Turkey and also found 8 age groups, dominated by those in age group 3 at 67.3% and by males at 73% and with maximum fork length at 390 mm and maximum weight at 720 g. Maximum life span is up to 15 years. The weight class 150-450 g dominated in Taham Dam in Iran (Iranian Fisheries Research Organization Newsletter, 54 & 55:4, 2008).

Food

Food items include mayfly and caddisfly larvae, other small organisms such as molluscs, and crayfishes, small fishes and frogs. Large fish feed mainly on other fish. Reputedly even fruit fallen in the water will be eaten. Trout eggs and fry are also eaten. Guts of Iranian specimens contained a wide variety of organisms including ants (presumably taken at the water surface), aquatic insects such as chironomids among others, crustaceans, filamentous algae, higher plant fragments, scales of cyprinids and the remains of a Paracobitis malapterura. Abdoli (2000) reports Ephemeroptera, Chironomidae and Trichoptera. In Taham Dam in Iran algae, higher plants, bivalve and insects were found (Iranian Fisheries Research Organization Newsletter, 54 & 55:4, 2008).

Reproduction

Öztaş (1989) examined reproduction of this species in a stream tributary to the Aras River in Turkey. Spawning begins at the end of May although most fish spawn in June. Water temperatures at this time are 12-18°C. Fecundity is up to 61,808 eggs and maximum egg diameter is 1.39 mm. Türkmen et al . (1999) found spawning between May and July in the Aras River proper with fecundity up to 17,187 eggs. Ünlü and Balcı (1993a; 1993b) found spawning to take place from May to late June in southeast Anatolia in a tributary of the Tigris River. Fecundity reaches 20,140 eggs and egg diameter 1.5 mm. Equations for the relationship between fecundity (F) and length (FL), weight (W) and ovary weight (GW) were given as F = 0.0458 FL2.3680 (r = 0.745), F = 270.96 W0.7912 (r = 0.761) and F = 1888.86 GW0.8163 (r = 0.775). In Sariyar Dam Lake near Ankara, spawning takes place between April and June (Ekmekci, 1996a). Ünver (1998) found fish in Lake Tödürge, Turkey spawning between May and July with a mean egg diameter of 0.65 mm (highest 1.04 mm) and a mean fecundity of 14,500 eggs (highest 28,664 eggs).

Iraqi fish attained sexual maturity a in 3 years at 20 cm length and 700 g weight, spawning in March and April and depositing eggs in shallow water on gravel (Al-Rudainy, 2008). In Azerbaijan, Abdurakhmanov (1962) gives spawning temperatures as 12-21°C, maximum fecundity as 118,000 eggs and maximum egg diameter as 1.8 mm. Spawning in Iran appears to take place in spring judging from egg development.

Around 40% of the fish in Taham Dam in Iran spawned in early June and around 30% did not spawn at all (Iranian Fisheries Research Organization Newsletter, 54 & 55:4, 2008).

Parasites and predators

Ergens and Gusev (1965) report the monogenean helminth Dactylogyrus prostae in this species from Bandar-e Shah (= Bandar-e Torkeman) on the Caspian Sea coast. The monogeneans Diplozoon paradoxum and D. megan are recorded from this species in the Tajan River, Mazandaran (Iranian Fisheries Research and Training Organization Newsletter, 6:7, 1994). Shamsi et al. (1997) report Clinostomum complanatum, a parasite causing laryngo-pharyngitis in humans, from this species. Mirhasheminasab and Pazooki (2003) list Ergasilus peregrinus, Tracheliastes polycolpus and Lernaea cyprinacea from this species in Mahabad Reservoir, the latter being the most dangerous parasite. Jalali et al. (2005) summarise the occurrence of Gyrodactylus species in Iran and record G. sp. from fish in the Safid River. Masoumian and Pazooki (1998) surveyed myxosporeans in this species in Gilan and Mazandaran provinces, finding Myxobolus minutus and M. muelleri. Pazooki et al. (2005) record Lamprolegna compacta, Ergasilus peregrinus and Lernaea cyprinacea from this species in waterbodies of Zanjan Province. Miar et al. (2008) examined fish in Valasht Lake and the Chalus River, Mazandaran and found the protozoan Chilodonella hexastica. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Ergasilus peregrinus, Ergasilus sp., Lamproglena compacta, Lernaea sp., Tracheliastes longicollis and Tracheliastes polycolpus on this species.

Economic importance

Listed as economically important in the Turkish Euphrates River (Özdemir and Şen, 1986).

Conservation

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in textbooks and for sport. Kiabi et al. (1999) consider this species to be of least concern in the south Caspian Sea basin according to IUCN criteria. Criteria include sport fishing, medium numbers, habitat destruction, widespread range (75% of water bodies), present in other water bodies in Iran, and present outside the Caspian Sea basin.

Ekmekci (1996a) recommends a minimum fishing size of 28 cm and a prohibition of fishing during the April-June spawning season for this Anatolian population.

Further work

The distribution and biology of this species in Iran needs study, as does the systematics of isolated taxa.

Sources

Iranian material: CMNFI 1970-0506, 6, 42.5-116.3 mm standard length, Gilan, Shalman River (37º08'N, 50º15'E); CMNFI 1970-0536, 3, 89.0-223.0 mm standard length, Gilan, Siah River estuary near Rudbar (36º53'N, 49º32'E); CMNFI 1970-0538, 7, 40.8-137.2 mm standard length, Gilan, Qezel Owzan River (36º44'N, 49º24'E); CMNFI 1970-0583, 6, 41.4-123.8 mm standard length, Gilan, Nahang Roga River (37º28'N, 49º28'E); CMNFI 1970-0589, 1, 142.0 mm standard length, Gilan, Safid River (37º12'N, 49º54'E); CMNFI 1979-0253, 33, 46.4-100.4 mm standard length, Markazi, Qareh Su River drainage (34º52'N, 50º49'E); CMNFI 1979-0255, 2, 90.7-90.8 mm standard length, Markazi, Bar River drainage (33º51'30"N, 50º23'E); CMNFI 1979-0276, 1, 59.7 mm standard length, Lorestan, Chamesk River (ca. 33º19'N, ca. 47º53'30"E); CMNFI 1979-0462, 12, 50.7-104.5 mm standard length, Markazi, Mazdaqan River (35º06'30"N, 49º40'30"E); CMNFI 1979-0469, 6, 86.0-127.5 mm standard length, Mazandaran, river west of Alamdeh (36º37'30"N, 51º35'E); CMNFI 1979-0474, 6, 71.3-80.6 mm standard length, Mazandaran, Tajan River (36º34'N, 53º05'E); CMNFI 1979-0482, 2, 112.2-160.1 mm standard length, Mazandaran, river between Minudasht and Dowlatabad (37º19'30"N, 55º31'E); CMNFI 1979-0493, 7, 85.2-114.5 mm standard length, Mazandaran, Tajan River drainage (36º19'N, 53º23'E); CMNFI 1979-0494, 12, 9.1-92.6 mm standard length, Mazandaran, Talar River tributray (36º21'N, 52º51'30"E); CMNFI 1980-0120, 2, 41.8-118.1 mm standard length, Mazandaran, Babol River at Babol Sar (36º43'N, 52º39'E); CMNFI 2007-0092, 5, 25.5-45.9 mm standard length, Azarbayjan-e Khavari, Zilber Chay (38º42'N, 45º16'E); CMNFI 2007-0100, 4, 41.2-171.8 mm standard length, Azarbayjan-e Bakhtari, Kalwi Chay near Piranshahr (ca. 36º44'N, ca. 45º10'E); CMNFI 2007-0101, 1, 162.0 mm standard length, Azarbayjan-e Bahktari, Tata'u River (ca. 36º54'N, ca. 46º07'E); CMNFI 2007-0105, 1, 96.9 mm standard length, Kordestan, Zarineh River basin (ca. 36º06'N, ca. 46º20'E); CMNFI 2007-0109, 1, 101.1mm standard length, Kordestan, Qeshlaq River basin (ca. 35º16'N, ca. 47º01'E); CMNFI 2007-0113, 1, 99.5 mm standard length, Kermanshahan, Qareh Su River tributary (ca. 34º25'N, ca. 47º01'E).

Comparative material: BM(NH) 1974.2.22:71-72, 2, 68.0-93.1 mm standard length, Iraq, Baghdad (33º21'N, 44º25'E); BM(NH) 1974.2.22:74, 1, 152.1 mm standard length, Iraq, Greater Zab near Aski Kalak (ca. 36º16'N, 43º39'E); BM(NH) 1974.2.22:75, 1, 154.0 mm standard length, Iraq, Qizilja River, Lesser Zab (no other locality data).

in Leuciscus?

Squalius latus
Keyserling, 1861

Flank scale

Common names

None.

[Zakaspiiskii elets or Transcaspian dace in Russian; Murgab dace].

Systematics

Squalius latus was described from the "Fluss. Heri-Rud bei Herat" (now in Afghanistan, then in Persia; Herat is at °'N, °'E).

Squalius transcaspiensis Berg, 1898 from "Habitat in flum. Tedschent, prope Aschabat in provincia Transcaspica" is a synonym. The type locality is presumably the Tedzhen River in Turkmenistan (not Iran as given in Eschmeyer et al. (1996)) although Ashkhabad is not on the Tedzhen River. Many syntypes are in the Zoological Museum of Moscow State University (MMSU) according to Eschmeyer et al. (1996). Possibly a subspecies of Squalius lehmanni (Brandt, 1852) according to Nikol'skii (1938) and Svetovidova (1967) or of Leuciscus leuciscus (Linnaeus, 1758) according to V. V. Kafanova (cited in Bogutskaya, 1994). Bogutskaya (1994) considers S. latus to be distinct.

Key characters

Dorsal and anal fin ray counts, both 7-9, lateral line scale counts under 48, and distribution identify this species in northeastern Iran.

Morphology

Dorsal fin branched rays 7-9, after 3 unbranched rays, anal fin branched rays 7-9 after 3 unbranched rays, pectoral fin branched rays 14-16, and pelvic fin branched rays 7-8. Pharyngeal teeth number 2,5-5,2; 2,5-5,1; 1,5-5,1; 3,4-4,2; 3,5-5,2; or 2,5-4,2. In specimens from the Tedzhen (= Hari) River, part of which is the Iran-Afghanistan border:- dorsal fin branched rays 7(12), anal fin branched rays 7(1), 8(6) or 9(5), pectoral fin branched rays 15(6) or 16(6), and pelvic fin branched rays 7(1) or 8(11). Lateral line scales 39(1), 40(1), 41(2), 42(2), 43(1), 44(2), 45(1), 46(1), or 47(1). A pelvic axillary scale is present. Scales bear a few anterior and posterior radii. Total gill rakers number 8(2), 9(1), 10(5) or 11(4) and reach the raker below when appressed. Total vertebrae 39(4), 40(6) or 41(3) (includes data from x-rays of ZISP 18361 and 11047). Pharyngeal teeth number 2,5-5,2(3) and have narrow cutting edges on each side of a shallow groove below a hooked tip and in larger fish some teeth are serrated below the tip. The mouth is oblique and extends back to the level of the nostrils, the lower jaw protruding slightly.

Sexual dimorphism

Unknown.

Colour

The back and upper flank are reddish-brown, the flanks yellowish and the belly silver. The dorsal and caudal fins are yellowish-brown and the other fins are reddish. The base of all fins becomes bright orange during spawning.

Size

Reaches 26.7 cm.

Distribution

This species is found in the Tedzhen River (= Harirud in Iran) and the Murgab River to the east in Afghanistan and Turkmenistan.

Zoogeography

This species is part of a complex of species in the genus Squalius and the related Leuciscus found from western Europe to Siberia. Its origins may lie in a Ponto-Caspian-Aralian refugium, becoming isolated and speciating in the eastern part of this area.

Habitat

Unknown in detail.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

None reported from Iran.

Economic importance

None.

Conservation

Numbers and habitat requirements are unknown and so a conservation assessment cannot be made.

Further work

The biology of this species needs study as well as it relationships to other Squalius.

Sources

Type material: There are no type specimens.

Iranian material: None.

Comparative material: ZISP 10357a, 1, 63.2 mm standard length, Turkmenistan or Afghanistan, Tedzhen River (no other locality data); ZISP 10359, 4, 93.4-130.5 mm standard length, Turkmenistan or Afghanistan, Tedzhen River (no other locality data); ZISP 10360, 6, 58.4-122.0 mm standard length, Turkmenistan or Afghanistan, Tedzhen River (no other locality data); ZISP 10419, 1, 107.6 mm standard length, Turkmenistan Iolotan near Merv (37º18'N, 62º21'E).

Squalius lepidus
Heckel, 1843

Common names

kavar or kawar.

[bara'an or bir-aan abiadh in Arabic; Tigris dace].

Systematics

Squalius lepidus was originally described from the "Tigris bei Mossul" (Heckel, 1843b).

Alburnus doriae De Filippi, 1865 and Alburnus maculatus Keyserling, 1861 are synonyms according to Coad (1982d; 1985h) but these are errors. Note also that Alburnus maculatus Kessler, 1859 from the Salghir River at Simferopol, Crimea, Ukraine preoccupies Alburnus maculatus Keyserling, 1861 and is a distinct species of Alburnoides (Bogutskaya and Coad, 2009).

The syntypes of Squalius lepidus are in the Naturhistorisches Museum Wien according to Krupp (1985c) under NMW 49342, 2 specimens, 229.6-245.5 mm standard length as measured by me and NMW 49343, 2, 88.0-107.8 mm standard length, collected by Th. Kotschy in 1843. Bogutskaya (1994), however, lists a lectotype (237.0 mm standard length) and 5 paralectotypes under NMW 49342, 2 paralectotypes under NMW 49343, and 1 paralectotype under SMF 847 (the paralectotypes measure 90-8-233.8 mm standard length). Eschmeyer et al. (1996) also lists 1 syntype under NMW 49344 and this specimen (164.8 mm standard length) appears in the 1997 Vienna card catalogue as a syntype along with the other 4 fish mentioned by Krupp (1985c). One syntype is in the Senckenberg Museum Frankfurt (SMF 847, formerly NMW) (F. Krupp, pers. comm., 1985). The catalogue in Vienna lists 6 fish in spirits and 1 fish stuffed.

Key characters

This species is distinguished from other Squalius by the elongate and pointed head with a projecting lower jaw, an adaptation for piscivory which develops early and is not evident in other Squalius even when these feed on fish as adults.

Morphology

Dorsal fin branched rays 8-10, usually 8 (Krupp, 1985c) or 9 (Bogutskaya, 1994)(8 in four syntypes, 9 in one), after 3 unbranched rays, anal fin branched rays 9-11, usually 9, after 3 unbranched rays, pectoral fin branched rays 14-18, and pelvic fin branched rays 8. Lateral line scales 42-50. Scales have few radii on the anterior and posterior fields. Gill rakers 7-12, reaching the one below when appressed (near junction of upper and lower arches) and sometimes curled. Pharyngeal teeth 2,5-5,2 or 2,5-4,2, strongly hooked and serrated. Total vertebrae 42-46. The projecting lower jaw fits into a notch in the upper jaw.

Sexual dimorphism

Unknown.

Colour

The back and upper flank are brown to bluish-brown or blackish, the flanks generally silvery, and the belly silvery-yellow. The upper flank, head and fins may be sprinkled with black spots. Flank scales have a spot at their base. All fins are reddish, with the pectoral and anal fins the brightest. The dorsal fin has a black margin. The caudal fin is blue-grey and its margin is blackish.

Size

Reaches 54.2 cm (Karabatak, 1997).

Distribution

Found in the Orontes (= Asi), Quwayq and Tigris-Euphrates basins as well as some lakes in central Anatolia (Beyşehir, Akşehir) and the rivers Ceyhan and Şeyhan in southern Anatolia.

Records from Iran are based on literature and include Lake Zaribar in Kordestan, Kermanshah, the Marakeh River, Luristan, and Dez River below Dez Dam (= formerly Mohammad Reza Shah Pahlavi Dam) (Wossughi, 1978). Also recorded from Mendeli on the Iran-Iraq border (Berg, 1949). Abdoli (2000) records this species from rivers of Khuzestan such as the Jarrahi, Marun, Karun, middle and lower Dez and middle Karkheh; and the Qom, Shur and Zayandeh rivers probably in error. A record from a canal near Gaz a few miles from Esfahan is probably an error (Keyserling, 1861).

Zoogeography

Durand et al. (2000) using cytochrome b suggest that this species is fully introgressed with L. cephalus mtDNA (= S. cephalus) and so question the taxonomic validity of this species. Morphological data contradicts this conclusion. Durand et al. (2000) conclude that their data does indicate that "L. lepidus and L. cephalus might have had different dispersion histories over the same geographical range" and that "L. lepidus introgression by the chub (L. cephalus) is ancient, explaining the complete sorting of the lepidus lineage".

Habitat

Generally unknown but found in rivers and lakes.

Age and growth

Karabatak (1997) studied fishes identified as this species from Lake Beyşehir in Turkey and recorded condition coefficients of 1.30-1.45 pre-spawning, 1.13-1.24 spawning and 1.24-130 post-spawning, with maximum values in the pre-spawning period coincident with maximum gonad development. Males and females showed similar length-weight relationships although females were slightly heavier because of their eggs. Çolak (1983) studied the age of this species in the Keban Dam, Turkey and found a maximum age of 8 years, that females grew faster and were longer than males, and growth was rapid until five years of age when it fell by almost half in each succeeding year. von Bertalanffy formulae, sexes combined, for two years were Lt = 87.92 (1-e-0.112(t+1.464)) and Lt = 65.64 (1-e-0.214(t+0.878)).

Food

Plant remains and fish scales have been found in gut contents. Fish appear to be the main diet item even in young from about 10 cm in length (Bogutskaya, 1994) although Al-Rudainy (2008) cites fish fry and aquatic insects for Iraqi fish, presumably smaller ones.

Reproduction

Karabatak (1997) studied fishes identified as this species from Lake Beyşehir in Turkey and found a peak spawning in mid to late June at 20.0-21.5°C with the highest gonadosomatic index on 15 May at 16.0-17.2°C. The spawning season could extend from late May to early July. Ünlü (2006) gives age at first maturity as 2-3 years in the Turkish Tigris River with spawning over sand, stone and gravel. Al-Rudainy (2008) cites sexual maturity at 3-4 years, 30 cm length and 3 kg weight in Iraq with spawning in March and April with eggs deposited in shallow water on gravel beds.

Parasites and predators

Williams et al. (1980) report the digenean helminth Allocreadium isoporum from this species in the Zayandeh River at Esfahan but the fish species was most probably misidentified. Barzegar et al. (2008) record the digenean eye parasite Diplostomum spathaceum from this fish.

Economic importance

Economically important in Turkey in Lake Beyşehir (Karabatak, 1997).

Conservation

The biology and distribution of this species in Iran need elucidation so a conservation assessment can be made.

Further work

The biology of this species needs study along with its systematic relationships with other Squalius species as morphology and molecular analyses are contradictory.

Sources

Some counts are taken from Bogutskaya (1994).

Type material: See above, Squalius lepidus (NMW 49342, NMW 49343 and NMW 49344).

Iranian material: None.

Comparative material: NMW 87786, 2, 329.0-381.5 mm standard length, Turkey, Karaviran at Konya (ca. 37º52'N, ca. 32º31'E); NMW 91149, 11, 69.6-115.9 mm standard length, Syria, Euphrates River at Jarabulus (36º49'N, 38º01'E); NMW 91620, 2, 91.6-95.6 mm standard length, Syria, Euphrates River at Jarabulus (36º49'N, 38º01'E); NMW 94441, 1, 175.7 mm standard length, Turkey, Palu, Murat River (38º42'N, 39º57'E).

Genus Tinca
Cuvier, 1816

The tench genus contains a single species found from Europe to Siberia including the Caspian Sea basin.

The characters of this genus include very small, elongate scales deeply embedded in slimy skin, pharyngeal teeth in a single row, all fins rounded and without spines, a short barbel at the terminal mouth corner, no keel on the belly, moderately long gill rakers, and short dorsal and anal fins.

Tinca tinca
(Linnaeus, 1758)

CMNFI 1979-0439, Gilan, Anzali Mordab

Common names

لاي ماهي (= lie, lai, lay or laay mahi, meaning bottom-dwelling or mud fish); nazi; tilkhos; saboni (= soapy?); hashtarkhan kapur.

[lil baligi in Azerbaijan; sew-zukgná in Armenia; lin' in Russian; tench, common tench, green tench].

Systematics

Cyprinus Tinca was described originally from European lakes. Ljabner et al. (2010) found the western and eastern phylogroups of tench were a single species under the biological species concept, although they were separate phylogenetic species. Tench from a western European and a Ponto-Caspian refugium came into contact by postglacial expansion after a separation of 750,000 years, and showed free interbreeding.

Key characters

The dark coloration, rounded fins and slimy body are characteristic of this species.

Morphology

Dorsal fin unbranched rays 2-4, usually 3-4 followed by 6-9, usually 8, branched rays, anal fin unbranched rays 3-4 and branched rays 5-9, usually 6-7, pectoral fin branched rays 13-18, and pelvic fin branched rays 7-10. Lateral line scales 70-120, gill rakers 10-16, usually 11-15, and vertebrae 35-44, usually 39-43. Anterior gill rakers may be difficult to distinguish from throat tubercles. Central rakers touch the one below when appresssed. Scales are very elongate ovals with a very anterior focus near the anterior scale margin. Radii are numerous on all fields. Pharyngeal teeth 4-5, 5-4, 5-3 or 4-4, more rarely 5-5, slightly to strongly hooked and expanded at the tip into a head supported on a narrow stalk. The oblique head has rounded edges and is concave in the middle. The gut is s-shaped with a small anterior loop. The chromosome number is 2n=48 (Klinkhardt et al., 1995).

Meristic values for Iranian specimens are:- dorsal fin branched rays 8(28); anal fin branched rays 5(1), 6(11), 7(15) or 8(1); pectoral fin branched rays 15(10), 16(13) or 17(5); pelvic fin branched rays 9(27) or 10(1); lateral line scales very embedded and obscured by skin and mucus, range about 82-95; total gill rakers 11(4), 12(7), 13(8), 14(7) or 15(2); pharyngeal teeth 5-4(5), 4-5(2) or 4-4(1); and total vertebrae 37(1), 38(7), 39(14) or 40(1).

Sexual dimorphism

The second pelvic fin ray in males is thickened and is accompanied by a muscular protuberance from the flank. Pelvic fins reach the anal fin in males.

Colour

The overall colour is a blackish-green, green or dark brown to bronze. Colour varies with habitat and is lighter in areas with less vegetation. The iris is red and the lips are yellowish-orange. Fins are grey to greenish or blackish. Golden, red and orange forms, some with black spots, may be bred in garden ponds and aquaria. Preserved fish have a dark body and fins with a lighter belly.

Size

Reaches 84.0 cm total length and about 8.5kg, possibly 10.0 kg.

Distribution

Found from the British Isles and the Iberian Peninsula across Europe to Siberia including southern drainages of the Black and Caspian seas. In Iran it is reported from the Anzali Mordab and Safid River, neighbouring rivers to the Gorgan River, and the Tajan and Babol rivers (Holčík and Oláh, 1992; Riazi, 1996; Abbasi et al., 1999; Kiabi et al., 1999; Abdoli, 2000; Daghigh Roohi and Mokhayer, no date; Abdoli and Naderi, 2009). Khara et al. (2005) record it from the Amirkelayeh Wetland near Lahijan.

Zoogeography

This widespread species has no close relatives among Eurasian cyprinids.

Habitat

Tench inhabit bays, small and shallow lakes, slow rivers and estuarine areas rich in vegetation. They are confined to the lower reaches of rivers in Iran and do not penetrate upstream (Berg, 1948-1949). Riazi (1996) reports that this species is native (resident) to the Siah-Keshim Protected Region of the Anzali Mordab. They are essentially inactive in winter. Tench are reported to bury themselves in mud during severe winters, lying dormant until spring. They can tolerate weak brackish water (to 12‰), acidic waters, low oxygen conditions and temperatures as high as 37°C for short periods. Preferred temperatures are 15-23.5°C (Brylińska, Bryliński and Bănărescu in Bănărescu, 1999). They remain alive for long periods when removed from the water. In Iran, adults have been caught in gill nets at 1-2 m (Annual Bulletin 1993-94, Iranian Fisheries Research and Training Organization, Tehran, p. 78-79, 1995).

Age and growth

Nezami Balouchi et al. (2004) studied of tench in the Amirkelayeh Lagoon and found age groups 1+ to 7+ years. Vetlugina (1992) studied the biology of tench in the Volga Delta using sections of dorsal fin rays for aging. Life span is 13 years or more. Females grow faster than males and comprise 90-100% of older fish. Growth rate is similar to fish from Dagestan, nearer Iranian waters. Tench begin to mature at 3 years (30-40% of fish are sexually mature) and by age 4-5, 60% are mature. In Dagestan they mature at 3-4 years (Shikhshabekov, 1977). Life span is up to 30 years elsewhere. Turkish populations in lakes formed by dams range up to 5 or 6 years of age (Altindağ et al., 1998; 2002; Benzer et al., 2009). These authors give age-length and age-weight relationships and condition factors. There is a substantial European literature on this fish.

Food

Diet comprises insect larvae, crustaceans, small molluscs, aquatic worms, vegetation and detritus. Chironomids may constitute as much as 91.8% of the diet in the second year of life while older fish favour snails and algae and macrophytes. This fish is probably a carnivore, switching to vegetation when animals are not available. Food may be picked out of vegetation or nosed out of mud as much as 13 cm deep using the snout (Brylińska, Bryliński and Bănărescu in Bănărescu, 1999). Nezami Balouchi et al. (2004) studied the diet of tench in the Amirkelayeh Lagoon and found 17 food groups including Odonata, snails, water plants, Trichoptera, Chironomidae, Hemiptera, Ephemeroptera, Perca fluviatilis, Diptera, Gammarus, Tubifex, plant seeds, Simulium, water bugs, water ticks, zooplankton and phytoplankton. Phytoplankton, snails and Hemiptera had the highest frequenceis at 68.5%, 65.7% and 34.0% respectively. This fish is an omnivore and diet varied with season age and sex.

Reproduction

Spawning occurs in shallow water with little current and abundant vegetation. Spawning aggregations can be seen in April in the Volga Delta and spawning takes place in May and June (Vetlugina, 1992). Tench in the Anzali Mordab have well-developed, 1.0 mm eggs in early June. Water temperatures at peak spawning are 20-26°C. In Dagestan, spawning takes place in June-July at water temperatures no lower than 19-20°C (Shikhshabekov, 1977). Each female is accompanied by 2-3 males and different males will fertilise the egg batches as they are released. Tench spawn 2 or 3 times at intervals of about 20-30 days, shedding eggs onto surface vegetation. Eggs are greenish, adhesive and up to 1.57 mm in diameter. Fecundity is up to 863,000 eggs (or 124,850 per kg of body weight) and increases with age, length and weight. Larvae have attachment organs which enable them to hang onto plants for the first few days of life. Maximum fecundity is 1,241,200 eggs in eastern Europe.

Parasites and predators

Daghigh Roohi and Mokhayer (no date) record the trematode Asymphylodora tincae from this fish in the Anzali Lagoon, the first record of this parasite in Iran. Khara et al. (2005) found parasites of this species in the Amirkelayeh Wetland were Raphidascaris acus, Camallanus lacustris, Asymphylodora tincae, Diplostomum spathaceum, Dactylogyrus sp., Caryophyllaeus fimbriceps, Lernaea sp. and Trichodina sp. Khara et al. (2006a) record the eye fluke Diplostomum spathaceum for this fish in the Amirkelayeh Wetland in Gilan, although it had the lowest abundance of 7 species examined. Sattari et al. (2004, 2005) surveyed this species in the Anzali and Amirkelayeh wetlands, recording Raphidascaris acus larvae and Camallanus lacustris. Khara et al. (2006b) record the trematode Asymphylodora tincae from this species in the Boojagh Wetland of the Caspian coast. Barzegar et al. (2008) record the digenean eye parasite Diplostomum spathaceum from this fish. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Lernaea sp. on this species.

Economic importance

Commercial catches are made in the Volga Delta where there are spring (April) and fall (September-October) fishing seasons. Up to 6,300 tonnes are taken using fyke nets, trap nets and seines. Holčík and Oláh (1992) report a catch of 540 kg in the Anzali Mordab in 1990.

The tench is a sought-after sport fish in Europe, putting on a strong fight for its weight (personal experience). There is an ornamental form, orange-yellow or reddish, which is kept in park ponds.

The roe or eggs of this species are particularly poisonous and it should be cleaned with care to avoid contamination of the flesh although it is not a common market fish in Iran (Halstead, 1967-1970; Coad, 1979b). Probable symptoms and treatment are given under the genus Schizothorax.

The tench is reputedly a "doctor fish" and other species are said to rub against its slimy body as a cure for injuries. Rubbed on humans, it is said to cure fever, headache, toothache and jaundice. In Iran, this species has been proposed as a component of polyculture as it feeds on snails, a host for diplostomiasis-causing parasites. A decrease in snail frequency varied from 43 to 82% in experiments using adult and juvenile tench (Iranian Fisheries Research Organization Newsletter, 37:3, 2003).

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use as food, in textbooks and because it has been introduced outside its natural range.

Conservation

Lelek (1987) classifies this species as intermediate in Europe (liable to be transferred to vulnerable or rare categories if their habitat deteriorates further). Kiabi et al. (1999) consider this species to be of least concern in the south Caspian Sea basin according to IUCN criteria. Criteria include commercial fishing, sport fishing, abundant in numbers, habitat destruction, widespread range (75% of water bodies), absent in other water bodies in Iran, and present outside the Caspian Sea basin.

There have been attempts to breed this species artificially in Iran to increase the recruitment rate and control loss (Annual Bulletin 1993-94, Iranian Fisheries Research and Training Organization, Tehran, p. 78-79, 1995; Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 39, 1996; Sirang, 1997). Brood fish weighed 0.2-0.45 kg and were kept in earthen ponds. Carp culture techniques were used with injection of gonadotropic hormones from carp pituitary at 3-8 mg per kg body weight applied in two doses. Water temperatures were 20-24°C, eggs were placed in saline carbamide solution after fertilisation to remove stickiness and washed in pure water. Swelled eggs measured 0.65-0.8 mm and were bright yellow to greenish-yellow. The number of eggs per spawner was 10,848-17,710 and 80-350 eggs per 1 g body weight. Fertilisation rate was 70-85%. Eggs were incubated for 2.0-2.5 days with a survival rate of hatched fry at 85-95% and size of newly hatched larvae at 4.5-5.0 mm.

Haematological changes were investigated after exposure to mercury, cadmium and lead by Lal Shah (2010), examining fish from Mogan Lake near Ankara, Turkey. Impaired haematological parameters resulted in hyperactivity, increased breathing, accelerated ventilation, surfacing and sinking, erratic swimming, inactivity, lethargy and convulsions.

Further work

The biology of this species has been well-studied elsewhere in its extensive range but not under Iranian conditions.

Sources

Iranian material: CMNFI 1970-0514, 2, 63.9-106.9 mm standard length, Gilan, Shafa River estuary (37º35'N, 49º09'E); CMNFI 1970-0553, 1, 70.9 mm standard length, Gilan, Sowsar Roga River (37º27'N, 49º30'E); CMNFI 1970-0580, 3, 28.0-36.2 mm standard length, Mazandaran, river near Iz Deh (36º36'N, 52º07'E); CMNFI 1979-0439, 1, 170.9 mm standard length, Gilan, Anzali Mordab (ca. 3727'N, ca. 4925'E); CMNFI 1980-0148, 14, 33.9-49.7 mm standard length, Gilan, Pir Bazar Roga River (37º21'N, 49º33'E); CMNFI 1980-0916, 6, 38.2-51.1 mm standard length, Gilan, Nahang Roga River (no other locality data); CMNFI 1993-0137, 1, 150.4 mm standard length, Mazandaran, Sari (36º34'N, 53º04'E).

Genus Tor
Gray1834

?

Much of the past literature on this genus appeared under Barbus (q.v.)

 

Tor grypus
(Heckel, 1843)

Kangir River, Tigris River basin, Ilam, courtesy of A. Mahjoor Azad

Kangir River, Tigris River basin, Ilam, ventral head, courtesy of A. Mahjoor Azad

Changule River, Ilam, Tigris River basin, courtesy of A. Mahjoor Azad

Changule River, Ilam, Tigris River basin, courtesy of A. Mahjoor Azad

Above photographs showing fleshy lips courtesy of Atabak Mahjoor Azad

Common names

شيربت (= shirbot, shirbod or shilbot), shaboot, سس ماهي (= sos or sas mahi), rumi, shebhe shirbot, سرخه (sorkheh, meaning= reddish, a local name in the Zohreh River - J. Gh. Marammazi, pers. comm., 1995), rumi (not heard of in Khuzestan).

[shabout or hamrawi in Arabic; large scaled barb, Tigris barbel].

Systematics

Howes (1987) places this species outside the genus Barbus sensu stricto as defined by him because it has the non-elongate lachrymal bone with a sensory canal running along the antero-dorsal border, a derived condition. Karaman (1971) and Ekmekçi and Banarescu (1998) placed it in the genus Tor (see Barbus) and it may belong in Naziritor Mirza and Javed, 1985 (M. R. Mirza, pers. comm., 6 December 2003). Al-Hassan (1984) looked at several "Barbus" species and found the electropherogram of this species to be distinctive, perhaps indicating that molecular studies could resolve the relationships of this species. This distinction was reiterated by Jawad (2003).

The type locality of Barbus Grypus is "Tigris bei Mossul" (Heckel, 1843b) and Krupp (1985c) records a syntype (dried) from the Naturhistorisches Museum Wien (formerly NMW) now in the Senckenberg Museum Frankfurt under SMF 2613, 375 mm standard length. One syntype is in the Museum für Naturkunde, Universität Humboldt, Berlin (ZMB 8788, not located February 2006). One syntype is in Vienna under NMW 54160, 2 are under NMW 54161 (280.9-318.9 mm total length as measured by me), and 1 is under NMW 91023 (Eschmeyer et al., 1996). The catalogue in Vienna lists 3 fish in spirits and 2 fish stuffed.

Labeobarbus Kotschyi Heckel, 1843 described from the "Tigris bei Mossul" has long been regarded as a synonym although Valiallahi (2000) resurrects this species. Krupp (1985c) records a syntype from the Naturhistorisches Museum Wien under NMW 49729, Th. Kotschy (188.8 mm standard length as measured by me). A dried specimen (NMW 59462) is also a syntype. Eschmeyer et al. (1996) also lists another syntype, NMW 91022. The catalogue in Vienna lists 1 fish in spirits and 2 fish stuffed.

Key characters

This species is identified by having two pairs of barbels, a strong, smooth spine in the dorsal fin, and less than 44 scales in the lateral line.

Morphology

The forehead is more rounded than in type material of kotschyi, although kotschyi types are smaller than grypus types which may account for this distinction. The mouth is inferior, horseshoe-shaped and has fleshy lips. The median lobe of the lower lip is well-developed in some individuals (such specimens were described as kotschyi - this form is rare in Khuzestan according to N. Najafpour, pers. comm., 1995) and intermediates can be seen) but not in others (grypus) (Karaman, 1971). The median lobe may extend back almost as far as the level of the rear margin of the lower lip or be distinctive with free lateral and rear margins but only extend back one third of this distance. The much fleshier lip structure in kotschyi (the upper lip can be reflexed for example) may be a form of hypertrophy seen in other cyprinid fishes (see Roberts and Khaironizam (2008) for further discussions on this feature)). The last unbranched dorsal fin ray is smooth and spine-like, with sharp edges but no serrations although serrations are weakly developed in young fish. Barbels are about equal in length. The gut has two anterior and two posterior loops.

Dorsal fin unbranched rays 4, branched rays 7-9, usually 8, anal fin unbranched rays 3, branched rays 5, pectoral fin branched rays 14-18 (including counts from Jawad (1975)), and pelvic fin branched rays 7-8, usually 8. Total gill rakers 16-22 (including counts by Jawad (1975)). Krupp (1985c) cites 13-17 gill rakers, presumably lower arch ones only. Gill rakers reach the second raker below or beyond when appressed, with large tubercles or branches on the inner surface in 2 rows alternating left and right. Lateral line scales 32-43. A pelvic axillary scale is present. Scales have a subcentral anterior, almost central, focus, numerous fine circuli and many radii on all fields with the exposed part of the scale tubercular. Pharyngeal teeth 2,3,4-4,3,2 in the literature, but see below, anterior teeth rounded, the most anterior one small and blunt, posterior ones spatulate with hooked tips. Total vertebrae 44-47 (Howes, 1987), 43-45 (Jawad, 1975) or 47(8), 48(7), 49(5) or 50(1) in fish seen by me (BM(NH) 1974.2.22:1283-1284, 1299-1315, 1317, 1323; 1920.3.3:1-18, 1874.4.28:24-6).

Ali et al. (1981) found differences in morphology for fish from Al-Therthar Reservoir and the Tigris River in Iraq, not by locality but by habitat type.

Iranian specimens have the following meristic data: dorsal fin branched rays 7(1), 8(20) or 9(1); anal fin branched rays 5(22); pectoral fin branched rays 15(2), 16(15), 17(4) or 18(1), and pelvic fin branched rays 7(1) or 8(21). Lateral line scales 32(4), 33(4), 34(4), 35(3), 36(3) or 37(3). Total gill rakers 16(4), 17(2), 18(3), 19(6), 20(4) or 21(2), with some evidence of higher counts in larger fish. Pharyngeal teeth usually 2,3,5-5,3,2(14) with variants 2,3,5-4,3,2(2), 2,3,4-5,3,2(2), 2,2,5-5,3,2(1) and 1,2,5-4,3,2(1), in contrast to literature reports of 4 main row teeth being typical.

Sexual dimorphism

Ali et al. (1981) found no sexual dimorphism in their Iraqi samples.

Colour

Overall colour has a pale rose to light orange effect, usually without other markings. The back is a dark olive-brown to blackish-green with the flanks pale rose, light orange to yellowish to silvery and belly silvery to milk-white. There may be an indistinct stripe along the mid-flank. Large fish have the upper flank darkened from the overall orange colour of the mid-flank and the lower flank scales are rimmed in white so they stand out. Lips are pale red. The operculum is golden. The pectoral, pelvic, anal and caudal fins are bright orange or pink at the base (perhaps white after preservation), distally blackish. Pectoral and pelvic fins may be dark overall with a reddish to reddish-brown tinge, and the leading edge of the pelvic fin pink. The anal fin may be a bluish-black distally. In some fish the caudal fin is black proximally and reddish distally. In large fish the pectoral, pelvic, anal and caudal are progressively darker in this order. The anal and pelvic fins, the pectoral fins less so, may be heavily pigmented with melanophores on rays and membranes so as to appear black in preserved fish. The dorsal fin is hyaline. The eye rim is yellow-green to lime-green. Young fish may have some scales darkened, giving a mottled effect and are more silvery on the flank than large fish. Their pectoral and pelvic fins are more orange and the anal and caudal fins are only slightly tinged with colour. The caudal fin carries a lot of grey. The smallest fish have a very faint fin colouration. Peritoneum black.

Size

Attains 96.0 cm and 9.7 kg in Dukan Reservoir, Iraq and 96.0 cm and 11.0 kg in Atatürk Dam Lake on the Euphrates River in Turkey (Al-Hakim et al., 1981; Oymak et al., 2008). Gruvel (1931) cited 1.5 m and 30 kg for Syria. Banister (1980) gives nearly 2 m and 100 kg but this may be confusion with B. esocinus although Krupp (1992) also cites almost 2 m. Reputedly reaches 60 kg in Lorestan (S. Nazeeri, pers. comm., 2000) and Ghofleh Marammazi (2004) found fish up to 20 kg in Khuzestan. Specimens reach 3 kg even in the small canals of the sugar-cane fields of Khuzestan.

Distribution

This species is found in the Tigris-Euphrates basin and the Orontes River basin. In Iran it is found in the Tigris River basin up to the Simarreh including marshes such as the Hawr Al Azim, in the Gulf basin in the Zohreh, Shapur, Helleh, Dalaki, Dasht-e Palang, Shur, Dozgah and Mand rivers and Lake Famur (= Perishan), although rare in the latter, and in the Hormuz basin in the Hasan Langi and Kul rivers (Berg, 1949; Gh. Izadpanahi, pers. comm., 1995; Marammazi, 1995; M. Rabbaniha, pers. comm., 1995; Maafi, 1996b; H. R. Alizadeh, pers comm., 2000; Abdoli, 2000).

Zoogeography

Karaman (1971) considers this species to have an Indian line of descent, placing it in the genus Tor which most subsequent authors restrict to the Indian subcontinent and southeast Asia.

Habitat

van den Eelaart (1954) and Al-Hamed (1966b; 1972) describe the habitat for this species in the Tigris River as distributed throughout the river and its tributaries. It is a strong swimmer. Al-Rudainy (2008) states that it can be found in the mid-water column in high current. Mature fish move upstream to the spawning grounds and spent fish descend to their original habitat. In summer under low water level conditions and high temperatures, the smaller fish remain in the lower reaches of rivers but the larger fish migrate up rivers and tributaries, returning in September and October when temperatures fall. This species may enter marshes on floods, favouring areas where there is fresh river water, but returns to rivers as it requires a higher oxygen concentration than most marsh residents. Heydarpour (1978) gives a temperature range of 9-31ºC for this species under culture conditions in Khuzestan.

Marammazi (1994) considers this species to be versatile in its habitats in the Zohreh River which drains to the northern Persian Gulf. It was found throughout the river in contrast to Mesopotamichthys sharpeyi which, being stenohaline, was restricted in its distribution. The form with a well-developed median lobe is said to occur in rocky habitats. This species is considered to be the dominant fish in the Karun and Zohreh rivers (Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 48, 1996; Iranian Fisheries Research and Training Organization Newsletter, 17:1, 1997). Ghofleh Marammazi (2004) found it in almost all water bodies in Khuzestan where it occurred under a wide range of temperatures and salinities. However its presence on the Khuzestan plain was for feeding while for reproduction it required more northerly areas with sandy or gravel substrates, high water flow, low temperatures and high oxygen content (Ghofleh Marammazi, 2000). Ramin (2009) records this species as the most abundant in the Karkheh River out of 37 species and subspecies.

Age and growth

Dorostghoal et al. (2009) found mean body lengths were 36.5-43.5 cm and mean body weights were 835.0-1012.0 g for their Karun River samples. Hashemi et al. (2010) examined fish from the southern Karun River in Iran and found a size range of 20-76 cm and 52-11170 g, growth was isometric, and growth and mortality parameters were L = 86.64, K = 0.27, t0 = -0.46, M = 0.5, F = 1.22, Z = 1.78 and E = 0.71. Relative yield per recruitment (Y'/R) was 0.037, relative biomass per recruitment (B'/R) was 0.29, exploitation ratio maximum sustainable yield (Emax) was 0.44, precautionary average target (Fopt) was 0.25 year-1, and limit (Flimit) as 0.331 year-1. The stock was overfished and fishing regulations are required.

Al-Hakim et al. (1976) studied some aspects of the biology of this species in Razzaza Lake, Iraq. Males are longer than females before maturation and shorter thereafter. Females reach 13 years and males 8 years of age and fish mature at 45-48 cm total length in their fifth year. Males mature earlier than females. Ali et al. (1981) found this species to mature at 3-5 years of age and 40-50 cm in the Al-Therthar Reservoir (about 65 km northwest of Baghdad) and the Tigris River (Kut Dam) in Iraq. Jiad et al. (1984) studied this species in the Al-Hindiya Dam in Iraq and found similar results to the studies cited above.

Al-Hakim et al. (1981) studied this species in the Dukan Reservoir, west of Sulaimaniyah, Iraq. Life span is 17 years for females and 11 years for males. Growth slows with age, and especially after maturity, and is fastest in the first year of life. 30% of males mature at age group 3 (39 cm) and all were mature at age group 6 (48 cm). Al-Hamed (1966a; 1966b; 1972) working with Tigris River populations in Iraq, found males to mature at about 45 cm and females at about 50 cm, with most fish mature in their fourth year and spawning at the beginning of their fifth year of life. Some fish mature in age group 3 and some as late as age group 5. Maximum age observed was 12 years. Males outnumber females, being two thirds of the fish on the spawning grounds. Al-Hakim et al. (1981) found all females are mature at 51 cm (age group 7) but only 10% at 42 cm. Males mature earlier than females and may grow faster and die younger.

Ali et al. (1981) found growth to be better in fish from Al-Therthar Reservoir compared to those from the Tigris River in Iraq. The fishing methods used, both commercially and experimentally, caught mature fish of 40-50 cm and 3-5 years of age.

Oymak et al. (2008) examined age and growth in the Atatürk Dam Lake on the Euphrates River in Turkey. Fourteen age classes were found with age classes 4-6 for females and 2-4 for males dominant. von Bertalanffy growth equations were given for males and females.

Growth in a polluted section of the Diyala River, Iraq is poor compared to other populations (Khalaf et al., 1984; Khalaf et al., 1985).

Food

Al-Hamed (1965) found this species to be a herbivore taking filamentous algae and higher plant parts. Incidental food items taken while feeding on plants include fish tissue and scales. Fallen ripe fruits from trees overhanging the water are also consumed as are cereal grains from loading docks. It may also take some small fishes. Iranian specimens contain filamentous algae, plant fragments and associated invertebrates. Ghofleh Marammazi (2000) considers it to be an omnivore.

Reproduction

In Dukan Reservoir, Iraq spawning takes place from the beginning of May until the end of June. van den Eelaart (1954) and Al-Hamed (1966b; 1972) studied the reproduction of this species in Iraq. Eggs are deposited on fine gravels overlying a layer of coarse sand in shallow, wide holes. Water depth varies from 30 to 150 cm. Egg diameter is 1.5 mm and fecundity up to 147,000. The spawning season on the Tigris River between Beled and Tigrit is late May to late June after an upriver migration in April. Fish appear on the spawning grounds in schools just before dark and remain there until shortly before midnight, making loud noises by splashing, jumping and chasing. After spawning, the fish return downriver but do not enter marshes as these are now too warm. Oymak et al. (2008) examined reproduction in the Atatürk Dam Lake on the Euphrates River in Turkey and found a sex ratio of 1:1.34 (females/males), with a spawning period in May to July, a fecundity up to 235,764 eggs and a mean egg diameter of 2.183 mm.

Its presence in areas of the Khuzestan plain is mainly for feeding while reproduction occurs in the northern parts of this province where there are sandy and gravel substrates, fast current, low temperatures and high oxygen content (Ghofleh Marammazi, 2004). A prolonged spawning season in fish from the Karun River, late April to early August with the highest gonadosomatic index June-July, was determined by Dorostghoal et al. (2009) using macroscopic and microscopic techniques. They also note that the fish migrate upstream for spawning in May.

Parasites and predators

Bykhovski (1949) report a new species of monogenetic trematode, Dactylogyrus pavlovskyi, from this species in the Karkheh River, Iran. Molnár and Jalali (1992) describe a new species of monogenean, Dogielius persicus, from this species in the Dez and Karun rivers of Khuzestan. Masoumian et al. (1994) describe two new species of Myxosporea from the gills of this species in the Karun River, Khuzestan, namely Myxobolus karuni and Myxobolus persicus. Molnár et al. (1996) report additional new species from this fish in Khuzestan, namely Myxobolus iranicus in the spleen and Myxobolus mesopotamiae in connective tissue of the caudal and pectoral fins. The latter myxosporean is also reported from Barbus rajanorum as is a new species Myxobolus shadgani infecting the gills - the identity of the host fish is unknown as Barbus rajanorum is not a distinct species (see under Luciobarbus pectoralis). Myxosporeans are potentially dangerous to fishes such as Tor grypus which may be used in fish culture in Khuzestan.

The monogeneans Dactylogyrus povlovskyi, D. barbioides, Dogielius persicus, Gyrodactylus sprostonae and Paradiplozoon sp. are recorded from this species in the Karun River with heavier infestations in spring and summer than in autumn and winter. These gill parasites caused no serious injuries but were thought to be important in respect of monitoring infestation levels on fish farms in Khuzestan (www.avz1.8m.com/fulltext.htm, downloaded 28 October 2002).

Ebrahimzadeh and Nabawi (1975) list a nematode species Philometra and Ascaridae from this species in the Karun River. Ebrahimzadeh and Kailani (1976) record parasites in the genera Myxosoma (protozoan) and Isoglaridacris (cestode) and also a nematode from Barbus (= Tor) grypus taken in the Karun River. Masoumian and Pazooki (1999) list Myxobolus persicus, M. karuni, M. mesopotamiae and M. iranicus from this species in various localities in Khuzestan. Moghainemi and Abbasi (1992) record a wide range of parasites from this species in the Hawr al-Azim in Khuzestan. Farahnak et al. (2002) record Contracaecum sp. and Anisakis sp. from this fish in Khuzestan Province. Peyghan et al. (2001) record Neoechinorhynchus sp. from fish from Khorramabad rivers. Peyghan et al. (2001) record Myxobolus karuni in 86.7% of fish in the Karun River at Ahvaz and Papahn et al. (2004) record the monogeneans Dactylogyrus pavlovskyi, D. barbioides, Dogielius persicus, Gyrodactylus sprostonae and Paradiplozoon sp. from the same locality, the latter two being first records for the host in Iran. Barzegar et al. (2008) record the digenean eye parasite Tylodelphys clavata from this fish. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Argulus sp. and Ergasilus sp. on this species.

Economic importance

An important food fish, with desirable taste (Al-Rudainy, 2008), comprising 23% of the total fish production in Iraq for example and forming the most important commercial fish there (Al-Hakim et al., 1981). Petr (1987) reports the catch for all Iraq in 1976 as 519 t. The weight at the Basrah fish market from October 1975 to June 1977 was only 3,330 kg however (Sharma, 1980) and Khalaf et al. (1984) rank it third in the inland wholesale trade of Iraq for the period 1967-1970.

This species is the preferred catch of anglers at Ahvaz in Khuzestan, with bread or potato as bait. There is a good demand for this species in local markets of Khuzestan (Ghofleh Marammazi, 2004). Peyghan et al. (2001) report that is is an economically important species with a good market value in the Khorramabad region.

Petr (1987) has suggested that this species be investigated for fish farming in Khuzestan. The Khuzestan Fisheries Research Centre at Ahvaz has experimented with this species in pond culture (Emadi, 1993a; Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 6, 1996).

Conservation

The stock of this species in the Gav Masiab River is severely reduced and only 3 fish were caught in western Iran in the Zagros rivers during a 4-year survey (J. Valiallahi, www.modares.ac.ir, downloaded 4 July 2000; pers. comm., 2001). Shirbot were considered to be on the verge of extinction in the Gav Masiab River of the Tigris River basin, through pollution, overfishing, dam building, aquaculture, and introduction of exotics (IranMania.com, 29 December 2006). Dadelahi Sohrab et al. (2009) found lead and cadmium levels in fish from the Arvand River were higher than acceptable by international standards. Dorostghoal et al. (2009) note that the species is caught on the spawning migration and while spawning and this accounts for its decline. Probably in decline in Turkey also (Fricke et al., 2007).

Further work

The biology of this species in Iran needs investigation as does its putative relationship to South Asian fishes in the genus Tor.

Sources

Type material: See above, Barbus grypus (NMW 54161) and Labeobarbus kotschyi (NMW 49729).

Iranian material: CMNFI 1979-0155, 1, 42.8 mm standard length, Fars, spring at Gavanoo (28º47'N, 54º22'E); CMNFI 1979-0291, 1, 31.4 mm standard length, Kermanshahan, Diyala River drainage (34º24'N, 45º37'E); CMNFI 1979-0356, 5, 22.5-40.7 mm standard length, Khuzestan, Karkheh River drainage at Hoveyzeh (31º27'N, 48º04'E); CMNFI 1979-0360, 1, 375.5 mm standard length, Khuzestan, canal branch of Karkheh River (31º40'N, 48º35'E); CMNFI 1979-0364, 1, 22.3 mm standard length, Khuzestan, river at Abdolkhan (31º52'30"N, 48º20'30"E); CMNFI 1979-0384, 2, 202.2-222.2 mm standard length, Khuzestan, Ab-e Shur drainage (32º00'N, 49º07'E); CMNFI 1979-0391, 1, 220.5 mm standard length, Khuzestan, stream in Marun River drainage (31º28'N, 49º51'E); CMNFI 1979-0392, 1, 62.4 mm standard length, Khuzestan, Zard River (ca. 31º32'N, ca. 49º48'E); CMNFI 1979-0395, 2, 32.7-38.0 mm standard length, Khuzestan, stream in Marun River drainage (ca. 30º57'N, ca. 49º51'E); CMNFI 1979-0402, 1, 80.7 mm standard length, Bushehr, Mand River 12 km north of Kaki (ca. 28º25'N, ca. 51º32'E); CMNFI 1991-0153, 1, 253.5 mm standard length, Khuzestan, Zohreh River (no other locality data); CMNFI 1993-0141, 1, 80.2 mm standard length, Bushehr, Dalaki River (29º28'N, 51º15'E); CMNFI 1995-009A, 4, Khuzestan, A'la River at Pol-e Tighen (31º23'30"N, 49º53'E); BM(NH) 1980.8.28:7, 1, 72.0 mm standard length, Khuzestan, Dezful (32º23'N, 48º24'E); ZMH 2508, 1, 343.3 mm standard length, Khuzestan, Karun River at Ahvaz (31º19'N, 48º42'E); ZSM 21864, 1, 157.7 mm standard length, Khuzestan, Dez River at Harmaleh (31º57'N, 48º34'E);

Comparative material:- BM(NH) 1874.4.28:24-26, 3, 231.2-254.7 mm standard length, Iraq, Tigris River at Baghdad (33º21'N, 44º25'E); BM(NH) 1920.3.3:1-18, 5, 104.8-196.0 mm standard length, Iraq, Basrah (30º30'N, 47º47'E); BM(NH) 1973.5.21:191, 1, 205.4 mm standard length, Iraq, Shatt-al-Arab; BM(NH) 1973.5.21:192, 1, 139.5 mm standard length, Iraq, Shatt-al-Arab; BM(NH) 1974.2.22:1283-1284, 2, 121.0-164.5 mm standard length, Iraq, Khalis (no other locality data); BM(NH) 1974.2.22:1299-1315, 9, 64.9-101.5 mm standard length, Iraq, branch of Khalis River (no other locality data); BM(NH) 1974.2.22:1317, 93.9 mm standard length, Iraq, branch of Khalis River (no other locality data); BM(NH) 1974.2.22:1323, 1, 160.1 mm standard length, Iraq, Basrah (30º30'N, 47º47'E); BM(NH) 1974.2.22:1328, 1, 161.9 mm standard length, Iraq, Basrah (30º30'N, 47º47'E); KU 10516, 1, 124.1 mm standard length, Iraq, Basrah (30º30'N, 47º47'E).

Genus Vimba
Fitzinger, 1873

This genus is found in the basins of the Baltic, Black and Caspian seas and has a single species. It is characterised by a compressed, moderately deep body with an inferior, crescentic mouth, a scaleless keel between the pelvic and anal fins, a scaleless groove in front of the dorsal fin and an evident keel behind it, pharyngeal teeth in a single row, short dorsal and long anal fin, gill rakers short, and scales moderate in size. Bogutskaya (1986) using skull morphology reaffirms the generic separation of Vimba Fitzinger, 1873 from Abramis Cuvier, 1816 although Howes (1981) considers it to be a synonym.

Vimba persa
(Pallas, 1814)

CMNFI 1979-0435, Gilan, stream 10 km west of Ramsar

Common names

سياه كولي (= siah kuli or siahkooli, meaning black fish, cooli, couli, coli, kooli or kuli being any small fish; in Gilaki), mahi siah kuli.

[garasol in Azerbaijan; chernospinka, Kaspiiskii rybets or Caspian vimba, both in Russian; southern white-eye, vimba].

Systematics

Cyprinus persa Gmelin, 1774 is a nomen nudum - see Kottelat (1997) - and was later made available by Pallas. It was described originally from "Persa; in lacubus ad Cyrum", i.e. the southern coast of the Caspian Sea in lakes of the Kura River system in Azerbaijan. It is distinguished by larger scales and usually fewer anal rays from V. vimba (Linnaeus, 1758) which it was formerly synonymised with or recognised as a subspecies of. Cyprinus Vimba was originally described from lakes of Sweden. Hänfling et al. (2009) found phylogenetically distinct mtDNA sequences for Caspian Sea basin samples, the Caspian clade having diverged from a western or Pontic clade 1-2 MYA at the beginning of the Pleistocene. Caspian populations could then rank as a separate species or subspecies although they consider further work involving western Caucasian populations is needed to support one conclusion over the other. Naseka and Bogutskaya (2009) recognise V. persa as a species.

Rahmani and Abdoli (2008) compared populations from the Gorgan River, Shirud and Anzali Lagoon and found morphometric and meristic differences between them.

A hybrid with Alburnus chalcoides was reported from the Safid River (Petrov, 1926).

Key characters

The snout projects over the lower jaw and in large fish is quite bulbous, there is a keel on the belly and on the back, and fin ray counts are distinctive.

Morphology

Western and southeastern populations in the Caspian Sea can be distinguished morphometrically, and represent different stocks.

Dorsal fin with 2-3 unbranched rays (always 2 in the subspecies persa (Berg, 1948-1949) but the first unbranched ray is minute and visible in x-rays in Iranian specimens) and 7-9, usually 8, branched rays, anal fin with 3 unbranched and 12-22 branched rays (16-18 in persa but see below), pectoral fin branched rays 11-18 and pelvic fin branched rays 7-10. Lateral line scales 47-64, 48-54 in persa (but see below). The lateral line runs below the midline of the caudal peduncle. Predorsal scales are small and crowded. A pelvic axillary scale is present. The naked ventral keel begins 0-3 scales behind the pelvic fin bases. Scales at the anal fin base form a sheath. The anterior scale margin is wavy and the posterior margin is crenualte. There is a central focus, numerous fine circuli and few anterior and posterior radii. Gill rakers 12-20, small and reaching the raker below when appressed. Pharyngeal teeth usually 5-5, with the largest teeth having long and narrow, flat to slightly concave crowns, and tips recurved or very slightly hooked. Vertebrae 38-45. Gill rakers 14-19. The gut is s-shaped. The chromosome number is 2n=50 (Klinkhardt et al., 1995) or 2n=52 (Reshetnikov, 2002).

Meristic values for Iranian specimens are:- branched dorsal fin rays 7(1) or 8(39); branched anal fin rays 16(3), 17(13), 18(18) or 19(6); branched pectoral fin rays 14(8), 15(22), 16(7) or 17(3); branched pelvic fin rays 8(10) or 9(30); lateral line scales 47(1), 48(4), 49(7), 50(10), 51(14), 52(1), 53(1), 54(1) or 55(1); total gill rakers 15(1), 16(2), 17(12), 18(14), 19(9) or 20(2); pharyngeal teeth 5-5(15), 5-4(4) or 4-5(1); and total vertebrae 41(3), 42(9), 43(24) or 44(3). Abbasi et al. (2004) found 149 Safid River fish to have mean values of 50.83 lateral line scales, branched dorsal fin rays 7.96 and branched anal fin rays 17.58.

Sexual dimorphism

Females are slightly larger than males of the same age and differ morphometrically on account of the eggs distorting body shape. The males become black on the back, reddish on the belly, their fins become red and the tips of the dorsal and caudal fins become dark, and they develop minute tubercles on the body during the spawning season (Kuliev, 1988; Abbasi et al., 2004). Females may also develop tubercles but to a lesser extent. Safid River fish showed differences in 2 meristic and 16 morphometric characters, especially body depth and lengths of dorsal, pectoral, pelvic and anal fins (Abbasi et al., 2004).

Iranian specimens have small tubercles lining the scale margins and larger tubercles over the whole head but particularly on the dorsal surface and upper sides. Fin rays bear small tubercles in files following the branching of the rays. The pelvic fin has weakly developed tubercles on its ventral surface as well as dorsally. The unbranched pectoral and pelvic rays bear several rows of tubercles.

Colour

The back is a reddish-brown to grey-blue, flanks are silvery and the belly yellowish. Paired fins are red at the base, pink distally. The anal fin base is red while other fins are grey to hyaline. Spawning fish develop black stripes along the dorsal and ventral body.

Size

Reaches 60.0 cm and 3 kg. The subspecies persa is smaller, to 30 cm.

Distribution

Found from central Europe to the Caspian Sea basin. In Iran it is recorded from the Aras to the Atrak rivers in the Caspian Sea basin including the Manjil Reservoir on the Safid River, the Anzali Mordab and Gorgan Bay, the Gorgan, Gharasu, Tajan, Babol, Haraz, Sardab, Tonekabon, Pol-e Rud and Safid rivers, and the southeast, southwest and south-central Caspian Sea (Kozhin, 1957; Nümann, 1966; Holčík and Oláh, 1992; Riazi, 1996; Abbasi et al., 1999; Kiabi et al., 1999; Abdoli, 2000; Abdoli and Naderi, 2009). It has also been introduced to Sistan.

Zoogeography

This species reaches its most south-easterly occurrence in Iran and has been given subspecific status. Its relationships lie with European taxa (see Abramis).

Habitat

Caspian vimba have a sparse distribution in the sea and are not fished there commercially. It is more common in Gilan than Mazandaran and Golestan coastal waters (Jolodar and Abdoli, 2004). The semi-migratory form enters fresh water or brackish water only for reproduction in spring. After spawning, it migrates to river mouths to feed until the next reproductive season (Kuliev, 1988). Riazi (1996) reports that this species is native (resident) to the Siah-Keshim Protected Region of the Anzali Mordab. S. Bazari Moghaddam (www.meeresschule.com/cgi-bin/abstracts/gastbuch.asp, downloaded 17 January 2005) records a migration into the Safid River in spring for reproduction. Feeding continues on this migration. Knipovich (1921) reports this species from depths of 36.6-53.0 m in the Iranian Caspian Sea. In fresh water it occurs in schools in the lower reaches of rivers, in deep water over stone and gravel bottoms. It may also occur in lakes over mud bottoms.

Age and growth

Maturity is attained in the second or third year of life, males maturing at age 2 in the Anzali region. In the Safid River migrating fish were 2-4 years old, predominately 3-year-old fish (S. Bazari Moghaddam, www.meeresschule.com/cgi-bin/abstracts/gastbuch.asp, downloaded 17 January2005). Four-year-olds predominate in the spawning population in Kyzylagach or Imeni Kirova Bay, Azerbaijan. Most spawning females are 16-23 cm (46%) and males 13-19 (42%). Large fish spawn first and the number of smaller fish spawning increases towards the end of the reproductive season (Kuliev, 1988; Shikhshabekov, 1979). Most fish on the spawning migration into the Anzali Mordab are 170-250 mm and ages 3-4 years (Holčik and Oláh, 1992). Fish on the spawning migration of the Safid River had a fork length of 116-208 mm and a weight of 21.1-116.1 g in males and 122-222 mm and 23.1-170.0 g in females. Spawning males were 2-6 years old and females 3-7 years (Abbasi et al., 2005). Maximum life span is about 15 years.

Food

Diet is aquatic insects, crustaceans, snails, worms and algae on muddy bottoms. Iranian specimens had zebra mussels and insect remains. S. Bazari Moghaddam (www.meeresschule.com/cgi-bin/abstracts/gastbuch.asp, downloaded 17 January 2005) reports oligochaetes, chironomids and Odonata in fish from the Safid River.

Reproduction

Kuliev (1988) and Shikhshabekov (1979) studied reproduction in the Kyzylagach Bay of the southwestern Caspian Sea and the waters of Dagestan respectively. The spawning migration begins in March or April at 10-13°C and spawning takes place at the end of April at 16-20°C, continuing until the end of May or into June.

Fish enter the Anzali Mordab of Iran in mid-January at a water temperature of 8-9°C, peaking from 21 April-10 May at 19-21°C (Holčík and Oláh, 1992). Khaval (1998) reports a spawning migration into the Safid River despite construction, sand removal and pollution. Fish from the Shafa River estuary in Iran caught on 10 April had highly developed eggs measuring 1.3 mm. Fecundity is up to 89,200 eggs per female, increasing with age and body size (elsewhere in Europe to 200,000 eggs with a diameter of 1.4 mm). Spawning is non-intermittent, in contrast to Black Sea vimbas. Eggs are deposited on gravel or stones where there is a current of 0.6-0.9 m/second. Concrete structures and flooded fields may be used as long as there is some current (Holčík and Oláh, 1992). The eggs may form a layer up to 10 cm thick. Initially attached to plants or stones, the eggs are later washed down between the plants or stones. Other fishes eat these eggs and mortality is high. Some fish deposit eggs in sandy shallows of bays or on the roots of reeds and bulrushes. The young migrate to the coastal zone of the Caspian Sea for the summer, moving to greater depths as winter approaches. At temperatures of 17-22°C, eggs incubate for 70-77 hours (Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 37-38, 1996).

Abbasi et al. (2005) found that the Safid River population started the spawning migration in March and this continued until July, peaking in mid-April to late May. Gonad weight for females increased with distance from the estuary. Spawning occurred from late May to late June, peaking in May at 18-29ºC water temperatures. Eggs were shed on pebble and gravel grounds 25-75 km from the estuary. The Disaam tributary was the major spawning site.

Parasites and predators

Jalali and Molnár (1990a) record the monogeneans Dactylogyrus cornoides and D. haplogonus from this species in the Safid River and Pazooki and Aghlmani (no date) record the trematode Asymphylodora kubanicum from Iranian specimens. Sattari et al. (2007) record the cestodes Caryophyllaeus laticeps and Caryophyllaeus fimbriceps, the digenean Diplostomum spathaceum and the monogenean Dactylogyrus extensus in this species in the Anzali wetland of the Caspian shore.

Economic importance

The vimba catch over the whole Caspian Sea basin was less than 100 tonnes per year in the 1980s (Kuliev, 1988). The catch by local fishermen in the Anzali Mordab region in 1990-1991 was 823 kg or about 8400 fish (Holčík and Oláh, 1992). They are caught in rogas and inflowing rivers of the mordab in late winter and early spring. In 1994-1995, the population of this species was noted as declining in recent years (Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 37-38, 1996).

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use as food and in aquaculture.

Conservation

Weirs are a problem for this species in Iran as they block the spawning migration, the fish massing below the obstruction, and causing re-absorption of eggs and sperm (Holčík and Oláh, 1992). Aquaculture of this species has been investigated in Iran; it can be bred semi-artificially using methods similar to that for Rutilus species (Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 38, 1996).

Lelek (1987) classifies this species as intermediate to rare in northwestern Europe.

Kiabi et al. (1999) consider this species to be near threatened in the south Caspian Sea basin according to IUCN criteria. Criteria include commercial fishing, sport fishing, abundant in numbers, habitat destruction, widespread range (75% of water bodies), absent in other water bodies in Iran, and absent outside the Caspian Sea basin. Mostafavi (2007) lists it as near threatened in the Talar River, Mazandaran.

Further work

The relationships of the Caspian Sea taxon to other Vimba needs investigation as do certain aspects of its biology in Iranian waters as it is under some threat.

Sources

Iranian material: CMNFI 1970-0531, 8, 44.4-70.5 mm standard length, Mazandaran, Larim River (36º46'N, 52º56'E); CMNFI 1970-0543A, 9, 38.9-87.4 mm standard length, Gilan, Caspian Sea at Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1970-0544, 1, 162.2 mm standard length, Gilan, Caspian Sea near Bandar-e Anzali (37º28'N, 49º27'E); CMNFI 1979-0431, 1, 145.0 mm standard length, Mazandaran, fish bazaar at Now Shahr (no other locality data); CMNFI 1979-0435, 1, 141.1 mm standard length, Gilan, stream 10 km west of Ramsar (36º57'N, 50º37'E); CMNFI 1979-0436, 5, 115.1-156.5 mm standard length, Gilan, stream 26 km west of Ramsar (37º02'30"N, 50º27'E); CMNFI 1979-0437, 1, 155.2 mm standard length, Gilan, Safid River, 2km west of Astaneh (37º16'30"N, 49º56'E); CMNFI 1979-0438, 2, 139.5-152.6 mm standard length, Gilan, Gholab Ghir River (37º27'N, 49º37'E); CMNFI 1979-0788, 2, 28.5-50.7 mm standard length, Mazandaran, Gorgan River (37º00'N, 54º07'E); CMNFI 1980-0120, 7, 47.8-65.5 mm standard length, Mazandaran, Babol River at Babol Sar (36º43'N, 52º39'E); CMNFI 1980-0121, 3, 139.5-152.2 mm standard length, Gilan, Shafa River estuary (37º35'N, 49º09'E); CMNFI 1980-0126, 1, 170.8 mm standard length, Gilan, Caspian Sea near Bandar-e Anzali (37º28'N, 49º27'E); CMNFI 1980-0127, 2, 180.3-182.1 mm standard length, Gilan, Caspian Sea near Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1980-0129, 1, 51.9 mm standard length, Mazandaran, Tajan River (36º49'N, 53º06'30"E); CMNFI 1980-0138, 10, 28.1-105.0 mm standard length, Gilan, Safid River estuary (ca. 37º28'N, ca. 49º54'E); CMNFI 1980-0142, 1, 180.1 mm standard length, Gilan, Nahang Roga River (no other locality data); CMNFI 1980-0908, 4, 87.8-135.7 mm standard length, Gilan, Safid River estuary (ca. 37º28'N, ca. 49º54'E).

 Cobitidae

This family of loaches, sometimes called sting-loaches, is found in Eurasia and Morocco and has about 26 genera with about 177 species (Berra, 2001; Nelson, 2006). Berra (2001) does not indicate the more southern distribution of this genus in Khuzestan and Fars provinces of Iran. Four species are recorded from Iran.

Anonymous (1988a) places Cobitidae on the Official List of Family-Group Names in Zoology (rather than the grammatically correct but unused Cobitididae) and Cobitis taenia is designated as the type species for the genus Cobitis (see also Kottelat (1986) for further information).

The body form is fusiform to rounded or elongate; the mouth is subterminal and has 3-6 pairs of barbels; the mental lobes of the lower lip have two parts: the anterior which is usually short and sometimes divided into lobules, and the posterior which is flap-like and longer and sometimes divided into 2 or more barbel-like extensions; there is 1 row of pharyngeal teeth; and there is an erectile spine in a groove below the eye (anterior in a non-Iranian genus). Iranian species have one pair of rostral barbels and a rounded or slightly emarginate caudal fin and belong to the subfamily Cobitinae. Menon (1992) considers that structural details of the bony covering of the swimbladder and the nature of the scales are only of use at the generic level. Lip structures, fin positions relative to one another and secondary sexual characteristics in males are important characters in differentiating species in India. Economidis and Nalbant (1996) discuss characters used in the study of these fishes and consider scales to be characteristic of each species along with colour pattern, sexual dimorphism, suborbital spine morphology, barbel and mental lobe morphology, and others. Black spots at the caudal fin base and four longitudinal pigment zones on the flank (Z1-4 or Gambetta's zones 1-4) are important in distinguishing and describing species. Males have 1-2 laminar projections on the dorsal surface of the anterior pectoral fin rays known as the laminae circularis or Canestrini's scales. Hybrid lineages are known, produced by gynogenesis and are nearly all-female. Males of bisexual lineages are sperm donors but the sperm only induces egg development and contributes no genetic material. The all-female lineages are therefore sperm parasites and have to occur in sympatry with one of the parental species in a hybrid complex. As a result, hybridogenous individuals in the complex are difficult to distinguish from the bisexual parent species on external characters. An unbalanced sex ratio in a population with more females than males is usually evidence that hybridogenous lineages are present (Kottelat and Freyhof, 2007). Maximum size is about 40 cm but most are much smaller.

The origins of this group of loaches may well lie at the end of the Eocene or in the early Oligocene in South China, spreading along a northern route through Europe and Siberia during the Oligocene-Miocene-Pliocene period and then later southwards into Southwest Asia (Sawada, 1982; Menon, 1987; 1992; Bănărescu and Nalbant, 1998; Šlechtová et al., 2008; Tang et al., 2008). A early Oligocene route also existed between the Anatolian landmass and Central Asia (Tang et al., 2008) and cobitids may have invaded the Euro-Mediterranean zoogeographic subregion at least five times independently based on cytochrome b data.

Some members of this family can live in oxygen-poor waters. They take in air at the surface, and pass it through the intestine where the mucosa absorbs the oxygen and carbon dioxide waste is released through the vent. As a consequence, they may be very sensitive to air pressure changes and become restless when it falls and can be used to predict the weather. Foods are mostly small insects, worms and crustaceans detected by the aid of the barbels on the habitat bottom. Some eat algal films or mats. Most species bury themselves in sand or mud during the day, emerging to feed at night. Movement is by undulations of the body, particularly marked in the more elongate species. A consequence of this form of movement is a reduction in fin size and variation. Reproduction involves the male chasing the female, entering vegetation and wrapping around the female as eggs are released and fertilised. Eggs swell and reach as large as 3.5 mm in diameter and as a result are retained in the vegetation although they are not adhesive.

A number of species are popular aquarium fishes, including the coolie or kuhli loaches and the weatherfish. None of the Iranian fishes are used in this fashion but they can be quite colourful. Cobitis taenia is a potential fishing bait for predatory fishes such as Sander lucioperca and has been examined experimentally for this purpose in Turkey (Kuşat et al., 1995).

Genus Cobitis
Linnaeus, 1758

These fishes are found in Europe, North Africa and Asia. There are 2 species in Iran. They are known generally as سگ ماهي (sag mahi meaning dog fish) in Farsi, the equivalent of loach in English. This name is not repeated under each Species Account. Also called لوچ (= louch in Farsi, from loach).

This genus is characterised by an elongate and compressed body, a usually bifid, erectile spine below the eye (sometimes hidden under the skin), 3 pairs of short barbels (4 at the snout tip and 2 at the mouth corners), minute scales cover the body (as many as 200 but they are seldom counted accurately), lateral line faint or indistinct, dorsal and anal fins small, caudal fin rounded or truncate, and swimbladder in a bony capsule with a free portion visible. Males have bony extensions of their pectoral fin rays, known as lamina circularis or scale of Canestrini, and no swellings of their body sides.

Polyploid unisexual, bisexual-unisexual complexes and gynogenetic forms of Cobitis exist in the basins of the Baltic, Black, Caspian and Mediterranean seas (Vasil'ev and Vasil'eva, 1996; Kottelat, 1997; Vasil'eva and Vasil'ev, 1998; Vasil'ev et al., 1999; Bohlen, 2001). The species are morpohologically undifferentiated and therefore require detailed study to resolve taxonomic and systematic problems. The composition of Iranian species has not been investigated.

The earliest fossil record is from the middle Miocene about 15MYA with divergence of Sabanejewia and Cobitis at 12-13MYA (Ludwig et al., 2001).

These loaches often bury themselves in mud to overwinter or escape predators. The spine under the eye when erected is an anti-predator device, discouraging swallowing by other fishes and birds. The fish is said to actively swing the head from side to side to prick predators.

Cobitis linea
(Heckel, 1849)

Common names

sagmahi-ye jonubi (= southern dogfish).

[southern spined loach]

Systematics

The type locality of Acanthopsis linea is "Bäche um Persepolis" according to Heckel (1846-1849b). Persepolis is at 29°57'N, 52°52'E in Fars.

The holotype described by Bănărescu and Nalbant (1966) is one of 7 specimens assumed to be syntypes. The catalogue in Vienna lists 1 specimen in one column and 6 in the adjacent column. The type series is in poor condition being dried with scales lost and colour mostly faded except for one specimen. Bianco and Nalbant (1980) cite a specimen as the holotype, a female, 67.0 mm standard length, housed in the Naturhistorisches Museum Wien (NMW 48560 - this includes all 7 fish) and the type locality as Persepolis, presumably following Bănărescu and Nalbant (1966). The dried specimens measure 41.2-62.9 mm standard length and the undried one 61.3 mm standard length. My measurement is at variance with these authors.

Placed in the subgenus Bicanestrinia Băcescu, 1962 by Bianco and Nalbant (1980) and originally described in the genus Acanthopsis Agassiz, 1832 (see Eschmeyer (1990) for further details on this genus). Bohlen et al. (2006) place this species in their Bicanestrinia lineage I using the cytochrome b gene. Bicanestrinia has a derived character state in the duplication of the Canestrini scale or lamina circularis on the pectoral fin of males, the primitive condition being a single lamina circularis as in, e.g. Cobitis s.s. The separation between Bicanestrinia and Cobitis s.s. occurred 12-17 MYA when the land connection between Central Europe and Anatolia broke.

Material identified as Cobitis turcica (Hanko, 1924) in Nalbant and Bianco (1998) is C. linea based on my examination of IZA 7829-30. Fricke et al. (2007) have it in the Tor (sic, meaning Kor) River system of Iran, the same error.

Key characters

This species is distinguished by the dark brown lateral spots being reduced or absent, males have 2 Canestrini's scales at the upper bases of the unbranched and first branched pectoral fin rays, and the laterocaudal branch of the suborbital spine is reduced or absent (although the spine itself is detectable by touch in even the smallest fish).

Morphology

Dorsal fin unbranched rays 2 and branched rays 6-7, anal fin unbranched rays 2-3 and branched rays 5, pectoral fin branched rays 7-9, and pelvic fin branched rays 5-6.

Meristic values for specimens examined by me are:- dorsal fin branched rays 6(16), anal fin branched rays 5(16), pectoral fin branched rays 8(6), pelvic fin branched rays 6(6), and total vertebrae 41(1).

Scales are embedded and have a reduced and eccentric focus. Johal et al. (2006) and Esmaeili and Niknejad (2006-2007) give scanning electron micrographs of the scales. The lateral line does not pass the end of the pectoral fin. The swimbladder capsule is globular but has ventral indentations anterolaterally on both sides. There are small keels on the upper and lower caudal peduncle. The anterior nostril is a short tube. The upper lip has fine furrows, the lower lip is thick and folded with a pair of medial lobes.

Sexual dimorphism

Males have Canestrini's scales as detailed above.

Colour

Overall colour is a yellowish-white with a golden iridescence especially on the operculum. Spots, blotches and dots are blue-grey or brown. The flanks are variably spotted and striped, the variability not related to sexual dimorphism. In some fish there is an upper flank, uniform stripe closely following the line of the back, below it a longitudinal series of irregular spots and blotches, a third stripe incompletely developed, and a fourth mid-lateral stripe of spots and blotches. In others the third and fourth stripes are an indistinguishable row of speckles. The mid-line of the back has a row of rounded spots numbering 7-11 (usually 8) predorsally, 0-3 (usually 2) at the dorsal fin base, and 5-11 (usually 8) postdorsally. The upper head is covered with many minute dots, extending onto the snout and upper head sides. The iris is golden-yellow. There is a blackish band from the eye to the mouth corner. There is a minute black spot on the caudal fin base. Fins are translucent. There are numerous dots scattered on the rays and membranes of the dorsal and caudal fins forming up to 9 irregular rows on the former and 8 more regular rows or bars on the latter, but still not very clearly defined. Spots on the dorsal edge of the caudal fin are clearly defined and regular; there are no spots on the ventral edge. The pelvic and anal fins are immaculate while the pectoral fin has some minor and irregular spots.

Size

Reaches 89.6 mm standard length.

Distribution

This species is found in the Kor River basin and the upper Kul River drainage of the Hormuz basin (Bănărescu and Nalbant, 1966; Bianco and Nalbant, 1980).

Zoogeography

This species may be related to Cobitis simplicispina Hankó, 1925 from Anatolia and other nominal species, all placed in the subgenus Bicanestrinia characterised by two Canestrini's scales. Members of this complex of species possibly reached western Asia from eastern Asia in the early Miocene. The disjunct distributions seen today were probably produced by Pleistocene climatic changes (Bianco and Nalbant, 1980).

Habitat

This species favours muddy bottoms. Specimens were caught in about 5-6 cm of mud, or at the foot of a muddy bank, with aquatic vegetation in the form of reeds.

Age and growth

Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 108 fish measuring 3.35-8.96 cm standard length. The a-value was 0.0089 and the b-value 3.060 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases).

Food

Unknown.

Reproduction

Adult females described by Bianco and Nalbant (1980) measuring 80.0-88.6 mm standard length have ripe eggs 0.8-1.1 mm in diameter. A specimen from the Lapui spring caught on 30 June carries eggs of this size.

Parasites and predators

Unknown.

Economic importance

This species is too small and rare to be of direct economic importance.

Conservation

Since this species is known only from a limited number of specimens, further studies should be undertaken to ascertain its abundance and distribution. It seems to favour muddy habitats and may have a restricted distribution in the rocky streams of Fars on this account.

Further work

See above.

Sources

Type material: See above (NMW 48560).

Iranian material:  CMNFI  1979-0292, 1, 69.8 mm standard length, Fars, Lapui Spring in the Kor River basin (29º48'N, 52º39'E); IZA 7829, 4, 41.9-85.5 mm standard length, Fars, Pulvar River 15 km north of Persepolis (29º59'N, 52º59'E); IZA 7830, 1, 27.1 mm standard length, Fars, springs of Kul River near Darab (no other locality data).

Cobitis taenia
Linnaeus, 1758

Common names

mahi roshtegar or roftegar (= dustman, cleaner or sweeper fish, presumably from the bottom-dwelling habit); loch or louch in Khuzestan (presumably from the English name, but also louch means a person with a squint); gel khorak in Khuzestan; sagmahi-ye sangi (= stone loach), sagmahi-ye juibari or sagmahi-e-jooibari, mar mahi (= snake fish).

[lakh mukhattat in Arabic; zagafgaziya iliskani in Azerbaijan; shchipovka or shchipovka zakavkazskaya or Transcaucasian spiny loach in Russian; spined or spiny loach, stone loach, spiny stone loach, weatherfish, spotted weatherfish, Siberian loach].

Systematics

Cobitis Taenia was originally described from Lake Mälaren, Sweden (Kottelat, 1997).

Cobitis taenia satunini Gladkov, 1935 may be the subspecies of the southern Caspian Sea basin distinguished by a truncate snout rather than gradually tapering to the tip as in European spined loaches as well as other characters listed by Gladkov (1935) including head depth comprising 63.5% of head length on average and 12.27% of standard length, and caudal peduncle shorter than the head (on average 16.27% of standard (?) length). However, it was described from western Transcaucasia, outside the Caspian Sea basin, and material from the Caspian coast of Iran has not been examined and compared with it. Berg (1948-1949) referred specimens from the upper Kura River basin to this subspecies but lacked material from the lower Kura, Lenkoran and the southern Caspian Sea coast.

Kottelat and Freyhof (2007) consider that the taxon in the southern Caspian Sea basin is not taenia, a northern European species, and this is followed by Mousavi Sabet et al. (2011) who refer to it as Cobitis sp.

The holotype of Cobitis taenia satunini described from "Kavkaz" and "nizov'ya r. Kintrish" (= lower Kintrish stream in the Caucasus; the Kintrish stream mouth is at 41°48'N, 41°46'E) is in the Zoological Museum of Moscow State University under MMSU P.2852 with a cotype (= paratype) under MMSU P.2317 (Eschmeyer et al. (1996) has MMSU P.2251, quoting Svetovidova (1978), which disagrees with Gladkov (1935)). The Zoological Museum of Moscow University (ZMMU; their acronym) has P-2852 as the holotype plus P-2313 as 2 paratypes (Pavlinov and Borissenko, 2001).

Key characters

Distinguished from other Cobitis and related taxa in northern Iran by having modally 14 branched caudal fin rays, large dark and obvious spots along the mid-flank, and above them speckles which tend to form a stripe and above these a row of spots (absent in S. aurata).

Morphology

Dorsal fin unbranched rays 2-3, branched rays 6-8, anal fin unbranched rays 2-3, branched rays 4-7, usually 5, pectoral fin branched rays 5-9, and pelvic fin branched rays 5-7. Vertebrae 40-46, in the south Caspian Sea 40. Some specimens are elongate while others are deeper bodied. Chromosome number is 2n=48-50 (Klinkhardt et al., 1995) but triploids and tetraploids are also known.

Meristics for Iranian specimens:- dorsal fin branched rays 6(29) or 7(51); anal fin branched rays 4(1), 5(78) or 7(1); pectoral fin branched rays 6(4), 7(35), 8(38) or 9(3); pelvic fin branched rays 5(6), 6(70) or 7(4); caudal fin branched rays 10(1, but deformed), 12(2), 13(7), 14(68), 15(1) or 16(1); total vertebrae ?.

Sexual dimorphism

The second ray of the male pectoral fin is thickened and there is an enlarged scale at the base (Canestrini scale). The body in front of the dorsal fin of males is not distended as in S. aurata.

Colour

The back is light brown. The flanks are pale yellow with a series of 10-20, usually 16-18, dark brown blotches or spots and the back bears a more or less distinct series of 12-21 dark spots. A small stripe of brown spots on the upper flank lies between the 2 series of dark spots. The spaces between these stripes are finely speckled with brown. The belly and lower head are yellowish white. The head is mottled with brown dots and there is usually a band from the eye to the snout tip. The head can have lime-green iridescent tints. Females have yellow-orange dorsal and caudal fins bearing dark spots while in males these fins are almost orange with dark spots. There is a distinctive dark spot or two at the upper base of the caudal fin, often of a crescent shape, although this spot may be absent occasionally. Dorsal and caudal fins have 3-4 rows of brown spots. The pectoral, pelvic and anal fins are whitish without dark pigment. The iris is silvery, slightly golden or orange.

Size

Reaches 15.0 cm.

Distribution

Found from England and the Iberian Peninsula across Eurasia to Japan. However, Kottelat and Freyhof (2007) give a more restricted distribution and exclude the Caspian Sea basin of Iran in their map. In the Caspian Sea basin of Iran, it is found from Astara to Gorgan Bay including the Anzali Mordab, the Aras, Safid, Pol-e Rud, Sardab, Takar, Nessa, Haraz, Babol. Chowbar, Qasemabad, Qa'emshahr, Tonekabon, Shirud, Tajan and Gharasu rivers, and from the upper Karkheh, the middle Dez, Kashkan, Qareh Chai, Simarreh and lower Gav Masiab rivers in the Tigris basin (Saadati, 1977; Roshan Tabari, 1997; Abbasi et al., 1999; Kiabi et al., 1999; Abdoli, 2000; Abdoli and Naderi, 2009). An anecdotal report from the Qareh Chai near Hamadan in the Namak Lake basin needs specimens for confirmation.

Zoogeography

Records from the Tigris River basin of Iran are based on literature (Saadati, 1977) and a single specimen (CMNFI 1979-0285). Mid-flank spots are less distinct than in C. taenia from the Caspian Sea basin of Iran of similar size being smaller and more numerous and the stripe on the centre of the back is continuous rather than spots as in Caspian fish. Identification is tentative. Cobitis taenia survived glaciations in at least three refuges in the Ponto-Caspian area, one remaining near its Black Sea refuge. C. taenia lineages may have a late Pleistocene origin (Culling et al., 2006).

Habitat

This species remains buried in sand, mud which is not too thick, or dense weed growths during the day, being active at night, and is mostly solitary. Swimming is by undulating motions over short distances. When concealed, the body is bent into an arch so only the head and tail protrude. It prefers cool, clear running waters. Along the Caspian shore it is found in the lower reaches of rivers (Jolodar and Abdoli, 2004).

Age and growth

This loach lives up to 5 years and is mature in its second year of life. Females dominated the population in the Safid River and growth was positively allometric (Patimar et al., 2010). Mousavi Sabet et al. (2011) examined fish from the Babol River and found the percentage of females was significantly higher than males, and mature females were longer than 45 mm total length and 2+ years old and mature males were longer than 35 mm and 1+ years old. Life span was 5+ years for females and 3+ years for males.

Food

Diet is small crustaceans such as ostracods, copepods and rotifers in the bottom mud or sand. A mouthful of mud or sand is taken in, chewed, food items extracted, and the residue expelled convulsively through the gill openings.

Reproduction

Fecundity reaches 5072 eggs, perhaps as high as 10,000 (Palicka, 1996), and egg diameter reaches 1.58 mm. Spawning takes place from April to June in slow to still water. Eggs are laid on sand, stones and vegetation in several batches. Eggs may be deposited on the roots of water plants cleared of debris by males rooting among them. Males use their enlarged pectoral fins to turn the female during spawning.

The population in the Safid River had egg diameters up to 1.02 mm, absolute fecundity reached 8111 eggs and relative fecundity up to 383 eggs/g (Patimar et al., 2010). Mousavi Sabet et al. (2011) found Babol River fish spawning from the beginning of May to late July at water temperatures of 19.1-24.6ºC. Average absolute and relative fecundities were2172 and 590 eggs with ranges of 734-3562 and 347 to 945 eggs, respectively. Egg diameters were up to 1.4 mm with an average of 0.58 mm.

Parasites and predators

This species is infected with Clinostomum complanatum, a parasite that can cause laryngo-pharyngitis in humans, in the Shirud of western Mazandaran Province (Shamsi et al., 1997).

Economic importance

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in textbooks and in aquaria. Palicka (1996) gives a short account of its aquarium care.

Conservation

Lelek (1987) classifies this species as rare in Europe. It was commonly caught in Iranian streams along the Caspian coast. Kiabi et al. (1999) consider this species to be of least concern in the south Caspian Sea basin according to IUCN criteria. Criteria include abundant in numbers, habitat destruction, widespread range (75% of water bodies), present in other water bodies in Iran, and present outside the Caspian Sea basin.

Further work

The biology of this species in Iranian waters has not been studied.

Sources

Iranian material: 

Cobitis turcica
(Hanko, 1924)

Nalbant and Bianco (1998) record this species from the "River Kor near Persepolis" but I identify this material as C. linea (q.v.). The species was described originally from Eregli in Anatolian Turkey and its presence in Iran needs confirmation. Erkakan et al. (1999) review the Turkish species of Cobitis but do not mention the occurrence of C. turcica in Iran.

Misgurnus
Lacepède, 1803

Misgurnus anguillicaudatus
(Cantor, 1842)

Recorded from the Karakum Canal and Kopetdag Reservoir of Turkmenistan by Shakirova and Sukhanova (1994) and Sal'nikov (1995), this exotic species from China may eventually reach the Tedzhen River and Caspian Sea basins of Iran. No Iranian record.

Sabanejewia
Vladykov, 1929

Sabanejewia is distinguished from Cobitis Linnaeus, 1758 by having 12 rather than 14 branched caudal fin rays, a stronger suborbital spine, developed mental lobes which may be unfringed or well fringed, large imbricated or unimbricated scales with a relatively large and central focus, and males have a protuberance on each side of the body in front of the dorsal and pelvic fins and lack the lamina circularis (a bony process at the base of the second pectoral fin ray)(see Vladykov (1929), Nalbant (1963, 1994), Sawada (1982), Vasil'yeva (1995b) and Perdices and Doadrio (1997, 2001) for further details). Krupp (1985c) does not consider Sabanejewia to be a distinct genus. Perdices et al. ((2003) using mtDNA demonstrate that Caucasian-Caspian lineages are the sister group of a Danubian-Balkan lineage. Tang et al. (2008) using the cytochrome b gene found that ancestral Sabanejewia might have been the first cobitids to cross Siberia and invade the Euro-Mediterranean zoogeographic subregion.

Sabanejewia aurata
(De Filippi, 1863)

Further illustrations

Sabaneajewia aurata; Sabaneajewia aurata; both after Berg (1948-1949)

Common names

sagmahi-ye khardar (= spiny loach), sagmahi-e-jooibari.

[gizili iliskan in Azerbaijan; opi-dzug in Armenia; peredneaziatskaya shchipovka or Hither Asia spined loach, zolotistaya shchipovka or golden spined (spiny) loach, both in Russian].

Systematics

Kottelat (1997) tentatively considers that this species is restricted to Iran and possibly adjacent waters and does not occur in Europe.

The lectotype of Cobitis aurata (MZUT N.674), designated by Tortonese (1961), is stored in the Istituto e Museo di Zoologia della R. Università di Torino and 2 paralectotypes (MSNG N.365) from the Collezione di Universitá di Genova are in the Museo Civico di Storia Naturale di Genova (Tortonese, 1940; Tortonese, 1961). Eschmeyer et al. (1996) give the paralectotypes' catalogue number as ?MSNG 42727 (ex Univ. Genoa 365).

The type locality is "un fiumicello presso Sartschem" (De Filippi, 1863; not 1865 as in Berg (1948-1949), Banarescu and Nalbant (1964) and Reshetnikov et al. (1997) and not 1862 as in Tortonese (1940) and Eschmeyer et al. (1996)). The type locality is possibly Sarcham-e Sofla (37°07'N, 47°54'E) in the Qezel Owzan River drainage of the Caspian Sea basin in Iran.

Cobitis Hohenackeri Brandt in Kessler, 1877 from the Kura River basin of Transcaucasia (Azerbaijan) is a synonym with 6 syntypes in the Zoological Institute, St. Petersburg (ZISP).

Key characters

Distinguished from other cobitids in northern Iran by having modally 12 branched caudal fin rays, large dark spots along the flank, and above them speckles which do not tend to form a stripe.

Morphology

Dorsal fin unbranched rays 2-3, branched rays 5-8, predominately 6 in the Caspian basin, anal fin unbranched rays 2-3, branched rays 4-8, usually 5, pectoral fin branched rays 5-9, predominately 7 in the Caspian basin (but see below), pelvic fin branched rays 4-8, usually 5 (but see below), and vertebrae 39-43.

Meristics for Iranian specimens:- ? check dorsal rays on x-rays, dorsal fin branched rays 6(11) or 7(4); anal fin branched rays 5(15); pectoral fin branched rays 7(1) or 8(14); pelvic fin branched rays 5(1) or 6(14); caudal fin branched rays 11(2) or 12(13); and total vertebrae ?.

Scales minute but visible to the naked eye, ca. 170-200. Dermal crest or adipose fins are variably developed behind the dorsal and anal fins. Barbels are longer than in C. taenia, the mouth corner barbels reaching back to the posterior eye margin. Karyotype is 2n=50 (Klinkhardt et al., 1995).

Sexual dimorphism

The second pectoral fin ray in males is not enlarged as in C. taenia but there is a lateral distension of the body in front of the dorsal fin in mature males.

Colour

The back is a brownish olive-green with darker marbling. The flanks are golden-brown with a row of dark brown spots, less conspicuous than those in C. taenia. These number 9-18, modally 11-13, in the Caspian basin. The upper row of spots is absent in contrast to C. taenia. The number of spots may be related to habitat, those fish from calm waters having few large spots. The back has several dark blotches along its mid-line. In the Caspian basin there are 7-16 back blotches, modally 9-10. There is a bar at the caudal base, sometimes with a small central gap. Fins are slightly pink.

Size

Attains 13.8 cm, but most fish are less than 10.0 cm.

Distribution

Found in the basins of the Baltic, Aegean, Black and Caspian seas and in the Tedzhen and Murgab rivers of Afghanistan and Turkmenistan according to most authors (Aliev et al., 1988; Nalbant and Bianco, 1998). A distinct subspecies is found in the Aral Sea basin (see below). Kottelat (1997) however considers that this species is restricted to Iran and possibly adjacent waters. It is found along the Caspian Sea coast of Iran including the Anzali Mordab and its tributaries, the Safid River at Kisom, lower Tonekabon, Chalus, Haraz and Babol rivers, Nakhurde (= ? Noqreh Deh), and in the Kashaf River of the Tedzhen River basin (Holčik and Oláh, 1992; Abbasi et al., 1999; Abdoli, 2000; Jolodar and Abdoli, 2004; Abdoli and Naderi, 2009).

The record of this species in the Tigris-Euphrates basin at Basrah, Iraq (BM(NH) 1920.3.5:9) in Nalbant (1963) and Bănărescu and Nalbant (1966), and repeated in Banister (1980), is probably an error of labeling (Bănărescu, 1973).

Zoogeography

The subspecies, Cobitis aurata aralensis Kessler, 1877, is reported from the Karakum Canal and the Uzboi lakes in Turkmenistan by Shakirova and Sukhanova (1994) and Sal'nikov (1995) and may well enter both the Tedzhen River and Caspian Sea basins of Iran eventually.

Habitat

This is a nocturnal species, hiding during the day under gravel and boulders of flowing rivers. If exposed, it will make jerking motions to the nearest cover. Some authors state that it also hides in sand. It prefers shallow and clear water. It occurs with C. taenia but is commoner in faster water in the upper and middle reaches of rivers, from 5 to 150 cm water depth. However it may form populations in still water left behind after floods.

Age and growth

Males tend to be slightly smaller than females.

Food

Bottom-dwelling invertebrates are the main food items including larval insects such as mayflies, dragonflies and caddisflies as well as nematodes, copepods, chironomids, fish eggs, algae and detritus.

Reproduction

Fecundity reaches 14,700 eggs and egg diameter 0.85 mm. Eggs are shed over plants from April to August and this species may spawn in batches.

Parasites and predators

Unknown.

Economic importance

None.

Conservation

Lelek (1987) lists this species as rare to vulnerable in Europe. Kiabi et al. (1999) consider this species to be conservation dependent in the south Caspian Sea basin according to IUCN criteria. Criteria include few in numbers, habitat destruction, limited range (less than 25% of water bodies), absent in other water bodies in Iran, and present outside the Caspian Sea basin. The 2000 IUCN Red List lists this species as DD (Data Deficient).

Further work

The biology of this species in Iran has not been examined and its systematics needs clarification.

Sources

Vasil'eva and Vasil'ev (1988) give details on variation in meristics and colour of this species.

Iranian material: CMNFI 1970-0508, 4, 39.2-57.1 mm  standard length; Gilan, Safid River at Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1970-0545, 1, 33.9 mm standard length, Gilan, Safid River (ca. 37º01'N, ca. 49º38'E), CMNFI 1979-0448, 1, 70.1 mm standard length, Azarbayjan-e Khavari, Ahar Chay (38º18'30"N, 48º22'E); CMNFI 1980-0131, 1, 57.4 mm standard length, Iran, Caspian Sea basin (no other locality data); CMNFI 1980-0132, 1, 49.8 mm standard length, Gilan, Safid River at Kisom (37º12'N, 49º54'E); CMNFI 1980-0155, 7, 38.5-63.5 mm standard length, Azarbayjan-e Khavari, Qareh Su at Ardabil (ca. 38º15'N, ca. 48º18'E).

Sabanejewia caspia
(Eichwald, 1838)

Further illustrations

Sabanejewia caspia; Sabanejewia caspia and suborbital spine; Sabanejewia caspia; all after Berg (1948-1949)

Common names

mahi roshtegar talaee (= golden dustman fish), rofteghar mahi, sagmahi-ye Khazari (= Caspian loach), sag mahi khardar (= spiny loach), mar mahi (= snake fish), sagmahi-e-jooibari.

[xazar iliskani in Azerbaijan; Kaspiiskaya shchipovka or Caspian spined (spiny) loach in Russian].

Systematics

This species was described from "in sinu mardofiensi prope castellum Lencoranicum" (i.e. the Caspian Sea in the ? Mardofiensi Gulf near the Lenkoran fortress). The type specimen is apparently lost (Vasil'yeva, 1995b).

Key characters

Distinguished from other cobitids in northern Iran by having modally 12 branched caudal fin rays, no large dark spots along the flank but a stripe, and above this stripe speckles which do not tend to form a stripe.

Morphology

Dorsal fin unbranched rays 2-3, branched rays 6-7, anal fin unbranched rays 1-2, branched rays 5-6, pectoral fin branched rays 6-7 (but see below) and pelvic fin branched rays 4-6. Vertebrae 41-42. A crest is well developed on the lower caudal peduncle but only posteriorly on the upper edge. Barbels are shorter than in C. taenia, the posterior ones reaching beyond the posterior eye margin. Karyotype is 2n=50 (Klinkhardt et al., 1995).

Meristics for Iranian specimens:- dorsal fin branched rays 6(83) or 7(1); anal fin branched rays 5(80) or 6(4); pectoral fin branched rays 7(6), 8(72) or 9(6); pelvic fin branched rays 4(3), 5(67) or 6(14); caudal fin branched rays 7(1, but deformed), 9(1), 10(3), 11(4), 12(74) or 13(1); total vertebrae ?.

Sexual dimorphism

The second pectoral fin ray in males is not enlarged as in C. taenia but there is a lateral distension of the body in front of the dorsal fin an mature males.

Colour

Live specimens, especially young, are almost transparent. The adult has a pronounced dark line along mid-flank indicating the separation of the upper and lower muscle masses. Above this line the upper flank is yellowish with irregular dark grey pigment or brown speckles near the back. The lower flank has irregular grey pigment or brown speckles. The mid-line of the back has a more or less pronounced dark line. The belly and lower head surface are pale yellowish without grey pigment. The dorsal and caudal fins have yellow-orange rays bearing 3-4 series of dark grey spots. The base of the caudal fin has 2 dark dots although these are not as marked as in Cobitis taenia. The pectoral, pelvic and anal fins are transparent though larger fish may have elongate grey spots along the rays. The iris is golden.

Note that Vasil'yeva (1995b), based on original data and that of Derzhavin (1934), found that the dark longitudinal band and the dark caudal fin base may not be pronounced.

Size

Reaches 9.2 cm standard length.

Distribution

Principally found in the southern Caspian Sea basin. Records from the northern Caspian Sea are apparently in error and the range is from the Kura to the Babol rivers including the Anzali Mordab at "Khalkai" for example, the Safid, Chowbar, Tonekabon, Chalus, Shahzadeh, "Laidschana", Meshedessera (= ? Babol Sar), Haraz, (Holčik and Oláh, 1992; Vasil'yeva, 1995b; Abbasi et al., 1999; Abdoli, 2000; Jolodar and Abdoli, 2004; Abdoli and Naderi, 2009).

Zoogeography

This species is endemic to the Caspian Sea basin (Nalbant and Bianco, 1998).

Habitat

This species is found in both fresh and brackish waters, in slow or still water with aquatic vegetation, in the lower reaches of rivers and near river mouths, and in brackish bays.

Age and growth

Unnown.

Food

Unknown.

Reproduction

Spawning takes place in April in the brackish lagoons of the Lenkoran, Azerbaijan. Up to 955 eggs of up to 0.85 mm diameter are carried by each female.

Parasites and predators

None reported for Iran.

Economic importance

None.

Conservation

Holčík and Oláh (1992) report the loss of this species from the Anzali Mordab where it was once found. Reasons for this loss are unknown. Kiabi et al. (1999) consider this species to be conservation dependent in the south Caspian Sea basin according to IUCN criteria. Criteria include possibly few in numbers, habitat destruction, limited range (less than 25% of water bodies), absent in other water bodies in Iran, and absent outside the Caspian Sea basin. Vulnerable in Turkey (Fricke et al., 2007).

Further work

The biology of this species in Iran requires study.

Sources

Iranian material: ?

Sabanejewia caucasica
(Berg, 1906)

? to write up

Reported from the Anzali Mordab and lower reaches of the Safid, Toenkabon, Chalus, Heraz and Babol rivers in Iran (Abdoli, 2000) and mapped from the Caspian coast of Iran at Babol by Kottelat and Freyhof (2007). Berg (1948-1949) notes that this species is found in the northern Transcaucasus and is not reported from the Kura River in the southwestern corner of the Caspian Sea. Its presence in Iran needs confirmation by specimens. Formerly in the genus Cobitis.

Nemacheilidae

Nemacheilidae, paracobitis, barbatula, schistura, triplophysa genus descriptions ? and search barbatula etc and change as needed to new genus

Materials to ID and add to Sources

Nemacheilid spp.

CMNFI 1970-0521, 1, ?, mm standard length, Gilan, Safid River near Lulaman (no other locality data) ?bergiana; CMNFI 1970-0545, 2, 34.7-35.6 mm standard length, Gilan, Safid River from Emamzadeh Hashem to Lulaman (ca. 37º01'N, ca. 49º38'E) ?bergiana; CMNFI 1970-0540, 1, 49.5 mm stanmdard length, ?; CMNFI 1970-0558, 4, ?mm standard length, Ghasemlou Chay ()?brandtii; CMNFI 1970-0559, 3, ?, mm standard length, Azarbaijan-e Bakhtari, Barunduz Chay (ca. 37º25'N, ca. 45º10'E); CMNFI 1970-0560, 3, ?, mm standard length, Azarbaijan-e Bakhtari, Mamiyand Chay near Mamiyand (ca. 36º59'N, ca. 45º39'E); CMNFI 1971-0341, 13, 28.6-44.3 mm standard length, ? ; CMNFI 1979-0026, 8, 22.8-46.6 mm standard length, Fars, Shapur River at Shapur (29º47'N, 51º35'E); CMNFI 1979-0027, 9, 31.0-41.3 mm standard length, Fars, Chehel Chashmeh (ca. 29º43'N, ca. 52º04'E); CMNFI 1979-0028, 5, 48.3-55.6 mm standard length, Fars, ?; CMNFI 1979-0059, 28, 32.8-57.0 mm standard length, Fars, Pulvar River 8 km south of Sivand (30º01'30"N, 52º57'E)?distinct taxonand se other fars fish; CMNFI 1979-0061, 3, 50.2-63.7 mm standard length, Fars, stream tributary to Pulvar River (30º04'N, 53º01'E); CMNFI 1979-0067, 1, 55.8 mm standard length, Fars, qanat at Zarqan (ca. 29º46'N, ca. 52º43'E); CMNFI 1979-0070, 2, 38.8-39.9 mm standard length, Fars, Pulvar River at Naqsh-e Rostam (29º59'N, 52º54'E); CMNFI 1979-0073, 10, 31.1-56.1 mm standard length, Fars, Mand River (ca. 29º42'30"N, ca. 52º01'30"E); CMNFI 1979-0111, 5, 43.2-52.7 mm standard length, Fars, stream on Shiraz to Bushehr road (29º37'30"N, 52º21'E); CMNFI 1979-0117, 2, 42.1-50.6 mm standard length, Fars, Pulvar River on road to Naqsh-e Rostam (29º59'N, 52º54'E); CMNFI 1979-0155, 2, 30.7-30.9 mm standard length, Fars, spring at Gavanoo (28º47'N, 54º22'E); CMNFI 1979-0157, 3, 34.8-37.7 mm standard length, Fars, qanat at Hadiabad (28º52'N, 54º13'E); CMNFI 1979-0167, 41, 20.6-48.3 mm standard length, Kerman, qanat at Bam (29º06'N, 58º20'E); CMNFI 1979-0168, 1, 47.2 mm standard length, Kerman, qanat at Shahabad (29º07'N, 58º16'E); CMNFI 1979-0169, 11, ? mm standard length, Kerman, qanat 10 km from Mahan (30º08'30"N, 57º17'E); CMNFI 1979-0170, 1, 45.0 mm standard length, Kerman, qanat at Baghin (30º12'N, 56º48'E); CMNFI 1979-0172, 18, 33.8-45.8 mm standard length, Kerman, qanat on Kerman to Bandar Abbas road (29º51'N, 56º14'E); CMNFI 1979-0184, 2, 27.7-28.4 mm satndard length, ?; CMNFI 1979-0186, 2, ? mm standard length, Hormozgan, stream at Sar Khun (ca. 27º24'30"N, ca. 56º25'E); CMNFI 1979-0192, 7, ? mm standard length, Fars, qanat 2 km east of Rostaq (28º26'30"N, 55º04'E); CMNFI 1979-0193, 1, ? mm standard length, Fars, river 8 km from Darab (28º45'N, 54º27'30"E); CMNFI 1979-0194, 2, 37.6-45.6 mm standard length, Fars, jube 15 km from Darab (28º45'30"N, 54º24'E); CMNFI 1979-0206, 3, 24.9-36.1 mm standard length, Fars, qanat 1 km from Runiz-e Pa'in (29º12'N, 53º40'E); CMNFI 1979-0208, 15, 31.1-58.8 mm standard length, Fars, qanat on road to Qatru (ca. 29º11'N, ca. 54º40'E); CMNFI 1979-0213, 7, ? mm standard length, Kerman, stream in Kharan River drainage (29º15'N, 56º25'E); CMNFI 1979-0219, 13, ? mm standard length, Kerman, jube 14 km west of Jiroft (28º37'N, 57º41'E); CMNFI 1979-0253, 7, 31.2-49.5 mm standard length, Markazi, Qareh Chay west of Baqerabad (34º52'N, 50º49'E); CMNFI 1979-0276, 14, 36.3-45.8 mm standard length, Lorestan, Chamesk River (ca. 33º19'N, ca. 47º53'30"E); CMNFI 1979-0284, 3, 38.7-41.9 mm standard length, Kermanshahan, Marek River at Mahidasht (34º16'N, 46º48'30"E); CMNFI 1979-0288, 1, 32.4 mm standard length, Ilam and Poshtkuh, Gangir River at Juy-e Zar Eivan (33º50'N, 46º18'E); CMNFI 1979-0292, 8, 29.0-45.6 mm standard length, Fars, Lapu'i spring (29º48'N, 52º39'E) ?distinct taxon; CMNFI 1979-0306, 11, 20.0-49.3 mm standard length, Kerman, qanat 33 km from Sirjan (29º13'N, 54º33'E); CMNFI 1979-0307, 4, 31.0-42.3 mm standard length, Kerman, river at Sartal (ca. 29º17'N, ca. 56º38'E)? distinct species; CMNFI 1979-0316, 1, ? mm standard length, Baluchestan, stream on road to Chah Bahar (26º48'N, 61º02'E); CMNFI 1979-0341, 10, 25.5-45.6 mm standard length, Kerman, Tahrud west of Bam (29º23'N, 57º52'E); CMNFI 1979-0365, 7, 32.0-40.5 mm standard length, Khuzestan, stream in Doveyrich drainage (32º25"N, 47º36'30"E); CMNFI 1979-0366, 1, 36.0 mm standard length, Khuzestan, stream 17 km west of Dehloran (32º45'30"N, 47º05'30"E); CMNFI 1979-0367, 1, 41.3 mm standard length, Khuzestan, Meymeh River 11 km north of Dehloran (32º44'30"N, 47º09'30"E); CMNFI 1979-0371, 2, 44.5-55.6 mm standard length, Khuzestan, stream in Karkheh River drainage (32º05'N, 48º19'E); CMNFI 1979-0374, 2, 41.3-41.7 mm standard length, Khuzestan, stream tributary to Bala River (32º40'N, 48º15'E); CMNFI 1979-0389, 1, 32.4 mm standard length, Khuzestan, Zard River 1 km south of Bagh-e Malek (31º31'N, 49º53'30"E); CMNFI 1979-0390B, 3, 29.1-39.4 mm standard length, Khuzestan, stream 3km south of Bagh-e Malek (31º29'N, 49º54'30"E); CMNFI 1979-0395, 1, 30.7 mm standard length, Khuzestan, stream in Marun River drainage (ca. 30º57'N, ca. 49º51'E); CMNFI 1979-0399, 1, 26.2 mm standard length, Fars, stream near Basht (30º19'30"N, 51º15'E); CMNFI 1979-0411, 3, 21.9-25.9 mm standard length, Hormozgan, Minab River near Rudan (27º24'N, 57º12'E)?cf bampurensis; CMNFI 1979-0419, 19, 32.4-58.7 mm standard length, Fars, stream 7 km from Rostaq (28º29'N, 55º01'E); CMNFI 1979-0423, 4, 43.2-45.7 mm standard length, Boyer Ahmadi-ye Sardsir va Kohkiluyeh-Fars border, river in Beshar River drainage (30º31'N, 51º31'E); CMNFI 1979-0447, 2, 51.2-59.8 mm standard length, Gilan, stream 7 km east of Namin (38º23'N, 48º28'E); CMNFI 1979-0448, 9, 31.1-62.0 mm standard length, Azarbayjan-e Khavari, Ahar Chay 8 km from Ardebil (38º18'30N, 48º22'E); CMNFI 1979-0450, 3, 38.7-40.9 mm standard length, Azarbayjan-e Khavari, stream near Kivi (ca. 37º37'N, ca. 48º17'E); CMNFI 1979-0452, 2, 42.1-42.8 mm standard length, Azarbayjan-e Khavari, Qezel Owzan River 6 km from Mianeh (37º23'N, 47º45'E); CMNFI 1979-0453, 1, 34.2 mm standard length, Zanjan, Zanjan River (37º06'N, 47º56'E); CMNFI 1979-0458, 1, 48.2 mm standard length, Markazi, Khar River 6 km north of Ab-Garm (35º47'N, 49º20'E); CMNFI 1979-0459, 4, ? mm standard length, Hamadan, stream 2 km south of Kazan (35º22'N, 49º02'E); CMNFI 1979-0462, 4, 32.0-45.6 mm standard length, Markazi, Mazdaqan River (35º06'30"N, 49º40'30"E); CMNFI 1979-0463, 2, 41.0-51.6 mm standard length, Markazi, Qareh Chay (34º53'N, 50º26'E); CMNFI 1979-0465, 1, 37.1 mm standard length, Markazi, Qom River at Neizar (34º18'30"N, 50º32'E); CMNFI 1991-0155, 1, 41.7 mm standard length, Hamadan, Gav Masiab River (34º12'N, 48º20'E); CMNFI 1991-0156, 1, ? mm standard length, Hamadan, Gav Masiab River (34º16'N, 48º10'E); CMNFI 1993-0125, 1, 47.8 mm standard length, Kermanshahan, Sarab-e Nilufar (34º24'N, 46º52'E); CMNFI 1993-0128, 1, 58.3 mm standard length, Kermanshahan, Sarab-e Sabz 'Ali Khan (34º25'N, 46º32'E) ?kermanshahensis q.v.; CMNFI 2007-0021, 11, 25.5-34.6 mm standard length, ?; CMNFI 2007-0037, 5, 45.2-54.9 mm standard length, Kerman, Hosseinabad and Gamatabad qanats at Bam (29º06'N, 58º21'E); CMNFI 2007-0038, 1, 39.0 mm standard length, Kerman, Mehtiabad qanat at Bam (29º06'N, 58º21'E); CMNFI 2007-0039, 6, 32.4-41.7 mm standard length, Kerman, Tahrud River (ca. 29º23'N, ca. 57º63'E); CMNFI 2007-0043, 9, 33.0-56.7 mm standard length, Kerman, qanat at Emamzadeh Sultan (ca. 29º40'N, ca. 56º45'E); CMNFI 2007-0047, 2, 36.8-58.2 mm standard length, Kerman, qanat at Hoshum (29º14'N, 56º19'E); CMNFI 2007-0051, 5, 28.6-36.3 mm standard length, Hormozgan, upper Kol River basin (28º19'N, 55º55'E); CMNFI 2007-0055, 3, 36.1-40.3 mm standard length, Hormozgan, stream in Minab River basin (27º47'N, 57º12'E); CMNFI 2007-0074, 10, 25.6-50.8 mm standard length, Markazi, Qareh Su 32 km west of Arak (34º03'N, 49º21'E); CMNFI 2007-0075, 6, 41.3-47.9 mm standard length, Hamadan, Malayer River 5 km from Malayer (ca. 34º17'N, ca. 48º47'E); CMNFI 2007-0081, 10, ? mm standard length, Zanjan, Zanjan River basin near Soltaniyeh (ca. 36º27'N, ca. 48º45'E); CMNFI 2007-0082, 7, ? mm standard length, Zanjan, Zanjan River basin near Zanjan (ca. 36º36'N, ca. 48º32'E); CMNFI 2007-0083, 2, ? mm standard length, Azarbayjan-e Khavari, Qaranqu River basin west of Sar Eskand Khan (ca. 37º25'N, ca. 46º55'E); CMNFI 2007-0084, 2, ? mm standard length, Azarbayjan-e Khavari, Talkheh River basin west of Sarab (ca. 37º56'N, ca. 47º19'E); CMNFI 2007-0085, 12, ? mm standard length, Azarbayjan-e Khavari, Talkheh River basin east of Sarab (ca. 37º56'N, ca. 47º41'E); CMNFI 2007-0086, 3, ? mm standard length, Azarbayjan-e Khavari, Qareh Su basin near Nir (ca. 38º02'N, ca. 48º00'E); CMNFI 2007-0089, 7, ? mm standard length, Azarbayjan-e Khavari, Ahar Chay at Ahar (38º28'N, 47º03'E); CMNFI 2007-0091, 13, ? mm standard length, Azarbayjan-e Khavari, Zilber Chay west of Marand (38º30'N, 45º23'E); CMNFI 2007-0093, 8, ? mm standard length, Azarbayjan-e Bakhtari, Qotur River south of Khvoy (38º30'N, 44º58'E); CMNFI 2007-0095, 2, ? mm standard length, Azarbayjan-e Bakhtari, Shahr Chay southwest of Reza'iyeh (ca. 37º27'N, ca. 44º56'E); CMNFI 2007-0096, 6, ? mm standard length, Azarbayjan-e Bakhtari, Qasemul River south of Reza'iyeh (ca. 37º25'N, ca. 45º10'E); CMNFI 2007-0097, 4, ? mm standard length, Azarbayjan-e Bakhtari, Baranduz Chay basin south of Reza'iyeh (ca. 37º16'N, ca. 45º08'E); CMNFI 2007-0099, 9, 24.6-43.8 mm standard length, Azarbayjan-e Bakhtari, Kalwi Chay west of Mahabad (ca. 36º35'N, ca. 45º25'E); CMNFI 2007-0100, 4, 44.2-58.4 mm standard length, Azarbayjan-e Bakhtari, Kalwi Chay near Piranshahr (ca. 36º44'N, ca. 45º10'E); CMNFI 2007-0103, 3, ? mm standard length, Kordestan, Zarineh River basin north of Saqqez (36º18'N, ca. 46º16'E); CMNFI 2007-0105, 13, ? mm standard length, Kordestan, Zarineh River basin south of Saqqez (ca. 36º06'N, ca. 46º20'E); CMNFI 2007-0106, 17, ? mm standard length, Kordestan, Qezel Owzan River basin near Divandarreh (ca. 35º52'N, ca. 47º05'E); CMNFI 2007-0107, 8, ? mm standard length, Kordestan, Qezel Owzan River basin near Bijar (ca. 35º54'N, ca. 47º20'E); CMNFI 2007-0115, 1, 41.5 mm standard length, Kermanshahan, Qareh Su basin (ca. 34º34'N, ca. 46º47'E); CMNFI 2007-0115, 2, 49.7-54.7 mm standard length, Kermanshahan, Qareh Su basin (ca. 34º34'N, ca. 46º47'E); CMNFI 2007-0116, 2, 40.6-44.8 mm standard length, Kermanshahan, Gav Masiab River basin west of Sahneh (ca. 34º28;'N, ca. 47º36'E); CMNFI 2007-0118, 15, 24.7-58.8 mm standard length, Kermanshahan, Bid Sorkh River between Sahneh and Kangavar (ca. 34º23'N, ca. 47º52'E); CMNFI 2007-0119, 8, 26.9-37.9 mm standard length, Kermanshahan, Gav Masiab River basin near Kangavar (ca. 34º31'N, ca. 48º03'E); CMNFI 2007-0121, 6, 32.0-54.1 mm standard length, Hamadan, Qareh Su basin north of Razan (ca. 35º25'N, ca. 49º02'E); BWC 1995-0002, 1, 41.6 mm standard length, ?; BWC 1995-0017, 3, 48.4-51.2 mm standard length, ?; BWC 1995-0020, BWC 1995-0030, 1, 22.8 mm standard length, ?; BWC 1997-0005, BWC 2000-0001, BWC 2000-0002, BWC 2000-0004, BWC 2000-0008, BWC 2000-0011, BWC 2000-0016, BM(NH) 1899.9.30:138-140, 4, 41.8-53.4 mm standard length, ?, Ula on the Zola Chai BM(NH) 1899.9.30:131-137, 7 (4 examined), 33.3-35.8 mm standard length, ?, Elinja Chai

The hillstream, mountain or river loach family has had various family names - see Menon (1987), Mirza (1989b), Hieronimus (1990; 1991), Ng and Lim (1991), Kottelat (1988, 1991), Nelson (1991), Thorne (1992), Anonymous (1993a) and Bănărescu and Nalbant (1995) for discussions. The Iranian species were classified in Cobitidae in older works and more recently in Homalopteridae and Balitoridae. Mirza (1989b), Nalbant (1998), Tang et al. (2006) and Šlechtová et al. (2007) consider that the correct name for this family is Noemacheilidae (in Mirza) or Nemacheilidae. Nemacheilids are aligned with the main stem of cobitoid fishes rather than homalopterids as proposed by Sawada (1982) on osteological grounds. Nalbant (1998) considers the similarities observed by Sawada between nemacheilids and homalopterids to be homoplasies. Homalopterids are more closely related to cyprinids and psilorhynchids.

The family is found throughout Eurasia with a single species in northeast Africa. There are about 60 genera and about 590 species (Berra, 2001; Nelson, 2006), with more being described regularly. Iranian species belong to the subfamily Nemacheilinae.

Conway et al. (2010) point out that although there are no morphological synapomorphies for the family, molecular data does support monophyly. Prokofiev (2010) gives osteological characters for the subfamily Nemacheilinae.

Most species in Iran were placed in the genus Nemacheilus Bleeker, 1863 but this name contains only species from Southeast Asia (Kottelat, 1997). Noemacheilus Kuhl and van Hasselt in van Hasselt, 1823 is a nomen nudum since there are no taxonomic characters accompanying the original description. The next available name is Nemacheilus Bleeker, 1863 (see Kottelat, 1987). Nemachilus Günther, 1868 is an incorrect spelling. Much of the earlier literature on Iranian members can be found under the name Nemacheilus or its variant spellings.

These loaches have been placed in several genera or subgenera including the following relevant to Iran: Orthrias Jordan and Fowler, 1903 (see Banarescu, Nalbant and Balik (1978); Orthrias = Barbatula Linck, 1790 of authors, see Bănărescu and Nalbant (1995) for reasons advocating the later name over the earlier one; Kottelat (1997) gives Linck as 1789), Adiposia Annandale and Hora, 1920 (see Annandale and Hora (1920); see also Bănărescu and Nalbant (1995) where it is synonymised with Paracobitis), Triplophysa Rendahl, 1933 and Hedinichthys Rendahl, 1933 (see Bănărescu and Nalbant (1966); see also Bănărescu and Nalbant (1995) where the latter is synonymised with the former), Oxynoemacheilus Bănărescu and Nalbant, 1966 (see Bănărescu and Nalbant, 1966; synonymised with Orthrias in Bănărescu and Nalbant (1995) and with Barbatula in Bogutskaya and Naseka (2004)), Paracobitis Bleeker, 1863 (see Bănărescu and Nalbant, 1966), Schistura McClelland, 1839 (see Mirza, Nalbant and Banarescu, 1981; regarded as polyphyletic by Bănărescu and Nalbant (1995)), and Seminemacheilus Bănărescu and Nalbant, 1995 (q.v.). Views on the generic validity of these names conflict between authors and between the same author at different dates (see Krupp, 1985c; Eschmeyer, 1990; Eschmeyer et al., 1996; Eschmeyer's "Catalog of Fishes"). Kottelat (1984) retained Nemacheilus until a revision of all species was complete but there has been a tendency to use the above genera and this is followed here. Prokofiev (2004, 2007, 2009, 2010) has revised some of the Nemacheilidae, in particular the group of nemacheiline loaches that lack the preethmoid I in the skull, using osteology and morphology. This includes most of the species found in Iran and is now being followed by such authors as Esmaeili et al. (2010).

The table below summarises some of the various allocations:-

?re-do

Species

Original genus^

Other generic allocations^

Current genus

angorae

Nemacheilus

Orthrias, Barbatula

Oxynoemacheilus

araxensis

?

?

Oxynoemacheilus

bampurensis

Nemacheilus

Schistura

Paraschistura

bergianus*check original ending

Nemachilus

Orthrias, Barbatula

Oxynoemacheilus

brandtii

Nemachilus

Orthrias Barbatula

Oxynoemacheilus

cristatus*check ending

Nemacheilus

Paracobitis, Schistura

Metaschistura

farsicus ?original spelling

?

 

Oxynoemacheilus

frenatus*check ending

Cobitis

Nemacheilus, Orthrias, Barbatula

 

iranica*

Paracobitis

-

Paracobitis

kermanshahensis

Noemacheilus

Orthrias, Barbatula

Oxynoemacheilus

kessleri

Nemachilus

-

Paraschistura

kosswigi

Turcinoemacheilus

-

Turcinoemacheilus

longicauda*check ending

Cobitis

Nemacheilus, Adiposia

Paracobitis

longipinnis

Ilamnemacheilus

-

Ilamnemacheilus

malapterura*check ending

Cobitis

Nemacheilus

Paracobitis

nielseni

Schistura

Schistura

Paraschistura

persa*check ending

Cobitis

Nemacheilus, Oxynoemacheilus, Orthrias, Barbatula

Oxynoemacheilus

rhadinea*check ending

Nemachilus

Adiposia

Paracobitis

sargadensis

Nemacheilus

Schistura

Paraschistura

smithi

Noemacheilus

-

Paracobitis

stoliczkai

Cobitis

Nemacheilus

Triplophysa

tongiorgii

Seminemacheilus

-

Seminemacheilus

vignai

Paracobitis

-

Paracobitis

* original suffix which may change if genus changes to masculine or to feminine; ^ Nemacheilus sometimes spelled Noemacheilus or Nemachilus by various authors

A number of nemacheilid species have been described from waters confluent with Iran, particularly from the Helmand River basin in Afghanistan. They have no Iranian records but are listed here as they may be relevant to revisionary studies.

The dating of the paper by Bănărescu and Nalbant as 1967 in various works may follow Nalbant and Bianco (1998). The Bănărescu and Nalbant paper states in Danish on page 186 "Reprints released the 31 December 1966" and this is presumably the correct date, as Art. 21.2 of the Code of Zoological Nomenclature states "Date specified. The date of publication specified in a work is to be adopted as correct in the absence of evidence to the contrary", and no evidence to the contrary has been presented (N. G. Bogutskaya, pers comm., 28 April 2011). This changing of dates here has no nomenclatural significance.

Members of this family in Iran are characterised by an elongate and weakly compressed and almost cylindrical body, head not compressed but rounded, scaleless or body covered in minute scales (too small for scale counts to be commonly or easily made), lateral line complete or incomplete, a small and inferior mouth, lips thick, fleshy and papillose, lower lip interrupted in the middle, 2 pairs of barbels on the snout and 1 pair at the mouth corners (8 pairs in some non-Iranian species), no collapsible spine under the eye (sometimes present in non-Iranian species but distinguishes members of the related Cobitidae in Iran), eyes small to minute, usually not visible from the underside of the head, reduced gill opening, short to moderate dorsal fin without spines, short anal fins, vent a short distance in front of anal fin origin, swimbladder enclosed partially or entirely in a bony capsule, certain osteological characters such as the shape of bones in the Weberian apparatus used in sound transmission from the swimbladder to the ear, gut short or long, a dermal crest or adipose fin may be present, caudal fin truncate, rounded or slightly forked, and often distinctive colour patterns of bars, stripes and blotches. Iranian species may lack scales, may have an adipose fin, and have a single unbranched ray leading the pectoral and pelvic fins.

Krupp (1985c) reviews prior works by P. M. Bănărescu and co-authors on Levantine nemacheilids and regards them as unsatisfactory. This calls into question works on Iranian species by this author. Krupp (1985c) lists characters important in studying Nemacheilus sensu lato and those which are individually variable or develop independently, much in contrast to characters favoured by Bănărescu. Morphometric characters can vary with nutritional status and ecological factors. Stable characters were head length, interorbital width, caudal peduncle length and depth, and predorsal length. Fin lengths are dependent on sex in some species, less so in others. Allometry is a problem in fin positions and measurements involving such characters can only be used when comparing fish of equal size. Mouth width and digestive tract shape are good characters but lip shape and development of the processus dentiformis are not. The swimbladder capsule form, including the presence or absence of a continuous collar between the two hemispheres and the shape of laminae, is an important character. Reduced laminae and wide recesses on the hemispheres are derived characters. The dorsal adipose fin development is stable in some species, variable in others, and is independently derived in different phyletic lines, thus being of limited value. The shortening and deepening of the caudal peduncle is derived in one Levantine species. Scale characters such as size and position of the focus and general scale structure are very variable and not characters easily quantified. Only specimens of the same size are comparable and numerous scales must be examined because aberrant ones are common. The lateral line length is a good character, although juvenile fish may have a shortened one. A reduced lateral line is a derived character. Colour patterns are subject to variation and both spotted and striped forms can be found within one species. Nevertheless, patterns can be important in distinguishing species. Thickening of pectoral fin rays is a derived feature but absence of this character is a symplesiomorph condition and cannot be used to relate species.

The species in this family are often difficult to identify and many literature reports are undoubtedly mis-identifications. While some of these may be corrected based on material deposited in museums, others have no voucher material and cannot be re-identified. Identification is problematical because scale counts are not available (too minute), fin ray counts are often very similar, and unique structures uncommon. Colour patterns can be used but are notoriously variable and many types in museums are decoloured making comparisons difficult. Morphometric characters require good series of adult fish of both sexes, from various localities in the species range, preferably even from the same locality taken over several years to allow for local variations in habitat which may conceivably affect shape. Menon (1987) considers that many species in this genus are from very similar habitats, the stressful one of running water, and have been constantly selected to fit this niche. Valid species resemble one another closely. Such characters as position of the anal opening, the dorsal fin origin and barbel length have been used in species definitions but Menon (1987) found these to vary with growth. Swimbladder structure depends on the habitat where the fish live and scale coverage on the physico-chemical nature of the water. Menon (1987) found lateral line character, number of branched dorsal fin rays, caudal fin shape, secondary sexual characters in males and, despite the above, body colour and anal opening position to be useful. Prokofiev (2010) reviewed the morphological classification of loaches and points out that changes in morphology occur with growth and species with wide ranges show large variations in morphology associated with the various biotopes. He gives extensive reviews of characters and their importance in defining genera and species, e.g. scale cover and scale characters are not considered important in phylogeny and are of only accessory import for genus and species diagnosis; colour, sexual dimorphism, adipose keel presence and extent, fin ray numbers, head and body shapes, abdominal axillary lobe presence and size, variations in the seismosensory system, nostril size and position, anal opening position, intestine shape, swimbladder and its capsule structure, and general osteology are all important characters.

Examination of Iranian species, where good series of fresh material was available, tend to confirm the observations of Krupp and Menon on characters. Position of the dorsal fin origin is variable within a species among morphometric characters used as distinctive, extent of the lateral line is also variable, there is marked sexual dimorphism, and colour patterns can be useful but also vary with the individual, the habitat and the temperature (fish kept in ice water have strong colour patterns while those preserved immediately from murky waters have faint patterns).

Two main problems exist in identifying certain Iranian loaches. These are determining appropriate characters which are not individually variable and which are apomorphic, and applying existing names to fresh material in comparison with poorly-preserved types.

A number of species remain to be described and are currently under study. Some named species are probably distinct taxa, e.g. "Nemacheilus" tigris (Heckel, 1843) (sagmahi-ye Dajleh) is recorded from the Karun River basin in Khuzestan (ZISP 24098) by Berg (1949) but specimens from this part of Iran differ from Heckel's types. The colour pattern on the fish figured by Berg (1949) from the Karun River in Iran is atypical according to Bănărescu and Nalbant (1966) - it has only 4-5 bars on the posterior part of the body. The type locality of Cobitis Tigris is "Flüsschen Kueik bei Aleppo" (Haleb, Syria) according to Heckel (1843b). Prokofiev 92009) places Cobitis tigris in the genus Paracobitis.

Sawada (1982) thought that this family dispersed by two routes from Southeast Asia, one through Siberia and one through South Asia to reach what is now Iran. Menon (1987) considers that the land mass between East Africa and the west coast of India has submerged only recently, probably simultaneous with the birth of the Ganges and Indus. Connections of the Pleistocene fore-deep of the Himalayas with the Tigris-Euphrates basin in what is now the Persian Gulf could have existed, allowing movement of "Nemacheilus" species along a continuous route from Yunnan to Anatolia. Menon (1987) further suggests a series of waves, spreading "Nemacheilus" westwards into Southwest Asia from a South China origin. The Triplophysa wave is the first wave of evolution, in which the earliest stock from Yunnan spread through Tibet in the late Miocene and early Pliocene before the major rise of the Himalayas. By the end of the Tertiary, particularly in the Pleistocene, the Tibetan Plateau had risen causing a dry and cold climate with increased solar radiation and torrential rivers. This change in the environment caused rapid evolution, leading to such taxa as Triplophysa and Hedinichthys. The rupecola wave took place in the late Pliocene along the southern face of the Himalayas through Iran to Anatolia and even northeast Africa. Some of the criticisms listed under the cyprinid genus Garra, whose distribution Menon (1964) also attributes to waves, may be apposite here too.

These are small fishes, up to about 200 mm in size although one species (not in Iran) reaches 482 mm (Prokofiev, 2010). They are quite secretive, hiding under stones or in mud. This common and stressful habitat may have led to a general similarity in body form among the various species. Some are known only from caves, including one Iranian species. Despite their small size, they are regarded as a delicacy in India (Hora, 1956). Barbatula angorae (and presumably other species) is a potential fishing bait for predatory fishes such as Sander lucioperca and has been examined experimentally for this purpose in Turkey (Kuşat et al., 1995). They are generally known as سگ ماهي (sag mahi meaning dog fish, but this is presumably the equivalent of loach in English), لوچ (= louch meaning loach) or mar mahi (= snake fish, presumably in reference to the elongate shape) in Farsi. These general names are not repeated below.

Genus Barbatula
Linck, 1790

Species in this genus were formerly placed in Orthrias (see Eschmeyer's "Catalog of Fishes" to track conflicting views). There are at least 18 species found mainly in western Asia with a few in Europe. The body is elongate, thick, and rounded or slightly compressed. The head is slightly depressed or compressed. Eyes are small and widely spaced. The caudal peduncle is relatively deep. The body is covered by scales at least on the posterior part of the body, the lateral line is complete or at least passes the middle of the body length, and the lateral line tubes do not penetrate the scales. There are no nasal barbels. The lower lips are moderately furrowed. The dorsal fin has 7-9, rarely 10, branched rays. The pelvic fins are inserted slightly behind the dorsal fin origin. The caudal fin is slightly emarginated to deeply forked. There is no dorsal crest on the caudal peduncle. The processus dentiformis (a tooth-like projection at the symphysis of the upper jaw) is weakly or moderately developed and lacks a corresponding gap in the lower jaw. The gut is short. Colour is variable, being barred, striped, irregular spots and blotches, or more or less regular rows of spots. The pectoral fin of males is broadened and thickened, and covered by tubercles in the spawning season. Tubercles also develop on the sides of the head.

 

Genus Ilamnemacheilus
Coad and Nalbant, 2005

This genus is characterised by a high, laterally compressed body; large head with small eyes and mouth; anterior lip lacking an interruption in the middle; posterior lip with widened mental lobes, small round papillae covering only the mental lobes, the rest of the lips being unfurrowed; the processus dentiformes absent; lateral line complete and terminating slightly before the posterior margin of the caudal peduncle; scales small with a quite large and eccentric focus, sparsely present on the rear half of the body; stomach syphonal and intestine straight without loops; gas bladder with two encapsulated chambers united by a short encapsulated duct; paired fins very long; dorsal fin long; and caudal fin well forked.

The type species is Ilamnemacheilus longipinnis by original designation and monotypy.

Ilamnemacheilus longipinnis
Coad and Nalbant, 2005

Common names

None.

Systematics

The holotype and only known specimen is CMNFI 1979-0366 (79-966 is a lapsus), 36.0 mm standard length, Iran, Meymeh River, formerly a tributary of the Tigris River, 17 km west of Dehloran and about 21 km east of the Iraqi border, 32º45'30"N, 47º05'30"E, 28 January 1978, B. W. Coad and S. Coad.

Key characters

Characters are those of the genus.

Morphology

Dorsal fin with 3 unbranched and 10 branched rays, anal fin with 2 unbranched and 5 branched rays, pectoral fin with 9 branched rays and pelvic fin with 5 branched rays. Total vertebrae 28 or 29 including the ural centrum (vertebral fusions present), and some centra have two neural and haemal arches. ?check vertebral counts against other loaches - very low ? fusions Other characters are listed above under the genus and Coad and Nalbant (2005) give some measurements.

Sexual dimorphism

Unknown.

Colour

The sole preserved specimen is a overall a pale brown with 3-4 indistinct greyish blotches in the middle of the second half of the body. All fins are pale but the caudal fin has faded greyish lines along the marginal rays. In life it was an olive-green overall with orange fins.

Size

Reaches 36.0 mm standard length.

Distribution

Endemic to Iran and found in Tigris River basin at a single locality (see above).

Zoogeography

An endemic genus in the Tigris-Euphrates basin (along with Turcinoemacheilus, not in Iran). This species may be related to an undescribed species from the Orontes River basin in Syria.

Habitat

The sample site was a small stream, 20 m wide with a maximum depth of 1 m. Altitude was 210 m. Capture depth was 30 cm in a medium current. The bottom was a mix of pebbles and mud with some encrusting algae. Water temperature was 14ºC, pH was 6.0 and conductivity was 1.65 mS. The cyprinids Cyprinion macrostomum and Garra rufa were caught with the loach.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

None.

Conservation

Known only from a single specimen, abundance and wider distribution unknown.

Further work

More collections are needed to record information on biology and distribution and provide a more detailed description.

Sources

The holotype and sole known specimen (see above).

Genus Metaschistura
Prokofiev 2009

This genus has a single, small species and the characters of the species are those of the genus. Prokofiev (2009) gives further osteological details.

Metaschistura cristata
(
Berg, 1898)

Common names

sagmahi-ye Torkomani or Turkmeny (= Turkmenian crested loach), sagmahi-ye kakoldar-e Torkomani.

[Turkmenskii grebenchatyi golets or Turkmenian crested loach in Russian].

Systematics

This species was described under Nemacheilus in Latin from "Habitat in flum. Tedschent, prope Aschabad, in provincia Transcaspica". The type locality is presumably the Tedzhen River in Turkmenistan although Ashkhabad is not on the Tedzhen River. Berg (1948-1949) notes "not in the Tedzhen!". Syntypes are reported to be in the Zoological Museum of Moscow State University (MMSU) by Eschmeyer et al. (1996).

Placed in the taxon Paracobitis by Bănărescu and Nalbant (1966) and later in Schistura by Bănărescu and Nalbant (1995) and by Nalbant and Bianco (1998).

Key characters

The dorsal fin branched ray count of 8 distinguishes this crested loach from other crested loaches in northeast Iran and adjacent regions. In addition the crest, or adipose fin, is shorter and thicker than in P. malapterura.

Morphology

Dorsal fin unbranched rays 2-6 (assumed to be usually 3) and branched rays 7-9, modally 8, anal fin unbranched rays 2-4, usually 3, and branched rays 5, pectoral fin branched rays 8-11, usually 9-10, and pelvic rays 6-8, usually 7. The dorsal fin origin lies midway between the snout tip and caudal base or nearer the caudal base. There is a fleshy pelvic axillary process but it does not extend beyond the end of the pelvic fin base. Caudal fin slightly emarginate in Turkmenistan but Hari Rud fish have a conspicuous fork. The two lobes of the caudal fin may be equal or either may be longer. The lobes are pointed or rounded. The dorsal adipose fin or crest begins shortly in front of the anal fin origin level and reaches the root of the caudal fin. There is a small gap between the dorsal fin insertion and the origin of the crest, and the depressed dorsal fin partially occupies this gap and the slight rise of the crest origin. It is strongly developed, high and thick. The crest is supported by 22-25 procurrent rays of the caudal fin. Large fish have scales posteriorly on the body. Scales are small, oblong and widely separated. Bănărescu and Nalbant (1966) did not find scales on fish from the Harirud (confirmed for these fish (ZMUC P 2798, 2799, 27100, a fourth specimen P 2797 is lost; these catalogue numbers corrected from Bănărescu and Nalbant (1966) where incorrect; 63.1-68.3 mm standard length) from Obeh, Afghanistan; and not found on 3 fish, 38.1-60.9 mm standard length from Sarakhs on the Hari Rud) and this population may be distinct. Scales may be completely absent or only on the crest (Prokofiev, 2009). The lateral line extends nearly to the caudal base. Caudal peduncle short, 5-6 times in standard length (6.1-6.9 for 3 Iranian fish, 38.1-60.9 mm standard length). Lips are thick and furrowed, the upper lip with a slight median gap and the lower lip widely interrupted. The dentiform process on the upper jaw is weakly developed. The anterior nostril has a large flap developed posteriorly and postero-dorsally. Large fish have bulging cheeks. The posterior part of the intestine is straight (Bănărescu and Nalbant, 1966) or with a single loop (Bănărescu and Nalbant, 1995).

Meristics for Iranian specimens:- dorsal fin branched rays 8(4), anal fin branched rays 5(4), pectoral fin branched rays 9(2), 10(1) or 11(1); and pelvic fin branched rays 7(4).

Sexual dimorphism

There is no sexual dimnorphism.

Colour

Somewhat similar to P. malapterura but with a dark spot at the anterior base of the dorsal fin, on the unbranched and first branched rays. A similar spot may be present at the anterior base of the anal fin on the first two rays. The anterior part of the body is dark and the posterior half (from the dorsal fin insertion to the tail base excluding any bar on the caudal fin rays) has 4-8 broad, brownish bands, separated by narrower light bands and reaching close to or to the ventral part of the body. Some smaller fish have anterior bands on the flank, the total number of bands reaching as high as 17. Bands vary in size from equal to twice the width of the pale interspaces. The last 3-4 bands extend onto the adipose fin. There is also a much darker and narrower band at the caudal base. Bănărescu and Nalbant (1995: Fig. 11A) illustrate a fish where this last band is only developed on the lower half of the caudal peduncle. The bar may also be composed of two blotches above and below the flank midline plus blotches on the bases of the outermost caudal rays both dorsally and ventrally. All fins have thin bands comprised of spots irregularly lined up on the fin rays. The caudal fin may have three such bands while this fin and others may have no clear bands. Some fish have pigment on rays of the dorsal fin almost forming a single broad band extending from mid-fin half way towards the margin. There may be a second smaller spot on the last unbranched dorsal fin ray above the one on the anterior base.

Size

Attains 8 cm.

Distribution

Found in streams on the northern slope of the Kopetdag, Turkmenistan. Some of these streams presumably have headwaters in Iran and the species may be found there although this has yet to be confirmed by specimens. Also reported from the drainage of the Tedzhen River in Afghanistan which forms part of the Iran-Afghanistan border as the Hari Rud or Hari River (Bănărescu and Nalbant, 1966). This species is confirmed for Iran from Sarakhs on the Hari Rud, and probably the Kashaf River, a Hari River tributary (Abdoli, 2000).

A record from Lake Topiatan in the Uzboi River drainage in Turkmenistan north of the Iranian border mentioned in Nikol'skii (1947) is housed at the Zoological Institute, St. Petersburg (ZISP 25788). Berg (1948-1949) corrects the locality as the Sumbar River, an Atrak River tributary in the Caspian Sea basin. This specimen was examined by me and it is not this species but P. malapterura.

Zoogeography

Bănărescu and Nalbant (1966) consider that the closest relative of this species is Nemacheilus tigris (sic) from the Tigris-Euphrates basin rather than other crested loaches such as P. malapterura and P. rhadinaea found in neighbouring river systems. Note that Paracobitis tigris was described from Aleppo (= Haleb, Syria) and its occurrence in the Tigris-Euphrates basin is in question (see family account).

Habitat

A rheophilic species but further details of habitat preferences are unknown.

Age and growth

Unknown.

Food

Gut contents of one fish, 60.9 mm standard length, from the Hari Rud are chironomid larvae.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

None.

Conservation

Population numbers or trends are unknown.

Further work

More collections are needed to record information on biology and distribution.

Sources

Iranian material: BWC 97-3, 3, 38.1-60.9 mm standard length, Khorasan, Hari River at Sarakhs (?); BM(NH) 1914.1.1:32, 1, 54.5 mm standard length, Khorasan, Kashaf River (no other locality data).

Comparative material: SMF 17133, 2, 34.4-98.3 mm standard length, Afghanistan, Hari River near Herat (34º21'N, 62º14'E); BM(NH) 1897.7.5:40, 51.4 mm standard length, Turkmenistan, Ashkhabad (no other locality data).

see also keys in Prokofiev for generic characters?

Genus Oxynoemacheilus
Bănăraescu and Nalbant, 1967
?cf with barbatula diagnosis

Members of this genus are found from Bulgaria east to Iran. There are 28 or more species with 7 recorded from Iran.

These small loaches are scaled, have a body with spots which tend to form bands or a stripe along the lateral line and there is no dark black spot on the base of the anterior dorsal fin rays. There is no caudal peduncle crest and no pelvic fin axillary lobe. Males do not develop inflated or muscular cheeks in the spawning season but tubercles are found on the sides of the head and on the pectoral fins but not as fine brush-like aggregations. Nostrils are closely spaced or slightly separated, the mouth is semilunar with simple lips which are almost smooth to plicate and the dentiform process is weak to moderate but sometimes absent (O. angorae). The lateral line is usually complete but if incomplete as in O. kermanshahensis it reaches beyond the level of the dorsal fin origin. The dorsal fin 7-8 branched rays and the anal fin 5 branched rays. Prokofiev (2009) lists various osteological characters such as 4 radial bones in the pectoral fin, usually 6 hypurals in the caudal fin support (O. brandtii has 5), the epural bone is present in the caudal fin support, the manubrium in the swimbladder is well-developed and the bony capsule is full divided by it, no preethmoid I, the supratemporal commissure is open from above, the fork of cst and cio is not fused to the skull, and others.


Oxynoemacheilus angorae
(Steindachner, 1897)

Common names

sagmahi-ye Angora, sagmahi-e-jooibari.

[lakh angorae in Arabic; Angor cilpaxcasi (or cilpagcasi), Lankaran cilpaxcasi (or cilpagcasi), both in Azerbaijan; Angorskii golets or Angora loach, Lenkoranskii golets or Lenkoran loach, both in Russian; otsidzug in Armenia; stone loach].

Systematics

Nemacheilus angorae was originally described from Angora (= Ankara, Turkey). Nemacheilus bergi Gratsianov, 1907 and possibly Nemacheilus bergianus Derzhavin, 1934 are synonyms.

Nemacheilus angorae lenkoranensis Abdurakhamanov, 1962 (incorrectly lenkoranica in Bănărescu and Nalbant (1966)) is described from "rivers of the Lenkoran coast; Lenkoranchai, Vilyazhchai, Kumbashichai, Tangyaru, Astarinka" in the southern Caspian Sea basin.

Orthrias angorae araxensis Banarescu and Nalbant, 1978 in Banarescu, Nalbant and Balik (1978), is described from the Aras River basin of Turkey (type locality given below) (this subspecies was formerly referred to as Nemacheilus angorae bureschi (Drensky, 1928) by Banarescu and Nalbant (1964) and Banarescu (1968)). Nalbant and Bianco (1998) and Fricke et al. (2007) elevate this taxon to a species.

Nemachilus angorae alasanicus Elanidze, 1983 was described from the upper reaches of the Alazani River at Alvani village, Georgia, a left bank Kura River tributary. It is here attributed to Elanidze (1983) since it is the only taxon listed in that book without an author.

The lower Kura River basin in Azerbaijan (of which the Aras is part) may have another subspecies in which about 40% of the fish have 7 branched dorsal fin rays, a shallower caudal peduncle than B. a. araxensis, and a colour pattern similar to B. angorae angorae (Banarescu, Nalbant and Balik (1978); see also description in Abdurakhmanov (1962) for measurements).

B. bergiana from Iran may be another subspecies of B. angorae. The resolution of the systematics of B. angorae and its subspecies or related species depends on obtaining large numbers of specimens of mature material from the whole range of this species complex. There are few, if any, good characters in this complex which can be used to separate the taxa; body proportions are especially prone to locally induced variation and even the number of dorsal fin rays is at least as variable within samples as between species.

Sixteen syntypes of Nemacheilus angorae from Tabakane-Sir and Tschibuk-Tschai, Turkey are in the Naturhistorisches Museum Wien according to Eschmeyer et al. (1996).

The holotype of Orthrias angorae araxensis, 62.0 mm standard length, from the "Kandili Karassu, oberes Araxes-Becken, Osttürkei" is in the Zoologischen Instituts und Zoologischen Museums der Universität Hamburg (ZMH 4827). Four paratypes, 45.0-65.2 mm standard length, from the same locality are under ZMH 5951 and 2 other paratypes, 51.2-59.3 mm standard length, also from the same locality are in the Institutul de Stiinte Biologice, Bucuresti, Romania (ISBB 2617). Five paratypes, 38.8-52.0 mm standard length, are from the "Oberlauf des Araxes Flusses bei Aras-Nehri, Hasankale" under ZMH 4826 and 3 paratypes, 41.0-45.0 mm standard length, from the same locality are under ISBB 2618 (Banarescu et al., 1978; Wilkens and Dohse, 1993).

A series of 18 fish in the Zoological Institute, St. Petersburg catalogued as ZISP 35701 from "Reka Lenkoran, Azerbaidzhan SSR" collected by Yu. Abdurakhmanov, 22.IX.1954 are probably the type series of Nemacheilus angorae lenkoranensis although they are not marked as such on the jar. One fish is a cobitid, 31.0 mm standard length (probably Cobitis taenia), while the rest measure 24.9-33.3 mm standard length.

Bănărescu and Nalbant (1966) follow Günther (1899) and describe specimens from the Lake Orumiyeh basin as Nemacheilus (= Oxynoemacheilus) persa (q.v.) but Abdurakhmanov (1962) and Saadati (1977) consider them to be close to B. angorae.

Key characters

Abdurakhmanov (1962) distinguishes two subspecies in Azerbaijan by the following key:-

1(2) Caudal peduncle depth in length 2.5-3.3, mean 2.9; dorsal fin with 7-8 branched rays; length to 80 mm (Kura and Aras river basins) B. angorae

2(1) Caudal peduncle depth in length 2.0-2.6, mean 2.3; dorsal fin with 8-9 branched rays; length to 60 mm (rivers of Lenkoran) B. angorae lenkoranensis

Morphology

Dorsal fin with 2-4, usually 2 according to Banarescu and Nalbant (1964) and Dadikyan (1986), unbranched and 6-9, usually 7-8, branched rays (strong mode of 7 in populations of B. angorae angorae from Anatolia while the mode for B. angorae araxensis is 8 according to Banarescu, Nalbant and Balik (1978) and for B. angorae lenkoranensis the range is 7-9 usually 8-9 according to Abdurakhmanov (1962)), anal fin with 2-3 unbranched and 4-6 branched rays, usually 5, pectoral fin with 8-12 branched rays, and pelvic fin branched rays 6-8. The dentiform process on the upper jaw may be present or absent. The lower lip is more fimbriate than the upper and is interrupted in the middle. Weakly developed median flaps are present. Scales are minute. The lateral line is almost complete (almost to the caudal fin base). Gut without a posterior loop. Caudal fin almost truncate (deeply forked in Lake Orumiyeh material examined by Bănărescu and Nalbant (1966) - other characters of this small sample of 3 fish include a long and shallow caudal peduncle, vent just in front of the anal fin, pelvic fin origin under that of dorsal fin, dorsal fin origin much nearer caudal fin base than snout tip, large eyes, lateral line almost complete ending just before the caudal fin, scales are distinct, especially in the posterior half of the body, and cover the whole body including the ventral part, the upper lip is nearly smooth and has a narrow median interruption, the lower lip is more furrowed and has a wider interruption, the posterior part of the intestine is straight, males have thickened pectoral fin rays with tubercles, flanks are spotted to reticulate, and there is a band at the caudal fin base). The subspecies B. angorae araxensis has body proportions practically the same as the type subspecies from Anatolia according to Banarescu, Nalbant and Balik (1978) but the caudal fin margin is almost straight and colour is different.

Sexual dimorphism

Mature males have a dense covering of tubercles on the head, body and fins. Pectoral fin rays 2-4, and to some extent 1 and 5, are thickened, widened and densely tuberculate. Pectoral and pelvic fins are longer in males than females.

Colour

Colour is variable but overall is greyish to yellow with orange tints in particular on the head. There are 5-6 spots along the mid-line of the back and about 20 brown to blackish spots along mid-flank, forming almost a stripe, especially in young. The dorsal spots may also form a stripe. The base of the caudal fin has a black bar. The dorsal and caudal fins have brown bars composed of small spots. The pectoral fin may also have bars, particularly in males. The other fins are hyaline. The subspecies B. angorae araxensis is darker in colour than B. angorae angorae, the flanks being covered in nearly black spots and fine dots almost as low as the belly. The spots are irregular or follow the lateral line almost to the caudal fin but there is no stripe as in the other subspecies.

Size

Reaches 8.5 cm.

Distribution

Found in the Black Sea basin, Aegean Sea basin and the Caspian Sea basin. In Iran, it is recorded from the the Anzali Mordab and the Masouleh River in that basin, upper Safid River basin (Qezel Owzan and Shahrud), the Qareh Su of the Aras River basin in the Caspian Sea basin, southern and eastern tributaries of Lake Orumiyeh (Zarineh and Tata'u, and Talkheh rivers), rivers of the Namak Lake basin (Karaj, upper Shur or Abhar, middle and upper Qareh Chai rivers, Jajrud), and possibly in the eastern Dasht-e Kavir basin (Bănărescu and Nalbant, 1966; Saadati, 1977; Holčík and Oláh, 1992; Abbasi et al., 1999; Abdoli, 2000; Jolodar and Abdoli, 2004; Abdoli and Naderi, 2009). Specimens from Lake Orumiyeh identified as Nemacheilus persa by Günther (1899) are probably this species. Iranian Caspian Sea and Namak Lake basin fish are probably O. bergianus. If O. angorae is restricted to Turkey, then the species for the western Caspian is lenkoranensis and for the Kura-Aras basin is araxensis.

Zoogeography

see (Bănărescu and Nalbant, 1966) for discussion, p. 155

Habitat

This species is found in both streams and lakes. In shallow running water it lives among stones and vegetation.

Age and growth

Heydarnejad (2009) gave the length-weight relationship for an Iranian sample as W = 0.0078TL3.055.

Food

Food items include chironomids, corixids and diatoms as well as the eggs of other fishes such as Alburnus hohenackeri. Abdoli (2000) lists Trichoptera, Ephemeroptera and Chironomidae.

Reproduction

Fecundity reaches 7000 eggs and egg diameter 0.84 mm. Spawning takes place from May to June.

Parasites and predators

Unknown.

Economic importance

None.

Conservation

None.

Further work

The systematics of this species complex needs to be resolved based on additional material and perhaps modern molecular and genetic techniques.

Sources

Oxynoemacheilus araxensis
(Banarescu and Nalbant, 1978)

The presence of this taxon in Iran needs confirmation. type locality??

?OxynoemacheilusBarbatula argyrogramma
(
Heckel, 1849)

Reported from the Tigris-Euphrates basin in Iraq but no Iranian record except by Saadati (1977). Its validity is doubtful (see Freshwater Fishes of Iraq for more information).

Oxynoemacheilus bergianus
(Derzhavin, 1934)

Common names

agmahi-ye Safidruds, sehkhareh (?), sagmahi-e-jooibari.

[Sefidrudskii golets or Sefidrud loach in Russian; Safidrud stone loach]. these names to brandtii?

Systematics

The type locality of this species in Latin from Derzhavin (1934) is "Systema fluminis Sefidrud" (= Safid River system). Berg (1948-1949) gives "Sefid-rud basin: Kisum village; Shah-rud R., falling into the Sefid-rud". The former is at Kisom at 37°14'N, 49°51'E or 37°12'N, 49°54'E in a gazetteer.

Bănărescu and Nalbant (1966) and Banarescu, Nalbant and Balik (1978) place this species as a subspecies of B. angorae. Nalbant and Bianco (1998) consider it to be a distinct species in Orthrias. Abdurakhmanov (1962) compares fish from Lenkoran (B. angorae lenkoranensis) with B. bergiana. Head length and depth, predorsal distance, body depth, caudal peduncle depth, dorsal fin height, and pectoral and pelvic fin lengths are all greater in lenkoranensis while caudal peduncle length, pectoral-pelvic fin distance, interorbital width and eye diameter are all greater in bergianus.

The problems of the systematics of such a widespread and variable taxon as B. angorae are briefly alluded to under that species and I have retained B. bergiana as a distinct species until the problem has received a full study.

The holotype of Nemacheilus bergianus is 41.6 mm standard length and is in the Zoological Institute, St. Petersburg (ZISP 25433) and is from "Basin R. Sefid-Rud, c. Kissum, River Shahrud, tributary Sefid-Rud", collected by A. N. Derzhavin, 20.V.1922. The specimen is faded but there is a figure in Berg (1948-1949, Fig. 619).

Key characters

This species differs from B. angorae by having a less deep caudal peduncle, 3.3 in caudal peduncle length according to ? (but my measurement is 2.6 for the holotype). Saadati (1977) states that the caudal peduncle is deeper in B. bergiana.

 

Character/Taxon

angorae

araxensis

bergiana

brandtii

lenkoranensis

           
           
           
           
           
           

Morphology

Dorsal fin with 3 unbranched and 8 branched rays, anal fin with 3 unbranched and 5 branched rays, pectoral fin with 9 branched rays and pelvic fin with 6-7 branched rays. Vertebrae 31. The pelvic fins are separated by almost the width of a pelvic fin base. Caudal fin slightly emarginate. The upper lip is indented at the mid-point with a bony projection underneath. The lower lip is interrupted and also has a bony projection underneath. There are no adipose fins. The flank is minutely scaled and the lateral line is well-developed along the whole flank (not as in Berg's drawing - ?check this).

Meristics for Iranian fish: dorsal fin branched rays 8(1), anal fin branched rays 5(1), pectoral fin branched rays 9(1) and pelvic fin branched rays 7(1) - based on type ?see my material and add

Sexual dimorphism

Unknown.

Colour

The flank has several irregular dark grey blotches and the back has transverse, dark, squarish spots. The back and upper flank have a pinkish tinge. The lower flank is pale yellowish or whitish. The belly and lower head are white. The iris is silvery on the lower part and golden on the upper, strikingly so. The dorsal, pectoral and caudal fins are pinkish with 2-3 rows of elongate, dark grey spots along the rays. The caudal fin spots are horizontal rather than forming bands as in Oxynoemacheilus brandtii. The upper base of the caudal fin is yellowish rather than dark as in B. brandtii. The pelvic and anal fins are colourless or slightly yellowish-pink without spots. The barbels are typically without pigment except occasionally at the base of the second pair.

Size

Reaches 6.8 cm with Jolodar and Abdoli (2004) giving 7 cm total length.

Distribution

This species is found in the middle Aras River and its tributary the Qareh Chai (?or is araxensis), the Safid, Shafa, Siyah, Shahrud and lower Qezel Owzan rivers of the Caspian Sea basin, and the Karaj, Abhar, upper Qareh Chai and upper Qom rivers of the Namak Lake basin (Saadati, 1977; Abbasi et al., 1999; Abdoli, 2000; Jolodar and Abdoli, 2004; Abdoli and Naderi, 2009)

Zoogeography

See family account.

Habitat

Unknown.

Age and growth

Females are mature at 5.0 cm.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

None.

Conservation

Kiabi et al. (1999) consider this species to be of least concern in the south Caspian Sea basin according to IUCN criteria. Criteria include few in numbers, habitat destruction, limited range (less than 25% of water bodies), present in other water bodies in Iran (sic), present outside the Caspian Sea basin (sic).

Further work

The biology and systematics of this species needs study.

Sources

Type material: Holotype of Nemacheilus bergianus (ZISP 25433), see above.

Iranian material: CMNFI 1970-0527, 8, 33.1-42.7 mm standard length, Gilan, Safid River near Kisom (37º12'N, 49º54'E); CMNFI 1970-0537, 24, 33.3-50.4 mm standard length, Markazi, Shah River near Manjil (36º44'N, 49º24'E); CMNFI 1970-0545, 1, 44.4 mm standard length, Gilan, Safid River (37º01'N, 49º38'E); CMNFI 1980-0132, 26, 19.4-44.8 mm standard length, Gilan, Safid River at Kisom (37º12'N, 49º54'E); CMNFI 1980-0154, 26, ? mm standard length, Markazi, Karaj River below village (35º47'N, 50º58'E); CMNFI 1980-0156, 80, 29.9-50.8 mm standard length, Markazi, Karaj River near village (35º47'N, 50º58'E).

Oxynoemacheilus brandtii
(Kessler, 1877)

Common names

sagmahi-ye Kura.

[Kur cilpagcasi in Azerbaijan; Kurinskii golets or Kura loach in Russian].

Systematics

A syntype of Nemacheilus Brandtii from the upper Kura River at Tbilisi (= Tiflis), Georgia is in the Natural History Museum, London (BM(NH) 1897.7.5:39, 19.6 mm standard length, small and decoloured), formerly in ZISP; other syntypes are in the Zoological Institute, St. Petersburg (ZISP) (Eschmeyer et al., 1996).

Nemachilus brandtii gibbusnazus is a subspecies with the author given as Elanidze in Elanidze (1983). The only apparently new taxon in Elanidze (1983) is Nemacheilus (= Oxynoemacheilus) angorae alasanicus (q.v.). This taxon is unique in this book as a species without an author name after it. On this basis it was attributed to Elanidze (1983). The date of N. b. gibbusnazus may also be 1983 but it may have been described in an earlier paper by Elanidze not yet located by me. Both these subspecies are not mentioned in Eschmeyer et al. (1996) nor in the online version (downloaded 26 August 2007). The distribution of N. b. gibbusnazus is given as "in fact found before in the R. Khrami (1947), then in the R. Kura (1962), in its lower course at Kukheti, in the upper reaches at Vardziya - Toloshi, in the R. Alazani - in the upper reaches at Alvani", all in Georgia and the drainage of the Kura River.

Placed in the genus Orthrias by Nalbant and Bianco (1998).

Key characters

This species is distinguished from B. angorae by having 3 unbranched and 8 branched dorsal fin rays (as opposed to 2 unbranched and 7-8 (usually 7) branched in B. angorae), longer and shallower caudal peduncle (caudal peduncle depth 2.2-3.0 in caudal peduncle length instead of less than 2.0 as in B. angorae), and a more forked caudal fin with more pointed lobes. The caudal peduncle is said to be somewhat shorter and much lower than in B. angorae araxensis (with which species it occurs in the Aras River basin): length of caudal peduncle 18.4-23.4% and depth 8.0-9.4% of standard length in brandtii, 18.1-21.8% and 10.2-12.7% in araxensis respectively. Other body proportions are given in Banarescu, Nalbant and Balik (1978). The colour pattern is much darker and the caudal fin margin is almost vertical in B. a. araxensis.

Morphology

Dorsal fin unbranched rays 3-4 and branched rays 7-9, usually 8, anal fin unbranched rays 2-3 and branched rays 5, pectoral fin branched rays 9-12, and pelvic fin branched rays 6-8. Scales minute. A weakly developed dentiform process on the upper jaw.

Sexual dimorphism

Unknown.

Colour

Typical fish from the Kura-Aras basin lack a stripe on the flank, having brown spots which are either large, more or less triangular and fairly well-defined or broken into many small speckles forming a reticulate pattern. Dorsal spots are better defined than in B. angorae especially behind the dorsal fin where they fuse completely or incompletely with the lateral flank spots to form bands. All spots are brown, never blackish as in B. angorae araxensis. There are several rows of speckles on the caudal fin and two rows on the dorsal fin.

Size

Reaches 8.5 cm.

Distribution

Found in the upper and middle Kura and Aras River basins, and presumably in the Iranian reaches of the latter. It is also reported from the Namak Lake basin by Saadati (1977) citing a manuscript report by V. D. Vladykov, ? confusion with bergianus and from the Arnar Chay, Azarbayjan.

Zoogeography

See family account.

Habitat

Unknown.

Age and growth

Age at maturity is about 2 years.

Food

Eggs of other fishes may be a food item as well as aquatic insects.

Reproduction

Fecundity reaches 17,409 eggs and egg diameter 0.95 mm.

Parasites and predators

Unknown.

Economic importance

None.

Conservation

Endangered in Turkey (Fricke et al., 2007).

Further work

The biology and systematics of this species need study.

Sources

Type material: Syntype of Nemacheilus Brandtii ((BM(NH) 1897.7.5:39), see above.

Oxynoemacheilus farsicus
(Nalbant and Bianco, 1998)

Common names

None.

Systematics

Originally described in the genus Orthrias. The species is named for Fars Province. The holotype is in the Department of Zoology, Naples University under IZA 7823. It measures 45.8 mm standard length (51.8 mm standard length when measured by me) and was collected from the "River Kor near Persepolis" in Fars on 30 May 1976 by P. G. Bianco (note that IZA is the acronym for Dipartimento di Scienze Ambientali, Universita' Degli Studi Dell'Aquila
L'Aquila, Italy, where the material was previously stored). Paratypes were all collected by P. G. Bianco and include 64 specimens, 39.0-57.0 mm standard length from the same locality as the holotype (IZA 7824), 61 specimens, 38.5-60.0 mm standard length from the same locality as the holotype but taken on 7 July 1975 (IZA 7825), 7 specimens, 42.0-51.3 mm standard length, same locality as the holotype but taken on 7 July 1975 stored in the Institutul Stiinte Biologice, Bucharest (ISSB 3451), 31 specimens, 43.2-56.3 mm standard length from the River Shur, tributary of the River Mand, near Dasht-e-Arzhan (Shiraz) 25 May 1976 (IZA 7826), 7 specimens, 47.0-57.0 mm standard length from the latter locality (ISBB 3446), 7 specimens, 47.0-58.0 mm standard length from River Qom, Qom Town 5 May 1975 (IZA 7845) and 7 specimens, 36.2-47.3 mm standard length from the latter locality (ISBB 3442). CMNFI material?

Key characters

This fish has long paired fins and a short, deep and compressed caudal peduncle. The flanks and back have roundish brown spots on a marbled background and the head and body are a pale yellow ventrally. Dorsal and caudal fins have rows of brown spots. The peritoneum is a dull brown. The dorsal fin has 3 unbranched and 8 branched rays, the anal fin 2 unbranched and 5 branched rays, the pectoral fin 2 unbranched and 9-11 branched rays and the pelvic fin 1 unbranched and 6-7 branched rays. It is related to B. brandtii but differs in in the deeper caudal peduncle and longer fins. Other species outside Iran have differing colour patterns.

Morphology

Dorsal fin branched rays 7-9 (90% with 8 rays in the type description, n=38), anal fin branched rays 4-5 (95% with 5 rays), pectoral fin branched rays 9-11 (69% with 10 rays) and pelvic fin branched rays 6-7 (65% with 7 rays). The lateral line is straight, on the mid-flank and ends just anterior to the caudal fin base. The cephalic canal system is well-developed laterally. The rear half of the body is scaled completely and the anterior half has scalation well-developed but not as closely spaced as posterior scales. Scales are oval with a large focus. Esmaeili and Niknejad (2006-2007) give scanning electron micrographs of the scales along with a description. The head is conical and the eye is small and centrally placed. The nostrils are just anterior to the eyes. The mouth has a moderate arch with long barbels. Lips and barbels have small tubercles and fine furrows. The upper lip has a median incision and the lower lip lacks mental lobes. The processus dentiformis in the upper jaw is well-developed but lacks a gap in the lower jaw.

Sexual dimorphism

Males have numerous fine tubercles on the dorsal surface of the pectoral fin rays in bands, and the anterior fn rays are expanded. There are fewer rays and less expansion when progressing proximally. There is a groove running anteriorly at a slight downward angle from the eye to the antero-ventral corner below the nostril, fading out under the anterior nostril. Near the eye and dorsal to this groove, fine tubercles form a narrow band. The sides and top of the head are finely tuberculate. The pelvic and dorsal fin rays have tubercles following the rays (not bands as in the pectoral fin). Fine tubercles line the margin of the scales.

Colour

The dorsal head is speckled brown and there is a dark-brown stripe from the eye to the snout tip, sometimes extending beyond the eye. The back has 7-11, modally 9, rounded dark-brown saddles. Similar blotches occur on the flank. The dorsal fin has 4 rows of elongate spots, the pectoral fin has up to 6 rows and the caudal fin has 4-5 rows. The anal and pelvic fins are immaculate.

Size

Reaches 67.7 mm standard length.

Distribution

Endemic to Iran and found in the basins of the Namak Lake basin (Qom River at Qom), Kor River basin (Kor River near Persepolis) and Gulf basin (Shur River near Dasht-e Arzhan in the Mand River drainage).

Zoogeography

Found in both endorheic and exorheic basins in central and southern Iran.

Habitat

Details are unknown.

Age and growth

Unknown. Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 34 fish measuring 3.15-6.77 cm standard length. The a-value was 0.0126 and the b-value 3.084 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases).

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

None.

Conservation

Population trends and numbers unknown.

Further work

More collections are needed to record information on biology and distribution.

Sources

Type material: The holotype (IZA 7823) and paratypes (IZA 7824, 7825, 7845) of Orthrias farsicus, see above.

?CMNFI material IZA 7824, 3 as gift, 36.5- , IZA 7826, 3 as gift, 34.0- the measurements are the smallest fish in these samples)

Oxynoemacheilus frenatus
(Heckel, 1843)

Common names

See genus account.

[lakh or telay (= bowed head according to Heckel (1843b) at Mosul), both in Arabic; banded Tigris loach].

Systematics

The type locality of Cobitis frenata is "Tigris", presumably at Mosul (Heckel, 1843b). Five syntypes are in the Naturhistorisches Museum Wien (NMW 48552) although the catalogue lists only 4 specimens. A lectotype designated by F. Krupp in 1984 is 70.0 mm standard length, the remaining specimens being small, 27.2-43.4 mm standard length.

Nemacheilus frenatus afrenatus Battalgil, 1942 described from "un petit ruisseau à Diyarbakir" is also from the Tigris River basin but in Turkey. This subspecies lacks the "frein à la bouche" (presumably the band across the snout) of B. f. frenata and its dorsal fin is higher than long (as measured at the base) while in B. f. frenata it is as high as long.

Bănărescu and Nalbant (1995) place this species in the genus Orthrias. Bănărescu and Nalbant (1966) consider this taxon to be a "doubtful species" but illustrate it in Bănărescu and Nalbant (1995: Fig. 20).

Key characters

The colour pattern is distinctive and there is no dermal crest or adipose fin behind the dorsal fin.

Morphology

Dorsal fin unbranched rays 2-3, branched rays 7-8, anal fin unbranched rays 2, branched 5, pectoral fin branched rays 10-13, and pelvic fin branched rays 6-7. Scales are present over the whole body but not readily visible without magnification. The anterior pectoral fin rays are thickened. Caudal peduncle thick (depth 80-90% of length) according to Saadati (1977). The bulb of the swimbladder capsule has large ovoid to circular perforations and the anterior and posterior lamina or wings are only moderately developed (Krupp, 1985c). Lips are not strongly plicate and the dentiform process is well-developed. The gut has a posterior loop.

Meristics for Iranian specimens: dorsal fin branched rays 8(1), anal fin branched rays 5(1), pectoral fin branched rays 10(1) and pelvic fin branched rays 6(1) based on lectotype.? add my material

Sexual dimorphism

Bands of tubercles are found on the pectoral fin rays of males, including the first, declining in breadth and extent on the smaller rays. Tubercles are also present on the pelvic and anal fin rays but are much less well developed. The head is covered in fine tubercles. Flank scales, particularly anterior ones, are lined anteriorly with tubercles. There is an elongate tuberculate swelling anterior to the lower eye margin on the snout.

Colour

Overall colour is yellowish, mottled with fine but irregular brown or black dots or blotches, some flank blotches being quite large. The rear of the body and the caudal fin in particular are mottled with brown, tending to form bars. A black band is continuous from the front of one eye, across the snout and round to the other eye. It may be diffuse on the snout or well-defined in fish from the same locality. The dorsal and caudal fins have thin but irregular bands made up of spots on the rays (up to 3 on the dorsal and 4 on the caudal fin), bands are faintly present on the anal and pelvic fins, and only a few are visible on the pectoral fins. The dorsal fin has an anterior basal spot at its origin, variably developed in individuals. There are distinct, dark spots at the base of the caudal fin above and below the body mid-line.

Size

Reaches about 9.2 cm (Heckel, 1843b).

Distribution

Found in the Quwayq and Tigris-Euphrates rivers. Abdoli (2000) records it from the upper Karun, middle and lower Dez, Kashkan and Simarreh rivers in the Tigris River basin of Iran.

Zoogeography

See family account.

Habitat

Known to inhabit both rivers and lakes, the environmental requirements of this species are unknown.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

None.

Conservation

The biology of this species, its distribution, numbers and habitat requirements are unknown so no comments on conservation can be made. Near threatened in Turkey (Fricke et al., 2007).

Further work

The biology and conservation status of this species need investigation.

Sources

Type material: Syntypes of Cobitis frenata (NMW 48552), see above.

Iranian material: BWC 95-30, mm standard length, .

Comparative material: BM(NH) 1974.2.22:1449-1477, 30, 37.7-55.5 mm standard length, Iraq, branch of Khalis River (); BM(NH) 1974.2.22:1478-1757, ca. 279, 19.7-53.3 mm standard length, Iraq, Khalis (); BM(NH) 1974.2.22:1758-1772, 15, 27.2-34.1 mm standard length, Iraq, Khalis (); BM(NH) 1972.2.22:1773-1776, 4, 28.3-34.0 mm standard length, Iraq, Khalis (); BM(NH) 1974.2.22:1777-1778, 2, 27.5-31.4 mm standard length, Iraq, Khalis ();.

Oxynoemacheilus kermanshahensis
(Bănărescu and Nalbant, 1966)

Common names

sagmahi-ye Kermanshah.

[Kermanshah loach].

Systematics

This species was tentatively placed in the subgenus Orthrias Jordan and Fowler, 1903 but is regarded as "aberrant" by Bănărescu and Nalbant (1966). However, Nalbant and Bianco (1998) later place it in Orthrias.

The holotype of Noemacheilus kermanshahensis (ZMUC P 2787, 46.4 mm standard length) and the 7 paratypes (ZMUC P 2788-94, 25.5-61.8 mm standard length) are stored in the Zoological Museum of Copenhagen (Nielsen, 1974). The type locality is "Kermanshah in the drainage of the Karun River, a tributary of the lower Euphrates, Western Iran" (Bănărescu and Nalbant, 1966). The type series was collected on 5 February 1937 by E. Kaiser. The type locality is poorly defined - the city of Kermanshah is on the Qarasu River and lies in the drainage of the Karkheh River which drains towards the Tigris River. Nalbant and Bianco (1998) correct their original type locality description to the Karkheh River drainage and "Quareh Su, Kermanshah, River Shimarek", probably referring to the Qareh Su-Simareh drainage of the upper Karkheh River.

Key characters

Bănărescu and Nalbant (1966) consider that this species differs from most South and West-Asiatic loaches by the longitudinal pigment pattern on the flank. Most other species have bars, vertical patches of pigment. The longitudinal arrangement is found in B. angorae but kermanshahensis has smaller eyes, a shorter lateral line (almost complete in angorae), a more anteriorly placed vent, and scale shape comprising an almost central focus, vertical sub-oval outline and radii on all fields widely and evenly spaced. Characteristically the caudal peduncle is short and deep, depth being 92.9-114.3% of length (Saadati, 1977).

Morphology

Dorsal fin with 3 unbranched and 7-8 branched rays, anal fin with 2 unbranched and 5-6 branched rays, and pelvic fin with 6-7 branched rays. Scales only on the posterior part of the body, well-developed on the caudal peduncle. The lateral line reaches the level of the middle or posterior half of the dorsal fin or over the front half of the anal fin. The origin of the dorsal fin is variable in relation to the snout tip and caudal fin base (Bănărescu and Nalbant, 1966). The caudal peduncle is short and deep and lacks a crest. The anus is somewhat anterior to the anal fin origin. Lips are thick and fringed, the lower lip being interrupted in the middle. The intestine is simple with the posterior part straight.

The type series counts are dorsal fin branched rays 7(8), anal fin branched rays 5(8), pectoral fin branched rays 8(3), 9(1), or 10(3), pelvic fin branched rays 6(3) or 7(6); vertebrae 38(6), 39(2) or 40(1).

Sexual dimorphism

Check my fish ? No sexual dimorphism was noted by Bănărescu and Nalbant (1966) as their large specimen is female and others immature.

Colour

The overall colour is yellowish with about 12, dark brown bars on the flank my fish - check this, see my fish?. A stripe on the mid-line of the back breaks up into spots posteriorly.

The colour pattern in alcohol-preserved specimens dating from 1937 is yellowish with 3 wide, brownish stripes along the flank. The dorsal stripe is continuous anteriorly but breaks up into spots posteriorly in most specimens. The central, mid-flank stripe is the widest and is continuous although width is variable. The ventral "stripe" (my quotation marks) is absent from smaller specimens and consists of small and irregular spots. The upper parts of the head have small, irregular brownish spots. The caudal fin has 3 rows of spots and the dorsal fin 2 rows of spots, apparently concentrated on the fin rays (Bănărescu and Nalbant, 1966).

Size

Reaches 6.3 cm standard length.

Distribution

Known only from Tigris River basin drainages of Iran (see above). Abdoli (2000) reports it from the Marun, upper Karun and lower Khersan, Dez, Simarreh, Qareh Su and Gav Masiab rivers.

Zoogeography

Relationships may lie with B. angorae according to Bănărescu and Nalbant (1966) based on colour pattern and general body shape, or with other species not found in Iran having stripes along the flank. This seems insufficient evidence on which to base relationships.

Habitat

my collection data ?

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

None.

Conservation

No measures are being undertaken for this poorly known species.

Further work

Studies should be carried out to determine the numbers of this species, its distribution and its ecological requirements in Iran to ascertain if conservation measures should be taken.

Sources

Type material: The holotype (ZMUC P 2787) and paratypes (ZMUC P 2788-94) of Noemacheilus kermanshahensis, see above.

Iranian material: CMNFI 1979-0285, 20, 32.0-52.6 mm standard length, Kermanshahan, Marek River (34º26'N, 46º37'E)checkID?; CMNFI 1979-0286, 1, 54.2 mm standard length, Kermanshahan, Ravansar River at Ravansar (34º43'N, 46º40'E)checkID?; CMNFI 1993-0128, 1, ? mm standard length, Kermanshahan, Sarab-e Sabz 'Ali Khan (34º25'N, 46º32'E); BWC 95-17, ? mm standard length, .

? genus(Kessler, 1877)

Reported from the Karakum Canal of Turkmenistan by Shakirova and Sukhanova (1994) and Sal'nikov (1995), this species may eventually reach the Tedzhen River and Caspian Sea basins of Iran. Originally described in the genus Nemacheilus. No Iranian record.

Oxynoemacheilus persus
(Heckel, 1849)

Common names

sagmahi-ye Fars (= Persian loach).

[Persian loach].

Systematics

The type locality for Cobitis Persa is "Quellen um Persepolis" according to Heckel (1846-1849b). Kähsbauer (1964) reports a syntype of this species in the Naturhistorisches Museum Wien under NMW 48567. It measures 47.6 mm standard length and is probably the holotype as the Vienna catalogue lists only 1 specimen (and the Vienna card catalogue examined in 1997 concurs). This specimen is in poor condition and not readily comparable to fresh material. It is scaled although scales are not imbricate, the caudal fin is broken off, the body is collapsed so its shape cannot be determined, and it is decoloured.

Kessler (1877) refers to a Heckel species Nemachilus persicus, presumably this taxon judging from Kessler's page number reference.

Günther (1899) recorded Nemacheilus persa from the Lake Orumiyeh basin ("Zola Chai near Ula") and the upper Aras River basin ("Elinja Chai") but these fish were probably B. angorae. Berg (1948-1949), Bănărescu and Nalbant (1966) and Nalbant and Bianco (1998) also refer Günther's material to Nemacheilus persa but Abdurakhmanov (1962) suggests that these fish are Nemacheilus angorae. Saadati (1977) places Lake Orumiyeh fish close to Nemacheilus angorae which makes more sense geographically. ? check my specimens and compare ? The dorsal fin origin is closer to caudal base than the snout tip in the Lake Orumiyeh fish, interorbital width is greater, the head is longer, the caudal fin less emarginate, and there are more and darker spots on the body (Saadati, 1977). Evidently new material from the Kor River basin wherein lies Persepolis, the type locality for Cobitis persa, and the Lake Orumiyeh basin are required to resolve this problem - the Lake Orumiyeh sample examined by Bănărescu and Nalbant (1966) comprised only 3 males.

Bănărescu and Nalbant (1966) place this species in their subgenus Oxynoemacheilus but later (Bănărescu and Nalbant (1995); and also Nalbant and Bianco (1998)) place it in Orthrias.

Key characters

?

Morphology

Dorsal fin with 2-3 unbranched and 8 branched rays, anal fin with 2 unbranched and 5 branched rays, pectoral fin with 8 branched rays, and pelvic fin with 6-7 branched rays. Lateral line almost complete. Scales are present on the anterior and posterior flank according to Banarescu and Nalbant (1964). Esmaeili and Niknejad (2006-2007) give scanning electron micrographs of the scales along with a description. The caudal fin is slightly emarginate to forked. The barbels are hair-like thin (Heckel, 1846-1849b).

Meristics for Iranian specimens: dorsal fin branched rays 8(1), anal fin branched rays 5(1), pectoral fin branched rays 9(1) and pelvic fin branched rays 6(1) based on type ?add my material

Sexual dimorphism

Banarescu and Nalbant (1964) report that pectoral fin rays 2-5 are widened and thickened in males. The appearance of a subocular pad in Bănărescu and Nalbant (1966:Fig. 3) is an error and was apparently meant to represent a small groove (Bănărescu and Nalbant, 1995).

Colour

The overall colour is yellowish with some brown blotches and 5-7 brown bars on the posterior body and irregular brown blotches on the anterior body. The dorsal and caudal fins are barred.

Size

Reaches 63.2 mm standard length.

Distribution

Found in the Kor River and Lake Maharlu basins (Esmaeili et al., 2011?) and, according to Bănărescu and Nalbant (1966), also in the Lake Orumiyeh basin and probably the whole of western and central Iran. Abdoli (2000) maps the Kor and Pulvar rivers.

Zoogeography

See family account.

Habitat

Unknown.

Age and growth

Unknown. Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 32 fish measuring 2.94-6.32 cm standard length. The a-value was 0.0211 and the b-value 2.784 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases).

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

None.

Conservation

?

Further work

?

Sources

Type material: The holotype of Cobitis persa (NMW 48567), see above.

Iranian material: CMNFI 1979-0019, 2, 28.9-33.7 mm standard length, Fars, Barm-e Baba Hajji (29º23'N, 52º40'E); CMNFI 2004-0004, ? mm standard length.

Genus Paracobitis
Bleeker, 1863

This genus comprises about 17, comparatively large species, some over 28 cm, in western Asia. Six species are reported from Iran.

The body is elongate, thick anteriorly and posteriorly compressed. The head is strongly depressed. Nostrils are closely spaced or weakly separated. The caudal peduncle is long, low and bears an elongate crest dorsally from the dorsal fin to the caudal fin and, often, a more or less well-developed ventral adipose crest. However, the crest is not supported by the procurrent rays of the caudal fin unlike in Metaschistura. The caudal fin is slightly emarginate or truncate, rarely rounded. There is a well-developed pelvic axillary lobe reaching beyond the vertical of the posterior edge of the pelvic fin base. Scales are present or absent anteriorly and the lateral line is complete. The processus dentiformis is usually strongly developed (weak to absent in P. vignai - ?check) but its notch on the lower jaw is reduced. Lips are smooth or grooved. The gut is short with a single loop. The colour pattern is variable comprising bars or irregular spotting. There is no intense black spot at the base of the anterior dorsal fin rays. Males and females show no external differences except males have more muscular cheeks. Prokofiev (2009) gives further osteological details of this genus including cst having a bony roof, the fork of cst and cio is fused to the skull in adults (the cst is the supratemporal commissure running across the top of the head at the rear and cio is the temporal part of the infraorbital head canal), the free part of the swimbladder is reduced or absent, right and left lobes of the bony capsule are fully divided by the manubrium, and the fifth trunk vertebra does not take part in support of the capsule. There are 4 radials in the pectoral fin skeleton and the caudal fin support has 6 hypurals and an epural is present. There is no preethmoid I bone in the skull.

Paracobitis boutanensis
(McClelland, 1842)

Probably described from the neighbourhood of the Bolan Pass, Helmand River drainage of Afghanistan in the Sistan basin according to Hora (1929) and Bănărescu and Nalbant (1966). No Iranian record.

Paracobitis cyri
(Berg, 1910)

Nemacheilus tigris cyri Berg, 1910 is described from "Fontes fl. Cyrus, Caucasus". This species is placed in the subgenus or genus Paracobitis by Banarescu (1968) and Prokofiev (2009) while Fricke et al. (2007) recognise the subspecies as a distinct species in Orthrias. Six syntypes of N. tigris cyri (31.4-51.5 mm standard length) are in the Zoological Institute, St. Petersburg (ZISP 13291) from "Okam village Gel'skaya Plain, Karka Oblast", K. Satunin, 6.IX.1901. This is in the upper Kura River basin of Turkey. No Iranian record.

Paracobitis ghazniensis
(Bănărescu and Nalbant, 1966)

Described from "Ghazni, on the Ghazni River, tributary of the Ab-i-Istadah Lake, Helmand drainage; East Afghanistan" (Bănărescu and Nalbant, 1966). Ghazni is at 33°33'N, 68°26'E in the Sistan basin. No Iranian record.

Paracobitis iranica
Nalbant and Bianco, 1998

Common names

sagmahi-ye irani.

[Iranian loach].

Systematics

The species is named for Iran. The holotype is 79.8 mm standard length (66.0 mm standard length when measured by me) and is from "River Qom near the town of Qom" collected on 6 May 1976 by P. G. Bianco and stored in the Department of Zoology, University of Naples (IZA 7831). The locality in the jar is "Qom River near Qom (a little salt river 3.5 p.p.t.) near Qom at the bridge, 950 m, 5 June 1976", the date being at variance with the published date. Paratypes number 5 specimens, 59.0-92.0 mm standard length (IZA 7832)(only two present in 2002 visit by me), and 4 specimens, 47.0-72.0 mm standard length (Institutul Stiinte Biologice, Bucharest, ISBB uncatologued). It may be related to P. malapterura and P. longicauda but differs in colour and the larger caudal peduncle keel.

Key characters

The body is elongate with a compressed posterior region and a depressed head. The caudal peduncle is elongate and has a long dorsal adipose crest. The body is scaled. The overall colour is yellowish-white with a row of dark grey spots on the mid-back and on mid-flank. Spots extend onto the adipose crest. A row of dark irregular spots is found on the posterior part of the ventral region. The holotype has a dorsal fin with 3 unbranched and 7 branched rays, an anal fin with 2 unbranched and 5 branched rays, a pectoral fin with 2 unbranched and 9 branched rays and a pelvic fin with 1 unbranched and 5 branched rays.

Morphology

The dorsal fin has 7-8 branched rays (90% with 7 rays, n=9), the anal fin has 5 branched rays in all specimens examined in the type description, the pectoral fin has 9 branched rays in all specimens and the pelvic fin has 6-7 branched rays (78% with 7 rays, n=9). The lateral line is straight and extends to the caudal fin base. The body is minutely scaled with a relatively eccentric and large focal zone. The head is conical with eyes in the anterior half. The nostrils are just in front of the eyes. The mouth has a strong arch with short barbels and well-furrowed lips interrupted in the middle. The processus dentiformis is well-developed. The stomach is syphonal and the intestine straight. The gas bladder capsule has a relatively short duct.

Sexual dimorphism

Unknown.

Colour

The head is covered by dark grey dots and all fins have small grey dots. See also Key characters.

Size

Attains 92.0 mm standard length.

Distribution

Known only from the Qom River near Qom in the Namak Lake basin, an Iranian endemic.

Zoogeography

This species and similar ones are widely distributed in Iran, although poorly collected, and their systematics and zoogeographical relationships are unknown.

Habitat

Details are unknown. The type series was collected in a small salt river (3.5‰).

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

None.

Conservation

Population trends and numbers unknown.

Further work

More collections are needed to record information on biology and distribution.

Sources

Type material: The holotype (IZA 7831) and paratypes (IZA 7832) of Paracobitis iranica, see above.

Paracobitis longicauda
(Kessler, 1872)

Common names

sagmahi-ye kakoldar-e sharqi (= eastern crested loach).

[vostochnyi grebenchatyi golets or eastern crested loach in Russian].

Systematics

May be a subspecies or synonym of P. malapterura. Has been placed in the genus Adiposia Annandale and Hora, 1920. Banarescu and Nalbant (1964) restrict P. longicauda to basins in Central Asia (e.g. the Amu Darya) and not Iran. Cobitis longicauda was originally described from the Ak-Darya in the Zeravshan River basin of Uzbekistan and the holotype is in the Zoological Institute, St. Petersburg under ZISP 2686 (Eschmeyer et al., 1996).

Bănărescu and Nalbant (1995) place this species in Paracobitis.

Key characters

This species is distinguished from P. malapterura by distinct scales and larger size.

Morphology

Dorsal fin unbranched rays 2-3, branched rays 7, anal fin unbranched rays 2-3, branched rays 5, pectoral fin branched rays 9-10,and pelvic fin branched rays 7. Caudal fin truncate. Large fish are scaled on the flanks, dorsal crest and belly, scales being visible to the naked eye in contrast to the few scales needing a microscope to be visible in P. malapterura and in P. rhadinaea. Caudal peduncle long, about 4.5-4.6 in standard length.

Sexual dimorphism

Colour

Colour is very similar to P. malapterura with spots sometimes loop-shaped and sometimes rounded.

Size

Reaches 20.0 cm.

Distribution

Found in the Tedzhen and Murgab rivers of Afghanistan and Turkmenistan and in the Aral Sea basin, and in the Tedzhen (= Hari) River basin of Iran.

Zoogeography

See family account.

Habitat

Unknown.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

None.

Conservation

This species is too poorly known in terms of biology and numbers for an effective conservation assessment.

Further work

The biology of this species in Iran needs study.

Sources

Comparative material: ZISP 2686, 1, 140.8 mm standard length, ?, Samarkand (); ZISP 4482, 1, 145.7 mm standard length, Zeravshan (?); ZISP 4483, 4, 107.2- 125.8 mm standard length, ?, Zeravshan (); ZISP 4484, 8, 62.2-96.7 mm standard length, ?, Zeravshan (); ZISP 10362, 4, 31.8-112.1 mm standard length, ?, Tedzhen (); ZISP 10963, 1, 93.7 mm standard length, ?, Samarkand (); ZISP 13300-13301, 3, 93.7-157.6 mm standard length, Pul-e Khatun (?); ZISP 14510, 3, 119.4-173.4 mm standard length, ?, Samarkand (); ZISP 33105, 9, 30.4-98.6 mm standard length, ?, Ak-Darya ().

Paracobitis malapterura
(Valenciennes, 1846)

Common names

sagmahi-ye kakoldar-e gharbi (= western crested loach), sagmahi-ye juibari or sagmahi-e-jooibari.

[lakh in Arabic; zapadnyi grebenchatyi golets or western crested loach in Russian].

Systematics

The type locality of Cobitis malapterura is given as "Syrie" in Cuvier and Valenciennes (1846) but has not been found there since (Berg, 1948-1949); it does occur in the Tigris-Euphrates basin however (Coad, 1991b). The specimen was collected by Aucher-Éloy who visited Iran and the specimen may in fact be from there. It is possible that the type locality is in the Caspian Sea basin of Iran although Banarescu and Nalbant (1964) give the Tigris-Euphrates basin which extends though Syria for the distribution of P. malapterura malapterura. Bănărescu and Nalbant (1995) places this species Paracobitis.

Cobitis longicauda Kessler, 1872 is possibly a synonym. Banarescu and Nalbant (1964) restrict P. longicauda (q.v.) to basins in Central Asia (e.g. the Amu Darya) and not Iran. Nemacheilus macmahoni Chaudhuri, 1909 has been advanced as a synonym or subspecies (Nikol'skii, 1947; Berg, 1948-1949; 1949; Banarescu and Nalbant, 1964) but see the review in Bănărescu and Nalbant (1966) and below under P. rhadinaea. Banarescu and Nalbant (1964) consider fish from Sistan and the Caspian Sea basin of Iran to be P. malapterura macmahoni.

Filippi's (1865) record of Cobitis merga (Krynicki, 1840) from "fiumicelli di Sartschem e di Sainkalé" was P. malapterura; these localities being in the Safid River basin near the falling of the Zanjan River into the Qezel Owzan of Safid Rud (Berg, 1948-1949) presumably at Sarcham-e Sofla (37°07'N, 47°54'E) and possibly Sa'in Qaleh (36°18'N, 49°04'E) in the Namak Lake basin.

Two syntypes are in the Muséum national d'Histoire naturelle, Paris under MNHN 3962 and B.3070 (formerly MNHN 3962) (Eschmeyer et al., 1996) and measure 125-145 mm total length (Bertin and Estève, 1948).

Key characters

The colour pattern is distinctive and there is a well-developed dermal crest or adipose fin behind the dorsal fin to the caudal fin base.

Morphology

Dorsal fin with 2-3 unbranched and 6-8 branched rays, anal fin with 2-3 unbranched and 5 branched rays, pectoral fin branched rays 10-12 (Nikol'skii (1947) gives 7-10) and pelvic fin branched rays 5-7. Caudal fin slightly emarginate. There is a well-developed dermal crest or adipose fin behind the dorsal fin to the caudal fin base. Scales are scattered on the posterior body in large fish but need magnification to be seen. The lateral line extends almost to the caudal fin. The dentiform process of the upper jaw is well-developed and fits in a lower jaw groove. The lips, especially the lower one, are strongly plicate. The eyes are small and widely spaced. Caudal peduncle short, 5.6-6.3 times in standard length. The gut is straight posteriorly.

Sexual dimorphism

The cheeks are distended in some fish and this is believed to be a sexual character.

Colour

The top and sides of the body and head are mottled or reticulated with grey and some yellowish pigment. The reticulations may be very fine, giving a more mottled appearance. The belly and lower head surface are white. The flank reticulations extend onto the caudal peduncle crest. When touching the dorsal margin of the crest, the reticulations make the crest there dark, otherwise the crest is a light creamy colour along the margin and, in some fish without reticulations reaching the margin, the whole edge is light. The lateral line is white, sometimes in marked contrast to the rest of the flank. The dorsal fin has darkly pigmented rays, sometimes broken into series of spots. The caudal fin has a series of 4-5 small spots elongated along the rays, the middle series being the blackest. The dorsal margin of the caudal fin may have 2-4 isolated spots. The pectoral fin has dark pigment along the rays or 2-3 series of small spots. The pelvic and anal fins have 1-2 series of grey spots and the pelvic fin may have only 1-2 spots. Pelvic and anal fins may be immaculate. At the front or along the base of the anal fin there is a broad pigmented band. There may be a dark, zig-zag bar at the caudal base, merging dorsally and/or ventrally with flank botches. The barbel bases are all darkly pigmented especially the second pair. The iris is silvery. Young fish have a less reticulated pattern with more blotches on the flank.

Size

Reaches 15.0 cm.

Distribution

This species is found in the Caspian Sea basin of Iran in rivers from the Safid to the Atrak including the Qezel Owzan, Sardab, Haraz, Ramian, Neka, Chalus, Tonekabon, Shahrud, Babol, Tajan, Karasu, Madar Su, Gorgan rivers, Sarchem near the Zanjan-Safid river junction, Bandar Gaz in Gorgan Bay (Laptev, 1934; Nikol'skii, 1947; Kiabi et al., 1994; 1999; Jolodar and Abdoli, 2004; Abdoli and Naderi, 2009; A. Abdoli, pers. comm.). Material from the Kor River and Esfahan basins may be this species.

Need to be checked and re-mapped?

Zoogeography

See family account.

Habitat

This species appears to favour deeper water and stronger current than other sympatric loaches.

Age and growth

Tabiee and Abdoli (2005) found a sex ratio of 4:1 (male:female) in the Zarringol River of Golestan Province. Average total length was 59.07 mm for males and 82.42 mm for females. Condition factor was 2.6 for females and 1.97 for males. Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 48 Iranian fish measuring 2.67-7.43 cm standard length. The a-value was 0.0126 and the b-value 2.628 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases). Another study in the Zarringol River by Patimar et al. (2009) found maximum ages of 3+ years for males and 4+ years for females, length-weight relationship was W = 0.020L2.62 for males, 0.002L3.81 for females and 0 .008L3.08 for the population, growth was positively allometric, and sex ratio was 1:1.27 in favour of females (cf. Tabiee and Abdoli, 2005). The difference with Tabiee and Abdoli (2005) who found negative allometric growth was attributed to environmental conditions changing between years of sampling.

Food

The Zarringol fish were carnivorous with chironomids making up 65.3% of the diet (Tabiee and Abdoli, 2005) or had a diet dominated by Plecoptera (72.4% and Trichoptera (20.8%) (Patimar et al., 2009).

Reproduction

The Zarringol River examined by Patimar et al. (2009) reproduced in April-June with highest GSI in May. Spawning was prolonged, over about 2 months. Absolute fecundity reached 1180 eggs with a maximum diameter of 2.8 mm.

Parasites and predators

A Diplostomum species is recorded from the muscles of this nemacheilid in the Shirud of Mazandaran (Alghmandi and Dalimi Asi, 2000).

Economic importance

Kiabi et al. (1999) consider this species to be of least concern in the south Caspian Sea basin according to IUCN criteria. Criteria include medium in numbers, habitat destruction, widespread range (75% of water bodies), present in other water bodies in Iran, and present outside the Caspian Sea basin.

Conservation

Little is known of the biology and population numbers so no conservation assessment can be made.

Further work

The systematics of this species and its relatives need further study.

Sources

?check for newer species from Bianco and Nalbant

Iranian material: CMNFI 1977-0510A, 2, 74.2-81.5 mm standard length, ?; CMNFI 1979-0252, 1, 53.7 mm standard length, Markazi, jube at Baqerabad (34º55'N, 50º50'E); CMNFI 1979-0253, 1, 58.9 mm standard length, Markazi, Qareh Chay west of Baqerabad (34º52'N, 50º49'E); CMNFI 1979-0481, 1, 77.9 mm standard length, Mazandaran, stream 3 km west of Ghalahleekesh (37º18'30"N, 55º31'E); CMNFI 1979-0486, 18, 12.7-63.2 mm standard length, Mazandaran, stream in Atrak River drainage (37º44'N, 56º18'E); CMNFI 1979-0489, 2, 27.4-45.3 mm standard length, Mazandaran, stream in Atrak River drainage (37º50'N, 55º53'E); CMNFI 1980-0154, 8, 45.8-80.8 mm standard length, Markazi, Karaj River below village (35º47'N, 50º58'E); CMNFI 1980-0156, 24, 28.4-63.6 mm standard length, Markazi, Karaj River near village (35º47'N, 50º58'E); CMNFI 1980-0160, 3, 35.4-40.0 m standard length, ? CMNFI 1991-0157, 2, 70.4-73.0 mm standard length, Mazandaran, Ramian River (36º58'N, 55º07'E); CMNFI 1993-0145, 2, ? mm standard length, Mazandaran, Qareh Su (no other locality data); CMNFI 1993-0155, 2, 53.4-58.7 mm standard length, Markazi, Sharra River near Khosbijan (34º07'N, 49º23'E); CMNFI 2007-0121, 5, 45.3-98.3 mm standard length, Hamadan, Qareh Su basin north of Razan (ca. 35º25'N, ca. 49º02'E); USNM 205921-22, 6, 25.9-34.6 mm standard length, Markazi, Baragon River (ca. 36º00'N, ca. 50º50'E); BWC 1995-0009, 2, 83.4-83.9 mm standard length, Gorgan River from Kiabi?

Comparative material: ZISP 25788, 1, 62.2 mm standard length, ?, Sumbar River at Aidere (?).

Paracobitis rhadinaea
(Regan, 1906)

Common names

See genus account. Fowler and Steinitz (1956) refer to a fish from Sistan known locally as mahrmahé (sic, presumably mar mahi, snake fish) and this may refer to this species which has an elongate body.

Systematics

Bănărescu and Nalbant (1995) and Nalbant and Bianco (1998) place this species in Paracobitis.

Nemacheilus macmahoni Chaudhuri, 1909 described from the "affluents (= delta, an error for effluents) of the Helmand" is a synonym according to Bănărescu and Nalbant (1966) who refute the opinions of Nikol'skii (1947) and Berg (1948-1949; 1949) who consider macmahoni to be identical to P. malapterura. Earlier Banarescu and Nalbant (1964) placed fish from Sistan and the Caspian Sea basin of Iran as Nemacheilus malapterurus macmahoni. P. malapterura has both lips strongly furrowed, pelvic fin origin under the dorsal fin origin rather than behind, better developed scales which are also present on mid-flank, and a colour pattern of numerous oblique bands.

P. rhadinaea is distinguished from macmahoni by Annandale and Hora (1920) in having an extremely short posterior diverticulum and minute vesicle in the swimbladder, by the absence of scales, a more elongate body, smaller, narrower and less flattened head, and by differences in the profile of the body.

A syntype of Nemacheilus rhadinæus (ZSI F1240/1) is in the Zoological Survey of India, Calcutta under the name Adiposia rhadinaea and the holotype of Nemacheilus macmahoni (ZSI F1222/1) is also there under the name Adiposia macmahoni (Menon and Yazdani, 1968). Two syntypes listed as Nemacheilus rhadinaeus from "Sistan" are in the Natural History Museum, London (BM(NH) 1905.11.29:28-29, 2, 137.8-209.1 mm standard length).

A specimen in the Zoological Institute, St. Petersburg (ZISP 24413, 76.5 mm standard length) is from "Helmand delta, northwest of Jalalabad, Seistan, XI 1918, Indian Mus. (Dr. Hora)" according to Berg (1949) and could be a syntype of Nemacheilus macmahoni (Eschmeyer's "Catalog of Fishes", downloaded 20 May 2008). However, this taxon was described from a single specimen ? . The jar bears a label on the outside reading "N. malapterurus XI 1918 Indian Museum S. L. Hora Delta of Helmand near Jalabad" and another label reads "Adiposia macmahoni Randa stream 4 mls N.W. of Jalabad Seistan. N. Annandale coll."

Adiposia Annandale and Hora, 1920 (type species Nemachilus macmahoni Chaudhuri, 1909, and including Nemacheilus longicaudus and N. rhadinaeus according to Annandale and Hora (1920)) is a synonym of Paracobitis.

Key characters

?

Morphology

macmahoni:- ? check The head is broad and flattened and the small eyes are dorsal. The dorsal profile is slightly convex behind the head but the dorsal and ventral profiles soon become straight and nearly parallel, or thicker in front and tapering behind the dorsal fin. Body depth 5.8 in total length. Caudal fin rounded. Oval scales are found on the posterior part of the body in adults (not in rhadinaeus in Regan (1906) and Annandale and Hora (1920)) or scales very small and deciduous all over the body. Nostrils are nearer to the eye than the snout tip. The two anterior pairs of barbels reach back to the nostril level and the posterior pair to the anterior or to middle or to the rear of the eye. The upper lip is minutely tubercular and the lower lip is widely interrupted in the middle. Scale radii are few and widely spaced, on all fields. Material identified originally as macmahoni has the pelvic origin behind that of the dorsal fin origin level, the caudal fin is slightly emarginate, the dorsal fin edge is straight, the anus is some distance in front of the anal fin, a well-developed adipose dorsal ridge runs from the dorsal fin to the caudal fin base, the lateral line is almost complete, scales are restricted to the last third of the body, being small, rounded and far apart, lips are almost smooth, and the intestine has a single loop posteriorly. The description of rhadinaeus is short. The snout is longer than the postorbital distance, body depth is 7-10 times in body length, head length 5.0-5.5 times in body length, the mouth cleft extends to below the nostrils, lower lip interrupted medially, outer rostral barbel as long as maxillary barbel reaching back to or beyond nostrils, no scales, dorsal fin origin nearer tip of snout than caudal fin base, caudal fin slightly emarginate, caudal peduncle 2.0-2.75 as long as deep, 5.0-5.3 in length of fish.

Dorsal fin with 2-3 unbranched and 7 branched rays. Anal fin with 2-3 unbranched and 5 branched rays. Pectoral fin branched rays 10 and pelvic fin branched rays 6-8. Scales are highly deciduous and not always present on old preserved material. The dorsal fin rounded. There is a well-developed post-dorsal fin crest and a slight ventral crest on the caudal peduncle. The pelvic fin origin lies just in front of the mid-point of the dorsal fin base. There is an adipose tissue flap at the pelvic fin base. The anterior nostril is a tube followed immediately by a horizontal slit. The bony upper jaw has a slight protuberance and the lower jaw is curved and not indented.

Meristic values for Iranian specimens including types and macmahoni are:- dorsal fin branched rays 7(33) or 8(1), anal fin branched rays 5(34), pectoral fin branched rays 9(2) or 10(32) and pelvic fin branched rays 6(2) or 7(32).

Sexual dimorphism

None reported, the pectoral fin being identical in both sexes in material identified as macmahoni (Bănărescu and Nalbant, 1966).

Colour

Living fish identified as macmahoni are pale olivaceous fading to silvery-white on the belly. The head and upper part of the body are irregularly spotted and darker. Some fish are pale yellowish without markings or with faint marbling. All fins are tinged a dull red, most evidently on the caudal fin, and are obscurely marked with small dark spots. There is a dark band at the caudal base. Preserved material identified originally as macmahoni is whitish with 9-21 irregular, brownish spots along the flank, other spots are present dorsally and smaller ones between the dorsal and lateral rows. There are 3 rows of minute spots on the dorsal and caudal fins and 2 rows on the pectoral fin (Bănărescu and Nalbant, 1966). Types of macmahoni are brown all over, darker dorsally, barbels a lighter brown, dorsal and caudal fins with darker bands, pectoral also slightly banded but anal and caudal uniform light brown. P. rhadinaea has large oblong or rounded dark spots on the back and sides, dorsal and caudal fins with small spots and red tinged, lower fins pale and immaculate although pectoral, and to a lesser extent pelvic, fins may be red tinged.

Size

Attains 28.8 cm as macmahoni.

Distribution

This species is probably restricted to the Sistan basin of Iran and presumably Afghanistan. Bănărescu and Nalbant (1966) place this species in the Atrak and Safid rivers of the Caspian Sea basin, the Abkhar River of central Iran, probably most of Iran, the Helmand drainage and the Tedzhen River, evidently confusing it with P. malapterura and P. iranica. Abdoli (2000) lists as questionably from the Bejestan, Kerman-Na'in and Dasht-e Lut basins, from the middle and lower Halil and Bampur rivers of the Hamun-e Jaz Murian basin, and from the Simish and the river to its north in the Mashkid River basin.

Zoogeography

The closest relatives of this species are P. malapterura and P. longicauda (q.v.) in Iran (Bănărescu and Nalbant, 1966).

Habitat

Annandale and Hora (1920) note that Adiposia macmahoni was healthy buried some inches in mud when cyprinids died in foul water above.

Age and growth

Unknown.

Food

Stomach contents include cyprinid fish remains and mayfly larvae (Annandale, 1921).

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

Unknown.

Conservation

?

Further work

?

Sources

Type material: Syntypes of Nemacheilus rhadinæus (BM(NH) 1909.11.29:28-29), see above.

Iranian material: CMNFI 1979-0222, 11, 16.8-28.8 mm standard length, Sistan, jube 2 km south of Lutak (30º46'30"N, 61º24'E); CMNFI 1979-0223, 1, 23.9 mm standard length, Sistan, ditch 1 km south of Lutak (30º45'N, 61º24'E); CMNFI 1979-0228, 3, 55.1-119.8 mm standard length, Sistan, ditch 1 km from Zabol (31º02'30"N, 61º31'E); CMNFI 1979-0229, 3, 87.4-115.5 mm standard length, Sistan, ditch 5 km from Zabol (31º03'N, 61º33'E); CMNFI 1979-0231, 1, 22.6 mm standard length, Sistan, jube 3 km from Zabol (31º01'N, 61º32'E); CMNFI 1979-0232, 2, 82.5-95.6 mm standard length, Sistan, jube 11 km from Zabol (ca. 30º58'30"N, ca. 61º36'E); CMNFI 1979-0233, 1, 68.7 mm standard length, Sistan, ditch at Deh Vazi (ca. 30º57'N, ca. 61º38'E); CMNFI 1979-0234, 1, 85.5 mm standard length, Sistan, effluent of Hirmand River near Zahak (30º54'N, 61º40'E); CMNFI 1979-0237, 2, 17.5-25.2 mm standard length, Sistan, ditch 18 km south of Zabol (30º53'N, 61º27'30"E); CMNFI 1979-0238, 2, 23.6-27.7 mm standard length, Sistan, ditch 11 km south of Zabol (30º57'N, 61º27'30"E); BM(NH) 1920.1.20:31-34, 4, 79.6-109.5 mm standard length, Sistan, northwest of Jalalabad (no other locality data); ZISP 24413, 1, 76.5 mm standard length, Sistan, Randa stream 4 miles northwest of Jalalabad (?).

Paracobitis smithi
(
Greenwood, 1976)

Dorsal view of head                                             Ventral view of head

   

Photographs courtesy of Amir Hosin Zalaghi, 11-19 May 2010

Common names

sagmahi-ye gharzi (= cave loach), mahi kurghar (= cave fish).

[blind loach].

Systematics

This species is named for Anthony Smith who collected the holotype. Accounts of Smith's searches for cave fishes in Iran are given in his two books (Smith, 1953; 1979). The holotype is in the Natural History Museum, London under BM(NH) 1976.6.28:1, was collected in April 1976, is apparently an immature male and is 35.5 mm standard length. Locality data is given below.

Greenwood (1976) placed this species in the catchall genus Noemacheilus (correctly spelled Nemacheilus) pending an adequate revision of the subfamily Nemacheilinae and since little purpose would be served by erecting a new genus for a fish with such features as eyelessness and depigmentation found in common with other unrelated cave-dwelling fishes. Greenwood (1976) considers its relationship to lie with species in the subgenus or genus Paracobitis as does Nalbant and Bianco (1998).

Key characters

The only eyeless, depigmented nemacheilid reported from Iran.

Morphology

Dorsal fin with 3 unbranched and 7 branched rays, anal fin with 3 unbranched and 5 branched rays, pectoral fin with 10 branched rays and pelvic fin with 5-6 branched rays. In specimens seen by me dorsal fin with 7(5) branched rays, anal fin with 5(5) branched rays, pectoral fin with 10(5) branched rays and pelvic fin with 5(5) branched rays. The caudal fin is unusual in having only 14(5) branched rays. Vertebrae 37-38. check on my fish? The barbels are short with the middle pair the longest. There are no scales. The lateral line is interrupted irregularly and there are more pores on the posterior half of the body. A long adipose fin is present dorsally, most obvious in young, and a weaker ventral ridge is present. The mouth is subterminal, lips are weakly to moderately papillose, and there is a horny ridge on the dentaries and on the upper jaw. The middle pair of barbels is the longest. The internarial flap is long. Gill membranes are broadly attached to the isthmus. The gut is short with a single transverse loop.

Sexual dimorphism

Unknown.

Colour

A dead white with the red of blood visible as a pale pink cast and a deeper red at the gills visible through the gill cover. Small, straw-coloured fat globules are visible under the skin of formalin preserved fish over the whole head and body with concentrations in the orbits and the fin bases, particularly the dorsal and anal bases. The viscera are visible through the body wall. There is no peritoneal pigment.

Size

Attains 64.5 mm total length.

Distribution

Found only at "Kaaje-ru" above the garden "Bagh-e Loveh", "Lowa" or "Levan" (probably Loven at 33°04'N, 48°37'E) which is about 4 km from kilometre 382 on the railway from Bandar Shapur and approximately 12 km north of the railway station Tang-e Haft. The stream below the cave locality is the "Ab-e Serum" which runs into the "Ab-e Zezar" which is a tributary of the Dez River. The locality is at 33°04'38.6"N, 48°35'33.1"E in Lorestan Province. Further locality details are given in Bruun and Kaiser (1948) and under the cyprinid Iranocypris typhlops.

Zoogeography

Endemic to Iran, its relationships to other species are unknown. It shares the cave habitat with another eyeless species, Iranocypris typhlops, a member of the family Cyprinidae. This blind cave species is placed in a world-wide context by Proudlove (1997a; 1997b).

Habitat

Known only from a well-like but natural outlet of a subterranean system. The outlet overflows to form a small stream from January to May (Smith, 1979) during the snow-melt period in the Zagros Mountains but in April to June this flow ceases (the precise timing of flow and its cessation is estimated from villager's comments and scientific visits and also varies with precipitation). The pictures above show flowing water in May 2010. The well area is about 5 by 3 m and gradually decreases as the year progresses. Divers descended to a depth of 60 feet (= 18.3 m) in 1977 in the "well" until the resurgence narrowed (Farr, 1977). A rope was let down by R. Mehrani (pers. comm., 2000) and reached 23 m before the rope ran out and yet it was not at the bottom. Smith (1979) reports divers descending to 60-70 feet (18.3- 21.3 m). The pool shelves deeply under the cliff rearwards but the whole pool surface is exposed to light. There is no vegetation in the pool except for some encrusting algae on the rocky sides. The shale fragments forming the outermost floor of the pool have a thin layer of mud on them which may contain algae.

It seems probable that a complex of flooded but narrow and inaccessible passages is the habitat of this species and the well is merely the surface manifestation of this complex (Bruun and Kaiser, 1948; Smith, 1978; Banister, 1992). There is a smaller pool (about 2 m across narrowing rapidly inside) and flowing exit stream lower down the gorge, about 50 m away from the main locality, where an Iranocypris typhlops was seen but not caught in December 2000 (Smith (1979) also tentatively reports sighting a fish here). This is assumed to be evidence of the interconnectivity of subterranean passages. The main pool was not flowing at this time. The stream from the smaller pool increases in flow downstream, possibly tapping more groundwater, and eventually has a moderate flow. No fish were seen in it. The stream falls over a high waterfall (estimated at 10-15 m high by Smith (1979) which seems about right) so the well localities are isolated from the local fishes in the main river. The main river houses Garra rufa and Nemacheilus species s.l.. The stream shows evidence of recent higher flow which tends to confirm overflow from the main well.

Sampling in December 2000 recorded a water temperature of 18.5°C, pH 7.5 and a conductivity of 334 µS. More photographs of the habitat can be seen under the account of Iranocypris typhlops (Cyprinidae).

A captive specimen attempted to climb out of a glass tank, almost its whole body being out of the water before it slid back. It did not hide from or react to light and spent most of the time resting on the tank bottom, moving along the bottom or more often swimming actively around the tank (Greenwood, 1976). Amir Hosin Zalaghi reports that this species swims faster than Iranocypis tyhplops (pers. comm., 10 August 2010).

 

 

Photographs courtesy of Amir Hosin Zalaghi, 11-19 May 2010

Age and growth

Unknown.

Food

Unknown but a captive specimen was fed on mosquito larvae.

Reproduction

Unknown.

Parasites and predators

Unknown but there are probably no predators in the cave environment.

Economic importance

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in textbooks.

Conservation

A fine of 10,000 rials is imposed specifically for illegal fishing of this species (Anonymous, 1977-1978), more recently 100,000 rials (N. Najfpour pers. comm., 2008). It is on the IUCN 1994 Red List of Threatened Animals as one of two rare fish species from Iran (see also Iranocypris typhlops) and is on the 2000 IUCN Red List as VU D2 (Vulnerable, acute restriction in its area of occupancy, also on subsequent Red Lists; see also Proudlove (2001)). Coad (2000a), using 18 criteria, found this species to be one of the top 4 threatened species of freshwater fishes in Iran.

B. Sandford (in litt., 1979) considered this fish to be endangered. The cave appeared to be a recently collapsed system and the network of fissures could be quite small. Coupled with recent collecting the number of extant specimens may be quite low.

This species is much rarer than the co-occurring Iranocypris typhlops, by at least an order of magnitude. Including the holotype, about 14 specimens are known to have been collected (see below; plus 2 in Muze-ye Melli-ye Tarikh-e Tabi'i, Tehran (MMTT 1227-1228) and 6 collected by students of A. Abdoli, Shahid Beheshti University, Tehran in 2001).

Further work

?

Sources

Type material: Holotype of Noemacheilus smithi (BM(NH) 1976.6.28:1), see above.

Iranian material: CMNFI 2007-0123, 5, 24.1-52.9 mm standard length, type locality, 28 January 1977.

Further information on the habitat is in the account of Iranocypris typhlops.

Paracobitis vignai
Nalbant and Bianco, 1998

Common names

sagmahi-ye Sistan.

[Sistan loach].

Systematics

The species is named for Professor Augusto Vigna Taglianti, La Sapienza University, Rome. The holotype is 89.0 mm standard length (86.5 mm standard length when measured by me) collected from "Nahr Taheri, Zabol, Seistan" on 9 October 1977 by A. Vigna and is deposited in the (Department of Zoology, University of Naples (IZA 7838). Paratypes are from the same locality and number 24 specimens, 35.0-78.0 mm standard length (IZA 7839) and 7 specimens, 44.0-49.0 mm standard length (Institutul Stiinte Biologice, Bucharest, ISBB uncatalogued). One specimen is in the American Museum of Natural History (AMNH 40946), presumably a paratype from one of the preceding series. CMN fish?

Key characters

This species is scaleless and has a deeply forked caudal fin. The dorsal adipose crest or keel on the caudal peduncle is well-developed. The anus is placed well anterior to the anal fin origin. The dorsal fin has 3 unbranched and 7 branched rays, the anal fin has 2 unbranched and 5 branched rays, the pectoral fin has 1 unbranched and 9 branched rays and the pelvic fin has 1 unbranched and 7 branched rays.

Morphology

Dorsal fin branched rays 6-8 (80% with 7 rays in original description, n = 20), anal fin with 5-6 branched rays (95% with 5 rays), pectoral fin with 8-10 branched rays (85% with 9 rays) and pelvic fin with 6-7 branched rays (95% with 7 branched rays). The lateral line extends to the base of the caudal fin. The body is slender and compressed, particularly posteriorly. The head is long and the eyes are small. The dorsal fin origin is at mid-body (snout tip to caudal fin base). The mouth is arched with strongly furrowed lips that have few papillae. Mental lobes are reduced. The stomach is syphonal and the intestine is straight. The gas bladder capsule has globular chambers and a short duct.

Sexual dimorphism

Unknown.

Colour

The head and body are greyish, lighter ventrally. Dark spots along the back and flank may be occasionally fused. Fine spots are scattered in a reticulated pattern on the body. The dorsal and caudal fins have dots in rows while other fins are colourless. The caudal fin base has a distinct blackish bar.

Size

Reaches 89.0 mm standard length.

Distribution

Endemic to Sistan.

Zoogeography

Nalbant and Bianco (1998) consider this species to be an epigean form of the blind cave fish P. smithi. The differences are in head shape and mouth and lip morphology, apart from the eyes and pigment loss typical of ipogean P. smithi.

Habitat

Details are unknown.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

None.

Conservation

Population trends and numbers unknown.

Further work

More collections are needed to record information on biology and distribution.

Sources

Type material: The holotype ((IZA 7838)) and paratypes (IZA 7839) of Paracobitis vignai, see above. + ?CMNFI paratypes

Genus Paraschistura
Prokofiev, 2009

This genus is recently described and not all species have been examined and ascribed to it or related genera. Many were formerly included in the genus Schistura McClelland, 1838.

The species in this genus are small and are distributed from interior water bodies of Turkmenistan and from Iranian Baluchestan east to the upper reaches of the Indus River in Afghanistan and Pakistan. Four species are reported from Iran.

The genus is characterised by an elongate body with dorsal and ventral profiles almost parallel. The belly is not rounded. The body is compressed anteriorly and the head is compressed or depressed. The caudal peduncle is short and moderately deep. The snout is usually blunt. The upper lip is furrowed and is continuous or has a slight median interruption. Lips are simple, being plicate to almost smooth. The processus dentiformis on the upper jaw is well-developed in many species and there is a corresponding notch on the lower jaw but the processus can be absent. Nostrils are closely spaced. The dorsal fin usually has 7 branched rays and the anal fin 5 branched rays. The caudal fin is slightly to deeply emarginated. There is no adipose crest or, if present, it is weakly-developed near the caudal fin and not supported by procurrent rays of the caudal fin. The lateral line is incomplete, may be very short and rarely reaches the end of the anal fin level. Scales are absent or rarely present but weakly developed on the caudal peduncle. A pelvic axillary lobe is present but is short and does not usually extend beyond the pelvic fin base.

Prokofiev (2009) lists various osteological characters that help diagnose the genus including the absence of a bony bridge between the parietal and pterotic bones which separates the species from Schistura sensu lato. There is no preethmoid I bone in the skull. Externally, there is no bony roof to cst, the fork of cst-cio is not ossified and it is not fused to the skull. The cst is the supratemporal commissure running across the top of the head at the rear and cio is the temporal part of the infraorbital head canal. The free part of the swimbladder is absent, the right and left lobes of its bony capsule are fully divided by the manubrium, and the fifth trunk vertebra does not take part in support of the capsule.

There is no sexual dimorphism. Colour pattern is distinctive with dark bars of varying number and form being present. The bar at the caudal fin base is usually much darker than anterior bars, and is never in the form of a spot. There is no stripe along the flank although a dark line is rarely present. A dark black spot or strip is always present at the base of the anterior dorsal fin rays.

Paraschistura alta
(Nalbant and Bianco, 1998)

Described from "Afghanistan, Kajkai, Helmand river drainage, north east of Girisk". No Iranian record.

?genus as sexually dimoprhic see ProkofievSchistura baluchiorum
(Zugmayer, 1912)

Possibly a synonym of S. bampurensis. The type locality is Panjgur on the Rakhshan River of the Hamun-i Mashkel basin in Pakistani Baluchistan and it is also recorded from Afghanistan in the Helmand River drainage. A cotype is in the Naturhistorisches Museum Wien under NMW 19851 and is 46.0 mm standard length. Not recorded from Iran by specimens although Abdoli (2000) reports it from the Simish River and a river just north of it, both in the Mashkel basin.

Paraschistura bampurensis
(Nikol'skii, 1899)

Common names

sagmahi-ye Bampur.

[Bampur loach]

Systematics

Publication date is given as 1900 in Berg (1949) and Bănărescu and Nalbant (1966). This may be correct if the volume appeared late as the volume of the publication is for the year 1899; but note that a footnote and the plate both bear the year 1899.

Nemacheilus baluchiorum Zugmayer, 1912 is possibly a synonym (Berg (1949) places this species in synonymy with Nemacheilus montanus - see below). Bănărescu and Nalbant (1966) and Nalbant and Bianco (1998) place S. bampurensis in the subgenus or genus Schistura.? read description carefully and comment

Berg (1949) placed S. bampurensis in Nemacheilus montanus (McClelland, 1839) but Bănărescu and Nalbant (1966) recognise bampurensis as distinct on its incomplete lateral line, focal zone of scales larger, dorsal fin positioned more anteriorly, deeper body, and colour pattern without regular bands (Bănărescu and Nalbant (1966: fig 12) for three pattern varieties). Observations were made by me on the faded syntypes of N. montanus from Simla, 62.8-68.7 mm standard length (BM(NH) 1860.3.19:118-119) as well as ZISP 8298, 9 specimens 25.3-51.1 mm standard length. Additional characters which distinguish bampurensis and montanus are as follows. The preorbital process is very strongly developed in N. montanus, being almost as deep as the eye and extending almost an eye length below the lower orbit margin, extending almost twice the distance of the process in bampurensis types and being less curved and close under the eye. N. montanus has a slight but obvious keel on the back before the caudal fin while bampurensis are humped there but usually not keeled. The caudal fin is dark at the base in montanus, not so in bampurensis or not as solid, wide and dark. The flank bars are oblique with the top forward in contrast to bampurensis, and are less numerous than in bampurensis (5-10 versus 11-22 total, from under dorsal fin insertion to caudal fin but excluding any caudal fin base bar). N. montanus types have faded bars but are clearly fewer.

The syntypes of Nemacheilus bampurensis are in the Zoological Institute, St. Petersburg under catalogue numbers ZISP 11698 and 11699 at the localities, dates and number of specimens in Latin as follows respectively:- "Kjagur prope urb. Bazman. 4. VII (6)" and "Kaskin prope urb. Bazman. 6.VII (4)" (Nikol'skii, 1899). Berg (1949) gives both these localities as between Bazman and Bampur. However, ZISP 11698 comprises 9 specimens, 35.1-44.6 mm standard length and ZISP 11699 comprises 4 specimens not listed as types on the jar, 36.8-44.5 mm standard length.

Key characters

The male has a characteristic, moveable protuberance directed downward on the preorbital bone at the antero-ventral corner of the eye. It lies close to the eye, extends slightly below the lower orbit, and curves partly around the orbit.

Morphology

Dorsal fin with 2-3 unbranched and 6-7 branched rays, anal fin with 2-3 unbranched and 4-5 branched rays. Pelvic fin branched rays 6-7. Lateral line incomplete, ending in front of the dorsal fin level. Scales well-developed but the anterior third to a quarter of the body is scaleless. The posterior nostril is ovoid, slanting postero-dorsally. Barbels are large and the third pair is usually the largest although in some fish the second pair is the largest (Saadati, 1977). The lower lip is divided, lips are corrugated, and the upper jaw process is rounded, overlapping the lower one. There is a fleshy pelvic axillary process. Caudal fin slightly emarginate.

The type series (ZISP 11698) has dorsal fin branched rays 6(3) or 7(6); anal fin branched rays 4(1) or 5(8); pectoral fin branched rays 9(8) or 10(1); pelvic fin branched rays 6(2) or 7(7); and flank bars 14(2), 15(3), 17(1), 19(2) or 22(1).

Meristic counts for Iranian material:- Dorsal fin branched rays 6(3) or 7(73), anal fin branched rays 4(1) or usually 5, pectoral fin branched rays 8(2), 9(59) or 10 (10), pelvic fin branched rays 6(7), 7(51) or 8(2), and flank bars 10(1), 11(7), 12(6), 13(12), 14(6), 15(8), 16(3), 17(1), 18(2), 19(2) and 22(1).

Sexual dimorphism

See above under Key characters. Males also have tubercles on the pectoral fin rays and on the operculum (see figure in Berg (1949)). The first branched pectoral fin ray is greatly expanded, 2-3 times broader than the second ray, which itself may be expanded a little.

Colour

Body yellowish to a light olive-green with 9-18 dark, brownish or chestnut-brown bands. The caudal fin has 3-4 dark wavy bars and a bar at its base, the dorsal fin 2-3 dark bars and a black spot at the anterior fin base. Fins are a light orange or pinkish, particularly the caudal fin.

Size

Reaches 5.3 cm.

Distribution

Berg (1949) and Abdoli (2000) record this species from the Dasht-e Lut basin without specific localities, questionably from the Tigris River basin (presumably based on Bănărescu and Nalbant (1966), see below), the middle and lower Bampur and Halil rivers of the Hamun-e Jaz Murian basin, and the Sarbaz and Nikshahr rivers of the eastern Makran. Specific localities include Kjagur and Kaskin near Bazman (Nikol'skii, 1899), both between Bazman and Bampur (Berg, 1949), Shur Ab near Kuh-i Birk (Berg (1949) and the Bampur River near Bazman.

Bănărescu and Nalbant (1966) report this species from Shapur, 12 km north of Kazerun and from Shah Bazan on a tributary of the Ab-i-Diz, and "probably most of Iran". This seemed inherently unlikely as the type locality is in Baluchestan and the material was later described as a new species, S. nielseni q.v.

Zoogeography

See family account.

Habitat

Unknown.

Age and growth

Unknown.

Food

Unknown.

Reproduction

A female measuring 4.8 cm and caught in late February carried fairly well-developed eggs (Berg, 1949).

Parasites and predators

Unknown.

Econmic importance

None.

Conservation

This species is reported from several localities in Baluchestan, including those remote from human influence, and does not seem to be in any danger.

Further work

The biology of this species needs to be studied.

Sources

Type material: Syntypes of Nemacheilus bampurensis (ZISP 11698 and 11699), see above.

Iranian material: CMNFI 1979-0312, 14, 24.9-41.5 mm standard length, Baluchestan, Bampur River 8 km west of Iranshahr (27º11'N, 60º36'E); CMNFI 1979-0313, 5, 31.1-39.9 mm standard length, Baluchestan, Bampur River at Bangharabad (27º20'N, 60º46'E); CMNFI 1979-0315, 17, 20.0-44.8 mm standard length, Baluchestan, Bampur River 2 km north of Karevandar (27º51'N, 60º46'E); CMNFI 1979-0317, 28, 19.9-45.8 mm standard length, Baluchestan, Sarbaz River at Bondan (26º35'N, 61º13'E); CMNFI 1979-0318, 1, 23.8 mm standard length, Baluchestan, Sarbaz River at Huvar (26º09'N, 61º27'E); CMNFI 1979-0323, 5, 20.7-31.4 mm standard length, Baluchestan, Sarbaz River (ca. 26º26'N, ca. 61º16'E); CMNFI 1979-0326, 12, 19.3-35.0 mm standard length, Baluchestan, stream in Oghin River drainage (ca. 26º35'N, ca. 60º02'E); CMNFI 1979-0327, 10, 21.2-37.4 mm standard length, Baluchestan, stream in Nikshahr River drainage (26º32'N, 59º57'E); CMNFI 1979-0329, 14, 21.1-33.4 mm standard length, Baluchestan, stream at Zaminbandan (27º02'N, 61º20'E); CMNFI 2007-0045, 6, 28.8-39.2 mm standard length, Kerman, Kharan River drainage at Baft (29º14'N, 56º38'E)checkID

?Schistura chrysicristinae
Nalbant, 1998

Described from the Tigris River basin in Turkey but no Iranian record.

Paraschistura kessleri
(
Günther, 1889)

Common names

sagmahi-ye Bejestan, sagmahi-ye Kessler.

[golets kesslera or Kessler's loach in Russian; sundali in Pakistan].

Systematics

Nemacheilus prashari Hora, 1933 is possibly a synonym, along with its subspecies (q.v.). Banarescu and Mirza (1965) have comparisons of Nemacheilus lindbergi (which is later regarded as a subspecies of Nemacheilus (= Parachistura) prashari, q.v.) with P. kessleri where most characters overlap in varying degrees. Bănărescu and Nalbant (1966) place P. kessleri in the subgenus Schistura and Nalbant and Bianco (1998) place it in the genus Schistura. Berg (1949) refers P. prashari to a subspecies of P. kessleri.

Nemacheilus kessleri turcomanus Nikol'skii, 1947 is reported from Turkmenistan east of the Iranian border in the Murgab River drainage (Kushk River near Kushk). Bănărescu and Nalbant (1966) consider that the differences are too slight to warrant a separate subspecies.

Berg (1948-1949) considered S. kessleri to be close to S. sargadensis, differing in having the snout slanting steeply in front of the eyes (rounded in sargadensis - this is generally true but exceptions occur with sharply downturned snouts in sargadensis examined by me), dorsal origin midway between the snout tip and the caudal fin base (further back in sargadensis - but see below, same in both species; and in a series of sargadensis of varying size small fish have a longer predorsal distance than dorsal origin to caudal base distance, in large fish a shorter predorsal distance, and in intermediate-sized fish an about equal predorsal distance, so this character appears to be highly variable and unreliable for separating these species), rudimentary caudal peduncle crest (weak to clearly pronounced crest - ?check on my sargadensis), fewer and less sharply pronounced flank bars (10-12 total, broad bars in kessleri seen by me while sargadensis has 13 or more thinner bars - ?check on my sargadensis), longer intestine (?), and pelvic fin origin just behind the level of the dorsal fin origin (under dorsal fin origin or slightly in advance - kessleri types seen by me had the pelvic fin origin almost exactly under the dorsal fin origin). Generally S. sargadensis agrees with the description of S. kessleri except for pigmentation (?I need to run type measurements I have against sargadensis noting that Ghazni fish may be a different species and comparing gut length/shape and bladder capsule (see Mirza et al. (1981)).

Nemacheilus kessleri was described from Nushki in the Pishin Lora River basin of Afghanistan (Menon, 1987). Four syntypes are in the Natural History Museum, London (BM(NH) 1886.9.21:177-180, 38.3-48.4 mm standard length) and 4 syntypes (now 3) are in the Zoological Survey of India, Calcutta (ZSI 11487-11490) (Eschmeyer et al., 1996). The holotype of Nemacheilus kessleri turcomanus is in the Zoological Museum of Moscow State University (MMSU P.5734) with apparently a paratype under MMSU P.5734 and another paratype under MMSU P.5735 (Eschmeyer et al., 1996). The Zoological Museum of Moscow University (ZMMU; their acronym) has 3 specimens under P-5734 and 1 specimen under P-5735, all syntypes (Pavlinov and Borissenko, 2001).

Key characters

Characterised by Menon (1987) as a member of the subgenus Schistura with 7 branched dorsal fin rays, no scales, forked caudal fin, short lateral line not reaching mid-body and no sexual dimorphism, and additionally by and Mirza et al. (1981) and Bănărescu and Nalbant (1995) as a member of the genus Schistura part of the kessleri-lindbergi group with the above characters and also small size, rather compressed head, emarginate or slightly forked caudal fin and a reduced dentiform process.

Morphology

Dorsal fin unbranched rays 2-3, branched rays 7, anal fin unbranched rays 2-3, branched rays 5, pectoral fin branched rays 8-11, pelvic fin branched rays 6-7, usually 7. Dorsal fin origin midway between the nostrils and the caudal fin base according to Berg (1948-1949) but the types examined by me clearly have the dorsal fin origin closer to the snout (predorsal distance when stepped back from dorsal fin origin overlaps the rays of the caudal fin). Scales are absent. The lateral line ends under the dorsal fin (above the tip of the pectoral fin in Menon (1987) ? check on fish, Menon quote is correct). No adipose fin but a rudimentary crest may be evident. Caudal fin slightly emarginate to quite deeply forked. Pelvic fin origin somewhat behind the level of the dorsal fin origin. The snout is blunt; in the types it falls gently at the nostrils in 3 fish, abruptly in 1 fish; and in 3 fish from Ghazni, Afghanistan abruptly in 2 fish and less so in the third, all apparently independent of size. Both lips are fringed, the upper with a slight interruption, the lower with a wide interruption. The dentiform process is reduced (Berg (1949), well-developed in Menon (1987) reduced in Ban and Nal 1995 ? check on fish). Barbels are variably developed or absent in some individuals, a condition illustrated by Hora (1933). The posterior part of the intestine has a single loop.

Sexual dimorphism

There is no sexual dimorphism.

Colour

The back is crossed by 10-12 irregular, brownish bands. There is a dark spot at the base of the 3 anteriormost dorsal fin rays and there are 1-2 bands composed of small spots on the distal part of the fin. The anal fin may have an anterior basal spot. There is a narrow blackish band at the caudal fin base and 2 narrower bands on the fin itself. The sides of the head are minutely spotted. Bănărescu and Nalbant (1966) consider that the fish reported by Berg (1949: fig. 54) from Kelat Margh in Neh-i-Bendan do not have the typical colour pattern of this species. Berg (1949) describes 6-8 chestnut-brown bars on the posterior part of the body, none anteriorly.

Size

Reaches 8.7 cm total length (Patimar et al., 2010).

Distribution

Found from the Lora River basin of Pakistan through Afghanistan to the Murgab River basin of Turkmenistan and in eastern Iran at such localities as Keljate-Marg in Zirckuch region and the Neh-i Bendan district (?Berg check localities for modern spelling). For Iran, Abdoli (2000) lists middle and upper Kashaf River of the Tedzhen River basin, questionably in the Bejestan basin, and in the Simish and Mashkid rivers, and river north of them in the Mashkid River basin. Nalbant and Bianco (1998) also cite the Mashkel Lake, northern Sistan.

Zoogeography

See family account.

Habitat

Unknown.

Age and growth

Patimar et al. (2010) examined fish from the Zanglanlou River in northeast Iran  and found maximum age for both sexes was 4+ years. Growth was positively allometric and sex ratio was 1:1.2 in favour of females.

Food

Unknown.

Reproduction

Patimar et al. (2010) found reproduction occurred in April-May with the highest GSI in April. Maximum egg diameter was 1.46 mm, aboslute fecundity reached 1246 eggs and up to 1285.71 eggs in relation to total weight.

Parasites and predators

Unknown.

Economic importance

None.

Conservation

?

Further work

?

Sources

Type material: Syntypes of Nemacheilus kessleri (BM(NH) 1886.9.21:177-180), see above.

Comparative material: BM(NH) 1944.4.1:7-9, 3, 42.0-57.6 mm standard length, Afghanistan, Ghazni (?).

Schistura lindbergi ?genus
(Banarescu and Mirza, 1965)

Found in the Mashkel (= Mashkid) River basin of Pakistan on the southeastern border of Iran (see S. prashari for further discussion). This species was described from a rivulet connected to the drainage of the Farah River, Siaw, between Farah and Dilaram, Afghanistan. No Iranian record although Nalbant and Bianco (1998) record a specimen from near the Iranian border in the "Mashkel Lake system". Originally described in the genus Noemacheilus.

Schistura namiri ?genus see Tigris/Euphrates list
(
Krupp and Schneider, 1991)

Recorded from the Tigris River basin in Turkey by Nalbant (1998) but no Iranian record. Originally described in the genus Nemacheilus.

Paraschistura nielseni
(Nalbant and Bianco, 1998)

Common names

sagmahi-ye Nielsen.

[Nielsen's loach].

Systematics

Note that P 27110 (33.7 mm SL, my measurement here and below) "Shah Bazan, W. Iran, 16. IV. 1937. E. Kaiser" is listed as the holotype in the Zoological Museum, University of Copenhagen (ZMUC), not 27109 as in the original, published description (observed in 1999). The Shah Bazar locality in the description is presumably a misprint for Shah Bazan. Paratypes are 27109 (25.8 mm SL) from "Shah Bazan..." etc as above (probably reversed with the holotype), and 27105 "lok 44" (30.7 mm SL), 27106 "lok 54" (12.7 mm SL), 27107 "lok 54" (29.5 mm SL), 27117 "lok 54" (37.3 mm SL) from "Shapur, 15.03.1937. E. Kaiser. W. Iran". The published description gives the locality of Shapur as "12 km north-west of Kazerun".

The size of one paratype (12.7 mm) is smaller than any cited in the type description. The catalogue number of one fish in a drawing in the original description (i.e. 27108) is not in that description nor in jars in ZMUC. There has evidently been some confusion over the type series.

Named for Dr. Jorgen G. Nielsen, Department of Ichthyology, Zoological Museum, University of Copenhagen.

This species was previously reported as Nemacheilus bampurensis by Bănărescu and Nalbant (1966) based on this type series.

Key characters

This species has a relatively short body with a short head and blunt snout. The lateral line is incomplete ending just anterior to the dorsal fin origin. Scales have a large, eccentric focus. The caudal fin is distinctly emarginate. The body is yellowish with 7-16 dusky brown bars. The dorsal fin has 2 unbranched and 7 branched rays, the anal fin 2 unbranched and 5 branched rays, the pectoral fin has 1 unbranched and 9 branched rays and the pelvic fin has 1 unbranched and 7 branched rays.

Morphology

Dorsal fin branched rays 7(20), anal fin branched rays 5(20), pectoral fin branched rays 7(19) or 8(1) and pelvic fin rays 7(1), 8(3), 9(15) or 10(1) based on my collection. Meristics from type series dorsal fin branched rays 7(5), anal fin branched rays 5(5), pectoral fin branched rays 9(4), 10(1), pelvic fin branched rays 6(1), 7(4), and vertebrae 33(1), 34(4), 35(1). The body is compressed, especially posteriorly. The eyes are small and positioned in the middle or anterior portion of the head. The mouth is slightly arched and the upper jaw has a reduced processus dentiformis. Lips are slightly furrowed and the mental lobes are reduced. The maxillo-mandibular barbels are the longest. The nostrils are near the eyes with a relatively long tube on the anterior opening. The dorsal fin origin is at the middle of the body or slightly posterior. The pelvic fin insertion is just below the dorsal fin origin. The caudal fin is emarginate or slightly forked with rounded lobes. The anterior part of the body is scaleless. The stomach is syphonal and there is a single intestine loop. The gas bladder capsule has two globular chambers connected by a relatively long duct.

See BWC00-8 for more details on colour and sexual dimorphism and general description ?

Sexual dimorphism

A 37.3 mm standard length specimen from the type series (ZMUC 27117) has the first pectoral fin branched ray broadened with wide band of tubercles and rays 2-4 decreasingly tuberculate. This male has a preorbital prolongation. Paratype ZMUC 27105 has tubercles on up to 6 branched rays of the pectoral fin in broad bands, gradually decreasing in extent on more medial rays.

Colour

The head and body are yellowish, darker and more intense along the upper flank. Bars on the body are sometimes irregular or interrupted and number 7-17. The caudal fin has a bar at its base. The anterior dorsal fin base has a round dark spot.

Size

Reaches 54.9 mm total length and 45.0 mm standard length.

Distribution

This species is endemic to Iran, known from the Tigris River and Gulf basins as in the type description cited above (and my fish?)

Zoogeography

Nalbant and Bianco (1998) regard this species as the westernmost member of the genus Schistura. See also family account.

Habitat

Unknown.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

None.

Conservation

Population trends and numbers unknown.

Further work

More collections are needed to record information on biology and distribution.

Sources

Type material: The holotype (ZMUC P 27110) and paratypes (ZMUC P 27105, 27106, 27107, 27109, 27117) of Schistura nielseni, see above and note possible confusion in numbers.

Iranian material: BWC 00-8, 54, 27.5-45.0 mm standard length, ?

All fish are badly faded and can't compare with drawings in original description.

Compare measurements of snout, interorbital and HL from Bănărescu and Nalbant (1966) and Nalbant and Bianco (1998) with my Khuzestan fish - snout and interorbital appear different from my cf. tigris fish NEED to check again with my 2000 material from Khuzestan - see counts e.g.

?genus and search prashari here Schistura prashari
(Hora, 1933)

Reported from the Mashkel (= Mashkid) River basin of Pakistan on the southeastern border of Iran and from the Farah River basin in the Sistan drainage of Afghanistan as Nemacheilus prashari lindbergi Banarescu and Mirza, 1965 (or as a distinct species, lindbergi, in Schistura or Nemacheilus in Mirza, Nalbant and Banarescu (1981) and Menon (1987)) and from the Helmand River basin in the Sistan drainage of Afghanistan as Nemacheilus prashari haarlovi Bănărescu and Nalbant, 1966 (Banarescu and Mirza, 1965; Bănărescu and Nalbant, 1966; Mirza, Bănărescu and Nalbant, 1969; Coad, 1981c; Mirza, Nalbant and Banarescu, 1981). Possibly a synonym of S. kessleri as suggested by Berg (1949). No Iranian record.

Paraschistura sargadensis
(Nikol'skii, 1899)

Common names

sagmahi-ye Sarhad.

[Turkmenskii golets or Turkmen loach in Russian; perhaps Sarhadd loach should be used in English, see below].

Systematics

The syntypes of Nemacheilus sargadensis are in the Zoological Institute, St. Petersburg under catalogue number ZISP 11700 with locality data in Latin as "Sija-Rischan in sargado. 20.VIII (6)" (Nikol'skii, 1899) and in Russian in Berg (1949) "Zia-rishan, Province of Sarhad in Kerman, S.W. Iran, border with Beluchistan, not far from Kuh-e Taftan volcano, 1 IX 1898, N. Zarudnyi". The catalogue data in ZISP give the date as 20.VIII. There are 31 fish, 31.0-53.7 mm standard length, in rather poor shape, some stained dark brown, others uniformally decoloured or drab. Sarhad is the "frontier" or "land at the upper boundary", an area of plains or broad valleys and isolated mountains around Khash, for example, and a term used generally for the mountainous plateau of Baluchestan: Saadati's (1977) statement that the Sarhad is unlocatable is in error.

Nemachilus turcmenicus Berg, 1932 is described from "einem Bache unweit von der Eisenbahnstation Gjaurs in Turkmenistan (Transkaspien)" near the Iranian border and alternatively from the Kelte-chinar River (Cherokh River) near Gyaurs (37°47'N, 58°44'E), Turkmenistan (Catalog of Fishes). Three syntypes, 31.7-33.8 mm standard length, are in ZISP 11064. Berg (1948-1949) synonymises turcmenicus with sargadensis but Bănărescu and Nalbant (1966) consider it to be a valid subspecies, adding a third, paludani, from the Kabul River drainage of Afghanistan. Later Mirza, Nalbant and Banarescu (1981) separate paludani as a distinct species in a different species group from S. sargadensis, since S. sargadensis has scales, an almost forked caudal fin, a feeble dentiform process and irregular bars on the flank, all in contrast to S. paludani.

Berg (1948-1949; 1949) and Saadati (1977) consider this species to be close to S. kessleri (q.v.), possibly a synonym. Nalbant and Bianco (1998) place this species in the genus Schistura.

Key characters

The caudal peduncle is deeper than in Schistura bampurensis and it lacks the protuberance under the eye in males.

Morphology

Dorsal fin with 2-5 unbranched and 6-8 branched rays, anal fin with 2-3 unbranched and 4-6, usually 5, branched rays, pectoral fin branched rays 7-10, and pelvic fin branched rays 6-7 for S. sargadensis turcmenica (Berg, 1932a; 1948-1949; Nikolaev, 1993). Body scaleless (scaled according to Mirza, Nalbant and Banarescu (1981) based on fish from the Kul River, Hormozgan ? check on my fish as this would distinguish from kessleri). Caudal fin moderately emarginate (deeply emarginate, almost forked in Mirza, Nalbant and Banarescu (1981)). Caudal peduncle depth 1.6-2.2 in caudal peduncle length. Dorsal fin origin nearer caudal base than snout tip and slightly behind the pelvic fin origin. A dorsal adipose fin may be present or absent. Dentiform process on upper jaw, if present, rudimentary.

Meristics for Iranian specimens:- branched dorsal fin rays 6(1) or 7(105), anal fin branched rays 5(106), pectoral fin branched rays 9(11), 10(84) or 11(11), pelvic fin branched rays 6(1), 7(94) and 8(11), and flank bars 11(12), 12(8), 13(13), 14(11), 15(8), 16(5), 17(5) or 18(5).

Sexual dimorphism

Unknown.

Colour

Body colour is brown backed with greenish tinges or grey, with 13-14 dark bands, in some fish hardly visible (and irregular in form according to Mirza, Nalbant and Banarescu (1981)). The bands are broad, slightly broader than the interspaces. A dark stripe runs along the midline of the back. The caudal fin has 2 dark bars and the dorsal fin an anterior dark spot at its base. A dark band is present at the caudal base. Young fish have a row of dark speckles along the lateral line.

Size

Reaches 6.5 cm.

Distribution

Found in the streams of the northern slope of the Kopetdag in Turkmenistan as Nemacheilus sargadensis turcmenicus, possibly extending into northern Iran. In southern Iran it is found in the Hormuz, Hamun-e Jaz Murian, Hamun-e Mashkid and Makran basins (Nalbant and Bianco, 1998) including at Tangeh Sarreh, Baluchestan, Shah Abbas qanat at Assadabad and the Kul River near Darab.

Zoogeography

See family account.

Habitat

Unknown.

Age and growth

Maximum age in the subspecies turcmenica does not exceed 1.5 years (Nikolaev, 1993).

Food

Important food items in the subspecies turcmenica are larval mayflies, caddis flies, chironomids, dragonflies and detritus (Nikolaev, 1993).

Reproduction

Eggs in the subspecies turcmenica reach 1.24 mm in diameter (Nikolaev, 1993).

Parasites and predators

Unknown.

Economic importance

None.

Conservation

?

Further work

?

Sources

Type material: Syntypes of Nemacheilus sargadensis (ZISP 11700), see above.

Iranian material: CMNFI 1979-0210, 37, 21.0-35.4 mm standard length, Kerman, river west of Istor (29º21'N, 56º08'E); CMNFI 1979-0215, 14, 27.7-51.7 mm standard length, Kerman, Kharan River drainage at Baft (29º14'N, 56º37'E); CMNFI 1979-0217, 6, 30.1-41.8 mm standard length, Kerman, Kharan River drainage (ca. 28º59'30"N, ca. 56º51'30"E); CMNFI 1979-0218, 8, 32.8-47.8 mm standard length, Kerman, Kharan River drainage (28º53'N, 56º55'E); CMNFI 1979-0220, 1, 37.3 mm standard length, Kerman, jube 2 km south of Jiroft (28º39'N, 57º43'E); CMNFI 1979-0221, 19, 24.3-46.8 mm standard length, Kerman, river in Halil River drainage (28º51'N, 57º52'E); CMNFI 1979-0308, 11, 22.2-40.1 mm standard length, Kerman, river 44 km from Baft (29º02'N, 56º50'E); CMNFI 1979-0338, 68, ? mm standard length, Baluchestan, Tahlab River drainage 8 km from Mirjaveh (28º58'N, 61º24'E); CMNFI 1979-0339, 5, ? mm standard length, Baluchestan, Tahlab River drainage 16 km from Mirjaveh (28º56'30"N, 61º21'E); CMNFI 1979-0340, 25, ? mm standard length, Baluchestan, jube at Ladiz (28º55'N, 61º18'E); OSU 4270 ?

Genus Seminemacheilus
Banarescu and Nalbant, 1995

This genus has 3 species in Turkey and Iran, with one of these species and Iranian endemic.

The body is deep, more so than in other Iranian genera. The snout is blunt and the eyes are small. Nostrils are widely spaced and equal in size. The processus dentiformis is weak to absent. A crest on the caudal peduncle is rudimentary near the caudal fin base. The lateral line is short and does not reach beyond the end of the pectoral fin tip level. There are no scales and no pelvic axillary lobe. There are 7, rarely 8, branched dorsal fin rays and 5 branched anal fin rays. The pelvic fin insertion is opposite the dorsal fin origin. Sexual dimorphism consists of the anterior rays in the pectoral fin having isolated and small tubercles (not patches) and male pectoral fins are larger than in females. Prokofiev (209) lists various osteological features including the manubrium in the swimbladder being absent or weakly developed and the bony capsule swollen. 4 elongated-cylindrical radial bones in the  pectoral fin, no preethmoid I bone in the skull and 5 hypurals but no epural in the caudal fin support, among others.

Seminemacheilus tongiorgii
Nalbant and Bianco, 1998

Common names

sagmahi-ye Hormuz.

[Hormuz loach].

Systematics

This species is named for Professor Paolo Tongiorgi, University of Modena. The holotype measures 23.7 mm standard length and is from " large water spring near Darab town, Kul river basin", collected on 2 June 1976 by P. G. Bianco and is stored in the Department of Zoology, University of Naples (IZA 801 but not located in a 2002 visit by me). One paratype, 20.0 mm standard length, is from the same locality (Institutul Stiinte Biologice, Bucharest, ISBB uncatalogued). It differs from the Anatolian species S. lendlii (Hankó 1925) mostly by colour pattern and possession of scales.

Key characters

This species has a short and deep body. The mid-flank has a row of small scales. Esmaeili and Niknejad (2006-2007) give scanning electron micrographs of the scales along with a description.The processus dentiformis is absent. The lips are furrowed, particularly the lower lip. The gas bladder capsules are strongly united. The dorsal fin has 3 unbranched and 8 branched rays, the anal fin has 3 unbranched and 5 branched rays, the pectoral fin has 2 unbranched and 9 branched rays and the pelvic fin has 1 unbranched and 6 branched rays.

Morphology

The head is wide with large eyes and nostrils just anterior to the eyes. The mouth is arched and the lower lip has large lobules but the mental lobes are reduced. The gut has a short oesophagus and the stomach and intestine have one loop. The gas bladder capsule valves are proximal. The lateral line is short and does not extend posterior to the level of the pectoral fin. The body is almost entirely scaleless except for minute scales on mid-flank.

Sexual dimorphism

Unknown.

Colour

The overall colour is yellowish-white. There is a row of grey blotches along the mid-flank and a row of grey dots on the posterior half of the belly. All fins have grey dots except the whitish pelvics. The peritoneum is silvery-white with scattered melanophores.

Size

Attains 59.3 mm standard length.

Distribution

Known from Iran in the upper reaches of the Hormuz basin near Darab, in the Kor River basin and in northwestern Iran (Nalbant et al., 2010; Esmaeili et al., 2011?).

Zoogeography

The limited distribution of this endemic and lack of information on relationships preclude zoogeographical analysis. See family account.

Habitat

Details are unknown.

Age and growth

Unknown. Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 45 fish measuring 2.95-5.93 cm standard length. The a-value was 0.0264 and the b-value 2.904 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases).

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

None.

Conservation

Population trends and numbers unknown.

Further work

More collections are needed to record information on biology and distribution.

Sources

Nalbant and Bianco (1998).

Genus Triplophysa
Rendahl, 1933

This genus contains loaches of the High Asian region numbering over 110 species. The region centres on the Tibetan plateau with some species found in north China to the east and in Central Asia to the west. The largest nemacheilines are found in this genus. One species is reported from Iran.

The body is elongate with a rounded belly and the caudal peduncle is slightly compressed or highly depressed, and is often very elongate or whip-like. The upper lip is deeply furrowed and there is no processus dentiformis on the upper jaw. The dorsal fin has 6-9 branched rays and is emarginate. There is no adipose crest on the caudal peduncle although a small ridge may be present in some species. The caudal fin is slightly to deeply forked, or is truncate. There is no pelvic axillary lobe. Scales are absent and the lateral line is usually complete. The swimbladder is reduced to two lateral parts and a posterior chamber which is rudimentary or with a secondarily developed bladder lying free in the abdominal cavity. A reduced swimbladder is an advantage in fast water, reducing buoyancy. The gut is short or long, a simple s-shape or a spiral. The colour pattern is usually mottled rather than banded. Males have a raised tuberculate area below the nostrils extending from the lip corner to the anterior eye margin, separated by a groove from a lower tuberculate area. Pectoral fin rays have thickened and tuberculate pads on their dorsal surface.

Prokofiev (2009) summarises osteological characters in his key and these include possession of 3 pectoral fin radials, if 4 are present then the 2 external ones are more or less dilated, flattened and at least partially overlie each other, and the presence of the preethmoid I bone in the skull. Prokofiev (2010) adds other osteological characters.

Triplohysa farwelli
(Hora, 1935)

Described from the "Helmand river, Afghanistan" in the Sistan basin but without any definite locality; it may be remote from the Sistan lowlands of Iran (Hora, 1935). Originally described under Nemacheilus. No Iranian record.

Triplophysa griffithii
(Günther, 1868)

Described erroneously as from Assam but the type locality is the Arghandab River near Kandahar in the Helmand River drainage of the Sistan basin (Hora, 1929; 1935; Bănărescu and Nalbant, 1966). Originally described in Nemachilus (sic). No Iranian record but see T. stoliczkai.

Triplophysa stoliczkai
(Steindachner, 1866)

Courtesy of Stephane Ostrowski.

Common names

sagmahi-ye Pamir.

[Tibetskii golets or Tibetan loach, Pamirskii golets or Pamirs loach, both in Russian; singhat in Pakistan; Stoliczka's loach].

Systematics

Often spelt stoliczkae, (e.g. in Annandale and Hora (1920)), the original spelling is Stoličkai, which becomes stoliczkai in Latin where accents on letters are not used. This species is placed in the genus Triplophysa by Menon (1987) and Bănărescu and Nalbant (1995).

Nemacheilus akhtari Vijayalakshmanan, 1950 described from "Farakhollum, about 10 miles South of Gardan Diwar, Helmund river" (Farakhulm at 34°31'N, 68°08'E), Afghanistan in the Sistan basin may be a synonym of Nemacheilus (= Triplophysa) griffithii (Günther, 1868) according to Bănărescu and Nalbant (1966) or T. stoliczkai according to Prokofiev (2007). There is no Iranian record.

This is a highly variable species originally described from streams around Lake Tsumureri in Rupshu Province of western Tibet and also found in the upper reaches of such rivers as the Hwang, Brahmaputra and Yangtse. It is reported further west from the Aral Sea basin, the upper Indus River basin, the upper Helmand River basin in Afghanistan and probably in Sistan in Iran. Prokofiev (2007) considers the systematics of the loaches in the stoliczkai group to be the most complicated in High Asian species. Berg (1948-1949) gives a series of figures which plainly show the variability in body form and pigmentation of this species. Such a wide distribution and variability may be indicative of several taxa being confused under this name.

Regan (1906) refers specimens from Sistan to Nemacheilus stenurus Herzenstein, 1888. Annandale and Hora (1920) consider stenurus not to be a distinct species but Hora (1922) compares stenurus with Nemacheilus tenuis (see below) and notes that stenurus has a continuous and entire lower lip (slightly indented in ZISP 7355 examined by me - wide interruption and greatly plicated in tenuis) and the dorsal fin origin is closer to the snout than the caudal fin base (equidistant or nearer the caudal base in tenuis). Specimens from Sistan labelled as Nemacheilus stenurus by Regan (1906) are then misidentified T. stoliczkai according to Annandale and Hora (1920). Since N. stenurus was described from the sources of the Yangtse Kiang in China and is a more recent name, it is probably not relevant to Iranian loaches.

Berg (1948-1949) places Nemacheilus tenuis Day, 1877 (publication dated 1876, apparently published 1877 according to Eschmeyer et al. (1996)) in stoliczkai as var. tenuis but notes that it occurs together with the type form in the Gunt River of the Pamirs in Tadjikistan as well as in Sistan, and thereby the implication is that this variety is not a subspecies (since subspecies do not occur together). Berg (1948-1949) also has the subspecies Nemacheilus stenurus uranoscopus Kessler, 1872 described from the Zeravshan River in Uzbekistan as possibly occurring in the Helmand River basin, presumably the upper reaches, but he includes Nemacheilus tenuis Day under this subspecies too.

Annandale and Hora (1920) initially place Sistan specimens in Nemacheilus stoliczkai but later Hora (1922) places them in Nemacheilus tenuis. Hora (1922) restricts the name stoliczkai to fish from the Indus River basin and considers that the various wide-ranging reports of stoliczkai refer to various other species. See also Vijayalakshmanan (1950) for comparisons with some nominal Afghan species in the Helmand basin.

The fish from Sistan examined by Annandale and Hora (1920) and assigned to T. stoliczkai are in two forms - one with a thin caudal peduncle, the stenurus type which they do not recognise as specifically distinct, and one with a thick caudal peduncle which they place in var. leptosoma Herzenstein, 1888 (note that varieties are not recognised by the International Code of Zoological Nomenclature). The former has a minimum caudal peduncle depth in caudal peduncle length of 5.3-7.8, mean 6.8 based on 3 fish from a table in Annandale and Hora (1920) and the one fish seen by me, and the latter has values of 2.9-3.6, mean 3.1 from the table in Annandale and Hora (1920). Annandale and Hora (1920) point out that this character is dimorphic but is neither related to sex nor "race" and that gut loop development also varies but independently of the caudal peduncle character. The specimen I examined from Sistan has a stenurus form of caudal peduncle (value 7.6), the dorsal fin origin is about equidistant between the snout tip and caudal base and so agrees with neither stenurus nor stoliczkai (see table in Vijayalakshmanan, 1950)), the head length in standard length (4.5) falls within the limits of stoliczkai (4.2-4.8) and not stenurus (4.9-5.5) (see table in Vijayalakshmanan, 1950)), and the lip grooves resemble another species, Nemacheilus akhtari Vijayalakshmanan, 1950, described from the Helmand River at Farakhollum about 10 miles south of Gardan Diwar in Afghanistan. The conclusion I reach here is that an individual fish in this group of species or forms can have a mix of characters used by authors to separate and define species.

The resolution of this problem, the correct name for Sistan loaches, other than the distinctive P. rhadinaea and P. vignai, requires extensive material from the whole range of the nominal species involved for dissection and comparison of gut shape, for analysis of sexual dimorphism and individual variability in such morphometric characters as caudal peduncle breadth and thickness and position of the dorsal fin, as well as in determining new and, hopefully, definitive characters. Conservatively, I refer Sistan fish to stoliczkai, the oldest available name, while recognising that tenuis may be a distinct species or subspecies and be the fish found in Sistan, or even that the fish in Sistan are an unnamed taxon or taxa. The Iranian and other material available to me does not permit a resolution of this wide ranging problem.

Syntypes of Cobitis stoliczkai are in the Naturhistorisches Museum Wien under NMW 48436 (5 fish), NMW 48439 (1) and NMW 50477 (4).

A syntype, or at least material examined by Day, of Nemacheilus tenuis is in the Naturhistorisches Museum Wien under NMW 48477 (Eschmeyer et al., 1996).

A syntype of Nemacheilus stenurus, 135.2 mm standard length, is in the Zoological Institute, St. Petersburg, Russia under ZISP 7355. Other types are ZISP 7256 (4), 7354 (3, now 2), 7355 (1), and BM(NH) 1891.10.7.33 (1, 64.9 mm standard length) (formerly in ZISP).

SEE Prokofiev (2007) for tenuis and stoliczkai and other data ?

Key characters

The elongate yet rounded caudal peduncle without an adipose fin distinguishes this species from P. rhadinaea and P. vignai the only other nemacheilids recognised in Sistan. The depth of the caudal peduncle just behind the end of the anal fin is the same as the width.

Morphology

Dorsal fin with 2-3 unbranched and 6-9, usually 7-8, branched rays, anal fin with 2-3 unbranched and 5 branched rays, pectoral fin with 8-12 branched rays, and pelvic fin with 6-8 branched rays. Lateral line complete and distinctive. Scales are absent. Caudal peduncle long and slender or short and deep (see above). Head length in standard length 4.0-4.8. Eye diameter in head length 4.8-5.5, mean 5.2 in Annandale and Hora (1922) for Sistan fish (note that Day (1876; 1878) gives eye diameter as 8 in head length but Steindachner (1866) in his original description gives 5.5). Snout blunt. Barbels are reported as the maxillary ones reaching behind the eye, the rostral ones shorter; the specimen from Sistan had the maxillary and longest rostral barbel about equal in length, the maxillary reaching back to about mid-eye level. Lips are rugose, deeply incised and may be fimbriated. The lower lip is fimbriate and interrupted in the middle. There is a narrow, elongate post-labial groove with well-marked ridges on each side extending back on the mid-line of the lower head (in one specimen seen by me). The pectoral fin tip is formed by the third and fourth branched rays. Dorsal fin origin nearer the snout tip than the caudal base in Iranian specimens or equidistant while elsewhere in the range of this species it is nearer the caudal base (but dorsal origin is individually quite variable in Nemacheilidae). Caudal fin slightly emarginate, lower lobe longer than upper. A well-marked groove between the anus and the anal fin is absent. The gut length is slightly longer than the fish itself with two loops. Chromosome number is 2n=50 for fish from Kyrgyzstan (Klinkhardt et al., 1995).

Sexual dimorphism

Mature males have the first 3-6 pectoral fin rays thickened and tuberculate. A raised tuberculate area below the nostrils extends from the lips to the eye and is separated by a groove from a tuberculate area below (see Fig. 2B in Bănărescu and Nalbant (1995)).

Colour

The flank bears irregular dark spots which may coalesce into a stripe, or is marbled dark green or black. There are dark brown saddles over the back. Fins have rows of pigment, best developed on the dorsal and caudal.

Size

Reaches 16.5 cm.

Distribution

Found from Tibet to northern Kashmir, Turkestan and Sistan. In Iran, it is reported from the Hirmand River delta of Sistan (Annandale, 1921).

Zoogeography

See family account.

Habitat

Unknown.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

None.

Conservation

Biology and numbers are poorly known so an assessment of conservation status cannot be made.

Further work

The biology of this species needs study.

Sources

Type material: Syntypes of Nemacheilus stenurus (ZISP 7355 and BM(NH) 1891.10.7.33), see above.

Comparative material: BM(NH) 1905.4.7:3, 1, 87.7 mm standard length, Sistan, Helmand River, collected by H. McMahon and from the Indian Museum, Calcutta (the size of this fish is at variance with the 6 specimens measured by Annandale and Hora (1920:181) but appears to be from the collections made by the Seistan Arbitration Commission of 1902-1904 and is possibly the seventh specimen referred to in the text. This specimen has been identified as Triplophysa griffithii by S. O. Kullander and has old labels as Nemacheilus stenurus and Nemacheilus tenuis).

Genus Turcinoemacheilus
Banarescu and Nalbant, 1964

This genus contains a single species and is only recently recorded from Iran. The characters of the genus are the same as the species.

Turcinoemacheilus kosswigi
Banarescu and Nalbant, 1964

Common names

None.

Systematics

Originally described from Kapozik Kadun, Hakkari, Turkey. The holotype is ZMH H1884 and 6 paratypes are ZMH H1885.

Key characters

The pelvic fin origin lies in front of the dorsal fin origin and the anus lies closer to the pelvic fins base than the anal fin base, distinguishing this species from all other Iranian nemacheilids.

Morphology

The body is of uniform depth and almost cylindrical. Dorsal fin with 2-4 unbranched rays and 7 branched rays, anal fin with 1-4 unbranched rays and 5 branched rays, pectoral fin with 7-9 branched rays, and pelvic fin with 5-7 branched rays. The lateral line is short with 18-19 pores, ending before the dorsal fin origin level. The lateral line pores are larger than in other species in Iran. The body is scaleless. Eyes are small and far apart. There is no processus dentiformis. All fins are small and more or less rounded. The dorsal fin origin is behind the level of the posterior margin of the pelvic fins. The caudal fin is weakly emarginate. The lips are simple and smooth, the head short and flat and there is no keel on the caudal peduncle. The greater distance between dorsal and ventral fin, longer snout length, lower body depth and narrow body width are viewed as adaptations to fast flowing streamlets (Golzarianpour et al., 2009). The pectoral fins are large and horizontal and act as vacuum suckers (and possibly the pelvic fins too) (Breil and Bohlen, 2001). There is a fairly large fleshy pelvic axillary lobe.

Sexual dimorphism

None.

Colour

Overall colour is brown to yellowish. The back has a brown line. There are a series of irregular flank blotches (ca. 12) extending from the back to the mid- or lower flank but not the belly. These are weakly expressed anteriorly or merge into the general dark pigmentation there. There is a bar or arc of dark pigmentation at the caudal fin base. Fins are not spotted.

Size

Attains 55.3 mm total length.

Distribution

Reported from the Sezar River, a tributary of the Dez River at 33°28'N, 49°03'E by Golzarianpour et al. (2009) and in the northern Gulf basin (Nalbant et al., 2010). Also found in the Tigris River and its tributaries and the upper Euphrates River basin including the Göksu River in Turkey (Breil and Bohlen, 2001; Golzarianpour et al., 2009).

Zoogeography

An endemic species in the Tigris-Euphrates basin, its relationships with other nemacheilids is uncertain.

Habitat

Gravel beds with a strong current are favoured, the slender body allowing this fish to exploit spaces between the gravel. Capture sites in Turkey had clear to muddy water. Aquarium specimens showed thigmotaxis, fixing themselves in place by erecting all fins and pressing against opposing structures. In a strong current, they attached to the substrate with the pectoral fins using suction. Fish even left the aquarium water, a few centimetres above the waterline, attached to the glass with only the tail tip in the water. Splashing with water moved the head and body but the pectoral fins remained attached (Breil and Bohlen, 2001).The Iranian specimens were caught at 1456 m altitude.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

None.

Conservation

Biology and numbers are poorly known so an assessment of conservation status cannot be made. It is widely distributed and probably more common than current knowledge indicates. Golzarianpour et al. (2009) point out that dam construction and human activities  may threaten the species.

Further work

The biology of this species needs study.

Sources

Based on Golzarianpour et al. (2009).

Bagridae

The bagrid catfishes are found in fresh waters of Africa and Asia. Some species attain 2 m in length. There are about 18 genera and about 170 species (Nelson, 2006). Only one species is known from Iran.

This family is characterised by a scaleless body; a depressed head and rounded to compressed elongate body; short dorsal fin with a strong, often serrated spine; a strong, serrated spine in the pectoral fin; a short to long adipose fin; anal fin short to long; caudal fin forked or deeply emarginate; a free margin to the gill membranes over the isthmus; the anterior and posterior nostrils are well separated; anterior nostrils tubular; mouth ventral and transverse or arched; 4 pairs of barbels with the nostril barbel on the posterior nostril, maxillary barbels can be very long, other barbels are mandibular and mental (chin); teeth on the prevomer, premaxillaries and mandible; and eyes often covered by skin. Maximum size is about 2 m.

These catfishes are generally nocturnal. Certain species are important food fishes and others are kept as pets in aquaria.

Genus Mystus
Scopoli, 1777

This is a catchall genus comprising numerous species in Asia. Roberts (1994) restricts the genus to 8 closely related species (see also Grant (2004)). Only 1 species is known from Southwest Asia including Iran. A history and usage of the name Mystus is given by Jayaram and Anuradha (1984).

This genus is characterised by an elongate body, rounded anteriorly and compressed posteriorly, a short and moderately depressed head, head smooth or rugose, an elongate cranial fontanelle extending posteriorly to the base of the occipital process and divided into anterior and posterior portions of nearly equal length by an epiphyseal bar (Roberts, 1994), small to moderate eyes set high and not visible from the ventral surface of the head, a free circular eyelid, a wide transverse, usually subterminal mouth, maxillary barbels very long, jaw teeth are in villiform patches, on the lower jaw as a curved or angular band interrupted at the mid-point, continuous and curved slightly in the upper jaw, total gill rakers 11-30, gill openings very wide and free from the isthmus, adipose fin high and very long, caudal fin deeply forked, upper caudal lobe often much larger than lower, 37-46 vertebrae about equally divided between abdominal and caudal ones, and branchiostegal rays 6-12.

Mystus pelusius
(Solander, 1794)

Common names

ابوزمير (abu-zummair or abu zumir, an Arabic name used in Khuzestan), mahi nish dor (an Abadani name, from www.abadan.net/abadanidictionary.html, downloaded 4 December 2003), sag mahi (= dog fish), گربه ماهي (= gorbeh mahi, meaning catfish), chamu.

[abu-zummair, abouz-zoumeir, abu-al-zamir, abu'l-zoumeir, zugzug in Aleppo, jahudi in Mosul, all in Arabic; Tigris mystus (Fricke et al., 2007)].

Systematics

Synonyms are Bagrus halepensis Valenciennes in Cuvier and Valenciennes, 1840 from the "Couiac, qui est la rivière d'Alep" (= Quwayq at Aleppo, Syria) (see Bailey (1951) on the publication date), Macrones aleppensis Günther, 1864, Macrones colvillii Günther, 1874 from "Bagdad", and Mystus misrai Anuradha, 1986 described from "Lake Antioche, Syria". Hypselobagrus Aleppensis is a new combination by Lortet (1883). Günther (1864) proposed Macrones aleppensis on page 75 but withdrew the name on page 431 when he realised the species was the Silurus pelusius of Solander.

No types of Silurus pelusius described from the "River Kowick" (= Quwayq) are known (Eschmeyer et al., 1996). A syntype of Macrones aleppensis, 127.8 mm standard length, from the "R. Coic, Aleppo, Syria" is in the Natural History Museum, London (BM(NH) 1955.6.25:1). Six syntypes of Macrones colvillii are also in London, ca. 175-234 mm standard length, from "R. Tigris nr. Baghdad" collected by Colville (BM(NH) 1874.4.28:6-8, 1875.1.14:19-21; Roberts (1994) agrees with my observations although Eschmeyer et al. (1996) give BM(NH) 1875.1.14:19-20 for the latter, i.e. 2 fish only).

The holotype of Mystus misrai is in the Muséum d'Histoire Naturelle, Geneva under MHNG 603.95, measuring 123.1 mm standard length, with 1 paratype under MHNG 2231.84 measuring 117.6 mm standard length, 1 paratype in the Zoological Survey of India, Calcutta under FF2315 and 1 specimen missing (Eschmeyer et al., 1996).

Roberts (1994) indicates that more material should be examined to compare fish with short barbels, weakly serrate dorsal spine, short adipose fin and a highland distribution (= pelusius) while the contrasting fish are colvillii.

Key characters

The 4 pairs of barbels, a strong spine in both the dorsal and pectoral fins, elongate and strong adipose fin are distinctive. The head tapers but is not as flattened as in Heteropneustes fossilis, and the mouth is subterminal.

Morphology

The dorsal fin spine is smooth on most of the outer edge and rough on the inner edge. Serrae can be weakly developed or absent, or may be well developed. The dorsal fin spine has 1-4 serrae or notches at the anterior tip and 5-9 along the rear margin, apparently not related to fish size. The pectoral spine is stronger than the dorsal spine and is serrated with 14-23 antrorse teeth on the inner margin, the number increasing with size. The maxillary barbel extends back to the pectoral fin origin or, rarely, as far as beyond the anal fin.

Dorsal fin with 1-2 spines and 7-8, usually 7, branched rays, anal fin with 6-10 branched rays, pectoral fin with 1 spine and 7-9 branched rays and the pelvic fin with 5-6, usually 5, branched rays. Al-Hassan and Hassan (1993) have shown asymmetry in pectoral ray and gill raker counts in samples of this species from the Shatt al Arab, Iraq, possibly due to environmental stress. Total gill rakers 10-18 (Roberts (1994) gives 12(1), 13(7), 14(11), 15(11), 16(8), 17(4) and 18(1), reaching the second raker or further when appressed. Total vertebrae 42-46 (Roberts (1994) gives 42(10), 43(16), 44(3), 45(2) and 46(5). The gut has a large stomach followed by an intestine with about 4 loops.

In specimens examined by me dorsal fin with 2(11) spines and 6(1) or 7(10) branched rays, anal fin with rays difficult to separate into branched and unbranched (perhaps 4-6 unbranched and 6-10 branched rays), pectoral fin with 1(11) spine and 7(3), 8(5), 9(2) branched rays and the pelvic fin with 5(11) branched rays. Total gill rakers 12(1), 13(2), 14(2), 16(2). One specimen with only 7 gill rakers, was possibly abnormal. Total vertebrae 42(1), 43(1), 44(2) and 45(1).

Sexual dimorphism

Unknown.

Colour

Pale brown to olivaceous overall with fins and belly lighter, on a predominant dark silver. Some fish may be silvery-grey overall when fresh. A dark shoulder spot is present. There may be a black spot at the base of the dorsal fin. The dorsal and anal fins have melanophores on the rays and membranes and so are darker than the other fins. The margin of the adipose fin is narrowly black. The caudal fin has a black margin. There may be 3 (sometimes 2), narrow, white stripes on the flank, one along and one each above and below the lateral line. The stripe below the dorsal and adipose fins is narrower than the others. Barbels are whitish, somewhat darker dorsally. Peritoneum silvery to light brown. Jayaram and Sanyal (2003) report an albino specimen from Baghdad.

Size

Reaches 171.2 mm standard length or 22.9 cm total length (Günther, 1874) but possibly to 30 cm total length (Firouz, 2005) or 33.0 cm (Al-Rudainy, 2008).

Distribution

Found in the Orontes, Quwayq, and Tigris-Euphrates basins. In Iran this fish is found in the lower reaches of rivers in the Tigris River basin of Iran such as the Karun, Karkheh and Jarrahi rivers, including too the Zohreh River of the northern Gulf basin (Abdoli, 2000). Also recorded from Fars near Darab in the Rudbal River drainage which flows to the Straits of Hormuz, possibly an accidental introduction, although Esmaeili and Coad (2005) point out that there is no evidence of fish introductions from Khuzestan to Fars. It may simply be very rare outside the Tigris-Euphrates basin in Iran.

Zoogeography

Jayaram and Sanyal (2003) consider that Mystus is derived from an African Bagrus-like ancestor and the genus spread from west to east.

Habitat

Niazi (1976) observed this species in rivers, marshes and brackish waters in Iraq although summer kill resulted from very low water levels and increased salinity.

Age and growth

Al-Hassan et al. (1991) aged this species using eye lenses and vertebrae for a population from the Qarmat Ali River north of Basrah, Iraq. Fish up to 20 cm total length were examined and three age groups were determined, with considerable overlap of lengths for each group. Al-Shami (1998) however found 7 age groups (0+-6+) for the same river using vertebrae to age fish 54-223 mm total length. The highest growth was found in the first year and no significant differences were found between males and females in growth rate. The L was 225.75 mm and the length-weight relationship log W = -4.7516 + 2.8173 log L. The relative condition ranged from 0.94 in December to 1.22 in May. Heydarnejad (2009) gave the length-weight relationship for an Iranian sample as W = 0.0277TL2.999.

Food

Roberts (1994) found eggs in the branchial chamber and stomach apparently identical with those from the ovary. Other stomach items were fish fin pieces and cyprinid fish scales. Aquatic insects, crustaceans, detritus and plant remains are also found in stomach contents of fish examined by me and Al-Rudainy (2008) also mentions fish. Al-Shami (1998) found mean feeding activity and intensity in the Qarmat Ali River, Iraq to be higher in spring and summer, declining in autumn and winter. This fish was carnivorous, taking mainly crustaceans but also insects, fishes, molluscs and aquatic plants. Hussein and Al-Shami (2001) also reported that fish in the Garma Canal, Iraq had a diet dominated by crustaceans (the isopod Sphaeroma annandalei, amphipods, the decapod Elamenopsis kempi, and the prawns Metapenaeus affinis and Atyaephyra desmaresti), followed by aquatic insects (chironomids, corixids and dytiscids), fish (Alburnus sp. and Aphanius dispar), molluscs (the gastropod Lymnaea tenera euphratica) and aquatic plants. Feeding occurred year round with a peak in May and a lowest value in November. Al-Shamma'a (2005) found shrimp and insects to form 47% by volume of the diet of this fish at Al-Fuhoud, Hawr al Hammar, Iraq.

Reproduction

The Qarmat Akli River fish attained maturity in the first year of life with the smallest mature male 92 mm long and the smallest female 72 mm. Eggs were laid in May and June with a fecundity range of 1156-25,833 eggs for fish 105-180 mm total length and 11.88-49.29 g in weight. Relative fecundity was 97.3-524.1 eggs/g (Al-Shami, 1998). Al-Rudainy (2008) gives a relative fecundity of up to 541 eggs/g in Iraq

Parasites and predators

None reported.

Economic importance

This species is of no economic importance. Anglers may catch it on hook and line in Khuzestan but, being scaleless, it is not eaten.

Conservation

This species appears to be relatively common, although not often caught in large numbers, and its conservation status has not been assessed.

Further work

See under Systematics.

Sources

Description also based on Anuradha and Jayaram (1985) and Anuradha (1986).

Type material: See above under Macrones aleppensis (BM(NH) 1955.6.25:1) and Macrones colvillii (BM(NH) 1874.4.28:6-8, 1875.1.14:19-21).

Iranian material:- CMNFI 1979-0087, 1, 162.2 mm standard length, Khuzestan, Karun River at Ahvaz (31º19'N, 48º42'E); CMNFI 1979-0368, 1, 75.6 mm standard length, Khuzestan, Karkheh River (32<º24'30"N, 48º09'E); CMNFI 1991-0153, 1, 123.3 mm standard length, Khuzestan, Zohreh River (no other locality data); CMNFI 1993-0133, 1, 152.6 mm standard length, Khuzestan (no other locality data); BM(NH) 1905.10.14:57, 1, 140.1 mm standard length, Bushehr, Jarrahi River 140 miles northwest of Bushehr (no other locality data); ZMH 2524, 1, 137.1 mm standard length, Kermanshah, Karasu-Gamasiab-Seymarreh (no other locality data); ZMH 4339, 2, 100.5-100.6 mm standard length, Khuzestan, Karun River (no other locality data); uncatalogued material, 1, 57.7 mm standard length, Fars, Cheshmeh Golabi, 15 km west of Darab (28º47'N, 54º22'E)(Esmaeili and Coad, 2005).

Comparative material:- CMNFI 1980-1036, 2, 161.0-167.8 mm standard length, Turkey, Elazig, Keban Dam near Elazig (38º41'N, 39º14'E); CMNFI 1987-0017, 1, 156.8 mm standard length, Iraq, Hawr al Hammar (no other locality data); BM(NH) 1912.5.2:7, 1, 172.5 m standard length, Iraq, Shatt al Arab (no other locality data); BM(NH) 1920.3.5:5-6, 2, 95.4-101.0 mm standard length, Iraq, Basra (30º30'N, 47º47'E); BM(NH) 1936.3.10:3, 1, 56.1 mm standard length, Iraq, Euphrates River at Nasiriyah (31º02'N, 46º16'E); BM(NH) 1974.2.22:1781-2, 1, 52.3 mm standard length, Iraq, Khalis (33º49'N, 44º32'E); BM(NH) 1974.2.22:1783-4, 1, 31.0 mm standard length, Iraq, Khalis (33º49'N, 44º32'E); BM(NH) 1975.5.16:6, 1, 155.2 mm standard length, Turkey, Elazig, Euphrates River, Keban Dam Lake (no other locality data).

Siluridae

The sheatfishes are found in Europe and Asia. There are about 11 genera and about 97 species (Nelson, 2006) with 2 reported from Iran. The phylogenetic relationships within the family are examined by Bornbusch (1995).

This family is characterised by a scaleless and elongate body; a moderately compressed head; a non-protractile mouth; teeth on the jaws and palate; 1-4 pairs of barbels (nasal barbels usually absent; maxillary barbels 1-2 pairs, sometimes vestigial or absent); nostrils separate, anterior ones tubular; 4-21 branchiostegal rays; gill openings very wide; dorsal fin short and spineless (usually fewer than 7 rays and sometimes absent); anal fin very long (41 or more rays) and may be confluent with the caudal fin; adipose fin absent; pectoral fin with a spine, often serrated; and pelvic fins small to absent. The largest species is found in Iran (Silurus glanis).

Genus Silurus
Linnaeus, 1758

These catfishes comprise about 5 species found from Europe to China and India.

This genus is characterised by an elongate body, rounded anteriorly but compressed posteriorly; a depressed head; 2-3 pairs of barbels, the maxillary barbels well-developed and often as long as, or longer than, the head; a large and terminal or superior mouth; teeth in bands on the jaws and roof of the mouth; nostrils well separated; eyes small and not visible from the underside of the head; a very short and spineless dorsal fin; no adipose fin; anal fin very long and united to the rounded caudal fin; pectoral fin with a strong serrated spine; and branchiostegal rays 12-15. The genus has been revised by Kobayakawa (1989).

Krieg et al. (1999) isolated microsatellite loci in both S. glanis and S. triostegus and found that the species diverged less than 20MYA and/or high levels of genomic conservation. Krieg et al. (2000) investigated mitochondrial DNA in S. glanis but found no consistent pattern of geographic structuring in European populations, evidence that gene flow and migration between populations were possible until quite recently. Their study also included S. triostegus and the data was diagnostic for the two species.

A general Farsi name for these fishes is گربه ماهي (= gorbeh mahi, meaning cat fish).

Silurus glanis
Linnaeus, 1758

Aras River Dam, courtesy of Asghar Abdoli (measuring fish)
2.25 m, ca. 90 kg, 5 October 1994

Safid River, 2 km west of Astaneh, 4 June 1978, caught in a large dip-net

Common names

mahi-e sebili, esbele or esbeleh in Gilaki (probably derived from sibil meaning moustache in reference to the barbels), esbele-ye orupaiye, Urupai or Europaiye; ispek (Floor, 2003), گربه ماهي روگاهي or gorbeh mahi rogahee, nake or naque in the Lake Orumiyeh basin, where they are known as "whales".

[naxa, nagka or nakki in Azerbaijan; loko in Armenia; som in Russian; European catfish, sheatfish, wels, wels catfish, Danube catfish].

Systematics

Silurus Glanis was originally described from lakes of Europe, Sweden. A skin is a syntype in the Natural History Museum, London under BM(NH) 1853.11.12:168 (Eschmeyer et al., 1996). An image of this type is available at http://acsi.acnatsci.org/base/image_show_wrapper.html?target=230943. Silurus chantrei Sauvage, 1882 is possibly a synonym (see under Silurus triostegus).

Key characters

This species differs from S. triostegus by having weaker and shorter teeth, the upper and lower jaws meet at an antero-dorsal position (dorsal and superior position in S. triostegus), a less serrate pectoral fin spine posteriorly, and a darker colour. Maxillary barbel length is much longer on average, although there is some overlap. Anterior mandibular barbels are alwayslonger than posterior mandibular barbels while in S. triostegus the posterior mandibular barbels are always longer (Ünlü and Bozkurt, 1996).

Morphology

The lower jaw is longer than the upper. Adults have one pair of maxillary barbels and two pairs of mandibular barbels, for a total of 6 barbels. The maxillary barbel is much longer than the head (equal to head length in Silurus triostegus). The pectoral fin spine is finely serrated or smooth on its inner surface and smooth on its outer surface. Vomerine teeth form a single broad patch, not two as in S. triostegus (Kobayakawa, 1989).

Dorsal fin branched rays 3-5, usually 4, anal fin branched rays 70-108, pectoral rays 12-18 with 1 spine (generally higher on average than in S. triostegus but still overlapping), and pelvic rays 1 unbranched followed by 9-14 branched rays (Coad and Holčík, 2000; Reshetnikov, 2002). Vertebrae 70-76 and total gill rakers 9-17 (counts of 9 and 10 may be lower arch rakers only). Total vertebrae 67-74. In specimens examined by me dorsal fin branched rays 3-4, anal fin branched rays 83-87, pectoral rays 15-16 with 1 spine, and pelvic rays 1 unbranched followed by 11-12 branched rays. Vertebrae 18-19 + 54-56 = 72-74. Total gill rakers 12, reaching the raker below when appressed. The gut has a large stomach and an intestine with 3 about loops. Chromosome number is 2n=60 (Ráb et al., 1994; Klinkhardt et al., 1995).

Sexual dimorphism

Abdurakhmanov (1962) reports that females have longer maxillary barbels, a longer postorbital length and a greater caudal peduncle depth than males in Azerbaijan.

Colour

The body is mottled with brown, green or dark grey, even ventrally, over the base colour. The back is dark, from olive-brown to a blue-black, the sides lighter and the belly greyish-white with bluish speckles. Fins are a dark red-brown to brown-violet. Paired fins have a yellowish streak in the middle. Iris yellowish with black speckles. This species can blend its colour with any bottom on which it lies in wait for prey (Fortunatova, 1961).

Size

Reaches legendary sizes of 5.0 m and 367 kg ( and possibly over 500 kg) but most are much smaller. In the Volga Delta females reach 1.75 m and 31 kg and males 1.95 m and 41 kg (Orlova, 1988); in Dagestan specimens up to 1.93 m and 41.3 kg are recorded although fish weighing 3.2-4.8 kg predominate (Shikhshabekov, 1978).

In the Caspian Sea commercial fishery of Iran, this species ranges in size from 41 to 186 cm and 0.6 to 42 kg (Farid-Pak, no date). Sohrabi (1996a) reports the larger fish in Iranian waters usually weigh 10-40 kg and depicts two fish from Gilan. One caught in 1995 weighed 27 kg, the other caught in 1994 weighed 62 kg. The record fish from Iran, caught by a Mr. Haratonian weighed 120 kg and was 2.2 m long. Eastwick (1864) bought a specimen 4.5 feet long (1.37 m) in the Safid River which had a 7 lb (3.2 kg) fish in its stomach. De Mecquenem (1908) reported that they reached 2 m in the Lake Orumiyeh basin and Anonymous (1977) reports fish from there at 400 lbs (= ca. 182 kg). Asghar Abdoli of the Agricultural and Natural Resources University, Gorgan kindly sent me a photograph of a specimen from the Aras Dam caught 5 October 1994 which was 2.25 m long and weighed about 90 kg.

Distribution

This species is found in Europe, Central Asia and Southwest Asia. In Iran it is found from the Aras River and Dam, along the whole Caspian coast from the Astara to the Atrak rivers including the Anzali Mordab and its Siah Keshim Protected Region, the Safid River, the Manjil Dam on the Safid River, the Amirkelayee Lagoon near Lahijan, Gorgan Bay, and from the southwest Caspian Sea and south-central Caspian Sea; and in the Lake Orumiyeh basin including the Nowruzlu Dam, Shapur-e Avval Reservoir, Mahabad, Gader Chai at Ocksa (Günther, 1899; Nedoshivin and Iljin, 1929; Berg, 1936; Nümann, 1966; Holčík and Oláh, 1992; Nejatsanatee, 1994; Riazi, 1996; Abbasi et al., 1999; Kiabi et al., 1999; Jolodar and Abdoli, 2004; Abdoli and Naderi, 2009). Also reported from the Karakum Canal and Kopetdag Reservoir in Turkmenistan (Aliev et al., 1988; Shakirova and Sukhanova, 1994; Sal'nikov, 1995) and may enter Iranian waters of the Tedzhen (= Hari) River basin.

Abdoli (2000) maps this species from lower Gorgan, Neka, Babol, Heraz, Chalus, Tonekabon, and Safid rivers, the Anzali Talab, along the Caspian coast, and in the middle Aras River, and in the lower Talkheh and Zarrineh rivers of the Lake Orumiyeh basin.

Also said to occur in the Tigris-Euphrates basin in Iraq but needs specimens for confirmation (see Freshwater Fishes of Iraq).

Zoogeography

The isolated populations of this wide-ranging species have not been studied. It may well have an origin in the Ponto-Caspian basin from which it has dispersed.

Habitat

This large species is found in the larger water bodies over soft bottoms and can tolerate brackish water (even for spawning) and moderately low oxygen levels. Warm, deep waters with slow current are preferred. It is active at night. Adults are solitary and found under overhanging banks or submerged trees. This catfish overwinters in aggregations on river beds. Resumed activity in spring depends on the local water temperature regime, probably as early as March in Iran. Riazi (1996) reports that this species is native (resident) to the Siah-Keshim Protected Region of the Anzali Mordab. Knipovich (1921) reports this species from depths of 23.8-25.6 m in the Iranian Caspian Sea. Khodabandeh and Shahriari Moghadam (2007) examined the ultrastructure of ionocyte cells which play an active role in osmotic regulation in this fish.

Movements, such as hunting for food, are stimulated by such environmental factors as temperature, solar radiation, air pressure and turbidity after rain. It is sensitive to extra-aquatic sounds. The head canal system is very sensitive and can track prey over distances up to 55 times the length of the prey and follow signals up to 10 seconds old (Kottelat and Freyhof, 2007).

Age and growth

Life span is 22 years for males and 16 years for females in the Volga Delta. Growth is most intensive in the first years of life. After maturity the annual increase in length is 5-7 cm. Relative weight increase is very high (30%) while length increase is 6-10%. June-July is the period of greatest length increase while weight increase takes place mainly in autumn. Maturity begins at 3-4 years, 57-66 cm and 1.3-2.3 kg but may be as early as 2 years, 51-52 cm and 1.2-2.2 kg or as late as the sixth year (Orlova, 1988). Maximum life span is 80 years.

Abbasi and Valipour (2005) found 9 age groups in the Anzali Lagoon with females making up 68.5% of fish caught. Total length of 95 fish was 19.8-186.5 cm and weight was 47.7-30,000 g. Nezami Balouchi et al. (2007) found age groups 1+ to 7+ in the Amirkelayeh Wetland with an average total length of 50.6 cm (range 32.5-73.0 cm) and an average weight of 944.2 g (range 200-3000 g).

Food 

Food in the Volga Delta includes such fishes as Cyprinus carpio, Abramis brama, Scardinius erythrophthalmus and, prior to the regulation of the Volga, Rutilus rutilus (andéor R. caspicus) and herrings (Clupeidae). Crustaceans now form part of the diet there (Orlova, 1988). At one time this catfish in the Volga delta ate 62-68% of its annual food in one month in spring when the Caspian roach (Rutilus rutilus - this may be R. caspicus) arrived on its spawning run. In the Kura region, commercial fishes such as Cyprinus carpio, Abramis brama, Rutilus rutilus (andéor R. caspicus), Aspius aspius, Sander lucioperca, Chalcalburnus (= Alburnus) chalcoides, Barbus (= Luciobarbus) brachycephalus or Barbus (= Luciobarbus) capito, and Silurus glanis make up 30.27% by frequency and 20.18% by weight, non-commercial species such as Scardinius erythrophthalmus, Tinca tinca, Cobitis spp., Rhodeus amarus, Pungitius platygaster, Atherina boyeri (= caspia) Alburnus alburnus (= hohenackeri), Blicca bjoerkna, and Caspiomyzon wagneri make up 50.68% and 33.79% respectively, marine fishes such as Clupeidae, Mugilidae, and Gobiidae make up 12.90% and 8.60% respectively, and crustaceans 25.64% and 17.09% respectively (Mamedov and Abbasov, 1990). In Azerbaijan, Abdurakhmanov (1962) reports Gobio gobio to comprise 22.4% of the diet, Caspiomyzon wagneri 15.7% and eggs 2.2%, Chalcalburnus (= Alburnus) chalcoides 10.2%, Alburnus alburnus (= hohenackeri) 10.2%, Cobitis taenia 9%, Barbus lacerta 7.8%, Barbus (= Luciobarbus) capito 6.7%, Chondrostoma oxyrhynchum 5.6%, Capoeta capoeta 3.4%, Blicca bjoerkna 2.2%, and loaches 4.5%.

In the Anzali Lagoon of Iran according to Abbasi and Valipour (2005), this species ate 78.6% bony fishes, 15.8% crustaceans, 4.13% insects, 0.9% amphibians and 0.5% bivalves. Carassius auratus dominated at 33.9%, followed by the crustacean Macrobrachium spp., at 14.22%, Neogobius kessleri (= Ponticola gorlap) at 4.59% and Proterorhinus marmoratus (= nasalis) at 2.75%. C. auratus dominated in spring and autumn, Macrobrachium spp. in summer and N. kessleri in winter. Consumption of fish increased with size, being 44.2% at 20-55 cm and 94.5% at 91-125 cm, Cannibalism was not observed and commercial fish stocks were not consumed. Catfish in the Amirkkelayeh Wetland fed principally on Tinca tinca and amphipods each at a frequency of 36.3%, along with minor amounts of Blicca bjoerkna, Pungitius platygaster, Perca fluviatilis, Proterorhinus marmoratus (= nasalis), Carassius auratus, Cobitis taenia, frogs, rats, birds, water bugs and beetles, plecopterans and odonatans (Nezami Balouchi et al., 2007). They were also cannibals.

Generally a wide variety of fishes is taken along with crayfish, frogs and even birds and small aquatic mammals. It is a voracious predator but stories of attacks on dogs and small children are more legendary than factual although human remains may be scavenged (Gudger, 1945b). Active feeding occurs at water temperatures above 8°C so winter feeding is minimal or absent. While feeding often occurs at night, catfish can be heard feeding in the evenings by the snapping of the mouth and tail strikes on the water. In cloudy water conditions they come into shallow water to take earthworms, grasshoppers and frogs washed in from nearby fields (Mihálik, 1982).

Young catfish feed on plankton, particularly Cladocera such as Daphnia, Chydorus, Alona and Bosmina among others. Later the diet involves mosquito larvae, larger crustaceans, organisms associated with the river bank, worms, snails and young fishes. Cannibalism occurs if food is short (Mihálik, 1982).

Reproduction

Behmanesh et al. (2009) found the maximum mean gonadosomatic index for Anzali Lagoon fish was in May and June, decreasing to September. Maximum values for gonad maturity were in April-May at water temperatures of 19.3-22.7°C. Spawning was from April to mid-June.

Non-intermittent spawning in Dagestan takes place in late May and continues to the middle of July when the eggs of females mature at a water temperature of 20-22°C. Spawning rarely occurs below 20°C. Males may actually have running milt 30-40 days earlier than this and also later, a longer potential spawning season. Adhesive yellow eggs are laid in depressions in weed beds, formed by the male pressing on the plants. Fecundity is up to 285,000 eggs with diameters around 2-3 mm (to 467,000 elsewhere). Elsewhere spawning may be intermittent (Shikhshabekov, 1978).

The male guards the incubating eggs, even during the day, moving his tail fin every 3-5 minutes to ensure adequate oxygen supplies. Nests in Europe may be on the fine roots of plants which hang freely in the water. The nest is in shallow, 40-60 cm, water. Males pursue females just under the water surface, an indication spawning will occur the same evening or the next day. Spawning usually occurs in the evening, often before a thunderstorm on warm and stifling days. The male nudges the female in the anal region, swims under her and may lift her so that her back is above water, the male wraps himself around the female for 10-12 seconds, the male and female separate and the female sinks slowly to the bottom and discharges eggs, the male following to release milt. This process can be repeated several times over 1.5-2.0 hours and the water around the nest is milky from sexual products. Eggs hatch after 2.5-3.0 days at 23-25°C. Larvae are light sensitive and die in direct sunlight ands also if water temperature falls below 13-14°C (Mihálik, 1982).

Parasites and predators

Mokhayer (1976b) records the digenetic trematodes Aphanurus stossichi and Bunocotyle cingulata, the nematode larvae Anisakis sp. and the nematode adults Cucullanus sphaerocephala, and the acanthocephalan Corynosoma caspicum. Ataee and Eslami (1999, www.mondialvet99.com, downloaded 31 May 2000) report the helminth Mazocea alaosa from the gastro-intetsinal tract of fish from the Anzali wetland. Masoumian et al. (2005) recorded the protozoan parasite Trichodina perforata from this species in the Aras Dam in West Azarbayjan. Khara et al. (2006a) record the eye fluke Diplostomum spathaceum for this fish in the Amirkalayeh Wetland in Gilan. Sattari et al. (2004; 2005) surveyed this species in the Anzali and Amirkelayeh wetlands, recording Raphidascaris acus, Raphidascaroides sp. and Eustrongyloides excisus. Barzegar et al. (2008) record the digenean eye parasite Diplostomum spathaceum from this fish. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Cyprinacea sp. and Ergasilus sp. on this species.

The Caspian seal, Pusa caspica, is a predator on this species (Krylov, 1984).

Economic importance

There is some opportunity for sport fishing for this species in the Anzali Mordab and Lake Orumiyeh basin where it will take spinners and spoons as well as frog live bait. It reputedly puts up a tremendous fight (Anonymous, 1977).

Nevraev (1929) reports on catches in various regions of Iran in the early years of the twentieth century. There were no evident trends of increase or decrease. In the Astara region from 1901-1902 to 1913-1914 the catch varied irregularly from 699 to 4031 fish, in the Anzali region from 1901-1902 to 1918-1919 the catch varied from 18,177 to 206,485 fish, in the Safid River region from 1899-1900 to 1917-1918 the catch varied from 3290 to 43,835 fish, in the Mazandaran region from 1906-1907 to 1913-1914 the catch varied from 5282 to 11,283 fish, and in the Astrabad region from 1902-1903 to 1912-1913 the catch varied from 3500 to 26,200 fish. The commercial catch in Iran from 1956/1957 to 1961/1962 varied between 4,913 kg and 37,630 kg (Vladykov, 1964) and from 1965/66 to 1968/69 varied from 11 to 31 tonnes (Andersskog, 1970) but in the 6 years from 1980 to 1985 catches were recorded by the Food and Agriculture Organization, Rome as respectively 2, 2, 0, 2, 3, and 0 tonnes. The catch has been as high as 107,593 kg for the Anzali region alone in 1934/1935 (Vladykov, 1964). Holčík and Oláh (1992) report a catch of 2663 kg in the Anzali Mordab in 1990, at 3.6% of the catch the sixth most important fish there, and from 1932-1964 reported catches varied from none to 12.6 tonnes annually. The Iranian fishery peaks in October and in April (Farid-Pak, no date). This species is of great commercial importance in Dagestan (Shikhshabekov, 1978). The roe has been used as a form of caviar and glue has been made from the swimbladder and bones. In the Lake Orumiyeh basin this fish was used as fertiliser since, being scaleless, it could not be eaten for religious reasons (De Mecquenem, 1908). Its export value is recognised and studies have been carried out on this species such as the effect of ascorbic acid and citric acid on the lipid stability and rancidity inhibition of fillets in frozen storage (Pourashouri et al., 2008a. 2008b; Pourashouri et al., 2009).

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use as food, for industrial processes (such as fertilizers, fish meal, pet food, novelty products although not specified for this species), in sport, and in textbooks. The swimbladders were used for isinglass in the Caspian Sea basin (Lönnberg, 1900a).

This species has been implicated in ichthyootoxism, the symptoms of which are summarised under the genus Schizothorax. The presence of venom associated with the pectoral fins (Coad, 1979b) needs definitive examination.

Conservation

Lelek (1987) classifies this species as rare to vulnerable in Europe because of habitat changes and angling pressure. Kiabi et al. (1999) consider this species to be of least concern in the south Caspian Sea basin according to IUCN criteria. Criteria include commercial fishing, sport fishing, medium numbers, habitat destruction, widespread range (75% of water bodies), present in other water bodies in Iran, and present outside the Caspian Sea basin. Vulnerable in Turkey (Fricke et al., 2007).

Further work

Population numbers need to be monitored carefully as it is fished for although it lacks scales. Populations in isolated basins have not been examined in detail for their relationships to see if they are distinct.

Sources

Kobayakawa (1989) revised the genus Silurus and his data are incorporated here. Mihálik (1982) reviewed the biology of this catfish.

Iranian material: CMNFI 1970-0509, 1, 131.2 mm standard length, Gilan, Safid River at Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1979-0685, 2, 63.9-107.7 mm standard length, Gilan, Safid River below Dehcha (ca. 37º22'N, ca. 50º06'E); CMNFI 1979-0788, 2, 293.5-335.6 mm standard length, Mazandaran, Gorgan River at Khvajeh Nafas (37º00'N, 54º07'E); CMNFI 1979-1236, 1, 269.0 mm standard length, Mazandaran, Gorgan River at Khvajeh Nafas (37º00'N, 54º07'E); CMNFI 1980-0123, 1, 225.3 mm standard length, Gilan, Safid River around Dehcha (37º22'N, 50º06'E); CMNFI 1980-0905, 1, 208.0 mm standard length, Mazandaran, Gorgan River at Khvajeh Nafas (37º00'N, 54º07'E); OSU 4278, 2, 227.2-280.2 mm standard length, Azarbayjan-e Bakhtari, Zarineh River (no other locality data).

Silurus triostegus
Heckel, 1843

NMW 92345, syntype, Tigris River near Mosul, ca. 582 mm standard length

Common names

esbele, and jirri, yeri, yery, yari or iry (= eel), esbele-ye beinolnahrein (= Mesopotamian catfish); sag mahi (= dog fish) in Khuzestan; gorbeh mahi (= catfish).

[djirri, jirri, girri or yerri (= eel) in Arabic; Mesopotamian catfish, Tigris catfish, Asian djirri].

Systematics

Silurus chantrei Sauvage, 1882, a species with 4 barbels, was described from the "Fleuve Koura à Tiflis (= Kura River at Tbilisi, Georgia) but was possibly based on material from the Tigris-Euphrates basin (Berg, 1948-49; Haig, 1952) and may well be a synonym of this species. Two syntypes of Silurus chantrei, 160-170 mm total length, are in the Muséum national d'Histoire naturelle, Paris under MNHN A.3932 (Eschmeyer et al., 1996).

Günther (1899), Banister (1980) and Hora and Misra (1943) considered that S. triostegus may not be distinct from S. glanis but Coad and Holčík (2000) detail differences.

The type locality of Silurus triostegus is the "Tigris bei Mossul" according to Heckel (1843) and the description was based on 4 specimens although the catalogue in Vienna lists 1 specimen in spirits and 2 stuffed specimens. The card index in Vienna in 1997 lists only NMW 92345 as a dried syntype. Coad and Holčík (2000) found only a single stuffed type, ca. 582 mm standard length. Eschmeyer et al. (1996) list 1 dried syntype of triostegus in the Senckenberg Museum Frankfurt (SMF 2623, formerly NMW) and this has a standard length of ca. 710 mm. An image of S. triostegus (SMF 2623) is available at http://acsi.acnatsci.org/base/image_show_wrapper.html?target=221708.

Key characters

This species differs from S. glanis by having robust and longer teeth (snaggly, catching on flesh), the upper and lower jaws meet at a dorsal and superior position (antero-dorsal in S. glanis), a distinctly and coarsely serrate pectoral fin spine posteriorly, and a lighter colour. Maxillary barbel length is about equal to head length while in S. glanis it is much longer on average, although there is some overlap. Anterior mandibular barbels (when present) are always shorter than posterior mandibular barbels while in Silurus glanis the anterior mandibular barbels are always longer (Ünlü and Bozkurt, 1996).

Morphology

Bears 4 barbels in some specimens as opposed to the usual 6 in S. glanis but there is evidence that the four-barbelled Silurus (sometimes placed in a distinct genus Parasilurus Bleeker, 1862, e.g. in Berg (1949), now synonymised with Silurus - see Eschmeyer (1990)) have 6 barbels when young and one pair of mandibular barbels is reabsorbed (see Haig, 1952). Specimens with one pair and with two pairs of mandibular barbels have been reported (Kobayakawa, 1989). Ünlü and Bozkurt (1996) record 4 mandibular barbels (2 pairs) in 3 specimens, 3 barbels in 1 specimen and 1 pair in another specimen for Turkish Euphrates fish. In Silurus triostegus, the adults apparently lose one pair of barbels (F. Krupp, in litt., 1992). Coad and Holčík (2000) found all S. glanis in their study had 4 mandibular barbels (2 pairs) while in S. triostegus 11 fish had 4 mandibular barbels and 12 fish lacked the posterior mandibular pair. There was no apparent trend in barbel loss associated with increase in body size.

The pectoral fin spine is strongly serrated on its inner surface and smooth on its outer surface. Vomerine teeth are in two patches, not one as in Silurus glanis (Kobayakawa, 1989), although Ünlü and Bozkurt (1996) record some specimens of S. triostegus with only one patch. The maxillary barbel reaches only to the end of the head, not much longer as in S. glanis. However, Ünlü and Bozkurt (1996) report that maxillary barbels are longer than the head and later that they reach the end of the head; it is suggested here that barbel length varies individually. The lower jaw is longer than the upper jaw. Teeth in both jaws are recurved, the band of teeth is wider than in S. glanis, and the teeth are stronger and longer. The eye is larger than in S. glanis.

Dorsal fin branched rays 3-4 (counts of 3 rays are more common than in S. glanis; 3 in ten fish and 4 in thirteen fish examined by Coad and Holčík (2000)), pectoral fin branched rays 11-14 with 1 spine, pelvic fin branched rays 8-13 after 1 unbranched ray, anal fin rays 77-94, vertebrae 16-17 + 52-53 = 69-70, and total gill rakers 12-17, reaching the one below when appressed (Coad and Holčík, 2000). The stomach is large, and apparently more elonagte than in S. glanis although this may be distortion due to food content. The intestine has about 3 loops.

Sexual dimorphism

Unknown.

Colour

The upper body is mottled pale yellow-brown and black. Overall colour may appear dark or light and yellowish. Generally much lighter than Silurus glanis. The belly and lower head are white with the belly having black spots. Maxillary barbels and margin of the lower jaw very dark brown.

Size

Reaches 1.5 m in the Syrian Euphrates (Gruvel, 1931) and to more than 2.0 m (Krupp, 1992) and more than 50.0 kg.

Distribution

This species is found in the Tigris-Euphrates basin including its Iranian portion in Khuzestan and such rivers as the Arvand, Bahmanshir, and Jarrahi, and the lower Karun, Karkheh and Dez (Marammazi, 1995; Abdoli, 2000).

Zoogeography

This species is presumably a relative of S. glanis but its closest affinities may lie with species to the east. Differentiation of the two species may have occurred around the Middle to late Miocene but this requires further study (Coad and Holčík, 2000).

Habitat

van den Eelaart (1954) reports this species from open and vegetated lakes and marshes and rivers in Iraq. The larger fish are mostly confined to rivers, entering marshes and lakes only on floods. The young have a greater tolerance of high temperatures and low oxygen.

Age and growth

Al-Abood (1989) found age groups 2 to 7 years for fish from a marsh area north of Basrah, Iraq in 6 weight groups from 300 to 3900 g. Al-Hassan and Al-Sayab (1994) examined 600 specimens from the Al-Hammar Marsh north of Basrah, Iraq for age using vertebrae and eye lens diameter and found 6 age groups. Oymak et al. (2001) describe 11 age groups in Atatürk Dam Lake, Turkey with females having higher L (202.85 cm versus 113.98) and lower K (0.046871 versus 0.101972) values than males. Males matured at age 3 and females at age 4.

Food

Fish are an important food including Liza abu and Acanthobrama marmid in the Iraqi marshes and presumably those across the border in Iran (Al-Shamma'a and Jasim, 1993; Ünlü and Bozkurt, 1996; Dawood, 1997; personal observations). Aquatic insects are also taken but fish predominate. The food in Hawr al Hammar, Iraq was predominately fish (Liza abu, Aphanius spp., Aspius vorax, Thryssa spp., Acanthobrama marmid, Silurus triostegus (young less than 16 cm were eaten by adults longer than 35 cm), Barbus (= Mesopotamichthys) sharpeyi, Heteropneustes fossilis and Cyprinus carpio) followed by shrimps (mainly Metapenaeus affinis), frogs (Rana esculenta) and crabs (mainly Sesarma boulengeri) with relative importance indices of 70.8, 16.3, 6.4 and 4.9 respectively (Al-Daham and Al-Seyab, 2000). Liza abu was the most important fish through most of the year (except July and August when absent)(relative importance index 42.0, followed by C. carpio at 11.5), in numerical abundance and total weight. During July the prey was B. sharpeyi and C. carpio and in August prey was restricted to C. carpio, presumably opportunistic feeding. Other fish species were mostly young of the year and of minor importance. The diet at Al-Fuhoud in the Hawr al Hammar was 70.7% fish by volume (Al-Shamma'a, 2005). Aquatic insects are also taken but fish predominated. Dawood (1997) also studied diet in the southern Hammar Marsh and found fish to be the most important prey year round while shrimps (Metapenaeus affinis) and molluscs were important  in certain months (mostly absent April-August). The disappearance of shrimps probably relates to their migratory pattern. Aquatic insects were found mostly in the spring. There is a reverse relationship between the presence of fish and shrimps. Fish and shrimps increased in the diet with increase in size while aquatic insects, molluscs and small crustaceans decreased with size. Fish species eaten were Liza abu, Barbus (= Carasobarbus) luteus, Alburnus sp., Cyprinus carpio, Thryssa hamiltoni, Heteropneustes fossilis, Aphanius sp., Gambusia holbrooki and Silurus triostegus. Frogs, detritus and aquatic plants were also found in the gut contents. Feeding occurred more at night and with another peak in late afternoon. The index of fullness values increased in April-September when water temperatures and metabolic rate rose.

Reproduction

Spawning takes place in March in Iraq (van den Eelaart, 1954; Al-Hassan et al., 1990) to May and June in Turkey (Oymak et al., 2001). The highest condition factors were found in April in Atatürk Dam Lake, Turkey, the mean egg diameter was greatest in May at 1.937 mm and fecundity attained 120,300 eggs (Oymak et al., 2001). Al-Rudainy (2008) gives sexual maturity in Iraq at 3 years, 50 cm total length and 1 kg weight, spawning in May and continuing for a few months, eggs being deposited on vegetation and egg diameters up to 4.2 mm.

Parasites and predators

Mortazaei et al. (2000) report an infection rate of 33.3% (22 of 6 fish) with the parasitic worms Proteocephalus sp. in this species from Khuzestan marshes.

Economic importance

This species forms 8.5% of the total catch in Iraq (Das et al., 1978), the total catch in 1976 being 691 t (Petr, 1987), but it is not a popular food. As a scaleless fish it is not eaten by Shi'a Muslims. It was found to have potential for use as a protein concentrate.

Conservation

This species is not commonly collected in Iran but this may be a consequence of habitats sampled and gear used. It may be under some threat as it is fished for, at least in neighbouring Iraqi waters.

Further work

Population numbers have not been examined for fish in Iran nor are its biology or relationships well known.

Sources

Kobayakawa (1989) revised the genus Silurus and his data are incorporated here as is the data of Ünlü and Bozkurt (1996) for Turkish Euphrates specimens.

Type material: See above, Silurus triostegus (NMW 92345).

Iranian material: CMNFI 1993-0133, 1, 192.9 mm standard length, Khuzestan, probably Karun River at Ahvaz (31º19'N, 48º42'E); SNM-YR 6421, 1, 325.6 mm standard length, Khuzestan, Karun River at Ahvaz (31º19'N, 48º42'E); ZSM 21832, 1, 406.0 mm standard length, Khuzestan, Dez River at Harmaleh (31º57'N, 48º34'E); ZSM 21833, 1, 425.4 mm standard length, Khuzestan, Dez River at Harmaleh (31º57'N, 48º34'E); uncatalogued, 1, 317.7 mm standard length, Khuzestan, probably Karun River at Ahvaz (31º19'N, 48º42'E).

Comparative material: BM(NH) 1874.4.28:3-5 and 1875.1.14:8 (same jar), 4, 236.7-511.5 mm standard length, Iraq, Tigris River near Baghdad (ca. 33º21'N, ca. 44º25'E); BM(NH) 1888.5.17:1, 1, 336.2 mm standard length, Iraq, Fao, presumably a fish market (no other locality data); BM(NH) 1893.6.23:26-28, 3, 254.3-425.7 mm standard length, Iraq, Fao (no other locality data); BM(NH) 1892.9.1:26, 1, 270.8 mm standard length, Iraq, Fao (no other locality data); BM(NH) 1920.3.3:167-176, 13, 123.1-490.9 mm standard length, Iraq, Basrah (30°30'N, 47°47'E).

Sisoridae

The sisorid or sucker catfishes are found in Asia as far east as Borneo. There are about 17 genera with about 112 species (Nelson, 2006). They are mostly small (as small as 2 cm) although some are very large (2 m). Five nominal species are reported from the Tigris-Euphrates basin in Southwest Asia but the diversity there is very limited, compared to India for example. A single specimen of a sisorid catfish has been caught in the Yeşil Irmak of Anatolian Turkey at Taşova (40°46'N, 36°20'E). This Black Sea drainage specimen calls into question the utility of characters used in identifying and defining species in Southwest Asia. Its characters are a mixture of features shared by G. armeniacus and G. silviae. A wide range of specimens of both sexes, various age groups and from various localities is not available to assess variation and resolve the species composition of the Southwest Asian fauna (Coad and Delmastro, 1985)).

The sisorid catfish family is characterised by a rounded to compressed body and a flattened head; short dorsal fin positioned anterior to the level of the pelvic fins; a spine in the dorsal fin (absent in some non-Iranian species) and in the pectoral fin; a well-developed adipose fin; a short anal fin; paired fins horizontal; gill membranes generally united to the isthmus; anterior and posterior nostrils close together; distinct nasal barbel present; 4-6 pairs of barbels; body with small tubercles (unculi); and a distinct thoracic adhesive apparatus in Iranian species but absent in some other species.

These catfishes are found in mountain streams where they use the adhesive apparatus to maintain position in the current. In Iran they are reputed to lie on their backs in the water to take a rest! They are also very resistant and can live for 6 hours wrapped in wet cloth, reviving when placed in an aquarium (R. Mehrani, pers. comm., 2000).

Genus Glyptothorax
Blyth, 1860

This genus is characterised by a flattened head, an adipose fin of moderate length, a short dorsal fin with a strong spine, the spine serrated anteriorly or posteriorly, or smooth, pectoral fin spine serrated posteriorly and in some with plicate skin ventrally, 4 pairs of barbels, maxillary barbels broadly based, an inferior and transverse mouth, villiform teeth on the roof of the mouth in two patches, eyes small and partly obscured by skin, gill openings wide, gill membranes joined to the isthmus, and an adhesive apparatus on the chest formed from plaits or folds of skin, often with a central depression.

The validity of the described Glyptothorax species in the Tigris-Euphrates has not been adequately resolved. Three other nominal species occur in addition to the two reported here from Iran, namely G. armeniacus (Berg, 1918), G. cous (Linnaeus, 1766), and G. steindachneri (Pietschmann, 1913) (see Coad and Delmastro (1985) for a partial discussion of this problem). Resolution of the taxa found in Iran awaits more extensive material for a better understanding of individual and species variation.

Aglyptosternon Bleeker, 1862 (and such mispellings as Aclyptosteron, Enclyptosternum and variants- see Eschmeyer (1990) for details) are synonyms of Glyptothorax (Li, 1986; Eschmeyer, 1990).

Confusion over the family placement of these fishes in various literature sources is reviewed in Banister (1980).

They are known generally as گربه ماهي (= gorbeh mahi, meaning cat fish) or arteshi in Farsi. Arteshi (meaning soldier-like) may be from their tank-like appearance or for their pigmentation which is said to resemble camouflage. Another general name is سگ ماهي (sag mahi, meaning dog fish). General names are not repeated below.

Menon (1954) considered that the members of this genus had spread westwards along the Himalayas as late as the early Pleistocene. A centre of origin in western Yunnan and the southern slopes of the Himalayas is advocated by Li (1986). This author suggests that a Pliocene movement occurred westwards and that, as well, the distribution of Glyptothorax was influenced by Pleistocene glaciations. Their entry route into the Tigris-Euphrates basin is given as along the Amu Darya system.

Glyptothorax armeniacus
(Berg, 1918)

Found in the Euphrates River basin southeast of Erzurum in Turkey but no Iranian record.

Glyptothorax cous
(Linnaeus, 1766)

Reported from the Tigris River basin in Iraq but no Iranian record. Species identity in the Tigris needs confirmation by specimens. See Banister (1980) for a brief history of the confusion surrounding the name of this species in the literature.

Glyptothorax kurdistanicus
(Berg, 1931)

Common names

gorbehmahi-ye Kordestan.

[Kordestan sisorid; Iran cat (Fricke et al., 2007)].

Systematics

This species was originally described in the genus Glyptosternum, an unjustified emendation of Glyptosternon McClelland, 1842 by Berg (1931a). See Eschmeyer (1990) for further details on this genus name.

The holotype of this species is in the Zoological Institute, St. Petersburg (ZISP 20780) and is in poor condition, the pectoral spines being damaged for example. It is an adult male. The collection date in Berg (1931a) is 10 July 1914 (or 27 June old style). The type locality is "in Kurdistan, at the village Germau (or Germav), at the height of 1500 m, during the works of the Turko-Persian delimitation commission. Germau (or Germav, Germaw) is situated in latitude 36°N south-east of Serdesht, on the western slope of the Sur-kei Range, in the basin of the river Bané, tributary to the Little Zab, which is tributary to the Tigris R.". Berg (1949) gives the variants Germab and Sar Dasht for the localities and Bané is probably Baneh. The village Germab could not be located in gazetteers or on maps but Sar Dasht (36°09'N, 45°28'E) and Baneh (35°59'N, 45°53'E) are evident and the locality is between them.

Key characters

Berg (1931a) separates this species from G. armeniacus by the broader than long adhesive apparatus which does not have pinnate lateral branches and these characters also contrast with G. silviae. The caudal peduncle is short (5.9-6.0 in standard length compared to 4.7-5.2 in G. silviae).

Morphology

Dorsal fin with 1 spine followed by 5-7 branched rays, anal fin with 2 unbranched rays followed by 7-9 branched rays (note that these fin ray counts in Berg (1931a) do not agree with his figure). Pectoral fin with 1 spine and 7-9 branched rays. Total gill rakers 7-9, moderately long and reaching beyond the base of the second raker below when appressed. The adipose fin is short, much shorter than the distance between the dorsal and adipose fins. There are oblique osseus striae under the skin of the upper surface of the first pectoral ray. Head and body covered with minute, elongate tubercles oriented longitudinally but without striae. Tubercles are also present on the base of the caudal fin rays, adipose fin, base of the dorsal and pectoral fins, on the pectoral spine along its whole length both dorsally and ventrally, a few on the base of the pelvic fin rays and few to none on the belly particularly anterior to the pelvic fins. Tubercles on the side of the head are more rounded. Berg (1931a) states that the upper jaw tooth patch has well-developed lateral rami, but these do not extend markedly from the main patch. The nasal barbel is short and does not extend back to the eye. The maxillary barbel is shorter than head length and the mandibular and mental barbels are progressively shorter. The gut is an elongate s-shape after a muscular stomach.

Sexual dimorphism

Unknown.

Colour

Overall colour grey to brown with large, obvious, round, black spots and blotches on the sides or with small round black spots about eye size. All fins with broad black central band and variably developed basal bar. Basal bar most evident on the caudal fin. Adipose mostly covered with a large dark spot but dorsal and posterior edges hyaline.

Size

Attains 267.2 mm total length.

Distribution

Found in the Tigris-Euphrates basin including that part in Iran. Abdoli (2000) has mapped it in the upper Karun, middle and lower Dez, middle and upper Karkheh, Kashkan, Simarreh and lower Gav Masiab rivers. Jawad et al. (2009) record it from the Garaf River of the lower Tigris River basin in Iraq

Zoogeography

The relationships of this species, as with other members of the genus, is presumably with the more diverse fauna to the east.

Habitat

Unknown in detail but it is assumed to be in rocky and gravelly rivers which provide hiding places.

Age and growth

Unknown.

Food

Unknown for Iran but Turkish specimens contained fish remains in the stomach.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

This species is not of any direct economic importance.

Conservation

This species is poorly known in Iran and may be rare enough to warrant conservation efforts.

Further work

See under G. silviae.

Sources

Type material: See above, Glyptosternum armeniacum (ZISP 20780).

Iranian material: None.

Comparative material:- BM(NH) 1974.2.22:1789, 346.6 mm standard length, Iraq (no other locality data); BM(NH) 1968.12.13:465-470, 4, 53.7-76.8 mm standard length, Syria, Euphrates at Mayadine (35º01'N, 40º27'E); SMF 23676, 4, 229.0-297.7 mm standard length, Syria, bei al-Hasaka (36º30'N, 40º44'E); SMF 23677, 2, 65.7-122.0 mm standard length, Syria, Wadi Furati (36º26'N, 40º52'E); ZMH 4430, 2, 129.8-133.3 mm standard length, Turkey, Kemaliye Karasu (no other locality data).

Glyptothorax silviae
Coad, 1981

Common names

gorbehmahi-ye jonubi or jonub (= southern catfish).

[Sylvie's sisorid, southern sisorid].

Systematics

The holotype is in the Canadian Museum of Nature, Ottawa under CMNFI 1979-0390A and measures 67.6 mm standard length. It is from "Khuzestan, stream 3 km south of Bagh-e Malek, tributary to Rud-e Zard or Ab-e Ala in the drainage of the Jarrahi River, 31°29'N, 49°54'30"E". Two paratypes from the same locality measure 44.0-51.5 mm standard length and are under CMNFI 1979-0390B, a third paratype measuring 42.3 mm standard length under CMNFI 1979-0389 is from a "stream tributary to Rud-e Zard or Ab-e Ala, 1 km south of Bagh-e Malek, in the drainage of the Jarrahi River, 31°31'N, 49°53'30"E", and a fourth paratype measuring 134.8 mm standard length under CMNFI 1979-0280 is from "Lorestan, river at "Pol-e Chubee" in Kashkan River drainage on Khorramabad to Kermanshah road via Nurabad (either Kaka Reza River at 33°43'N, 48°15'E or its tributary at 33°47'N, 48°12'E)".

Key characters

The head and body dorso-laterally lack striated or elongate tubercles (present in G. kurdistanicus). The thoracic adhesive apparatus is longer than wide (the reverse in G. kurdistanicus). The caudal peduncle is long (4.7-5.2 in standard length, 5.9-6.0 in G. kurdistanicus).

Morphology

Dorsal fin spines are smooth and number 2, branched rays 6; anal fin branched rays 8; pectoral fin branched rays 7-9; pelvic fin branched rays 5; total gill rakers 6-9; retrorse pectoral fin spine teeth 7-16, the number increasing with size of the fish; and total vertebrae 35-38. The adipose fin is long, its length being about equal to the distance between the dorsal fin insertion and the adipose origin (0.9-1.1). The pectoral fin is short and does not extend back to the pelvic fin origin. The caudal peduncle is deep (depth 47-62% of caudal peduncle length). The "sucker" or thoracic adhesive apparatus has pinnate lateral branches and is markedly longer than wide with a wide and long central depression. The head and body are finely papillose, particularly on the ventral surface. Anterior to the adhesive apparatus the ventral head surface is strongly papillose, becoming less developed laterally. The upper lip is much more strongly papillose than the weakly papillose lower lip. The diploid chromosome number is 2n=52 and the karyotype consists of 9 pairs of metacentric, 14 pairs of submetacentric and 3 pairs of subtelocentric chromosomes. The arm number is 98 (Esmaeili et al., 2009).

Sexual dimorphism

Unknown.

Colour

The body is nearly immaculate but in live fish is mottled light lime- green, grey-green, brown or grey. All fins have a central black bar on a salmon-pink, peach or yellow coloured background. The thoracic adhesive apparatus is pink due to an underlying vascular supply. The eye is red.

Preserved fish may have very few scattered brown or black spots dorsally and laterally on an otherwise immaculate body. The overall body colour is brown becoming pale brown or cream on the belly. The base of the caudal fin has a wide black bar separated from a second distal bar by an unpigmented section of the fin rays. The central-most 4 rays of the caudal fin are variably black in the otherwise unpigmented bar. The postero-dorsal and postero-ventral corners of the caudal lobes are not pigmented but the margins of the lobes are black. The central portion of the adipose fin is black with the margins unpigmented in smaller specimens. There is no black pigment on the basal part of this fin. The paired fins and the anal fin are unpigmented distally but become yellowish with fleshy tissue proximally and then brown at their bases. A central band is not well-defined in small specimens. A light patch is found on the back at the dorsal fin origin and at the dorsal fin insertion. The dorsal fin is darkly pigmented and a central black band is apparent although poorly defined. The adult female is generally darker than the smaller male fish in the type series such that the caudal fin bars are not as well defined, The adipose fin is dark brown and the light patches at the dorsal fin are not distinctive. However the bars on the dorsal, anal and paired fins are more obvious.

Size

Reaches 13.5 cm standard length.

Distribution

Known only from rivers draining to the Persian Gulf in southwestern Iran. Abdoli (2000) has also mapped it in the upper Karun and middle to lower Khersan, and middle to lower Dez rivers in the Tigris River basin and in the Mand and Shur rivers of the Gulf basin (the latter tributary to the Dasht-e Palang River).

Zoogeography

This species is known from waters in southern Khuzestan and from a single specimen from the Gulf basin. This latter is probably a relict of the late Pleistocene when the Tigris-Euphrates flowed down a drained Gulf receiving tributaries now isolated by the post-Pleistocene rise in sea level (Coad and Krupp, 1983).

Habitat

Their ability to use their sucker for clinging to objects can be seen in plastic jars where small specimens will adhere to the sides out of the water.

Age and growth

Unknown. Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 10 fish measuring 5.60-10.66 cm fork length. The a-value was 0.0164 and the b-value 2.975 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases).

Food

Unknown.

Reproduction

The largest specimen in the type series is a female bearing eggs and was caught on 6 July. The breeding season is probably the summer months.

Parasites and predators

Unknown.

Economic importance

This species is not of any direct economic importance.

Conservation

Collections with an electroshocker in 1995 showed this species to be common in the Rud-e Zard.

Further work

The distribution of this and related species should be determined by further field work using electroshocking equipment to extract specimens from under rocks. The validity of the nominal Tigris-Euphrates basin species, including this one, needs examination using large series of adult and young which are not yet available in collections. Variation in critical characters is poorly known because of this shortage of specimens. Molecular and chromosomal techniques may provide additional characters.

Sources

Type material: See above, Glyptothorax silviae (CMNFI 1979-0390A, CMNFI 1979-0390B, CMNFI 1979-0280 and CMNFI 1979-0389).

Iranian material: Type material.

Glyptothorax steindachneri
(Pietschmann, 1913)

Reported from the Tigris River basin at Mosul in Iraq. The two syntypes in the Naturhistorisches Museum Wien have not been located (1997 visit) and the brief description is without figures or details of the thoracic adhesive apparatus. Its validity is in question. Not reported from Iran.

Heteropneustidae

The stinging or airsac catfishes comprise a single genus with about three species found naturally from Pakistan through India to Thailand.

The family is characterised by an elongate and compressed body with a flattened head; the mouth is small and transverse with fleshy, papillated lips; villiform teeth present on the jaws and vomer; 4 pairs of barbels present (nasal, maxillary and 2 mandibular); the anterior nostril is tubular and the posterior nostril a slit; gill openings wide and gill membranes free from the isthmus; air sacs are present (see below); swimbladder very small; the dorsal fin is short and spineless; no evident adipose fin; very long anal fin confluent with the caudal or separated from it by a notch; pectoral fin with a strong and venomous spine; skin scaleless; and branchiostegal rays 7.

These fishes can live in stagnant water by breathing air. They are dangerous to man since the pectoral spine harbours a strong venom. Stinging catfishes nonetheless are an important food in the native range.

Genus Heteropneustes
Müller, 1840

The only genus in the family, its characters are given above.

Heteropneustes fossilis
(Bloch, 1794)

Common names

eshlambo or abu shalambo (note variants on this word are used for catfishes and mudskippers); dudeh, doodeh or dood in Khuzestan (= smoke, perhaps because it is blackish); bu shalambo in Khuzestan; گربه ماهي نيش زن (= gorbeh mahi-e nishzan); گربه ماهي هندي (= gorbeh mahi hendi, meaning Indian catfish).

[samaka, abu-al-hukum, abu al-hakim, abu-alhaka, samma, samaka samma, djirri lasseye or jamhoori (latter at Baghdad in reference to the then new republic or jamhooria (F. Kedairy, in litt., 21 December 2005)) in Arabic; singhi in Pakistan; Indian stinging catfish].

Systematics

Silurus fossilis was originally described from India. A syntype of this species is in the Museum für Naturkunde, Universität Humboldt, Berlin under ZMB 3074 (Eschmeyer et al., 1996).

Key characters

The 4 pairs of barbels, short and spineless dorsal fin, absence of an adipose fin, and the long anal fin are distinctive. The head is small and very flattened and tapers both dorsally and ventrally to a terminal mouth.

Morphology

There are two, tubular air sacs extending from the gill cavity almost to the caudal peduncle, enabling this catfish to breathe air. On capture, air from these sacs may escape and cause a peculiar squawking sound. The anatomy and function of these organs was reviewed by Datta Munshi (1993).

Dorsal fin with 6-8 rays but no spine, anal fin rays 60-79, pectoral fin branched rays 7-8 after a strong spine serrated on its inner margin, and pelvic fin branched rays 5-6. Fin rays are difficult to count without dissection or x-rays because of the fleshy and heavily pigmented nature of the fins. Iranian specimens generally fall within the ranges cited above from literature sources as far as can be determined. Spine serrations are more notch-like than toothed. Al-Hassan et al. (1990) have demonstrated that the level of asymmetry in pectoral fin ray and total gill raker counts increases with fish length. Gill rakers are elongate, reaching adjacent raker 5-7 when appressed and number about 25. Barbels are elongate, the snout barbel being the shortest at about head length, the inner mandibular barbel being head length or longer, and the mouth corner and outer mandibular barbel being much longer than the head. The gut is elongate with several posterior coils.

Zakaria (1964) gives details of the pectoral fin spine anatomy. Singhkohli and Goswami (1987) and Kaul and Rishi (1987) describe abnormalities in this species including an upturned tail, a forked tail and forked barbels.

Sexual dimorphism

Unknown.

Colour

Yellow or leaden to dark green, grey-brown, rust-brown or even black, occasionally with two yellowish stripes. The flanks may also be spotted. The barbels are darker than the adjacent body. The eye is yellow. Young specimens are reddish and have a pale belly with numerous melanophores.

Size

Males reach 24.2 cm, females 34.4 cm in India (Datta Munshi and Choudhary, 1996).

Distribution

First recorded from Iraq for 1960 by Khalaf (1961) and Zakaria (1964) when a strange fish was reported to have inflicted a "painful bite" on several victims. The species spread northward and also eastward into Iran from southern Iraq. One collection from Dezful (see below) is dated February 1960 so the spread into Iran must have been very rapid or the original Iraqi introduction some years earlier than documented. It is now known from the Turkish Tigris River near Diyarbakır, presumably dispersing upstream from southern Iraq (Ünlü et al., 2010).

Found from Turkey and Iraq eastward but not continuously through South Asia to Vietnam. It is common in rivers and marshes of Khuzestan including the Dasht-e Azadegan and the Arvand Dez, Karkheh, Karun and Jarrahi rivers (Abdoli, 2000). Berra (2001) omits their Middle Eastern distribution as they are thought to be introduced.

Zoogeography

An Iraqi biologist told me that this species was introduced to Iraq for mosquito control (sic) by local authorities although no one would later admit to it. A more reasonable assumption is that it was introduced to eat the snail Bulinus truncatus, a vector for the human parasite causing schistosomiasis. It was ineffective in this regard (L. A. J. Al-Hassan, in litt., 1995; Jawad, 2003). There has probably been no natural, large scale migration from Pakistan as envisaged by Banister and Clarke (1977) and Banister (1980). Some Sumerian names may refer to this species but this is by no means certain (Sahrhage, 1999).

Habitat

It is common in rivers, marshes, ponds and canals and is found in both fresh and slightly brackish waters. Al-Daham and Bhatti (1977) found this species to suffer a 25% mortality over 72 hours at 10.25‰  sea water. It was most abundant in polluted and stagnant areas in the lower Diyala River where it dominated catches or was the only fish present (Khalaf et al., 1986; Biro et al., 1988). It is common in swamps and can survive temperatures up to 39.8°C (Pethiyagoda, 1991). Specimens survived 3-6 hours in air. It air breathes every 3-5 minutes but the frequency varies with time of day and weather conditions. On hot and calm days it visits the surface more frequently than during a heavy shower. S. Cowton (pers. comm., 23 August 2005) has observed schools of this species gaping at the surface in the artificial lake around Al Faw Palace in Baghdad, presumably in response to high temperatures and low oxygen. Individuals were also seen rapidly swimming straight up to the surface, gulping and diving straight back down again. On especially sultry days it may float or swim near the surface. In the dry season of India it can live in semi-liquid mud or at the bottom of fissures where the mud has cracked (Jayaram, 1980). It makes nest holes in the sides of ponds about 1 foot below the water surface in the form of anastomosing tubes with several exits. Up to 364 fish can be found in one complex of holes (Datta Munshi and Choudhary, 1996). Tekrival and Rao (1999) report its aquarium preferences and habits as 22-25°C, pH 7.5-8.5, alkaline water, predator, not too bright lighting, bottom dwelling with stones, roots and crevices preferred and cave brooding reproduction. Zakaria (1964) noted aquarium specimens producing audible squeaks when excited and preferred the darker side of an aquarium.

Age and growth

Khalaf et al. (1987) gave length-weight relationships for Diyala River, Iraq fish in autumn as W = 9.12 L2.95 (r = 0.98, n = 58) and in spring W = 0.11 L2.15 (r = 0.84, n = 66) and condition factor was 0.0012 in autumn and 0.08 in spring. Islam et al. (1982) give the following length-weight relationships for fish from the Ashar Canal, Basrah, Iraq:- log W = -6.35211 + 3.53226 log TL (r = 0.93543), log W = -5.96765 + 3.42353 log SL (r = 0.93687), and log W = -1.35223 + 2.04705 log GL (r = 0.87876) (TL = total length, SL = standard length, GL = girth length). Tabrez Nasar (1993) studied populations in India and gives length-weight relationships, log W = 1.7661 + 3.035 log L for one population and log W = 1.805 + 2.615log L for another. The coefficient of condition varied from 1.582 to 2.151, mean 1.89. A pond population did not grow as well as a natural one. Life span was up to 4 years in Iraq at Qarmat Ali using ocular lens diameter and vertebral rings (Al-Hassan et al., 1992) and may be 4+ years in India (Datta Munshi and Choudhary, 1996).

Food

Al-Daham et al. (1977) studied the diel feeding of this species in the Ashar Canal, Basrah, Iraq. Two feeding peaks were observed - at 0500 hours and 1700 hours, dawn and dusk, but stomachs examined around the clock had food in them. Aquatic plants and detritus are the bulk of the diet, followed by entomostracans and aquatic insect larvae. Also present are fish parts, molluscs and non-aquatic organisms. Cannibalism is reported in India for young fish (Jayaram, 1980). Khalaf et al. (1987) studied this species in the Diyala River, Iraq and found young fish to take chironomids and worms while larger ones ate fish. However all sizes take aquatic insects in spring. There is some competition with Barbus (= Mesopotamichthys) sharpeyi, a commercial species (Jawad, 2003). In a study of the recovering Hawr al Hammar, diet was 47.2% insects, 22.1% shrimps and 20.8% fish, in the Hawr al Hawizah 33.9% shrimps, 25.8% fish, 20.8% insects and 19.2% snails and in the Al Kaba'ish (= Chabaish) Marsh 51.2% shrimps, 26.4% fish, 12.0% insects and 10.4% snails (Hussain et al., 2006).

Chili macaroni, corned beef casserole, mixed vegetables and salad are dietary items at Camp Liberty, a former palace in Baghdad, where American soldiers feed leftovers from a distinguished visitors dining hall to catfishes, apparently this species (www.estripes.com, downloaded 7 September 2006).

Small Iranian specimens contained insects in their guts including Notonectidae and Diptera larvae. Abdoli (2000) lists diatoms, Chlorophyceaae, fish remains, Corixidae, Hemiptera, cyclopoid Copepoda, termites, Isopoda, Chironomidae, Oligochaeta and Rotifera.

Reproduction

Sexual maturity in India is reached when fish are about 1 year old, at 8.5 cm for males and 12.0 cm for females. Fecundity reaches 12,000 eggs (Haniffa et al., 2008). Al-Rudainy (2008) reported sexual maturity at 2-3 years, 12 cm length and 40 g weight in Iraq. Spawning there took place continuously from June to August with eggs deposited in batches on vegetation. Females take care of the fry.

Eggs are laid in a shallow depression excavated by both the male and female in mud or sand. Eggs are light green. They hatch in about 2 days in Sri Lanka. The parents guard the eggs and young until the young fish are about a month old and able to look after themselves. Singh Kohli and Goswami (1987) describe spawning behaviour in aquaria after hypophysation using pituitary glands of Indian major carps. A pair of males circled each other in a figure 8 pattern until one established dominance. The dominant male chased the female, swimming underneath her or obstructing her path, and touching barbels. The male tried to bite the female in the chases and shivered its whole body while making lateral passes. The male arcs its body into a u-shape, the female touches the male's genital papilla and the pair remain motionless for 2-5 seconds. The female jerks and separates from the male releasing the eggs which settle to the bottom. The pair rest before mating agian. About 40-200 eggs are released after each mating. Mating acts number 20-100 and always occur near the surface of the aquarium. Spawning is more complete and egg fertilisation is better when there is one female and two males, the other male acting as a stimulator, with the spawning male quarrelling with the non-spawning male between mating with the female. Datta Munshi and Choudhary (1996) report similar behaviour. The male nudges the genital region of the female with his head, occasionally shaking it from side to side. Eventually the female is aroused and nudges the male genital region. This female action was necessary for mating to occur. Mating did not happen when spawners were of different sizes. Once the female has her snout below the male genitalia, the male twists his body to place his snout below her genitalia. The fish remain motionless for 2 seconds, then the male vibrates his body and the female convulses and releases eggs. This can happen 30-50 times for each couple at 2-3 minute and then later 5-10 minutes or longer intervals. About 100-150 eggs are extruded, the number decreasing over time. If more than one male is present, mating only occurs after one establishes dominance. Males may eat eggs. Spawners mate in the water column or near the surface.

Parasites and predators

Jalali et al. (2005) summarise the occurrence of Gyrodactylus species in Iran and record G. fossilis in fish from the Karun River.

Economic importance

An important food fish in India and Sri Lanka, where its flesh is reported to have invigorating qualities. Some fish are exported from Sri Lanka for the aquarium trade. V. D. Vladykov reported (in litt., 22 July 1963) that he had seen this species in pet shops in Tehran, on sale at about $2.00 each.

The pectoral spine can cause a serious wound because of the toxin content of the epidermal cells covering the spine. The histology of the pectoral spines was described by Bhimachar (1944) based on Indian material and the toxin was found to have both neurotoxic and haemolytic effects. The toxin is fatal to frogs (within 15-20 minutes of subcutaneous injection of glycerinated venom) and to other fishes.

Zakaria (1964) reported severe swelling involving the whole arm from a hand sting in Iraq. The swelling and pain recede after about a day but the puncture wound can take about two weeks to heal and some pain can be felt when applying pressure to the wound site up to six weeks later. Caras (1964) (probably based on a report in Farsi in Game and Nature, Tehran, ca. 1961) recorded a diminutive black fish found in the Shatt al Arab which reputedly killed 28 people with a venomous bite (sic). Death was said to be swift. This was presumably a garbled report on this species. Verbal and newspaper reports from Tehran (V. D. Vladykov, in litt., 26 August 1961) maintain that this species could cause death to cattle and humans although Vladykov (in litt., 30 September 1963) considered fatal cases "not well proved". I was stung in the thumb by this fish in Iran with no effect (although I did devote considerable time and effort into squeezing and sucking blood from the puncture site!). Freshly caught or netted fish swing the head from side to side and thus are active envenomators (despite knowing this I was still stung). Treatment is symptomatic and some relief can be obtained by immersing the sting site in water as hot as can be withstood and applying a meat tenderiser. These treatments serve to coagulate the protein toxin. The wound should be cleaned to avoid secondary infections such as tetanus (Halstead, 1967-1970; Coad, 1979b).

R. Beck (pers. comm., 2000) reports that this fish is now present in the Syrian Euphrates, its tributaries, and in irrigation canals. Incredible numbers occur near town sewage outlets and in weed beds. It is known to consume eggs of Tor grypus, a preferred food species

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in aquaculture, as food, in textbooks and because of its venomous nature.

Conservation

This is an exotic species and requires no conservation although it is is listed as Vulnerable in its natural habitats (Haniffa et al., 2008a).

Further work

The distribution of this species should be mapped as it is potentially hazardous to humans.

Sources

Iranian material:- CMNFI 1979-0087, 1, 189.0 mm standard length, Khuzestan, Karun River at Ahvaz (31º19'N, 48º42'E), CMNFI 1979-0359, 5, 96.9-114.2 mm standard length, Khuzestan, Karkheh River at Hamidiyeh (31º29'N, 48º26'E); CMNFI 1980-0909, 4, 113.2-165.0 mm standard length, ? Khuzestan (no other locality data); BM(NH) 1980.8.28:4-5, 2, 91.7-98.0 mm standard length, Khuzestan, Dezful (32º23'N, 48º24'E); ZSM 27369, 3, 111.9-139.0 mm standard length, Khuzestan, Karun River near Ahvaz (ca. 31º19'N, ca. 48º42'E).

Comparative material: BM(NH) 1962.7.26:80-83, 4, 127.8-189.0, Iraq, Baghdad (33º21'N, 44º25'E); BM(NH) 1974.2.22:1785, 1, 89.8 mm standard length, Iraq (no other locality data); BM(NH) 1974.2.22:1786-1788, 3, 102.3-165.3 mm standard length, Iraq (no other locality data); ZSM 19455-56, 2, 123.2-127.7 mm standard length, Iraq, Tigris River near Amara (ca. 31º43'N, ca. 47º06'E).

Salmonidae

This family comprises the salmons, trouts, charrs and whitefishes and contains 11 genera and about 66 species found in cooler water in the Northern Hemisphere (Nelson, 2006). The distribution map in Berra (2001) is too extensive in the south of Iran. There are only two species native to Iran but several others have been introduced with varying degrees of success. Whitefishes may be placed in their own family, Coregonidae, but views differ (see Nelson, 1994; 2006).

Members of this family are characterised by numerous, deeply embedded scales on the body but not the head; an adipose fin; a lateral line with relatively numerous and often quite small scales; 7-20 branchiostegal rays; 3 upturned vertebrae at the tail fin; a large swimbladder; usually numerous pyloric caeca (11-210); gill membranes free from the isthmus; a small dorsal fin with few rays (less than 17); a pelvic axillary process; young usually with parr marks (bars along the flank); and a tetraploid karyotype.

Family members are important in fish culture in all the cooler waters world-wide.

Genus Coregonus
Linnaeus, 1758

This genus is characterised by small, almost toothless mouth, 115 or fewer scales in the lateral line, gill rakers often long and slender, there is a single flap between the nostrils, and no bright and complex colour patterns in adults and no parr marks in young, forked caudal fin. Numerous named species in North America and Eurasia (FishBase gives 217 in Coregonus alone, August 2007), variously recognised as full species or not.

Coregonus lavaretus
(Linnaeus, 1758)

Common names

safid mahi (= white fish at Karaj Dam), safid mahi juibarye (= brook or rivulet white fish), koregon or coregon.

[European whitefish, powan, houting, skelly, gwyniad, lavaret, pollan].

Systematics

Salmo lavaretus was originally described from Lake Bourget, France (Kottelat, 1997). There are numerous subspecies and infrasubspecific names for populations of this fish in northern Europe (Berg, 1948-1949). The origin of the Iranian specimens is unknown so it is not clear which subspecies or species they belong to and accordingly information on biology is of a general nature.

Key characters

This species is the only one in its genus reported from Iran and is recognisable by the generic characters above.

Morphology

The mouth is small, inferior and lacks evident teeth. Lateral line scales are 84-105. The dorsal fin has 3-5 unbranched and 8-13 branched rays, the anal fin has 3-5 unbranched and 11-15 branched rays and the caudal fin has a strong fork. Gill rakers are sparse and short or long and dense, denticulated or not, and number 21-56, with means and form varying between subspecies. Vertebrae number 56-64. The chromosome number is 2n=80 (Klinkhardt et al., 1995).

Sexual dimorphism

Males may bear rows of elongate tubercles on the scales.

Colour

The back and upper head is a dark bluish-grey, blue-green or dark green, flanks are greenish-grey to silvery and the belly is silvery to yellowish-white. Fine speckles may be apparent on the body. The snout is black. The iris is white. The dorsal, anal, adipose and caudal fins are dark while the pectoral and pelvic fins are only dark at the tips.

Size

Attains 97.0 cm and 10.0 kg.

Distribution

The native distribution of this species is in northern Europe and Siberia, in particular the drainages of the Baltic, North, Barents and White seas with isolated populations in central Europe in mountainous areas.

Fingerlings were released into the Karaj and Latian reservoirs near Tehran from 1965-1968 after hatching from eggs imported from Europe (Armantrout, 1980). Walczak (1972) reports some existing still in the Karaj Reservoir but none were found in the Latian. It was also introduced to the Manjil Dam on the Safid River (Griffiths et al., 1972) but this reservoir is drained to remove excess silt and no fishery exists (J. Holčík, pers. comm., 1992). There is no evidence of reproduction. Saadati (1977) states that they were established in both reservoirs. Also reported from the Farahnaz Reservoir, Markazi. Abdoli (2000) depicts the Karaj River and the Abhar River as harbouring this species.

Zoogeography

An exotic species not naturally occurring in Iran.

Habitat

This species is typically an inhabitant of large and deep lakes where its oxygen requirments are quite high. However it is tolerant of warm water and even a measure of pollution. Spawning may occur in rivers tributary to the lake. Anadromous forms occur but are rare in full seawater.

In lakes during the day, it is found at depths of 20-30 m or on the bottom if water is shallower. At night it may rise as far as the surface following the migrating plankton or to the waters edge in the shallows. Schools form in sub-littoral areas during the spawning season and strandings may occur.

Age and growth

Maximum age attained is 20+ years although in some populations most fish are 1+ to 3+ years old while in others most fish are 7+ to 10+ years old. Under good conditions young can attain 10-12 cm after one year and 15-20 cm after two years of life. Maturity is attained at 3-5 years and 25-35 cm.

Food

Plankton is the principal food item, with benthic crustaceans being taken in brackish water and crustaceans, molluscs and insect larvae in rivers. This species cannibalises its own eggs and eats the eggs of other fishes. Feeding may vary through the year, planktonic crustaceans being taken in summer and benthic invertebrates in winter.

Reproduction

There is a spawning migration which may occur in summer but more usually peaks in autumn. Spawning takes place in summer, autumn or winter, varying with the subspecies or form. Summer spawning takes place in deeper water than winter spawning. In lakes, the adult males enter the spawning grounds at dusk, these grounds being gravelly shoals off headlands or on offshore reefs. Some populations spawn over sand or even mud. Females move in each night as they ripen. The yellow eggs are 2-3 mm in diameter and slightly adhesive. They stick to gravel and are protected from predators by falling in crevices between the gravel. Up to 82,250 eggs may be laid. Eggs incubate for 90-100 days during winter at an optimum temperature of 6°C or less.

Parasites and predators

None reported from Iran.

Economic importance

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in aquaculture and as food in Europe and Russia. It has been widely introduced to reservoirs.

Conservation

No conservation of this exotic species is advisable.

Further work

The survival of this species in Iran should be verified.

Sources

Based on general European literature.

Iranian material: None.

Comparative material: BM(NH) 1986.11.14:1-3, 3, ca. 312-330 mm standard length, England, Cumbria, Lake Haweswater (no other locality data).

Genus Oncorhynchus
Suckley, 1861

Members of this genus are found naturally in the Northern Pacific Ocean and its drainages, migrating between fresh and salt water. Some live permanently in fresh water. There are about 12 species and two have been successfully introduced to Iran.

This genus is characterised by a large mouth, in adults extending back to the level of the posterior eye margin or further, scales are small, ca. 100 or more in the lateral line, lateral line scales are as large or larger than scales in adjacent rows and overlap with scales in front and behind, evident teeth in adults, teeth are present on the head and shaft of the vomer bone in the roof of the mouth, usually dark parr marks in young but adults without snow-white leading edge to lower fins, and usually black spots on the body or caudal fin or both.

These salmons spawn once and then die. They are important commercially as food and as sport fishes.

Oncorhynchus gorbuscha
(Walbaum, 1792)

Introduced to the former Soviet Caspian Sea basin in 1963 but not observed subsequently (Baltz, 1991). No Iranian record.

Oncorhynchus keta
(Walbaum, 1792)

Common names

azad mahi keta (= keta free fish or keta salmon, azad mahi being the Farsi for trout or salmon), mahi-ye azad-e chum.

[chum salmon, dog salmon, keta, summer chum].

Systematics

Salmo keta was originally described from rivers of Kamchatka, Russia.

Key characters

This species is distinguished by having 124-153 lateral line scales, 13-17 principal anal fin rays, flanks and caudal fin without distinct black spots, short, stout gill rakers 18-26 and pyloric caeca 135-249.

Morphology

Dorsal fin with 10-14 principal rays, pectoral rays 14-16 and pelvic rays 10-11. Scales are horizontal ovals with a central focus and no radii. Circuli are fine but are lost or only partly developed on the flesh-covered exposed part of the scale. The gill rakers reach just beyond the first to just beyond the second raker when appressed. The pelvic axillary scale is very elongate. The gut is s-shaped. The chromosome number is 2n=74 (Klinkhardt et al., 1995).

Sexual dimorphism

Breeding males develop hooked jaws and large teeth and a slight hump in front of the dorsal fin.

Colour

Marine fish are steel-blue on the back and upper flank, silvery on the flank and silvery-white on the belly. The upper flank and back may have fine black speckles but no spots. The pectoral, pelvic, anal and caudal fins have dark edges. Spawning males in fresh water are dark olive to black above, greyish-red to brick-red on the flank with greenish to purplish bars or blotches and a dark grey belly. Calico is the characteristic colour of dominant males. The anal and pelvic fins are often tipped with white. Females are less strongly marked. Young chum are iridescent, mottled green on the back and silvery iridescent green on the flanks and belly. There are 6-14 parr marks which do not descend much below the lateral line and are narrower than the space between them. Fins are clear to white.

Size

Reaches 120.0 cm and 20.8 kg.

Distribution

Found from Alaska to California on the Pacific coast of North America and also in eastern Arctic Siberia, the Beaufort Sea and south to the Sea of Japan, the widest distribution of any Pacific salmon.

This species was introduced to the former Soviet Caspian Sea in 1962-1970 in an attempt to offset losses of Salmo caspius which was cut off from its spawning grounds by dams. It was considered as ideal because it spawns in the lower reaches of rivers and dams would not affect its spawning migration, it returns to spawn after 2 or 3 years and so some year-classes could escape detrimental conditions in any one year, and because it produces downstream migrants at a smaller size and age than the native salmonid (Magomedov, 1970; 1978; Baltz, 1991). 7.5 million fertilised eggs were transported from the Amur River and Sakhalin Island in the former Soviet Far East and incubated at the Samur Fish Farm, Dagestan. 5,850,000 fry were released.

Walczak (1972) reported capture of a specimen in Iranian waters in 1971, presumably a stray from the former Soviet stocking programmes. Holčík and Razavi (1992) record only two specimens from Iran, one in 1964 and one in 1972, taken between Bandar Anzali and the Astara River on the Iranian coast of the Caspian Sea. They also seem to indicate another specimen taken after 1972. Jolodar and Abdoli (2004) report it from Astara and the Anzali regions, indicating no large scale spread.

Zoogeography

A species introduced to Iranian waters by man.

Habitat

In North America, chum salmon enter streams on the spawning migration and often travel less than 150 km, stopping at the first barrier as they are not strong jumpers. Some fish even spawn in tidal areas. However some rivers, like the Yukon, have a run which travels about 3200 km and takes from early June to the end of September. A migratory speed of up to 115 km/day has been recorded with bursts of speed to 4.6 m/s. Most runs are in the fall but some are in the summer. Very rarely a chum will become trapped in a lake if an outlet stream dries up but normally adult chum are only found in fresh waters on the spawning run.

Age and growth

Reproduction occurs at ages 1+ to 2+, with spawning fish at 2+ predominating in Caspian populations from former Soviet waters. In their native streams in the Far East fish are in their fourth and fifth years of life. 1500 adults were caught returning to one river to spawn in the Caspian in 1966, considered impressive by former Soviet biologists. Accelerated growth and maturity in transplanted fish is well documented (Magomedov, 1970). There is a 15C° difference between the Caspian and native water temperatures for spawning and egg development. Life span in Canada is about 7 years, perhaps 9 years, with fish in British Columbia maturing at 1-6 years with age 3 fish dominant. They spend 2-7 years at sea before returning to spawn. Males grow faster and larger than females.

Food

Food in the sea includes crustaceans, worms, molluscs, squids, jellyfish and fishes. Young fish in the sea take zooplankton. Adults on the spawning run do not feed. Young fish in fresh water eat aquatic insects, such as chironomids, mayflies and caddisflies, as well as crustaceans, worms and terrestrial insects.

Reproduction

Transplantations into the former Soviet Caspian Sea basin were not very successful because of a lack of suitable spawning streams (McNeil, 1979). Spawning runs into some former Soviet streams were recorded from the first half of September to the end of October, the same periods as in the native habitat (Magomedov, 1970). Males arrived first, the mass run was composed more of females and there were more males at the end of the run. Since a single male will spawn with several females, early male arrival on the spawning grounds may promote successful fertilisation. The absolute fecundity of a 3+ salmon was 2739 eggs in the Caspian.

Females excavate a redd by lying on their sides and lashing the tail. The redd is a trough up to about half a metre deep and up to 3.2 m long and 2.1 m wide bordered by a ridge of gravel at the downstream end. In some cases no redd is excavated and eggs are shed over and between boulders. Females may excavate more than one redd and males may spawn with more than one female. A female and one dominant male lie in the redd, gape their mouths, vibrate and release eggs and sperm. The dominant male may have several accessory males accompanying him. The female dislodges gravel at the upstream end of the redd to cover the eggs. The orange eggs are up to 7.8 mm in diameter (perhaps 9.5 mm when fertilised) and each female may shed up to 7779 in the Pacific basin. The adult fish die after spawning and may live only a week after first entering fresh water. The fry emerge from the gravel in March-May in North America, some remaining for several weeks in fresh water or immediately migrating to the sea.

Parasites and predators

Various insects, fish, birds and mammals prey on both young and adult chum.

Economic importance

It is too rare to have any economic importance in Iran. Total catches in the North Pacific Ocean have been as high as 69.2 million fish annually. In Japan it is very important in an ocean ranch industry.

Conservation

The presence of this species in Iranian waters is the result of former Soviet attempts to acclimatise it to the Caspian Sea as a potential replacement for declining stocks of Salmo caspius. The stocking programme lasted from 1962 to 1979 and rivers along the Dagestan coast showed mass spawning but this became rare. Pollution by wood from forestry operations in Dagestan, absence of suitable gravel beds, low salinity in the Caspian Sea and heavy surf on the Dagestan coast may be responsible for the heavy mortality (Magomedov, 1978; Holčík and Razavi, 1992). As an exotic species, there is no need for conservation of chum salmon.

Further work

Reports of this species in Iranian waters should be documented.

Sources

Bakkala (1970) summarised the biology of this species in North American waters and Salo in Groot and Margolis (1991) over its whole range.

Iranian material: None.

Oncorhynchus kisutch
(Walbaum, 1792)

Introduced to the Caspian Sea basin but not subsequently observed and no Iranian records (Holčík and Razavi, 1992).

Oncorhynchus mykiss
(Walbaum, 1792)

Lorestan fish farm specimen, north of Aleshtar (golden form), 3 December 2000

Common names

قزل آل (= gazalala, ghezel ala, qezel ala or kizil ala; probably in confusion with native Salmo caspius, these words meaning literally red spots or red spot fish but used for trout and salmon species in Iran),
قزل آلاي رنگين كمان (qezel ala-ye ranginkhaman, meaning rainbow trout).

[mykiss in Russian; rainbow trout, steelhead trout].

Systematics

Salmo mykiss was originally described from Kamchatka, Russia.

Formerly known as Salmo gairdneri Richardson, 1836 (see Bailey and Robins, 1988; Smith and Stearley, 1989). Placed in the genus Parasalmo Vladykov, 1972 by Reshetnikov et al. (1997) and Mednikov et al. (1999) but Osinov (1999) has reservations since allozyme data show specific level differences with Oncorhynchus although further studies were deemed necessary.

Key characters

This species is distinguished by having 100-161 lateral line scales, 8-12 principal anal fin rays, the vomer bone in the roof of the mouth has teeth on its head and shaft, no red spots on the body but only small dark ones and radiating rows of black spots on the dorsal and caudal fins, and no teeth at the tongue base.

Morphology

Dorsal fin principal rays 10-12, pectoral rays 11-17 and pelvic rays 9-10. Gill rakers 16-22. Pyloric caeca 27-80. The chromosome number is 2n=58-62 (Klinkhardt et al., 1995).

Sexual dimorphism

Breeding males have an elongated snout, the lower jaw is hooked and the roof of the mouth is white.

Colour

Overall colour is very variable. Stream fish are darker and more colourful (rainbows) than lighter, silvery lake or sea fish (steelhead). Some sea-run and lake fish have small orange to red marks below the lower jaw. The back and upper flank are steel-blue, greenish, silvery-olive or even brown, flanks and belly are silvery, grey, white or yellow-green. The side of the head and the flank are characteristically pink. The flank has a broad pink to red or lilac stripe with small black spots. The adipose fin has a black margin and a few spots. Pectoral, pelvic and anal fins may have a few spots and are dusky without any strong markings. Pectoral and pelvic fins are often orange-red. Spawning fish are very dark and the flank stripe is dark red or purple. The young have 5-13 dark, oval parr marks centred on the lateral line with the spaces between the marks wider than the marks. There are 5-10 parr marks on the back in front of the dorsal fin. The upper flank has some dark spots. The dorsal fin is tipped white or orange and has a dark leading edge, sometimes broken up into spots. The adipose fin has a black margin. The anal fin has a white or orange tip. Some adults in streams do not lose their parr marks. A golden form occurs rarely on fish farms in Lorestan and elsewhere is farmed specifically.

Size

Reaches 122.0 cm, possibly 150 cm, and 35.0 kg as sea-run or lake fish but smaller in streams.

Distribution

Found in western Canada and from Japan and Alaska to Mexico. Widely introduced outside this natural range world-wide in suitable waters. They have been introduced to the former Soviet part of the Caspian Sea in 1973 and several hundred returning adults were reported in 1975 (McNeil, 1979; Baltz, 1991).

Rainbow trout were introduced to Iran about 1966 for hatchery production as a commercial product (MacCrimmon, 1971; 1972). They are now widespread in Iran, stocks being maintained by hatchery introductions and sometimes natural reproduction, wherever temperature regimes are suitable in the higher reaches of the Alborz and Zagros mountains (Walczak, 1972; Anonymous, 1977; B. Sandford, in litt., 1979; Y. Keivany, in litt., 1992; Ghorbani Chafi, 2000; personal visits to fish farms).

B. Sandford considered few populations became established because of competition with large populations of omnivorous cyprinids which also take fish eggs, presumably of trout too. Sandford cited viable populations in the Madar Su, a small stream in the former Mohammad Reza Shah Park (see below), Tar Lake in the Alborz (also recorded from Tar Lake by Riahi (1996)), Gahar Lake in the Zagros and the Qara Chai west of Hamadan. Introductions to Neuer or Neur Lake near Ardabil, Ghorighol Lake, Rebeshahr and the Ab-e Bazuft were all failures (although Walczak (1972), Saadati (1977) and R. Mehrani (pers. comm., 2000) report several successful stockings including the Ab-e Bazuft, Gahar Lake, Namrud, Dez River and Jajehrud).

Also reported from the Gorgan and Haraz rivers and Gorgan Bay (the latter escapees from cages)(Kiabi et al., 1999), Madar Su, Golestan National Park (Kiabi et al., 1994), from the Farahnaz Reservoir, Markazi, Namrud in Semnan, Golestan River, Karaj River, Amir Kabir Dam (www.iran-doe.org/Special/Alborz.htm), Shah Abbas Dam west of Esfahan, Darius-e Kabir Dam north of Shiraz, Karaj River, Zayandeh near Esfahan, Rebeshahr southeast of Yasuj (Anonymous, 1977; Y. Keivany, in litt., 1992). An attempt to culture this species in cages in the Avan River, 7 km from Alamut, in 1994 failed and fish escaped (Nialmir, 2001). Also in the Sardab Rud (Jalali et al., 2005).

An attempt to establish a run of steelhead, the migratory form, in the Chalus River from the Caspian Sea was apparently unsuccessful (Armantrout, 1980).

This species was stocked in the 1970s by the Iranian Department of the Environment in the Doogh or Madar Su, head stream of the Gorgan River in Golestan National Park and "Neur" Lake in East Azarbayjan near Ardabil in the Caspian Sea basin; in Ghorigol Lake (east of Tabriz) and the Liqvan Chay in the Lake Orumiyeh basin; the Karaj River, the Jajrud, the Namrud, Tar Lake northeast of Tehran, Lake Lasem near Tehran, the Lar River, probably in the Chashmeh Do Barare, Ab Kharsang, Varangarud and Baragon near Tehran, all in the Namak Lake basin; the Karun River, Mohammad Reza Shah Dam, Rebeshahr River southeast of Yasuj, the Ab-e Bazuft in 1975 (established), Gahar Lake in the headwaters of the Dez River and its outlet stream (established, caught by A. Abdoli in 1995), and an isolated section of the Dez River formerly fishless (established), all in the Tigris River basin; the Shah Abbas Dam west of Esfahan, in the Zianrud (presumably the Zayandeh River of the Esfahan basin) where a reproducing population was established; in the Dorudzan (Dariush-e Kabir) Dam near Shiraz (104,000 in Esfand 1350)(Surber, 1969; Anonymous, 1977; Armantrout, 1980; Petr, 1987; Abdoli, 1993b).

Qanats in Markazi Province were stocked with 930,000 trout (presumably this species) in 2006 (www.iranfisheries.net, downloaded 28 July 2006).The qanats were in Shazand (2 qanats), Arak (4), Delijan (1) and Khomein (1).

Abdoli (2000) lists the Dasht-e Kavir and Kerman-Na'in basins generally, the upper Kashaf River in the Tedzhen River basin, the Gorgan, lower Neka, middle and lower Babol, and Heraz rivers in the Caspian Sea basin; the upper Talkheh, Tatavi and Zarrineh rivers in the Lake Orumiyeh basin, the Abhar stretch of the Shur and the upper and middle Karaj rivers of the Namak Lake basin; and the Khersan, upper Dez and Kashkan rivers of the Tigris River basin, and the upper and middle Zayandeh River of the Esfahan basin.

Zoogeography

An exotic species introduced as a food fish.

Habitat

Rainbows are found in rivers or streams where there are pools and riffles. Some live in lakes and are called Kamloops trout in Canada while others run to sea and are called steelheads. They can tolerate temperatures up to 24°C, warm for a trout, but prefer temperatures below 20°C. Sea-run fish spend about 1-4 years usually in inshore waters at middle to surface depths after 1-4 years in fresh water. Some fish (half-pounders) return to streams after a few months at sea. Summer steelhead have spent one winter at sea and return in summer (April-October) to spawn next spring while winter steelhead are larger and return from December to April peaking in January to spawn in March and April. Aquaculture of trout in Iran is affected by drought conditions reducing water flow to farms. Production was higher in 1999 compared to the previous year but this was because 82 new farms were opened; average production per farm fell (Foghi, 2004).

Age and growth

Life span varies with habitat, up to 11 years in some lake fish but only 3-4 years in many streams and small lakes. Growth varies with habitat including such factors as length of sea, stream or lake life, years before spawning, available food supplies, latitude, altitude, temperature regime, competition with other salmonids, and so on. Ageing these fish may be difficult because of the complicated life history pattern of stream and lake residency. Maturity is also variable with habitat. Some males mature at 9 months in fish introduced to warm southern waters and some females only at 8 years, but generally maturity is reached at 3-5 years in Canada, for example, with males maturing a year earlier than females. Neur Lake fish had a growth rate of 17 lbs (7.72 kg) in 3 years, more than doubling in size, because of the excellent food supply of shrimp. In Tar Lake, where food resources were poor, mostly surface insects, fish grew to only 3-5 lbs (1.4-2.3 kg) in 2 years (Anonymous, 1977). In the Karaj Reservoir, one-year-old fish with a length of 8-10 cm grew to 25 cm in one year although the available food was only plankton.

Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 111 Iranian fish measuring 27.5-57.0 cm standard length. The a-value was 0.0161 and the b-value 3.044 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases).

Food

Food includes plankton, crustaceans, aquatic and terrestrial insects, snails, leeches, salmon eggs, and other fishes. The fish eaten enhance growth and the species taken depends on what is available. In Lorestan, this species thrives in rivers where Gammarus is numerous, e.g. near Dow Ab-e Aleshtar. Fish and squid are taken at sea along with some crustaceans. Abdoli (2000) lists Trichoptera, Plecoptera, Chironomidae, Ephemeroptera, Ceratopogonidae, Odonata, Simuliidae, Coleoptera, Decapoda and Amphipoda as food in Iran.

Reproduction

Spawning takes place from March to August but is usually in spring. North American Great Lakes fish may spawn from late December to late April. Water temperature for spawning usually exceeds 10°C but may be 5-13°C. A female excavates a redd by lying on her side and thrashing her tail. Redd excavation occurs during the day and night and dimensions are usually longer and deeper than the female's body. A male courts a female by rubbing his snout and body against her, by vibrating, by swimming over her in the redd and by pressing against her. Several males are found around each female but one male is dominant. The spawning act lasts 5-8 seconds with the pair parallel in the redd pressed together, both fish gape, arch and vibrate. Other males may shed sperm. The female covers the eggs with gravel by dislodging it from the upstream end of the redd. Most spawning takes place in the morning and evening and nests may be abandoned during the day. Females construct several redds and may spawn with several males. Eggs are orange or pink, 5.0 mm in diameter and up to 12,749 in number. However, egg numbers are usually a few hundreds to thousands per female. The eggs hatch in about 8 weeks and fry generally emerge in June to August from spring spawnings. Repeat spawning can occur for up to 5 years.

Nematollahi and Azari Takami (2002) studied the quality and quantity of seminal fluid in cultured male broodstock specimens of this species at the Karaj Fish Farm. Differences were found between stocks in sperm motility and concentration.

Parasites and predators

A wide variety of other fishes and birds feed on this species and there have been extensive studies on parasites and diseases of this commercially important species. Akhlagi (1999, www.mondialvet99.com, downloaded 31 May 2000) reports infectious pancreatic and haemotopoitic necrosis in fish from Bovir Ahmadi va Kohkiluyeh Province but not in Fars Province. Liver and intestinal submucosa degeneration and kidney necrosis were observed in diseased fish. Prearo and Ghittino (1993) record a case of lipid liver degeneration in cultured trout. Asadzadeh Mangili (2000) records ichthyobodosis in 3 cm fry from a farm in Bovir Ahmadi va Kohkiluyeh. The mortality rate was 40% and infection rate with Ichthyobodo was 100%. Rainbow trout fed on raw marine fishmeal powder in Fars Province were found to be more exposed to vibiosis (pathogenic Vibrio anguillarum)(Ghazi and Akhlaghi, 1998). Diplostomiasis (infection with Diplostomum spathaceum) is reported from cultured trout in West Azarbayjan (Asadzadeh Manjili and Ghorbanzad, 1999). Zorriehzahra (2002) reports on diseases in farmed fry and Zorriehzahra et al. (2002) record enteric redmouth disease in farmed fish around Tehran. Akhondzadeh et al. (2002) and Akhondzadeh Basti and Zahrae Salehi (2003) show that the psychotropic pathogen Listeria monocytogenes is found in market and fish farm samples of this species. Soltani and Rostami (1997) record a mortality level of about 16% for farmed trout in northern Iran infected with a Cytophaga/Flexibacter-like bacterium. Soltani et al. 1999) report a yersinosis-like infection in farmed Iranian rainbow trout. Jalali et al. (2005) summarise the occurrence of Gyrodactylus species in Iran and record G. derjavini from fish in the Sardab-rud. Asadzadeh Mangili (2001) describes gas bubble disease from fish farms in Kermanshah Province. Mortality occurred mostly in fish at 100-200g and symptoms included inappetition, skin darkening, aggregation of fish at inlets and outlets in the ponds, and swimming near the surface. In 20% of the samples, unilateral or bilateral exophthalmia was observed. High day-night fluctuations in the oxygen and carbon dioxide content of the pond water was observed through excessive plant growth. Abdi et al. (2006) recorded the first mortality from acanthocephalosis with Pomphorhynchus laevis in fish farms in Maku, West Azarbayjan. Ebrahimzadeh Mousavi et al. (2007) isolated 12 species of fungi from eggs in Mazandaran hatcheries. Barzegar et al. (2008) record eye parasites from this fish including the protozoan Ichthyophthirius multifilis and the digenean Diplostomum spathaceum. Miar et al. (2008) examined fish in Valasht Lake and the Chalus River, Mazandaran and found the protozoans Trichodina trutta, Ichthyophthirius multifilis and Chilodonella hexastica as well as the metazoan Gyrodactylus derjavini. Khosravi et al. (2010) found 5 species of Saprolegniaceae fungi on eggs in two hatcheries in Kermanshah Province, three species for the first time in Iran. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Lernaea sp. on this species.

Economic importance

This species is commercially farmed in Iran for local sale. Experimental studies in the 220 ha "Neur", "Neuer" or "Noor" Lake near Ardabil have shown very good production of this species. A growth rate of 17 lbs (7.72 kg) in 3 years was reported (Anonymous, 1977). A yield of 160 kg/ha was obtained and a catch of 35,000 kg reported (Saadati, 1977) but this was unusual in that there were no other fish and food supplies were extensive. The lake is subject to winter-kill and requires stocking and/or helixers to maintain open water in winter (Bullock, 1971; Nehring, 1973b; 1973c; 1973d; 1974b; 1974c; 1974d; 1974e; 1974f; 1975b; 1975c; 1975d; 1975e; 1975f; 1975g; Nehring et al., 1974; Abbasi, 1974; 1975; Boettcher, 1974a; 1974b; Sanford, 1975; Harrington, 1976).

Kohnehchahry and Heydarpur (1973) outline methods of raising trout using submerged cages which they believe would be suitable for Iranian waters. Cage culture has been tried in Gorgan Bay, the fish escaping during several storms and numbering in the millions (B. Kiabi, pers. comm., 1994; www.ramsar.org/ram_rpt_37e.htm, downloaded 28 July 2000). They have also been cage cultured in Valasht Lake, Mazandaran where 12 cages were expected to produce 22 tonnes of fish (Abzeeyan, Tehran, 5(5):IV, 1994). Nezami et al. (2000) give a total production of fish farms in Gilan, Mazandaran and Golestan provinces (along the Caspian Sea) as 851 t.

The "Kelardasht" or Kalerdasht" Fish Farm (= Shaheed Bahonar Trout Farm of Shilat, the Iranian Fisheries Company, near Kelardasht) has an annual production capacity of 2,500,000 fingerlings of 2 g. Various other fish farms in Iran produce rainbow trout for stocking (Surber, 1969; Krasznai, 1987; Petr, 1987). More recently, Bartley and Rana (1998b) give a figure for the Kelardasht Farm of 3 million fingerlings from a pool of 2000 brood fish at a cost of 600 million rials. White (1988) reported that 60 ha of trout ponds and raceways produced 1000 t a year (see also under Cyprinus carpio). The largest fish farm in Lorestan produced 500 t a year in 2000. Edwards (1989) reported production of 1750 t from 20 licensed farms with a further 1000 t from unlicensed farms. Fingerling "trout" production (probably rainbow trout) in government hatcheries was 0.57 million in 1984, 1.81 million in 1985, 1.57 million in 1986, 3.02 million in 1987, 0.50 million in 1988, 4.23 million in 1989, 4.34 million in 1990, 1.90 million in 1991, and 2.00 million in 1992 (Emadi, 1993a). Fingerling production in 1996 was 6 million (Bartley and Rana, 1998a). Production of this trout increased 27% per year betwen 1991 and 1996, the raceway area for trout increased from 80,000 to 166,000 sq m between 1992 and 1996 and production increased from 775 to 1900 t (Rana and Bartley, 1998a). For comparison, the figures for carps (probably Chinese carps) exceeded 100 million and for mahi safid (Rutilus frisii kutum) it was 145 million. However there are also private commercial hatcheries producing about 750 t of trout each year. The Jajrud Hatchery near Tehran produced 1 million fingerlings as early as 1968 as well as several thousand marketable sized fish (Surber, 1969); and there has been a considerable expansion of hatchery capacity in Iran since that time. Rainbow trout production is only about 5% of total aquaculture production although value per metric tonne is U.S.$14,000 while for carp it is U.S.$10,000. Trout production in Ilam Province was estimated at 3.3 million fish based on 21 farms (Tehran Times, 8 August 2005, downloaded 20 January 2006).

Rana and Bartley (1998a; 1998b) outline trout aquaculture methods and problems in Iran. Most culture of this trout takes place in the Alborz and Zagros mountains which are cooler because of altitude. Farms are small, producing less than 50 tonnes per year. Raceways are used for breeding, rearing of larvae and growing out. Eyed eggs may be imported from Scotland and Norway but this may be banned in future because of disease risks. Increased Iranian expertise and costs may also favour production of eyed eggs in Iran. Survival to the eyed stage attains 80% and to the alevin stage 70%. The private sector produced 10 million fingerlings in 1996. The Manzanarbad Trout Farm near Kelardasht sold 2 million fingerlings in 1997 and produced 25 t for local consumption from 2000 sq m of concrete raceways (Bartley and Rana, 1998b). The Mahi Sera Fish Farm at Karaj produces market sized fish from 20,000 brood stock (Bartley and Rana, 1998b). These are replaced every 2-3 years to maintain growth rates. About 5000 fish are saved each year as next year's brood stock based on size, form and colour. Milt from 4 males is used to fertilise eggs from one female. In 1997, the farm produced 185 t of fish each weighing about 250 g (= 800,000 fish). Sale price is 10,000-12,000 rials/kg. The production cycle is 9-10 months at 13°C.

Trout are fed commercial pellets with a conversion efficiency of 1:1.1-1.4 (wet:dry weight). Growth to marketable size (about 30 cm and at least 225 g) takes 9-14 months depending on temperature (2-13°C) at different localities. Feed deficiencies, poor genetic stocks and low temperatures contribute to slow growth in some farms. Production in raceways is nevertheless about 15kg/cu m, a good efficiency. Cost of fingerlings to grow out farms is 120 rials for a 1 g fry, 250 rials for a 5 g fry and 15 rials/g for heavier fish (Bartley and Rana, 1998b).

There is no consistent monitoring of released fish so the success of stocking is unknown. The Dorudzan Reservoir about 100 km from Shiraz has received 400,000 fry in 1970, 2 million in 1972, 300,000 from Australia in 1974, 100,000 in 1984 and 100,000 in 1985 (Petr, 1987). Experimental fishing in 1976 gave a poor return, perhaps through competition for space with native Capoeta species, heavy poaching during migration into the inflowing river, and lowering of the reservoir level. Eggs are no longer imported and the supplies are spawned in Iran (Emadi, 1993b).

Artificial breeding of this species at the Shahid Motahary Fish Hatchery in Yasuj has been carried out 4-5 months earlier than normal using changes in light regime and hormones. The object of this work was to produce fingerlings throughout the year, increase production and save on production costs (Abzeeyan, Tehran, 7(5):IV, 1996). Dehghani and Akbari (1997) have successfully carried out gonadectomies on this species under field conditions.

A production of 5.5-8.0 tonnes of rainbow trout in rice paddies during winter has been recorded (Iranian Fisheries Research Organization Newsletter, 22:2, 2000). Production in Ilam Province was estimated at to be 3.3 million fish by the end of Iranian year 2005 (http://agriculturenews.faorne.net, downloaded 11 April 2006). However, some farms are not economical in Ilam (Rezaei and Darvishi, 2007).

Culture of this species in earthern ponds at Bafgh in Yazd Province used running, saline ground water unsuitable for drinking or agriculture. Production was 2-4 tonnes/ha in 0.40-0.75 ha ponds with salinity at 10-15‰, well water temperature 24ºC reduced to usually less than 20ºC, with a water replacement of 5-10% per day for a period of 5-6 months in autumn and winter. The highest growth performance was from fish fed 35% protein, 430 Kcal/100g energy diet and 20.6% lipid with P/E ratio of 81.4 mg protein/Kcal energy. Aeration had no significant effects on growth (Iranian Fisheries Research Oraganization Newsletter, 25:4, 2000; 54 & 55:4, 2009; Ahmadi and Alizadeh, 2004; Bahabadi, 2006). A total of 500,000 fingerlings in fish ponds were released in a two month period in 2005, expected to result in 220 t of product (www.growfish.org/Iranianreport.html). Gonad development was found to be accelerated in brackish water, males maturing two months earlier than in fresh water and ovarian development also being accelerated in Yazd fish (Fahlati Marvast et al., 2003; Fahlati Marvast et al., 2006).

Edwards (1989) gives details on culturing this species and the problems to overcome. Abtahi et al. (2002) found clove oil had no significant difference with MS222, another anaesthetic used in fish farms. Sajedi et al. (2003) have examined mtDNA in Iranian hatchery stocks to measure inter- and intra-populational diversity as an aid in fishery management by selection of the best broodstocks. Studies on the best foods and conditions for this trout in Iran have been undertaken by various authors. Kenari and Mirzakhani (2005) examined the effects of using Artemia urmiana from Lake Urmia (= Orumiyeh) enriched with n-3 HUFA, a fatty acid not present in the brine shrimp, as a larval food. Growth and survival was better than with non-enriched shrimp or with artificial feed. All-female diploid and triploid rainbow trout have been produced in Iran, such trout being unable to reproduce if escaping from fish farms (Johari, 2005). Ahmadi et al. (2005) used the Caspian Sea amphipod, Pontogammarus maeoticus, in rainbow trout food as a means of improving muscle pigmentation, the reddish colour from carotenoids in this dietary item enhancing market value. Carotenoid deposition was higher in males (R. Khodarahami in 5th International Symposium on Sturgeon, Iranian Fisheries Research Organization, 9-13 May 2005, Ramsar). Alavi Yeganeh et al. (2004) studied the use of gammarid powder as a dietary item and its effects in reducing stress in trout larvae, finding a 10% supplement to be effective. Supplementary levels of 2 and 4% of the amphipod were effective in improving flesh colour. Fani (2006a) found fishmeal to have higher digestible and metabolizable energy than soybean meal as feed. Fani (2006b) examined the use of oil by-product from soybeans on weight gain, feed conversion ratio and flesh chemical composition, recommending 10% oil by-product be added to the diet. Shadnoush (2006) showed that corn meal could be used as a nutrient in trout diet, improving weight and length. Kalbassi et al. (2004) induced tetraploidy using heat shock. Johari et al. (2006) investigated production of all-female trout using sex-reversed males and found this to be one of the best methods. Zamani et al. (2006) showed that chromosome manipulation (triploidy) had no effects on digestive enzyme activity.

The decline in caviar production has led to a project in Iran to use trout eggs, test being made on egg tenderness, colour, smell taste and shelf life (Iranian Fisheries Research Organization Newsletter, 58 & 59:1, 2009).

In addition to these reports, there have been a wide variety of studies in Iran on methods and treatments relevant to culturing rainbow trout, briefly summarised here. Others may be searched for in the Bibliography by common or scientific name of the fish. Sharifian (1998) studied increases in cortisol hormone levels in trout as a measure of stress caused by handling and induced spawning; Akhlaghi (2000c) on immunology of suspected viral diseases in cultured trout, namely infectious haematopoitic and pancreatic necrosis; Akhlagi et al. (2000) on using powdered clove tree as an anaesthetic; Mirvaghefi et al. (2000) on hydrogen peroxide for fungal control on eggs; Alizadeh et al. (2001) on the effects of dietary energy levels on growth and carcass composition in trout maintained in brackish water; Nafisi Behabadi et al. (2001) on using poultry by-product meal as a dietary replacement for fish meal fed to trout maintained in brackish water; Soltani et al. (2001) on the anaesthetic effect of clove flower extract under various temperatures and pH; Vahdati et al. (2001) on optimal conditions for phagocytic activity of granulocytes; Vahdati et al. (2001) on crowding stress and haematological parameters; Vahdati et al. (2001) on the effect of cortisol on phagocytic activity of macrophages; Akhlagi (2002) on vaccination against streptococcosis; Azari Takami et al. (2002) and Dorafshan et al. (2006) examined induction of gynogenesis by UV radiation; Dorafshan et al. (2002) induced spawning by use of a GnRH analogue; Soltani et al. (2002) on fry mortality syndrome in farmed fish in Tehran and Lorestan provinces; Yakhchali (2002) on the abundance of the parasite Ichthyophthirius multifilis on coldwater fish farms (which produce trout mostly); Fallahi et al. (2003) on isolating and identifying infectious haematopoietic necrosis virus from farmed fish; Farhangi et al. (2003) on the use of natural zeolites for reducing ammonia toxicity; Johari et al. (2003) on sperm traits and fertilisation performance of normal males and neomales; Ebrahimi (2004; 2005) on the deleterious effects of copper, cadmium and zinc, pollutants, on sperm anatomy and motility; Lorestany et al. (2004) on activating solutions and their effects on sperm motility and fertilisation rate; Sharifrohani (2004) on the use of plant essential oils as an antifungal agent for fish eggs in aquaculture; Sheikhi Moghaddam et al. (2004) on the use of alvita (sodium di-acetate) as a fungicide and bactericide; Bazyar Lakeh et al. (2005) on the effect of dietary astaxanthin on fertilisation rate; Azari Takami et al. (2005) on the nutritional effects on trout larvae of vitamin C-enriched Artemia urmiana nauplii in relation to growth, survival and environmental stress; Esmaeili et al. (2006) induced meiotic gynogenesisi by thermal shocks, the yield being best using shock 35 and 50 minutes after fertilisation; Fallahi et al. (2006) on the serological diagnosis of infectious haematopoietic necrosis disease in fry; Gholipour et al. (2006) found a density of 62 fish per square metre was optimal in concrete ponds for weight gain and feed conversion; Hosseini Najd Geramei and Irani (2006) studied the effects of photoperiod on growth, survival and and feeding of larvae; Johari and Kalbassi (2006) on alterations in the red blood cells of triploid trout; Mirzakhani et al. (2006) studied the feeding of n-3HUFA enriched Artemia nauplii as food for larvae used to increase resistance to environmental stress from pH and temperature; Mirzakhani and Mahmood Abad (2006) found feeding n-3HUFA enriched Artemia nauplii to larvae increased resistance to environmental stress (pH and temperature); Niksirat et al. (2006) found that unfertilised eggs can be held in coelomic fluid outside the body for at least 48 hours without any other treatment;  Rohani et al. (2006) on using Geranium herbarum essence to control fungi on eggs, Safari and Boldaji (2006) examined use of canola meal as a partial replacement for fishmeal to reduce cost of diets; Sear et al. (2006) evaluated sperm in relation to weight, length and condition factor of the fish; Tala et al. (2006) on gonad morphology after administration of 17α-methyl testosterone, female trout becoming functional males; Tala et al. (2006) on histological changes to the gonads after application of 17α-methyl testosterone, ranging from functional gonads to intersexes and sterility; Yarahmadi and Moghadasi (2006) on use of decapsulated cysts of Artemia urmiana in the larval diet; Zorriehzahra (2006) on fry mortality syndrome from 52 fish farms and hatcheries across Iran; Emtiazjou and Sheykhi Moghadam (2007) on the use of Alvita (sodium di-acetate) for removing fungi from eggs as an alternative to the carcinogen malachite green; Geramy et al. (2007) on photoperiod on growth, survival and feeding parameters in larvae; Hosseinie Najd et al. (2007) on initial different foods on subsequent growth of larvae; Kalbasi and Lorestani (2007) investigated the effect of various solutions on the duration of sperm motility, coelomic fluid without blood being the best; Khajeh and Peyghan (2007) evaluated blood serum parameters in fish cultured in earthen ponds and showed age-related changes; Latifi et al. (2007) found that citric acid retarded deterioration of trout during refrigerated storage; Lorestani et al. (2007) correlated various sperm quality measures of broodstocks with fish of different ages; Meshkat Rouhani et al. (2007) found that a lower protein content (40%) when the carbohydrate to lipid ratio in diet is 0.5 gave the most favourable conditions for growth; Mohagheghi Samarin et al. (2007) on the effect of temperature on oocyte quality and therefore stripping time; Naem et al. (2007) on treatment of diplostomiasis using praziquantel; Nafisi Bahabadi (2007) found that high energy feeds for trout reared in brackish water were more effective and commercial; Rezaei et al. (2007) on bacterial changes and the biogenic amines found in farmed trout stored on ice, Soltani et al. (2007, 2007) on vaccination against, and immunological responses to, Streptococcus iniae infection; Naji et al. (2007) on the effects of 17ß-estradiol valerat on sex reversal on sac fry larvae; Rahmati Leeshei et al. (2007) on the effect of phytase enzyme on  digestibility of  four vegetative foods; Safari and Boldaji (2007) on dietary lipid level effects on growth, feed utilisation and composition; Tabandeh and Akhlaghi (2007) on efficacy of conventional disinfectants on isolated Streptococcus iniae from diseased fish, sodium iodide and sodium hypochlorite being most efficient although biofilms lowered  efficiency; Varirizadeh et al. (2007)  on using the gonadotropin releasing hormone agonist (GnRHa) in emulsified form to stimulate onset of ovulation; Yousefian and Rezvani (2007) found that abnormalities were not significantly higher in brother-sister matings than controls; Zamani et al. (2007) on the activity of various digestive enzymes at initiation of exogenous feeding in comparison to Salmo caspius, activity being higher in rainbow trout; Zargarian et al. (2007) on using 2000 units/kg diets of phytase with at least 35% replacement of fish meal by soybean meal in farming trout; Akhlaghi and Sharifi Yazdi (2008) identified Yersinia ruckeri, the causative agent of enteric redmouth disease, in cultured fish from Fars Province; Alevi Yaganeh et al. (2008) on use of Gammarus powder as a dietary supplement for larvae; Alishahi (2008) on vaccinating trout against the ciliate Ichthyophthirius multifilis; Faghani et al. (2008) on the immunostimulatory effects of alginic acid and anti-streptococcus vaccine  on growth rate, feed conversion ratio, condition factor and survival rate; Johari et al. (2008) on red blood cell alterations in triploids; Karbassi et al. (2008) on the environmental management of aquaculture in raceways at Sarab Gerdu; Manouchehri and Akrami (2008) on a new automatic sorter for large amounts of fish; Mohagheghi Samarin et al. (2008) on eyeing and hatching rates under cold temperatures; Motalebi et al. (2008) on production of aflatoxins by molds in improperly stored fish meal used as food for trout; Nafisi Bahabadi and Soltani (2008) on dietary energy levels and feeding rates in fingerlings; Safari and Boldaji (2008) on replacing soybean meal with canola meal in diets; Shadnoush et al. (2008) on using acorn meal as a pellet binder and stabiliser; Shafaeipour et al. (2008) on the effects of dietary canola meal on physiology and biochemistry as it can be used to replace substantial levels of fish meal in food; Yousefian (2008) on genetic parameters of growth; Yousefian and Nejati (2008) on inbreeding depression shown as variations in percentage egg hatchability, survival of fry and weights; Zargar et al. (2008) on the distribution of infectious haematopoietic necrosis in five provinces; Iranian Fisheries Research Organization Newsletter (56:42, 2008) on use of the growth hormone somatotropin to increase body weight, daily weight gain and length; Akbari et al. (2009) on feeding larvae with HUFA and vitamin C enriched Artemia urmiana nauplii; Fazaeili et al. (2009) on deposition of pigments from plants in various tissues (feed conversion ratio and specific growth rate were affected by the experimental diets); Gashti (2009) determined that machine trimming of fillets produced less waste than hand trimming, although hand trimming was faster; Heydarnezhad and Purser (2009) assessed food anticipatory behaviour in fish hand-fed in small raceways; Imani et al. (2009) on feeding deprivation and re-feeding, showing that fish could tolerate and compensate for starvation; Jafarian et al. (2009) on the increase in growth performance and feeding efficiency in larvae fed probiotic larvae with Daphnia magna meal; Lorestani et al. (2009) on the interaction of male broodstock age and different dilutions on the production rate of eyed eggs; Mahmoudzadeh et al. (2009) on the effect of synthetic and alfalfa pigments on tissue pigmentation (feed conversion ratio and specific growth rate were affected by the experimental diets); Mousavi et al. (2009) on the antifungal activity on trout eggs of a combination of essential oils from various herbs; Mousavi et al. (2009) on identification of 7 species of Saprolegniaceae fungi from infected trout eggs; Omran and Hedayatifard (2009) on fatty acid composition; Pooramini et al. (2009) on use of yeast as a probiotic and its affect on growth, survival and carcass quality; Rafaei et al. (2009) on the bioaccumulation of copper, zinc iron and manganese in eggs during incubation, found to be lower than EPA standards; Shafaeipour et al. (2009) on effects of canola meal on growth, body composition and biochemical parameters and its use to replace fish meal in diets; Soltani et al. (2009) on the use of Zataria multiflora (Persian thyme) essential oil and its effects on egg hatchability and survival of larvae compared with hydrogen peroxide and malachite green used as anti-fungal agents; Soltani et al. (2009) on the experimental pathology of Streptococcus sp.; Iranian Fisheries Research Organization Newsletter (56:2, 2008 and 58 & 59:2, 2009) on culturing trout using Silo tanks which are easier to maintain and have higher production than cement tanks; Tavakoli and Akhlaghi (2009) on changes in serum and blood factors after infection with the pathogen Aeromonas hydrophila; Akbary et al. (2010) comparing live food and an artificial diet on survival, growth and body composition in larvae, the former two factors being better with live food; Dadgar et al. (2010) on cottonseed meal as a total replacement for soybean meal; Haghparast et al. (2010) on the antioxidant properties of various sodium salts on refrigerated sticks; Moogouei et al. (2010) on the effects of physcio-chemical parameters on growth in a raceway culture system; Zorriehzahra et al. (2010) on hematological and biochemical parameters of fry in western Mazandaran, helpful in diagnosing some diseases; etc.

The Department of the Environment has arranged fishing matches for this trout in the Lar Dam since 1994 (www.iran-doe.org/Special/LAR.htm, downloaded 29 December 2000).

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in aquaculture and aquaria, as food, in sport and in textbooks. Rainbows have been used extensively as research animals as their requirements are well known and they are readily available from hatcheries. Hatchery fish often have reduced or absent fins and deformed mouths. Farmed fish are sold frozen world-wide, the most important trout in this regard. Rainbow trout are one of the top few sport fishes in North America and of great commercial importance because of the money spent on gear, accommodation, transport, etc. by anglers in pursuit of this fish. Many books and articles have been written on the methods and joy of catching this trout. The flesh is excellent eating fresh or smoked and may be red if food is mostly invertebrates or white if food is fishes.

Conservation

Alborz trout have escaped and made their way to the Caspian Sea but the numbers and impact are unknown (Bartley and Rana, 1998b). This species is an exotic and does not need conservation but escapees may affect survival of native species.

Further work

Bartley and Rana (1998b) make various recommendations for the aquaculture of this species in Iran. The establishment of reproducing populations in the wild should be carefully monitored because of the dangers for native species posed by this exotic. Triploid all-female trout should be considered.

Sources

Sources are summaries of biology in North America such as Scott and Crossman (1973). Observed on trout farms in Iran but there is little data on escaped Iranian populations.

Iranian material: no preserved material.

Genus Salmo
Linnaeus, 1758

Members of this genus comprise about 25 nominal species found in North America and Eurasia including the Black, Caspian and Aral seas basins and the upper, and cooler, reaches of neighbouring basins in Iran and Turkey including the Euphrates River, and the Orumiyeh and Namak lakes (Berg, 1948-1949). There is a single described species, native to Iran. These fishes have teeth on the shaft and head of the vomer bone in the roof of the mouth (may be lost with age), the jaw is long, reaching to the posterior eye margin or beyond, scales are small usually more than 100 in the lateral line, colour is pale silvery with dark markings or spottings, anal fin rays (all counted) number 15 or less (16 or more in Oncorhynchus), and there are various unique osteological characters.

These are famous sport fishes and were once caught in commercial quantities. Their biology has been studied extensively and numerous books and papers have been written about them.

Salmo salar
Linnaeus, 1758

Introduced to the Caspian Sea (Mamaev, 2002). No Iranian record.

Salmo caspius
Kessler, 1877

 

Stuffed specimen, Bandar-e Anzali

Liqvan Chay trout, ca, 27.0 cm total length, 4 October 1994, courtesy of Asghar Abdoli

 

Liqvan Chay trout, ca, 27.0 cm total length, 4 October 1994, courtesy of Asghar Abdoli

Common names

آزاد ماهي (azad mahi or mahi-ye azad = free fish, used in Farsi for trout and salmon), ماهي آزاد (= mahi azad), azad mahi qezelala-ye daryacheh khazar (= Caspian Sea trout), mahi azad Darya-ye Khazar, mahi azad setareh-i Darya-ye Khazar, all for the subspecies caspius; قزل آلا (= gazalala, ghezel ala or kizil ala, meaning red spots), قزل آلاي خال قرمز (= qezelala-ye khalqermez; the Turkic qezel ala is used in Farsi for trout and hence this last means "trout with red spots" although literally it is a tautonym, "red spots with red spots"), mahi azad-ye kezel ala-ye ilbumi (= native salmon trout), qezel ala khalqermez yabumi, qezel ala-ye juibary (= brook trout), all apparently for freshwater residents; mahi azade khal sorkh; qezel ala-ye Liqvanchai (for trout in the Lake Urmia basin).

[gizilxalli or ala-balyk for freshwater residents and gizil balig or kizil-balyk for sea-run fish in Azerbaijanian; Kaspi azatmahysy or kumja in Turkmenian; ala-balukh Armenia; Kaspiiskii losos' or Caspian salmon for sea-run fish, and ruch'evaya forel or brook trout and pestrushka for freshwater residents in Russian; alabalik in Turkish; massialé in Kurdish; brown trout, brook trout, river trout, Caspian trout].

Systematics

Salmo Trutta was originally described from European rivers. Salmo caspius Kessler, 1877 was described from the Bozh'em-Promysl (or Bozhii Promysel) fishing grounds on the Kura River, Azerbaijan with 3 syntypes in the Zoological Institute, St. Petersburg (ZISP). Salmo spurius Pallas, 1814 from the Terek River in the Caspian Sea basin, Russia and Salmo lacustris var. Romanovi Kavraiskii, 1896 from Lake Tabiszchuri in Transcaucasia are usually regarded as synonyms of Salmo trutta. Salmo trutta ezenami Berg, 1948 is a subspecies from Lake Ezenam in Dagestan (ZISP 28356)(Dorofeyeva and Salmanov (2001)) though it may be deserving of species rank (Reshetnikov et al., 1997). Salmo trutta ciscaucasicus Dorofeyeva, 1967 (see also Dorofeyeva (1997) and Dorofeyeva and Salmanov (2001)) is described from the Keyranchay (ZISP 26244) and is found in drainages of the western shore of the Caspian Sea, except for the Kura River and, presumably, excepting Iranian rivers.

The native brown trout (freshwater populations) and Caspian salmon (sea-run populations) of Iran were both usually referred to Salmo trutta caspius Kessler, 1877 in the Caspian Sea and Namak Lake basins (Saadati, 1977). Derzhavin (1929b) examined two small fish from the Karaj River of the Namak Lake basin and recognised them as typical "brook trout" (Salmo trutta). Nümann (1969) considers that southern Alborz trout originate from Caspian stocks. Some authors call the species Salmo fario caspius Kessler, 1877, e.g. Dadikyan (1986). Fricke et al. (2007) and Naseka and Bogutskaya (2009) regard caspius as a full species.

Tortonese (1954), Kuderskii (1974) and Kottelat (1997) discuss the confused nomenclature of this species. Boulenger (1896), Berg (1948-1949) and Derzhavin (1929b) consider Namak Lake basin trout to be Salmo trutta macrostigma (Dumeril, 1858), originally described from Algeria as a distinct species. The Namak trout were supposedly derived from Mediterranean populations via Persian Gulf drainages but Saadati (1977) dismisses this on the grounds of a close similarity between Namak and Caspian trout and the apparent absence of trout from the Zagros Mountains and Persian Gulf drainages of Iran. Osinov (1988) could not determine a Mediterranean route based on electrophoretic evidence.

In the Lake Orumiyeh basin, apparently restricted to the "Lighvanchai" (probably the Liqvan Chay, the town of Liqvan being at 37°50'N, 46°26'E), is a population of brown trout which has been referred to as a distinct subspecies (Anonymous, 1977). It has not been formally described. Saadati (1977) considers all Iranian trout to be S. t. caspius, with those in the Namak Lake basin being a distinct "race", with some slight differences to typical S. t. caspius (see below), and those trout in the Lake Orumiyeh basin could be a new subspecies.

Earlier authors consider the Caspian basin species to be the same as the Atlantic salmon, Salmo salar Linnaeus, 1758, but blood serum electrophoresis (Ostroumova, 1970) support its position as Salmo trutta (and presumably caspius) as does karyology and osteology (Dorofeyeva, 1965; 1967) and enzyme electrophoresis (Osinov, 1984; 1988). Osinov and Bernatchez (1996) consider trout from the Caspian Sea basin to be part of their "Danubian" grouping, brown trout from the Black, Caspian and Aral sea basins as opposed to an "Atlantic" grouping from the Baltic, Barents and White sea basins (and also a tributary of the upper Volga River in the Caspian Sea basin), based on allozyme and mtDNA studies. Vera et al. (2011) concur. Novikov et al. (2008) examined allozyme variability in trout populations from Iran and found them to be similar but diverging significantly from other populations in the Caspian Sea basin. Turan et al. (2009) described two new species of trouts from northern Anatolian streams, one resident and the other migratory. A similar situation may occur in Caspian streams where there are the two life history types, although the authors caution that each situation should be investigated individually. They advocate a more cautious treatment of trout taxonomy as such results have important implications for management and conservation. Vera et al. (2011) found two main clusters, using microsatellite data, connected by gene flow among river basins, presumably by anadromous fish.

A artificial hybrid of Salmo trutta caspius with Oncorhynchus mykiss is reported on by Pourgholam and Noruzy Moghadam (1996).

Key characters

The dark-spotted back, light halos around some of the dark-coloured flank spots, caudal fin not or only weakly spotted, teeth on the vomer shaft, and only 9-15 total anal fin rays are distinctive. ? from S. trutta?

Morphology

Salmanov (1990) has demonstrated that brook, lake and sea forms of trout from the Caspian Sea basin differ in head and body proportions and in fin shapes although no populations in Iran were considered or given taxonomic status. Osinov (1988) maintains that migratory and non-migratory populations are the same species.

In the Caspian Sea basin dorsal fin with 3-5 unbranched and 7-12 branched rays, anal fin with 2-5 unbranched and 6-10 branched rays, pectoral fin with 10-14 branched rays, and pelvic fin with 6-9 branched rays. Lateral line scales 111-132. Gill rakers 16-23. Pyloric caeca 26-61. Vertebrae 55-61. Scales in lateral series for S. caspius 140-160, vertebrae 59-61 (Derzhavin, 1929b; Behnke, 1965). See also below under Zoogeography. The chromosome number is 2n=78-84 (Klinkhardt et al., 1995), 2n=80 (Annual Report, 1995-1996, Iranian Fisheries Research and Training Organization, Tehran, p. 61, 1997; Kalbassi et al., 2006) or 2n=80-82 (Dorofeeva, 1998). The chromosome formula is 14M + 10SM + 56T with NF=104 (Kalbassi et al., 2006).

Meristic values for Iranian specimens are:- dorsal fin branched rays 9(1), 10(5) or 11(1), anal fin branched rays 8(7), pectoral fin branched rays 12(7), pelvic fin branched rays 7(1) or 8(6), and total gill rakers 17(3), 18(3) or 19(1).

The Liqvan Chay fish are similar to S. caspius in all except colour, gill rakers being somewhat higher (18-22, mean 20.2) and lateral series scales being higher (132-154, mean 142) but sample sizes were small.

Sexual dimorphism

Abdurakhmanov (1962) reports on two populations form Azerbaijan where he found head length, predorsal distance, snout length and lower jaw length to be greater in males and postorbital length greater in females for both populations, while body depth, dorsal and anal fin heights, and pectoral and pelvic fin lengths are greater in males and eye diameter, pelvic-anal fin distance, head depth and interorbital width are greater in females for one population.

Colour

A specimen from the Anzali Mordab had a dark brown-grey back and top of the head with some round black spots, silvery-yellowish sides with 3-4 longitudinal rows of small red spots with whitish halos, a large black spot on the preoperculum and several smaller spots on the operculum. The belly and lower head surface were white with minute dark melanophores. The dorsal fin was a pale grey with 3-4 horizontal rows of black spots and 1-2 series of 4-8 ellipsoid red spots vertically. The adipose fin was overall yellowish with some grey pigment while the upper half of the fin was more orange in colour. The pectoral, pelvic and anal fins were a pale yellowish or nearly colourless with grey pigment. The anterior 2-3 rays were coloured orange. The caudal fin was greyish with the upper and lower marginal rays dark orange. All fins lacked spots except one red spot observed on the dorsal fin of a larger fish (19.8 cm total length). Smaller fish (13.7-16.2 cm fork length) had 12-14 parr marks. The red flank spots are recognised in the name "kizil ala". Generally in the sea, this species has numerous, dark, x-shaped spots on silvery flanks although some individuals have almost none. The fish caught by Holmes (1845) mentioned below had a dirty olive-green back, dark brown and red spots on the flanks, a roseate tint to the flanks, and a golden belly. The migratory form was dark blue on the back, silvery on the flanks and belly, and had dark spots on the flanks.

Prosek (2003) illustrates in colour a trout from the Tigris River in Turkey, from the Caspian Sea basin in Turkey and Armenia, and from the Liqvan Chay.

The Liqvan Chay population has 54-400 red to orange spots along its flank (compared to only 30-50 generally in European brown trout) giving it a ruby-red sheen (Anonymous, 1977). Only one Liqvan Chay fish had less than 100 red spots based on 13 fish examined by Saadati (1977) compared to a range of 27-134 for 13 S. t. caspius from the Shah Neshin River near Ardabil, where only 2 fish had more than 100 spots. The spotting pattern consists of profuse black and red spots and is distinct from all other Salmo trutta sensu lato which have fewer and larger spots (Saadati, 1977). Colour photographs, courtesy of Asghar Abdoli of the Agricultural and Natural Resources University, Gorgan, of 3 fish from the Liqvan Chay dated 4 October 1994 with a total length of 27.0 cm showed two fish with about 87 red spots while the other had about 200 (accurate counts not possible because of body curvature and silvery reflections). There was considerable variation in spot size among the three fish, the one with most spots having generally smaller spots which give the sheen referred to above, while the other two fish have more discrete and larger spots. It is unknown whether the Liqvan Chay population has been contaminated with other stocks of Salmo trutta.

Size

The sea-run Caspian trout attains a larger size than freshwater populations. Attains 51 kg and 1.5 m but most seen in Iran were 10-15 kg (Walczak, 1972). Length reaches 118 cm and weight 21 kg in Iranian samples but such large fish are rare in Iran (Farid Pak, 1968b). Most Iranian catches are 55-105 cm and 1.8-12.7 kg (Farid-Pak, no date). Holmes (1845) reported one fish caught at the mouth of the "Mazzur" River weighing 16 lbs (7.3 kg) and measuring 38.5 inches total length (0.94 m). Nümann (1969) reports fish almost 1 m long in the Karaj Reservoir in 1967. A fish 78 cm long and weighing 8 kg was worthy of note in 1996 (Anonymous, 1996c).

Distribution

The Caspian salmon is found in the Caspian Sea and enters Iranian rivers to spawn as well as being resident in Gilan, Mazandaran and Golestan provinces (Sheil, 1856; Nedoshivin and Iljin, 1929; Kozhin, 1957; Nümann, 1966; R. J. Behnke files; Armantrout, 1980; Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 28, 1996; Abbasi et al., 1999; Abdoli, 2000). Formerly known from the Anzali Mordab and its tributaries (Holčík and Oláh, 1992). Reported prolifically from the Seh Hazar, Chalus and Babul rivers (Fortescue, 1920). Found in the Safid River, Kargan River near Hashtpar, Chalus and Babol rivers. Rare in the Aras (Berg, 1948-1949). Abundant in the upper reaches of the Gorgan (G. S. Karelin cited in Berg (1948-1949) although Karelin only visited the lower Gorgan himself). Reported as S. t. caspius from the Sardab and Tonekabon rivers, Gorgan Bay, the southeast Caspian Sea, southwest Caspian Sea and south-central Caspian Sea and as S. t. fario from the Tajan, Babol, Haraz, Sardab, Tonekabon, Pol-e Rud, and Safid rivers (Kiabi et al., 1999; Abdoli and Naderi, 2009). Jolodar and Abdoli (2004) give its distribution in the Caspian basin as in upstream waters from the Aras to the Tajan, absent in the Atrak and Gorgan rivers.

Also found in the Jajrud (Ouseley, 1819-1823), in the Karaj River (Fortescue, 1924; Derzhavin, 1929b; Berg, 1949), and from the vicinity of Damavand (? Damavand River) (Fraser, 1825), and numerous other localities in the Namak Lake basin (see Armantrout (1980) for localities from the 1970s and Abdoli (2000) for more recent general distributions). However Nümann (1969) seems to indicate that fish in the Karaj River were stocked about 100 years previously from Caspian Sea rivers. Reports from the Zagros Mountains in Iran are uncertain and those from mountains near Kerman even more so! (Walczak, 1972). These last two records have not been confirmed by specimens and with introduction of exotic material may now be impossible to verify. Fraser (1834) reports that "Trouts are found in several of the streams of Azerbijan and Kurdistan". It may be worth noting that Heckel (1843b:995) reports that the collector Theodor Kotschy "did not find trout....in the mountains of Kurdistan. Since we know our collector's diligence, we doubt that trout occurs there" but in (1846-1849a:254) Heckel states "we must add one trout (Salmo), which tastes excellently according to the report presented by our traveller; that trout occurs relatively frequently in the mountains of Kurdistan, but has not been seen by us". This is merely confusing: did Kotschy later find trout or change his account, or did Heckel recall Kotschy's accounts incorrectly?

The Liqvan Chai in the Lake Orumiyeh basin contains a distinctive trout and Liqvan Chay trout were stocked in the Ab-e Bazuft of the upper Karun River (Tigris River) basin in the autumn of 1975 but this failed (B. Sandford, in litt., 1979; Prosek, 2003).

Zoogeography

The origin of Iranian trout is probably from the north via glacial lakes during the last or earlier ice ages and via the Volga River system or, possibly, from the west via the Black and Mediterranean sea basins (Tortonese, 1954; Nümann, 1969; Kuderskii, 1974; Osinov, 1984; Farid-Pak, 1991). The initial colonisation of the Caspian Sea by this trout apparently occurred much earlier than the late glacial period on electrophoretic evidence (Osinov, 1988). The question of a northern origin remains open however on present biochemical evidence (Osinov and Bernatchez, 1996). The division between their Atlantic and Danubian groupings occurred about 0.5-0.9 million years ago based on their data, perhaps 0.7-2.0 million years ago when the division of European and North American groups of salmonids are taken into account. These authors note that populations within the Caspian Sea basin have unique gene pools, suggestive of reproductive isolation, and these fish are not simply the subspecies S. trutta caspius. Bernatchez (2001) considers all Caspian trout to belong to the Danubian or Ponto-Caspian lineage, one of five lineages, and that the centre of origin was probably from drainages associated with the Cauacasus region of the Black Sea. The major demographic expansion of this lineage probably occurred about 270,000-290,000 years ago. Bernatchez (2001) considers that populations of each sea basin occupied by the Danubian lineage should be recognised as distinct evolutionary lineages but he does not give them names.

Trout in the Zagros Mountains may be Salmo trutta macrostigma (Dumeril, 1858), if this is a valid subspecies, while some authors consider Zagros trout to be S. t. caspius on zoogeographical grounds but pure samples of this population have never been examined systematically (Behnke, 1965). Boulenger (1896) and Berg (1948-1949; 1949) consider trout in the upper Euphrates and in the Namak Lake basin to be S. t. macrostigma. Namak Lake trout are then Pliocene relicts of this Mediterranean subspecies from an invasion via the Persian Gulf region. However, the Namak Lake fishes generally show evident affinities with the Caspian Sea basin ichthyofauna and the trout themselves are very similar in appearance.

The Namak Lake basin trout are generally held in Iran to have been planted by the royal family in the nineteenth century (Saadati, 1977). The royal family had sporting camps on the Lar River and the "Varang-e Rud", a tributary of the Karaj River. Saadati (1977) compares 30 specimens from each of these two localities and found characters to be similar except for pyloric caeca (45-58, mean 51 in the Karaj sample, 35-46, mean 39 in the Lar sample) and in scales in lateral series (131-146, mean 138) and 121-134, mean 127 respectively). He concluded that Karaj River fish are native, of relatively recent origin from the Caspian Sea basin, and only slightly divergent.

Saadati (1977) also reports on a population of trout in the Mordugh Chai (probably Mordaq Chay) in the Lake Orumiyeh basin. This population was typical S. caspius with only 17-38 red spots on the flank. It may have been of recent origin via a headwater transfer as the tributaries to the Caspian Sea basin are found nearby. A human agency may have been involved, always a complicating factor in the distribution of fish which are of sporting or commercial interest.

Habitat

Mahi azad live in the Caspian Sea proper and migrate up rivers to spawn. Moosari (1996) describes migration into the Tonekabon River of Iran. Young mahi azad stay in rivers for 2 years. There are also land-locked populations such as that of the Lar River near Damavand Mountain and this species may also be found in cool to cold lakes. Areas with clean gravel are required for reproduction. Temperatures above 15°C cause egg mortality. Adults can survive up to 29°C, 18-24°C is the optimum range and 12.4-17.6°C the preferred range. Atai Mehr et al. (2006) found juveniles to survive salinities of 0 to 12.5 g/l over 120 hours, indicative of their adaptability to changing environments. The best weight and water salinity for release of cultured juveniles was 10 g and 8.0-12.5 g/l.

In the sea, they inhabit coastal areas at 40-50 m with distinct stocks centred on the basins of the major rivers. Migrations occur from Iranian shores to Dagestan (Caspian Sea Biodiversity Database, www.caspianenvironment.org).

Age and growth

The Liqvan Chay population females only begin to mature in their third year and only 50% of males are mature in their second year. This population appears to be in good condition compared to other populations, e.g. in the Lar River, and these figures probably represent more natural values. A population estimate was 1246 fish per km compared to 11,380 per km in part of the Lar River where fish had concentrated to avoid severe drought. The life span in the Lar is 6 years (Nehring, 1975a).

There are stocks of dwarf males in Caspian rivers which do not descend to the sea.

Farid Pak (1968b) found the smallest sexually mature fish in the Caspian Sea was 53 cm long and the largest 105 cm. Most fish in the commercial catch of the Caspian Sea were 5-9 years old in the 1950s (Farid-Pak, no date). Caspian Sea fish may mature as early as 1 year for males under riverine conditions but most fish mature at 3-9 years, depending on the river (Caspian Sea Biodiversity Database, www.caspianenvironment.org). Afraei et al. (2000) examined 190 fish from the Tonekabon River and found age groups 0+ to 4+. Most fish were in the 2+ age group. Maximum size was 175 mm and 84.5 g. Condition factors were 1.268 for males and 1.257 for females. See also below under Reproduction.

Niksirat and Abdoli (2009) and Abdoli et al. (2011) found a decline in length from 77.6 cm in 1947 to 59.8 cm in 2007 (22.4%), in weight from 4880.2 g to 2486.8 g (49.5%), in absolute fecundity from 7041.8 in 1947 to 4526.1 in 1973 and 2941.2 in 1986, and in relative fecundity from 1451.4 per kg body weight in 1947 to 1372.6 in 1973 and 1199.8 in 1986 (only latter value significantly different in relative fecundity). Size classes over 75 cm have disappeared from the population over the past 60 years and mature samples were less than 50 cm. Various factors cause such declines, including fishing mortality and overfishing of prey such as Clupeonella spp. (and their loss from the exotic comb jelly), an important food which helps produce more eggs. Kheyrandish et al. (2010) examined fish from six rivers in Mazandaran and found five age classes (0+ to 4+) with the most frequent being 1+ and 2+. Condition factor ranged from 0.58 to 1.47. Females dominated over males in 5 of 6 rivers, the highest sex ratio being 6.7 females:1 male in the Pajimiane River and 0.8:1 in the Khojirood. There were significant correlations between fish length and depth and fish length and width of rivers surveyed.

Nümann (1969) found Alborz trout to reach 26 cm at the end of 3 years, corresponding to the growth rate for central European fish.

Food

Freshwater populations are known to eat mayfly larvae in Iran as well as other aquatic insects and crustaceans and terrestrial insects taken at the surface. Larger specimens elsewhere are known to take fish and crayfish, and more rarely frogs, salamanders and rodents and this probably occurs in Iran too. Tonekabon River fish (Afraei et al., 2000) fed most intensively in spring and least intensively in autumn. The main prey was Simulium, Plecoptera and Ephemeroptera.

In the sea, young feed on amphipods, mysids and shrimps while adults take common and anchovy kilkas, silversides and shads.

Reproduction

This fish runs up rivers in October-November and March-May in Iran (Roux, 1961b). Holčík and Oláh (1992) record migrations into the Anzali Mordab and a run up the Pasikhan and Siahdarvishan rivers from late September to the end of December. A second run occurred in April-May but was much smaller. Holmes (1845) reports "salmon" in rivers of Gilan from June to the end of October. Bartley and Rana (1998b) report the spawning run on the Tonekabon River to be from September to November and spawning to be from November to December. Spawning occurs on riffles of sand and gravel beds. Spawners are 8-12 years old and are mature at 700 g. About 1500 eggs are produced for each kilogramme of female. The fish examined from the Tonekabon River (Afraei et al., 2000) had an absolute fecundity of 168-379 eggs, average 268 eggs. Maximum egg diameter is 6.1 mm, number of eggs per gram reaches an average of 14.7 and absolute fertility reaches 13,468 eggs in Iranian fish which are considerably smaller than Kura River fish in Azerbaijan (Farid Pak, 1968b).

Alborz trout are ripe at 2-3 years while Caspian Sea fish spend 2-3 years in rivers, then 2 years in the sea before re-entering rivers to spawn at 60-70 cm. Spawning can occur more than once in a life span (Nümann, 1969).

In the Kura River of Azerbaijan there are two races. One enters the river in October and travels no higher than the middle reaches 600-700 km from the mouth, is sexually mature and does not weigh more than 12 kg, and spawns in the same year. The second begins to enter the Kura in October but most fish run in November and December, this race spawns in the upper reaches about one year later (8-11 months) after a long migration of over 1000 km, and attains 51 kg. There are two races also in Iranian rivers (Berg, 1959). The Kura salmon may have up to 45,000 eggs of up to 6.5 mm diameter. Kura salmon spawn only once between ages 5 and 9 years. Some Kura salmon become smolts and migrate to sea in the first year of life but most descend in their second year.

Parasites and predators

Mokhayer (1976b) records the protozoan Trichodina from the gills of trout (? this species) in the Karaj River, the cestodes Eubothrium crassum and E. rugosum, the acanthocephalan Corynosoma caspicum, and the annelid Piscicola geometra. Jalali et al. (2005) and Malmberg et al. (2007) summarise the occurrence of Gyrodactylus species in Iran and record G. derjavini from fish in the Sardab-rud. Sattari et al. (2005) surveyed this species in the Chesli and Khorma rivers, recording Cystidicoloides ephemeridarum. Sattari et al. (2004) record the nematode Cystidicoloides ephemeridarum sp. from this species in Gilan.

Economic importance

In a survey of the Lar River in June, RaLonde and Walczak (1970b) found 22 fishermen had caught 222 trout with an average length of 19.85 cm, comparable with an earlier survey by Nümann (1964). The largest fish was 27 cm long. The catch was 2.56 fish per fisherman per hour, a good rate of success (but see older records below). This rate and the production of trout in this river was expected to decline drastically with construction of a dam on the Lar. Sport fishermen took 50,000 trout from the Lar River in 1967 (Surber, 1969). Malek-Eizadi (1993) gives a recent account, in Farsi, of the Lar trout and apparently confirms its decline. Floor (2003) cites older records of fishing in the Lar River.

Azad mahi were caught in weirs, nets and with long, 3-pronged forks year round but the principal seasons were spring and autumn (Holmes, 1845; Floor, 2003). The catch in the Safid River region was largest from February to April, with a maximum in the middle of March. In 1912-1913, a total of 1180 fish were caught in the Safid River, Anzali and Astara regions. Weights reached 10-12 kg, average 7.5 kg (Nedoshivin and Iljin (1929) cited in Berg (1948-1949)). Fortescue (1920) reported 1500-2500 "salmon" from the Seh Hazar at Shahsevar, averaging 8-10 lbs (3.63-4.54 kg) each and seldom exceeding 18 lbs (8.17 kg). Nevraev (1929) reports on catches in various regions of Iran in the early years of the twentieth century. In the Astara region from 1901-1902 to 1913-1914, the catch ranged in numbers from 236 to 1563 and in the Safid River region from 1916-1917 to 1917-1918 the range was 185-663 fish. Yearly catches of mahi azad in the Anzali region alone have reached as high as 3,037 kg in 1939/1940 and as low as 90 kg in 1946/1947 before commercial fishing was prohibited (Vladykov, 1964; Walczak, 1972). The Food and Agriculture Organization, Rome reported only 1 tonne of "salmonoids" caught in Iran for each of the years 1983 to 1985, presumably the Caspian salmon. Occasional catches are taken in sturgeon nets of fish 1-4 kg in the Anzali region and elsewhere along the Iranian coast (Holčík and Oláh, 1992). Many fish are taken by poachers in traps and nets used to block spawning streams. Salehi (2008a) notes a decline in catches of this species from 13 t in 1995 to less than 3 t in 2005 and most of the catch is based on stocking rather than natural reproduction.

There is a state supported stocking programme but growth rate is slow. High summer temperatures (28°C) in the Caspian Sea would affect cage culture and survival (Rana and Bartley, 1998a).

Salmo trutta is reported to be ichthyootoxic although there are no reports for S. caspius (Coad, 1979b).

Conservation

Pietro delle Valle, who traveled in Iran from 1616 for 7 years, reported trout from rivulets in Ardabil in the Caspian Sea basin (see Pinkerton, 1758-1826, volume IX:84), a city not now noted for so salubrious an environment for salmonids.

Ouseley (1819-1823) dined while at Tehran on dried and salted azad mahi two feet long (0.61 m) from the Caspian Sea for breakfast one day in November 1811 and on fresh trout (later referred to as "kizl-áleh") from the Jajrud for dinner.

Vigne (1842) recorded catching six or seven dozen a day in the Lar River near Tehran, Charles Murray reported in 1858 that the stream "abounds so much in trout that I frequently kill 50 in an hour with a fly" (recorded in Wright (1977)), Valentine Baker (cited in Prosek (2003)) caught 50-60 trout a day up to 4 lb in weight, and Mounsey (1872) caught 450 fish up to 1.5 lbs (0.68 kg) in 3 days using flies. Anderson (1880) found that the Lar River "abounded in trout". They were still numerous there in the 1970s (Nehring, 1975a). However this population was unusual in that growth rates and population densities fluctuated drastically from year to year, probably because spawning habitat and nursery areas were the only adequate features of this river. Features lacking were feeding areas, stable banks and stream bed, good plant cover along the banks to supplement productivity, good physical and chemical water quality and water levels, and presence of predators as controls on excess reproduction. As a result, recruitment of young-of-the-year was high and led to interspecific competition, earlier sexual maturity, and stunting. Overgrazing in the Lar Protected Area (formerly a National Park) has affected the survival of trout in this region since this reduces the availability of insect food, and in 2000 this situation was exacerbated by the prolonged drought (Imamai, 2000).

Numbers of this species have declined drastically as evidenced by catch records. The catch decreased 100 times since 1959 to the early 1970s and smaller fish taken in beach seines were used by fishermen to eat as their market value was low. In the 1970s, the Shahsavar River was assessed as the best in Iran for this species as the river was in good condition. However, even this river was blocked in at least two areas for capturing fish and almost all adults were taken for "culture" by Shilat. A take of 300 fish provide as many eggs as 20 should because of poor treatment (15°C, Saprolegnia infections). The Anzali Mordab was surveyed half a mile from a guard station and 11 gill nets were found, all illegal, in a half mile stretch in half an hour (Carl Bond Archives, Oregon State University, Corvallis). Niksirat and Abdoli (2009) report gill nets used in estuaries catch migrating fish illegally and they are also caught as bycatch in the Rutilus frisii kutum fishery, but not released. About 2-3 times the the legal fishery for artificial reproduction is taken by poaching.

Attempts to increase natural reproduction by raising young to a size more likely to survive have been attempted in Iran (Andersskog, 1970). The "Kelardasht" (or Shaeed Bahonar) Fish Farm (part of Shilat, the Iranian Fisheries Company) on the "Sandar" River has a production capacity of 100,000 fingerlings of 15-20 g but this has never been achieved (Woynarovich, 1985; Krasznai, 1987). It takes about 2 years before fingerlings reach 15-20 g and 10-15 cm, large enough to release. Brood stock are collected from the Tonekabon River during the September to November spawning migration. Spawners are 8-12 years old and mature at 700 g. New brood stock are collected each year although stock from previous years are kept as backup. In 1997, 500 fish were caught in a 3:1 ratio of males to females. Eggs from two females are mixed with milt from two males. Egg yield is about 1500 eggs/kg. Growth rate is slow in this species, compounded by low rearing temperatures of 2-17°C. Fingerlings take 10 months to reach 10 g (Bartley and Rana, 1998b). In a study of the Tonekabon River migration pattern, the following conditions are found to be favourable for release of cultured fish: release size should be at least 15 g, release point 500-700 m away from the estuary, release time March and October, and for one-year-old fish smolt release is favoured over parr as the former would migrate to the sea (Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 28, 1996). Fingerling production by government hatcheries was 0.01 million in 1986, 0.05 million in 1988, 0.10 million in 1989, 0.16 million in 1990, and 0.20 million in 1991 and in 1992 (Emadi, 1993a). Fingerling production in 1995 was 0.8 million and in 1996 was 0.42 million (Bartley and Rana, 1998a; 1998b). In 2002 and 2003, 344,000 and 325,000 fingerlings were released at a cost of 2125 and 2288 million rials respectively (Salehi, 2008a). The "Gharasoo" Research Station in Sari is using cage culture of this species (Madbaygi, 1993b). Smolts weighing 25 g on average were released into cages and fed on locally prepared food (Iranian Fisheries Research and Training Organization Newsletter, 4:5, 1994). Culture of a rapid-growing triploid sea trout has been studied in Iran (Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 32, 1996; Annual Report, 1995-1996, Iranian Fisheries Research and Training Organization, Tehran, p. 67, 1997). Sayyad Borani et al. (2006) studied the effects of weight on the osmoregulatory ability of juveniles in an attempt to determine the ideal weight for release (see also above under Habitat). Weight classes were 5, 10, 15 and 20 g and the three larger classes were capable of osmoregulation in Caspian Sea water. Mojazi Amiri et al. (2005) detailed the histology of the digestive tract from hatching to parr stage in order to better understand the digestive system capabilities and enhance rearing. Asaeian et al. (2006) studied brood fish taken from the early mid and late migrations, finding those from the early period were more suitable for offspring production, passing through smoltification more rapidly. However, they noted that fish from all periods should be used to maintain genetic diversity. Bahrkazemi et al. (2006) traced the histochemical changes in the digestive tract from hatching to parr stage. Javaheri Baboli et al. (2006) examined the effects of n-3 HUFA enriched Artemia nauplii as a starter food and showed it was no better than newly hatched Artemia. Sarvi et al. (2006), Hatef et al. (2007), Niksirat et al. (2007) and Sarvi Moghanloo et al. (2007), have studied cryopreservation of sperm, sperm and seminal plasma composition, and in vitro storage of unfertilized ova in Iran as means to help conserve wild populations. Research into osmotic regulation determined that fingerlings 10 g or above have a higher chance of survival (Iranian Fisheries Research Organization Newsletter, 53:4, 2008). Ataei Mehr et al. (2007) studied the effects of weight and salinity on the number and size of Malphigian bodies in juvenile kidneys which has relevance for release of cultured fish in fresh and salt water. Saber et al. (2007) found that for larvae a diet with 50% protein and 4600cal/g energy was appropriate. Jamalzadeh et al. (2008) established haematological and serum biochemical indices for smolts, juveniles and breeders. Zamani et al. (2007) found differences in activity of some digestive enzymes in the stomach, pyloric caeca and intestine of parr and of smolt. Bahrekazemi et al. (2009) studied the relation between egg viability, egg and ovarian fluid composition and time of stripping in fish at the Kelardasht Hatchery. Eyeing and hatching rate declined with over-ripening time  although larval abnormalities remained constant. The best time to take eggs was up to 10 days post-ovulation. Mahmoudi et al. (2009) fed different levels of dietary nucleotides showed positive effects on growth performance and a decrease in liver demolition. Ramezani (2009) fed fish experimental diets and found a better feed conversion ratio and specific growth rate at lower protein levels, 50% protein supporting maximum growth. Bahre Kazemi et al. (2010) investigated the use of biochemical and histological parameters as egg quality biomarkers. Hajirezaee et al. (2010a; 2010b) examined the chemical properties of seminal fluid and its effect on sperm motility, and effects on semen characteristics of various stripping frequencies.

Salehi (2008a) carried out a cost factor analysis for fingerling production at the Kelardasht Hatchery in Iran and found labour costs to be 52% and feed 16%. The average cost of production of a fingerling was U.S.$0.84 (6269-7123  rials), more than that for sturgeon (1753-2028) and kutum (54-121). Higher costs are probably associated with lack of broodstock, lower production quantities and the scale of the hatchery. The rate of return at age 3-4 years was estimated at 0.5% of those released annually.

RaLonde and Walczak (1970b) and Walczak (1972) lists the following reasons for decline of this species: lowering of the Caspian Sea level making spawning migrations difficult, habitat changes including siltation, pollution, irrigation dams, and unscreened irrigation intake canals, and poaching. Both adults and freshly released fingerlings are heavily poached (Petr, 1987). In the Lar and Karaj rivers about 1960 professional fishermen took trout using nets, chemicals and explosives until laws regulating seasons and equipment were passed and game wardens hired to enforce them (Surber, 1969). Streams running into the Caspian have their lower 1-3 km dried up by May through irrigation demands and trout are unable to reach the sea. Stronger enforcement of laws by game guards, education of the public, particularly local people, habitat protection and improvement, and a rational exploitation system are required to protect this species. Firouz (1974; 1976) reported that the sea-run form was a major commercial species 25 years prior to his account but the deteriorating environment coupled with dynamiting, netting and trapping severely reduced populations. This fish is now completely absent from the Anzali Mordab (Holčík and Oláh, 1992). Waters where populations survived were designated "Protected Rivers" in an effort to manage this species effectively. There are special licence requirements, bag limits and seasons which anglers must observe. A fine of 10,000 rials is imposed specifically for illegal fishing of this species (Anonymous, 1977-1978).

The Liqvan Chay population has been confined to this single river by, it is believed, destruction of habitat through agriculture and domestication of sheep and goats (Anonymous, 1977). De Mecquenem (1908) said trout were abundant in the upper reaches of rivers in the Lake Orumiyeh basin. In 1975, 500 specimens of this subspecies were transplanted into the Ab-e Bazoft near Shiraz in an effort to conserve it (Anonymous, 1977).

Caspian salmon used to enter the Kura River basin, with 20% running up the Aras River on the Iranian border. Most spawning beds on the Aras became inaccessible in the 1950s with the construction of the Bagramtapinskaya Dam. The Mingechaurskaya Dam on the Kura cut off 80% of that rivers spawning beds in 1951. The former Soviets built salmon hatcheries to offset the losses but these proved to have a low efficiency. The young fish had a high mortality because of release at unsuitable high temperatures, release in poor feeding areas, and exposure to water intakes. In addition, the released fish were too small or too few to undergo significant smoltification and migration to ensure subsequent reproduction. Under natural conditions only a small proportion of juveniles undergo smoltification (Bakshtanskii et al., 1973).

Kiabi et al. (1999) and Jalali et al. (2009) consider S. caspius to be critically endangered in the south Caspian Sea basin according to IUCN criteria. Criteria include sport fishing, few in numbers, habitat destruction, medium range (25-75% of water bodies), absent in other water bodies in Iran, and absent outside the Caspian Sea basin. Kiabi et al. (1999) also consider S. t. fario to be vulnerable in the south Caspian Sea basin according to IUCN criteria. Criteria include sport fishing, medium numbers, habitat destruction, medium range (25-75% of water bodies), present in other water bodies in Iran, and present outside the Caspian Sea basin. Coad (2000a), using 18 criteria, found this species to be one of the top 4 threatened species of freshwater fishes in Iran. Nezami et al. (2000) consider this species to be endangered because of overfishing, habitat destruction and spawning ground degradation. Mostafavi (2007) lists it as vulnerable in the Talar River, Mazandaran.  Critically endangered in Turkey (Fricke et al., 2007). Jalali et al. (2009) recommend reduction in pollution, waters where fish are released should be designated Protected Rivers, cage culture should be enhanced, spawners from different populations should be used to enhance genetic diversity in restocked populations, and gamete storage protocols could be applied to commercial aquaculture or to a biological conservation programme..

Osinov and Bernatchez (1996) note that populations within the Caspian Sea basin have unique gene pools, suggestive of reproductive isolation, and these fish cannot be managed as a single subspecies, S. trutta caspius. Re-introductions and artificial maintenance of native populations could lead to loss of diversity.

Further work

Togan et al. (1995) show that two Turkish populations of brown trout are genetically distinct, at a level often found between species. Genetical analysis of suspected surviving pure populations of Iranian Salmo caspius should be carried out to determine which stocks should be receive special attention for conservation.

Sources

Iranian material:- CMNFI 1970-0551, 5, 109.9-166.3 mm standard length, Gilan, Ghaleh River near Fowman (37º13'N, 49º19'E); CMNFI 1979-0086, 2, 76.5-145.5 mm standard length, Mazandaran, Hasanabad River tributary to Chalus River (no other locality data); CMNFI 1980-0133, 3, 116.8-139.2 mm standard length, Markazi, Karaj fish hatchery (no other locality data); CMNFI 1980-0158, 1, 93.3 mm standard length, Markazi, Polurd River at Damavand Mountain after Ab-e Ali (35º51'N, 52º04'E); CMNFI 2007-0125, 3, 91.2-169.3 mm standard length, Markazi, Luniz River in Karaj River basin (no other locality data); CMNFI 2007-0126, 7, 95.2-158.3 mm standard length, Azarbayjan-e Khavari, Aldarvish River, Sabalan Mountain (no other locality data); BM(NH) 1877.7.5:1, 7, 168.1-194.7 mm standard length, Tehran (no other locality data); BM(NH) 1908.8.7:27-28, 2, 129.0-145.7 mm standard length, north slope of Elburz Mountains near Tehran (no other locality data).

Salmo trutta
Linnaeus, 1758

 

Common names

See above under Salmo caspius where general names for trout will encompass this species.

[Brown trout, German brown trout, European brown trout, sea trout, brownie, Loch Leven trout, Von Behr trout, spotted trout and liberty trout].

Systematics

See above under Salmo caspius.

Key Characters

?This species is a relative of the Atlantic Salmon and is distinguished from it by the upper jaw extending beyond the eye in adults (and below rear half of the eye in young rather than the centre in young), the gill cover has many spots, dorsal fin principal rays usually 10-14, and orange to rusty-red spots are often present on adult flanks.

Morphology

Principal anal fin rays 9-12, pectoral rays 13-14 and pelvic rays 9-10. Lateral line scales 110-136 and pyloric caeca 30-60. Males develop a hooked lower jaw when spawning and the brown colour becomes more intense and golden.

Sexual dimorphism

Males develop a hooked lower jaw when spawning and the brown colour becomes more intense and golden.

Colour

Overall colour is a light to golden brown with silvery flanks and a white to yellowish belly. The back, flanks, side of the head and dorsal and adipose fins bear black or dark brown spots, often with a lighter halo of orange, pink or red, and the flank has pink or red spots. Only brown trout have both light and black spots on the flanks. The caudal fin may have spots restricted to its upper lobe but lacks the overall radiating spots of rainbow trout. The adipose fin is orange to orange-red, the only family member with this colouration. Sea run or lake fish are more silvery, obscuring some of the spots. Spots may be x- or y-shaped. The red flank spots usually have blue halos. Young have 7-14, narrow parr marks and a few red spots along the lateral line. The adipose fin is orange with a light margin.

Size

Reaches 1.4 m and reputedly 50 kg.

Distribution 

The trout is found throughout Europe, in North Africa and east to the Aral Sea basin. It is also recorded from the upper Euphrates basin (Berg, 1949).

Brown trout were artificially planted in Gahar Lake of the upper Dez River of the Tigris River basin where viable populations existed in the 1970s in both the upper and lower lake, and more recently (B. Sandford, in litt., 1979; R. Mehrani, pers. comm., 2000). European brown trout were planted in the Caspian Sea and Namak Lake basin and established south of Dorud in the Zagros, and in the Zayandeh River dam but their origin is unknown (Coad and Abdoli, 1993). Trout were also introduced to the Karun River basin and the Zayandeh River Dam (Y. Keivany, in litt., 1992). Trout are also recorded from the the Lake Orumiyeh basin in the upper Talkheh, Zarreineh and Tatavi rivers (Abdoli, 2000) but whether these are introduced is not certain.

Zoogeography

This species is introduced in Iran. 

Habitat

Brown trout are mostly stream and river dwellers although some are in lakes and ponds. They can tolerate warmer and more turbid waters than brook trout and only the tainbow trout is more tolerant among salmonids. Brown trout may survive in areas no longer suitable for Brook Trout because deforestation has increased stream temperatures and agriculture and industry have increased pollution and turbidity. Ideally there should be overhanging and submerged vegetation, coarse gravel and cover such as logs, boulders and undercut banks, growth is better in alkaline waters (20-200 mg L-1, total dissolved solids 800 mg L-1 for culturing this species), a depth of 7-58 cm is needed for spawning, a winter flow of less than 15 cm sec-1 is required, preferred temperatures are 10.0-17.6°C (other reports give 21.1°C and temperatures up to 24°C are tolerated) and for spawning 6.7-8.9°C, dissolved oxygen is optimal at 7-9 mg L-1 and pH range tolerated is 5.0-9.5 although the optimal range is 6.8-7.8.

Age and Growth

Life span is over 18 years with maturity attained at 2-4 years. Çetinkaya (1999) found the fish in the Çatak stream, of the upper Tigris River, Turkey to have an age range of 1-8 years, a fork length of 8.4-39.0 cm, a weight of 6.7-756 g, and a sex ratio 2.45:1 male:female. This population may not be the same taxon as introduced populations in Iran.

Food

Food is aquatic and terrestrial insects, crustaceans, molluscs, various fishes, frogs, salamanders and even small mammals. Most food is taken as drift, the trout positioned in slower water behind a rock and darting out into faster current to seize prey. Predators include other fishes, birds, water-snakes and otters. Çetinkaya (1999) found the diet of fish in the Çatak stream, of the upper Tigris River, Turkey to be Trichoptera, Ephemeroptera, Gammarus, chironomid pupae and larvae, and Plecoptera. This population may not be the same taxon as introduced populations in Iran.

Reproduction

Spawning occurs from October to January depending on locality, usually at 7-9°C but as low as 2°C or as high as 14°C. It usually takes place in gravel stream riffles but may occur on rocky shores of lakes. The female excavates a redd into which the spawning pair deposit eggs and sperm while gaping and quivering over a 4 second period. The female covers the redd with gravel after spawning to protect the eggs. Subsequent spawning occurs, usually after a 10 hour interval. Occasionally a community redd is excavated by a number of fish spawning close together. Eggs are up to 5.0 mm in diameter and each female can contain 20,865. Young may spend about 2 years in their natal stream before going to a lake but some fish remain permanently in streams.

Çetinkaya (1999) found the fish in the Çatak stream, of the upper Tigris River, Turkey, to be sexually mature at age 4 and 16-17 cm fork length with an individual fecundity of 2349 eggs.This population may not be the same taxon as introduced populations in Iran.

Parasites and predators

None known in Iran.

Economic importance

This trout is a valuable and popular sport fish in Europe, The benefits of stocking brown trout include its difficulty of capture (and thus appeal to anglers), its longer contribution to the fishery than brook trout and rainbow trout, better survival and growth than other trouts, and its tolerance of warmer waters. There is an extensive European literature on angling methods and it is reputed to be a wilier fish than brook trout, for example. They tend to bite best in the late evening or at dawn, especially when large. These trout are caught on worms, crayfish, lures and flies. Brown trout have white to pink flaky flesh and are excellent eating. Flesh colour changes with age, to pink, and is related to diet.

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in aquaria and aquaculture, as food, in sport, in textbooks and because it has been widely introduced outside its natural range. Salmo trutta is reported to be ichthyootoxic although there are no reports for Iran or for S. caspius (Coad, 1979b; see also under the genus Schizothorax for symptoms of this egg poisoning).

Conservation

None required fro an exotic species.

Further work

The distribution and biology of this trout in Iran needs detailed investigation to separate it from native taxa.

Sources

Based on general accounts of the species.

Genus Salvelinus
Richardson, 1836

The charr genus comprises many species in Europe, northern Asia and North America whose systematics have not been fully worked out. FishBase lists 83 nominal species (August 2007). The charrs have numerous small scales 105 or more in lateral series), teeth on the jaws, palatines and tongue, teeth are present on the head but not the shaft of the vomer bone in the roof of the mouth, scales in the lateral line are smaller than surrounding scales and have little or no overlap with scales before and behind, the body has light pink, red or cream spots, and the lower fins have snow-white leading edges.

Salvelinus fontinalis
(Mitchill, 1814)

Common names

ماهي آزاد چشمه اي (= mahi azad cheshmehi or azad mahi cheshmehi, meaning spring free fish, i.e. spring salmon, free fish being used for salmon and trout species in Farsi), qezel ala-ye juibary (= brook trout).

[brook trout, brook charr, speckled trout].

Systematics

Salmo fontinalis was originally described from the vicinity of New York city.

Key characters

This species is characterised by 109-132 lateral line scales, 8-13 anal fin principal rays, light-coloured spots on the body, teeth on the head of the vomer bone in the roof of the mouth, pectoral, pelvic and anal fins with a white leading edge followed by contrasting black, truncate caudal fin, dorsal and caudal fins have wavy, dark lines and blotches and the back has dark or light green or cream, worm-track markings (vermiculations).

Morphology

Dorsal fin principal rays 9-14, pectoral rays 10-15 and pelvic rays 7-10. Scales are minute, horizontal and irregular ovals with a central focus, few circuli and no radii. The pelvic axillary scale is well-developed. Gill rakers 13-22, reaching the second adjacent raker whem appressed. Pyloric caeca 20-55. The gut is s-shaped. The chromosome number is 2n=84 (Klinkhardt et al., 1995).

Sexual dimorphism

Spawning males develop a hooked lower jaw or kype.

Colour

The back is olive-green to dark brown or blackish fading to a silvery-white belly. Flanks have a red to yellow tint. Flank spots are pale but there are also small, red spots with blue halos. The pectoral, pelvic and anal fins are yellow, orange, or reddish behind the white and black leading edges. Sea-run fish have a blue-green back and silvery flanks with a purplish tinge. Spots are obscured except for the red ones. Brook trout in large lakes are also more silvery than stream resident fish. The jaw tips and the roof of the mouth are blackish. Spawning males are much brighter in overall colour and have an orange-red lower flank and upper belly, bordered below by black on each side which delimits the white belly. Young have 6-12 brown parr marks, the widest equal to eye diameter, and small red, yellow or blue flank spots. The white leading edge to the lower fins is apparent.

Size

Attains a reputed 86.0 cm and 8.0 kg, possibly 9.39 kg, but most are smaller.

Distribution

In North America this species is found from the shores of Hudson Bay and Labrador south in marine waters to Cape Cod in the east and Georgia in the Appalachian Mountains, west through all the Maritime provinces, Québec and Ontario (except the extreme west) to northeast Manitoba. Widely introduced to western Canadian provinces, the U.S.A., South America, Europe, Asia and Australasia.

A private hatchery on the Jajrud imported over 1 million brook trout eggs which were raised to fingerling size only for most to be lost in a flood in 1968. Some were planted in the Jajrud and in the Latian Reservoir in the Namak Lake basin. Survival remains unknown. Also recorded from the Sardab and Chalus rivers of the Caspian Sea basin (Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 26, 1996) but this is possibly a misidentification for Oncorhynchus mykiss.

Zoogeography

An exotic species from North America, not closely related to native Iranian salmonids.

Habitat

Brook trout are found in cool waters of streams and lakes, usually less than 20°C, adults preferring 14-16°C. Spawning requires temperatures below 15°C, and below 9°C for optimal hatching. The upper lethal limit is 25°C for adults and 20°C for newly hatched fish. pH range is 4.1-9.5. Pools, underneath banks, under overhanging bushes or behind rocks are favoured spots. During summer months they retreat to deeper water, to about 8 m, in lakes. Some populations run to sea in Hudson Bay and Atlantic Canada. They stay in coastal waters, not moving more than a few kilometres from their natal stream. Populations in the North American Great Lakes live and feed mostly in the lake and run up natal streams to spawn.

Age and growth

Maximum life span is over 20 years but most reach only 5 years in North America. Maturity is attained at 2-3 years, some males at 1 year. Stunting is common in small streams while sea run fish grow faster than freshwater ones. Optimum growth temperatures are 10-19°C.

Food

Food in North America includes aquatic and terrestrial insects, molluscs, various fishes, frogs, and even snakes, mice, voles and shrews. Stream-dwelling fish feed heavily on drifting aquatic organisms during spring run-off but in summer as drift decreases surface insects become important. Most feeding occurs in the early morning and late evening although some food is taken throughout the day. Diet shifts in response to competition with other species. Sea run fish take various marine invertebrates and fishes. Sea run adults in spring and summer eat crustaceans and fish in lower estuarine areas while young are in the upper estuary eating crustaceans and insects. During the fall in the river adults eat little and during winter back in the estuary consumed mostly crustaceans. There is thus a division of food resources between young and adults.

Reproduction

Spawning in North America takes place from August to December, earlier in the north and later in the south. Sea-run charr enter their natal stream in spring and summer even though spawning occurs in fall. Each year they spend 1.5-3 months feeding in the sea. The spawning ground is usually gravelly streams but may be lake shoals if there is some current or spring outflow to keep eggs oxygenated. Spring flows are preferred even in streams. Males arrive on the spawning ground first and defend a territory. Both sexes will rush at other fish entering the redd area. The female cleans a redd of debris by turning on her side and lashing her tail. Redd depth between stones is tested by inserting the anal fin. Redd construction may take up to 2 days with work being carried out both by day and night. Courtship involves gentle pushes, touches and strokes of the female by the male. The female is ready to spawn when she crouches in the redd with her genital area between the stones. The male arches his body and may press the female against the redd bottom, both fish vibrate and eggs and sperm are shed. Accessory or sneaky males may rush in to shed sperm. The female lashes her tail to push eggs into the gravel and then dislodges gravel with her anal fin to cover the eggs to depths as great as 20 cm. Yellow-orange eggs are up to 5.0 mm in diameter and number up to perhaps 17,000 per female although averages range from the low hundreds to a few thousand. Both sexes may spawn again with other fish. The eggs develop over winter, taking 165 days at 2.8°C but only 47 days at 10°C. Temperatures above 11°C will kill the eggs.

Parasites and predators

Brook trout are cannibals on their eggs and young and are eaten themselves by other fishes, water snakes, turtles, various birds and otters.

Economic importance

Brook trout are very popular sport fish in North America caught on lures, live baits and flies. These trout are easier to catch than Salmo trutta sensu lato and take a wider range of lures. They fight well but are often quite small and do not leap spectacularly like some other members of the salmon family.

The ready availability of this salmonid has made it a useful experimental fish for various physiological, biochemical, toxicological and other studies. Some reared stocks however show deformed or lost fins and distorted mouths.

Conservation

This introduced species is not in need of conservation but in fresh water their preference for cool and clear water makes them susceptible to loss if waters are dammed, channelised and polluted or if banks are eroded deforested and overgrazed, conditions not uncommon in Iran.

Further work

The survival of stocks of this species in Iran should be verified and the effects of this exotic on native species researched.

Sources

Based on general summaries on North American populations such as those in Scott and Crossman (1973) and Becker (1983) as no Iranian material to hand. Iranian populations have not been studied.

Genus Stenodus
Richardson, 1836

The inconnu genus contains a single species found in waters draining to the Arctic Ocean in Eurasia and North America and in the Caspian Sea.

Its characters are essentially those in the species description; a small bone lacking a head canal on the outer side of the lower jaw at the joint of the articular and dentary is distinctive. Branchiostegals number 8-11.

Stenodus leucichthys
(Güldenstaedt, 1772)

Common names

safid mahi (= white fish), آزاد ماهي (azad mahi or free fish), azad mahi Volga (= Volga free fish, or Volga salmon, free fish being used in Farsi for salmon), زيبا (= ziba), mahi-ye ziba or ziba mahi (= beautiful fish, in northwest Gilan, e.g. at Astara), mahi-e-azad-e-ziba, inkonu.

[belorybitsa in Russian; nelma or azatmahy in Turkmenian; inconnu, conny, white salmon, Caspian inconnu].

Systematics

Salmo leucichthys was originally described from the Volga and Ural rivers, Caspian Sea and Kamchatcka, Russia.

The type subspecies in the Caspian Sea basin is the one found in Iran. The subspecies Stenodus leucichthys nelma (Pallas, 1773) of White Sea, Siberian and northwestern North American drainages is more common.

Key characters

The characters of the species are those of the genus.

Morphology

Dorsal fin branched rays 8-13 after 2-6 unbranched rays and anal fin branched rays 9-16 after 2-5 unbranched rays. Lateral line scales 88-121, scales above the lateral line 8-13, and scales below the lateral line 10-12. There is a pelvic axillary scale. Scales bear numerous fine circuli, the focus is slightly subcentral posterior, the anterior scale margin is wavy but irregular in outline and the scale is generally rounded. There are no radii. There is a single flap between the nostrils.Gill rakers number 17-27 and reach the fourth adjacent raker when appressed. Pyloric caeca number 191-193 and the gut is s-shaped. Vertebrae number 64-70.

Sexual dimorphism

Males develop tubercles on the head and sides of the abdomen during spawning.

Colour

Overall colour is silvery without spots.

Size

Reaches 1.5 m and 28 kg, rarely 40 kg, possibly to 48 kg. On the coast of Dagestan the average size was 7.6-10.8 kg in the 1920s (Berg, 1948-1949).

Distribution

The subspecies is restricted to the Caspian Sea and its drainage. Rare in Iranian waters with 5 specimens caught and preserved from 5-7 km east of Bandar-e Anzali in the Caspian Sea from 1984-1990. Several dozen fish have been caught each year in Iran, starting in the late 1960s, all from between the mouths of the Safid and Astara rivers (Holčík and Razavi, 1992). Reported from the southeast Caspian Sea, southwest Caspian Sea and south-central Caspian Sea (Kiabi et al., 1999; Abdoli and Naderi, 2009).

Zoogeography

This species is found naturally in the Caspian Sea but was generally believed to inhabit only the northern, and rarely the western, parts to the north of 40°N. Rare specimens were reported by Kazancheev (1981) from the coasts of Dagestan and Azerbaijan in early summer. It probably penetrated into the Caspian Sea from the Arctic Ocean basin during the last glacial period via ponded lakes.

Habitat

Inconnu live in the Caspian Sea proper at depths less than 60-65 m (optimally 25-45 m) and migrate up rivers to spawn, in the Volga for 3000 km. Fat content of fish at Astrakhan at the Volga mouth was 26% of total weight but by the time fish reached the spawning grounds in the Ufa River this had fallen to 14%, and after spawning to 1.5% for females and 1.6% for males. Off Iran they are caught mostly above depths of 25 m from October until the end of December. Some are caught in beach seines in shallow water (Holčík and Razavi, 1992) but elsewhere are known from depths down to 46 m. It is said to move into the central and southern Caspian in summer and returns north when water temperatures fall below 8-10°C in the first half of September (Berg, 1948-1949). This contrasts with the catch records for Iran. Fingerlings tolerate temperatures up to 25ºC in culture ponds but in the sea it is found at temperatures below 20ºC (Caspian Sea Biodiversity Database, www.caspianenvironment.org).

Age and growth

Life span is at least 12 years in the Volga population for females and 8 years for males. Iranian fish were up to 6 years old (Holčík and Razavi, 1992). Spawning females are predominately 6-8 years and males 5-6 years old (Letichevskiy, 1981). Females on the Volga spawning grounds are 85-105 cm, 5.5-9.5 kg while males are 75-100 cm, 3.5-7.5 kg.

Food

There is no feeding on the spawning migration, the fish relying solely on their stored reserves of fat. Fat declined in one study from 21% before, to 2% after, spawning. In the Caspian Sea it feeds on fishes such as gobies (Gobiidae), Rutilus rutilus (presumably R. caspicus), Atherina boyeri (= caspia), young Sander, common and anchovy kilkas, and shads. In the sea, adults feed on kilka and silversides (97-99%)(Caspian Sea Biodiversity Database, www.caspianenvironment.org). Young feed on zooplankton 7-10 days after hatching but are predatory in their second month. Fry descend to the Caspian Sea to feed until they are sexually mature.

Reproduction

Spawning takes place in October and November at 0.2-6.0°C after an ascent up north Caspian rivers. The Volga River run peaks in December and March. Fecundity reaches 390,000 eggs of up to 2.4 mm diameter. Eggs average 26% of the female's weight. The eggs are slightly adhesive and shed on the river bottom. Spawning occurs only once or twice in the life cycle of most fish although exceptional females may spawn 3 times. Spawning occurs at intervals of 2-3 years. Incubation takes 180-200 days and hatching takes place in spring. The young immediately descend to the sea.

Parasites and predators

Eggs are eaten by Gobio species and by Lota lota.

Economic importance

This species has no economic importance in Iran but was a significant member of former Soviet fisheries, taken both in the sea and on its spawning migration. Up to 110,000 fish were taken annually in the Volga-Caspian region. It has a rich, delicate flesh with a fat content of 18-26% and is a favoured smoked product (Caspian Sea Biodiversity Database, www.caspianenvironment.org).

Conservation

This species was listed as threatened in Southwest Asia by Holloway (1976) and endangered in the Caspian Sea by Fricke et al. (2007). Kottelat and Freyhof (2007) state that it is maintained only by stocking and is on the brink of extinction. The population of this species, spawning principally in the Volga, declined drastically after regulation of this river. In the 1960s, the population was estimated to be only 2000 fish but it has recovered to 17,000 fish through artificial rearing (Mina, 1992). In the two decades 1961-1980, fish hatcheries in the Volga Delta raised 72.4 million inconnu. Gravel spawning grounds were also constructed (Letichevskiy, 1981). It has been proposed for inclusion in the "Red Book of the U.S.S.R." which forms the basis for measures to protect species (Pavlov et al., 1985). Lelek (1987) classified it as endangered. Kiabi et al. (1999) consider this species to be data deficient in the south Caspian Sea basin according to IUCN criteria. Criteria include commercial fishing, sport fishing, few in numbers, bodies), limited range (less than 25% of water bodies), absent in other water bodies in Iran, and present outside the Caspian Sea basin.

Further work

Further records of this species should be recorded to document the presence and spread of this fish in Iranian waters.

Sources

Morphology based on Holčík and Razavi (1992) for Iranian specimens.

Iranian material: None.

Comparative material: BM(NH) 1878.12.26:23, 1, 199.5 mm standard length, Bremen (no other locality data); BM(NH) 1899.7.25:22, 1, 215.5 mm standard length, Russia, River Obi (no other locality data).

Esocidae

The pikes, pickerels and muskellunge are found in fresh waters of the Northern Hemisphere. They are moderate to large-sized fishes, up to 1.4 m. There are only 5 species with 1 reported in Iran.

The family is characterised by a flattened, elongate, duck-billed snout; dorsal and anal fins far back on the body near the tail; no adipose fin; teeth on the tongue and on the basibranchial bones behind the tongue are small; jaws have large teeth; branchiostegal rays 10-20; nasal bones are present; the swimbladder is connected to the gut by a duct; intermuscular bones are forked or y-shaped; no fin spines; pelvic fins are abdominal; cycloid scales; the infraorbital sensory canal on the head has 8 or more pores; gill rakers are present as sharp denticles in patches; no pyloric caeca; the lateral line is complete; and the forked caudal fin has mostly 17 branched rays.

Pikes are predators on other fishes aided by the posterior dorsal and anal fins which facilitate rapid darts forward. They are important sport fishes, much sought after by anglers for their fighting ability, but are not very good eating because of the intermuscular bones.

Genus Esox
Linnaeus, 1758

The characters of this genus have been outlined above under the family.

Esox lucius
Linnaeus, 1758

Common names

ordak mahi (= duck fish from the snout shape), shok, shuk or shook (in Gilaki), shook chehkhab, chekab.

[durnabaligi in Azerbaijan; shchuka in Russian; pike; northern pike].

Systematics

Esox Lucius was originally described from Europe.

Key characters

The broad and flat snout and the dorsal and anal fins set far back on the body are distinctive.

Morphology

Dorsal fin principal rays 15-19, about 6-10 unbranched and 13-18 branched, principal anal rays 12-16, about 4-8 unbranched and 10-15 branched, pectoral rays 11-17 and pelvic rays 7-13. The number of branched rays may be size-related as in smaller fish more anterior rays in the dorsal and anal fins are not branched. Lateral line scales 105-148, pored scales 42-56, but difficult to count accurately. Each scale on mid-flank is a rounded rectangle. The anterior margin is indented where 1-2 radii terminate. The radii split the scale so that the segments overlap. Circuli are very fine and the focus is posterior. Gill rakers are broad and spinulose, embedded in the arch skin with the tips of the spinules protruding. There are 9-11 pores on the lower jaws (usually 5 on each jaw). Vertebrae 56-65. The chromosome number is 2n=50 (Klinkhardt et al., 1995). The gut is an elongate s-shape.

Meristic values for Iranian specimens are:- dorsal fin branched rays 14(7) or 15(9); anal fin branched rays 11(1), 12(8) or 13(7); pectoral fin branched rays 14(9), 15(5) or 16(1); pelvic fin branched rays 8(1), 9(7), 10(6), 12(1) or 13(1); pores on each lower jaw 5(22).

Sexual dimorphism

There is no obvious sexual dimorphism. Attempts have been made to sex pike by characters of the urogenital region, but these are hampered by seasonal variations. Abdurakhmanov (1962) reports on fish from Azerbaijan where head length is greater in males while predorsal distance and interorbital width are greater in females.

Colour

The overall colour is dark with light spots, although there is variation over the vast range of this species in the details. The back and upper flank are dark green, olive-green or brownish, fading to a whitish belly. The flank has 7-9 rows of greenish, yellow to whitish blotches along it. Scales have a golden tip. The head sides have wavy, golden or yellow blotches and lines and the eyes are bright yellow to golden. The dorsal, anal and caudal fins are green, yellow, orange or pale red, blotched and barred irregularly with black. The pectoral and pelvic fins are dusky to orange. Young have 8-12, wavy, white or yellow bars which become the bean-shaped blotches in adults as they gradually break up. There is a gold to green stripe along the middle of the back in some fish but others are completely dark green. There is a stripe below the eye. The peritoneum is silvery.

Size

Attains 1.75 m and about 48.0 kg, possibly to 2.13 m and 65.78 kg, despite legends of pike up to 5.0 m (Tsepkin, 1986).

Distribution

Across northern Eurasia and northern North America. Iranian populations are found in the Caspian Sea basin, from the Anzali Mordab and the Sia Keshim Protected Region to Gorgan Bay and its tributaries such as the Karasu, with other rivers including the Safid, Tajan, Babol and Haraz, and the Atrak River basin, and the Amirkelaye Lagoon near Lahijan (Derzhavin, 1934; Berg, 1936; Armantrout, 1980; Holčík and Oláh, 1992; Nejatsanatee, 1994; Riazi, 1996; Abbasi et al., 1999; Kiabi et al., 1999; Jolodar and Abdoli, 2004; Abdoli and Naderi, 2009). However, Abdoli and Naderi (2009) do not report it from the Atrak River.

This species has been introduced to Valasht Lake near Marzanabad, Avan River and Evan Lake northeast of Qazvin, the upper Karaj and middle Shur (Abhar stretch) rivers of the Namak lake basin, the Avan River near Alamut (in 1956), Ghorigol Lake near Tabriz, Marivan Lake in Kordestan, and the Haft Barm Lakes west of Shiraz (Anonymous, 1977; Petr, 1987; Niamir, 2001).

Across northern Eurasia and northern North America. Iranian populations are at the southern edge of the range for this species. Atrak River basin (Berg, 1936). Geluga, Anzali Mordab, Gorgan Mordab (Armantrout, 1980 and USNM 205941 at 37 30'N, 49 20"E). Anzali Mordab, its outlets and in the "Khalkai" (Hol…ik and Oláh, 1992). Gorgan Bay incl. Karasu and other tributaries (Derzhavin, 1934). Siah-Keshim Protected Region of the Anzali Mordab (Riazi, 1996). Introduced to Valasht Lake near Marzanabad, Evan Lake northeast of Qazvin, Ghorigol Lake near Tabriz, Marivan Lake in Kordestan and the Haft Barm Lakes west of Shiraz (Anonymous, 1977; Petr, 1987). Ameerkalaye Lagoon near Lahijan (Nejatsanatee, 1994) ?? not checked in gazeteer??Introduced to the Avan River 7 km from Alamut in 1956 (Nialmir, 2001).

Reported from the Tajan, Babol, Haraz, and Safid rivers, and the Anzali Mordab (Kiabi et al., 1999). Anzali Talab and Safid River (Abbasi et al. (1999).

Abdoli (2000) records from lower Babol, Heraz, Chalus, Tonekabon and Safid rivers; upper Karaj and middle Shur (Abhar stretch).

Zoogeography

This widely-distributed species reaches its southern range limit in Iran.

Habitat

Pike are solitary and are found in lakes and rivers where the water is still or flowing slowly as well as marshes and ponds. They are found only in the lower reaches of rivers along the Iranian shore and do not penetrate upstream (Berg, 1948-1949). O'Donovan (1882) reported that a small stream in the Atrak River drainage had many large pike lurking under bushes, stupefied by foul water, and that the Cossacks in his escort caught many of them by striking with the point of their sabres or simply whisked them out of the water by the tail. Large numbers of Rutilus frisii were seen here too and presumably the pike thrived on this food source. Riazi (1996) reports that this species is native (resident) to the Siah-Keshim Protected Region of the Anzali Mordab. They are found in the more brackish areas of the Caspian Sea, at least in Kizlyar Bay of the north Caspian, at salinities up to 4‰ under the influence of fresh water from the Volga River. Here they are found in thickets of soft and rigid vegetation, as solitary predators but also feeding beyond the vegetation limit in the sea. They aggregate only in spring for spawning and in autumn prior to wintering on the bottom (Stolyarov and Abusheva, 1997).

In fresh water, vegetation is heavy and the water warm but they usually retire to deeper, cooler water at the height of summer. Temperatures above about 30°C are usually fatal to pike. However pike are active in winter, as anglers can testify, and at this time can tolerate dissolved oxygen concentrations lower than 0.1 mg/l. pH range is 5.0-9.5 although they have been recorded as spawning at 4.2-4.4 but embryos are then malformed (Mann, 1996). Summer distribution is usually within 300 m of shore and less than 4 m deep. On windy days, pike retreat offshore in surface waters. Pike tolerate brackish water, up to about 7‰ for reproduction and 10‰ for feeding and growth. Reproduction requires living or dead vegetation (either aquatic or flooded terrestrial vegetation) in shallow, still waters protected from strong winds. Vegetation is also important for recruitment of young pike. Pike deposit their faeces at specific locations far from their usual feeding area as the faeces contain alarm pheromones recognised, and avoided by, prey species.

Age and growth

Life span is up to about 26 years but is less than half this in fast-growing southern populations. Some aquarium fish have lived 75 years. Maturity, like growth, varies with latitude and habitat, and also with quality of food. Higher temperatures may inhibit growth. Males mature at 1-6 years (30-46 cm) and females usually at 2-6 years, rarely at 1 year (31-63 cm). Females grow larger, faster and live longer than males. Growth is best at 19-21°C and is very efficient.

Nezami Balochy et al. (2005) examined this species in the Zibakenar-Kiashar Bojagh lagoon on the Caspian coast of Iran. The 122 fish were 17.7-74.0 cm long, average total length being 33 cm, and weight was 38-1100 g (average 307.3 g). The age groups were 0-9 years. Nezami et al. (2004; 2006) examined  this species in the Amirkelayeh Lagoon and found ages 1+ to 6+ years, average total length 44.8 cm (range 15.6-63.0 cm) and average weight 717.9 g (24-1700 g). Valipour (1998) investigated the pike in four areas of the Anzali Lagoon and found relatively fast growth with fish above 2 years of age mature with an average length of 32 cm.

In the Kizlyar Bay of the north Caspian Sea, males mostly mature at age 2+ with a body length of 36-40 cm while females are 3+ and 45-50 cm. Some males are mature in the first year of life at 26-30 cm and some females in the second year of life at 33-36 cm. Maximum age is 11 years in this population (Stolyarov and Abusheva, 1997). In Lake Aksehir, Turkey most males and females are mature at 2 years of age (Karabatak, 1988). Altindağ et al. (1999) give growth features for a population in a Kesikköprü Dam lake, Turkey where females reached age 5 and males age 4.

Food

Food is initially zooplankton and aquatic insects but fish begin to predominate at 5.0 cm after about 1 month's growth. Over 90% of the diet of adults is fish, but frogs, crayfish, mice, muskrats and ducklings are taken. Both sexes fast during spawning but females have rations 1.5-2.3 times as much as males in summer and winter. The daily ration was high from May to August with a June peak and very low in winter in North America.

Pike have a highly mobile eye which enables them to spot prey in almost any direction and have sighting grooves along the snout to facilitate their judgement of depth and distance. Food is seized after a rapid dart from concealment. Cylindrical fish like perch are preferred over more deep-bodied species as being easier to swallow. The prey capture process can be summarised as follows: eye movements towards prey, turning of body towards prey, stalking, darting, capture, rotating prey head first in mouth, and swallowing. Prey is sucked into the mouth which is opened just as the pike reaches its prey. Prey size is usually about one-third to one-half the length of the pike.

Valipour (1998) found feeding intensity and growth coefficient of this species in the Anzali Lagoon decreased with age. Fish in the 0+ age group fed mainly on zooplanktonic mysids while older fish took Carassius auratus, Hemiculter leucisculus, Rhodeus amarus and larvae of Alburnus chalcoides. Apparently, the pike is not restricting populations of commercial species in the lagoon but has an important role in controlling non-commercial exotics such as Carassius auratus and Hemiculter leucisculus. Abdoli (2000) lists Gambusia holbrooki, Carassius auratus, Hemiculter leucisculus, Liza saliens, Atherina boyeri (= caspia) and Alburnus charusini (= Alburnus hohenackeri) as food items for Iranian pike. The Zibakenar-Kiashar Bojagh lagoon fish had Odonata (14%) as the most frequent food, followed closely by Syngnathus caspius (13.8%) and Ponticola gorlap (13.4%). Other fish eaten included Gambusia holbrooki and pike. Nezami et al. (2004) and Nezami Balouchi

. (2006) examined  this species in the Amirkelayeh Lagoon and found the diet to be 24% Tinca tinca, 16% Proterorhinus marmoratus (= nasalis), with Esox lucius, plecopterans, frogs and water beetles at 8%, and Syngnathus caspius, Carassius auratus and Gammarus at 4%. Diet varied with season, and age and sex of the pike.

In the Kyzylagach or Imeni Kirova Bay of Azerbaijan Abramis brama, Cyprinus carpio, Rutilus rutilus (presumably R. caspicus), Rutilus frisii, Vimba vimba (= V. persa), Mugilidae, and Atherina boyeri (= caspia) are taken (Kuliev, 1989). Pike are cannibals when food is short. They compete with other piscivorous fishes, such as Sander lucioperca and Perca fluviatilis, for food. In the Kizlyar Bay of the north Caspian, the food of 2-month-old pike 3-4 cm long is mainly Rutilus rutilus  (presumably R. caspicus) fry, of pike 9-15 cm long this species plus fry of Cyprinus carpio and Scardinius erythrophthalmus, and of adults mainly Cyprinus carpio, Rutilus rutilus  (presumably R. caspicus) and Clupeonella cultriventris (= caspia) (during its spawning migration in April and May), Scardinius erythrophthalmus, Tinca tinca, Blicca bjoerkna, Alburnus alburnus (= hohenackeri), Cobitis sp., some gobies, frogs and crayfish. Sturgeon fry were once an important diet item but this declined with the decline in sturgeon numbers (Stolyarov and Abusheva, 1997).

Reproduction

The spring spawning migration in Dagestan begins at ice melt and spawning takes place from mid-March to the beginning of April when water temperatures reach 5-7°C. One batch of eggs is laid over a period of 10-15 days (Shikhshabekov, 1978). Spawning runs occur in the late evening and early night, several days before spawning occurs. Spawning takes place during the day, often in mid-afternoon, in shallow bays or flooded fields just after ice melt, generally in late March to May. Water temperatures on the spawning run are as low as 1.1°C. Spawning itself takes place at temperatures several degrees warmer than this, up to 17.2°C. Each female is accompanied by 1-2 males as she swims over vegetation in the shallow water. Both sexes roll to bring their genital regions close together, vibrate and release 5-60 eggs and the sperm. Tail sweeps scatter the eggs. This 3-10 second process is repeated many times each day. Eggs are amber, up to 3.4 mm in diameter after fertilisation, adhesive and each female can produce up to 595,000, although usually much less. In the Kizlyar Bay of the north Caspian Sea, the oldest females produce 365,000 eggs (Stolyarov and Abusheva, 1997). Eggs hatch 12-14 days later and the fry attach to vegetation by an adhesive head gland until the yolk sac is absorbed 6-10 days later.

The Kizlyar Bay population begins to spawn in late February or early March, sometimes under ice or immediately after ice melt at 4-6°C. Eggs are laid at depths of about 0.5 m and may dry out during water surges caused by northwestern winds although the increase in sea level has lessened this. Eggs adhere to vegetation but fall off after 2-3 days but do not die because of the low water temperatures and favourable oxygen conditions. Eggs hatch in 7-18 days depending on water temperature (Stolyarov and Abusheva, 1997).

Parasites and predators

Eslami et al. (1972) found helminths in 78.9% of 109 pike examined from Iran, a very high rate of infestation. The species encountered were Triaenophorus crassus, Raphidascaris acus and Contracaecum osculatum baicalensis. This latter parasite can infest man if fish is eaten smoked, salted or fried at temperatures below 50°C. Mokhayer (1976b) records the digenetic trematode Rhipidocotyle illense, the nematode larva Eustrongylides excisus, and the acanthocephalan Acanthocephalus lucii. Molnár and Jalali (1992) report the monogenean Tetraonchus monenteron from pike in Lake "Sama" in the Alborz Mountains. Ataee and Eslami (1999, www.mondialvet99.com, downloaded 31 May 2000) report Asymphylodora tinca from the gastro-intestinal tract of fish from the Anzali wetland. Naem et al. (2002) found the monogenean trematode Tetraonchus monenteron on the gills of this species from the western branch of the Safid River. Khara et al. (2004) looked at pike from the Amir Kelaieh Lagoon and recorded Raphidascaris acus, Camallanus lacustris, Eustrongylides excisus, Triaenophorus crassus, Trichodina sp., Tetraonchus monenteron, Diplostomum spatheceum, Lernaea sp., Argulus sp. and Piscicola sp. and atrributed the diversity to its piscivorous diet. Khara et al. (2006a) record the eye fluke Diplostomum spathaceum for this fish in the Amirkalayeh Wetland in Gilan. Sattari et al. (2002) and Sattari (2004) records the presence of the nematode, Eustrongylides excisus. This parasite can damage muscles in commercial species and render them unsuitable for sale. Sattari et al. (2005) surveyed this species in the Anzali, Amirkelayeh and Boojagh wetlands, recording Raphidascaris acus, Eustrongyloides excisus and Camallanus lacustris. Khara et al. (2006b) record the nematode Raphidascaris acus from this species in the Boojagh Wetland of the Caspian coast. Khara et al. (2007) examined fish from the Chamkhaleh River and recorded Raphidoscaris acus, Camallanus lacustris, Diplostomum spathaceum, Tetraonchus monenteron, Triaenophorus crassus, Corynosoma strumosum and Lernaea sp. Sattari et al. (2004; 2007) record the nematodes Camallanus lacustrisRaphidascaris acus and Eustrongylides excisus, the digeneans Rhipidocotyle illense and Diplostomum spathaceum and the monogenean Tetraonchus monenteron in this species in the Anzali wetland of the Caspian shore. Khara et al. (2004) recorded parasites from fish in the Amir Kelayeh Lagoon including Raphidascaris acus, Camallanus lacustris, Eustrongylides excisus, Triaenophorus crassus, Trichodina sp., Tetraonchus monenteron, Diplostomum spathaceum, Argulus sp., Piscicola sp., and Lernaea sp., a diversity governed by its piscivorous diet.  Barzegar et al. (2008) record the digenean eye parasite Diplostomum spathaceum from this fish. Khara et al. (2008) found the eye parasite Diplostomum spathaceum in this fish from Boojagh Kiashar Wetland in Gilan. Miar et al. (2008) examined fish in Valasht Lake and the Chalus River, Mazandaran and found the metazoans Rhaphidoscaris acus and Tetraonchus menonteron. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Lernaea sp. on this species.

Young pike are eaten by various other fishes including adult pike, birds and even large aquatic insects.

Economic importance

Nevraev (1929) reports a catch for the 1901-1902 to 1913-1914 period in the Anzali region was 1150 to 20,529 fish. Holčík and Oláh (1992) report a catch of 5836 kg in the Anzali Mordab in 1990, at 7.8% of the catch the fourth most important fish there, while annual reported catches from 1932-1964 varied from none to 98 tonnes. However it is not a favoured food fish in Iran (Vladykov, 1964).

There is some opportunity for sport fishing for this species in the Anzali Mordab and potentially in various lakes around the country where it has been introduced (Anonymous, 1977). Anglers catch this species in rivers along the Caspian shore such as in the Shazdeh River at Babol Sar (Noorbakhsh, 1993b). It is an important sport and commercial fish in other parts of its range. The catch in Turkey in 1981 was 796 tonnes and for the inland waters of the former U.S.S.R. in 1975 it was 16,101 tonnes. Stolyarov and Abusheva (1997) report a commercial stock of 2500 t with a recommended catch of 800-850 t in Kizlyar Bay in the north Caspian in the early 1990s.

The eggs or roe of this species are very poisonous as fresh extracts injected intravenously into rabbits have caused respiratory distress, convulsion and death within one hour (Halstead, 1967-1970). Curiously, Anzali lagoon "perk" eggs have been studied as a replacement for caviar (Iranian Fisheries Research Organization Newsletter, 58 & 59:1, 2009).

Conservation

Raat (1988) gives details of conditions which should exist to facilitate pike reproduction and growth, including such factors as vegetation, water levels, eutrophication, pollution, prey availability, intra- and inter-specific interactions, fishery, stocking, concentration of dissolved solids, pH levels, and temperature regimes. Vladykov (1964) noted a fish kill in the Anzali Mordab on 11 June 1962 where water chestnut (Trappa natans) had caused an oxygen deficiency in the shallow water.

Gilan Fisheries Research Station has cultured pike in earthern ponds. Pike caught in autumn and winter were injected with gonadotropic hormone from carp (4-7 mg/kg). Eggs were stripped after 48-72 hours and incubated for 10 days at 8-11°C and 7-8 days at 10-15°C. Absorption of the larval yolk sac took approximately twice as long as the incubation period.

Ramin (1999) reports on a project involving artificial spawning and raising of fry in earthen ponds. Spawning temperatures were 8-15°C from 4 February to 20 March. Males and females were a minimum of 3 years old and a maximum of 5 and 6 years respectively and weighed 0.75-4.0 kg. Eggs composed 10-20% of body weight, incubation lasted 120 degree days and yolk sac absorption 160-180 degree days. The rate of fertilization was 45-85% and swelled eggs were 2.5-3.5 mm. Absolute fecundity was 22,400-112,000. Survival from larvae to fingerlings in chicken manure enriched ponds was 20-22% over 50 days with growth to 7.4 g and 8.5 cm on average.

Pike from the Anzali Lagoon have been cultured with Chinese carps and were found to be mature in less than a year (Iranian Fisheries Research Organization Newsletter, 56:2, 2008).

Kiabi et al. (1999) consider this species to be conservation dependent in the south Caspian Sea basin according to IUCN criteria. Criteria include commercial fishing, sport fishing, medium numbers, habitat destruction, medium range (25-75% of water bodies), present in other water bodies in Iran, and present outside the Caspian Sea basin. Mostafavi (2007) lists it as conservation dependent in the Talar River, Mazandaran. Endangered in Turkey (Fricke et al., 2007).

Further work

The distribution and population numbers of of this species needs documentation in Iran.

Sources

Crossman and Casselman (1987) give a bibliography and Raat (1988) and Craig (1996) give synopses of biological data on this extensively studied species. General biology and characters are based on world-wide data. Sohrabi (1996b) gives an account of this species in Farsi.

Iranian material: CMNFI 1970-0510, 1, 250.3 mm standard length, Gilan, Golshan River (37°26'N, 49°40'E); CMNFI 1970-0535A, 1, 329.6 mm standard length, Gilan, Pir Bazar Roga (37°21'N, 49°33'E); CMNFI 1970-0542, 1, 39.0, mm standard length, Gilan, Old Safid River estuary (37°23'N, 50°11'E); CMNFI 1970-0543A, 1. 136.2 mm standard length, Gilan, Caspian Sea at Hasan Kiadeh (37°24'N, 49°58'E); CMNFI 1970-0553, 1, 136.7 mm standard length, Gilan, Sosar Roga (37°27'N, 49°30'E); CMNFI 1970-0579, 9, 71.9-111.2 mm standard length, Gilan, Old Safid River estuary (37°23'N, 50°11'E); CMNFI 1971-0343, 3, 63.8-74.7 mm standard length, Gilan, Langarud at Chamkhaleh (37°13'N, 50°16'E); CMNFI 1979-0685, 1, 149.2 mm standard length, Gilan, Safid River around Mohsenabad below Dehcha (no other locality data); CMNFI 1980-0123, 2, 126.3-209.3 mm standard length, Gilan, Safid River around Dehcha above Mohsenabad (no other locality data); CMNFI 1980-0138, 2, 148.2-164.3 mm standard length, Gilan, Safid River estuary (ca. 37°28'N, ca. 49°54'E); CMNFI 1993-0147, 1, 106.0 mm standard length, Iran (no other locality data).

Lotidae

The Lotidae or cuskfishes may be included within the cod family Gadidae. The family is found in temperate to cold marine waters in the Northern Hemisphere, with only one species always resident in fresh water. There are about 5 species in the family. Maximum size exceeds 1.5 m.

Cuskfishes ods are recognised by the single barbel usually present under the chin; 1-2 dorsal fins and 1 anal fin; the caudal fin usually extends around the dorsal and ventral tip of the caudal peduncle and is rounded; no fin spines; wide gill openings with the branchiostegal membranes free or narrowly attached to the isthmus; 6-8 branchiostegal rays; vomer bone in the roof of the mouth toothed; swimbladder not connected to the auditory capsules and with 2 slender, anterior processes; scales small and cycloid; an obvious lateral line; and the egg with an oil globule. Eggs and larvae are usually pelagic.

Genus Lota
Oken, 1817

This genus has a single species found in North America and Eurasia, rarely in Iran. The characters of the species are therefore the same as for the genus.

Lota lota
(Linnaeus, 1758)

Common names

mahi charb (= fat or greasy fish), lot.

[nalim in Russian; burbot, eelpout].

Systematics

Gadus Lota was originally described from European lakes. Iranian specimens are presumed to be the type subspecies with a long and low caudal peduncle and high meristic counts (Pivnička, 1970). Subspecies defined on numbers of pyloric caeca are not accepted since there is clinal and size variation (Kottelat, 1997).

Key characters

The chin barbel and 2 dorsal fins, both lacking spines, are distinctive.

Morphology

First dorsal fin rays 7-16, second dorsal fin rays 60-94, anal fin rays 58-86, pectoral fin rays 15-24, pelvic fin rays 5-10, gill rakers 4-12, short, rounded and spinulose, discrete or almost reaching the base of the adjacent raker when appressed, pyloric caeca 21-67, and vertebrae 50-72. The head is flattened, the anterior body rounded and the posterior body compressed. Scales on the body are minute, cycloid, embedded, with few circuli and no radii, and extend onto fin bases and gill covers, and to the level of the nostrils on the head. The second pelvic fin ray is elongated. The anterior nostril is a barbel-like tube. The gut has numerous pyloric caeca, and anterior and posterior loops. The chromosome number is 2n=48 (Klinkhardt et al., 1995).

Sexual dimorphism

Unknown.

Colour

Overall colour is a yellowish to olive-green to dark brown and may be black, with large light blotches on the head, body and vertical fins. The belly is yellow to white and may be finely spotted. All fins are strongly mottled. Mottling is most evident in younger fish; adults tend to become uniformly dark.

Size

Reaches 1.52 m and 34 kg.

Distribution

Found across northern Eurasia and northern North America. Recorded from the lower Safid River in 1921 (Derzhavin, 1934; Berg, 1948-1949) and possibly waters in Gilan but apparently rare in Iran. Berra (1981) omits the distribution in Iran and the southern Caspian Sea basin of Azerbaijan.

Zoogeography

Populations in the Kura River of Azerbaijan may be disjunct from northern populations but their characters have not been thoroughly examined. Abbasov (1980) did not report this species from the Aras River on the northern border of Iran so the Iranian fish, if not strays, may be disjunct even from the southern populations in the Kura River.

Habitat

This species favours cool, clear rivers and lakes where it is most active at night. It hides under rocks and plant roots or in holes in river banks during the day. Young fish remain in shallow weeded areas or rocky streams. It is found only in the lower reaches of rivers along the Iranian shore and does not penetrate upstream (Berg, 1948-1949). This species can tolerate 6‰ and so may enter the Caspian Sea near river mouths. Embryos of this species suffer a 50% mortality if pH rises to 8.0 (Mann, 1996). Fat reserves in the liver are used up in summer when this species is less active.

Age and growth

Maturity is attained at age 2-7 years, varying with habitat, rarely at 1 year for males. Growth is faster in southern populations and maturity is earlier than for northern populations. Maximum life span is 22 years.

Food

Food when young is mostly aquatic insects, crustaceans and molluscs while older burbot feed voraciously on fishes and their eggs and on frogs. Adults do not feed during the spawning period. Most feeding occurs at night. It is reported to be cannibalistic. Glycogene and fat are stored in the liver during spring and autumn feeding periods, allowing the fish to cope with high water temperatures in summer when the fish is less active and feeding is low, even enabling some growth and gonad development (Kottelat and Freyhof, 2007).

Reproduction

Spawning takes place from November to March, even under ice, at temperatures of 0.5-4.0°C on gravel, sand or hard bottoms. Successive matings occur with the male and female swimming head down along the bottom until the male rotates and pushes his belly against hers, releasing eggs and sperm in a cloud. The female beats her tail, mixing the gametes and dispersing the fertilised eggs. One or two females may be surrounded by many males in a spawning ball. The eggs are easily moved by water currents because of a large oil globule in the yolk but gradually sink and lodge in interstices in gravel and sand. Up to 5 million eggs are produced with a diameter about 1.9 mm. About 80 "day degrees" are needed for the eggs to hatch and since they are laid in winter this takes some weeks, e.g. about 6 weeks at 2°C.

Parasites and predators

None reported from Iran.

Economic importance

This species is of economic importance in the former U.S.S.R. and the flesh is said to be excellent as is the liver. The eggs have been used as caviar but are also reported as toxic (Halstead, 1967-1970; Coad, 1979b; see under the genus Schizothorax for symptoms of ichthyootoxism). However it is too rare in Iran to be a food fish and a potential health hazard.

Conservation

Classified as rare to intermediate by Lelek (1987) for Europe. Abdurakhmanov (1962) gives data on only one specimen from the southern Caspian Sea basin, indicative of its rarity. It may be extirpated from Iranian waters.

Further work

Anglers and commercial fishermen should report any captures of this rare species in Iranian waters. Specimens should be preserved for comparison with northern populations.

Sources

Counts are taken from Pivnička (1970).

Mugilidae

The mullets or grey mullets are found world-wide in temperate to tropical coastal waters readily entering estuaries and even resident in freshwaters. There are about 17 genera and 72 species but only 3 species are native to Iran in fresh and brackish waters and a further 2 species have been successfully introduced (the latter not mapped in Berra (2001) because they are exotics). Other species are recorded as entering the rivers of southern Iran from the Persian Gulf and Sea of Oman although identification is not always certain (see Marine List in Checklists in the Introduction). Maximum size is about 0.9 m.

This family is characterised by a compressed to subcylindrical body with a somewhat flattened head; moderate sized scales which may be cycloid but are ctenoid in most adults and extend onto the top and sides of the head; faint or no lateral line along the flank but pits or grooves on scales contain the sense organs; the eye may have an adipose or fatty eyelid forming a vertical, slit-like opening; vertebrae usually 24, rarely as high as 26; wide gill openings; gill rakers long and slender and increasing in number with growth; upper elements of the gill arch are specialised as a pharyngobranchial organ,;5-6 branchiostegal rays; a spiny and short first dorsal fin (4 spines), the second dorsal with 1 unbranched ray or spine and 6-10, usually 8, branched rays; anal fin with 2-3 spines and 8-12 branched rays; an abdominal pelvic fin with 1 spine and 5 soft rays; pelvic bones connected to the postcleithrum by a ligament; mouth transverse and small; teeth on jaws short, weak and flexible, and the lower jaw may be toothless; the stomach wall is strongly muscled (gizzard-like); and the gut is very long and coiled.

Mullets are schooling fish which feed on microscopic algae and the minute animals associated with the algae. They grub, gulp or suck (hence "mugil") bottom deposits, spitting out some of the debris and extracting nutrient from the remainder. The long gill rakers filter the food, the strong gizzard-like stomach crushes it and the long intestine (about 7 times body length) aids in digestion. Their bottom feeding leaves long patches of disturbed sediment readily visible from a distance. Eggs, larvae and young mullet are pelagic. Adults are found in large schools in coastal waters or on tidal flats. These fishes are very important economically as food eaten fresh, smoked or canned, as bait, and as cultured fish in ponds. The flesh is oily and rich but has few bones. Mugilid species in Khuzestan are thought to be the intermediate hosts of Heterophyidae flukes found in humans and carnivores (Massoud et al., 1981).

Recent studies have been carried out on culturing Mugil cephalus and Liza aurata in brackish water, in inland pools and salt and freshwater culture ponds in the northern and central parts of Iran (Emadi, 1993a; Iranian Fisheries Research and Training Organization Newsletter, 11:6, 1996; Azari Takami et al., 1997a). In April 1994, 20,000 Mugil cephalus fry weighing 0.5 g were imported from Hong Kong as part of this programme (Iranian Fisheries Research and Training Organization Newsletter, 4:8, 1994). Azari Takami et al. (1997a) successfully raised fry of Liza aurata and L. saliens at Bafgh-Yazd in central Iran in earthen ponds supplied with brackish well water. Fry in the 3-10 g range were cultivated for 100 days at 2200 fry/ha with pond fertilisation and with fertilisation and complementary feeding, giving 100 g and 120 g fish respectively. Experiments with different stocking densities and mixes of the two species were also carried out. Second year L. saliens reached 165 g and third year L. aurata reached 630 g and sexual maturity.

Experiments have been carried out on fishing methods for these fishes in Iran, comparing encircling gill nets with fixed gill nets and with purse seines (Annual Report 1992-93, Iranian Fisheries Research and Training Organization, Tehran; Annual Report 1993-94, Iranian Fisheries Research and Training Organization, Tehran). In 1994-1995, the total mullet catch was 4145 t, 85% of which was L. aurata. Catch per unit effort decreased by 50% over the preceding three years (Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 37, 1996).

Soltani and Rahanandeh (2001) describe a case of gastric bloat in Mugil capito in the Caspian Sea. Mugil capito Cuvier, 1829 is a synonym of Liza ramada (Risso, 1827), a species not known from the Caspian Sea.

The general name for mullets in Farsi is kefal or kafal.

Pillay (1972) gave a bibliography on mullets and Thomson (1997) reviewed the family, giving further details of anatomy than summarised here.

Genus Liza
Jordan and Swain, 1884

This genus is characterised by thin to moderately thick, terminal upper lip without papillae, the lower lip is directed forwards and is thin-edged, teeth are setiform, ciliiform or absent in the upper lip, ciliiform or absent in the lower lip, there is a symphysial knob to the lower jaw and the lower jaws meet at a 90º angle or more, the maxilla is bent down over the premaxilla and is either uniformly curved or is s-shaped, the maxilla end is visible when the mouth is closed, the anteroventral edge of the preorbital bone is serrate, weakly concave or kinked and ventrally it is broad and squarish, an adipose eyelid is present sometimes but is not well-developed being a narrow rim around the eye at all ages, the pharyngobranchial organ has two valves, pyloric caeca number 2-14, predorsal scales are unicaniculate, and the pectoral axillary scale is weak or absent. Thomson (1997) lists further characters but the genus is distinguished easily from Mugil (q.v.), the only other genus from Iran.

Liza abu
(Heckel, 1843)

 

Further illustrations

Liza abu, lateral view of head of above; Liza abu, dorsal view of head of above; both after Berg (1949)

Common names

بياح (biah), biah zury, zuri, kafal, shochy, do'kelki in Khuzestan (= with two sails, presumably referring to the two dorsal fins), ? derbak.

[maid, khishni, hishni, abu-khraiza, hosoon, hashsoun, abu-khraiza, or abu sukkanejn (= father of two anchors in allusion to the toothed suborbital bone according to Heckel (1843b)), all in Arabic; minghaj in Pakistan; abu mullet, freshwater mullet].

Systematics

Mugil pseudotelestes Pietschmann, 1912, described from the "Schatt el Arab bei Basra (Aschar)" (Ashar is at 30°31'N, 47°50'E), is probably a synonym (Coad, 1991b) based on the original description. Mugil hishni Misra in Hora and Misra, 1943 described from "Rivers and Hors, Iraq" (hors or hawrs are marshes) is also a synonym based on the description, an opinion concurred in with Ingham (no date) and Özdilek (2003). Thomson (1997) considers hishni to be a synonym and possibly pseudotelestes although he considers the description of this species insufficiently detailed. Mugil abu zarudnyi Berg, 1949 was described from "Ser-i-pul, 30 km from Malamir, between Deh-i-dez and Malamir, region of the Bakhtiars and Lurs, upper course of the Karun R., south-western Ian, 17 III 1904, N. Zarudnyi". Thomson (1997) considers this subspecies to be of doubtful validity given the variability of this species. Different stocks exist in the Tigris, Euphrates and Orontes rivers as evidenced by morphology (Turan et al., 2004).

Randall (1995b) places this species in the genus Chelon Rose, 1793; Thomson (1997) does not.

Material of Mugil abu, housed in the Naturhistorisches Museum Wien, is under NMW 9224-9230 (7 fish) and 67868 (2) and are syntypes. The type locality is the "Tigris bei Mossul" according to Heckel (1843b) and the catalogue lists 4 specimens.

Mugil (Liza) abu zarudnyi Berg, 1949 was described from Iranian tributaries of the Tigris River basin. The 5 syntypes of zarudnyi are in the Zoological Institute, St. Petersburg (ZISP 24336), measure 69.1-80.7 mm standard length. The date in the catalogue is 4.II.1904 (old style).

The holotype of Mugil hishni is in the Zoological Survey of India, Calcutta under ZSI F13626/1 with 1 paratype under ZSI F13627/1 (Menon and Yazdani, 1963; Eschmeyer et al., 1996).

Key characters

The high lateral scale count, long pectoral fins reaching almost level with the first dorsal fin origin when folded back (note fin tips often frayed, especially in preserved material, so not as apparent), short pectoral axillary scale, thin lips, 3 anal fin spines and 8 branched rays, relatively strong spines in the first dorsal and anal fins, and peg-like or setiform teeth (not tricuspid) in the upper jaw only, distinguish this species from other species in the genus Liza and from other mullets.

Morphology

The eye has fatty tissue covering as far as part of the iris. Lips are thin and the lower lip has a pronounced knob. Peg-like teeth are obviously present only on the upper lip, the lower lip having scattered ciliate teeth. The end of the maxilla has an s-shaped bend and its tip is exposed. The anterior edge of the denticulate preorbital is angular, bending down at the corner of the mouth. The pectoral axillary scale is weakly developed. Occasional antero-dorsal scales have double canals. Pectoral fin length is 75-78% of head length and this fin extends back to a level almost at the first dorsal fin origin. First dorsal fin spines 3-4, usually 4, second dorsal fin soft rays 5-10, usually 8 after 1-2 spines, anal fin spines 3 followed by 5-10, usually 8 soft rays, pectoral fin branched rays 13-17 and pelvic fin branched rays 5-6, usually 5. Vertebrae 21-25. Fusions and minor vertebral anomalies were described by Al-Hassan (1987) and Al-Hassan and Naama (1986b) from Iraqi fish. Lateral scales 39-53, usually 44-50. Scales are strongly ctenoid on the exposed part and the fish feels rough to touch when rubbed from tail to head. The embedded part of the scale has fine circuli. The focus is very posterior. Scale shape is rectangular with the anterior margin vertical or somewhat wavy. The anterior upper and lower corners are square or even pointed. The dorsal and ventral scale margins are straight and the posterior margin is rounded. There are a few radii (as few as 4) from the focus to the anterior margin and, where these intersect the margin, it may be slightly indented to form a wavy edge. There are 4 pyloric caeca and the gut is elongate and coiled. Gill rakers are short than the gill filaments and number 44-55. They bear teeth on the internal edge. Islam and Al-Nasiri (1978) give morphometric characters. Hoda (1978) describes the larva (as Mugil hishni) in Iraq. 2n = 50 (Balasem et al. (2000). Thomson (1997) notes that this is the most variable species in the family.

Al-Hassan (1984b) showed differences in counts of vertebrae and dorsal fin rays between fish from Basrah and the Karkheh River in Iran. Turan et al. (2004) also noted differences in morphometric data for stocks from the Orontes (= Asi), Tigris and Euphrates rivers of Turkey, attributing this to phenotypic adaptation as the stocks were genetically homogenous. Thomson (1997) notes that this is the most variable species in the family. Jawad (2004b) studied asymmetry in this species from the Shatt al Arab at Basrah and suggested pollution could be a cause. Jawad and Öktener (2007) record lordosis (axial spinal curvature) in this species in the Atatürk Dam Lake, Turkey. Jawad et al. (2010) describe a specimen from Karkheh River branch near Susangerd with a malformed caudal fin.

The nominal subspecies zarudnyi has 3-4 first dorsal fin spines, 1-2 spines and 5-7 soft rays in the second dorsal fin, the anal fin has 3 spines and 5-9 soft rays, usually 9, and scales in lateral series 41-46. The larger scales distinguish it from the typical form at Mosul in Iraq (Berg, 1949). My observations on the types show 4(4) first dorsal fin spines, 1(2) or 2(2) second dorsal fin spines and 7(3) or 8(1) soft rays, 7(1), 8(1) or 9(2) anal fin soft rays, 5(4) pelvic fin rays, 13(1) or 14(3) branched pectoral fin rays, and 39(1), 43(1), 44(1) or 45(1) scales in lateral series. One specimen is unusual in having a mix of abnormally low counts and normal counts: 3 first dorsal fin spines, 5 second dorsal fin soft rays, 5 anal fin soft rays, 5 pelvic fin rays, 15 pectoral fin rays, and 43 lateral series scales.

Meristic values for Iranian specimens, excluding the above: first dorsal fin spines 4(36); second dorsal fin soft rays 7(4), 8(31) or 9(1); anal fin soft rays 8(28) or 9(8); pectoral fin branched rays 13(1), 14(17), 15(13) or 16(4); pelvic fin branched rays 5(35) or 6(1); lateral series scales 44(5), 45(5), 46(5), 47(8), 48(2), 49(4), 50(1) or 51(1); and total vertebrae ?.

Sexual dimorphism

Unknown.

Colour

The back is a light brown or greyish or olive-green with the flanks and belly silvery or yellowish-silver. There may be an indistinct silvery stripe or 2 grey stripes along the flank. Generally the upper flank is dark, the lower flank pale, and the two clearly demarcated. The upper part of the lower flank may bear alternating light and dark stripes but these are sometimes poorly expressed. The second dorsal and caudal fins are dusky to brownish and lower fins are yellowish-white to hyaline. Scale margins are covered in small black spots. The top of the eye is lime-green and reddish-brown just below but still silvery around the pupil generally. Young fish have a reddish colour at the base of the pelvic, anal and caudal fins and have an evident mid-lateral stripe against a pale body.

Size

Attains 26 cm total length and 0.15 kg.

Distribution

Found in the Tigris-Euphrates and Orontes (= Asi) (Özdilek, 2003) river basins including the Tigris River basin in Iran in rivers and marshes such as Dez, Karkeh and Karun rivers and Shadegan Marsh, in lower reaches of rivers draining to the Persian Gulf, and in Pakistan. In Iran it is also recorded from Parishan (= Famur) Lake but it was not captured in a 1995 survey. A record from the Barm-e Shur near Shiraz is probably an introduction as this species was not caught there in the 1970s by me (Gh. Izadi, pers. comm., 2001).

Abdoli (2000) maps this species in the lower Hasan Langi, Kul, Gowdar and Mehran rivers at the Straits of Hormuz in the Hormuz basin; the lower Mand River and the lower Zohreh (= Hendijan) rivers in the Gulf basin, the Arvand, lower Karun, lower Karkeheh and the Simarreh rivers in the Tigris River basin. It is also reported from the upper Karun River as M. a. zarudnyi.

Zoogeography

As a freshwater member of a genus with many species in the sea, its origins lie with these marine relatives.

Habitat

This mullet is a freshwater species found in rivers, streams, channels, canals and drains, lakes, reservoirs and ponds, on fish farms of Iraq and neighbouring countries, occasionally entering estuaries. It usually occurs in schools. Epler et al. (2001) found it to be the dominant species of fish in lakes Habbaniyah, Tharthar and Razzazah, Iraq, comprising 72% of all fish collected. This was one of the most abundant species in the recovering marshes of southern Iraq in 2005-2006 (at almost 36% of 16,199 fishes collected)(Hussain et al., 2006) and in the marshes in the 1980's (Hussain and Ali, 2006). Nasir and Na'ama (1988) and Hussain and Naama (1989) report it from the Khawr az Zubayr in a marine environment, probably a consequence of human-induced environmental changes. It is the most abundant species in autumn in the Karun River, Iran, comprising 44.2% of the catch (Mokhayer, 1981c).

van den Eelaart (1954) reporting on Iraqi populations found this mullet in the surface waters and submerged vegetation of lakes and marshes, preferably where there is a gentle water flow. In December-January it enters rivers and deeper waters, especially in very cold winters. Ahmad et al. (1983; 1985) have shown this species can withstand abrupt increases of temperature up to 30°C and salinity up to 10‰ for 24 hours under experimental conditions. Mortality in water at 15‰ is low. Salinities up to 30‰ and temperatures up to 35°C are tolerated, presumably if increases are gradual. Ahmed et al. (2002) stated that it prefers salinities not exceeding 2‰ (later 5.6‰) but can survive abrupt transfer to 15‰. S. Cowton (pers. comm., 23 August 2005) has observed schools of this species gaping at the surface in the artificial lake around Al Faw Palace in Baghdad, presumably in response to high temperatures and low oxygen. It is often the dominant species left in small pools when marshes dry up, as in the Shadegan Marshes of Iran observed by B. W. Coad.

Age and growth

Al-Nasiri and Islam (1978) studied the age and growth of this species from commercial catches in the Shatt al Arab, Iraq. Specimens were aged by examining the otoliths and three age groups determined with 0+ fish being 8.9-10.9 cm, 1+ fish being 9.9-14.8 cm and 2+ fish being 13.6-18.2 cm total length. Length-otolith formulae were given. Al-Yamour et al. (1988) examined a population of this species in the Al-Daoodi Drain, Baghdad and found an age span up to 7 years using scales to age fish. Older age groups (5 and 6) grow more slowly than Diyala River fish, perhaps because of higher salinity in the Drain and the enriched conditions in the Diyala River due to sewage. The length-weight relationship was W = 0.034 L2.6, for males W = 0.062 L2.396,and for females W = 0.031 L2.642. The condition factor was 6.308 for males and 3.155 for females. Khalaf et al. (1986) studied a population in the Diyala River, Iraq which was polluted with sewage from the Rustamiyah treatment plant. Khishni is the second dominant species from December to March after Chondrostoma regium and in August the dominant species in catches. Fish were heavier and in better condition in the more polluted areas, but probably not as good for human consumption! Growth declines after age group 5 with few fish in age group 6. Length-weight relationships were W = -3.39 L2.64 for males and W = -2.96 L2.50 for females. Mhaisen and Yousif (1989) examined a population of khishni in Mehaijeran Creek, a side branch of the Shatt al Arab south of Basrah, Iraq. Only 3 age groups (0+ to 2+) are reported since the creek water level is affected by the tide and older fish tend to move into the deeper waters of the adjacent Shatt al Arab. Growth is faster in the creek than in other Iraqi locations. The length-weight relationship was log W = 2.794 log TL - 1.64840 for males and log W = 3.337 log TL - 2.25728 for females. Mhaisen and Al-Jaffery (1989) report 6 age groups for fish from Babylon Fish Farm west of Hilla, Iraq with a total length-weight relationship log W = 2.96057 log L - 11.19599 (and for standard length log W = 2.87574 log SL -10.24769). Fish were in good condition as they were taking food meant for cultured carp. The relative condition factor was 0.98-1.04, similar to values obtained by other authors. The von Bertalanffy growth equation was TLt = 409.6 {1 - exp [-0.06314 (t - 3.13868)]}. Al-Shamma'a et al. (1995) examined fish from the Tigris River at Al-Za'faraniyah, Iraq and found log total weight = -2.033 + 3.119 log total length (r= 0.955). Epler et al. (2001) found the oldest age groups to be 4+ in Iraqi lakes Habbaniyah and Razzazah and 5+ in Lake Tharthar. The mean condition factor was 0.97, 1.05 and 1.09 in lakes Habbaniyah, Tharthar and Razzazah respectively. Syzpuła et al. (2001), studying age and growth in the lakes Habbaniyah, Tharthar and Razzazah in 1981 and 1982, found this species grew fastest in Lake Tharthar. A decrease in length growth was noted in the second year of life and a decrease in body mass in the last years of life.

The effect of starvation over 35 days on the proximate chemical composition of this species was studied by Yesser et al. (1999). They found a sharp decline in lipid content (4.38 to 0.98%), a slight drop in protein and an increase in both moisture and ash. Both condition factor and viscera somatic index declined gradually, 1.2 to 0.83 and 10.18 to 3.63% respectively. Body weight fell by 30.59%. Lipid reserves in muscle were therefore an important form of energy storage.

Ünlü et al. (2000) report age groups of 1+ to 4+ in the Turkish Tigris River. Hussain et al. (1987) found condition factor to range from 1.12 to 1.64 for different length groups for Karkheh River fish from Iran near the Iraq border and Hawr al Hawizah.

Food

Al-Nasiri et al. (1977) studied the feeding ecology of this species in the Shatt al Arab, Iraq. Aquatic plant parts and phytoplankton are food items but the vast bulk of material is sand grains and organic detritus, the former presumably ingested while searching for detrital food. Phytoplankton is made up of diatoms (50%), green algae (36%) and blue-green algae (14%), with diatoms the most abundant numerically. Plant parts are fragments of Vallisneria leaves and rarely leaves and stems of Potamogeton and Polygonum. Islam et al. (1981) reported on the seasonal patterns of feeding in Rashdiyah Reservoir, Baghdad. However it should be noted that only 10 fish were examined for each month. Aquatic plant parts, organic debris and phytoplankton are dominant foods followed by zooplankton and aquatic insects. The diet is diverse, 48 species of Chlorophyceae, 60 species of Bacillariophyceae, 9 species of Myxophyceae and 3 of Euglenophyceae. Heavy feeding occurs in late winter, spring and fall with peaks in April and November. Empty stomachs are more common in summer. Islam and Khalaf (1983) studied diel feeding patterns in the same reservoir and found peaks at 0600 and 2100 hours. Naama and Muhsen (1986) studied the feeding periodicity of this species in the Hawr al Hammar, Iraq and found the main food to be algae, mixed with incidental sand grains. There is a single feeding peak each day at 1730 hours and feeding stops at 0200 hours. This species is a day feeder. In another study of the recovering Hawr al Hammar, diet was 76.49% algae and 20.3% diatoms with amounts of crustaceans and plants being less than 10% each, in the Hawr al Hawizah 74.3% algae and 22.5% diatoms, with amounts of plants and crustaceans being less than 10% each, and in the Al Kaba'ish (= Chabaish) Marsh 75.8% algae and 20.0% diatoms with plants and crustaceans at less than 10% each (Hussain et al., 2006). Ahmed and Hussain (1982) examined the food of young fish near Basrah, Iraq and found organic detritus to be an important food followed by phytoplankton. The smallest fish are consuming eggs in large quantities. Al-Shamma'a and Jasim (1993) studied feeding in the Hawr al Hammar in Iraq during the flood period and reports small quantities of adult insects, chironomid larvae, trichopteran larvae, other insect larvae, molluscs and worms, while copepods, cladocerans, and rotifers are important foods. This variation from the diet studies reported above may be due to the flood not allowing diatoms and algae to settle out and become available as food. Typical foods and gut contents such as organic detritus, algae, diatoms, plant tissues and sand grains are also present. Sand grains may help in trituration of food and therefore are not an accidental inclusion. Al-Shamma'a et al. (1995) examined fish from the Tigris River at Za'faraniyah and found phytoplankton ranked first in the diet followed by detritus. Debris comprised more than 37% of stomach contents on the points method. Epler et al. (1996) found detritus to be the main food in three Iraqi lakes. Epler et al. (2001) found detritus to be 83.7% of the diet in Lake Tharthar, mineral parts (presumably sand) 12.7% and plants 3.6%. In Lake Habbaniyah detritus was 73.0%, mineral parts 16.5% and plants 8.4%. In Lake Razzazah detritus was 84.4%, mineral parts 14.7% and plants 0.9%. Detritus was the principal food year round with no seasonal variations. Hussain and Ali (2006) examined feeding relationships among fishes in the Hawr al Hammar and found this species to be a herbivore, 39.6% of the diet being plants and algae, with 18% detritus.

Hussain et al. (1987) examined the food of khishni in the Karkheh River, Khuzestan, where it is the dominant species in terms of numbers caught, by two orders of magnitude. However capture techniques would affect this assessment. Food was similar to that reported from Iraqi fish. Phytoplankton is the most important food for fish 3.0-3.9 cm long, followed by organic detritus, while in larger fish this order is reversed. Silt is an important content of the gut in all fishes studied. Eighteen species of Bacillariophyceae, 6 Chlorophyceae and 4 Cyanophyceae species were recorded from gut analyses. Abdoli (2000) reports Daphnia, Hemiptera, Navicula, Nitzschia, Amphora and Cymbella species for Iranian specimens.

Reproduction

Marammazi (1994) considered it likely that this species spawns twice each year in the Zohreh River, Iran draining to the northern Persian Gulf. Chelemal et al. (2009) studied reproduction in fish from Dasht-e Azadegan west of Ahvaz. Females predominated over males (2.7:1) and maximum gonadosomatic indices were found in March for both sexes with a minimum in August, suggesting spawning in April with a spawning season extending from February to June. Details were given on histological development of the ovaries with spawning females having eggs 470.3μ in diameter.

van den Eelaart (1954) gave the spawning season in Iraq as end of February to mid-March. In the Al-Daoodi Drain at Baghdad and the Hawr al Hammar of southern Iraq spawning starts in March and continued through May (Al-Yamour et al., 1988). Mean number of eggs reaches 59,793, increasing with size and weight. In the study by Khalaf et al. (1986), all age group 1 fish are mature and females outnumber males in age groups 2 to 6 although this may be due to sampling bias. Ünlü et al. (2000) report maturity at age 1 in the Turkish Tigris River based on gonad development, with no spawning in the period August to February. Mhaisen and Yousif (1988) examined a population of khishni in Mehaijeran Creek, a side branch of the Shatt al Arab south of Basrah. Fish are ripe from January to March and females partly spent in April and spent in May. The fish spawn once. All males and females were mature at 16 cm in the creek while females mature at 13 cm in the Hawr al Hammar. Some males matured at 10.5 cm and some females at 10.6 cm in the creek. Naama et al. (1986) studied reproduction in the Hawr al Hammar and found a prolonged breeding period from November to April during which two batches of eggs could be shed. Maturity is attained at the end of the first year of life for both sexes when they are about 10 cm total length, with all males mature at 16 cm and all females at 13 cm. Egg diameter is up to 0.75 mm. Epler et al. (1996) record a relative fecundity of 359,873 to 756,118 eggs for fish of age groups 1+ to 3+ from three lakes in Iraq. Mature eggs were found from January to May in one lake and from November to March in another, maturity being two months earlier in saltier water. Daoud et al. (1998) examined reproductive biology in fish from the Tigris River at Al-Khadhmia north of Baghdad. They found a sex ratio of 1:1.45 (male:female), significantly different from 1:1, a breeding season from December to March, sexual maturity in all fish longer than 10 cm total length, smallest mature male 10.4 cm, smallest mature female 10.6 cm, absolute fecundity 12,033-63,836 eggs and relative fecundity 1237 eggs/g. Epler et al. (2001) studied reproduction in lakes Tharthar and Habbaniyah and found both sexes to achieve maturity in the first year of life at 14.2 cm. Spawning occurred in May and fecundity was 652-791 thousand eggs/kg body mass. Eggs are shed on, and adhere to, vegetation.

Rashid (19940 gives details of the histological changes in the ovaries for fish from the Qarmat Ali canal, along with several indices. The gonosomatic index showed no significant change from October to December but increased from 3.03 in December to 9.43 in January and 16.4 in February, decreasing subsequently suggesting a gradual spawning took place in March and into April. The hepatosomatic index showed a significant increase in February to 3.34 and a significant decrease in March to 1.32, suggesting the liver was involved in vitellogenin synthesis necessary to oocyte development. The somatic index increased from 0.65 in October to 1.15 in December as the fish lay down reserves, decreasing to 0.53 in February and rising throughout March and April.

Parasites and predators

Bykhovski (1949) reports a new species of monogenetic trematode, Ancyrocephalus fluviatilis, from this species in the Karkheh River, Iran. Mokhayer (1981c) reports a high infestation rate with larvae of the nematode Contracaecum in Karun River fish. This has health consequences for humans if fish are eaten raw, or are inadequately salted, smoked or cooked. There has been no report yet of this parasite from humans in Iran.

Moghainemi and Abbasi (1992) record a wide range of parasites from this species in the Hawr al-Azim in Khuzestan. Mortazaei et al. (2000) report the worm Neoechinorhynchus tylosuri in 54.8% of fish examined from Khuzestan marshes. Farahnak (2000) and Farahnak et al. (2002) record Anisakidae from this species in Khuzestan. Barzegar et al. (2008) record the digenean eye parasites Diplostomum spathaceum and Tylodelphys clavata from this fish. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Caligus lacustris, Lamproglena sp. and Ergasilus sp. on this species.

Economic importance

It appears in stores as a regular food fish in Ahvaz, Khuzestan, and is an important food fish in southern Iraq (Al-Nasiri and Islam, 1978) with 212,850 kg marketed at Basrah from October 1975 to June 1977 (Sharma, 1980). It forms about 29% of the total annual catches of freshwater fishes in southern Iraq and about 7.4% of the total annual catch for the whole country despite its small individual size (Mhaisen and Yousif, 1989; Mhaisen and Al-Jaffery, 1989). Its caloric value is 162.42 Kcal/100g (Mahdi et al., 2005). Salim (1962) records this species jumping into boats in Iraq when attracted by lights. It is important on fish farms in Iraq where it competes for food with cultured carp (Al-Shamma'a and Jasim, 1993) and Mhaisen and Al-Jaffery (1989) believed it could be farmed successfully because of its high growth rate. Taken also in the subsistence fisheries of the Indus River in Pakistan.

van den Eelaart (1954) gave the fishing season for this species as December-January (peaking in January) in rivers, and December-March (January) for lakes and marshes in Iraq.

Balasem et al. (2000) showed this species to be more sensitive to pollution than other Iraqi species tested, namely Barbus (= Carasobarbus) luteus, Cyprinus carpio, Carassius carassius (sic, probably C. auratus) and Gambusia affinis (sic, probably G. holbrooki). The LC50 of L. abu to copper after 72 hours exposure was 0.4 p.p.m. while the other species showed no mortality at this level. L. abu was also more sensitive to arsenic than these species and also Barbus (= Mesopotamichthys) sharpeyi, Barbus (= Luciobarbus) xanthopterus, Barbus (= Tor) grypus and Ctenopharyngodon idella (Ali et al., 1999). Wahab (1999) also tested the toxicity of four organophosphate insecticides to young fish, namely Dursban, Superacid, Malathion and Nogos with LC50 at 0.102. 0.125, 0.875 and 1.2 5 mg/L respectively. Salinity has a significant effect on the toxicity. This mullet has also been used to assess the impact of gas oil from oil spills in the Shatt al Arab on ionic regulation, on the chloride cells and on the gills (Faddagh et al., 2004). It was found to be sensitive to this form of pollution and suitable as an indicator species.

Conservation

Mhaisen and Yousif (1988) recommend banning fishing from mid-January to mid-May during the spawning season and instituting an increase in fishing net mesh size to 23.2 mm.

Further work

Variation in this species should be re-examined to determine if nominal species and subspecies are valid. Biology and stock assessments for Iranian waters need to be carried out.

Sources

Type material: Liza abu zarudnyi (ZISP 24336).

Iranian material: CMNFI 1979-0240, 3, 57.1-70.7 mm standard length, Fars, Lake Famur (ca. 29º31'N, ca. 51º50'E); CMNFI 1979-0304, 12, 21.5-50.0 mm standard length, Fars, Lake Famur (ca. 29º31'N, ca. 51º50'E); CMNFI 1979-0352, 5, 11.4-22.6 mm standard length, Khuzestan, marsh 57 km east of Abadan (30º33'30"N, 48º48'E); CMNFI 1979-0353, 3, 15.7-18.3 mm standard length, Khuzestan, Karun River (30º22'30"N, 48º17'E); CMNFI 1979-0354, 6, 18.4-25.2 mm standard length, Khuzestan, Karun River tributary (30º31'N, 48º19'E); CMNFI 1979-0355, 3, 16.7-29.4 mm standard length, Khuzestan, Karun River tributary (30º35'N, 48º22'E); CMNFI 1979-0360, 12, 17.7-35.8 mm standard length, Khuzestan, canal branch of Karkheh River (31º40'N, 48º35'E); CMNFI 1982-0369, 11, 39.3-57.9 mm standard length, Khuzestan, Karkheh River (no other locality data); CMNFI 1993-0149, 2, 106.0-142.2 mm standard length, Khuzestan, Karun River (no other locality data); ZSM 25387, 4, 56.6-90.9 mm standard length, Khuzestan, Karun River at Ahvaz (31º19'N, 48º42'E); ZSM 25700, 6, 41.0-57.4 mm standard length, Khuzestan, fish pond near Ahvaz (no other locality data); uncatalogued 1, 169.8 mm standard length, Khuzestan (no other locality data).

Comparative material: BM(NH) 1971.4.2:1, 1, 130.2 mm standard length, Iraq, pond 25 km from Mosul (no other locality data); BM(NH) 1973.5.21:179-184, 6, 41.4-51.4 mm standard length, Iraq, Tigris River at Jadriyah, Baghdad (no other locality data); BM(NH) 1974.2.22:10-13, 4, 107.3-119.5 mm standard length, Iraq, Najab Bazar (no other locality data).

Liza aurata
(Risso, 1810)

Common names

kefal-e auratus, mahi kefal-e talaee (= golden mullet fish), kefal-e zarin, kafal.

[gizili kefal in Azerbaijanian; gatykelle, singil, orsbalyk in Turkmenian; singil' in Russian; golden mullet, golden grey mullet, long-finned mullet].

Systematics

Mugil Auratus was originally described from Nice, France. No major relevant synonyms.

Key characters

This species is distinguished from its relative in the Caspian Sea by each head and antero-dorsal flank scale having only one pit or groove (occasionally double), 6-11 pyloric caeca of about equal length or gradually becoming longer from ventral to dorsal, scales on the snout ending anteriorly as a single row of small scales, and the oral edge of the preorbital bone is moderately concave. The upper jaw reaches posteriorly to a level with the posterior nostril, the only species in the genus with this character. Young have two vertical dark lines at the origin of the caudal fin rays and a herring bone pattern on the flanks (Harrison in Miller, 2003).

Morphology

The first dorsal fin has 4 strong spines, the last weakly developed compared to the first three. The second dorsal fin has 1-2 weak spines (more resembling unbranched rays than spines) and 6-9, usually 7 or 8 soft rays (the second unbranched ray or spine may be branched near the tip - the mode of 8 may include this ray in literature sources). The anal fin has 3 spines and 7-10, usually 9 branched soft rays. The pectoral fin has 2 unbranched rays and 13-17 branched rays and the pelvic fin has 1 spine and 5 branched rays. Scales number 40-49 in lateral series. There is no pectoral axillary scale. There is an elongate first dorsal fin axillary scale and a pelvic axillary scale. Scales have an almost vertical anterior margin with the anterior dorsal and ventral corners square to pointed. This margin is slightly indented where radii intersect it. The dorsal and ventral scale margins are parallel and the posterior margin is rounded. The focus is posterior and the posterior field has weak ctenii. Circuli are fine and numerous and radii are limited to the the anterior field, numbering up to 13 in fish examined here but probably size dependent. The adipose membrane of the eye is rudimentary. The gill rakers are fine and numerous, apparently finer and more numerous than in Liza saliens, exceeding 100 in the larger fish examined here. Raker number is probably size dependent. Rakers are slender, compressed and serrated medially. When appressed, a raker reaches the thirteenth raker below. The gut is elongate with several anterior and posterior loops after a muscular stomach. The chromosome number is 2n=48 (Klinkhardt et al., 1995).

Meristic values for Iranian specimens are:- first dorsal fin spines 4(30); second dorsal fin with 1(3) or 2(27) "spines" or unbranched rays and 6(1), 7(27), 8(1) or 9(1) branched rays; pectoral fin unbranched rays 2(30) and branched rays 15(7), 16(20) or 17(3); pelvic fin branched rays 5(30); lateral line scales 41(2), 42(4), 43(8), 44(3), 45(8), 46(4) or 47(1); and total vertebrae ?.

Sexual dimorphism

Unknown.

Colour

The body is a dusky grey to blue-grey with a silvery flank and belly. The back and upper flank have a series of dark to golden stripes. There is a golden blotch on the operculum. The peritoneum is dark brown to black. See also Key characters.

Size

Commercial catches in Iran during the 1950s weighed 0.3-0.8 kg, sometimes as much as 1.5-1.8 kg (Farid-Pak, no date). Reaches 59 cm and 2.5 kg on the coast of Turkmenistan. The Fisheries Research Centre in Mazandaran caught an exceptional specimen of this species weighing 1.5 kg and measuring 54 cm long (Iranian Fisheries and Research Training Organization Newsletter, 3, 1994).

Distribution

Found in the Mediterranean and Black seas and from the British Isles south to South Africa.

First reported from Iranian waters in 1933 by Shukolyukov (1937a) and by Dmitriev (1946), this species and Liza saliens became acclimated to the Caspian Sea over a period of 30-35 years and are regarded as naturalized from the latter half of the 1960s onward in Soviet waters (Marti, 1940; 1941; Khoroshko, 1982). It was first introduced to the Soviet Caspian Sea in 1930-1931 (Baltz, 1991). Kaplin in Mandych (1995) indicates that these grey mullets entered from the Black Sea during a Caspian transgression but this is incorrect.

In Iran, this species is found in the lower reaches of rivers along the Caspian coast, the Anzali Mordab and its outlets where numbers will probably increase with increasing salinity (Holčík and Oláh, 1992), the Safid River, Gorgan Bay, the southeast Caspian Sea, southwest Caspian Sea and south-central Caspian Sea (Abbasi et al., 1999; Kiabi et al., 1999; Abdoli and Naderi, 2009).

Zoogeography

This species is an exotic in Iran.

Habitat

This mullet inhabits the sea and enters the lower reaches of rivers, and is occasionally found in nearby lakes. It is recorded from depths of 5-700 m (Caspian Sea Biodiversity Database, www.caspianenvironment.org). Eggs develop in the open sea, larvae migrate to the coast and young feed along the shore and in bays. It winters along the Iranian shore in coastal waters, has a feeding migration in surface waters beginning in March to shallow areas of the middle Caspian and, by September-October, mature fish migrate over deeper water (300-700 m) of the middle and south Caspian to spawn. Feeding ceases at 6-8ºC and death occurs below 1.5ºC. The optimum temperature is 23-25ºC but young can be found in shallow water at 37.5ºC. It occurs from fresh water to salinities of 57‰ with mass mortalities recorded at 65‰ (Caspian Sea Biodiversity Database, www.caspianenvironment.org). Amini Ranjbar and Sotoudebnia (2005) assessed heavy metal (Zn, Cu, Pb and Cd) concentrations in fish from the Caspian Sea in Mazandaran and found higher than normal levels of Pb and Cd in muscle tissues.

Age and growth

Life span is up to 12 years in the Caspian Sea (Khoroshko, 1982). Maturity is attained at 38-45 cm and an age of 3-4 years for males and 5-6 years for females (RaLonde and Walczak, 1972). Fish taken from commercial catches in Iran are mostly 3-7 years old, 30.2-49.1 cm long and weigh 354-1266 g (Razivi et al., 1972). Fazli (1998) studied this species in the southeastern Caspian Sea of Iran and found that both scales and opercula could be used in ageing, although the  latter were better. Maximum growth occurred at 3 years of age. The b value in the relation between length and weight was 3.019 and sex ratio was 1:1.3 with females dominating. Ghaninejad et al. (2010) sampled fish from along the Iranian coast captured in 2007-2008. The overall male/female sex ratio at 1:1.22 differed significantly from 1:1 although only fish from Golestan were significantly different, size range was 19.0-50.2 cm, mean 32.7 cm, weight was 67 g to 1475 g, mean 411 g, age range was 2 to 10 years with mean age 4.42 years, age groups 3,4 and 5 years constituted 62% of the age composition, peak spawning was in October off Gilan but in November off Mazandaran and Golestan, average absolute fecundity was 700,881 eggs with a range of 200,112 to 2,282,862 eggs, and length of females at 50% maturity was estimated at 28.4 cm.

Food

This species is reported to enter the Anzali Mordab to feed from January to March (Holčík and Oláh, 1992), a result of the increasing salinity of this lagoon. Food items are small benthic invertebrates and detritus with some insects and plankton. Most stomach contents contain numerous sand grains. Adults scrape periphyton from rocks, silt and artificial structures. The predominant food items found by Ghadirnejad and Ryland (1996) from fish taken in the southern Caspian Sea were bivalves, foraminiferans and calanoid copoepods. Other food items were ostracods, eggs, nematodes, Nereis and cyclopoid copepods. Juveniles fed on ostracods and copepods while adults preferred bivalves, foraminferans and Nereis. This fish consumed more foraminferans than its relative and less copepods.

Reproduction

This mullet spawns in more northerly waters of the Caspian than its relative Liza saliens. Spawning begins in the middle of July in the middle Caspian and ends in the south Caspian from the middle to the end of October. Intensive spawning in the middle Caspian takes place in August-September at 20-22°C and a transparency of 7-11 m. Spawning depth is in the upper 0.5 m. The main spawning areas are at least 50-60 miles (80-96 km) from the coast (Avanesov, 1972) and this prevents the spawning area being easily exploited. Fecundity exceeds 4.4 million eggs in the Caspian Sea (Khoroshko, 1982); in Mazandaran average absolute fecundity reaches 772,350 eggs (Abdoli et al., 1996). The eggs are pelagic and have a diameter of 0.9 mm. In Iran, Gorgan Bay is believed to be a very important nursery and this is the major species caught by fishermen in winter (www.ramsar.org/ram_rpt_37e.htm, downloaded 28 July 2000). Shabanipour (1995) and Shabanipour and Heidari (2004) carried out histomorphological studies on the ovary of this species in Iran. Fazli (1998) found populations in the southeastern Caspian Sea of Iran spawned in September-November based on the gonadsomatic index, with most spawning in October. Males are ready to reproduce earlier than females. Absolute fecundity was 586,165 eggs. Fazli et al. (2008), again examining populations in the southeastern Caspian Sea, report a reproductive season of October to December, a male:female ratio of 1:1.42, sexual maturity at 26.0 cm fork length, mature gonads in 20% of fish at age 3, 63% at age 4, 88% at age 5 and 97% at age 6, individual absolute fecundity from 113,386 to 1.47 million eggs with a mean of 451,963. They concluded that the overlap between the fishing and spawning seasons required a delay of one month in the former to protect stocks. Heidari et al. (2009) studied late oogenesis in fish from Anzali. Recently spawned oocytes were lemon in colour and up to 0.8 mm in diameter.

Parasites and predators

Mokhayer (1976b) records the trematode Saccocoelium obesum from this species on the Iranian coast. This is a parasite of the intestine, with contamination rates of 90.4%, 88.6%, 81% and 34% for autumn, winter, spring and summer respectively. Gills carry Microcotyle mugilis at a rate of 2.0-2.3% and fish are also infected with Ancylodiscoides (sic) at a rate of 19.6%, 43%, 72% and 43.3% for autumn, winter, spring and summer respectively (Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 27-28, 1996; Annual Report 1993-94, Iranian Fisheries Research and Training Organization, Tehran, p. 49-50, 1995). Naem et al. (2002) found species in the monogenean treamtode genus Ligophorus on the gills of this species from the western branch of the Safid River.

This species is eaten by Sander spp., Silurus glanis and Huso huso.

At Ziba-kenar beach, Gilan in February 2004, this species showed erratic swimming and belly-up posture. A post-mortem showed gas accumulation and distension in the swimbladder, a yellowish liver, liquefaction of the gall bladder, excess micro-sand in the the caecum and hyperaemia of the intestine. The cause is thought to be an infectious agent closely related to a betanovirus or nodavirus (Iranian Fisheries Research Organization Newsletter, 38:3, 2004; Zorriehzahra et al., 2005).

Economic importance

The catch in Iran for this species and Liza saliens from 1956/1957 to 1961/1962 varied from 166,197 kg to 960,282 kg (Vladykov, 1964) and from 1965/66 to 1968/69 varied from 490 to 1916 tonnes (Andersskog, 1970). Vladykov (1964) and RaLonde and Walczak (1972) cite figures for the Bandar-e Anzali region for Liza aurata from 1937/38 to 1961/62 of 14 kg to about 113 t and for 1969/70 a catch of 1,085 t was recorded, while for the whole Northern Shilot (Fisheries Company) in the period 1965/66 to 1968/69 catches varied between 548 t and 1916 t. Holčík and Oláh (1992) report a catch of only 3 kg in the Anzali Mordab in 1990. The Food and Agriculture Organization, Rome records the following catches for the 6 years from 1980 to 1985 for mullets in Iran, presumably from the Caspian Sea:- 150, 400, 1500, 2733, 2135, and 2200 t respectively. Moghim et al. (1994) give a total biomass of 2400 tonnes for this species in the southern coastal area of the Caspian Sea with a maximum sustainable yield of 960 tonnes (figures for Liza saliens are 7000 tonnes and 2900 tonnes respectively). Caspian Sea catches of this species in Iran for the year 1993-1994 was 22% of the bony fish catch with Liza saliens comprising only 4% (the major part of the catch was safid mahi, Rutilus frisii, at 62%)(Annual Report 1993-94, Iranian Fisheries Research and Training Organization, Tehran, p. 83, 1995). Catches in former Soviet waters of the Caspian Sea are about three-quarters or more this species and one-quarter Liza saliens (Khoroshko, 1982). The main season for fishing in the Caspian Sea off Iran is from December to February, peaking in January (Farid-Pak, no date). Abdoli et al. (1996) note that beach seiner cooperatives started fishing a month earlier than usual in 1993, coinciding with the spawning season in October. Annual catch variations occur in Iranian waters, this species having an abundance of 86.5% in 1994 decreasing to 38% in 1995 (Iranian Fisheries Research and Training Organization Newsletter, 9:5, 1995). The catch of this species declined from 76.9% to 41.2% of the total mullet catch from 1993-1994 to 1994-1995 through overfishing of juveniles (Iranian Fisheries Research and Training Organization Newsletter, 11:7, 1996). Abdolmalaki (2001) records a catch per unit effort for beach seining as 93 kg in 1991, falling to 33 kg in 1997 through overfishing. The mean length of the catch in Guilan decreased from 1991 to 1997 while in Mazandaran it increased. Fazli and Ghaninejad (2004) give the year 2001 as that of the maximum mullet catch in their survey of the years 1993-2001, the catch increasing through this period. Catch per unit effort in each seine declined from 114 kg in 1993 to 43 kg in 1996, increasing to 66.4 kg and 78.4 kg in 2000 and 2001. For the years 2000 and 2001, L. aurata had a mean fork length of 32.7 and 32.3 cm, a weight of 418.3 and 419.8 g, and a K value of 0.61 and 0.93. They concluded that the harvest of this species is at a sustainable level. Heydatifard et al. (2002) state that its catch is greater than that of other bony fishes (presumably excluding kilka) in the southern Caspian Sea. S. M. E. J. Zorriehzahra (pers comm., 19 January 2011) noted a decline in catches from 6442 mt in 2002 to 2400 mt in 2010.

Hosseini et al. (2004) determined the shelf life of this species on ice in Iran was 10 days. Mazandaran fish have an average fat composition of 9.25%, 72% being unsaturated. Omega-3 fatty acids constitute 18.7% of the total fatty acids with a significant ratio to omega-6 fatty acids of 1.97 (Bedayatifard et al., 2002). Akhondzadeh Basteh et al. (2006) found the bacterial pathogens Listeria monocytogenes and Staphylococcus aureus in salted Liza aurata and Vibrio haemolyticus in fresh and salted L. auratus. Hedayatifard and Yousefian (2009; 2010) determined the comparative fatty acid composition in filleted fish that were fresh or dry-salted and found this fish to be one of the best sources for omega-3 essential fatty acids compared to other fishes in the Caspian Sea basin. Razavilar and Tavakoli (2006) studied the the prevalence of human toxigenic Clostridium botulinum and the need for food safety control measures.

Conservation

The successful introduction of this species and Liza saliens in the Caspian Sea has apparently led to a decline in various clupeid species (Alosa) and of Sander marinum (Baltz, 1991). Beach seines with mesh sizes of 28 mm, 33 mm and 36 mm caught non-standard fish as 43.3%, 35% and 2.2% of the total catch respectively. Seines with a cod end of at least 36 mm are recommended for stock protection (Iranian Fisheries Research and Training Organization Newsletter, 6:8, 1994) (note the report names the fish as Mugil cephalus but the catch was probably a mix of Liza aurata and L. saliens). Contamination of this species in Iranian waters with Pb, Ni and Zn was examined by Fazeli et al. (2005). The highest contamination was in the southwest part of the Caspian followed by the south centre and the southeast. The highest concentrations observed were 17.51 mg/kg for lead, 6.23 mg/kg for nickel and 647.28 mg/kg for zinc. Pazooki et al. (2009) found the heavy metals cadmium and chromium in fish from Bandar Anzali, more in the skin than in muscle and more in females than males, but not enough to constitute a health hazard.

Further work

As an important fisheries species, population numbers and biology should be carefully monitored on an ongoing basis.

Sources

Distinguishing characters based on Trewavas and Ingham (1972); Sakri (1993) gives an account in Farsi on the spawning and catch of mullets in the Iranian Caspian.

Type material: None.

Iranian material: CMNFI 1970-0509, 1, 124.2 mm standard length, Gilan, Safid River (37º24'N, 49º58'E); CMNFI 1970-0535, 1, 137.2 mm standard length, Gilan, Shafa River (37º35'N, 49º09'E); CMNFI 1970-0543, 3, 33.5-38.5 mm standard length, Gilan, Caspian Sea at Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1970-0543A, 4, 33.1-60.5 mm standard length, Gilan, Caspian Sea at Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1970-0563, 19, 52.8-87.0 mm standard length, Gilan, Caspian Sea, Kazian Beach (ca. 37º29'N, ca. 49º29'E); CMNFI 1970-0565, 4, 52.2-93.0 mm standard length, Gilan, Safid River estuary (ca. 37º28'N, ca. 49º54'E); CMNFI 1970-0581, 2, 147.7-151.5 mm standard length, Gilan, Caspian Sea near Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1970-0587, 1, 78.9 mm standard length, Mazandaran, Babol River (36º43'N, 53º39'E); CMNFI 1971-0326A, 20, 30.0-46.0 mm standard length, Gilan, Caspian Sea near Bandar-e Anzali (37º28'N, 49º27'E); CMNFI 1979-0430, 1, 42.0 mm standard length, Mazandaran, river 1 km east of Now Shahr (36º39'N, 51º31'E); CMNFI 1979-0470, 19, 29.4-57.4 mm standard length, Mazandaran, stream 21 km west of Alamdeh (36º35'N, 51º43'E); CMNFI 1979-0471, 3, 56.7-65.8 mm standard length, Mazandaran, Caspian Sea 14 km west of Alamdeh (36º35'N, 51º48'E); CMNFI 1979-0686, 2, 59.1-87.3 mm standard length, Gilan, Safid River near Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1980-0117, 2, 99.7-114.5 mm standard length, Gilan, Golshan River (37º26'N, 49º40'E); CMNFI 1980-0127, 6, 29.5-36.9 mm standard length, Gilan, Caspian Sea near Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1980-0146, 1, 135.7 mm standard length, Mazandaran, Gorgan Mordab at Ashuradeh-ye Kuchak(36º50'N, 53º56'E); CMNFI 1980-0149, 1, 38.4 mm standard length, Gilan, Chabak River (37º21'N, 49º50'E).

Liza saliens
(Risso, 1810)

Common names

kefal-e saliens, kafal-e poozehbarik, kafal.

[sivriburun kefal in Azerbaijanian; vitibutum or gatykelle in Turkmenian; ostronosik in Russian; sharpnose mullet, small mullet, grey mullet, leaping mullet].

Systematics

Mugil Saliens was originally described from Nice, France. No major relevant synonyms.

Key characters

This species is distinguished from its relative in the Caspian Sea by each head and antero-dorsal flank scale having 2-7 or more grooves (mostly 2 grooves), pyloric caeca in two groups, 3-5 short and 3-4 long (total 6-9), scales on the snout ending anteriorly as numerous rows of small scales, and the oral edge of the preorbital bone is deeply notched. Young lack the vertical dark lines at the origin of the caudal fin rays and the herring bone flank pattern seen in L. aurata (Harrison in Miller, 2003).

Morphology

The first dorsal fin has 4 spines, the last one weakly developed compared to the first three. The second dorsal fin has 1-3, usually 1or 2, spines (more resembling unbranched rays than true spines) and 6-9, usually 8 or 9 soft rays (the second unbranched ray or spine may be branched near the tip - modes of 8 or 9 in literature may include this ray). The anal fin has 3-4, usually 3, spines and 7-10, usually 9, soft rays. The pectoral fin has 2 unbranched rays and 14-17 branched rays and the pelvic fin has 1 spine and 5 branched rays. Scales number 42-50 in lateral series. There is no pectoral axillary scale. There is an elongate first dorsal fin axillary scale and a pelvic axillary scale. Scales have an almost vertical anterior margin with the anterior dorsal and ventral corners square to pointed. This margin is slightly indented where radii intersect it. The dorsal and ventral scale margins are parallel and the posterior margin is rounded. The focus is posterior and the posterior field has weak ctenii. Circuli are fine and numerous and radii are limited to the the anterior field, numbering up to 10 in fish examined here but probably size dependent. The gill rakers are fine and numerous, but not as fine and numerous as in Liza aurata, exceeding 70 in the larger fish examined here, and probably size dependent. When appressed, a raker reaches raker 6 or 7 below. Rakers have a serrated medial edge. Vertebrae 23-25. The gut is elongate with 2 posterior loops after a muscular stomach. The chromosome number is 2n=48 (Klinkhardt et al., 1995).

Meristic values for Iranian specimens are:- first dorsal fin spines 4(30); second dorsal fin with 1(6), 2(22) or 3(2) "spines" or unbranched rays and 6(1), 7(20) or 8(9) branched rays; pectoral fin unbranched rays 2(30) and branched rays 14(2), 15(27) or 16(1); pelvic fin branched rays 5(30); lateral line scales 42(2), 44(6), 46(6), 47(8), 48(4) or 49(4); and total vertebrae ?.

Sexual dimorphism

Males are smaller than females.

Colour

The back is dark grey to blue-grey or grey-brown, the belly pale to silvery or yellowish, and the silvery-grey flanks have about 7 bluish stripes. There is a golden blotch on the operculum. The peritoneum is brown to black.

Size

Commercial catches in Iran during the 1950s weighed 0.3-0.8 kg, sometimes as much as 1.5-1.8 kg (Farid-Pak, no date). Reaches 40.0 cm.

Distribution

Found in the Mediterranean and Black seas and along the Atlantic coast to South Africa.

First reported from Iranian waters by Dmitriev (1946), this species was introduced to the Soviet Caspian Sea in 1930-1931 (Baltz, 1991). In Iran it is reported from the lower reaches of such rivers as the Gorgan, Gharasu, Tajan, Talar, Babol, Haraz, Sardab, Pol-e Rud, and Safid, the Anzali Mordab and Gorgan Bay, the southeast Caspian Sea, southwest Caspian Sea and south-central Caspian Sea (Roshan tabari, 1987; Abbasi et al., 1999; Kiabi et al., 1999; Abdoli and Naderi, 2009).

Zoogeography

This species is an exotic in Iran.

Habitat

This is a pelagic species found mostly in inshore areas and entering lagoons and rivers. It is found in surface waters over depths of 5-700 m at salinities of 4-13‰. It migrates to the southern Caspian in autumn when water temperatures fall (Caspian Sea Biodiversity Database, www.caspianenvironment.org). Generally it tolerates fluctuations of 5-27Cº and 11-28‰ although fry need time to adjust to lower salinities (Harrison in Miller, 2003).

Age and growth

Life span is up to 10 years in the Caspian Sea (Khoroshko, 1982). Males mature at 3 years and females at 4 years (Caspian Sea Biodiversity Database, www.caspianenvironment.org). The maximum growth rate in fish sampled between Babolsar and Bandar-e Torkeman in the Iranian Caspian was observed in fish age 3 years. The operculum gave better ages than scales (Fazli, 2000). Patimar (2008a) examined fish in Gorgan Bay and found maximum ages of 6+ for males and 7+ for females. Both sexes grew allometrically (b = 2.478 for males, 2.545 for females) and rapidly during their first year of life, reaching more than 40% of their growth. Age and growth were in contrast to other populations in the southeast Caspian Sea. The von Bertalanffy growth functions were Lt = 30.415(1-e-0.275(t+0.645)) for males and Lt = 34.832(1-e-0.211(t+1.009)) for females. Males predominated at smaller sizes and females at larger sizes but see below.

Food

Diet is very similar to that of Liza aurata, comprising detritus, periphyton and small benthic organisms Stomach contents contain sand grains in Iranian fish. The predominant food items found by Ghadirnejad and Ryland (1996) from fish taken in the southern Caspian Sea were bivalves, foraminiferans and calanoid copoepods. Other food items were ostracods, eggs, nematodes, Nereis and cyclopoid copepods. Juveniles fed on ostracods and copepods while adults preferred bivalves, foraminferans and Nereis.

Reproduction

Spawning starts at the end of May and the beginning of June in the southern Caspian Sea and continues until the end of September and the beginning of October, most intensively between June and August off the Turkmenistan coast according to Avanesov (1972). Iranian fish spawned in July-August and males ripen earlier than females (Fazli, 2000; Yousefian et al., 2003). Yousefian et al. (2003) give details of oogenesis, detailing the stages in gonad development. Spawning occurs over depth ranges of 5-700 m but most spawning in the southeastern Caspian is at 5-7 m. Temperatures are between 17 and 29°C with the most intensive spawning when sea surface temperature is 25-29°C. Water transparency is 1-9 m. The greatest aggregations of eggs are found 5-7 miles (8-11 km) from shore in the eastern Caspian. Fecundity exceeds 2.1 million eggs in the Caspian Sea (Khoroshko, 1982). Egg diameter is up to 0.8 mm. Females predominate in Iranian samples examined by Fazli (2000) at a ratio of 1:3.14. Patimar's (2008a) Gorgan Bay fish reproduced in May-July with an average fecundity reaching 389,790 eggs in 7+ fish and relative fecundity 4281 eggs/g. The low absolute fecundity contrasted with other populations in the southeast Caspian Sea.

Parasites and predators

This species has a worm parasite in the intestine, Saccocoelium obesum, with contamination rates of 62%, 10.7%, 12% and 22.2% for autumn, winter, spring and summer respectively, lower than in Liza aurata. Gills carry Microcotyle mugilis at a rate of 2.0-2.3% and fish are also infected with Ancylodiscoides (sic) at a rate of none, none, 1.6% and 4% for autumn, winter, spring and summer respectively, much lower than in Liza aurata (Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 27-28, 1996; Annual Report 1993-94, Iranian Fisheries Research and Training Organization, Tehran, p. 49-50, 1995).

Economic importance

The main season for fishing in the Caspian Sea off Iran is from December to February, peaking in January (Farid-Pak, no date). Some catch information is summarised above under Liza aurata which is 5-6 times more abundant in catches. However a report on the 1994-1995 catch showed a severe reduction of 2500 tonnes, as Liza saliens replaced L. aurata as the main part of the catch. L. aurata declined from 76.9% of the mullet catch in 1993-1994 to 41.2% in 1994-1995, the cause being overfishing of juveniles (Iranian Fisheries Research and Training Organization Newsletter, 11:7, 1996). Moghim et al. (1994) give a total biomass of 7000 t for this species in the southern coastal area of the Caspian Sea with a maximum sustainable yield of 2900 t (figures for Liza aurata are 2400 tonnes and 960 tonnes respectively). Annual catch variations occur in Iranian waters, this species having an abundance of 13.5% in 1994 increasing to 62% in 1995 (Iranian Fisheries Research and Training Organization Newsletter, 9:5, 1995). Abdolmalaki (2001) records a catch per unit effort for beach seining as 78 kg in 1991, falling to 15 kg in 1997 through overfishing. The mean length of the catch in Gilan and Mazandaran did not decrease from 1991 to 1997, the former in contrast to Liza aurata. Fazli and Ghaninejad (2004) give the year 2001 as that of the maximum mullet catch in their survey of the years 1993-2001, the catch increasing through this period. Catch per unit effort in each seine declined from 114 kg in 1993 to 43 kg in 1996, increasing to 66.4 kg and 78.4 kg in 2000 and 2001. For the years 2000 and 2001, L. saliens had a mean fork length of 27.5 and 25.1 cm, a weight of 224.7 and 179.1 g, and a K value of 0.91 and 0.71.

Conservation

See above under Liza aurata. Mercury levels at 0.0102-0.108 mg kg -1 w.w. in this species from the Iranian shore were lower than the WHO guideline of 0.5 mg kg-1 w.w. (Agah et al., 2007).

Further work

As an important fisheries species, population numbers and biology should be carefully monitored on an ongoing basis.

Sources

Distinguishing characters based on Trewavas and Ingham (1972).

Type material: None.

Iranian material: CMNFI 1970-0510, 34, 38.1-72.8 mm standard length, Gilan, Golshan River (37º26'N, 49º40'E); CMNFI 1970-0507, 9, 23.6-46.8 mm standard length, Gilan, Caspian Sea at Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1970-0508, 1, 50.6 mm standard length, Gilan, Safid River at Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1970-0509, 4, 31.9-66.8 mm standard length, Gilan, Safid River at Hasan Kiadeh(37º24'N, 49º58'E); CMNFI 1970-0528, 16, 43.6-64.4 mm standard length, Mazandaran, Tajan River estuary (36º49'N, 53º06'30"E); CMNFI 1970-0535, 1, 153.3 mm standard length, Gilan, Shafa River (37º35'N, 49º09'E); CMNFI 1970-0539, 43, 25.1-102.0 mm standard length, Gilan, Caspian Sea off Bandar-e Anzali (37º28'N, 49º27'E); CMNFI 1970-0543A, 2, 76.7-79.9 mm standard length, Gilan, Caspian Sea at Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1970-0549, 1, 48.1 mm standard length, Gilan, Qareh Su near Alb Imamzadeh (no other locality data); CMNFI 1970-0565, 3, 91.8-108.8 mm standard length, Gilan, Safid River estuary (ca. 37º28'N, ca. 49º54'E); CMNFI 1970-0577, 13, 22.1-38.8 mm standard length, Gilan, Caspian Sea near Astara (ca. 38º26'N, ca. 48º53'E); CMNFI 1970-0581, 4, 41.0-71.0 mm standard length, Gilan, Caspian Sea near Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1970-0586, 8, 27.3-50.4 mm standard length, Mazandaran, Gorgan Mordab at Ashuradeh-ye Kuchak(36º50'N, 53º56'E); CMNFI 1970-0587, 165, 15.7-47.9 mm standard length, Mazandaran, Babol River at Babol Sar (36º43'N, 52º39'E); CMNFI 1970-0590, 11, 28.6-54.0 mm standard length, Mazandaran, Shesh Deh River near Babol Sar (ca. 36º43'N, ca.52º39'E; CMNFI 1979-0470, 17, 16.6-43.4 mm standard length, Mazandaran, stream 21 km west of Alamdeh (36º35'N, 51º43'E); CMNFI 1979-0471, 1, 59.1 mm standard length, Mazandaran, Caspian Sea 14 km west of Alamdeh (36º35'N, 51º48'E); CMNFI 1979-0477, 1, 16.7 mm standard length, Mazandaran, Gorgan Mordab at Bandar-e Shah (36º54'N, 54º02'30"E); CMNFI 1979-0788, 1, 22.7 mm standard length, Mazandaran, Gorgan River at Khadje Nafas (37º00'N, 54º07'E); CMNFI 1980-0120, 9, 48.8-73.2 mm standard length, Mazandaran, Babol River at Babol Sar (36º43'N, 52º39'E); CMNFI 1980-0126, 2, 217.5-244.2 mm standard length, Gilan, Caspian Sea near Bandar-e Anzali (37º28'N, 49º27'E); CMNFI 1980-0127, 5, 22.8-52.8 mm standard length, Gilan, Caspian Sea near Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1980-0129, 18, 28.3-64.2 mm standard length, Mazandaran, Tajan River estuary (36º49'N, 53º06'30"E); CMNFI 1980-0136, 13, 35.9-76.8  mm standard length, Mazandaran, Fereydun Kenar River estuary (36º41'N, 52º29'E); CMNFI 1980-0146, 2, 88.8-96.6 mm standard length, Mazandaran, Gorgan Mordab at Ashuradeh-ye Kuchak(36º50'N, 53º56'E); CMNFI 1980-0149, 10, 36.0-83.2 mm standard length, Gilan, Chabak River (37º21'N, 49º50'E); CMNFI 1980-0157, 77, 25.7-50.1 mm standard length, Mazandaran, Gorgan River estuary (36º59'N, 53º59'30"E); CMNFI 1993-0144, 2, 78.4-92.2 mm standard length, Mazandaran, Neka Power Plant (36º51'48'N, 53º23'24"E).

Liza subviridis
(Valenciennes, 1836)

Common names

biah or biah arabi at Bushehr, garriz at Bandar Abbas; meid or maid (for small specimens); kefal posht sabz.

[biag, meid or maid, biah akhtar, beyah akhtar in Arabic; greenback mullet, greenback grey mullet, silver mullet].

Systematics

Originally described from Malabar, India. Synonyms are Mugil dussumieri Valenciennes in Cuvier and Valenciennes, 1836 and Mugil jerdoni Day, 1876. Placed in the genus Chelon Rose, 1793 in Randall (1995b).

Key characters

The preorbital bone does not occupy the whole space between the eye and upper lip as in all other mullets.

Morphology

The head is wide and flattened with head length 23-27% of standard length. There is a well-developed adipose eyelid, covering about half of the iris, but this is absent in young fish. The lower lip has a weak to marked symphysial knob (authors differ) and both lips are thin. The upper jaw extends back on a level with the anterior nostril to the anterior eye rim. Upper jaw teeth are in several rows and ciliiform lower jaw teeth in one row. The preorbital bone is strongly notched anteriorly and only occupies three-quarters of the space between the eye and upper lip. Scales are regularly arranged and are dense on the second dorsal and anal fins. The second dorsal fin origin lies over the anterior half of the anal fin base. The pectoral axillary scale is rudimentary or absent. The pectoral fin does not reach back on a level with the first dorsal fin origin and is 74-76% of head length.

The first dorsal fin has 4 spines, the second dorsal fin has 1 spine and 8-9, usually 8, soft rays and the anal fin has 3 spines and 8-9, usually 9, soft rays. Scales in lateral series 27-32. Transverse scales 10-11, usually 11. Pyloric caeca 4-5.

Ahmed and Hussain (2003) describe the larvae of this species from Khawr az Zubayr and the Shatt al Arab estuary. Al-Hassan and Madhi (1989) studied enzyme polymorphisms in populations from the Shatt al Arab, Khawr az Zubayr and Kuwait but found no evidence of sub-population differentiation. A single stock is found in the northern Persian Gulf.

Sexual Dimorphism

Unknown.

Colour

The back is dark to light green or greyish-green, the flanks silvery to white and the belly white. There may be 3-7 blackish stripes along the flank but these may not always be obvious. The caudal fin is bluish and has a black margin, other fins are hyaline.

Size

Attains 39.5 cm total length and 270 g (Al-Hassan and Al-Seyab, 1992).

Distribution

Found in the Persian Gulf and eastwards to India, China, northern Australia and Polynesia. Reported from the Shatt al Arab and Hawr al Hammar, Iraq near Iranian Khuzestan by Al-Hassan et al. (1989), Al-Hassan and Al-Seyab (1992) and Hussain et al. (1997; 2001). Records of Liza dussumieri from the Tigris River at Amara and Baghdad, the Euphrates River and the Shatt al Arab by Kennedy (1937), Khalaf (1961), Mahdi (1962), Al-Nasiri and Hoda (1976) and Hussain et al. (1989), if correctly identified, are referred to this species (Thomson and Luther in Fischer and Bianchi, 1984).

Recorded from the lower Mand River in Bushehr Province at 28°14'N, 51°51'E, 28°11'N, 51°32'E and 28°07'N, 51°24'E by M. Rabbaniha (pers. comm., 1995) and from Khuzestan in the Zohreh and Bahmanshir rivers and the Hawr al Azim by Najafpour (1997). Records from Iran could be L. vaigiensis (q.v.). No specimens have been examined by me to confirm these anecdotal and literature records.

Zoogeography

A widely distributed marine species, this mullet also enters and is resident in fresh waters but is dependent on reproduction of marine populations to maintain the freshwater ones.

Habitat

This species lives in the sea, particularly in coastal waters, lagoons and estuaries, but regularly enters fresh water. At a freshwater station on the Shatt al Basrah Canal, Iraq with salinities up to 3.5‰, Al-Daham and Yousif (1990) found this species to be dominant, comprising 59.6% by number and 40.0% by weight. Eggs of this species and L. klunzingeri were most abundant in the inner part of the Shatt al Arab estuary in February with larvae most abundant in February and April. The temperature range for several sites was 12.5-22ºC and salinity 32-38‰. Temperature is the most important factor influencing spawning (Ahmed and Hussain, 2000). This was one of the more abundant species in the recovering Hawr al Hammar of southern Iraq in 2005-2006 and in the marshes in the 1980's (Hussain and Ali, 2006).

Age and Growth

In the Shatt al Basrah Canal of southern Iraq, populations of this species are comprised of age groups 1 to 6 but most fish are in age groups 1 and 2 (Al-Daham and Wahab, 1991). Age 2 dominates from February to April and age 1 from June to January. Females are slightly longer in each age group and have a longer life span than males. The oldest females are age 6 and 303 mm and the oldest males age 4 and 251 mm. Males are less numerous than females with a sex ratio 1.0:1.4. Maturity is attained at age 1 and the smallest mature male is 137 mm and the smallest mature female is 142 mm. Ali et al. (1999) examined the stock of this species in Khawr Abdullah and found the infinity length (L) to be 34.3 cm total length (TL), the growth coefficient (K) 0.31, annual total mortality (Z) 1.06, natural mortality (M) 0.70 and fishing mortality (F) 0.366. The exploitation rate (E) was 0.34 indicating that the stock was slightly exploited during September 1989-August 1990. The length-weight relationship was W = 0.0000208 L2.985. Mohamed et al. (1998) found, for fish from the northwest Arabian (= Persian) Gulf, a length-weight relationship of W = 1.34 x 10-5 L2.961, condition factor ranged from 0.98 to 1.21 and the growth model was Lt = 308 (1-e-0.225 (t + 0.618)).

Ali et al. (1999) summarise annual growth of this species from their own and other studies as follows, attributing differences to differing methodologies:-

Author

sex

L

K

1

2

3

4

5

6

Wahab (1986) (Shatt al Basrah)


29.4
40.2

0.39
0.151

14.6
15.5

19.3
20.5

22.1
23.0

24.0
25.2

-
27.3

-
29.0

Al-Hisnawi (1990) (Khawr az Zubayr)


25.2
48.9

0.474
0.141

11.4
11.35

16.55
16.78

19.84
20.92

21.85
24.31

-
27.82

-
30.5

Mohammed et al. (1998) (Northwest Arabian Gulf)

♂ + ♀

30.8

0.225

9.4

13.6

17.0

19.7

21.9

23.7

Ali et al. (1999) (Khawr Abdullah)

♂ + ♀

34.3

0.31

8.9

15.5

20.37

23.97

26.65

28.64

Muhsin (1988) determined protein, lipid and ash content of fish taken monthly from the marine Khawr az Zubayr and found fluctuations to follow the reproductive cycle. Lipid content of the liver increased in winter and of the carcass in winter and spring, the pre-spawning period, along with increases in the gonadosomatic and hepatosomatic indices. The latter indices peaked in March and decreased sharply until June. Total protein decreased during winter and spring from the  carcass and in spring from liver and ovaries as they were used for energy and to support maturation of the ovaries.

In Malaysia, maturity is attained at 95-115 mm for males and 105-115 mm for females (Chan and Chua, 1980).

Food

Food items are algae, diatoms and detritus extracted from ingested mud and sand. Al-Hassan and Madhi (1987) include higher plants as part of the diet and Al-Hassan and Al-Seyab (1992) copepods for Iraqi waters. Mohamed et al. (1998) found this species to be a detritivore in the northwest Persian Gulf. Hussain and Ali (2006) examined feeding relationships among fishes in the Hawr al Hammar, Iraq and found this species to be a herbivore, 75.2% of the diet being plants and algae. In another study of the recovering Hawr al  Hammar, diet was 80.84% algae and 15.66% diatoms with amounts of crustaceans and plants being less than 10% each (Hussain et al., 2006).

In Malaysian estuaries and coastal areas, Chan and Chua (1979) working in Malaysia found this species to feed only on zooplankton when less than 12 mm standard length, becoming bottom feeding at 16-20 mm on zooplankton, diatoms and detritus and by 24 mm zooplankton was absent but filamentous algae was added where available. The proportion of the various food items varied with the locality.

Reproduction

Spawning occurs in the sea, with all eggs released at once in a Malaysian population (Chan and Chua, 1980). Al-Daham and Wahab (1991) were unable to locate eggs and larvae in fresh waters of Iraq. They believe that adults leave the Shatt al Basrah Canal to spawn in the sea. However, Al-Hassan and Madhi (1987) give a reproductive season of February through April in the Shatt al Arab, Iraq and Ahmed and Hussain (2003) give March to June in the offshore area near the mouth of the Khawr az Zubayr and Shatt al Arab. Muhsin (1988) determined April-June to be the spawning season, with mature ova released in May, in the Khawr az Zubayr (see above). Male and female gonads are best developed in February to March and spawning is deduced to have occurred from March to May, possibly offshore in the Persian Gulf. In Malaysia, spawning can occur from June to November and in the off-season (Chan and Chua, 1980). In Iraq, fry 27-40 mm in length are captured from April to June in fresh water. The adult mullet return to fresh water after spawning. Fecundity was 133,224-295,065 eggs for fish 182-243 mm total length, this being higher than in Malaysia at 40,000-145,000. Fecundity is somewhat lower than in other mullets but may be attributed to year round spawning at some localities (Chen and Chua, 1980). Al-Hassan and Madhi (1987) give a fecundity of 549,278 eggs with egg diameter up to 0.65 mm and Al-Hassan and Al-Seyab (1992) give 580-590,000 eggs for their large specimen from Hawr al Hammar.

Parasites and predators

Unknown.

Economic importance

This species forms part of coastal fisheries in shallow waters and is caught with a variety of nets, particularly seines, stationary traps, gill nets and cast nets. It forms a substantial part of brackish water fisheries in southern Iraq, being the most abundant species in estuaries, and economically the most important mullet (Al-Daham and Wahab, 1991). Ali et al. (1999) gave a catch of 35 t in Khawr Abdullah, Iraq, almost 6% of the marine catch. Mohamed et al. (1998) gave a value of 3.4% of the commercial catch for this species with catch rates in the northwest Arabian (= Persian) Gulf of 0.15 kg/h in June to 0.75 kg/h in October. This species is potentially important in fish culture as it is herbivorous. Consequently ponds can be fertilised inexpensively and plant food supplements such as rice bran and peanut meal can be given. Chan and Chua (1979) recommend that fish for stocking should be at least 30 mm standard length as their bottom feeding habit is then most suitable for pond culture.

The proximate chemical composition measured in fish from the Basrah fish market was moisture 76.16%, protein 16.82%, fat 5.1% and ash 1.82% and the general condition was a maximum of 1.78 (Ali et al., 2004). It is characterised as a fatty fish according to a lipid content 9-14% by wet weight of muscle in autumn (Hantoush et al., 1999).

Conservation

Population numbers and trends are unknown so conservation requirements cannot be ascertained.

Further work

Materials of freshwater mullets should be collected in Iranian waters and studied for their identity and biology.

Sources

Description based on literature sources, particularly Blegvad and Løppenthin (1944), Fischer and Bianchi (1984), Talwar and Kacker (1984) and Thomson (1997).

Liza vaigiensis
(Quoy and Gaimard, 1824)

Common names

[massiyat or biah in Iraq; square-tail mullet, diamond-scaled mullet, diamond-scale grey mullet, brown-banded mullet].

Systematics

Originally described from Waigeo, Indonesia. Placed in the genus Ellochelon Whitley, 1930 in Randall (1995b).

Key characters

This species is uniquely characterised in this area of the northen Indian Ocean by the caudal fin being only slightly emarginate. All other local mullets have an obviously forked tail fin.

Morphology

The preorbital bone is notched anteriorly and is wide, filling the space between the eye and the upper lip. The head is broad and flattened on top, with head length 24-30% of standard length. The eye has adipose tissue around its margin. Lips are thin but the lower lip has a high symphysial knob. There are ciliate teeth on the lips, in 1-2 rows on the upper lip and 1 row on the lower lip. Larger fish lose the upper lip teeth at >250 mm standard length and the lower lip teeth at ca. 100 mm standard length. About half of the anal fin base lies anterior to the level of the second dorsal fin origin. The pectoral axillary scale is rudimentary or absent. The pectoral fin does not reach back on a level with the first dorsal fin origin. The second dorsal and anal fins are densely scaled. Hoda (1978) describes the larva in Iraq.

First dorsal fin spines 4, second dorsal fin with 1 spine and 7-9 soft rays, anal fin with 3 spines and 8 soft rays, pectoral fin with 15-18 branched rays, lateral series scales 24-29, transverse scales (from first dorsal fin origin to pelvic fin origin) 9-11, and predorsal scales 15-16. Gill rakers 40-61.

Sexual Dimorphism

Unknown.

Colour

The back is olive brown to pale olive, the flanks silvery and the belly white with yellowish tinges. Upper flank scales have characteristic brown blotches which appear to run together to form about 5 stripes. The iris has yellow patches. Fin margins are dusky to black but the body of each fin is yellowish. Young fish have black pectoral fins but in adults the lower half is yellow. The caudal, anal and pelvic fins are a bright yellow in young fish while the second dorsal fin is dark on its distal half.

Size

Reaches 63 cm.

Distribution

This species is widespread in the tropical Indo-Pacific Ocean from Africa to Japan and Tahiti including the Persian Gulf. It also enters fresh waters. Reported from the Shatt al Arab, Iraq near Iranian Khuzestan by Al-Nasiri and Hoda (1976). A single specimen from Khuzestan, Iran confirms its presence in fresh water. Akbari (1998) records it from coastal and estuarine waters in Hormozgan. Other records from Iranian rivers draining to the Persian Gulf (e.g. Dez, Karun, Dalaki and Mand rivers) may be this species but could also be L. subviridis.

Zoogeography

A widely distributed marine species, this mullet also enters and is resident in fresh waters but is dependent on reproduction of marine populations to maintain the freshwater ones.

Habitat

This species is found in coastal waters over sand, in lagoons and over reef flats and commonly enters fresh waters. It apparently enters fresh water during the rainy season elsewhere in its range, ascending only to tidal limits.

Age and Growth

Unknown.

Food

Food items are algae, diatoms and detritus whih are ingested along with associated sand and mud.

Reproduction

Fry are known to occur along the sea coast.

Parasites and predators

None reported from Iran.

Economic importance

This mullet is common in commercial catches along the west coast of India. It is caught in castnets, stakenets and beach seines.

Conservation

Population numbers and trends are unknown so conservation requirements cannot be ascertained.

Further work

Materials of freshwater mullets should be collected in Iranian waters and studied for their identity and biology.

Sources

Description based on literature sources, particularly Fischer and Bianchi (1984), Talwar and Kacker (1984) and Thomson (1997).

A single specimen from Khuzestan, Iran was examined measuring ?.

Genus Mugil
Linnaeus, 1758

This genus is characterised by a strongly flattened head, a transverse mouth with the lateral cleft short, thin lips, lower jaw with a symphysial knob and lower jaws meeting at symphysis with an acute angle, the maxilla end is not visible when the mouth is closed, the anteroventral edge of the preorbital is serrate and straight and the ventral edge is slender and pointed, no true teeth in the jaws, a very well-developed adipose eyelid reaching to, or nearly to, the pupil, two pyloric caeca, pharyngobranchial organ with a single valve, no vomerine or palatine teeth, scales cycloid or feebly ctenoid, and pectoral axillary scale long and pointed. Thomson (1997) and Harrison in Miller (2003) give a wide range of generic characters but since only one species is found in Iranian waters, and that rarely, the species description suffices here.

Mugil cephalus
Linnaeus, 1758

Common names

kefal or kafal, biah-e sarpahn (= flathead mullet), كفال خاكستري (= kafal-e khakestari).

[biah, biyah, biaha Ramadiyah, anubah or asfatiya in Arabic; striped mullet, loban, common, grey, jumping, flathead grey, sea, or river mullet; mighach, minghaj or karul in Pakistan].

Systematics

Mugil Cephalus was originally described from the Atlantic Ocean at the shores of Europe, entering rivers. No major relevant synonyms.

Key characters

This species is distinguished from Liza species by having only 2 pyloric caeca, the posterior end of the maxilla is straight rather than curved down behind the premaxilla and visible behind the corner of the closed mouth, adipose tissue on the eye reaches the pupil in adults, and the tip of the jaw end is on the line of the gape.

Morphology

There is a prominent adipose eyelid which leaves only a narrow slit over the pupil and extends twice as far behind the eye as in front of it.

First dorsal fin with 4 spines, second dorsal fin with 1-2 spines and 6-9, usually 8or 9, soft rays, anal fin with 3 spines and 7-9, usually 8, soft rays (below 5 cm there are 2 spines and 9 soft rays), pectoral fin branched rays 14-17, and pelvic fin branched rays 5.

Lateral series scales 36-46 with 3-5 on the caudal fin, transverse scale rows 14-15, predorsal scales 24-26 and cheek scales in 3-4 rows. Scales on the head extend forward to the tip of the snout. Scales are found on the caudal fin rays and at the bases of the anterior dorsal and anal fin rays. The pectoral axillary scale is very long, about one-third the length of the fin, and there is a pelvic axillary scale. Scales bear an anterior mid-margin notch, the anterior corners are very square and sharp, upper and lower margins are parallel and the posterior margin is rounded. There are few radii (as few as 5) running from a subcentral posterior focus to the notch. Circuli are numerous and fine and the exposed scale surface is coarse. Gill rakers 24-80, long, fine and crowded, count increasing with growth of fish. Pyloric caeca number 2 and the gut is long with numerous loops, complexly coiled. The chromosome number is 2n=48 (Klinkhardt et al., 1995).

There is a distinct axillary scale at the pectoral fin, 33-36% of pectoral fin length. The pectoral fin reaches the tenth postopercular scale but not the level of the first dorsal fin origin; folded forward it reaches the eye but not the pupil.

A single Iranian specimen has 4 first dorsal fin spines, 8 soft second dorsal fin rays, 8 anal fin soft rays, 15 pectoral fin branched rays, 5 branched pelvic fin rays, 43 scales in lateral series, and 24 total vertebrae.

Sexual dimorphism

Females grow larger and weigh more than males.

Colour

Back bluish-grey to olive-green or greyish-bown, flanks silvery and belly silvery to white. Each scale has a dark spot which line up to give the impression of 6-12 indistinct brownish or grey stripes. The upper rear corner of the operculum has a golden spot. There is dark purple blotch at the pectoral fin base. The dorsal and pectoral fins are a dark blue-grey while the anal and caudal fins are a yellowish-green. The dorsal and caudal fins have dusky margins. Young are a bright, iridescent silver. Estuarine and freshwater fish may be duller overall than marine fish.

Size

Attains 1.2 m and 7 kg.

Distribution

Found world-wide between about 51°N and 42°S. Introduced into the Caspian Sea in 1902, it has not been observed there since and there appear to be no valid Caspian records for Iran (Dmitriev, 1946; Baltz, 1991). The introduction probably failed because the pelagic eggs were not buoyant at the low Caspian salinities (Baltz, 1991) although Harrison in Miller (2003) notes spawning has been induced in fresh water. However Kiabi et al. (1999) report it from the southeast Caspian Sea and the species is being cultured at Sari on the Iranian coast of the Caspian Sea (Iranian Fisheries Research Organization Newsletter, 40 & 41:3, 2004; and see below). Kolodar and Abdoli (2004) report it from the Gomishan Lagoon but only in farms there.

Reported from the Shatt al Arab, Iraq near Iranian Khuzestan by Al-Nasiri and Hoda (1975b). Akbari (2002) records this species from creeks and coastal waters of Hormozgan. It enters rivers of southern Iran from the Persian Gulf and Sea of Oman, notably the Sarbaz River.

Zoogeography

A widely distributed marine species, this mullet also enters and is resident in fresh waters but is dependent on reproduction of marine populations to maintain the freshwater ones.

Habitat

This mullet is found in surface waters of the sea and frequently ascends rivers. It is common in coastal and estuarine waters. It favours marine waters where the average monthly temperature is above 16°C and where summer temperatures rise above 18°C. Temperature ranges of 5-37°C are tolerated. It can live in salinities up to 126‰. It is a fast swimmer and may leap out of the water.

Age and growth

Age at maturity for both sexes varies with water temperatures, at 1 year in the warmest waters, such as Florida, and as late as 6-8 years in the Black Sea for example. Black Sea fish were 31-37 cm long at maturity while in the Gulf of Mexico females were 25.8 cm and males 24.0 cm and larger. Life span lies between 11 and 21 years. Growth rate varies with locality and ceases in warm temperate waters during mid-winter.

Food

Diet is comprised of microscopic organic matter taken in with mouthfuls of sand. The sand probably helps grind up food in the gizzard-like stomach. The food items are diatoms, blue-green and green algae, foraminifers, small crustaceans and detritus, and sometimes plankton. Rarely, swarming marine worms are eaten rather than microscopic items, perhaps as an additional energy source prior to spawning. Juveniles eat invertebrates.

Reproduction

In the sea, spawning occurs in surface waters near the edge of the continental shelf over deep water after a migration from rivers. In the Sea of Marmara, Turkey, this takes place in October and in southern India in October-May. Eggs and larvae drift with ocean currents until they are 20-30 mm about 2-3 months after hatching. Fry make their way into estuaries and juveniles live in estuaries and lower reaches of rivers. Each female is attended by several males during spawning. The males nudge and press against the female's abdomen. The fish may quiver and cease swimming momentarily, sometimes rising to the surface. Fertilisation is external. Spawning occurs once yearly and some females only spawn in alternate years after first maturity. Fecundity is up to 4,800,000 non-adhesive, straw-coloured eggs and egg diameter reaches 1.08 mm (some reports cite 7 million eggs). Hatching occurs in about 2 days.

Parasites and predators

Eslami and Anwar (1971) record the cestode Caryophyllaeus fimbriceps from this species on the Caspian coast of Iran. In the absence of valid records for this fish in Iranian Caspian waters, the host may have been another mullet species.

Economic importance

Fisheries take place around coasts world-wide as the mullet merge into migratory schools. In Hormozgan, is is caught in coastal waters by cast net and stake-net but only at 1.33% by weight and 0.19% by frequency compared to other edible fishes and with a frequency of 5% compared to other mugilids. World-wide, it is sold fresh, frozen or salted and the roe is made into a type of caviar. This species is also widely cultured, for example in Israel and India, with yields up to 2434 fish per hectare. It has been reported as being ciguatoxic (intermittently poisonous through feeding on toxic food) (Bagnis et al., 1970) and as being ichthyoallyeinotoxic (hallucinogenic fish poisoning) (Halstead, 1967-1970; 1978). The latter is a sporadic and mild form of poisoning. This species is rare in Iranian freshwaters and unlikely to be health hazard there (Coad, 1979b).

Fingerlings from Taiwan, Egypt and Hong Kong have been cultured in enclosed ponds in Mazandaran and the Gomishan Wetland (A. Matinfar, pers. comm., 1995; Annual Report 1993-94, Iranian Fisheries Research and Training Organization, Tehran, p. 43-44, 1995; Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 31, 1996; Annual Report, 1995-1996, Iranian Fisheries Research and Training Organization, Tehran, p. 39, 42, 1997; Iranian Fisheries Research and Training Organization Newsletter, Tehran, 18:5, 1997; Iranian Fisheries Research Organization Newsletter, 40 & 41:3, 2004; Iranian Fisheries Research Organization Newsletter, 53:3, 2008). 20,000 fingerlings, weighing about 0.5 g, were imported to the Caspian Sea Ecology Research Center in Sari in 1993. Fry were cultured for 21 days on plankton grown by manuring ponds with wheat and rice bran and soya. Growth in salt water averaged 235 g and 28.4 cm in the first year and 544 g and 40 cm in the second year. Growth in fresh water averaged 216 g and 28.5 cm in the first year and 668 g and 38 cm in the second year. Experiments on the mass production of Nannochloropsis oculata algae used in the culture of rotifers and Mugil cephalus larvae have been carried out in Golestan Province (Iranian Fisheries Research Organization Newsletter, 37:3, 2003). Mirhashemi Rostami et al. (2008) has examined mortality factors in fry from 9 year old broodstock in Iran.

Sharifpour et al. (2003), Ghelichi et al. (2004), Yeganeh et al. (2005), Mirhashem Rostamy et al. (2006) and Tehrani (2006) give further details on culturing this mullet in Iran. Peak spawning was mid to late January with the best time for artificial breeding using hormones being December. Up to 2.6 million eggs could be produced from a female with fertilisation varying from 10 to 95% and hatching rate between 0.008 and 88.9%. Ghelichi and Jorjani (2004) state that this species is expected to play an important future role in fish culture in Iran. They studied induced spawning in this species and found 17-α hydroxyprogesterone to be effective. Gelichi et al. (2007) studied oocyte maturation after stimulation with hormones and also found 17-α hydroxyprogesterone to be the most effective steroids in final maturation.

Conservation

Population numbers and trends are unknown so conservation requirements cannot be ascertained.

Further work

The biology and frequency of occurrence of this species in fresh waters of Iran is unknown.

Sources

Thomson (1963) gives a synopsis of the biology of this species and the above account is based mostly on his work. Distinguishing characters are based on Trewavas and Ingham (1972).

Iranian material: OSU 4281, 1, 234.3 mm standard length, Baluchestan, Kalani, Sarbaz River (25º17'N, 61º24'E).

Atherinidae

The silversides or sand smelts are found in coastal areas of temperate to tropical seas and in fresh water. There are about 60 species (Nelson, 2006) with only one reported from the Caspian Sea and Iran. Most are small fishes with a maximum length of 60 cm.

These small, silvery fishes have a moderately elongate body; usually cycloid and moderately large scales; no lateral line but sometimes a pit on each scale; small teeth in the jaws and sometimes on the roof of the terminal, upwardly-directed mouth; wide gill openings with branchiostegal membranes free of the isthmus; 5-6 branchiostegal rays; gill rakers usually long and slender; two well-separated dorsal fins, the first with 3-10 unbranched but flexible rays, and the second with 1-2 unbranched and the rest branched rays; anal fin long; pectoral fins high on the flank; no pyloric caeca; egg membranes with filamentous outgrowths; back bluish to greenish with small melanophores but translucent; and a silvery stripe along the flank, often distinctively outlined with black.

Silversides can occur in vast schools in inshore waters and are an important item in the diet of other fishes. They have been used as bait but are too bony to be much used as food. Their food is plankton. Eggs are large and greenish. The sticky egg filaments entangle with plants, rocks or sand as anchors until hatching.

Genus Atherina
Linnaeus, 1758

Members of this speciose genus are found in fresh and brackish waters with a single representative in Iran. The record of Atherina hepsetus Linnaeus, 1758 in the Caspian Sea by Quignard and Pras in Whitehead et al. (1984-1986) is an error (Vasil'eva, 1994).

The body is compressed with a rounded belly, the mouth large and terminal, jaws large, reaching back to the anterior eye level or beyond, sides of the upper jaw are straight and premaxillaries protractile, the dentary bone has a high central portion, a preopercular notch is absent, rows of setiform teeth on the jaws, vomer and palatines, cycloid scales extending onto the head, short pectoral fins, vent nearer the pelvic fin origin than the anal fin, numerous gill rakers, 5-10 flexible rays in the first dorsal fin, and the second dorsal fin is similar in length to the anal fin with its origin above the anal fin.

Atherina caspia
Eichwald, 1831

Common names

shisheh mahi (= glass fish), گل آذين ماهي (gol azin mahi, meaning flower decoration fish), atrinka (from the Russian).

[aterinka or xazar aterini in Azerbaijan; kaspi aterinasy in Turkmenian; Kaspiiskaya aterinka or Caspian silverside in Russian; Caspian sand smelt, Boyer's sand smelt, big-scale sand smelt].

Systematics

Formerly called Atherina mochon pontica natio caspia Eichwald, 1831 or Atherina mochon caspia in Caspian Sea literature. Atherina mochon Cuvier, 1829 was described originally from Ivasa and Atherina presbyter pontica Eichwald, 1831 from near Odessa, Ukraine. The Caspian Sea taxon was also known as Atherina boyeri Risso, 1810. Atherina Boyeri was originally described from the sea shore and lower course of rivers in the Département du Var, France. Reshetnikov et al. (1997) give the date for this species as 1826. Atherina presbyter var. caspia Eichwald, 1831 described from "in Caspii maris littore australiore, sinu balchanensi" (i.e. southern shore of the Caspian Sea, Balkhan Bay) was considered a synonym. No types are known. Naseka and Bogutskaya (2009) consider caspia to be a full species.

Syntypes of A. boyeri are in the Muséum national d'Histoire naturelle, Paris under MNHN A-4342 (2) and MNHN B-860 (1) (Eschmeyer's "Catalog of Fishes", downloaded 29 August 2007).

Bamber and Henderson (1985) demonstrated how the morphology of this species around Britain varies according to temperature and salinity during embryonic development. Isolation of populations results in local selection and random genetic drift and thus recognisable morphologies but these were not recognised as distinct. Kottelat (1997) reviews literature reports of variation in this species in the Mediterranean and Black seas and neighbouring fresh waters and concludes that lacustrine populations should be called A. boyeri while marine populations are A. mochon. The status of Caspian populations is not commented on.

The silverside in the Caspian Sea was referred to as Atherina boyeri caspia by Savenkova and Asanov (1991) and by Vasil'eva (1994; 1996b) and this seemed a reasonable step as the population is isolated from other populations in the Atlantic Ocean and Mediterranean Sea. The subspecies was characterised by a reduced number of infraorbital bones (3 bones in Caspian Sea fish as opposed to 4 in Black Sea fish; apparently infraorbital bones 2 and 3 fuse), a reduced number of gill rakers (generally 19-27 as opposed to 27-37), and the form of the maxilla (the inferior margin in Caspian Sea fish is always smooth while in the Black Sea and Sea of Azov a "wing" is usually developed). However Kiener and Spillman (1972) allocated Caspian populations to this subspecies on the basis of larger size, hardly an adequate criterion. Vasil'eva (1994) makes the allocation based on a reduced number of gill rakers, reduction of the number of bones around the eye to three (preorbital, infraorbital and postorbital) and a smooth lower margin to the maxilla without a wing protuberance.

Dobrovolov and Ivanova (1999) studied two non-enzymatic proteins and 11 enzymes for putative Atherina boyeri and A. mochon pontica. They concluded that these are distinct species and indeed the Black Sea fish are a distinct species, A. pontica. They diverged 2.316MYA. These authors did not examine Caspian Sea material. Miller in Miller (2003) followed a conservative approach, regarding the various named and wide-ranging populations as representing a single polymorphic species.

Key characters

The two dorsal fins, cycloid scales, pectoral fin high on the flank and the vent remote from the anal fin are characteristic.

Morphology

First dorsal fin spines 5-10; second dorsal fin with 1-2 spines and 8-15 soft rays; anal fin with 1-2 spines and 9-18 soft rays; pectoral branched rays 10-17, usually 12-15; pelvic fin with 1 spine 5-7, usually 5, branched rays; lateral series scales 37-53, possibly to 61, see Vasil'eva (1994); total gill rakers 19-27 in the Caspian Sea, to 39 elsewhere, spinulose on the interior surface, and long and reaching about 7 rakers along the arch when appressed; and vertebrae 39-52. The anus is 4-5 scales in advance of the anal fin. Scales are higher than broad, with slight indentations on the otherwise straight dorsal and ventral margins, a rounded posterior margin, and a wavy to rounded anterior margin with a protuberant central point. There are no radii. Circuli are restricted to the anterior third of the scale with a central and vertical roughened area posterior to the circuli, presumably made up of fragmented circuli. The focus is central. The lower jaw symphysis fits into a notch in the upper jaws. The haemal arches of the anterior 4-7 caudal vertebrae are expanded around the swimbladder. Gut s-shaped. The chromosome number is 2n=48 (Klinkhardt et al., 1995).

Meristic values for Iranian specimens are: first dorsal fin spines 8(28), 9(5) or 10(1); second dorsal fin soft rays 11(10), 12(21), 13(2) or 14(1); anal fin soft rays 13(7), 14(15), 15(9) or 16(3); pectoral fin branched rays 12(1), 13(12), 14(14) or 15(6); pelvic fin branched rays 5 (34); caudal fin branched rays 13(1), 15(30), 16(2) or 17(1); lateral series scales 50(1), 51(10), 52(5), 53(3), 54(8), 55(3), 56(3) or 57(1); predorsal scales 17(1), 18(2), 19(12), 20(8), 21(3), 22(3), 23(1), 24(3) or 26(1); caudal peduncle scales 12(31), 13(2) or 14(1); transverse scales from anal fin antero-dorsally 10(2), 11(6), 12(23) or 13(3); total gill rakers 24(3), 25(4), 26(14), 27(8), 28(4) or 29(1); total vertebrae 45(1), 46(19), 47(7) or 48(7).

Sexual dimorphism

Females tend to be larger than males and there are some differences in morphometrics and meristics (Ghoorbanalidoost et al., 2003).

Colour

General colour is given in the family account. The lateral band is strongly developed and bright in Caspian Sea specimens. The belly is white, fins are pale to translucent grey. Lagoon specimens are brownish or grey-brown on the back. The peritoneum is brown to black, eggs being encased in a black peritoneum while the abdomen wall is a light brown.

Size

Reaches 11.8 cm and 17.6 g in the Atrak River (Savenkova and Asanaov, 1991), to 14.5 cm standard length in the Caspian Sea basin generally, and elsewhere to 19.0 cm standard length (Henderson and Bamber, 1987). Kiener and Spillman (1972) found a maximum of 15.4 cm in their Caspian Sea sample.

Distribution

Found from the shores of Europe to the Black and Caspian seas, and in the Uzboi Valley of Turkmenistan. In Iran, it is reported along the whole Caspian Sea coast, and from the the Safid River, Talar River, Anzali Mordab, and its outlet the Sowsar Roga seasonally, the Bandar Anzali breakwater and beach, Farahbad, Mazandaran, and Gorgan Bay (Abbasi et al., 1999; Holčík and Oláh, 1992; Roshan Tabari, 1997; Kiabi et al., 1999; Abdoli and Naderi, 2009).

    

Zoogeography

This species probably entered the Caspian Sea from the Black Sea during Khvalyn transgression (10-70,000 years B.P.) via the Kumo-Manych Depression (Kosarev and Yablonskaya, 1994). Berg (1948-1949) contends that it entered the Caspian from the Black Sea in post-glacial times while most other Caspian fishes are relicts of earlier transgressions or migrants from northern waters.

Habitat

The Caspian silverside is a schooling fish found at depths exceeding 100 m but is concentrated at 10-20 m. Salinity tolerance is high, up to 77‰ (Miller in Miller, 2003), up to 60‰ in the Atrak River (Savenkova and Asanov, 1991). Reproduction can occur up to 42‰> while preferred levels are 3-12‰. A wide temperature range is tolerated, 0-31ºC (Miller in Miller, 2003). It is also found in lagoons and river mouths, and enters rivers to spawn, against currents up to 1.2 m/sec. It is the dominant species in the Gomishan Wetland in spring, summer and autumn (Patimar et al., 2009). Holčík and Oláh (1992) report its apparent recent occurrence in the Anzali Mordab in response to increased salinity there. It is also known from fresh water in Lenkoran.

Age and growth

Maturity is attained in the first year and life span varies from 1 to 5 years, 4-5 years usually in the Caspian Sea basin (Henderson and Bamber, 1987). Miller in Miller (2003) summarises growth and population dynamics from populations outside the Caspian Sea. Ghoorbanalidoost et al. (2003) found populations in the south Caspian Sea to be 73.82 mm long on average and 3.15 g in weight, with age  groups 1-3 years, average 1.7 years, total length and length-weight relationship W = 0.00000615L3.02 and a sex ratio of male:female = 0.47:0.53. Heydarnejad (2009) gave the length-weight relationship for an Iranian sample as W = 0.0326TL3.033.

Food

Savenkova and Asanov (1991) report plankton, eggs and juvenile fishes to be food items in the Atrak River of Turkmenistan. Some populations also eat benthic organisms such as amphipods, worms and molluscs. Iranian specimens contained encysted cladocerans and beetles in their guts. It has a trophic plasticity, adapting to whatever conditions obtain (Miller in Miller, 2003).

Reproduction

Spawning in this species is intermittent and occurs along the coast from May to July, peaking in mid-May to mid-June (Savenkova and Asanov, 1991) or from April to August (Henderson and Bamber, 1987). Preserved Iranian fish samples have relatively large eggs from at April to September, e.g. 1.4 mm on 27 April, 1.6 mm on 14 May. Savenkova and Asanov (1991) also studied the annual spawning migration into the Atrak River (lower reaches in Turkmenistan, upper reaches shared with Iran). Fish are caught in a fish ladder 18-19 km from the sea and in the river. The first schools appear in the Atrek mouth as early as mid-February at water temperatures of 8-10°C but the mass migration takes place in mid to late March and the first half of April at 14-22°C. Spawning occurs in March, April and May but most intensively in April at flood water temperatures of 13.7-23.0°C. There is a larger migration in high-water years. Sex ratio is about 1:1. A female may be able to spawn 5-6 times in one season so that egg numbers and diameters vary within each individual for different generations of eggs. Egg deposition in the Caspian Sea is associated with the alga Cladophora to which the eggs are attached or entangled by long filaments (up to 15) and this plant is present in the Atrak. Egg diameters are up to 2.0 mm (3.5 mm in Europe, but possibly these are water hardened) and the light yellow eggs number up to 568 (Gon and Ben-Tuvia, 1983) although the Caspian Sea Biodiversity Database (www.caspianenvironment.org) gives 5500 eggs per female (presumably total seasonal fecundity). Only one gonad develops. Larvae are pelagic but may school close to shore.

Henderson and Bamber (1987) and Gon and Ben-Tuvia (1983) review the reproductive biology of this sand smelt. It can rapidly adjust its life history to a range of environments, from fresh water to coastal water.

Parasites and predators

The Caspian seal, Pusa caspica, is a predator on this species (Krylov, 1984) as are the larger fishes such as Stenodus leucichthys and Alosa saposchnikowii (Lönnberg, 1900a).

Economic importance

This species is food for a number of other fishes including such economic species as sturgeons, Sander and the predacious shads. It has been caught as a bycatch in kilka seine nets and used in fishmeal production. In Europe it has been sold fresh or canned.

Conservation

Kiabi et al. (1999) consider this species to be of least concern in the south Caspian Sea basin according to IUCN criteria. Criteria include abundance in numbers, widespread range (75% of water bodies), absent in other water bodies in Iran, and present outside the Caspian Sea basin.

Further work

The biology of this species in Iranian waters has been investigated by Patimar (1995) (not seen).

Sources

Some meristics are taken from Altun (1999) and Trabelsi et al. (2002).

Iranian material: CMNFI 1970-0507, 11, 22.2-56.3 mm standard length, Gilan, Caspian Sea at Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1970-0509, 5, 49.5-101.7 mm standard length, Gilan, pond at Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1970-0543A, 7, 52.9-83.1 mm standard length, Gilan, Caspian Sea at Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1970-0563, 18, 39.8-112.5 mm standard length, Gilan, Kazian Beach (ca. 37º29'N, ca. 49º29'E); CMNFI 1970-0581, 5, 47.6-54.8 mm standard length, Caspian Sea near Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1971-0343, 2, 65.9-72.5 mm standard length, Gilan, Langarud at Chamkhaleh (37º13'N, 50º16'E); CMNFI 1979-0081, 3, 74.1-77.3 mm standard length, Mazandaran, Caspian Sea, 3 km west of Chalus (36º41'N, 51º24'E); CMNFI 1980-0146, 4, 92.1-103.6 mm standard length, Mazandaran, Gorgan Mordab at Ashuradeh-ye Kuchak (36º50'N, 53º56'E); and CMNFI 1980-0160, 2, 49.2-73.8 mm standard length, Iran, Caspian Sea basin (no other locality data).

Adrianichthyidae

Genus Oryzias
Jordan and Snyder, 1906

Oryzias latipes
(Temminck and Schlegel, 1846)

Introduced to the Tedzhen River basin and Karakum Canal in Turkmenistan (Aliev et al., 1988; Sal'nikov, 1995) but no confirmed Iranian record. It is reported as reproducing in Iran by the Food and Agriculture Organization, Rome (www.fao.org/scripts/acqintro/query/retrive.ide, downloaded 28 July 1999) but this record is presumably in error and needs confirmation by specimens.

Cyprinodontidae

The tooth-carps, killifishes or pupfishes are small fishes found in fresh, brackish and sea water. There are 9 genera and about 105 species found in tropical to warm temperate climates almost world-wide. Iran has 7 described species, with more to be discovered.

This family is characterised by a moderately elongate and compressed body, rather cyprinid-like but with jaw teeth (hence the common and scientific names), body and head covered with scales, no spines in the fins, no barbels, the mouth is very protractile and armed with comb-like teeth, the lower jaw is strong and robust as the dentary is expanded medially, 4-6 branchiostegal rays, gill membranes free from the isthmus, lateral line absent or reduced to points, caudal fin rounded or truncate, pectoral fins set low on the body (in contrast to Poeciliidae), dorsal and anal fins short, no adipose fin, no pyloric caeca, and no gonopodium (egg layers, in contrast to the livebearer subfamily in the Poeciliidae).

Wildekamp (1993) gives a general account of cyprinodontid species. These fish can be maintained in aquaria, and are very popular with aquarists. Specific requirements where known are given under each species. Moderate to hard fresh water is used and for water with a total hardness less than 10dGH, sea salt can be added (about one teaspoon per 10 litres). Summer temperatures of 20-25°C are good and although higher temperatures make the fish more active, they also age the fish more rapidly. Winter temperatures below 20°C imitate nature and are recommended. Weekly partial water changes are suggested and live and frozen fish foods and even flake foods are eaten. Breeding occurs throughout the year in aquaria kept above 15°C, eggs are deposited on fine-leaved plants (thread algae, Java moss or yarn mops) or even on aquarium filter sponges and gravel. Egg development takes 8-20 days, fry take about 2 days to absorb their yolk sac and will then consume baby brine shrimp. These fish can be kept in outdoor ponds in milder climates (http://ark.aka.org/Eurasian.htm, downloaded 1 December 2003).

Genus Aphanius
Nardo, 1827

These tooth-carps are found around the shores of the Mediterranean, in Southwest Asia and as far as northeastern India and Somalia. There are about 23 known species.

Sethi (1960) advocated placing Aphanius in a separate family, Aphaniidae, but this did not find general acceptance.

The genus Aphanius Nardo, 1827 has been used for these tooth-carps for many years. However, Lazara (1995) designated Lebias fasciata Valenciennes, 1821 as the type species for Lebias Goldfuss, 1820, making Lebias a subjective senior synonym of Aphanius. On this basis Lebias must be used rather than Aphanius but Lazara's type species designation is invalid (Kottelat and Wheeler, 2001). Since Lazara (1995) could have chosen another species as type species, the change involves a large number of species and these species are threatened and listed in various legislation, a petition is before the International Commission on Zoological Nomenclature to suppress Lazara's designation (Kottelat, 1997; Kottelat and Wheeler, 2001; Wildekamp, 2001; Villwock et al., 2002). Wildekamp et al. (1999) present evidence that Lebias is a synonym of Cyprinodon.

Literature on these Mediterranean and Southwest Asian fishes may appear under either of these names or under Cyprinodon Lacepède, 1803, the latter now restricted to American species.

Bănărescu (1995) disagrees with Parenti's (1981) relationship of Aphanius to South American Orestias (see also Parker and Kornfield (1995), Stevenson (1997) and Costa (1997) for conflicting views).

This genus is characterised by a thick oval body, large to moderate cycloid scales, the head flattened on top, a small, superior mouth with tricuspid teeth, the upper jaw bordered by the premaxillaries only, lateral line system present only on the head, dorsal fin positioned somewhat posteriorly with1-2 unbranched rays and 7-13 branched rays, anal fin rays 1-2 unbranched and 7-14 branched, dorsal and anal fins larger in males than in females, dorsal fin inserted opposite the anal fin origin (in contrast to Gambusia), and colouration of males and females distinct. Seyfali et al. (2002) has 2n=48 for fish from the Damghan basin which Coad and Abdoli (2000b) suggest may be a distinct taxon. Kamal et al. (2009, 2009) studied some biological characteristics of the Damghan population.

Aphanius is the only genus in the family currently recognised in Iran. However Parenti (1981) distinguishes derived members of the genus Aphanius as `Aphanius' without formally describing a new genus. One of the distinguishing features of `Aphanius' is the reduction of cephalic sensory pores to neuromasts, a character found in A. mento, A. sophiae and A. vladykovi in Iran.

Wildekamp et al. (1999) review the Turkish species which show scale reductions not seen in Iranian populations. Hrbek et al. (2002) outline the historical biogeography of the species complex in central Anatolia, Turkey using mtDNA, testing the hypothesis of geographic speciation driven by early Pliocene orogenic events that may well be paralleled in Iran. Hrbek and Meyer (2003) studied the phylogeny of Eurasian tooth-carps using mtDNA and observed a western Tethys Sea clade (all species except those listed below) with a middle Oligocene divergence into Iberian Peninsula and Atlas Mountains, and Turkey and Iran sections. Late Miocene orogenic events were correlated with a large amount of genetic differentiation in Turkey (and presumably Iran). An eastern Tethys Sea clade (dispar, ginaonis, mento, sirhani) had an Oligocene divergence into a freshwater clade inhabiting the Arabian Peninsula and neighbouring areas and a euhaline clade inhabiting coastal areas from Pakistan to Somalia. Speciation is predominately vicariant-based but ecological factors played a significant role.

These fishes are known generally as gour-e khar in Farsi (= literally "striped donkey" which means zebra here although usually in Iran this term refers to the wild ass or onager), kapurdandandar or kopurdandandar (= tooth-carp), or even آفانيوس (= Aphanius). Saefali (1999) is a recent study of these fishes in Farsi.

Iranian Aphanius away from coastal drainages are thought to be relicts of the Tethys Sea (Kosswig, 1955a; 1967; Bianco, 1995), having been trapped by the rising Iranian Plateau, rather than invaders from the coast (Krinsley, 1970). Fossil tooth-carps, Aphanius kirgisicus and A. longipinnis have been described from Kyrgyzstan in Miocene deposits, far inland from the distribution of living species (Yakovlev, 1959). Priem (1908) describes a fossil cyprinodont from the Miocene of Iran.

The systematics of the Iranian populations would repay careful study using biochemical and genetic techniques coupled with aquarium studies on behaviour and cross breeding. Certain populations, isolated from others in discrete drainage basins, appear to be different on the basis of colour patterns yet show little morphological variation, e.g. the Damghan population, see Coad and Abdoli (2000b), Saefali (1999) and Seyfali et al. (2002), the most northerly and easterly population in Iran characterised by large spots in females; populations in Fars outside the Lake Maharlu and Kor River basins; and populations in the Namak Lake basin - all yet to be investigated in detail (Coad, 2000b). Since these patterns are used in mate recognition, the populations could be distinct taxa. Huber (1996) has noted in cyprinodonts from Africa that many taxa can only be separated on the basis of colour in life, other characters overlapping. He suggests a cyprinodont species definition which includes possession of at least one stable phenotypic characteristic, and this could be colour of the male fish. He does recommend that genetic isolation be demonstrated where possible by karylogy, biochemical techniques or breeding experiments.

Seifali et al. (2003) compare A. sophiae and A. vladykovi meristically and morphometrically. Saifali et al. (2004) found a suite of morphometric characters to be different between samples of A. vladykovi and fish from Qomy-Abad in Tehran Province, the Namak Lake basin.

The mosquitofish (Gambusia) is a niche competitor and may well eliminate these tooth-carps. Generally, Aphanius prefers springs, lakes, marshes, sea shores and hot springs. Flowing water is usually avoided but they can be found near the source of springs flowing into salt lakes, constrained by hypersaline waters downstream. However they are tolerant of high salinities. They occur in schools but males are aggressive to other males. Males are brightly coloured, often striped, while females are subdued with spots or faint bars.

A related Mediterranean species is reputed to be ichthyootoxic but this has not been demonstrated for Iranian species (Coad, 1979b). Symptoms of this egg poisoning are summarised under the genus Schizothorax.

These small fishes are used in the aquarium trade as they tolerate a wide range of temperatures and salinities and are particularly colourful. As early as 1912, J. P. Arnold in Germany noted the survival of imported Iranian Aphanius (possibly A. persicus) in aquaria at temperatures of 14-27°C, with a variety of plants such as Myriophyllum, Nitella, Riccia, Cabomba, Salvinia and filamentous algae, with daphnia, cyclops, enchytraeids and red mosquito larvae as food, and egg laying apparently preferred on the Salvinia with 8-12 days for hatching at 27°C.

Aphanius sp. from the Namak Lake is recorded as having the parasite Clinsotomum complanatum (Hosseinie, 1987). Aphanius from Damghan were infected with the trematode Tetracotyle sp. (Gholami et al., 2011). 

Kamal et al. (2007; 2009) and Bakhtiyari et al. (2011) give some life history data on the Damghan and Shour River of Eshtehard (west of Tehran) populations. Food in the Shour River was mostly chironomids while crustaceans including Daphnia were eaten at Damghan. Stable environmental conditions at Damghan allowed more prey selection and a higher condition factor. The Damghan fish lived longer and had a higher egg diameter while Shour River fish had a higher L and a shorter reproductive period, again probably reflecting environmental constraints..

Aphanius dispar
(Rüppell, 1829)

Male

Common names

gour-e khar, kopurdandandar-e balehbolband (= tooth-carp with striped fin), kapurdandan-e balmband.

[harsun. batrikh or batrikh motakayer in Arabic; nambal in Pakistan; high-finned pupfish, Arabian killifish, mother of pearl fish].

Systematics

A detailed synonymy is given by Villwock et al., (1983), Wildekamp et al. (1986) and Wildekamp (1993). Hoedeman (1951) proposed a new genus for this species, Aphaniops, based on the absence of a dermal sheath or genital pouch around the anterior anal fin rays, only 8-9 dorsal fin rays in contrast to 10-14 rays in Aphanius and 7-8 pelvic fin rays in contrast 5-7 rays. It has not found general acceptance.

The type subspecies is found in all Iranian drainages (Krupp, 1983) although Aphanius dispar stoliczkanus (Day, 1872) has been recorded in earlier literature as the subspecies of these drainages except the Tigris River basin for which Aphanius dispar richardsoni (Boulenger, 1907) is reported (Berg, 1949). Aphanius dispar richardsoni is limited to the Dead Sea valley of Israel and western Jordan by Villwock et al. (1983) and Wildekamp (1993). Cyprinodon stoliczkanus Day, 1872 is regarded as a synonym of Aphanius dispar dispar by Krupp (1983) while Berg (1949) places Bampur River, Baluchestan fish in this subspecies as a southern representative of Aphanius dispar with 6-9, rarely 10 dorsal fin rays as opposed to 9-10 rays in Aphanius d. dispar.

Hrbek and Meyer (2003) note that, based on their mtDNA study, the monophyly of A. dispar is strongly rejected and it does not constitute a species in terms of the phylogenetic species concept. Reichenbacher et al. (2009) found that isolated populations in Saudi Arabia may be capable of evolving into new species in a short period of time, based on otolith morphology - evidently there is still much to be learned about this nominal species.

Hybrids with Aphanius dispar richardsoni are reported from Israel (Goren and Rychwalski, 1978).

The lectotype of Lebias dispar as designated by Villwock et al. (1983) is in the Senckenberg Museum Frankfurt under SMF 821 with paralectotypes SMF 1988 (10). Further paralectotypes in the Natural History Museum, London under BM(NH) 1860.11.9:152 listed as from Abyssinia (3 fish, 44.1-49.6 mm standard length) (Eschmeyer et al., 1996) but a label in the jar states that "status as types doubtful, types in SMF range from 26.1-35.8 mm SL, locality in description and types given as 'Red Sea'".

Types of Cyprinodon stoliczkanus Day, 1872 are in the Zoological Survey of India, Calcutta (ZSI 1477, ZSI 1478), the Natural History Museum, London (BM(NH) 1889.2.1.2065-74, originally 21 fish, now 14 fish, 14.4-29.4 mm standard length when examined in September 2007) and the Australian Museum, Sydney (AMS B.7730-7731) (Whitehead and Talwar, 1976; Eschmeyer et al., 1996; Ferraris et al., 2000). This species was originally described from a "stream at the village of Joorun, and also at Lodai, along the edge of the Rann" (= the salt water Rann of Cutch, Sind, India).

The lectotype of Cyprinodon richardsoni as designated by Krupp and Schneider (1989) is in the Natural History Museum, London under BM(NH) 1856.5.2:4 with paralectotypes under BM(NH) 1856.5.2:5 (8 fish) (12.9-25.0 mm standard length, the largest being the lectotype) (Eschmeyer et al., 1996; personal observations).

Krupp (1983) did not find the variation in teeth numbers between different populations and subspecies observed by Berg (1949).

Key characters

The colour pattern is distinctive.

Morphology

Scales along the flank 24-35. Scales are squarish with an almost vertical anterior margin and protruding anterior corners, parallel dorsal and ventral margins and a rounded posterior margin with tiny teeth. The anterior margin has a small central protuberance with shallow indentations above and below or almost straight margins to each anterior corner. Radii are present on the anterior field, numbering 10-20 but mostly 14-16, and are almost horizontal and parallel. The exposed part of the scale has dimples rather than circuli. Circuli are few. The focus is subcentral posterior. There is no pelvic axillary scale. Total dorsal fin rays 7-11, usually 9-10, total anal fin rays 8-12, usually 10-11 (the number of unbranched rays in the dorsal and anal fins varies from 1-3), total pectoral rays 12-18 and total pelvic rays 6-7, usually 7. Vertebrae 24-29. Al-Hassan (1982b) reports on abnormalities in the vertebrae including loss of the posterior part of the centrum and fusions. Teeth are tricuspid with the central cusp concave at its tip and only slightly longer than the lateral cusps (Goren and Rychwalski, 1978), although in some fish seen by me the tip is rounded. There are 12-20 teeth per jaw (Krupp, 1983). Gill rakers 11-20, modally 13-16, reaching the second adjacent raker when appressed. Some gill raker counts are difficult to make accurately as those at the anterior arch end are minute and those at the dorsal end are partially concealed in flesh. The gut is coiled ?see my ginaonis paper. Reichenbacher et al. (2007) give a description of otolith morphology. Chromosomes number 2n=48 (Klinkhardt et al., 1995; Esmaeili et al., 2008). Karyotype 16Sm + 32St and arm number 32 (Esmaeili et al., 2008).).

Meristic values for Iranian specimens:- total dorsal fin rays 8(1), 9(39) or 10(10); total anal fin rays 9(2), 10(42) or 11(6); total pectoral fin rays 14(2), 15(16), 16(28), 17(3) or 18(1); total pelvic fin rays 6(6) or 7(44); lateral series scales 24(2), 25(5), 26(11), 27(17), 28(7), 29(2), 30(1), 31(2), 33(2) or 35(1); total gill rakers 15(13), 16(16), 17(16) or 18(3); and total vertebrae ?.

Sexual dimorphism

Males have longer fins than females and are more brightly coloured. The dorsal fin is twice as long in the male and reaches the caudal fin when appressed. When expanded it is widely flared and distinctive as is the enlarged anal fin.

Colour

Breeding males are brown-grey, grey or black-brown with iridescent blue-white flank spots and white and brown to light orange or light blue, irregular, narrow bars. The head has blue and orange tinges and in particular there is an orange spot on the operculum postero-dorsally. Lips are blue-white. The flank over the pectoral fin has electric blue spots. The anterior belly becomes blue with pearl spots. Scales have a dark margin. The dorsal fin is spotted light blue on a bright orange background and is barred. Barring may be irregular and in overall view in breeding males appears as speckling. Pectoral, anal and pelvic fins are lemon-yellow. The anal fin has barring on the posterior 4 rays in breeding males. The caudal fin has 2-3 dark and light blue alternating bars, the last bar being yellow. The bars are crescent-shaped, with concave side posterior. Males outside the breeding season are less brightly coloured with silvery on the flanks with a grey or black-brown back and irregular flank bars. Male specimens from the Bampur River basin in Iran have a body speckled silvery-white to light blue, weak tail bars, blue and orange tinges to the head, an orange spot on the operculum and an orange-yellow edge to the anal fin.

Young have pale brown flank bars on the caudal peduncle. Some young may have a few spots on the flank rather than bars. Females are brown-grey to silvery with 8-20 narrow flank bars and a dark-brown back. The flank bars are dark brown and may have a tinge of orange, the interspaces silvery. Flank bars may number as few as 7 (Talwar and Jhingran, 1991). Females also have the orange operculum spot and the blue spots over the pectoral fin. Female fins are hyaline. The peritoneum is brown to black.

Ross (1980) notes that Arabian specimens are brighter in waters with algae and plants; those found over sand and mud are duller, an effective camouflage. He also records variant colour patterns with blue tails, or an overall gold colour, or with green or turquoise flank spots.

Size

Reaches 8.0 cm, but this may be an error as Krupp (1983) points out. He found fish only to 5.2 cm standard length although in Carpenter et al. (1997) 8 cm is given as a maximum. Wildekamp (1993) records fish to 7.0 cm for males and 6.5 cm for females in total length. Fish from Ain Al Adhari, Bahrein reach 7.3 cm total length. Generally fish in fresh water are smaller than marine specimens.

Distribution

Found from Egypt, eastern Sudan and Ethiopia, and in the Red Sea through southern Southwest Asia and east to Rajasthan. Also in the eastern Mediterranean Sea (Kornfield and Nevo, 1976). The subspecies Aphanius dispar richardsoni is found in the Dead Sea rift valley.

A specimen in the Naturhistorisches Museum Wien from the hot spring at Ginao (NMW 13799) may be mislabelled as several other collections from this hot spring are all clearly A. ginaonis.

This species is found in Iranian fresh waters from the Tigris basin to the Makran basin in water bodies draining to the Persian Gulf and Sea of Oman, islands in the Gulf such as Qeshm (OSU 8120), and in the endorheic Hamun-e Jaz Murian basin. It is reported from the Shapur and Dalaki rivers (Gh. Izadpanahi, pers. comm., 1995), Gudar River east of Anveh, Mehran River below Bastak, Sarageh River north of Bandar Abbas, Galashkerd Lake near Minab, and the Kol. Shur, Hasan Langi and Ghale-gazii rivers in Hormozdgan. Abdoli (2000) illustrates distributions in the Arvand, lower Zohreh (= Hendijan), lower Hilleh, Mand, Kul, Hasan Langi and Minab rivers, lower reaches of rivers in the Makran basin from the Jagin to Bahu Kalat, the lower Halil and Bampur Rivers in the Hamun-e Jaz Murian basin, and the endorheic Mashkid River.

Zoogeography

Villwock et al. (1983) contend that an ancestor of A. dispar was present in the middle Miocene Transgression connecting the Mediterranean Sea to the Indian Ocean. The mesohaline conditions, which developed in the Mesopotamian basin, were a good prerequisite for the development of A. dispar since, apart from some euryhaline species, there would be no competitors. A. dispar spread from Mesopotamia through the Persian Gulf to India and the Red Sea. During the Pleistocene, it colonised inland waters.

Habitat

Highly tolerant of salinity, this species is found from the sea to fresh water habitats, including landlocked basins. Slightly saline waters in aquaria seem to discourage parasites (Andrews, 1983). Salinities as high as 145‰ are tolerated (Lotan, 1971) and this fish can be moved abruptly from fresh to hypersaline water. Adaptation to high salinity waters involves a reduced osmotic permeability of the gills and an increase in transportation of sodium chloride by the intestine (Skadhauge and Lotan, (1974). Lotan (1969, 1971, 1973) and Skadhauge and Lotan (1974) describe sodium, chloride and water balance, and osmoregulation in this species. Plaut (1999) found that salinities above ~250% seawater seem to decrease swimming capabilities and routine activity rate and a decrease in resting metabolic rate indicated stress. This species has been reported to survive in waters 38.4°C in Oman and only began to die at 45.1-46.0°C (Haas, 1982). Fish survived up to 40°C in aquaria without acclimation (Lewis, 1970). Populations survive in isolated sections of streams with little food because of histological changes in the stomach (Ba-Omar et al. (1998). Aphanius dispar occurs in shallow water and among vegetation over sand, rock or soft detritus bottoms.

Frenkel (1995) describes the effects of environmental factors on growth and reproduction in this species in Israel. Frenkel and Goren (1997) showed fish to be tolerant of a wide range of environmental variables as measured against the gonado-somatic index (GSI) and ovary maturation stages. The GSI was not affected by temperature over a range of 18 to 37°C while maturation increased from 18 to 27°C but remained the same between 27 and 37°C. For salinities between 0‰ and 56‰ differences were only found in mean GSIs at the two extremes and at 0‰ ovaries contained only primordial germ cells. GSIs and maturation stages only differed in feeding experiments when fish were deprived of food or fed at equal or greater than 1% of body weight per day. A decrease in photoperiod from 14L:10D (14 hours light, 10 hours dark) to 10L:14D caused a decrease in oocyte maturation stages and ovaries were no longer suitable for spawning. In aquaria, a pH of 7.5-8.5, temperatures above 27°C and water hardness greater than 200 p.p.m. and the addition 4 heaped tablespoons of kosher salt per ten gallons of water (38 litres) favoured reproduction (Mackowiak, 1988). Alkahem (1989) found 50% mortality in 96 hours when fish were exposed to pH levels of 3.5-3.9.

Feulner (1998) reports that United Arab Emirate fish "hover" in the water column with the tail slightly curved to one side, presumably a feeding behaviour. Males and females in aquaria swam in separate schools (Al-Daham et al., 1977).

Age and growth

Sexual maturity in ideal conditions may be attained as early as 7 months but full size takes a year. The length-weight relationship in southern Iraq was W = 1.2301 x 10-4 L2.4544 for males and W = 2.8357 x 10-6 L3.5112 for females. The average condition factor for males was 1.8900 and for females 1.6952. For a given length group males are heavier than females and condition factors are higher for males (Huq et al. 1977). Muhsin (1987) studied the effect of differing food supply on the weight and chemical composition in females of this species taken from the Al-Asafiah River, Basrah, Iraq but examined in tanks. Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 41 Iranian fish measuring 2.37-3.56 cm total length. The a-value was 0.0012 and the b-value 3.214 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases).

Food

Iranian specimens have guts packed with filamentous algae and detrital remains. Al-Daham et al. (1977) found food in southern Iraq to be predominately filamentous algae and diatoms, with some copepods, rotifers and, in one instance, a small winged insect. In contrast, Shafi and Shalli (1986) report a diet of beetles, ephemeropteran nymphs, rotifers, filamentous algae and plants in southern Iraq. Younis et al. (2001b) found Shatt al Arab, Iraq fish fed on amphipods and copepods (29.9%), organic detritus (11.6%), and algae and diatoms (7.1%). In aquaria, Al-Daham et al. (1977) found this species to prefer animal foods in contrast to Aphanius mento which preferred plant foods. Al-Akel et al. (1987) have observed the selective feeding behaviour of this species at Al-Khurj, Saudi Arabia. Certain species of filamentous algae are selected over others, e.g. the blue-green alga Oscillatoria, the green algae Ulothrix, Spirogyra and Crucigenia, the desmid Cosmarium, and the diatoms Cocconeis, Diatoma, Navicula and Rhoicosphania, and algal spores and zygotes. Haas (1982) observed this species in Oman to pick at rocks and other substrates, take items at the surface, chase small fish and eat insect larvae. Gut contents are mainly desmids and diatoms, some filamentous algae and rarely insect larvae and one snail. This species is an omnivorous surface feeder (Carpenter et al., 1997).

Reproduction

A female in spawning condition approached a male and appeared to feed on a plant or plant-encrusted rock in fish studied by Haas (1982) in Oman. Feulner (1998) reports fluttering of the tail by male fish. The male places his chin on her nape and 2-3 eggs are spawned rapidly. The female presses her vent against the plant or rock vertical surface, the male arches his back into an s-shape and presses his body and anal fin against hers and clasps her with his dorsal fin. Eggs are released singly with a quiver. The time elapsed from chin on nape to egg release is as short as 1-2 seconds. Reproduction occurs throughout the day and throughout the year in Oman with 69% of females having ripe eggs (up to 41) in April-June. Peak spawning in southern Iraq is April to June when only a small proportion of eggs in the single ovary are fully developed eggs (Al-Daham et al., 1977). Shafi and Shalli (1986) record up to 73 mature eggs in southern Iraq specimens with a diameter of 220 microns. Breeding season was May-July. In Saudi Arabia mature and developing oocytes are observed in fish during the whole year. The egg cycle may be more than one year with spawning in March and April (El-Hawawi and Al-Imam, 1984). On the Mediterranean coast of Egypt, spawning occurs from March to September with a peak in July and August. Maximum egg diameter is 2.2 mm and size at maturity for females is 30 mm total length (Lotan and Ben-Tuvia, 1996). Iranian specimens have large eggs (2.0 mm) on 16 March but young fish (8.9 mm standard length) were caught on 26 November suggesting reproduction is almost year round.

Spawning areas have some water flow and vertical surfaces of plants or algae-encrusted rocks. Males defend a territory about 30 cm wide against other males and will display with erect fins if approached. Parallel swimming, head to head confrontations, rapid circling, and flank bites were observed by Haas (1982). Two males will swim and chase each other in circles until one swims away with fins depressed. Males will change their territory after one day defending it. In Israel, males of Aphanius dispar richardsoni do not show any territorial behaviour (Goren and Rychwalski, 1978). This fish will spawn in a few months of birth in aquaria (Ross, 1987). Three pairs of fish yielded about 40 eggs per day in aquaria and these eggs hatched in two weeks at 27°C (Mackowiak, 1988).

Parasites and predators

This species is reported to be heavily infested with metacercariae of the trematode Clinostomum complanatum in eastern Saudi Arabia, up to 86.6% incidence with up to 41 metacercariae per fish, the infestation being higher in females than males (Kalantan et al., 1987).

Economic importance

Al-Akel et al. (1987) suggest the use of this species as a control agent for filamentous algae.

Steven (1913) reported that this fish was responsible for the absence of mosquito larvae from streams in Karachi which otherwise appeared to be good breeding places. Christophers and Shortt (1920-1921) thought that tooth-carps in southern Mesopotamia were a significant factor in lowering the malaria rate, presumably mostly this species. Sharma and Al-Daham (1979) compared the efficiency of this tooth-carp with the mosquitofish for mosquito control under experimental conditions. The tooth-carp is more efficient when males and females were together than mosquitofish since males of the latter expend time and energy defending territory. Mosquitofish are more active in pursuit of mosquito larvae than the tooth-carp when filamentous algae was present. Cloudy conditions and lower temperatures slow consumption and both species prefer pupae to larvae of mosquitos, with females taking more pupae than males. The tooth-carp could consume nearly twice the amount of mosquitos than the mosquitofish over a given period of time. Overall the tooth-carp compares favourably with the mosquitofish as a destroyer of the malaria-carrying mosquito. Homski et al. (1994) under laboratory conditions found that Aphanius dispar is more successful than the mosquitofish in preying on the third and fourth instars and on pupal stages of mosquitos while the mosquitofish, with a smaller relative mouth size, is more successful on the first two instars. Small fish of both species eat first instars exclusively. However, Aphanius dispar eats more second instar larvae under a vegetation cover since the larger Aphanius are more capable of penetrating shallow water than mosquitofish. Aphanius dispar occurs in shallower water and among more vegetation than the open water mosquitofish. The two species can be used to complement each other in control of mosquitos.

Ataur-Rahim (1981) found that this species will eat mosquito larvae in water storage tanks in Riyadh, Saudi Arabia. Fifteen fish consumed all larvae at a mean density of 5/100 sq cm water surface area in a tank 3 by 2 m in a day. One fish, 5.2 cm long, ate 61 mosquito larvae. Haas (1982) fed this species mosquito larvae in the presence of abundant algae as an alternative food. An average of 96 larvae are eaten each day. Etemadfar et al. (1983) studied the potential use of this species for malaria control in southern Fars, Bushehr and Hormozgan provinces. Louis et al. (1988) detail a control strategy against malaria using this species in Djibouti as do Fletcher et al. (1992) in Ethiopia and Fareed and Baban (1995) in Yemen. Fletcher et al. (1992) showed a decrease in sites harbouring mosquito larvae from 34% without the fish to 1.6% with the fish stocked. Chandra et al. (2008) briefly reviews the use of this species in biocontrol.

The studies of Frenkel and Goren (1997) in Israel show that the reproduction of this species can be controlled such that it has potential for mass production and could be used in mosquito control. Carpenter et al. (1997) state that this species is used in mosquito control. Ross (1987) gives details of aquarium care and breeding of this species.

Conservation

Al-Johany and Yousuf (1993) show that Gambusia affinis has a wider range of temperature tolerance than this species and is better suited for a desert habitat in Saudi Arabia. The exotic Gambusia is therefore a threat to A. dispar. Al-Kaheh-Al-Balawi et al. (2008) found that Saudi Arabian populations are declining from reduced availability of food, habitat degradation, chemical contamination, introduction of exotics and exploitation and some of these factors doubtless obtain in Iran.

Further work

This widespread species has been reported from a number of springs in southern Iran and, like A. ginaonis, may be distinct at some level. Molecular work would be useful in determining the taxa involved and evolution in hot springs. The utility of this species in mosquito control in smaller water bodies of Iran where it also occurs naturally could be explored. Al-Daham et al. (1977) found that this species in aquaria ate mosquito larvae "eagerly" compared to "very eagerly" for Gambusia holbrooki and "not eaten" for Aphanius mento.

Sources

Type material:

Iranian material: NMW 13799, 1, ?; CMNFI 1979-0129, 1, 31.3 mm standard length, Fars, spring about 2 km north of Farrashband (28º54'N, 52º04'E); CMNFI 1979-0137, 106, 8.9-37.7 mm standard length, Fars, stream outside Lar (27º40'N, 54º32'30"E); CMNFI 1979-0138, 1, ? mm standard length, Fars-Hormozgan border, Rasul River drainage (ca. 27º32'N, ca. 54º58'30"E); CMNFI 1979-0139, 2, ? mm standard length, Fars-Hormozgan border, Rasul River drainage (ca. 27º25'30"N, ca. 54º59'E); CMNFI 1979-0140, 3, ? mm standard length, Hormozgan, Kul River drainage (27º14'N, 55º46'30"E); CMNFI 1979-0141, 6, ? mm standard length, Hormozgan, Kul River at road bridge (27º17'30"N, 56º03'30"E); CMNFI 1979-0142, 10, ? mm standard length, Hormozgan, Baghu River at Baghu (27º17'N, 56º28'E); CMNFI 1979-0143, 118, ? mm standard length, Hormozgan, Hasan Langi River drainage (27º21'N, 56º50'30"E); CMNFI 1979-0145, 1, ? mm standard length, Hormozgan, Geru River (26º55'N, 57º01'30"E); CMNFI 1979-0148, 21, ? mm standard length, Hormozgan, Sarzeh River (27º30'30"N, 56º15'30"E); CMNFI 1979-0151, 45, ? mm standard length, Hormozgan, Shur River drainage near Kahkom (28º02'N, 55º53'E); CMNFI 1979-0177, 37, ? mm standard length, Hormozgan, Sarzeh River drainage (27º33'N, 56º14'E); CMNFI 1979-0179, 13, ? mm standard length, Hormozgan, Sarzeh River drainage (27º36'N, 56º15'E); CMNFI 1979-0181, 9, ? mm standard length, Hormozgan, Kul River drainage (27º17'30"N, 56º03'30"E); CMNFI 1979-0182, 31, ? mm standard length, Hormozgan, stream on road to Bandar-e Lengeh (27º14'N, 55º46'30"E); CMNFI 1979-0185, 24, ? mm standard length, Hormizgan, stream in Rasul Rievr drainage (27º06'N, 55º45'E); CMNFI 1979-0187, 2, 26.2-33.2 mm standard length, Hormozgan, Sar Khun oasis (27º23'30"N, 56º26'E); CMNFI 1979-0312, 7, ? mm standard length, Baluchestan, dam on Bampur River (27º11'N, 60º36'E); CMNFI 1979-0313, 1, ? mm standard length, Baluchestan, Bampur River at Bangharabad (27º20'N, 60º46'E); CMNFI 1979-0328, 18, ? mm standard length, Baluchestan, jube 2 km south of Bampur River near Bampur (27º10'30"N, 60º21'E); CMNFI 1979-0352, 2, ? mm standard length, Khuzestan, marsh in Jarrahi River drainage (30º33'30"N, 48º48'E); CMNFI 1979-0354, 8, ? mm standard length, Khuzestan, Karun River tributary (30º31'N, 48º19'E); CMNFI 1979-0355, 1, ? mm standard length, Khuzestan, Karun River tributary (30º35'N, 48º22'E); CMNFI 1979-0362, 2, ? mm standard length, Khuzestan, jube in Karkheh River drainage (31º42'N, 48º33'E); CMNFI 1979-0379, 4, ? mm standard length, Khuzestan, Dez River (32º12'N, 48º27'E); CMNFI 1979-0381, 1, ? mm standard length, Khuzestan, stream west of Shushtar (ca. 32º10'N, ca. 48º35'E); CMNFI 1979-0384, 7, ? mm standard length, Khuzestan, Ab-e Shur drainage (32º00'N, 49º07'E); CMNFI 1979-0403, 25, check size range ?19.7-37.0 mm standard length, Bushehr, stream 23 km south of Kaki (ca. 28º11'N, ca. 51º43'E); CMNFI 1979-0406, 48, 15.6-35.8 mm standard length, Hormozgan, stream near Bandar-e Charak (26º48'N, 54º18'E); CMNFI 1979-0407, 3, ? mm standard length, Hormozgan, stream 30 km from Barvedun (26º51'N, 54º37'E); CMNFI 1979-0408, 106, 14.6-48.4 mm standard length, Hormozgan, Mehran River (27º04'N, 54º35'E); CMNFI 1979-0410, 10, ? mm standard length, Hormozgan, Mehran River (26º53'N, 55º17'E); CMNFI 1979-0414, 12, ? mm standard length, Hormozgan, pool on road to Tiab (27º05'N, 57º02'E); CMNFI 1979-0415, 1, ? mm standard length, Hormozgan, stream 18 km south of Genu (27º17'30"N, 56º20'E); CMNFI 2007-0053, 4, ? mm standard length, Hormozgan, Sarzeh River (ca. 27º36'N, ca. 56º15'E); CMNFI 2007-0057, 16, ? mm standard length, Hormozgan, Mehran River 4 km below Bastak (ca. 27º05'N, ca. 54º05'E); CMNFI 2007-0058, 3, ? mm standard length, Fars, headwaters of Gowdar River (ca. 27º24'N, ca. 54º16'E); OSU 8129, 1, 54.5 mm standard length, Hormozgan, Qeshm Island salt spring (no other locality data); BWC95-30, uncatalogued, 3, ?, Khuzestan, Kupal (31º15'N, 49º10'E); BWC98-6, uncatalogued, 24, ?, Hormozgan, Ab-e Garm-e Khamir (ca. 26º59'N, ca. 55º35'E); BWC2000-6, uncatalogued, ?, ?, Khuzestan, Dez River (32º14.663'N, 48º20.112'E); BWC2000-15, uncatalogued, ?, ?, Khuzestan, stream south of Shushtar (31º41.867'N, 48º59.581'E).

Comparative material:- BM(NH) 1920.3.3:193-202, 12, 21.0-30.5 mm standard length, Iraq, Basra (30º30'N, 47º47'E); BM(NH) 1920.3.6:9-10, 3, 17.1-26.8 mm standard length, Iraq, Basra (30º30'N, 47º47'E); BM(NH) 1949.7.21:1-11, 12, 18.4-42.4 mm standard length, Iraq, Bahr-el-Milk (32º35'N, 43º50'E); BM(NH) 1974.5.22:30, 1, 35.2 mm standard length, Iraq, Basra (30º30'N, 47º47'E); BM(NH) 1974.5.22:31-32, 2, 37.4-44.4 mm standard length, Iraq, Al Faw ().

Aphanius ginaonis
(Holly, 1929)

Male

Common names

gour-e khar, kopurdandan-e Genu or kapurdandan-e Geno (= Genow tooth-carp).

[Holly's pupfish, Genow pupfish, Gheno pupfish].

Systematics

Cyprinodon ginaonis was originally described from "Hei$e Quelle vom Djebel Ginao nordlich von Bender Abbas, südöstliches Persien". This is the Ab Garm-e Ganow at 27°26-28'N, 56°18-20'E, north of the Iranian port of Bandar Abbas at the Straits of Hormuz.

Berg (1949) places this species in the synonymy of Aphanius dispar stoliczkanus (see A. dispar above) and Villwock et al. (1983) regard it tentatively as a synonym of Aphanius dispar. Wildekamp (1993) is of the opinion that it may be a subspecies of Aphanius dispar. Hrbek and Meyer (2003) using mtDNA found this species to be deeply nested within the A. dispar clade. Villwock (2004) has used cross-breeding experiments that demonstrate this taxon and A. dispar are comparable to intraspecific crosses in other taxa. However, fin ray counts are non-overlapping and Reichenbacher et al. (2007; 2009) demonstrate differences in otolith morphology and affirm its validity as a species. Holly (1929b) reports a specimen of Cyprinodon dispar from the same spring. I was unable to verify the co-occurrence of two species in the hot spring through my own samples. Holly's specimen of A. dispar is in the Naturhistorisches Museum Wien (NMW 13799) but there are no field notes to confirm its locality with accuracy. It may have been collected in an adjacent stream. A. ginaonis is regarded here as a good species on morphological grounds (see below).

Four syntypes are in the Naturhistorisches Museum Wien (NMW 13800-13803). Holly (1929a) in his original description refers to 3 syntypes. One of these 4 fish was selected as a lectotype by F. Krupp (NMW 13800) and measures 21.8 mm standard length while the remainder measure 17.8-23.2 mm standard length.

Key characters

This species is distinguished from Aphanius dispar, the only other tooth-carp in this part of southern Iran by its locality and by having fewer total dorsal fin rays (5-7 as opposed to 8-11, mostly 9-10 in A. dispar from neighbouring drainages).

Morphology

Total dorsal fin rays 5(16), 6(38) or 7(6). Gut coiling is highly variable between individuals and is complex (illustrated in Coad (1980b)). Reichenbacher et al. (2007) give a description of otolith morphology. Chromosome number 2n=48, karyotype 14Sm + 34St and arm number 31 (Esmaeili et al., 2008). Esmaeili and Gholami (2007) give details of scale ultrastructure.

Saefali (1999) has D4-7, Pelvic5-7, Pect 13-17, A6-10, ll25-32,vert 26-30, bars10-22, gr13-21 SEE DATA SHEETS

Sexual dimorphism

Females have a higher mean numbers of anal fin rays, total vertebrae and flank bars, and longer pelvic fins (Coad, 1980b). Seifali and Sheidai (2001) found rostrum to pelvic fin distance, rostrum to anal fin distance, weight, total distance (? total length), head depth, head distance (? length) and body depth to differ between the sexes. Esmaeili and Gholami (2007) found that female scales were larger than those in males.

Colour

Males have a brighter colouration than females although colouration is individually variable within sexes. The back is mottled black. The dorsal, anal, pectoral and pelvic fins are a light orange. Flank bars are alternately black tinged with orange and grey-white, light blue or light orange. The side of the head anteriorly is iridescent light blue to green and the opercle is dorsally orange, postero-ventrally blue. The iris is iridescent green and the anterior ventral head surface is light blue. The peritoneum is black.

Preserved specimens have two bars on the male caudal fin with the fin margin hyaline. These bars are usually straight. Some fish have 1-3 bars or a forked bar. Females lack these bars and their caudal fin rays are lightly speckled. The male dorsal fin has 4-5 horizontal rows of spots, fading dorsally. These are not always well-developed and then consist of irregular, black pigmentation on the rays and membranes. The dorsal fin pigmentation in females is weak. The anal and pelvic fins have little or no pigment. The pectoral fins of males carry pigment concentrated on the rays, weakly speckled in females. Flank bars in females vary from weak to obvious, the light intermediate bars not as highly coloured as in males nor as well defined.

Size

Reaches 4.5 cm total length.

Distribution

This species was described by Holly (1929a) from the Ab Garm-e Ganow at 27°26-28'N, 56°18-20'E (various reports give differing latitude-longitudes for this well-known locality, hence the range). Ab Garm literally means "hot water" in Farsi and is used to denote a hot spring. Ganow means "foul water". The spring lies at an altitude of about 135 m (Moghadam, 1974) on the slopes of the Ganow Mountain (= transliterated as Ginao in German), hence its scientific name.

Zoogeography

The occurrence of sympatric A. dispar would confirm the species distinction of this species.

Habitat

The species is common within the bounds of the hot spring stream, numbering in the thousands in the 1970s, but no population trends have been recorded (but see below). Fish are found particularly along the stream margin and in many minor subsidiary springs which emerge a few metres from the main spring. Some fish are found in water as shallow as 1 cm but temperatures are 37-40°C (when shade air temperature was 20°C at 1400 hours on 27 January). The main spring itself issues from the ground at 30 litres second-1 and 41°C and drains as a stream about 10-15 m wide. A fault, over which the spring stream pours, isolates the fish from those downstream. Coad (1980b) summarises water chemistry and Reichenbcher et al. (2009) give a recent description of the habitat. The water is clear and colourless but there is a strong sulphur odour. The stream bed is composed of stones and pebbles covered by lime-green to dark blue-green algae. Moghadam (1974) gives details of the algae species.

Age and growth

Essentially unknown although fish are larger later in the year (11.8-31.4 mm standard length in January and 15.6-34.7 mm in March). Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 33 fish measuring 1.70-3.17 cm standard length. The a-value was 0.0210 and the b-value 3.375 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases).

Food

Gut contents are the lime green algae typical of the hot spring and chironomids.

Reproduction

Temperature is constant and daylength may then be important in triggering reproduction, although fishes caught on 27 January and 21 March have mature eggs of similar size (ca. 1.6-1.7 mm) suggesting an extended reproductive period.

Parasites and predators

Unknown.

Economic importance

None except related species are important in the aquarium trade and this is a potential economic use.

Conservation

Coad (1980b) listed this species as rare although it should be classed as vulnerable under the revised IUCN Red List Categories (IUCN, 1996) according to Coad (1998a).

The site is easily accessible by an asphalt road and is close to the large city of Bandar Abbas. Two hammams (bath houses) are in operation, one for men and one for women, the spring water is drained off to irrigate date palms, and local people also use the site for bathing, washing kitchen utensils and food containers, and washing clothes so there is input of soap and food debris (personal observations; A. R. Zinaei and H. R. Esmaeili, Payam-e Noor University, Bandar Abbas, 6 August 1997). The area is not under direct management by environment officials although in 1999 press reports referred to planned research studies aimed at preventing extinction of this taxon (IRNA, 29 September 1999) and the spring does lie in the Genu Protected Area (Biosphere Reserve) described by Zehzad et al. (1997).

The habitat could deteriorate if too much soap and food debris enter the stream. A single catastrophic event, such as a chemical spill, water diversion or construction activity, could eliminate the species. Recent construction has severely affected the population with only 1-2 specimens being caught and the population being replaced by A. dispar (H. R. Esmaeili, pers. comm., September 2007).

Reichenbcher et al. (2009) suggest that A. dispar specimens have been added to the spring by local people to increase the population size. Otolith differences between samples taken in 201 and 2009 suggest hybridisation.

Local people using the spring should be educated on its ecological significance and alternative facilities for bathing and washing established. Roadside access to the spring should be prevented. The population structure and reproductive regime of the fishes could be analyzed to determine if they are susceptible to abrupt changes.

Further work

This species differs from A. dispar in a number of characters but this may merely be a consequence of the unusual environment of the hot spring. Cross-breeding experiments would be very revealing and would demonstrate or disprove reproductive isolation and hybrid fertility. The population is very important as the only species in a unique habitat, lending itself to studies of speciation, adaptation and variation in response to high temperatures.

Sources

Type material: see above, NMW 13800-13803.

Topotypes: CMNFI 1979-0175, 77, 11.8-31.4 mm standard length. check size range; CMNFI 1979-0416, 7, ? mm standard length, Hormozgan, Ab Garm-e Ganow below falls (ca. 27º26'N, ca. 56º20'E); CMNFI 1979-0417, 44, ? mm standard length, Hormozgan, Ab Garm-e Ganow, side springs and edge of mainstream (ca. 27º26'N, ca. 56º20'E); CMNFI 2007-0054, 13, ? mm standard length, Hormozgan, Ab Garm-e Ganow (ca. 27º26'N, ca. 56º20'E);

Aphanius isfahanensis
Hrbek, Keivany and Coad, 2007

Common names

كپوردندان اصفهان (= kapurdandan-e Esfahan or Esfahan tooth-carp).

Systematics

The holotype is in the Canadian Museum of Nature, Ottawa under CMNFI 2004-0001, male, 25.0 mm SL, Esfahan Province, Zayandeh Rud (Zayandeh River) at Varzaneh Bridge, 32º25'32"N, 52º39'14"E, 1 July 2002, Y. Keivany and S. Asadollah and paratpes are under CMNFI 2004-0002, 18 males, 20.8–30.9 mm SL, 18 females, 22.0–38.4 mm SL, of 49 total (13 not used in meristic and morphometric analyses, total range in size 12.1-38.4 mm standard length), same locality as holotype; AMNH 233639, 1 male, 25.2 mm SL, 1 female, 21.6 mm SL, same locality as holotype; MRAC 2004-05-P-01-02, 1 male, 25.2 mm SL, 1 female, 21.6 mm SL, same locality as holotype; GenBank accession numbers AY593488, AY593489, AY593497, and AY593498.

Key characters

Distribution and colour pattern identify this species. Males can be distinguished from those of all other Iranian species by having distinct black edge on the dorsal, anal, and pelvic fins. The dorsal fin is covered with a high density of black blotches. Females can be distinguished from females of A. sophiae and A. vladykovi by having flank-bars rather than spots. It can also be distinguished from A. persicus by less well-defined bars terminating at a mid-flank stripe and a relatively light gray stripe at the caudal-fin base rather than a black spot or blotch. Flank-bars are also characteristic of females of Aphanius dispar, however.

Morphology

Morphometric data are given by Hrbek et al. (2007) but these characters cannot be used for a simple distinction from other Aphanius in Iran but only in multivariate space. This species is clearly distinguished at the genetic level from all other species of Aphanius. It has 82 molecular apomorphies – 19 transversions, two transversions/transitions (depending on comparison), and 61 transitions – that show fixed character state differences to homologous characters analyzed in A. sophiae, A. persicus, and A. vladykovi from Iran. Thirty-seven of these character states are also apomorphies when compared to A. anatoliae, A. danfordii, A. villwocki, A. asquamatus, and A. fasciatus from Turkey.

Dorsal fin rays 11-14, anal fin rays 10-13, pectoral fin rays 13-16, pelvic fin rays 4-6, lateral line scales 25-29, total gill rakers 10-13, precaudal vertebrae 10-13 and caudal vertebrae 15-18. Chromosome number 2n=48, karyotype 12Sm + 36St and arm number 30 (Esmaeili et al., 2008).

Sexual dimorphism

See colour below.

Colour

Male flank bars number 8-12, mean 10.2, significantly less than in A. sophiae at 10-21, mean 14.4. The bars are broad with interspaces about equal or slightly narrower. The bars extend from behind the head to the tail. Anterior bars fade on the belly, whereas, posteriorly on the caudal peduncle, they encircle the body. Dorsally, the head is dark gray and the body is lighter but still heavily pigmented with melanophores; the belly lacks pigmentation. The sides of the head are densely speckled with melanophores, more thinly on the ventral side; in most specimens the chin is darker than the rest of the ventral head surface. The eye is bounded ventrally and postero-ventrally by a thin line of black pigment. The defining male coloration is the black margins of the pelvic, anal, and dorsal fins. The dorsal and anal fins may present a halo effect, the margins being so dark in relation to the rest of these fins. The tips and outer margin of the pelvic fin are blackish. The anal fin has a broad, blackish margin with the rest of the fin light cream-colored. The dorsal fin has the blackest margin. The rest of the dorsal fin is variably blotched, the blotches being much lighter than the fin margin. Most specimens have a contrasting pigmentless area just below the fin margin. The pectoral fin has sparse pigmentation along rays and ventrally on the interradial membranes but lacks the concentrated black pigmentation seen on the pelvic fins. The caudal fin rays and membranes are sparsely pigmented, and the whole margin may be blackish but in most fish pigment is restricted to the upper and lower margins, the lower margin only, or is absent. Large females have a grayish dorsal surface to the head, a lighter back and upper flank covered with scattered melanophores, and a mid-flank stripe terminating on the caudal peduncle in a blackish, short stretch covering up to three scales (Fig. 2b). This short stretch of pigment is present in all females, faint in some, rarely forming a blotch and in some small fish tapering anteriorly. The flank stripe may be broken into a series of blotches in some smaller females, or it may be continuous as in large fish. Starting anterior to the belly there is a ventrolateral series of thin bars (up to ten) separated by cream colored interspaces 1–3 bars wide. These are absent in some smaller females, which may only have blotches at various levels present in this region. Even some large females have faint flank pigmentation so that bars and the stripe are weakly expressed. At about the origin of the anal fin, the flank bars may continue onto the caudal peduncle in regular form or become irregular, breaking up into blotches. Anteriorly the bars terminate ventrally at about the level of the lower edge of the pectoral fin. The belly and lower head have sparse pigment although the chin and sides of head are speckled with melanophores. The eye is bounded ventrally and posteroventrally by a thin line of black pigment. Fins lack any distinctive pigment pattern. Fin rays are outlined with melanophores, and interradial membranes of the caudal and anal fins have melanophores at varying degrees of density. The dorsal fin has the most interradial pigmentation, particularly near, but clear of the fin base.

Size

Attains 38.4 mm standard length.

Distribution

The type locality is near the town of Varzaneh on the lower reaches of the Zayanadeh River, about 30 km upriver from the terminal sump, the Gav Khuni marsh.

Zoogeography

An estimate of a 4.8 MYA divergence of A. isfahanensis and A. sophiae + A. persicus is given by Hrbek et al. (2007). This divergence time is in accord with the hypothesis of a near-simultaneous diversification of ~5 MYA of organisms occupying different geological units of central Iran.

Habitat

The type locality had a water temperature of 27ºC, pH was 6.7, the water was brackish, conductivity was 10.9 mS, dissolved solids were 5450 ppm, dissolved oxygen was 12.3 mg/L, river width was 50 m, and capture depth was 0.5 m. Current was slow, and there was no cover. Gambusia holbrooki was captured at the same locality. Bleher (20110 notes  that the type locality was dry at Varzaneh Bridge and only found specimens in an artificial circulating spring channel 50 m long in southern Esfahan.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Gholami et al. (2009) record the trematode Diplostomum spathaceum from the eyes of this fish.

Economic importance

None at present but a potential aquarium species. Breeding and aquarium conditions are similar to A. mento although it is reputedly difficult to breed.

Conservation

Known only from the type locality, which is not protected, this species needs a conservation assessment.

Further work

The biology of this species requires study.

Sources

Type material: See above, CMNFI 2004-0001, CMNFI 2004-0002, AMNH 233639, MRAC 2004-05-P-01-02.

Aphanius mento
(Heckel, 1843)

Male

Common names

gour-e khar, kopurdandandar-e Arvand (= Arvand River tooth-carp), kopurdand-e Irani.

[batrikh qabras in Arabic; Persian pupfish, Persian minnow, Black Persian minnow, Persian killie, Arvand pupfish, chin killifish, pearl-spotted killifish].

Systematics

Lebias Cypris Heckel, 1843 is a synonym. Krupp (1984a) gives a detailed synonymy of this species but considered cypris to have priority. Klee (1967) and Lazara (1989) point out that Garman (1895), in a work dated in July of that year, as first revisor placed cypris in mento. This decision has priority over Gaillard (1895), a work with no exact date of publication, in which mento was placed in cypris, according to the Zoological Code of Nomenclature (Ride et al., 1985). The type locality for both Lebias mento and Lebias cypris is "Mossul" according to Heckel (1843b).

Hybrids with Aphanius dispar richardsoni are reported from Israel (Goren and Rychwalski, 1978).

Parenti (1981; 1984) places this species in the genus "Aphanius", i.e. distinct from true Aphanius without defining and naming a new genus. This genus is more closely related to the Anatolian Kosswigichthys Sözer, 1942 and the South American Orestias Valenciennes in Cuvier and Valenciennes, 1846 than true Aphanius. Hrbek and Meyer (2003) note that, based on their mtDNA study, this species diverged from the ancestor of the A. dispar clade at an early date, 26.87±1.31 MYA. They speculate that the ancestor of mento invaded the northern perimeter of the Arabian plate as it was abutting Laurasia some time after the separation of their eastern and western clades of Aphanius (see above) before the closing of the Tethys Sea. It later spread through the present-day Tigris-Euphrates and into the Levant and southern Turkey.

The 5 syntypes of Lebias cypris are in the Naturhistorisches Museum Wien (NMW 59598) with 1 specimen selected as a lectotype by Krupp and Schneider (1989), the other 4 being paralectotypes. The syntypes of Lebias mento are (NMW 62105 (4 fish), 59042 (5) and 22624-22632 (9). Eschmeyer et al. (1996) list NMW 21699-704 (6) and NMW 59832 (21) as possible syntypes. The Vienna catalogue lists 5 specimens of Lebias cypris and 8 of Lebias mento.

Key characters

The adult colour pattern is distinctive and dorsal fin ray counts are usually higher than in A. dispar.

Morphology

Scales along side of body 23-28, total dorsal fin rays 9-14, usually 12-14, total anal fin rays 9-13, pectoral rays 12-16, and pelvic rays 4-6, usually 5. Frontal scalation is E-type (Hoedemann, 1958). Flank scales are squarish to a vertical oval. The former shape has a vertical anterior margin which may be wavy or slightly convex, upper and lower anterior corners rounded but square-cut, dorsal and ventral margins parallel, and the posterior margin rounded. The oval scales have all margins rounded. The focus is central to subcentral posterior, circuli are fine although coarser on the posterior field, radii are restricted to the anterior field and are almost horizontal and parallel. There is no pelvic axillary scale. Total gill rakers 11-15, rakers being spinulose and reaching beyond the adjacent raker when appressed. Vertebrae 25. Teeth are tricuspid with a long and pointed central cusp (Goren and Rychwalski, 1978). There are 14-16 teeth on the lower and 10-12 on the upper jaw (Berg, 1949). The gut is s-shaped. Reichenbacher et al. (2007) give a description of otolith morphology. Chromosome number 2n=44 or 48 (Wildekamp, 1993; Klinkhardt et al., 1995).

Sexual dimorphism

See below under Colour.

Colour

Adult, breeding males are a dark blue-black to dark brown or almost black with iridescent blue-white to silvery spots regularly-arranged on the fins as curved lines, and irregularly on the body (sometimes as irregular vertical bars and sometimes the spots are vertically elongate) (Richter, 1989; preserved material from Iraq; Goren and Rychwalski, 1978). The edges of the gill cover are orange-red (Wildekamp et al. (1999). Male colour fades in low light conditions and in winter males have silvery flanks with a dark brown back. Spots are silvery-blue on the upper flank and are not as numerous as in the spawning male. Females are grey-brown or grey-white to silvery with large golden blotches or silvery to blue spots and dark dots. Scales along the mid-flank usually have a dark border (Wildekamp et al. (1999). Fins in females are hyaline. Body colour is reportedly heightened in brackish water (Grimes, 1974). The peritoneum is silvery with dense but fine melanophores.

Size

Reaches 8.4 cm total length or more (Güçlü and Küçük, 2008).

Distribution

Found in the Orontes (= Asi) and Tigris-Euphrates basins, the Levant in coastal and Dead Sea basins, western Jordan, and in southern Turkey in Mediterranean basins as well as in central Turkey. Tigris-Euphrates records are relatively rare (see map in Wildekamp (1993)). The species was described from Mosul in northern Iraq and there are records from the region of Basrah in southern Iraq near the border with Khuzestan. Wildekamp (1993) has one record apparently mapped in southern Iran near the border with Iraq. This is the only record from Iran although Abdoli (2000) indicates the Arvand River on a map.

Zoogeography

See Systematics above and under the genus.

Habitat

This species has been reported to survive in waters up to 38°C and as low as 0°C. pH at 7.0-8.5 has been reported as ideal for aquaria maintenance. Aquarists report it as a robust species favouring hard and alkaline water with some salt added (Thiermann, 1978; Kostich, 1979; Phillips, 1985). Grimes (1974) considers it to be quarrelsome in aquaria like other Aphanius and Allen (1988) also noted that males are very aggressive. It inhabits fresh or slightly brackish water of springs, streams and lakes, usually near shore where males establish territories in vegetation. It generally prefers habitats dense in vegetation.

Age and growth

Life span is about 3 years with maturity in aquaria at 4-6 months. The length-weight relationship in southern Iraq was W = 6.0001 x 10-5 L2.6403 for males and W = 2.3028 x 10-4 L2.235 for females. The average condition factor for males was 2.0499 and for females 2.3121. The condition factor increases with increase in length in females and decreases in males (Huq et al. 1977). Güçlü and Küçük (2008) examined a population of this species in a spring in the Mediterranean Sea basin of Turkey and found fish in age groups 0 to 7. They also give length-weight relationships and a von Bertalanffy growth equation.

Food

Al-Daham et al. (1977) found filamentous algae and diatoms were the most important foods in southern Iraq. Shafi and Shalli (1986) report a diet of beetles, ephemeropteran nymphs and algae in southern Iraq. Aquarium specimens prefer meaty foods such as brine shrimps over flake foods (Grimes, 1974) but flake food is taken (Allen, 1988). Curiously, Al-Daham et al. (1977) for their southern Iraqi fish in aquaria found this species to prefer plant food. Males approached food items individually and prevented other fish from taking the food. Females were also solitary though less aggressive than the males. However Gambusia holbrooki out-competed male A. mento for food, on one occasion even snatching a Gambusia embryo from the mouth of an A. mento. Güçlü and Küçük (2008) found their Turkish fish to feed principally on Gammarus sp and Palaemon sp.

Reproduction

Anderson (1966), Lutman (1982) and Richter (1989) observed spawning in aquaria. There is a dominant male which is more deeply and richly coloured than other males. The dominant male defends a spawning territory, fighting other males. Males will defend separate spawning mops if these are made available in aquaria. The dominant male will pursue a female and places his larger dorsal fin over the her back. Several spawnings will follow and females will spawn with more than one male. Spawning can occur near the surface. These fish can spawn at six months of age. Peak spawning in southern Iraq is April to June when only a small proportion of eggs in the single ovary contained fully developed eggs (Al-Daham et al., 1977). However, Shafi and Shalli (1986) state that it breeds in southern Iraq in May-July. There are up to 71 highly-adhesive, eggs of 170 microns diameter. Aquarium spawned eggs hatch in 7-14 days, depending on temperature, 10 days at 25°, 3 days in the 80s°F, or 6-7 days in the low 70s°F (accounts vary). Allen (1988) found egg production to be higher in aquaria when the equivalent of seven tablespoons of salt was added per gallon at a water temperature of 25°C. A minimum of 20 eggs per day were produced.

Byniak (1979) describes the effects of day length and temperature on the reproductive cycle of females in this species in Israel.

Parasites and predators

Unknown.

Economic importance

An aquarium fish with breeding details given by Zipay (1961), Anderson (1966), Lutman (1982), Richter (1989), Semeit (1999) and www.killi-data.org/ (downloaded 21 September 2007). The species is hardy and fry are easily raised, given large enough tanks and an easily defensible area for the male to guard eggs. However, populations stop breeding in aquaria after 6-8 generations (www.killi-data.org/, downloaded 21 September 2007). Blaustein and Byard (1993) show that this species will prey on mosquito larvae in a laboratory situation and has potential in control of this vector of malaria, at least in vegetated habitats.

Conservation

The presence of this species in Iran needs confirmation by field work and until this is done and its distribution, population numbers and biology worked out, its conservation status cannot be assessed. Endangered in Turkey (Fricke et al., 2007).

Further work

See above.

Sources

Iranian material: None.

Comparative material: BM(NH) 1920.3.3:203-222, 22, 24.9-36.0 mm standard length, Iraq, Basra (30º30'N, 47º47'E); BM(NH) 1949.7.21:12, 24.4 mm standard length, Iraq, Bahr-el-Milh Lake east of Karbala (32º35'N, 43º50'E); BM(NH) 1981.10.6:19-23, 5, 26.6-31.8 mm standard length, Iraq, Qarmat Ali, Basra (30º30'N, 47º47'E); BM(NH) no catalogue number, 4, 19.7-31.7 mm standard length, Iraq, Basra Liwa (no other locality data); uncatalogued, 1, 31.2 mm standard length, Iraq, Khawr az Zubayr (no other locality data). ?Iraq fish from Hussain

Aphanius mesopotamicus
Coad, 2009

 

 

Male paratype 21.7 mm SL above, female holotype below

 

 

 

From Basrah, Iraq (ZISP 25393) after Berg (1949). Female (above), 30 mm, and male, 29 mm.

 

Common names

None.

[Mesopotamian tooth-carp].

Systematics

The holotype is a female, 29.3 mm SL from Khuzestan, canal branch of Karkheh River, 31º40’N, 48º35’E (CMNFI 1979-0360A) and paratypes are 37 specimens, 14.6-29.1 mm SL, same locality as above (CMNFI 1979-0360B) and 6 specimens, 15.1-20.5 mm SL, from Khuzestan, Karkheh River branch at Abdolkhan, 31º52’30”N, 48º20’30”E (CMNFI 1979-0364). The species is named for Mesopotamia (the land between the rivers) referring to the Tigris-Euphrates basin where the species is found.

Key characters

This species is distinguished by distribution and by pigmentation, males having clear margins to the unpaired fins, no bars on the caudal fin and 10-15 clearly defined flank bars, females bear irregular blotches or spots on the flank.

Morphology

Total dorsal fin rays 11-13, total anal fin rays 10-12, branched pectoral fin rays13-15, branched pelvic fin rays5-6, lateral series scales 26-29, total gill rakers 10-14, precaudal vertebrae 11-13 and caudal vertebrae 14-17.

Sexual dimorphism

See colour description.

Colour

Males have clear margins to the unpaired fins, no bars on the caudal fin and have 10-15 clearly defined flank bars. The dorsal surface of the head and the upper flank are more heavily pigmented with melanophores than more ventral areas. The belly and lower head are unpigmented. The chin and snout have dense melanophores and a rim of melanophores underscores the eye in both sexes. The dorsal, anal and caudal fins in males have wide clear margins. This is also seen in the material from Basrah, Iraq (BM(NH) 1982.9.2:326-328). The material from Basrah figured by Berg (1949) has distinct light margins in some printed versions of the figures. The caudal fin is darker just proximal to the clear margin, lighter in mid-fin and dark again at the base. The dorsal fin has irregular pigmentation on the membranes and, to a lesser extent, on the rays. The pigmentation may involve an overall darker colour in contrast to the light margin or may have some pattern to it. The pattern is often elongate and short blotches with no regular arrangement and sometimes may appear as up to 5 wavy and oblique bands. Dark pigmentation is found just behind the first ray on the fin membrane. The anal fin is darkest just proximal to the clear margin. Up to the last 6 membranes of the anal fin are dark and this pigment may be broken up in as many as 4 elongate bars along each membrane. A similar pattern is found in some dorsal fins and the general effect on both fins is that the postero-dorsal (anal fin) and postero-ventral (dorsal fin) parts of these fins are the darkest. The dorsal, anal and caudal fins generally have more pigment on the membranes than the rays and in some this is quite distinctive, making the rays stand out.

The pectoral and pelvic fins in males are generally clear or somewhat milky and opaque and lack melanophores. The distal parts of the membranes between the last 5 rays of the pectoral fin and the small membrane area of the pelvic fins can be pigmented. Males have flank bars circling the caudal peduncle and reaching the anal fin base but fading ventrally on the lower part of the anterior flank, not reaching the ventral margin of the belly and becoming progressively shorter and less distinct the more anterior they are. Bars are 2-5 times broader than the pale interspaces.

Females have a similar head and dorsal and ventral body pigmentation but it is much lighter than in males. Fins have little or no pigmentation. The proximal third to half of the dorsal fin rays and membranes, particularly the anterior ones, have pigment in some fish but this is weakly expressed compared to the condition in males. Some fish have a few faint melanophores lining the anal fin rays.

The most distinctive feature in females is a spot, oval to lozenge-shaped, at the central base of the caudal fin. Its concentration of melanophores is much higher than any other feature. The flank in females can have up to 14 thin, dark, wavy and irregular vertical patches of pigment. These patches may be interrupted in their vertical extent and are weakly expressed anteriorly. They fade ventrally, ending above the lower edge of the caudal peduncle and above the anal fin base, and are absent on the lower anterior flank. The patches are light and not as contrasting with the lighter interspaces as the bars found in males. Patches are thin, half to one third of the width of the interspaces. Very often the patches are broken up into spots and elongate blotches of various sizes, and a regular barred appearance is not usual. The spots and blotches are all smaller than the eye size by at least half. Material from Basrah, Iraq figured by Berg (1949) has the spots emphasised but material from Basrah (BM(NH) 1982.9.2:326-328) examined for this study has a more blotchy appearance and spots are not well-defined.

Size

Attains 29.3 mm standard length.

Distribution

Found in the southern Karkheh River basin of Iran and at Qarmt 'Ali at the northern part of Basrah on the Shatt al Arab, southern Iraq. Possibly also at Shalili-ye Bala or Shalili-ye Pa'in (ca. 31º58'N, 48º53'E), near Shushtar, Karun River basin of Iran from a record in Berg (1949) from Shellali (ZISP 25446).

Distribution map of Aphanius in Mesopotamia and adjacent areas. Solid squares type series of A. mesopotamicus (lower one CMNFI 1979-0360A and B, upper 1979-0364), open squares additional, non-type material of A. mesopotamicus (BM(NH) 1982.9.2:326-328 and ZISP 25393 (lower left) and ZISP 25446 (adjacent to type series)), open circles A. vladykovi, stars A. isfahanensis, solid circles A. sophiae, and diamond A. persicus (map modified after Hrbek et al. (2006)).

Zoogeography

This species is found in the Tigris River basin and its morphology shows evidence of relationship with species of internal basins in Iran.

Habitat

The habitat of the new species is known only from field notes made at the time of capture in Iran on the Khuzestan plains. The two localities are a river and a canal branching from that river. The 25 m wide river had a water temperature of 22°C at 1205 hours, pH 6.0, conductivity 1.0 milliSiemens, a mud bottom and the principal plant materials were rushes and reeds. The 30 m wide canal had a water temperature of 15°C at 0930 hours, pH 6.0, conductivity 1.8 milliSiemens, a mud bottom and the principal plant material was filamentous green algae.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

None recorded in Iran.

Economic importance

None at present in Iran but a possibly a species of interest to aquarists.

Conservation

This species is known only from museum collections and recent efforts to capture fresh material were unsuccessful. These attempts should be repeated and the numbers and biology of the species assessed.

Further work

The biology and continued presence of this species in Iranian waters requires study.

Sources

Based on Coad (2009).

Type material: See above.

Comparative material: BM(NH) 1982.9.2:326-328, 4, 20.2-25.3 mm SL, Iraq, Qarmat `Ali, Basrah (30°34’N, 47°46’E).

Aphanius persicus
(Jenkins, 1910)

Male

Female

check male has bars on tail!

Common names

gour-e khar, kopurdandan-e Irani (= Iranian tooth-carp).

[Persian pupfish or toothcarp, but see also A. sophiae].

Systematics

Cyprinodon blanfordii Jenkins, 1910 and Cyprinodon pluristriatus Jenkins, 1910 are potentially synonyms. All are synonymised with A. sophiae by authors (see Coad, 1996a). Hanko (1924) placed Cyprinodon blanfordi and C. persicus in Cyprinodon chantrei Gaillard, 1895 but this is an error.

Jenkins (1910) described Cyprinodon blanfordii ("East of Shiraz, South Persia"), C. persicus ("Spring on the edge of Shiraz Lake, Southern Persia") and C. pluristriatus ("East of Shiraz, stream running to Fussa, Southern Persia, 5,000 feet"). The first nominal species has so generalised a locality as to be of very uncertain provenance. It could conceivably be in the Shiraz valley, the Kor River valley or outside either of these (see below under A. sophiae where it is argued that is is a synonym of this species). The second nominal species is clearly from the valley of Shiraz and its salt lake Maharlu, an endorheic basin. The last nominal species was found near Fussa (= ? Fasa) and, based on maps, is probably in the drainage of the Mand River which flows to the Persian Gulf.

The valid name for Lake Maharlu basin Aphanius is given here as persicus. The types in Calcutta have not been examined by me. Cyprinodon blanfordii has page priority but the locality is vague and the fish have a pigment pattern description and illustration which is spotted on the flank with a lozenge-shaped spot at the caudal fin base, thus appearing to be a female A. sophiae. Cyprinodon pluristriatus appears last in the descriptions and may be an available name for Aphanius outside the Lake Maharlu basin in rivers draining to the Persian Gulf should these prove distinct.

Material described and illustrated by Villwock (1960) as A. sophiae are this species.

Three syntypes of Cyprinodon blanfordi (ZSI F9416 to ZSI F9418), two syntypes of Cyprinodon persicus (ZSI F9403 and ZSI F9404) and four syntypes of Cyprinodon pluristriatus (ZSI F9408 to ZSI F9411) are in the Zoological Survey of India, Calcutta (Menon and Yazdani, 1968), although Jenkins (1910) has ZSI F9408-9412 for C. pluristriatus.

Key characters

Females of this species are barred where all other Aphanius species in this area of southern Iran have spotted females. The distribution is apparently limited to the Lake Maharlu basin near Shiraz.

Morphology

Although populations are isolated in springs and streams around Lake Maharlu, Coad (1996a) found them to be relatively homogenous.

Lateral line scales 24(3), 25(34), 26(69), 27(176), 28(100) or 29(22). Scales above the lateral line 4-6, scales between lateral line and the anal fin 4-8, scales between the lateral line and the pelvic fin 5-9, and scales around the caudal peduncle 12-17. Esmaeili et al. (2007) determined a chromosome number of 2n=48 with 11 pairs of submetacentric and 13 pairs of subtelocentric chromosomes and an arm number of NF=70. Karyotype 16Sm + 32St and arm number 32 (Esmaeili et al., 2008).

Total dorsal fin rays 8(1), 10(8), 11(97), 12(204), 13(89), or 14(7); total anal fin rays 9(3), 10(111), 11(225), 12(64) or 13(3); total pectoral fin rays 13(9), 14(82), 15(211), 16(94), 17(9) or 18(1); total pelvic fin rays 4(11), 5(181) or 6(213); total gill rakers 9(5), 10(51), 11(220), 12(108), 13(19) or 14(3); abdominal vertebrae 9(1), 10(1), 11(27), 12(355), or 13(22); and caudal vertebrae 12(1), 14(16), 15(285) or 16(104).

Sexual dimorphism

Most apparent in colour and pigmentation detailed below. Females are longer and heavier than males (Esmaeili and Shiva, 2006).

Colour

"The basic colouration is bright shining silvery white. The entire body, with the exception of the dorsal crest and the belly, is tigered with small brown spots; in some specimens these spots seem to be arranged in three irregular longitudinal rows of which the uppermost extends as a band across the operculum and to the eyes; all fins are uniformly yellow with a reddish border. In males the spots are more dense, in females they are paler and scarcer, only the 2-3 at the base of the caudal fin are dark brown" (Heckel (1846-1849b) on A. sophiae). "The colour of this small fish with its enviable crystal teeth is uniformly brown, at least it is now in ethyl alcohol, with a silvery-white throat and belly; laterally on the tail there are a few scattered, darker spots, and a deeper black spot is located on the last scales anteriorly of the caudal fin. All fins are blackish without any markings" (Heckel (1846-1949b) on A. crystallodon). "Silvery, brownish on back, brown on top of the head, with numerous small spots of brownish, arranged in three or more, more or less irregular, longitudinal series, the median of which, on the lateral line, commonly ends in a black spot on the middle base of the caudal. On some this median series is entirely composed of blackish spots. Rarely the spots are somewhat confluent into longitudinal bands. Brownish specimens show a longitudinal band of silvery along the middle of the flank. The form described as A. crystallodon has spots only towards the base of the caudal, around the black spot usually ending the median series" (Garman, 1895).

The flank in females bears numerous alternating light and dark bars, the light bars varying in width from about one half to twice that of dark bars. The bars gradually merge with background pigmentation anteriorly on the flank and while clearly defined on the rear flank are difficult to distinguish anteriorly.

The caudal fin spot in females can be oval, tear-drop shaped or elongate but is usually in the form of a lozenge. Occasionally, single, smaller, dark, subsidiary spots may be found antero-dorsally and antero-ventrally to the basal spot or scattered spots may be found irregularly before and behind the basal spot.

Males have a pigmentation very similar to that of A. sophiae and the description here is identical. Some minor observed variation is attributed to variation in size and maturity of the fish compared. Males have light flank bars half the width or much narrower than alternating dark bars. The margins of the dorsal, anal and caudal fins are clear while the rest of these fins is dark. Some fish have up to 3, but usually 2, thin, light bars on the basal half of the caudal fin; these are generally larger fish. The margin of the lower half of the pectoral fin has concentrations of pigment on the membranes such that this area is darker than the rest of the fin. The anal fin is darkest posteriorly where pigment is concentrated on membranes. The distal third of the fin is pigmented to form a dark band, becoming lighter proximally. The dorsal fin is the darkest fin (except for the clear margin) and the anterior base is the darkest part of the fin. Bands are not always evident but pigment spots are large proximally. Some fish have 2, sometimes 3, thin light bars at the base separated by thin dark bars and paralleling the body profile while others have none. The dorsal fin base may have instead a series of lighter spots, sometimes irregular and not paralleling the body profile.

Size

Reaches 59.5 mm standard length (Esmaeili and Shiva, 2006).

Distribution

This species is restricted to the Lake Maharlu basin near Shiraz.

Zoogeography

This species is found only within the endorheic Lake Maharlu basin, Coad (1996a) suggests that tooth-carps in inland waters may have risen with the post-Pliocene uplift of the Zagros Mountains rather than being the result of relatively recent inter-basin dispersal.

Habitat

This species is found in fresh streams and springs and in springs of varying saline content or saline influence from hypersaline chloride Lake Maharlu. The lake at 124‰ is an impassable barrier to dispersal but is very shallow and dries out periodically, e.g. in 1967 (Cornwallis, 1968a). The larger springs and their streams can then meet and transfer fishes on the dried-out lake bed. Some springs are 27 km apart along the lake margin and have probably had no contact or fish exchange over many generations. This is especially the case for the smaller springs which emerge from the ground with a diameter of only 1 m and restricted flow. When lake water rises, lower springs are inundated with hypersaline water and the fishes are killed. The spring is presumably recolonised from a higher spring when lake level falls.

Fish placed in pure lake water die within minutes. Springs discharging directly into the lake contain tooth-carps but the fish do not venture into the lake. Some fish, if disturbed or deliberately harassed, will swim out for a short distance into the hypersaline lake water but they blanch and rapidly retreat to fresher water (Coad, 1996a).

Esmaeili and Shiva (2006) found the bottom of these water bodies to be muddy but the water was clear and slow-running. Conditions in October at three different sites were 16.9-19.0ºC, pH 6.70-6.74, dissolved oxygen 3.96-6.11 mg/l, nitrate 0.9-1.6 mg/l, nitrite 0.029-0.062 mg/l, phosphate 0.35-0.65 mg/l and ammonium 1.55-2.60 mg/l.

Age and growth

In the studies of Esmaeili and Shiva (2006), the sex ratio was 1.67:1 for females:males, highly significant, the age groups were 0+ to 3+ years, and positive allometric growth was determined (b value significantly greater than 3).The condition factor for females was highest in February and March and lowest in September while in males it was lowest in December and increased until February. Fishes smaller than 25 mm had an equal sex ratio, suggesting selective predation on males, or better survival or longevity of females. Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 62 fish measuring 1.86-4.27 cm standard length. The a-value was 0.0222 and the b-value 3.395 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases).

Food

Unknown but presumably similar to other tooth-carps.

Reproduction

Esmaeili and Shiva (2006) found the gonadosomatic index in females to increase from November to June, decreasing slowly from late June to November. The reproductive period lasted six months. Males had two peaks, in April and August. Egg diameters reached 1.71 mm, absolute fecundity ranged from 45 to 250 eggs (average 115.7 eggs) and relative fecundity was 21.6 to 244.1 with a mean of 90.01 per gram body weight. Even small fish had hydrated eggs and, with the extended reproductive period, shows adaptation to an unstable habitat. Monsefi et al. (2009) found that this species is a batch spawner in the Barm-e Shur spring with a spawning period from April to November. This species lives in unstable environments such as temporary lagoons or very small pools and batch spawning of relatively large eggs gives a greater chance of survival.

Parasites and predators

Mokhayer (1989) reported metacercariae of the eye fluke, Diplostomum spathaceum from Aphanius sophiae, probably this species, in Iran. The fluke can cause complete blindness and death in commercially important species of fish. This fish was also infested with yellow grub, Clinostomum complanatum. González-Solís et al. (1997) report Contracaecum sp. larvae from this species in the Lake Maharlu drainage, Fars.

Economic importance

This species has no current economic importance although it would make an excellent aquarium fish. Villwock (1959) gives details of maintaining what seems to be this species under aquarium conditions, gradually transferring them from a mixture of salty and magnesium sulphate water to stagnant tap water. They prefer some admixture of sodium chloride and magnesium sulphate which matches their natural habitat (2% synthetic sea water plus a tablespoon of epsom salts per 25 litres water). Aquatic plants such as Myriophyllum and Cabomba are also necessary for breeding as the eggs are attached to them but artificial substitutes can be used. Yellowish transparent eggs are 1.5 mm and are deposited usually individually, rarely in small groups. Larvae feed on Artemia salina nauplii and adults on daphnia, red mosquito larvae and tubifex. Male-male interactions include mock fights with spread fins and close swimming. The loser hides in the algae and is not strongly pursued by the victor. These interactions, and those between sexes, are not strongly developed and aquaria can be stocked at 15-20 fish per 25 litres. Y. Keivany notes (pers. comm., 2004) that this species is difficult to maintain in aquaria.

Conservation

The populations are found in a variety of springs and streams around Lake Maharlu. Some of these localities are threatened by water abstraction, road construction and pollution but a number of localities are small and/or saline and not under threat.

Further work

Studies on aquaria maintenance would enable it to become established in the aquarium trade and help its conservation.

Sources

Type material: None seen.

Iranian material: CMNFI 1979-0017, ?47, ?17.9-35.5 mm standard length, Fars, stream at Pol-e Fasa (29º29'N, 52º38'30"E); CMNFI 1979-0018, 75, ? mm standard length, Fars, Barm-e Shur (29º28'N, 52º41'30"E); CMNFI 1979-0033, 1, ? mm standard length, Fars, Barm-e Shur (29º28'N, 52º41'30"E); CMNFI 1979-0034, ?124, 14.8-38.7 mm standard length, Fars, spring on shore of Lake Maharlu (29º27'N, 52º44'E); CMNFI 1979-0035, 17, ? mm standard length, Fars, spring south of Barm-e Shur (29º28'N, 52º41'30"E); CMNFI 1979-0038, 16, ? mm standard length, Fars, spring near Barm-e Shur (29º28'N, 52º41'30"E); CMNFI 1979-0039, 91, 15.9-37.9 mm standard length, Fars, spring on shore of Lake Maharlu (29º27'N, 52º44'E); CMNFI 1979-0041, 61, 20.8-41.6 mm standard length, Fars, spring on shore of Lake Maharlu (29º23'N, 52º48'E); CMNFI 1979-0042, 28, ? mm standard length, Fars, spring on shore of Lake Maharlu (29º23'N, 52º48'E); CMNFI 1979-0046, 5, 18.1-32.2 mm standard length, Fars, qanat at Barm-e Dalak (ca. 29º35'N, ca. 52º38'E); CMNFI 1979-0047, 41, 11.3-32.1 mm standard length, Fars, spring source of Ab-e Paravan marshes (ca. 29º34'N, ca. 52º42'E); CMNFI 1979-0048, ?45, ?15.3-39.9 mm standard length, Fars, spring on shore of Lake Maharlu (ca. 29º32'N, ca. 52º48'E); CMNFI 1979-0049, 12, 17.2-38.5 mm standard length, Fars, spring on shore of Lake Maharlu (ca. 29º32'N, ca. 52º48'E); CMNFI 1979-0050, 27, 18.3-28.9 mm standard length, Fars, spring on shore of Lake Maharlu (ca. 29º31'30"N, ca. 52º49'30"E); CMNFI 1979-0051, 3, ? mm standard length, Fars, spring on shore of Lake Maharlu (ca. 29º30'N, ca. 52º52'E); CMNFI 1979-0052, 16, ? mm standard length, Fars, spring on shore of Lake Maharlu (ca. 29º30'N, ca. 52º52'E); CMNFI 1979-0064, 10, ? mm standard length, Fars, spring on shore of Lake Maharlu (ca. 29º31'30'N, ca. 52º49'30"E); CMNFI 1979-0065, 8, ? mm standard length, Fars, spring on shore of Lake Maharlu (ca. 29º31'N, ca. 52º50'E); CMNFI 1979-0066, ?31, 15.6-42.9 mm standard length, Fars, spring on shore of Lake Maharlu (ca. 29º32'N, ca. 52º48'E); CMNFI 1979-0110, 2, ? mm standard length, Fars, lake in Park-e Shahr, Shiraz (29º38'N, 52º32'E); CMNFI 1979-0112, 24, 11.9-45.9 mm standard length, Fars, stream at Pol-e Fasa (29º29'N, 52º38'30"E); CMNFI 1979-0118, 12, ? mm standard length, Fars, Barm-e Shur (29º28'N, 52º41'30"E); CMNFI 1979-0119, 75, ? mm standard length, Fars, steram at Pol-e Fasa (29º29'N, 52º38'30"E); CMNFI 2004-0006 (GenBank AY593484, AY593493), 15, ?22.2-41.8 mm standard length, Fars, Nasrabad Spring near Imamzadeh Ibrahim (29º35'09"N, 52º39'08"E);

Aphanius sophiae
(Heckel, 1849)

Male

Female

Common names

gour-e khar, kopurdandan-e safiyeh or kopurdandan-e sufieh (= sufi or sophy tooth-carp).

[batrikh sophiae in Arabic; Persian minnow, Cypris pupfish, but this seems inapt].

Systematics

The type locality of Lebias Sophiae is in "lauen Salzquellen bei Persepolis" according to Heckel (1846-1849b). Lebias punctatus Heckel, 1849 listed from "Nemek-Deria oder Salzsee, in welchen sich unter Schiraz die Quellen des Saadi ergiessen" or "the salty Maharlu Lake below Shiraz into which the Saadi springs flow" (Heckel, 1846-1849b) and Lebias crystallodon from "grossen Salzsee Nemek-Deria, unter Schiraz" (Heckel, 1846-1849b) are synonyms (Günther, 1859-70; Garman, 1895; Gaillard, 1895; Hanko, 1924; Berg, 1949; Coad, 1996a). The original catalogue handwritten by Heckel in Vienna gives the type locality as "Salzquellen bei Persepolis" which is in agreement with Heckel's publication. Lebias punctatus and Lebias crystallodon are described from localities in the Shiraz valley wherein lies the salt lake Maharlu. The catalogue is more equivocal for these two nominal species, listing only "Nemek Deria". The labels in jars containing L. crystallodon and L. punctatus were written under Franz Steindachner's curatorship and are not original (H. Ahnelt, in litt., 1987). Nemek Deria (= salt lake in Farsi) is a general term employed for the type of terminal water body seen in both the Maharlu and Kor River basins. The situation was further confused by Berg (1949) who listed the locality of punctatus as "Lake Niriz in Shiraz". Lake Niriz (= Neyriz or Bakhtegan) is the terminal sump for the Kor River.

All material of Aphanius from the Kor River basin examined fresh by me comprises fish with bars along the flank and fish with spots. The barred fish are males and the spotted fish females as confirmed by dissection. Males and females do not segregate in the field and it seems very unlikely that the material examined by Heckel from several collections would be comprised solely of males (I found sample sizes as low as 5 to contain both males and females). The males and females may have been separated and allocated different collection localities. The material described as L. punctatus is then female L. sophiae and I strongly suspect that it was misunderstood as coming from the Shiraz valley and the locality in Heckel (1846-1849b) is incorrect. All female Aphanius from the Shiraz valley are barred and the name punctatus is singularly inappropriate and would not be applied to them (see illustrations in Villwock (1960; 1977)). L. crystallodon could be a female L. sophiae with weak spotting on the flanks, again mislabelled as from the Shiraz valley. Günther (1866) considers crystallodon to be a post-spawning female of L. sophiae and Gaillard (1895) also considers it to be a female L. sophiae. Berg (1949) places L. sophiae as the male and L. punctatus and L. crystallodon the females of L. sophiae.

Cyprinodon blanfordii Jenkins, 1910 may well be a synonym of A. sophiae (see above under A. persicus for other views). The locality is vague, "East of Shiraz, South Persia" and could lie in the Lake Maharlu basin (occupied only by A. persicus, the females of which are barred on the flank), the Kor River basin (A. sophiae only), or outside either of these, in rivers draining to the Persian Gulf. The illustration shows a finely-spotted fish with a lozenge-shaped larger spot at the tail base, pigmentation found in A. sophiae. Cyprinodon pluristriatus Jenkins, 1910 is often referred to the synonmy of A. sophiae but is found "East of Shiraz, stream running to Fussa, Southern Persia, 5,000 feet". Assuming this to be Fasa and using topographic maps, the locality is in the Mand River basin draining to the Persian Gulf. The illustration and description refer to male fish and fall within the parameters of A. sophiae, a species thought to be restricted to an endorheic basin. It may be an available name for a taxon occurring outside the endorheic Kor River basin where A. sophiae is endemic.

Parenti (1981; 1984) places Aphanius sophiae in the genus "Aphanius", i.e. distinct from true Aphanius without defining and naming a new genus. This genus is more closely related to the Anatolian Kosswigichthys and the South American Orestias than true Aphanius.

The syntypes of Lebias sophiae are in the Naturhistorisches Museum Wien and include a wide range of material of which the following is part:-

Lectotype and paralectotypes (as labelled in 1997): NMW 14496 (6 fish measuring 25.8-30.4 mm standard length, one of which measuring 29.3 mm standard length is the lectotype as designated by F. Krupp, 24 October 1984 according to a jar label), NMW 22616-22623 (8, 26.1-29.0 mm standard length), NMW 60327 (8, 25.2-31.1 mm standard length), NMW 68283 (8, 18.9-27.6 mm standard length, partially dried and distorted), NMW 75067 (7, 24.7-27.4 mm standard length), and probable syntypes NMW 14760 (67) and NMW 15056 (7). The catalogue in Vienna lists 40 fish in one column and 12 fish in the adjacent column, the latter possibly meant to be the type series as it is written more boldly but which fish these were cannot now be ascertained. All these fish are males.

Eschmeyer et al., (1996) list syntypes under NMW 14496 (6), NMW 33616-23 (8) (not noted above), NMW 60327 (8), NMW 68283 (8), NMW 75067 (7) and also 9 fish in the Museum für Naturkunde, Universität Humboldt, Berlin under ZMB 31377 (not noted above). The Berlin material is listed as ex 16140, is all male and 20.3-26.7 mm standard length.

Syntypes of Lebias punctatus comprise the following material:-

Lectotype and paralectotypes (as labelled in 1997): NMW 76509 (7 fish, measuring 29.9-36.0 mm standard length, one of which measuring 36.0 mm standard length is the lectotype as designated by F. Krupp, 24 October 1984 according to a jar label), NMW 15070 (5, ca. 29.9-35.6 mm standard length, dried and distorted), NMW 15156 (14, 19.2-32.1 mm standard length), NMW 59837 (5, 26.6-35.7 mm standard length), and probable syntypes NMW 59609 (6, ca. 21.4-34.7 mm standard length, smallest dried). The Vienna catalogue lists only 12 fish opposite this name. These fish are all females.

The holotype of Lebias crystallodon measuring 40.1 mm standard length is NMW 15175 as the description and catalogue agree there is only one fish. The specimen is decoloured and in poor condition.

Key characters

The large, dark, usually lozenge-shaped spot at the caudal fin base in females with fine spotting on the flanks is distinctive for tooth-carps in this area of southern Iran.

Morphology

Lateral line scales 25(1), 27(7), 28(14), 29(17), 30(12) or 31(2). Scales above the lateral line 4-7, scales between lateral line and the anal fin 5-8, scales between the lateral line and the pelvic fin 6-9, and scales around the caudal peduncle 15-20.

Total dorsal fin rays 11(1), 12(11), 13(27), 14(12) or 15(2); total anal fin rays 10(1), 11(18), 12(27) or 13(7); total pectoral fin rays 14(1), 15(7), 16(31), 17(10), 18(3) or 19(1); total pelvic fin rays 5(9) or 6(44); total gill rakers 10(2), 11(28), 12(21) or 13(2); abdominal vertebrae 11(11), 12(37), or 13(5); caudal vertebrae 15(3), 16(33) or 17(17); and total vertebrae 27(8), 28(29) or 29(16).

Cephalic sensory pores are reduced to a series of neuromasts. Older literature on chromosome number was 2n=48 but is based on fish from outside the Kor River basin to which the species is restricted here (Klinkhardt et al., 1995). Esmaeili et al. (2007) determined a chromosome number of 2n=48 with 14 pairs of submetacentric and 10 pairs of subtelocentric chromosomes and an arm number of NF=76. Karyotype 8Sm + 40St and arm number 280 (Esmaeili et al., 2008).

Sexual dimorphism

Males have more bands on the flank and are darker and more intensely coloured than females. Male dorsal, anal and caudal fins are a deep black with narrow, silvery-white borders and the anal fin has a few silvery spots at its base. Females have brown dorsal, anal and caudal fins with 3-4 rows of black spots and the anal fin also has 2 cross rows of white silvery spots near the base. The caudal fin spot in females can be oval, tear-drop shaped or elongate but is usually in the form of a lozenge. Occasionally, single, smaller, dark, subsidiary spots may be found antero-dorsally and antero-ventrally to the basal spot or scattered spots may be found irregularly before and behind the basal spot. Male flank bars 10(4), 11(1), 12(2), 13(3), 14(5), 15(4), 16(6), 17(3), 18(1) or 21(1).

Eleven of 21 morphometric characters are significantly different between the sexes including head length, head and body depths, fin lengths, and caudal peduncle shape (Coad, 1998i).

Colour

"The body colouration is basically dark brown; the body, with the exception of the forward dorsal portion and the belly, is marked by 12-17 white, silvery shining vertical lines or narrow bands. Those individuals which have the most bands and are also darker and more intensively coloured seem to be males; their vertical fins are deep black with narrow silvery-white borders, and only the anal fin has a few silvery spots at the base. The lighter individuals which are also slightly higher and wider, we believe to be the females; they have brown vertical fins with 3-4 cross rows of black spots; their anal fins are marked with 2 cross rows of white silvery spots at the base" (Heckel, 1846-1849b).

The caudal fin spot in females can be oval, tear-drop shaped or elongate but is usually in the form of a lozenge. Occasionally single, smaller, dark, subsidiary spots may be found antero-dorsally and antero-ventrally to the basal spot or scattered spots may be found irregularly before and behind the basal spot.

The flank spots in females are much lighter than the caudal base spot and are from a half to much less in size, usually much smaller than half size. Spots are usually rounded but can be oval and vertically elongate. Spots are independent of scale arrangement on the lower half of the flank, particularly posteriorly and may form 2-3 poorly defined to distinct longitudinal rows. Upper flank scales have a crescent of pigment in mid-scale, leaving the margin and anterior base mostly free of pigment, or pigment may fill each exposed scale surface almost entriely except for the posterior margin, and thus appear as spots. Females have purplish tints on mid-flank and yellowish tints on the lower flank, upper flank, head top and chin.

Males have a pigmentation very similar to that of A. persicus and the description here is identical. Some minor observed variation is attributed to variation in size and maturity of the fish compared. Males have light flank bars half the width or much narrower than alternating dark bars. The margins of the dorsal, anal and caudal fins are clear while the rest of these fins is dark. Some fish have up to 3, but usually 2, thin, light bars on the basal half of the caudal fin; these are generally larger fish. The margin of the lower half of the pectoral fin has concentrations of pigment on the membranes such that this area is darker than the rest of the fin. The anal fin is darkest posteriorly where pigment is concentrated on membranes. The distal third of the fin is pigmented to form a dark band, becoming lighter proximally. The dorsal fin is the darkest fin (except for the clear margin) and the anterior base is the darkest part of the fin. Bands are not always evident but pigment spots are large proximally. Some fish have 2, sometimes 3, thin light bars at the base separated by thin dark bars and paralleling the body profile while others have none. The dorsal fin base may have instead a series of lighter spots, sometimes irregular and not paralleling the body profile.

Size

Reaches 5.6 cm total length.

Distribution

This species is restricted to the Kor River basin in Fars.

Zoogeography

As noted in the genus account, Aphanius species in Southwest Asia are regarded as relicts of the Tethys Sea. The mountain populations of A. sophiae (and A. vladykovi) may well have risen with the post-Pliocene uplift of the Zagros Mountains. Since Aphanius species have a low dispersal ability (Kosswig, 1967) it is considered unlikely that these small fishes dispersed from lowland populations into an existing, high mountain range represented today by the Zagros.

Bobek (1963) suggests that there may have been an outflow from the Kor River basin to the Persian Gulf at the south-east corner which was cut off at the end of the Pleistocene by alluvial fans. Krinsley (1970) maintains that any outlet was closed by the late Pliocene. Evidence for the isolation time of Kor River basin tooth-carps is equivocal.

Habitat

Tooth-carps identified as this species has been reported to survive in waters up to 39°C and salinities as high as 130‰. In aquaria lives at 5-37°C. It is found in springs, streams and small pools and lakes in fresh and saline waters.

Age and growth

Unknown. Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 64 fish measuring 2.18-4.64 cm standard length. The a-value was 0.0343 and the b-value 2.787 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases).

Food

Unknown in detail.

Reproduction

Unknown. Incubation is said to be 10 days at 25°C but this is a general citation for Aphanius spp. in aquaria.

Parasites and predators

Unknown.

Economic importance

This species has importance as an aquarium species, being colourful and hardy.

Conservation

Construction of irrigation canals and drying of some springs because of man-induced changes in the water table may threaten populations of this species.

Further work

The distribution of this species and its population numbers should be documented by field work. Molecular and chromosomal techniques would prove useful information for relating this species with others from neighbouring basins and throughout central Iran. Its biology is poorly known under natural conditions.

Sources

Type material: See above, Lebias sophiae (NMW 14496, 22616-22623, 60327, 68283, 75067, 14760 and 15056, Lebias crystallodon (NMW 15175), and Lebias punctatus (NMW 76509, 15070, 15156, 59837 and 59609).

Iranian material:- CMNFI 1979-0025, 1, 31.6 mm standard length, Fars, Kor River at Marv Dasht (29º51'N, 52º46'30"E); CMNFI 1979-0059, 30, 18.1-39.3 mm standard length, Fars, Pulvar River 8 km south of Sivand (30º01'30"N, 52º57'E); CMNFI 1979-0061, 3, 23.2-32.1 mm standard length, Fars, stream tributary to Pulvar River (30º04'N, 53º01'E); CMNFI 1979-0062, 6, 24.0-42.0 mm standard length, Fars, spring 17 km south of Sa'adatabad (30º05'N, 53º00'E); CMNFI 1979-0067, 1, 25.7 mm standard length, Fars, qanat at Zarqan (ca. 29º46'N, ca. 52º43'E); CMNFI 1979-0071, 6, ? mm standard length, Fars, qanat stream on road to Ramjerdi (ca. 30º00'N, ca. 52º38'E); CMNFI 1979-0091, 3, 23.2-32.1 mm standard length, Fars, stream tributary to Pulvar River (30º04'N, 53º01'E); CMNFI 1979-0117, , mm standard length, ; fish? CMNFI 1979-0155, 2, ? mm standard length, Fars, spring at Gavonoo (28º47'N, 54º22'E);ID? CMNFI 1979-0292, 6, 26.5-34.4 mm standard length, Fars, Lapu'i spring near Zarqan (29º48'N, 52º39'E); CMNFI 1979-0342, 1, 39.4 mm standard length, Fars, Kor River at Band-e Amir (29º49'N, 52º51'E); CMNFI 1979-0498, 6, 17.4-25.2 mm standard length, Fars, spring in Kor River basin (30º05'N, 52º27'E); CMNFI 2004-0003 (GenBank AY593483, AY593492), 47, ?20.7-28.8 mm standard length, Fars, Abdolmahdi Spring (30º06'12"N, 52º58'38"E); CMNFI 2004-0004 (GenBank AY593482, AY593492), 38, ?19.5-34.0 mm standard length, Fars, Malasskuh Spring (29º52'04"N, 52º29'20"E); uncatalogued (GenBank AY593481, AY593490), 6, 25.7-37.0 mm standard length, Fars, Dolatabad Spring (29º43'05"N, 52º50'11"E);

Aphanius vladykovi
Coad, 1988

Male

Female

Common names

gour-e khar, mahi-e gour-e khari (= "striped donkey" or zebra fish), kopurdandan-e Zagros (= Zagros tooth-carp), mahi parchami (= flag fish).

[Zagros pupfish].

Systematics

The type locality is in the Shahrestan-e Bakhtiari va Chahar Mahall in a large pool, 3 km west of Boldaji at 31°57'N, 51°01'E. The male holotype, 36.6 mm standard length, is in the Canadian Museum of Nature, Ottawa under CMNFI 1979-0247 with 35 male and 16 female paratypes from the same locality under CMNFI 1979-0247A and 1 male paratype from a stream 3 km east of Boldaji at 31°55'N, 51°05'E under CMNFI 1979-0248. Villwock (2004) has used cross-breeding experiments that demonstrate this species, A. sophiae and A. dispar are genetically isolated and distinct species.

Key characters

This species is distinguished from all other Aphanius in Iran and Southwest Asia by the higher number of normal-sized lateral line scales (36-47). Other members of Aphanius, and the related Kosswigichthys, have 35, usually 30, or less in the lateral series. The high number of scales is matched only by the genus Anatolichthys Kosswig and Sozer, 1945 of southwest Anatolia. Anatolichthys is polymorphic for scale counts which reach as high as 55 and shows scale loss, perhaps associated with the bitter, salty lakes in which it lives (Coad, 1988i). Male dorsal fin colour is much darker and the anal fin is light compared to A. sophiae, which most closely resembles A. vladykovi. Females lack the typical large spot at the base of the caudal fin found in A. sophiae.

Morphology

Scales are relatively small and numerous over the whole body as exemplified by scale counts. Scales are regularly arranged, embedded and imbricate. Anterior flank scales are a vertical oval with circuli and radii restricted to the posterior field. Numbers of circuli and radii are body and scale dependent and so are fewer in this species than in A. sophiae. There are up to 21 teeth in the upper jaw and teeth are in a single row in each jaw. Gill rakers are short, just reaching the adjacent raker when appressed. All fins are rounded distally and the anal fin is enclosed by a fleshy sheath, most apparent in large females but is also present in males. The gut has a single large loop. Cephalic sensory pores are reduced to a series of neuromasts. Keivany (2003) gives details of cephalic osteology. Pazooki et al. (2008) and Mardani Karani et al. (2008)  examined interlocality morphological variation, finding one of four populations to be more distinctive and 55 of 72 characters different respectively. Esmaeili et al. (2009) give a chromosome number of 2n = 48 and a karyotype of 8 submetacentric and 40 subtelocentric chromosomes with an arm number of 28.

Lateral line scales 36(2), 37(8), 38(15), 39(6), 40(10), 41(6), 42(3), 43(1), 46(1) or 47 (1). Scales above the lateral line 6-9, scales between lateral line and the anal fin 8-12, scales between the lateral line and the pelvic fin 11-15, and scales around the caudal peduncle 20-25.

Total dorsal fin rays 11(4), 12(21), 13(27) or 14(1), total anal fin rays 11(13), 12(25) or 13(15), total pectoral fin rays 14(4), 15(18), 16(25) or 17(6), total pelvic fin rays 5(9), 6(42), 7(1) or 8(1), total gill rakers 10(8), 11(41), or 12(4), and total vertebrae 27(2), 28(24), 29(26) or 30(1). Saefali (1999) records 8-11 gill rakers, 9-14 anal rays, and 9-15 dorsal rays.

Sexual dimorphism

Colour is sexually dimorphic and is outlined below. Ten of 21 morphometric characters are significantly different between the sexes including head length, head and body depths, fin lengths, and caudal peduncle shape (Coad, 1998i). Females attain larger sizes than males (76 mm versus 58 mm total length (Keivany and Soofiani, 2004a; b)).

Colour

Live adult males bear creamy bars on yellowish flanks. The pelvic fins and the edge of the caudal fin are yellow, the anal fin has an orange-yellow edge and the pectoral fins are orange-yellow. The dorsal fin is white with a wide blue-black band in the centre and a narrow blue-black band at the base. In some males the basal band is absent. Live adult females have a slight bluish tinge to their flanks and flank spots are brown.

In preserved specimens, the dorsal fin bands are dark and remain darker in preservative than the pigmentation seen in Aphanius sophiae which is more diffuse, tending to several horizontal rows of speckles, and which is not formed into a strongly distinct band or bands. Other fins are lightly pigmented, being finely speckled with melanophores with the dorsal and pectoral fins the darkest. The anal fin is fleshy and not hyaline but pigmentation is light and not dark as in A. sophiae. There is some tendency to formation of one or two narrow bars on the anal fin, paralleling the base, but these are weakly developed or absent in most fish. A mid-dorsal stripe is not apparent, perhaps because the dorsal flanks and back are darker than in females. Flank bars are light alternating with darker broad bars. The lighter bars become broader posteriorly. There is a light, wide bar at the base of the caudal fin which is not included in the count of male flank bars. Male flank bars 9(3), 10(8), 11(8), 12(10), 13(3), or 14(5). The sides of the head and upper and lower jaws are pigmented with scattered melanophores while the underside of the head is not. The belly is also free of extensive melanophores.

The most characteristic feature of females is the large number of scattered spots on the flank, extending from behind the head to the base of the tail. These spots do not extend to the dorsal flank or the back. The distinctive, large and often lozenge-shaped spot on the central caudal fin base of A. sophiae is absent. The upper back and flank are more heavily pigmented than the flanks (except for the flank spots). The lower surface of the head and the belly are melanophore free as in males and the sides of the head and the upper and lower jaws are heavily pigmented. The dorsal fin is finely pigmented on the rays and membranes and is generally darker than the caudal fin which is also finely pigmented. The other fins have very little pigment but all are fleshy and not hyaline. There is some suggestion of a thin horizontal stripe near the base of the dorsal fin in the largest specimens but this is never well developed.

Teeth are hyaline or discoloured with brown in patches. The peritoneum is dark brown.

Size

Reaches 7.6 cm (Keivany and Soofiani, 2004b).

Distribution

This species is found near Boldaji and in the Chaghakor Wetland in the upper reaches of the Karun River basin. Abdoli (2000) indicates its presence on a map in the upper Marun River and the upper Khersan River (the latter in the Karun basin).

Zoogeography

The origins of this mountain species have been discussed above under the account for A. sophiae.

It is found only in the uppermost reaches of the Karun River basin which drains to the head of the Persian Gulf. In this locality it is isolated from populations of Aphanius in the lowlands by a series of tangs which block gene flow. Oberlander (1965) in studying the origin of Zagros drainage patterns found that streams did not follow the line of least resistance but often incise the higher parts of mountain ridges which are anticlines composed of Miocene Asmari limestone. This situation arises because the Zagros ridges were eroded by a superposed drainage pattern already antecedent to the exhumation of overlying sediments of later Miocene marls and evaporites. The trend of the Zagros ridges is in a NW-SE direction and streams often cross this direction at right angles. This has resulted in the development of the tangs, narrow clefts in the Zagros ridges with very swift waters, which may be up to 2400 m deep with vertical sides rising from the water surface for 300 m, effective barriers to small fishes.

Habitat

This species is found high in the Zagros Mountains (the type locality lies at an altitude of about 2380 m). It is found in small pools, streams and marshes, in fresh water. The type locality is an artificially dammed pool about 300 m wide with cloudy water and a mud and pebble bottom. Capture depth was 40 cm. The shore was grassy and the pool contained large amounts of Myriophyllum and marginal rushes. At 1515 hours on a warm, windy and sunny day, water temperature was 29°C, pH was 6.5 and conductivity was 0.2 mS. A second locality in Coad (1988i) was a mud-bottomed stream 3-4 m wide with pools up to 1 m deep. At 1545 hours water temperature was 22°C, pH was 6.2 and conductivity was 0.45 mS. Current was slow and the stream had moderate amounts of Myriophyllum. Keivany and Soofiani (2004b) reporting on the habitat at the Madar-Dokhtar spring near Gandoman, found water conductivity of 240-280 μS, pH 6.9-8.5 and total dissolved solids125-138 mg/l. The bottom was muddy, interspersed with pebbles, and water is clear (but cloudy in the river). Myriophyllum and Potamogeton were used for spawning. Fish survive 30‰ and 30°C in aquaria.

Pazooki et al. (2008) found females to be more numerous than males, and the sexes lived in mixed schools. Occasionally larger males would chase smaller males but then both would rejoin the school. Males are territorial and defended against fish and beetles up to twice their size. The species preferred areas free from vegetation. These authors found the species easily tolerated to 33°C in aquaria for long periods, and the optimum temperature for the species is probably 21-24°C.

A mass mortality of this species was observed by A. Abdoli in late March or early April 1994 at Chagh-khor wetland near Boldaji.

Bagheri (1999) describes the macrofauna and environment of the Chaghakhour Lagoon where this species occurs.

Age and growth

Keivany and Soofiani (2004b) show maximum age is 2+ years. Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 319 fish measuring 1.46-4.64 cm standard length. The a-value was 0.0309 and the b-value 3.062 (a b-value < 3 indicating a fish that becomh increaseand a b-value >3 indicating a fish that becomes more rotund as length increases).

Food

Abdoli (2000) reports Daphnia and Gammaridae as food. The main food in the Madar-Dokhtar spring is Gammarus, but a specimen from a river also had some algae, Cyclops sp., snail eggs and fish larvae. Mouth structure would lead to the conclusion that this species is a surface feeder but it concentrated on the dominant crustacean in the Madar-Dokhtar spring (Keivany and Soofiani, 2004b).  Pazooki et al. (2008) found chironomids, dipterans, Diaptomus, Daphnia, diatoms and filamentous algae in guts and considered this species to prefer an animal diet.

Reproduction

Mature males chase the female in aquaria, pushing her into a corner, and showing pectoral fin flipping and shivering. Eggs were laid in a batch of 3-10. Absolute fecundity in the Madar-Dokhtar spring examined by Keivany and Soofiani (2004b) was 220-650 eggs, relative fecundity was 45-155 per gram body mass and working fecundity (free and ripe eggs) was 110 eggs absolute and 73 relative. Size of ripe eggs was 0.8-1.2 mm. Eggs are adhesive and attch to plants in small patches, from 3 to 30. Early April was the spawning peak. Eggs hatched at 9-13 days at 21-22ºC. Young reached 11.0-27.2 mm after three months in an aquarium.

Parasites and predators

Barzegar et al. (2004) examined this species for parasites in fish from the Beheshtabad river in Chahar Mahall va Bakhtiari Province and found Lernaea cyprinacea. Keivany and Soofiani (2004b) found it to be susceptible to trichodiniasis and ichthyophthiriasis in aquaria. Barzegar et al. (2008) record eye parasites from this fish including the digeneans Diplostomum spathaceum, Tylodelphys clavata and Ornithodiplostomum sp. Raissy (2008) also investigated parasites in this species from the Gandoman Lagoon. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Lernaea cyprinacea on this species.

Economic importance

This pretty little fish has great potential as an aquarium species (Coad and Keivany, 1998; Keivany and Soofiani, 2004b). It has been maintained in an aquarium for several months, tolerating temperatures of 30°C but probably favouring 20-22°C. pH values were 8-9. Water changes with their associated variations in conditions had no affect on the fish. The pupfish survived 3-4 hours longer than a Capoeta species when the latter was treated with formalin in a shared tank for a Saprolegnia infection and the tank was inadvertently left for 6-8 hours without a water change. Male pupfish are very aggressive and frequently attack other fishes of the same size or even larger. They ate all the coexisting guppy larvae in the aquarium, even those aged 2-3 weeks. The pupfish was fed crushed, dried freshwater shrimp and sometimes live shrimp although they had difficulty handling live shrimp. They were also observed biting and destroying vegetation in the aquarium, although it is unclear whether they actually ate any. It is now available in aquarium stores in Germany (Oliver Lucanus, pers. comm., 23 January 2004) and is kept during summer months in garden ponds, reproducing there successfully (Thomas Schulz, pers. comm., 14 January 2005).

Conservation

The distribution of the species is apparently quite limited, in waters neighbouring the towns of Boldaji and Shahr-e Kord. This should be confirmed by extensive field work in this poorly explored area of the Zagros Mountains. A limited distribution renders this species liable to significant loss or even extinction if habitats are disturbed or destroyed. The type locality, it should be noted, is an artificially dammed pond.

Significantly, a mass mortality of this species was observed by A. Abdoli in late March or early April 1994 at Chagha Khur wetland although the cause is unknown. M. Raissy (pers. comm., 1 July 2009) reports parasitic infections (7 species, including the digenean Ornithodiplostomum in most organs) as a cause for depopulation in Choghakhor (Chagha Kur) lagoon.

An additional threat is escapes of cultured rainbow trout (Oncorhynchus mykiss). Trout culture is being rapidly developed in this region and the trout are known predators on native fishes.

Further work

The distribution of this species in the Zagros needs to be clarified.

Sources

Type material: See above, CMNFI 1979-0247, CMNFI 1979-0247A and CMNFI 1979-0248.

Iranian material: CMNFI 2004-0005, ?, ?, mm standard length, ; uncatalogued (GenBank AY593486, AY593495), 4, 25.6-37.0 mm standard length, Chahar Mahall va Bakhtiari, Ebrahimabad (31º52'30"N, 51º10'10"E); uncatalogued (GenBank AY593487, AY593496), 2, 21.5-28.7 mm standard length, Chahar Mahall va Bakhtiari, Taqanak Bridge (32º12'35"N, 50º49'29"E); uncatalogued (GenBank AY593485, AY593494), 5, 31.0-49.8 mm standard length, Chahar Mahall va Bakhtiari, Madar-Dokhtar spring (31º52'12"N, 51º08'29"E).

Poeciliidae

Poeciliids, including the livebearers, are found in fresh and brackish waters from the eastern United States south through the Caribbean to northeastern Argentina and in Africa. There are about 37 genera with about 304 species (Nelson, 2006) and one or two species of livebearer has been widely introduced around the world as a control agent for malaria and is found in Iran, along with a released aquarium species.

This family is characterised by an anteriorly rounded and posteriorly compressed body and a depressed head; large cycloid scales on the head and body; supraorbital pores with neuromasts in fleshy grooves; a small, terminal mouth with the lower jaw projecting; teeth incisor-like or villiform on the jaws; gill membranes free from the isthmus; 4-6 branchiostegals; no spiny fin rays; a small dorsal fin; pectoral fins set high on the side (in contrast to Cyprinodontidae anterior pelvic fins; no adipose fin; and the anal fin in males is developed into a copulatory organ with rays 3-5 modified while females have a rounded anal fin (in livebearers only, the species found in Iran, in contrast to Cyprinodontidae).

The gonopodium is used for transferring sperm packets into the female. It is moveable to the side or forward to allow copulation to occur. The sperm packets release sperm when placed in the female and some can be stored for future use. Details of gonopodium anatomy are important in identifying and classifying species. The gonopodium is composed primarily of the third, fourth and fifth anal rays with various hooks, serrae and spines. Young are born alive, a condition known as ovoviviparity where the eggs develop and hatch in the mother.

Livebearers are found in habitats from mountain streams to brackish coastal marshes and river mouths. Food is mostly encrusting algae and the associated, small invertebrates. Males tend to be smaller than females and more brightly coloured. These colours are best seen during the courtship display.

Some species are important in the aquarium business, such as the guppy and swordtail, while the mosquitofish has been used world-wide as a predator on aquatic mosquito larvae, the adult fly being a carrier of malaria. They have also been used extensively in genetics research, research on tumours and in immunology. Some livebearers are all-female species and egg development is stimulated by spermatozoa from another species, without any genetic contribution. Young are identical to the mother. This unusual form of reproduction is called gynogenesis. Others have superfetation where eggs are at different developmental stages within the mother and are born over a period of several days rather than all at once.

Genus Gambusia
Poey, 1854

This genus is characterised by large scales, short dorsal and anal fins, the anal fin in males anteriorly placed and modified into a long intromittent organ (or gonopodium), dorsal fin inserted behind the anal fin origin (in contrast to Aphanius, Cyprinodontidae), and dorsal and caudal fins are spotted.

Gambusia affinis
(Baird and Girard, 1853)

Gambusia holbrooki was considered to be a subspecies of this species. Iranian mosquitofish were G. holbrooki but it is possible that some populations consist of G. affinis although none were seen by me or by Holčík and Oláh (1992). Shakirova and Sukhanova (1994) report this species from the Atrek River in Turkmenistan, which flows into Iran, as well as Kopetdag streams on the northern border of Iran. However identification of these taxa may be confused. The source of most (? all) Middle East mosquitofish is Europe and Kottelat and Freyhof (2007) record that there is no confirmed presence of G. affinis in Europe.

Gambusia holbrooki
Girard, 1859

Common names

گامبوزيا (= gambusia).

[gambuzi, zoory, or zurri in Arabic; gambuziya in Azerbaijan; gambuziya in Russian; mosquitofish; eastern mosquitofish (for holbrooki, western mosquitofish for affinis); plague minnow, in reference to its ecological impact].

Systematics

Gambusia holbrooki was originally described from eastern Florida and South Carolina, U.S.A. Wooten et al. (1988) distinguish the eastern and western mosquitofish in the U.S.A. based on morphology, biochemistry and distribution. However, extensive hybridization occurs in the native habitat and the composition of all the introduced Iranian populations has not been studied. Introduced populations world-wide are generally referred to the taxon holbrooki.

Key characters

Males are easily recognised by the anal fin rays 3-5 being specially modified into an elongate gonopodium for intromittent fertilisation. G. holbrooki has 8 dorsal and 11 anal total fin rays while G. affinis has 7 dorsal and 10 anal total fin rays (Walters and Freeman, 2000).

Morphology

Dorsal fin with 5-9 rays, usually 7 (with the last two counted as one), anal fin with 7-11 rays, usually 10, and pectoral fin with 11-14 rays. Lateral scale rows number 26-33. In males, the posterior edge of the joints of the first elongate ray is serrated (smooth in affinis), the cusps of the posterior branch of the second elongate ray are short and almost straight (long and curved in affinis), and the apical hook of the posterior branch of this ray is short with 2-3 joints (very long with 4-6 joints). Vertebrae 28-34. The karyotype is 2n=48. Detailed counts on Iranian specimens were not made; examination of cleared and stained material indicates that the species in Iran and Iraq is G. holbrooki.

Sexual dimorphism

Males reach a smaller adult size than females (see below). Males have the end of the anal fin base well ahead of the beginning of the dorsal fin. Females lack a gonopodium and the end of the anal fin is under the beginning of the dorsal fin. In addition various morphometric characters differ widely (see Abdurakhmanov (1962) for details).

Colour

Males are translucent grey to light olive with a blue, green or purplish sheen on the sides and opercle. The back is olive-brown to yellowish-brown and the belly silvery or yellowish. A dark bar passes through the eye. The iris has a purple sheen. The flanks may appear spotted as pigment margins the scales to form a diamond pattern on the body. Dorsal and caudal fins are spotted and a dusky, light tan but other fins are clear. Adult females have a large, triangular, bluish-purple blotch on the lower flank behind the pectoral fin (called the gravid spot). The black peritoneum can be seen through the body wall.

Size

Reaches 63.0 mm in females and 45.4 mm in males (Tabibzadeh et al., 1970a; Vargas and de Sostoa, 1996), perhaps to 8 cm in the largest females (Reshetnikov, 2002).

Distribution

The natural distribution is from New Jersey southward to northern Mexico but it has been introduced to all continents except Antarctica. This species was first introduced to the Ghazian marshes of the Caspian littoral of Iran in 1922-1930. Some mosquitofish may have spread from the Lenkoran District of Azerbaijan, reaching the Safid Rud in 1937 (Shukolyukov, 1949). From the Ghazian stock, additional samples were collected in 1966 for introduction around the country via raising ponds to over 3000 permanent water bodies (Tabibzadeh et al., 1970a, 1970b; Spillman, 1972; Emadi, 1996c). Many of these contained fishes but some were previously fishless or at least now appear to contain only mosquitofish, e.g. springs at Kahurak and Hormak in Sistan and Baluchestan Province. Over 1.5 million fish were distributed in 1969 alone (Tabibzadeh et al., 1970b). This species is now the most widely distributed fish in Iran (Abbasi et al., 1999; Kiabi et al., 1999; Abdoli, 2000).

Zoogeography

This species is an exotic, originating in eastern and southern North America and introduced to Iran from stocks introduced to Italy and from Baku in Azerbaijan.

Habitat

Mosquitofish are normally inhabitants of clear and weedy streams and ditches, weedy margins of large rivers and lakes, marshes and brackish coastal lagoons, usually over mud or sand bottoms. They prefer more open waters with less vegetation than Aphanius dispar, for example. Males and females in aquaria swam together in a mixed school in contrast to the sexually segregated schools of Aphanius dispar (Al-Daham et al., 1977). In the Anzali Mordab, dense schools are found in surface waters of areas covered by submerged and floating vegetation while deeper water is fishless (Holčík and Oláh, 1992). In Iran, it is one of two most abundant species in Caspian wetland areas along with Carassius auratus (Iranian Fisheries Research and Training Organization Newsletter, 19:4, 1998).

Al-Daham and Bhatti (1977) found G. affinis, probably this species, to have a complete tolerance of 10.25‰ sea water and 90% of fish survived 20.5‰ for 24 hours and even as high as 58‰. This species has been reported to survive in waters up to 42°C and as low as 0.5°C. Summer air temperatures of 45°C and winter temperatures several degrees below zero are survived as long as the water is deep enough (Tabibzadeh et al., 1970a). Upper and lower thermal tolerance increases with body size (Al-Habbib and Yacoob, 1993). At temperatures below 15-18°C, mosquitofish grow but do not mature or breed. They prefer a temperature of 31°C.

Muddy and polluted conditions, acid to alkaline water, and dissolved oxygen less than 1 mg/l are survived by this species. pH range survived is 4.46-10.2 and dissolved oxygen levels as low as 0.2-0.4 mg/l as long as water surface access is available. They can take advantage of the oxygen rich surface layer as they position their bodies with flattened head and back immediately adjacent to t\he air-water interface (Lewis, 1970).

Their success in Iran and other waters world-wide is attributed to the following factors according to Meffe and Snelson (1989):- 1) abundant in original range, 2) polyphagous, 3) short generation time, 4) a single female can colonise since livebearer, 5) broad physiological tolerances, 6) closely associated with man, 7) high genetic variability, 8) specialized reproduction with moderate numbers of advanced young several times a year, and 9) high aggression levels.

Age and growth

Numbers of each sex in a population vary between localities, mostly females predominate over males. Males are more sensitive to temperature extremes, starving and overcrowding and this will affect such things as size and age at maturity. Males do not grow much after sexual maturity is attained but females have indeterminate growth, although growth slows as energy is put into egg production. The average condition factor for males was 1.2369 and for females 1.0906 in southern Iraq. The condition factor slowly decreases as length increases in the male and vice versa for females (Huq et al. 1977). Sexual maturity can be reached in only 2-3 months but such fish die before winter. Minimal size for maturity is 16-28 mm total length. In the laboratory under ideal conditions sexual maturity can be attained in 3 weeks. Life span seldom exceeds 15 months, although the range is 6-24+ months depending on when in the spawning season the fish was born. In Spain, maximum ages are 2+ for females and 0+ for males (Vargas and de Sostoa, 1996). Fish born late in the year may not mature until the following year.

Growth to maturity can be so rapid that a 50-fold increase in numbers can occur over a 10-week period. Densities may reach over half a million fish per hectare. Under ideal conditions it is theoretically possible for 10 pregnant females to produce 5 million fish in 6 months.

Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 35 Iranian fish measuring 1.98-3.27 cm standard length. The a-value was 0.0190 and the b-value 3.214 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases).

Food

Generally this species is regarded as an opportunistic omnivore. Swanson et al. (1996) refer to this fish as an aggressive predator and Al-Daham et al. (1970) for southern Iraqi fish considered it to be a carnivore, feeding on aquatic and terrestrial insects including spiders, ants, beetles and mites, and also filamentous algae. Females are cannibals. Food includes diatoms, algae, worms, crustaceans including zooplankton, insects, snails, other fishes and amphibian larvae. The mosquitofish is dependent on sight to detect and attack prey. Selection of zooplankton can lead to an increase in phytoplankton numbers (and thus affect water quality) and to changes in size frequencies of zooplankton populations.

Food is taken at or near the surface and by grazing on plants and rocks. Tabibzadeh et al. (1970a, 1970b) record diet as molluscs, aquatic insects and various forms of mosquito larvae. They report up to 94 pupae or 104 mosquito fourth-stage larvae eaten per day by this fish and effective clearance of springs and marshes. Large-sized mosquitofish took more than 500 second instar mosquito larvae per day and medium- and large-sized mosquitofish consumed 22-64 fourth instar larvae per day in an Iraqi study under laboratory conditions (Mohsen et al., 1989). Abdoli (2000) lists Diptera and Chironomidae as food items. Meffe and Snelson (1989) report predatory behaviour on the young of other species of fish and females attack other fishes, shredding fins and sometimes causing mortality.

Reproduction

Na'ama and Al-Hassan (1989) report G. affinis (presumably G. holbrooki) to have a brood size up to 48 from Baghdad specimens and 51 from the Basrah area, Iraq. The smallest reproductive female was 1.7 cm total length. Weight is the most accurate predictor of brood size. In Iran, Tabibzadeh et al. (1970a) found that females mature and reproduce within 2 months at temperatures above 15°C. In less favourable conditions this may take 8-10 months. A minimum photoperiod of 12.5-13 hours is necessary to stimulate reproduction although year-round temperatures above 20°C can offset this light requirement. About 17 days after fertilisation, the female gives birth to as many as 428 live fish over a period of about 1 month after 3-8 weeks gestation. Each fertilisation can give 2-3 broods and each female can produce up to 9 broods during her life, although 2-5 is more usual.

The breeding season extends from April to November in favourable conditions, May to September in more temperate conditions. Several months pass between successive spawnings. Eggs are up to 1.8 mm in diameter when mature and embryos are about 6-8 mm at birth. The young are protected within the female and are independent when born, with resulting low mortality. Environmental breeding requirements are simple and with low mortality results in the successful spread of this exotic.

The small male approaches the female from behind and with a rapid motion inserts his gonopodium tip into the female. The gravid spot is a releasing stimulus. It indicates a receptive female and also is a target for the male gonopodium. Sperm are transferred in a spermatophore. During mating the gonopodium is angled forward at 140-150°.

Most males copulate without courtship. When courtship occurs it involves the male assuming an s-shaped position with its body and vibrating in front of, or at the side of, the female. Males are aggressive and dominate smaller males to restrict access to females. Females may be inseminated by several males and sperm can be stored to be used up to 10 months later. A single transplanted female can populate a new habitat.

Parasites and predators

Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Lernaea sp. on this species (as G. affinis).

Economic importance

Introduced widely to combat malaria by consuming the aquatic larvae and pupae of the carrier mosquito. However to be effective, this fish must have aquatic plants cleared from its habitat. The mosquitofish is also a predator on the eggs and young of other, native fishes and a competitor for food, and it alters ecosystems by greatly reducing rotifer, crustacean and insect populations allowing phytoplankton to increase dramatically (Myers, 1965; Hurlbert et al., 1972; Tabibzadeh et al., 1970a; Edrissian, 2006; Chandra et al., 2008). Native species might well be more effective predators on mosquito larvae (see account of Aphanius dispar and Ahmed et al. (1988) and Beidas and White (1982)). Muhaisen et al. (1985) studied control of this species in Iraq as larvae on fish farms and found potassium permanganate, copper sulphate, acriflavine and malachite green to be effective.

Haas and Pal (1984) point out that Gambusia may favour schistosomiasis, a human parasite, by altering the ecology of freshwaters, perhaps by reducing the numbers of natural predators on vectors of this parasite. They also note that it is considered a pest by fish farmers.

Tabibzadeh et al. (1970a, 1970b), Motabar (1978) and Swanson et al. (1996) give methods for raising and distributing mosquitofish. Also used as a research species and sometimes seen in aquaria. Al-Nasiri and Sharma (1978) used this species in Iraq to study the toxicity of the agricultural insecticide Ekatin 25 to fish and Mohsen et al. (1989) studied the predatory efficiency and tolerance to mosquito larvicides.

Conservation

The most widespread fish in Iran and, as an exotic, not in need of any protection. However this distribution undoubtedly affects the conservation status of native species since mosquitofish have a strong impact in small and non-diverse habitats such as the small springs found throughout Iran. Muus and Dahlstrøm (1999) mention that it tends to oust native toothcarps.

Further work

The effects of this species in competition with native fishes should be investigated although it is so widespread that it could never be eradicated.

Sources

Meffe and Snelson (1989) give the ecology and evolution of the family and Rauchenberger (1989) the systematics of the genus. Swanson et al. (1996) and Pyke (2005, 2008) review biology, culture and mosquito control and are a source for general information above.

Iranian material: The following is a sub-sample of Iranian material as this species is very common and material extensive (see map for distributional information): CMNFI 1970-0515, 2, 20.6-30.2 mm standard length, Gilan, Shafa River near estuary (37º35'N, 49º09'E); CMNFI 1970-0519, 5, 14.1-19.5 mm standard length, Gilan, Chelvand River (ca. 38º18'N, ca. 48º52'E); CMNFI 1970-0590, 3, 25.1-29.2 mm standard length, Mazandaran, Shesh Deh River near Babol Sar (ca. 36º43'N, ca. 52º39'E); CMNFI 1971-0343, 1, 26.0 mm standard length, Gilan, Langarud at Chemkhaleh (37º13'N, 50º16'E); CMNFI 1979-0019, 26, 15.4-29.0 mm standard length, Fars, Pol-e Fasa (29º29'N, 52º38'30"E); CMNFI 1979-0025, 2, 28.7-33.8 mm standard length, Fars, Kor River at Marv Dasht (29º51'N, 52º46'30"E); CMNFI 1979-0026, 1, 21.6 mm standard length, Fars, Shapur River at Shapur (29º47'N, 51º35'E); CMNFI 1979-0126, 53, 14.9-33.1 mm standard length, Fars, Dasht-e Arjan springs (29º37'N, 51º58'E); CMNFI 1979-0173, 13, 24.7-39.6 mm standard length, Hormozgan, qanat at Hajjiabad (28º19'N, 55º54'E); CMNFI 1979-0192, 2, 24.9-26.4 mm standard length, Fars, qanat 2 km east of Rostaq (28º26'30"N, 55º04'E); CMNFI 1979-0226, 65, 23.1-39.2 mm standard length, Sistan, 3 km east of Kuh-e Khajeh (30º57'N, 61º17'E); CMNFI 1979-0230, 36, 7.2-44.4 mm standard length, Sistan, Jehil-e Puzak (ca. 31º15'N, ca. 61º42'E); CMNFI 1979-0283, 1, 35.9 mm standard length, Kermanshahan, Qareh Su basin (34º21'N, 47º07'E); CMNFI 1979-0288, 7, 26.8-29.4 mm standard length, Ilam and Poshtkuh, Gangir River (33º50'N, 46º18'E); CMNFI 1979-0309, 5, 15.4-32.8 mm standard length, Kerman, Fahraj River (28º57'N, 58º42'E); CMNFI 1979-0328, 2, 24.3-24.8 mm standard length, Baluchestan, jube near Bampur (27º10'30"N, 60º21'E); CMNFI 1979-0360, 13, 17.8-25.9 mm standard length, Khuzestan, Karkheh River canal (31º40'N, 48º35'E); CMNFI 1979-0478, 2, 17.8-28.2 mm standard length, Mazandaran, ditch 29 km from Eimar (37º09'N, 54º36'E); CMNFI 1979-0498, 2, 22.1-35.8 mm standard length, Fars, spring on road to Dariush Dam (30º05'N, 52º27'E); CMNFI 1980-0122, 25, 14.3-26.8 mm standard length, Mazandaran, Nerissi River (36º38'N, 52º16'E); CMNFI 1980-0128, 7, 23.6-35.6 mm standard length, Mazandaran, Qareh Su (36º49'30"N, 54º03'30"E). CMNFI 1980-0132, 5, 18.4-30.2 mm standard length, Gilan, Safid River at Kisom (37º12'N, 49º54'E); CMNFI 1980-0145, 75, 32.6-40.4 mm standard length, Gilan, Bandar-e Anzali (37º28'N, 49º27'E).

Comparative material: BM(NH) 1974.5.2:38-40, 3, 17.3-25.1 mm standard length, Iraq, Baghdad (33º21'N, 44º25'E); BM(NH) no catalogue number, 6, 10.7-37.5 mm standard length, Iraq, Baghdad Liwa (no other locality data); BM(NH) no catalogue number, 4, 28.3-38.8 mm standard length, Iraq, Baghdad Liwa (no other locality data); BM(NH) no catalogue number, 8, 16.8-32.5 mm standard length, Iraq, Kut Liwa (no other locality data); BM(NH) no catalogue number, 2, 20.3-21.9 mm standard length, Iraq, Hilla Liwa (no other locality data).

Genus Xiphophorus
Heckel, 1848

This genus comprises the platyfishes and swordtails and is found in Mexico and central America with 25 or more species. these are small fishes, 14 cm or less, with several members and hybrids important in the aquarium trade. They are also used in genetic studies. They have a gonopodium in males as in other poeciliids with a unique structure in detail (Rosen, 1960). Male swordtails may have a characteristic elongate lower caudal fin lobe, the "sword" but this is not always developed in all species and all individuals.

Xiphophorus hellerii
Heckel, 1848

aquarium swordtails from Wikimedia Commons

Male and female specimens from Jahrom

Male and female specimens from Jahrom

Common names

dom shamshiri.

[swordtail, green swordtail]

Systematics

The trivial name is often spelled helleri.

Key characters

The swordtail is characteristic along with details of the gonopodium.

Morphology

Dorsal fin branched rays 11-15, anal fin branched rays 8-10, pectoral fin branched rays 12-13 and pelvic fin branched rays 6. Lateral line scales 26-30. Vertebrae usually 30-31.

The swordtail is medium to large with a long straight caudal appendage. The terminal segment of gonopodial ray 3 is produced into a crescent-shaped hook and blade, pointed distally  Ray 4a curves strongly backward over the blade at an angle greater than 90°. Distal serrae of ray 4p are reduced in size and number, and proximal serrae are rather slender; The terminal segment of ray 5a is produced into a claw, several times larger than the distal serrae of ray 4p.

scale rows along the middle of the body are green to blue; belly is white and there is a s in all populations (regional forms). In a few variants, there can be one or two additional lines above and one or two below this line. In almost all populations, the dorsal fin has spots or flecks of red. This coloration can also appear in the caudal fin. The sword of males is bright yellow and edged more broadly in black below than above. Old fish (3 to 4 years old) can have a totally black sword from the caudal fin all the way to the tip of the sword.

Sexual dimorphism

The male sword may not always develop in aquarium specimens but usually appears at 3-4 weeks of age. Mature females have a gravid spot in front of the anal fin.

Colour

The overall colour is olive-grey to olive-green. There is a light red, dark red dark, violet or dusky brown longitudinal line or rarely a zig-zag stripe on the greenish-yellow flank, and there may be 1-2 reddish stripes above and 1-2 reddish stripes below. Scale rows along the mid-flank are green to blue, edged with brown and net-like in appearance. The dorsal fin is spotted red or brownish in median and below the margin rows. Fins are generally a yellow-green. The belly is white. The sword is a bright yellow or orange-yellow edged more broadly in black below than above. The black lower edge is carried forward onto the caudal peduncle. Fish 3 to 4 years old can have a totally black sword. Aquarium specimens may be red, green, black and albino, and various combinations of these.

Size

Attains 16.0 cm total length, females larger than males.

Distribution

The native distribution is from southern Mexico to Guatemala, Honduras and Belize. Introduced specimens were caught in Iran at Jahrom (28º29'N, 53º31'E) and Kashan 34º04'N, 51º31'E) (Esmaeili et al., 2010).

Zoogeography

An exotic species presumably released from aquaria.

Habitat

In its native habitat this species is found in streams and rivers but can also occur in canals, ponds, lagoons, ditches and in warm springs. The Iranian specimens were caught in a small pool, part of a qanat system, and in a spring. Vegetation can be none to common. The water may be clear to polluted and  current none to moderate. Adults prefer current in streams while the young remain in quieter shore areas. It occurs at both low and relatively high elevations, up to ca. 1500 m. In aquaria this species prefers slightly alkaline, medium hard water and temperatures in the 22-28ºC range.

Age and growth

Sexual maturity is reached after eight to twelve months in some reports, three months in others. Life span exceeds 4 years.

Food

Food is crustaceans, insects, worms and plant matter.

Reproduction

This species is a livebearer and is easy to breed in aquaria. Females can produce more than 180 young at one time, at intervals of about a month. Adult males can be aggressive to other fishes. A well-developed male sword stimulates females to mature and inhibits maturity in younger males. The female swordtail can undergo sex reversal, becoming  male under certain environmental conditions.

Parasites and predators

None recorded in Iran.

Economic importance

None at present in Iran but a possible source of aquarium specimens. It is a popular aquarium species, easy to maintain and is sold world-wide and it has been used in genetic studies. There are various cultivated forms of this species, hybrids with related species. It does adversely affect native species when introduced.

Conservation

None required as an exotic. 

Further work

The biology of this species in Iranian waters, and its effects on native species, requires study. It should be extirpated, if possible.

Sources

General aquarium literature and Esmaeili et al. (2010).

Gasterosteidae

Sticklebacks are found in marine and fresh waters of the cooler parts of the Northern Hemisphere. There are about 8 species of which one is native to Iran and another has been introduced.

This family of small fishes is characterised by a compressed, fusiform body; teeth in bands in each jaw but none on the tongue or palate; a protractile mouth; 3 branchiostegal rays; no postcleithrum; no scales but a series of plates along the flank variably developed, sometimes absent; 2 or more (usually 3-16) isolated spines in front of a soft dorsal fin (usually 6-14 rays); and a pelvic fin with a strong spine and only 0-2 soft rays.

The species in this family have been studied extensively (Coad, 1981f; Paepke, 1983; Wootton; 1976; 1984; Paepke in Bănărescu and Paepke, 2002). There is variation in colour, body form, spine numbers and development, and plate numbers. These variations in anatomy are matched by variations in biology such as habitat, feeding and reproduction. Variation in behaviour, biology and in speciation makes these small fishes, which have no commercial value, particularly important. Some of the variation is owing to environmental factors while some has a genetic basis. Several books have been devoted to them and thousands of scientific studies. Sticklebacks make excellent aquarium fishes. Their reproductive behaviour is complex, involving courtship and nest building. Some populations are anadromous and enter fresh water to breed.

Genus Gasterosteus
Linnaeus, 1758

This genus is characterised by having a modal count of 3 isolated strong spines in front of the soft dorsal fin, the body usually has strong bony scutes along the flank developed as a keel on each side of the caudal peduncle, a strong spine in the pelvic fin, and a complex reproductive behaviour.

The threespine stickleback, and related populations which act as good species but have not been formally named, have been studied extensively for their reproductive and other behaviours and for the insights that morphological variation and genetics throw on evolution.

Gasterosteus aculeatus
Linnaeus, 1758

Common names

سه خار seh khar (= threespine), سه خاره (= seh khareh or sehkhareh), mahi seh khareh.

[trekhiglaya kolyushka or threespine stickleback in Russian; threespine stickleback].

Systematics

Gasterosteus aculeatus was originally described from Europe. The complex systematics of sticklebacks are not reviewed here. The Iranian specimens appear to be of a "marine" form with well-developed scutes on the flanks but their origin is not known.

Key characters

The 3 strong dorsal spines, strong pelvic fin spine with a single cusp at the base and the row of plates along the flank are distinctive.

Morphology

First dorsal fin comprised of usually 3 spines (2-4), second dorsal fin with 1 spine and 7-14 soft rays, anal fin with 1 spine and 6-13 soft rays, pectoral fin with 8-11, usually 10, branched rays and pelvic fin with 1 spine and 1 soft ray. Gill rakers 14-27, elongate and reaching as far as the fourth adjacent raker when appressed. Vertebrae 27-36. Plates along the flank 30-37; the Iranian specimens have a complete row of flank plates and a well-developed caudal peduncle keel, other populations may have an incomplete row or no plates at all. This species is very widespread and extremely variable over this range in the characters listed above and may well exceed the limits cited, e.g. dorsal fin spines are absent in some fish but the Iranian fish are of the typical marine form. The chromosome number is 2n=42 (Klinkhardt et al., 1995). The gut is short and s-shaped.

An Iranian specimen had 3 dorsal fin spines and 10 soft rays, 1 anal fin spine and 7 soft rays, 10 pectoral in rays and 1 spine and 1 soft ray in the pelvic fin. Scutes, including those forming the caudal peduncle keel, numbered 32. Total gill rakers numbered 19.

Sexual dimorphism

Breeding males develop a red belly and throat, blue sides, light blue back and have bright blue or turquoise eyes.

Colour

Marine populations are more silvery on the flanks than freshwater ones which are more olive. Generally the back is green-brown, olive or grey to blue-black, flanks olive to silvery and the belly silvery-white. Fins are generally clear. The peritoneum is silvery with large, widely spaced melanophores. Stream-dwellers have an irregular pattern of dark spots on the flanks and the back is an iridescent dark green.

Size

Reaches 11.0 cm.

Distribution

Found around the Northern Hemisphere in fresh and marine waters. Reported as an introduction to Iran by Abdoli (1993a, 1993c) and Coad and Abdoli (1993b), this species is found at the Neka Power Plant in Mazandaran near Behshahr, the Tajan River, the lower Gorgan River and Gorgan Bay, Bandar-e Torkeman, Gomishan Lagoon, the lower Babol, Heraz, Chalus, Tonekabon, Langarud, Polarud and Safid rivers, the Anzali Talab and the southeast Caspian Sea, the middle to upper Kashaf River in Khorasanin the Tedzhen (= Hari) River basin, and the upper Kal Shur and upper Jomein.rivers in the Dasht-e Kavir basin (Abbasi et al., 1999; Kiabi et al., 1999; Abdoli, 2000; K. Abbasi, pers. comm., 2001; Jolodar and Abdoli, 2004; Niksirat et al., 2006; Abdoli and Naderi, 2009). It is also caught by herring fishery ships on the coast of Iran (K. Abbasi, pers. comm., 2001).

Zoogeography

The exotic origin of the Iranian population is not known but the complete row of plates suggests a marine population. Sal'nikov (1995) reports this species from the southeastern Caspian Sea and moving into the Atrek River in Turkmenistan in large numbers. He refers to it as the Black Sea-Azov three-spined stickleback but it is unclear whether by this is meant its origin, its dispersal route or simply the nearest natural habitat for the species.

Habitat

These sticklebacks inhabit inshore coastal waters, lakes, ponds, rivers and streams, and may be anadromous. Marine and lake fish can be pelagic. They are often found among algae. In Iran, this species is caught at 30-100 m by herring fishery ships (K. Abbasi, pers. comm., 2001).

Age and growth

Maximum life span is a little over 3 years although some fish probably live only 1 year and a few months, dying after they spawn. Paepke in Bănărescu and Paepke (2002) is a recent review of literature.

Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 500 Iranian fish measuring 3.60-7.20 cm standard length. The a-value was 0.0090 and the b-value 3.428 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases). Patimar et al. (2010) studied this exotic in the Gomishan Wetland of the southeast Caspian Sea. Maximum ages were 2+ years for males and 3+ years for females, and the length-weight relationship was W = 0.0042TL3.711 for immature fish, 0.0095TL3.1328 for males and 0.0075TL3.4678 for females. The sex ratio was 1:2.63 in favour of females.

Food

Food is various crustaceans, aquatic and terrestrial insects, snails, worms, fish eggs and fry including their own species, and a wide variety of other available organisms taken both on the bottom or pelagically. Copepods and chironomids tend to dominate as food items. Again Paepke in Bănărescu and Paepke (2002) is a recent review of literature.

Niksirat et al. (2010) found stomach contents in the southeast Caspian Sea comprised gammarids, Nereis sp., chironomids, fishes, oligochaetes, fish eggs and hirudineans. Gammarids dominated in December and January and were replaced by Nereis in February. The preference shifted to chironomids in July and to fishes in October.

Reproduction

Spawning occurs from April to October, varying with locality over the wide range of this species. The male parental cycle at one site in Canada lasts 9-15 days with female interspawning intervals of 19 days. Males and females only complete one spawning here, though laboratory studies show males capable of 5 reproductive cycles and females of producing a clutch of eggs every 3-4 days. Harsh physical conditions are probably the cause. The male builds a barrel-shaped nest in shallow, sandy areas from plant fragments glued together on the bottom with kidney secretions. The nest is in an open area but near vegetation. The nest has an opening at each end. The male has a complex courtship dance with zig-zag motions and a leading motion to the nest. A responsive female adopts a submissive head up position, which also reveals the egg-swollen belly. The male pokes his snout at the nest to indicate its position to the female, tipping his head sideways to display the bright red throat. The male jabs the female with his snout through the nest wall after she enters to stimulate egg release. He then follows the female through the nest to fertilize the eggs and drives the female away. Several females may spawn in one nest which can contain up to 1026, yellowish 1.8 mm diameter eggs. The male guards and fans the eggs and guards the fry.

Patimar et al. (2010) in their Gomishan Wetland study found reproduction to occur in March-June, peaking in April. Large yolk-filled eggs with a mean diameter of 1.317 mm (maximum 1.96 mm) numbered 128-885. Small white opaque eggs were also found in the ovaries with a mean diameter of 0.549 mm and numbered 311-4709. The mean absolute fecundity was 1241.69 eggs and mean relative fecundity was 535.24 eggs/g body weight. Niksirat et al. (2010) found the breeding season in the southeast Caspian Sea began in March and lasted 5 months. Condition factor somatic condition factor and hepatosomatic index increased at the beginning of the spawning season and decreased as it progressed.

Parasites and predators

Many fishes and birds, and even snakes, seals and small mammals, feed on sticklebacks despite their protective spines which are locked erect when they are disturbed. Niksirat et al. (2006) record Corynosoma strumosum in this species from the Gomishan Lagoon.

Economic importance

The threespine stickleback has received much attention for its evolutionary interest, behaviour and utility as an experimental fish. They may be predators on fish eggs and larvae, and competitors for food, of more commercially important species. The Baltic Sea area has had sufficient numbers to support an oil extraction industry (Paepke in Bănărescu and Paepke, 2002). In the Iranian parts of the Caspian Sea this exotic may compete with juvenile sturgeons for food and its spread may account for the disappearance of the native Pungitius platygaster (Niksirat et al., 2010).

Conservation

Lelek (1987) classifies this species as intermediate to rare, locally vulnerable in Europe. As a recently recorded exotic species in Iran it is still comparatively rare but should not be accorded any status.

Further work

The spread of this exotic species should be monitored and its effects on native fishes observed.

Sources

Iranian material: CMNFI 1993-0144, 1, 57.3 mm standard length, Mazandaran, Neka Power Plant (36º51'48"N, 53º23'24"E).

Genus Pungitius
Coste, 1848

This genus of sticklebacks is found in North America and Eurasia with about 3-5 species (opinions vary) but only 1 in Iran (Paepke in Bănărescu and Paepke, 2002).

It is distinguished by a series of 7-12 small spines (much smaller than in the genus Gasterosteus) in front of the dorsal fin, arranged alternately to the left and right, the pelvic fin has 1 spine and 0-1 soft rays, and the skin is naked or has small bony plates in a row along mid-flank.

Pungitius platygaster
(Kessler, 1859)

Common names

nohkhar or nokhareh (= ninespine), mahi seh khareh nama (= threespine-like fish), mahi khardar (= spiny fish).

[tikan balig in Azerbaijan; malaya yuzhnaya kolyushka or southern ninespine stickleback in Russian; Ukrainian stickleback].

Systematics

Gasterosteus platygaster was originally described from Odessa and Aleshki on the Dnieper in the Ukraine.

Gasterosteus pungitius var. Kessleri Yakovlev, 1870 and Gasterosteus pungitius var. niger Yakovlev, 1870 both from lakes near Astrakhan, Volga River delta, Russia, Gasterosteus platygaster var. caucasicus Kessler, 1877 from Transcaucasia, and Pygosteus platygaster nuda Berg, 1905 from Lake Charkhal in the Ural River valley, are synonyms. Syntypes of the latter are in the Zoological Institute, St. Petersburg (ZISP 10613) (Eschmeyer et al., 1996; Kottelat, 1997).

Münzing (1969) retains this taxon as a distinct species but suggests that it may only be a subspecies of Pungitius pungitius (Linnaeus, 1758). Ziuganov and Gomeluk (1985) studied hybridisation of this species and P. pungitius under experimental conditions and consider that they are at most subspecifically distinct. They found 9 out of 19 morphological characters distinguish this species from P. pungitius as well as a differing ecology. However there were no ethological, nor presumably genetical, isolating mechanisms since F3 hybrids were fertile. Ziuganov (1991), Keivany (1996), Keivany and Nelson (1997) and Keivany et al. (1997) recognise P. platygaster as a distinct species since it is 100% distinguishable from other species in the genus. However, Keivany and Nelson (2004) found that P. platygaster lay within a polychotomy with subspecies of P. pungitius. P. platygaster is retained as a full species until the two taxa come into contact naturally, which will test the hypothesis of Ziuganov and Gomeluk (1985).

Key characters

The small spines usually alternating left and right in front of the dorsal fin are distinctive.

Morphology

Characterised by 22-34 bony plates along the flank, perhaps a response to predators (Ziuganov and Gomeluk, 1985). The first 4-12 plates are large (Keivany and Nelson, 1998). There is no caudal peduncle keel. The plates in young fish are minute fragments of bone and difficult to see, especially posteriorly. The body form is less elongate than the related Pungitius pungitius. Dorsal fin spines slightly curved, 7-11 (in Iranian specimens 8(7), 9(11)) and branched rays 6-11, anal branched rays 6-10, pectoral fin rays 9-11 (usually 10), and pelvic fin with 1 serrate spine and 0-2 soft rays, usually 1. Rarely up to 4 dorsal fin spines may follow the midline and not alternate markedly in Iranian fish. The last spine in the series is usually slightly longer than the others. Vertebrae number 29-31 (in Iranian specimens 29(9), 30(9)) and gill rakers 7-13, reaching the base of the second raker below when appressed. The gut is short and s-shaped. The chromosome number is 2n=42 (Klinkhardt et al., 1995).

Sexual dimorphism

Colour at breeding is the most evident sexual dimorphism.

Colour

Females are olive with greenish-brown stripes, patches and mottling even during spawning. Males are jet black and develop a blue-white colour on the posterior side of the ventral spines as a stimulus for leading the females. The specimens described as var. kessleri were light grey and var. niger were black, perhaps non-spawners and spawners. The peritoneum is silvery.

Size

Attains 7.0 cm standard length.

Distribution

Found in the Black, Caspian and Aral seas and their basins. Reported from the southeast Caspian Sea, southwest Caspian Sea and south-central Caspian Sea, the Anzali Mordab, Ameerkalaye Lagoon near Lahijan, and lower reaches of rivers such as the Safid, Shakhzaderud (lower Babol), lower Karasu and Gorgan (Derzhavin, 1934; Holčik and Oláh, 1992; Nejatsanatee, 1994; Abbasi et al., 1999; Kiabi et al., 1999; Abdoli and Naderi, 2009). Keivany (1996) considers it to be rare on the southern Caspian Sea coast. Jolodar and Abdoli (2004) report it from brackish portions of the Caspian Sea and rivers falling into it, being more numerous in the Gomishan Lagoon.

Zoogeography

Pungitius platygaster aralensis (Kessler, 1877) is reported from the western Uzboi lakes in Turkmenistan (Shakirova and Sukhanova, 1994) and with canal construction projects may ultimately reach the Caspian Sea basin.

Habitat

Described as a benthivore living a cryptic life in weed beds (Ziuganov and Gomeluk, 1985). It can be found in fresh and brackish waters.

Age and growth

In the Volga delta, mature adults are 3.2-5.8 cm long (mean 4.0 cm) from April to July and young-of-the-year are 1.2-2.8 cm long (mean 2.3 cm). Life span appears not to exceed 2 years as only sexually mature adults are caught in spring (Fortunatova, 1961) although a few fish may reach 3 years and maturity may be attained shortly before reaching 1 year (Paepke in Bănărescu and Paepke, 2002).

Food

Diet is small invertebrates including chironomids. Eggs of commercially important fishes may be eaten.

Reproduction

Reproduction can occur at high temperatures, e.g. 28°C, at which eggs of P. pungitius die. Spawning begins at 17°C. In the Volga delta spawning begins in late April and continues to the middle of June but is most intensive in the middle of May. Dense schools are formed which disperse after spawning with the fish becoming secluded in the heavy vegetation which develops at this time (Fortunatova, 1961). Fecundity is up to 90 eggs. Males build nests out of the remains of the previous year's vegetation glued together with kidney secretions (Fortunatova, 1961). A female indicates readiness to spawn by a head-up posture near the nest. The male performs a zig-zag dance and leads the female to the nest entrance by angling his head down at 60°, flashing his blue-white pelvic spines and moving by short jerks. The female enters the nest, the male taps her caudal peduncle to stimulate egg deposition, she leaves and the male enters the nest to fertilize the eggs. The leading part takes only 10 seconds.

Parasites and predators

The Caspian seal, Pusa caspica, is a predator on this species (Krylov, 1984) as are such fishes as Perca fluviatilis, Esox lucius, Silurus glanis, Aspius aspius and Sander lucioperca (Fortunatova, 1961) and Neogobius melanostomus, water snakes, pond turtles, frogs and fish-eating birds (Ziuganov, 1991).

Economic importance

None; although Paepke in Bănărescu and Paepke (2002) consider it important as a predator on mosquito larvae (and hence a biological control for malaria) in smaller water bodies.

Conservation

Holčík and Oláh (1992) report the loss of this species from the Anzali Mordab, where it was once abundant, through a change in the environment to fresh water from brackish, siltation, oxygen depletion and loss of spawning grounds. Keivany (1966) considers it to be rare on the southern Caspian Sea coast while Kiabi et al. (1999) consider this species to be of least concern in the south Caspian Sea basin according to IUCN criteria. Criteria include medium numbers, widespread range (75% of water bodies), absent in other water bodies in Iran, and present outside the Caspian Sea basin.

Further work

The distribution and biology of this species in Iranian waters needs further investigation.

Sources

Descriptions of breeding and some meristic characters are taken from Münzing (1969) and Ziuganov and Gomeluk (1985).

Iranian material: CMNFI 1970-0552, 1, 16.0 mm standard length, Gilan, Sowsar Roga (37º27'N, 49º30'E); CMNFI 1970-0554, 12, 17.5-22.0 mm standard length, Gilan, Pir Bazar Roga (37º21'N, 49º33'E); CMNFI 1970-0567, 2, 17.7-21.4 mm standard length, Gilan, Pir Bazar Roga (37º21'N, 49º33'E); CMNFI 1970-0591, 2, 15.8-17.4 mm standard length, Gilan, Pir Bazar Roga (37º21'N, 49º33'E); CMNFI 1979-1215, 1, 22.3 mm standard length, Gilan, Sowsar Roga (37º27'N, 49º30'E).

Syngnathidae

The pipefishes and seahorses are found world-wide, mostly in tropical and warm-temperate waters, with some brackish and freshwater species. There are 52 genera and about 232 species (Nelson, 2006) but only one species of pipefish is reported for Iranian fresh waters. Maximum length is about 65 cm. The body of pipefishes is characteristic being very thin and very elongate, and enclosed in bony rings as a form of armour. The body is divided into a trunk and a tail, the tail being prehensile in seahorses. The first trunk ring has the pectoral fin base and the last has the anus in it. The snout is elongate with a small toothless mouth at the tip. Gills are tufted and the gill opening is small. There are 1-3 branchiostegal rays. There is a single dorsal fin without spines, the pelvic fins are absent and the pectoral, anal and caudal fins may be absent too. The caudal is always small when present. The anal fin is always small with only 2-6 rays. Those without a caudal fin may have the tail prehensile, able to grasp and hold onto objects. Pipefishes and seahorses have only one kidney, on the right side. Males usually have a pouch on the belly formed from two skin folds which meet at the mid-line. The female attaches eggs to the male belly or places them in a pouch (or marsupium). Here the eggs develop and eventually leave after about 4-6 weeks through a slit or pore so the male "gives birth".

Pipefishes and seahorses are found mostly in shallow coastal areas and in estuaries but also occur down to about 400 m and in the open ocean often associated with floating seaweed. They are slow-moving because of their armour and easily picked up by hand. Propulsion is by undulating the dorsal and pectoral fins. Since they cannot outswim most predators, they are often very well camouflaged by colour, body form and by appendages which disrupt the body outline. Their food is small crustaceans sucked into the tube-like snout by a sudden expansion of the buccal cavity. Many species have a life span of only about 2 years.

Dried seahorses and pipefishes are commonly sold as curios and some family members have been used as medicines or aphrodisiacs in the East. They are popular aquarium fishes. Certain species are endangered by this collecting for curios, medicines and aquaria.

Genus Syngnathus
Linnaeus, 1758

Members of this genus are found in all seas and there are about 31 species. A single species is found in the Caspian Sea and Iranian waters.

The genus is characterised by a brood pouch in the caudal area of males, running forward to about the origin of the dorsal fin. The eggs develop in isolation from the sea, oxygen diffusing from the "pregnant" male. Dorsal, caudal, anal and pectoral fins are present. The caudal and anal fins are very small.

Syngnathus caspius
Eichwald 1831

Common names

ney mahi or naymahi (= reed fish, i.e. as thin as a reed), ney mahi darya-ye Khazar (= reed fish from the Caspian Sea), سوزن ماهي (= suzan mahi, meaning needle fish).

[xazar iynabaligi in Azerbaijan; Kaspiiskaya igla-ryba or Caspian pipefish in Russian; short-snouted pipefish, blackstripe pipefish].

Systematics

The Caspian pipefish has appeared under several scientific names. Lueken (1967) demonstrated that Syngnathus nigrolineatus caspius Eichwald, 1831, the taxon generally referred to in the literature as occurring in the Caspian Sea, is a synonym of Syngnathus abaster. Syngnathus abaster was originally described from Nice, France.

Other taxa of relevance are Syngnathus ponticus Pallas, 1814 described in part from mouths of rivers falling into the Caspian Sea, Syngnathus nigrolineatus Eichwald, 1831 described from "Hab. in sinu Bacuensi, Murdofiensi Caspii maris", i.e. Baku Bay, and also at Odessa on the Black Sea, Syngnathus caspius Eichwald, 1831 described from "Hab. in sinu balchanensi Caspii maris" (Balkhan Bay in the Caspian Sea), and Syngnathus bucculentus Rathke, 1837 described from Sevastopol and Feodosiya on the Black Sea, all considered as synonyms of S. abaster. However, Naseka and Bogutskaya (2009) retain S. caspius as the species in the Caspian Sea.

Key characters

The extremely thin, elongate body encased in bony rings is unique in Iran.

Morphology

Dorsal fin rays 22-43, the count increasing with age; anal fin rays 3, pectoral fin rays 9-15, and total caudal fin rays 8-11. Trunk rings 14-18; tail rings 28-42, and rings under the dorsal fin 5-12. A dorsal ridge ends near the rear of the dorsal fin and a continuation of the mid-lateral ridge becomes the dorsal ridge posteriorly. A mid-belly ridge extends back to the anus where it terminates. As a result the anterior part of the body from the dorsal fin forward has 7 longitudinal ridges, reduced to 4 posteriorly.

Meristic values for Iranian specimens are:- 30(3), 31(6), 32(9), 33(18), 34(13), 35(7), 36(1), 37(1), 39(2), 40(4), 41(3), 43(1); anal fin rays uniformly 3; pectoral fin rays 11(14), 12(22), 13(10), 14(8), 15(1); total caudal fin rays 9(1), 10(51), 11(2). Trunk rings 15(46), 16(18), 17(4); tail rings 33(3), 34(7), 35(28), 36(13), 37(5), 38(6), 39(4), 40(2), rings under the dorsal fin 6(1), 7(28), 8(24), 9(2); and brood pouch rings 15(1), 17(2), 18(3), 19(1), 20(3), 21(1), 22(2), 23(1). Total vertebrae 52(3), 53(2), 54(2), 55(4), 58(1), 59(1).

Sexual dimorphism

Males have a brood pouch under the tail.

Colour

Overall colour is brown to green with dark or light spots and bars arranged in a reticulate fashion on each ring. Pigment is best developed on the dorsal half of the head and body. The ventral part of the tail becomes darkest posteriorly near the tail fin. The reticulate pattern of darker pigment can form a consistent pattern of arcs bounding the plate junctions along both the upper and lower flank such that a double row of dark ovals with light centres is apparent. Behind the dorsal fin where there is no central ridge on the flank, there is a single dark oval at each plate junction. While some fish have this regular pattern of dark ovals, others are more irregular with the ovals filled in with pigment leaving only very small clear patches in the centre. Various other patches of clearer or lighter pigmentation form part of the general reticulate pattern. Additionally, large dark spots or pigment concentrations may be present along the edge of the uppermost ridge anteriorly and small dark spots may be scattered along the back and flanks.

The dorsal fin base can be darkly spotted or striped. The fin itself may be almost immaculate but a few melanophores line the rays. The pectoral fin base has a dark line of merged spots and the rest of the fin is almost immaculate although the rays can be very lightly lined with melanophores. The anal fin is immaculate. The caudal fin is dark on the rays and membranes, in particular on the central part.

Size

Reaches 23 cm total length.

Distribution

Found in the Mediterranean, Black and Caspian seas and the west coast of the Iberian Peninsula. In Iran, it is recorded from the Anzali Mordab and neighbouring rivers such as the Sowsar Roga and at Ab Kenar village, at Bandar-e Anzali and Anzali beach, the Amirkelayee Lagoon near Lahijan, Gorgan Bay, lower reaches of Caspian Sea rivers such as the Safid, Rud-e Sera, Lalarud and Babol, the southeast Caspian Sea, southwest Caspian Sea and south-central Caspian Sea, including the deltas of some rivers (Derzhavin, 1934; Holčík and Oláh, 1992; Paateemaar, 1993; Nejatsanatee, 1994; Kiabi et al., 1999; Jolodar and Abdoli, 2004; Abdoli and Naderi, 2009).

Zoogeography

Berg (1948-1949) contends that this species entered the Caspian from the Black Sea in post-glacial times while most other Caspian fishes are relicts of earlier transgressions or migrants from northern waters.

Habitat

This pipefish can live and reproduce in water at a salinity of 59.5‰ (Zenkevitch, 1963) and is euryhaline. It is found over sandy or muddy bottoms usually where there is vegetation or detritus, down to about 5 m. Campolmi et al. (1996) consider this species to be a major component of nearshore meadows in Sicily. Juveniles have a benthic distribution for at least 4 weeks after release from the marsupium in Portugal (Silva et al., 2006).

Age and growth

Growth is faster in the first year of life as in the second year energy is used for reproduction. Life span is slightly over one year in northwest Sicily (Campolmi et al., 1994; 1996) and 17 months in the Po River Delta, Italy (Franzoi et al., 1993), or up to 4 years generally. Male:female sex ratio is 1:2.18 in Sicily (Campolmi et al., 1994; 1996) but about 50:50 in the Po Delta (Franzoi et al., 1993). Maturity in the Mauguio Lagoon in southern France is reached as early as 3-4 months and almost all individuals are mature after their first winter (Tomasini et al., 1991). Heydarnejad (2009) gave the length-weight relationship for an Iranian sample as W = 0.0099TL3.543.

Food

Campolmi et al. (1994; 1996) found their northwest Sicily population to feed mainly on zoobenthos, 70% being harpacticoids, ca. 18% or less gammarids and caprellids, and ca. 10% or less isopods. Prey hidden in vegetation was taken on account of the short and conical snout which was less effective than the longer snout of other species which feed on more active pelagic prey. Franzoi et al. (1993) in their study found harpacticoids associated with algae to be the most important food item, with up to 87.5% of the diet comprised of only 3 species in 1 genus, Tisbe.

Reproduction

Broods are found from April to October with the first breeding males appearing in March in Sicily, the Po Delta and southern France (Campolmi et al., 1994; 1996; Franzoi et al., 1993; Tomasini et al., 1991).

Reproductive behaviour starts with mutual flickering movements in fish observed in Portugal. This comprised rapid and vigorous bends of the body with both males and females approaching the opposite sex  (Silva et al., 2006). If the opposite flickered in response, the next phase was rapid side-by-side movements while swimming more or less parallel. Flickering increased in frequency and the female's genital papilla protruded. The distended genital papilla is placed in the anterior area of the marsupium of the male, the folds in this area being separated and swollen. The mated pair then ascend slowly in the water column, vibrating the dorsal fin and rotating a few times. Eggs are transferred and the  female retreats to the substratum while the male continues swimming with violent body contractions - these help pack the eggs in the posterior end of the marsupium. Later, spawning occurs again up to three times by the same pair although females may mate with up to 3 different males in less than 30 minutes.

Egg numbers in the Po Delta are 104±40 in females and 109±27 in males. Eggs are yellowish to bright orange and translucent with diameters usually 1.0-1.4 mm, up to 1.8 mm (Tomasini et al., 1991) or 1.09-2.06 mm in Portugal (Silva et al., 2006). Young first appear first in May and are present until August in the Po Delta. This species is a batch spawner in northwest Sicily and southern France and males can incubate several broods during the breeding season since incubation lasts about 1 month (Tomasini et al., 1991; Franzoi et al., 1993; Campolmi et al., 1994; 1996). Females in the Po Delta probably breed only once but the female may parcel out the same egg batch among many males. A male may have several batches of eggs in different stages of development in the brood pouch at any one time. This distribution among several males increases survival (Tomasini et al., 1991). Incubation at 20-21°C in aquaria is about 15 days and a new mating occurs 8 days after the litter (Tomasini et al., 1991). In Portugal, development lasted 24-32 days at 18-19ºC or 21 days at 21-22ºC (Silva et al., 2006). The female potential fecundity per breeding act in southern France averages 21-23 ovocytes while the male brood pouch can accommodate 36-52 eggs. As a consequence the female matures rapidly between two breedings and the species is polygynous (Tomasini et al., 1991). Males carry 10-64 eggs in Portugal (Silva et al., 2006). New-borns are about 13.5 mm long in the Po Delta (Franzoi et al., 1993).

Broods are found in males from Iran collected on 4 May, the same sample having males with eggs and males with minute young pipefish. A sample from 22 September has eggs just about to hatch while males taken on 3 August, 28 September and 17 December contain neither eggs nor young. Females with large eggs have been caught in Iran on 17 April and 7 July while on 3 August, 19 September, and 12 and 17 December females had only small eggs. The 17 April sample contained eggs of dissimilar sizes. Iranian fish appear to have a similar breeding season to those from other localities and probably have batch spawning too.

Parasites and predators

The Caspian seal, Pusa caspica, is a predator on this species (Krylov, 1984).

Economic importance

None.

Conservation

The ecology of this species has not been studied in detail in Iranian waters and its population is unknown rendering an assessment of its conservation status difficult. Kiabi et al. (1999) consider this species to be of least concern in the south Caspian Sea basin according to IUCN criteria. Criteria include abundant in numbers, widespread range (75% of water bodies), absent in other water bodies in Iran, and present outside the Caspian Sea basin.

Further work

The significance of this species in the ecology of the Iranian coastal region and its biology have not been determined.

Sources

Paateemaar (1993) gives an account of this species in Iran in Farsi. Data were taken from Lueken (1967) and Dawson in Whitehead et al. (1984-1986).

Iranian material: CMNFI 1970-0507, 1, 91.3 mm standard length, Gilan, Caspian Sea at Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1970-0508, 1, 89.4 mm standard length, Gilan, Safid River at Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1970-0531, 1, 108.1 mm standard length, Mazandaran, Larim River mordab (36º46'N, 52º58'E); CMNFI 1979-0535, 1, 126.3 mm standard length, Gilan, Shara River estuary (37º35'N, 49º09'E); CMNFI 1970-0543A, 5, 104.6-144.8 mm standard length, Gilan, Caspian Sea at Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1970-0554, 13, 40.4-95.6 mm standard length, Gilan, Pir Bazar Roga (37º21'N, 49º33'E); CMNFI 1970-0563, 1, 134.9 mm standard length, Gilan, Caspian Sea, Kazian Beach (ca. 37º29'N, ca. 49º29'E); CMNFI 1970-0567, 7, 52.9-99.5 mm standard length, Gilan, Pir Bazar Roga (37º21'N, 49º33'E); CMNFI 1970-0575, 1, 94.3 mm standard length, Gilan, Pir Bazar Roga (37º21'N, 49º33'E); CMNFI 1970-0587, 3, 131.6-140.0 mm standard length, Mazandaran, Babol River near Babolsar (36º43'N, 52º39'E); CMNFI 1971-0343, 3, 104.6-109.0 mm standard length, Langarud at Chamkhaleh (37º13'N, 50º16'E); CMNFI 1979-0077, 8, 128.4-149.2 mm standard length, Mazandaran, Caspian Sea beach at Now Shahr (36º39'N, 51º31'E); CMNFI 1980-0130, 9, 59.3-97.8 mm standard length, Mazandaran, river near Iz Deh (36º36'N, 52º07'E); CMNFI 1980-0136, 16, 76.6-100.5 mm standard length, Mazandaran, Fereydun Kenar River estuary (36º41'N, 52º29'E).

Mastacembelidae

The spiny eel family is found principally in fresh waters of tropical Africa and eastwards to Korea and Malaysia. The distribution mapped by Berra (2001) extends too far into central Iran. There are about 73 species. Maximum length is less than 1 m.

The family is characterised by an very elongate, compressed or sub-cylindrical body; minute cycloid scales or body naked; a non-protractile mouth; gill opening a slit; an elongate snout with a sensitive tip flanked by tubular anterior nostrils, the posterior nostrils are hence far to the rear; long dorsal and anal soft fins (30-131 rays), dorsal soft fin preceded by usually numerous isolated spines (9-42); anal fin preceded by 2-3 spines; pectoral fin present with 12-27 rays but pelvic fin absent; caudal fin short and confluent with the dorsal and anal fins or only separated narrowly; 60-110 vertebrae; no pseudobranchiae; 2 pyloric caeca; and swimbladder present.

Some species are food fishes and they regularly appear in the aquarium trade as they are brightly coloured with distinctive patterns. They may burrow in mud and even survive some drying in ponds through their air-breathing ability. Their eel-like shape is reflected in a wriggling behaviour when handled and some are known to move backwards to impale the hand. They are found at high altitudes as well as in lowlands, in both still and running waters, often in rocky crevices or in vegetation near banks, and they emerge particularly at night to feed on prey. The rostral appendage is used to detect the prey by touch and the prey is rapidly inhaled.

Genus Mastacembelus
Scopoli, 1777

This spiny eel genus is found from the Levant to Southeast Asia. There is 1 species in Southwest Asia. Heckel (1846-1849a) misspells the genus name Mastacacemblus.

The genus is characterised by a moderate to large size, an elongate and compressed body, inferior mouth with a narrow cleft, small eyes, the absence of eye spots on the soft dorsal fin, the absence of striations under the snout, 6 branchiostegal rays, preopercle spiny or not at its postero-ventral corner, preorbital spine present, scales minute and cycloid, elongate swimbladder, lateral line present, minute jaw and palate teeth, and a rounded caudal fin.

Mastacembelus mastacembelus
(Banks and Solander, 1794)

Common names

مارماهي (= mar mahi, meaning snake fish), marmahi-ye khardar (= snake fish with spines or spiny snakefish), مارماهي شاخدار (= mar mahi shakhdar).

[salbouh abu-el-sian, salbu-al-sayan, saebouh abou siyan; abu salmabah; simack, englisi or englese at Aleppo, marmaritch or marmarij at Mosul, all in Arabic; Mesopotamian spiny eel, Euphrates spiny eel].

Systematics

Ophidium Mastacembelus was originally described from "Aleppo" and from the "River Kowick" (= Quwayq) in the book by Russell (1794).

Ophidium Simack Walbaum, 1792 (non-binomial), Rhynchobdella haleppensis Bloch and Schneider, 1801 and Mastacembelus syriacus Gronow in Gray, 1854 are synonyms. Mastacembelus aleppensis Günther, 1861 is an unjustified emendation of haleppensis (Eschmeyer et al., 1996). Wheeler (1956) and Sufi (1957) discuss the names of this species more fully.

Three syntypes of Ophidium Mastacembelus from "Aleppo" are in the Natural History Museum, London under BM(NH) 1955.6.25:4-6, measuring 289.5-544.0 mm standard length.

Çakmak and Alp (2010) found morphological differences between river and reservoir populations in Turkey, but none for meristic traits. Some differences appear to be associated with habitat type, a thicker and longer caudal peduncle in river populations may enhance swimming ability in faster water and longer lower jaws in river fish could be a feeding adaptation, for example.

Key characters

The eel-like body with 30-35 short, sharp dorsal spines, long soft dorsal and anal fins and the unique flexible snout flanked by tubular nostrils distinguish this species from all other Iranian fishes.

Morphology

The mouth gape extends back as far as the anterior eye margin or somewhat forward of this point. The posterior nostril is slit-like. The structure of the elongated eye, typical of streamlined forms, has been described by Jasim (1998) based on Iraqi specimens. The regular, mosaic pattern of the retina is associated with fishes that search for their food and the double cone structure may be associated with detection of moving prey.

Soft dorsal rays 68-90, soft anal rays 70-90 after 3 spines, pectoral rays 18-24 and total vertebrae 85-88. The penultimate spine in the dorsal fin is the longest and the central anal spine is the longest. Iranian specimens had 31(2), 32(6) or 33(4) dorsal fin spines, 19(5) or 20(3) pectoral fin branched rays, and total vertebrae 86(1), 87(1). The diploid chromosome number is 2n=48, arms number NF=88 and there are 11 metacentric, 9 submetacentric and 4 subtelocentric chromosome pairs (Esmaeili et al., 2006).

Scales minute but covering the whole body, under the eye, below the posterior nostril and between this nostril and the maxilla. Each scale has a central to anterior focus with radii on all fields and an oval shape. There is a strong preorbital spine under the eye, present in some fish but concealed under the skin or absent in others. Teeth form broad bands in both jaws with the outermost teeth the largest. There are no gill rakers but spinulose patches lying flat on the arch. The gut is an elongate s-shape.

Sexual dimorphism

Unknown.

Colour

The body is blotched and barred, often forming a reticulate pattern, or a series of mid-flank blotches most evident posteriorly and sometimes running together as a stripe anteriorly. Flank blotches may form up to 17 bars running from the dorsal to the anal fin across the flank. The back is blackish to brown, olive, greyish or blue-grey, the lower flank is spotted yellow or is yellow overall and the belly is white to yellowish. A series of about 20-24 black to blackish-brown, oval spots ringed with a lighter brown follow a dark, broad but irregular stripe on the head and anterior back in the mid-line. Dorsal, anal and caudal fins are yellowish with the dorsal and caudal fins finely barred and the anal fin continuing the pattern on the adjacent body. The anal fin may be almost immaculate. The soft dorsal fin may have vermiculations rather than bars. The pectoral fins are yellowish and are finely barred. The peritoneum is brownish, with numerous fine melanophores. Small fish (about 7.7 cm total length) can be an almost uniform grey-brown to brown-green, with yellowish brown on the fins and the tail region, and fin spots are dark to absent. There is a thin bar extending vertically down or obliquely back from the eye.

Size

Reaches 76.9 cm total length (Gümüş et al., 2010), probably larger to almost 1 m.

Distribution

Found in the Quwayq, Orontes and Tigris-Euphrates basins. In Iran its is recorded from the Tigris River, Gulf and Kor River basins, including lakes Zaribar and Marivan, and the Fasa, Dez and Dalaki rivers (Löffler, 1957; Abzeeyan, 5(5):III, 1994; Gh. Izadpanahi, pers. comm., 1995).

The record of Mastacembelus armatus (Lacepède, 1800) reported by Mokhayer (1981b) from the Kor River basin is probably a mis-identified Mastacembelus mastacembelus.

Abdoli (2000) maps this species from the Kor and Pulvar rivers of the Kor River basin; the middle and lower Shur River tributary of the Dasht-e Palang, the upper Mand and Qara Agaj, the middle and upper Hilleh and the upper Zohreh rivers of the Gulf basin; the Jarrahi, lower Karun, middle and lower Dez, Karkheh, Simarreh and lower Kashkan rivers of the Tigris River basin.

Zoogeography

This species is now known to occur outside the Tigris-Euphrates and Quwayq basins in Southwest Asia in contrast to Banister's assertion (1980). The distribution of this species is not, however, continuous across Iran as shown in a figure by Travers (1984b) mis-quoting Coad (1979, actually 1980a). It appears to be absent from the saline rivers draining to the Straits of Hormuz and from Baluchestan.

Habitat

This species is known from both lotic and lentic environments (Pazira et al., 2005). Sufi (1957) described mastacembelids as usually lurking in rock crevices or among stumps of plants near the bank and I have observed them at Lake Parishan (= Famur) inhabiting crevices of a submerged rock wall. They may be able to survive desiccation by burying themselves in mud. They are not commonly caught with nets and may be mostly nocturnal in habits. In the Marun River below Behbehan, Khuzestan this species is very common, commoner then any other site sampled in this province, possibly the result of pollution-enriched water (field sample; H. R. Alizadeh, pers. comm., 2000). In areas under human influence in Lorestan, such as the lower reaches of rivers and near cities, it is more common than in higher, pristine waters.

Age and growth

In the Helleh and Dalaki river basins of southern Iran, the condition factor of this species was 0.162-0.458 (mean 0.296) for females and 0.162-0.386 (mean 0.289) for males. Condition factor = 0.374 - 0.004 total length. Life span was up to 6 years although most fish were 3 years or younger. Females grew rapidly to age 3, after which annual growth decreased. von Bertalanffy length-at-age equations were Lt = 873.4 (1-exp{-0.082[t + 1.488]}) for females and Lt = 923 (1-exp{-0.081[t + 1.464]}) for males (Pazira et al., 2005). Eroğlu and Şen (2007) examined fish in Karakaya Dam Lake, Malatya, Turkey and found 9 age groups with males in age group 4 forming the majority of the population. Males generally outnumbered females in all age groups. Gümüş et al. (2010) found males up to 21 years and females to 9 years in fish from Atatürk Dam Lake on the Turkish Euphrates River. The length and weight frequency distributions were significantly different between sexes in this population and other growth parameters were given. It was suggested that differences in life span between Iranian and Turkish fish could be due to habitat differences (lotic and lentic habitats) or fishing pressure.

Food

The flexible snout is used for sniffing out food but the eye structure suggests a visually feeding fish also. Food is assumed to include invertebrates but two fish from Iran contained fish scales and fish skeletal remains. Other species are known to eat fish eggs and fry. Food in the Hawr al Hawizah, Iraq in 2005-2006 was 55.0% shrimps and 45.0% fish and in the Al Kaba'ish (= Chabaish) Marsh entirely fish (Hussain et al., 2006). Pala et al. (2010) found fish from Karakaya Dam Lake, Turkey to contain plant material (predominately 16 taxa of Bacillariophyta), cladocerans, copepods, and fish.

Reproduction

Fish taken on 26 November have small but developing eggs, suggestive of spring spawning. Al-Rudainy (2008) states that Iraqi fish reach sexual maturity at 2 years, 25 cm length and 125 g in weight, and spawn in June and July in shallow muddy water among rocks. Şahinöz et al. (2006) give details of development of embryos and artificial breeding, based on fish taken from Ataturk Dam Lake in Turkey. They obtained a fertilisation rate of 80%, egg diameters reached 2.015 mm, and hatched larvae were observed at 85 h after insemination. Eroğlu and Şen (2007) in their Turkish population found males to mature at age 2, females matured at age 1 (although in most fish females mature later because of the demands of egg production), spawning took place mostly in June-July and eggs numbered up to 27,944 and reached 2.1 mm in diameter.

Parasites and predators

Predators might find this species difficult to swallow. The row of dorsal spines are very sharp and can severely lacerate the hand when this fish is picked up carelessly. Mokhayer (1981b) records a heavy infestation with Contracaecum larvae in Lake Parishan near Kazerun. Akhalaghi (2001d) found the nematode Anguillicola crassus in swimbladders of this fish from Parishan Lake. Jalali et al. (2002) and Jalali and Barzegar (2006) record Diplostomum spathaceum, Trichodina pediculus, Ichthyophthirius multifilis, Mastacembelocleidus heteranchorus, two species of Argulus, Lernaea cyprinacea, a Polyonchobothrium species and an Ancyrocephalus sp. from this species in Lake Zarivar. Barzegar et al. (2008) record the digenean eye parasite Diplostomum spathaceum from this fish. Jalali et al. (2008) examined this fish in lakes Zerivar and Parishan and the Helleh River and found the protozoans Trichodina pediculus and Ichthyophthirius multifilis (Zarivar), the monogenean Mastacembelocleidus heteranchorus (all three localities), the digenean Diplostomum spatheceum (Zarivar), the crustaceans Argulus foliaceus, Argulus sp., Lernaea cyprinacea and Lernaea sp. (all Zarivar), the cestode Polyonchobothrium sp. (Zarivar), and the nematode Contracaecum sp. (Parishan). Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Lernaea sp. , Lernaea cyprinacea,  Argulus foliaceus and Argulus sp. on this species.

Economic importance

Russell (1794) reported that this fish was "found in great abundance" and "esteemed a lighter and more delicate food" than the eel at Aleppo but it does not seem to be so common in Iran or used generally as food. Heckel (1846-1849a) gives the common name "englisi" because it was highly prized by Europeans at Aleppo. Foreign soldiers have caught this species by angling at Baghdad (www.carpecapio.com, downloaded 26 August 2005). Harlioğlu and Yilmaz (2011) note its local commercial value in Turkey, analyse its chemical composition, and recommend it for human consumption as a good source of nutrition.

Conservation

This species appears to thrive in polluted areas and is not commonly caught or utilised. Its conservation status has not been assessed in field studies but it is probably not under any threat. Critically endangered in Turkey (Fricke et al., 2007).

Further work

The biology of this species needs further investigation. An assessment of its conservation status could be carried out as its somewhat cryptic nature makes its numbers difficult to determine.

Sources

Type material:- See above (BM(NH) 1955.6.25:4-6).

Iranian material: CMNFI 1979-0029, 1, 148.0 mm standard length, Fars, Dalaki River (no other locality data); CMNFI 1979-0075, 1, 252.0 mm standard length, Fars, Mand River at Pol-e Kavar (29º11'N, 52º41'E); CMNFI 1979-0124, 1, 436.0 mm standard length, Fars, Mand River at Shahr-e Khafr (28º56'N, 53º14'E); ZSM 21831, 1, 584.5 mm standard length, Khuzestan, Harmaleh on Dez River (31º57'N, 48º34'E); ZSM 26629, 1, 154.4 mm standard length, Kordestan, Lake Zaribar (35º32'N, 46º08'E); uncatalogued: 1, 227.5 mm standard length, Fars, Mand River outside Jahrom (28º36'N, 53º37'E); 1, 219.0 mm standard length, Khuzestan, Rud Zard, (31º22'N, 49º43'E); 3, 105.0- ca. 420.0 mm standard length, Khuzestan (no other locality data).

Comparative material:- BM(NH) 1875.1.14:7, 2, 341.8-569.1 mm standard length, Iraq, Baghdad (33º21'N, 44º25'E); BM(NH) 1912.5.2:8, 1, 314.3 mm standard length, Iraq, Shatt-al-Arab (no further locality data); BM(NH) 1920.3.3:297-300, 4, 134.5-289.6 mm standard length, Iraq, Basra (30º30'N, 47º47'E); BM(NH) 1920.3.5:7, 1, 103.8 mm standard length, Iraq, Basra (30º30'N, 47º47'E); BM(NH) 1922.3.25:1, 1, 424.0 mm standard length, Iraq, Baghdad (33º21'N, 44º25'E); BM(NH) 1936.3.10:4, 1, 40.0 mm standard length, Iraq, Nasiriyah (31º02'N, 46º16'E); BM(NH) 1974.2.22:1798, 1, 380.5 mm standard length, Iraq, Basra (30º30'N, 47º47'E); BM(NH) 1974.2.22:1807-1808, 2, 132.5-414.7 mm standard length, Iraq, Mosul (36º20'N, 43º08'E); BM(NH) 1968.12.13:440-442, 3, 178.9-351.2 mm standard length, Syria, Euphrates River at Mayadine (35º01'N, 40º27'E).

Percichthyidae

Genus Morone
Mitchill, 1814

Morone saxatilis
(Walbaum, 1792)

Introduced to the Caspian Sea basin by Soviet authorities (McNeil, 1979) but no Iranian record.

Centrarchidae

 Genus Lepomis
Rafinesque, 1819

Lepomis macrochirus
Rafinesque, 1819

The bluegill sunfish, a North American species, was introduced to the Yengi Kand Reservoir of the Namak Lake basin in 1969 from the U.S.A. (Andersskog, 1970; Anonymous, 1970a; Saadati, 1977; see also under Pimephales promelas) but the population perished through winterkill (R. J. Behnke, in litt., 1979). The locality mentioned in Saadati (1977), the Golpayegan Reservoir, is an error (R. J. Behnke, in litt., 1979). Hoffman (1970) mentions that a disease-free certification was required for importation of this species to Iran, indicating that the introduction was deliberate.

Genus Micropterus
Lacepède, 1802

Micropterus salmoides
(Lacèpede, 1802)

This North American species was introduced to the Yengi Kand Reservoir in the Namak Lake basin and possibly the Tigris River basin in the Ardakan-Yasuj area (Saadati, 1977; A. Shiralipour, pers. comm., 1978; see also under Pimephales promelas). The Namak Lake basin population died because of winterkill (R. J. Behnke, in litt., 1979) and the survival of the Ardakan-Yasuj population is not known. Its Farsi name is khorshid mahi baleh kuchek (= small-finned sunfish) or bas dehan or dahan bozorg (= largemouth bass). Heidinger (1976) summarises the biology of this species.

Percidae

The perch, darters, pike-perches and their relatives comprise a family of mostly freshwater species found across the Northern Hemisphere. There are 10 genera and about 201 species total with about 187 in North America alone (Nelson, 2006). Maximum size approaches 1 m but many species (the darters particularly) are small.

They are characterised by ctenoid scales; a dorsal fin with an anterior spiny portion and a soft rayed posterior portion; an anal fin with 1-2 spines (rather than 3 as in related families) and a few soft rays; pelvic fins thoracic in position, with 1 spine and 5 soft rays; branchiostegal membranes not attached to the isthmus; branchiostegal rays 5-8; teeth on the jaws, vomer and palatines in patches, sometimes with canine teeth; and the operculum has a sharp spine.

Perches are found in warm southern waters to subarctic ones, in both flowing and still water. Some larger species are commercially important while smaller species make attractive aquarium fishes. The small darters of North America rival coral reef fishes for colour when in breeding condition. Perches have a variety of reproductive strategies which include broadcasting, stranding, burying, attaching, clumping and clustering. During the breeding season tubercles develop, particularly on the male. These may be on the body, fins or head and are used to maintain contact and enhance grip between males and females during the spawning act.

Genus Perca
Linnaeus, 1758

This genus comprises 2 species, one found in North American and one in Eurasian, fresh waters.

The body is compressed, scales are small and ctenoid, cheeks and gill covers are scaled, the opercular bone carries a single flat spine, the preopercle is serrated posteriorly and has spikes ventrally, there are no canine teeth, branchiostegal rays 7, the lateral line does not continue onto the caudal fin, and the body usually has strong bars.

Perca fluviatilis
Linnaeus, 1758

Perca fluviatilis from Wikimedia Commons.

Perca fluviatilis from Wikimedia Commons.

Common names

mahi-ye khardar, bacheh suf (= baby suf), mahi suf rudkhanehi Astrakhan (= Astrakhan river suf fish, presumably an old name at this Russian locality), سوف حاجي طرخان (= suf-e Haji Tarkhan meaning Astrakhan suf, an old name no longer in use), suf-e rudkhaneh'i (= river suf), hashtarkhan suf.

[xanibaligi in Azerbaijan; okun' in Russian; perch, European perch, Eurasian perch, river perch].

Systematics

Perca fluviatilis was originally described from Europe.

Collette and Bănărescu (1977) refute earlier workers who maintain that this species and the North American yellow perch (Perca flavescens (Mitchill, 1814)) were the same or at best subspecies, e.g. see Svetovidov and Dorofeeva (1963) and Čihar (1975) for opposing views. Collette and Bănărescu (1977) base their conclusion on the observation that the predorsal bone is anterior to the first neural spine in fluviatilis rather than extending between the first and second neural spines as in flavescens. Other characters also exist to separate the two species. Data on the North American perch cannot therefore be used uncritically as a summary of biology for the Iranian perch.

Key characters

Characters of the genus serve to identify the single, distinctive species.

Morphology

Lateral line scales 40-78; scale rows above lateral line 7-10; rows below lateral line 12-22; and predorsal scales 10-21. Scales have very fine circuli, few anterior radii, a posterior focus and a markedly incised anterior margin where about 5-7 radii terminate. The exposed part of the scale is coarse and is the base for ctenii, best developed on the margin. Dorsal fin spines 12-18; dorsal soft rays 8-17, usually 12-15, after 0-5, usually 2-3 spines; anal fin soft rays 6-11 after 1-3 spines; pectoral rays 9-17; and pelvic rays 4-6, usually 5 after 1 spine. Gill rakers 23-25, reaching between the third and fourth rakers below when appressed usually but variable in length with diet, shortest when feeding on fish, longer when food is zooplankton. The gut is s-shaped with a large anterior loop and there are 3 pyloric caeca. Vertebrae 38-44, and gill rakers 14-29. The chromosome number is 2n=48 (Klinkhardt et al., 1995).

Meristic values for Iranian specimens are:- lateral line scales 59(3), 60(2), 61(2); scale rows above lateral line 9(7); rows below lateral line 17(4), 18(2), 19(1); scales between lateral line and pelvic fin 6(2), 7(5); predorsal scales 10(1), 11(2), 12(1), 13(2), 14(1); and caudal peduncle scales 24(1), 25(1), 26(1), 27(3), 28(1); dorsal fin spines 14(2), 15(1), 16(4); dorsal soft rays 13(2), 14(5) after 2 spines; anal fin soft rays 8(3), 9(3), 10(1) after 2 spines; pectoral rays 11(2), 12(2), 13(3); pelvic rays 5(7) after 1 spine; vertebrae 40(2), 41(3), 42(1).

Sexual dimorphism

Males have longer paired fins than females and are brighter in colour. Females are larger than males of the same age.

Colour

Colour can be affected by diet, especially in the fins which are reddest when feeding on certain crustaceans, and by habitat depth but generally the colour is stable. Fish from along the shore in weedy habitats are greenest, those in open water a pale yellow, and at depth are darker. The body is an overall greenish-yellow with 5-9 black bars on the flanks. The first dorsal fin is grey with black markings on the membranes. The first spine is often black and deep black membranes are evident between spines 1 and 2 and the last 4 to 5 spines. The second dorsal fin is greenish-yellow with melanophores on the rays and membranes, the pectoral fin yellowish and other fins pinkish to yellow to silvery-white. Paired and caudal fins have much sparser melanophores than the second dorsal fin. The lower part of the caudal fin is orange to red. Peritoneum silvery and speckled with melanophores.

Size

Reaches 68.0 cm and possibly 10.4 kg but most are much smaller than this.

Distribution

Found from the British Isles across northern Eurasia to eastern Siberia. Their presence in the Caspian Sea basin of Iran is their most southerly natural distribution. Also introduced to South Africa, Australia and New Zealand.

In Iran it has been reported from the Anzali Mordab (= Talab) and its outlets, Ab Kenar and Siah Keshim Protected Region in the Anzali Talab, at Bandar Anzali, Bandar Anzali beach, the lower Safid River and the Caspian Sea near Bandar Anzali (Derzhavin, 1934; Holčík and Oláh, 1992; Riazi, 1996; Abbasi et al., 1999; Abdoli, 2000; Abdoli and Naderi, 2009). Jolodar and Abdoli (2004) restrict its presence to the Anzali lagoon and rivers draining into it. Anderson (1880) reports perch to be abundant in the Lar River near Tehran, but this is probably a misunderstanding at this early date.

Zoogeography

Its closest relative is found in North America and they were once thought to be the same species on both continents.

Habitat

Distribution is limited by an inability to survive a temperature of 31°C for more than a few hours, by an inability to tolerate salinities above about 10-12‰ and by avoidance of waters with an oxygen level of less than 3 ml/l. The upper lethal temperature is 33.5°C (Collette et al., 1977). It is not found in the Caspian Sea proper because of the latter limitation. Fresh water is required for spawning. Riazi (1996) reports that this species is native (resident) to the Siah-Keshim Protected Region of the Anzali Mordab and it is also reported from swamps near Hendeh Khaleh in Gilan at about 37°23'N, 49°28'E.

Optimal conditions are large, weed-free, moderately deep, mesotrophic waters with food fish such as Rutilus rutilus readily available. Turbidity is a limiting factor for this species which depends on sight to feed. It is found only in the lower reaches of rivers along the Iranian shore and does not penetrate upstream (Berg, 1948-1949). Nevertheless it can be found in both running and still water and in both small and large water bodies.

Perch are a schooling fish, arranged by size and age. Schools form in the morning and disperse as dusk falls. Schools usually number about 50-200 fish but schools in the thousands are reported. There is a nocturnal resting area and perch move from it to a diurnal active area. The perch may move short distances within a lake and in large water bodies over 90 km but show strong homing tendencies. Seasonal movements are between feeding, spawning and overwintering areas.

Different morphs are found in some areas, depending on habitats: one small, slow- growing, dark and gregarious, feeding on small crustaceans, and found in reed beds, the other large, fast-growing, light and solitary, feeding on fish, and found in open waters.

Populations in the Safid River were supposedly increased after construction of the dam which reduced water flow and raised temperatures.

Age and growth

Maturity in males is usually attained in the second year of life (at only 5-12 cm long) with females maturing 1-2 years later (at 12-18 cm or larger). However some males may mature during their first year or as late as their third. Females grow slightly faster than males after the first 1-2 years. Eutrophication may reduce the age of first maturity because of increased growth rates. Perch in different habitats within the same water body, e.g. weeds beds as opposed to open water, will show different growth rates and body forms. Growth over the whole range of the species varies markedly. Generally fish at age 2 which are greater than 20 cm total length are characterised as having very good growth, moderate growth would be fish at age 3 greater than 16 cm total length while a very poor growth would be evidenced by all fish in the population being less than 16 cm total length. Life span is up to at least 21 years and under artificial conditions up to 27 years, perhaps even 50 years. A maximum age of 11 years is given for a Volga delta population examined by Makarova (1986).

Nezami et al. (2004) found fish in the Amirkelayeh Lagoon on the Caspian coast of Iran were in age groups 1+ to 6+, a total length of 9.5-33.5 cm and a weight of 10.5-350.0 g. Heydarnejad (2009) gave the length-weight relationship for an Iranian sample as W = 0.0145TL3.011.

Food

Food for small perch is zooplankton such as rotifers, switching to insect larvae, crustaceans, molluscs and leeches with growth (larger than about 20 mm). Growth is enhanced if fish and crayfish are available. Fish predominate in the diet at a range of sizes between 10 and 25 cm. In the Caspian basin, Rutilus rutilus, Blicca bjoerkna, Pungitius platygaster and gobies (Gobiidae) are eaten (Makarova, 1986). Some slow-growing perch may feed on plankton until 2-3 years old. Cannibalism is common. Maximum feeding levels occur in summer and by autumn has fallen to a maintenance level (Collette et al., 1977; Popova and Sytina, 1977). Feeding is a daylight and highly visual activity. Feeding is more effective in shoals as the perch attempts to seize other fish by the head and, if an individual perch misses, then other members of the shoal have an opportunity to seize the prey. Large perch lie in wait for passing prey items and then dart out to seize them. Unlike northern pike, perch will pursue a prey item if it tries to escape.

In the Amirkelayeh Lagoon diet varied according to age, season and sex, and comprised a wide variety of organisms such as water bugs, odonates, gammarids, plant materials, chironomids, Tinca tinca, hemipterans, Perca fluviatilis, snails, Syngnathus caspius, Gambusia holbrooki, Pungitius platygaster, dipterans, branchiopods, trichopterans, tubifex, frogs and shrimps. The species is an omnivore here and a cannibal.

Reproduction

Spawning occurs in the spring in shallow water, 0.5-3.0 m deep, end of March to early June in the Volga Delta (Makarova, 1986). In Dagestan spawning is from the end of March to the beginning of April and lasts 10-15 days (Shikhshabekov, 1978), elsewhere only 2-3 days. Water temperatures are around 11°C on the Volga (Lönnberg, 1900b) and above 8°C in Dagestan (Shikhshabekov, 1978) but can occur under ice at 4°C. Up to 80% of the spawning population in the Volga Delta is female (Makarova, 1986). There is a spawning migration from deepwater resting areas to shallow spawning areas. Males precede females onto the spawning ground by days or weeks and remain behind after spawning. Brackish-water populations migrate into fresh water. Spawning itself can occur by day or by night.

Although temperature is the major factor affecting the timing of spawning, the occurrence of spring floods is significant in some populations as it gives access to inundation zones of large rivers.

Eggs are twisted around plants, roots and logs in an egg-strand, a cylindrical, hollow, twisted structure up to 3.75 m long, 3.8 cm thick and 8 cm wide. This structure offers protection from predators, fungal infections, desiccation, mechanical damage and smothering in the mud. Egg diameters reach 2.5 mm and fecundity 300,000 eggs. Fecundity increases with age and depends on food supply as in most fish species. As many as 15-25 males queue up to fertilize the egg-strand, following the female as she twists around the logs and plants, rubbing against the plants to void the eggs. The female drives the males away from the egg-strand after fertilisation and may guard the egg-strand for some hours.

Parasites and predators

Sander lucioperca, Esox lucius and Lota lota are predators on perch in the Caspian basin and doubtless other large fishes and birds take this species. Cannibalism begins as early as the fry stage when fish only 2.1 cm long eat smaller fry.

Mokhayer (1976b) records the protozoan Trypanosoma percae from this species in the Caspian Sea basin, the nematode larva Eustrongylides excisus and the annelid Piscicola geometra. Khara et al. (2006a) record the eye fluke Diplostomum spathaceum for this fish in the Amirkalayeh Wetland in Gilan. Sattari et al. (2002) and Sattari (2004) records the presence of the nematode, Eustrongylides excisus. This parasite can damage muscles in commercial species and render them unsuitable for sale. Sattari et al. (2004; 2005) surveyed this species in the Anzali and Amirkelayeh wetlands, recording Raphidascaris acus, Eustrongyloides excisus and Camallanus lacustris. Khara et al. (2005) examined this species in the Amirkalayeh Wetland and found its diversity of parasites to be less than other predatory species such as Esox lucius. Parasites recorded were Camallanus lacustris, Diplostomum spathaceum, Lemaea sp., Argulus sp., and Dactylogyrus sp. Sattari et al. (2007) record the nematode Eustrongylides excisus and the digenean Diplostomum spathaceum in this species in the Anzali wetland of the Caspian shore. Barzegar et al. (2008) record the digenean eye parasite Diplostomum spathaceum from this fish. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Achtheres percarum on this species.

Economic importance

Holčík and Oláh (1992) report a catch of only 15 kg in the Anzali Mordab in 1990. This species has been studied for aquaculture in Iran.

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in aquaria and aquaculture, as food, in sport, in textbooks, in experiments and because it has been widely introduced outside its natural range. It has been implicated in ichthyootoxism (Coad, 1979b), the symptoms of egg poisoning being summarised under the genus Schizothorax.

Conservation

Illegal fishing and non-standard nets threaten stocks of this species (Annual Report, 1995-1996, Iranian Fisheries Research and Training Organization, Tehran, p. 55, 1997). Kiabi et al. (1999) consider this species to be vulnerable in the south Caspian Sea basin according to IUCN criteria. Criteria include commercial fishing, sport fishing, medium numbers, habitat destruction, limited range (less than 25% of water bodies), absent in other water bodies in Iran, and present outside the Caspian Sea basin.

Further work

The status of populations of this species in Iran should be carefully monitored in view of the threats outlined above.

Sources

General biology is based on Thorpe (1977) and Craig (1987), and there is extensive European angling literature on this species.

Iranian material: CMNFI 1970-0510, 1, 156.8 mm standard length, Gilan, Golshan River (37º26'N, 49º40'E); CMNFI 1979-0685, 1, 71.5 mm standard length, Gilan, Safid River around Mohsenabad below Dehcha (no other locality data); CMNFI 1980-0123, 1, 68.9 mm standard length, Gilan, Safid River around Dehcha above Mohsenabad (no other locality data); CMNFI 1980-0127, 1, 152.8 mm standard length, Gilan, Caspian Sea near Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1980-0148, 2, 118.2-142.1 mm standard length, Gilan, Pir Bazar Roga (37º21'N, 49º33'E).

Genus Sander
Oken, 1817

This genus is found in both North America and Eurasia and contains 5 species. There are 3 species in the Caspian Sea basin, 2 of which are reported from Iran. The Shahid Beheshti hatchery on the Safid River breeds the third species, S. volgense (Gmelin, 1789), a northern Caspian Sea species, according to Raymakers (2002) but it is unclear if these have been released and have established populations in the southern Caspian Sea.

The genera Stizostedion Rafinesque, 1820 and Lucioperca Schinz in Cuvier, 1822 are junior synonyms of Sander (see Kottelat, 1997). The pike-perches are elongate and compressed, have large jaws reaching back beyond mid-eye level, canine teeth on the jaws and palatines, the preopercle is serrated posteriorly and has spines ventrally, the opercle has a weak, flat spine at its postero-dorsal corner, cheeks are naked or scaled only dorsally, there are 7-8 branchiostegal rays, adult gill rakers are densely denticulated tubercles, in young denticulated rods, and the lateral line continues onto the caudal fin with accessory lateral lines on the upper and lower caudal lobes. Divergence between North American and Eurasian members of this genus may have occurred in the middle to late Miocene or in the Pliocene (Billington et al., 1990; 1991; Faber and Stepien, 1998).

Sander lucioperca
(Linnaeus, 1758)

Sander lucioperca from Wikimedia Commons.

Sander lucioperca from Wikimedia Commons.

Common names

سوف (= suf), سوف معمولي (= suf-e ma'muli or soof-e-maamooli, meaning common suf), sibey(ak) in Gilaki, ? sevideh.

[sif, caysifi or quaysifi in Azerbaijanian; adaty silebalyk in Turkmenian; sudak in Russian; pike-perch, European pikeperch, zander].

Systematics

Perca Lucioperca was originally described from European lakes. This may be the fish referred to as sevideh from the Langarud Mordab by Holmes (1845). However the word may be sepideh which means "the white one" and may in fact refer to Rutilus frisii, the safid mahi. Akbarzadeh et al. (2007, 2009) identified three distinct populations in the southern Caspian Sea using a truss analysis with morphometric characters (body heights and caudal peduncle measures) showing better discrimination than meristic characters. Gharibkhani et al. (2009) examined the population genetic structure of fish from the Talesh coast, Anzali Wetland and Chaboksar coast and found them to be genetically differentiated, with consequences for conservation.

Key characters

This species is separated from S. marinum by the dorsal fins being close together, obviously much less than the eye diameter apart, the anal fin spines are not closely joined to the soft rays, the interorbital width is less than, or equal to, the vertical eye diameter in adults, the upper jaw extends rearwards on a level behind the posterior eye margin in adults (under the rear of the eye in young), there are usually more than 18 soft rays in the dorsal fin, and the spiny dorsal fin bears large spots.

Morphology

Lateral line scales 75-150 (this wide range from various literature sources presumably includes counts of scales in the lateral line, and other counts which are of smaller scales to one side of the lateral line; Berg (1948-1949) gives a count of 80-97 for example, which is inherently more reasonable). Scales above the lateral line 12-17 and scales below the lateral line 16-24. Scales are incised on the anterior margin where about 6-9 radii terminate, although not as incised as in Perca fluviatilis. The focus is posterior. Ctenii are well-developed. First dorsal fin spines 11-17; second dorsal spines 1-4 and soft rays 16-27, usually 19-24. Anal fin spines 1-3, soft rays 9-14. Pectoral fin branched rays 11-18 and pelvic fin branched rays 5. Stubby, spinulose gill rakers number 10-17 and vertebrae 40-48, usually 45-48, mode 46. There are 4-9 pyloric caeca. The gut is relatively short but has a long anterior loop. The chromosome number is 2n=48 (Klinkhardt et al., 1995; Miri Nargesi et al., 2007) although the latter authors found a different karyotype formula in Iranian fish compared to other populations (2n=1m + 13sm + 4st + 6a).

Meristic values for Iranian specimens are:- lateral line scales 82(1), 86(1), 89(1), 90(1), 91(3); scales above the lateral line 13(1), 14(4), 15(1); scales below the lateral line 21(2), 22(1), 24(2), 25(1); caudal peduncle scales 33(3), 34(3); first dorsal fin spines 13(3), 14(4); second dorsal soft rays 20(2), 21(2), 22(1), 23(2); anal fin soft rays 11(4), 12(1) 13(2); pectoral fin branched rays 13(2), 14(2), 15(3); pelvic fin branched rays 5(7); and vertebrae 45(3) or 46(3).

Sexual dimorphism

Females have a much lighter, whitish belly than that of males which is marbled blueish in the spawning season. The genital papilla of females protrudes.

Colour

The back and flanks are green to blue-grey to brown-black, the belly is white to bluish and fins are yellow-grey. The dorsal and caudal fins have rows of black spots on the membranes, largest and most distinctive on the spiny dorsal fin. Other fins are pale yellow. The eye is silvery because of reflection from the tapetum lucidum. Young have 8-13 brown to blackish-brown bars but these usually fade with maturity. The peritoneum is silvery to brownish in preserved fish.

Size

Up to 1.5 m and 20.0 kg. The Iranian commercial catch in the 1950s was 24-60 cm long and weighed 1.6-2.7 kg, declining to 0.6 kg (Farid-Pak, no date).

Distribution

Found in the basins of the Baltic, Aegean, Black, Caspian and Aral seas. In Iran reported in the Aras River and along the whole Caspian coastal plain from Astara to the Atrak River including the Ameerkalaye Lagoon near Lahijan, the Gorgan, Qareh Su, Neka, Babol, Haraz, Safid and middle Aras rivers, as well as in the southeast Caspian Sea, southwest Caspian Sea and south-central Caspian Sea, Gorgan Bay, Anzali Mordab and Siah Keshim Protected Region (Nedoshivin and Iljin, 1929; Derzhavin, 1934; Kozhin, 1957; Griffiths et al., 1972; Holčík and Oláh, 1992; Nejatsanatee, 1994; Riazi, 1996; Abbasi et al., 1999; Kiabi et al., 1999; Abdoli, 2000; Jolodar and Abdoli, 2004; Abdoli and Naderi, 2009).

This species has also been stocked in the Zarreinerud, 70 km upstream of Miandoab in 1971 (Griffiths et al. (1972), to Valasht Lake near Marzanabad, Evan Lake northeast of Qazvin, Ghorigol Lake near Tabriz, Marivan Lake in Kordestan and the Haft Barm Lakes west of Shiraz (Anonymous, 1977). Although this species has been introduced to the Manjil Reservoir on the Safid River (Griffiths et al. (1972), this reservoir is drained to remove excess silt and no fishery exists (J. Holčík, pers. comm., 1992). Introductions to reservoirs in Khuzestan did not survive (M. Al-Mukhtar, pers. comm., 1995).

It is also found in the Karakum Canal and Kopetdag Reservoir of Turkmenistan (Sal'nikov, 1995) and may eventually reach the Iranian Tedzhen (= Hari) River basin.

Zoogeography

The relationships of this species are discussed under the genus.

Habitat

Generally this species is found in small schools near sandy and stony bottoms in deeper water of rivers. Ideally there should be some concealment. It can also live in reservoirs. This species has both freshwater and semi-anadromous forms in the Caspian Sea basin. It has a limited migratory behaviour such that morphologically distinct stocks may exist in larger water bodies. The population of the Anzali Mordab represents a separate stock (Hydrorybproject, 1965). Riazi (1996) and Karimpour (1998) report that this species migrates into the Siah-Keshim Protected Region of the Anzali Mordab and is also resident there. Most movements of this species are within 10-20 km although distances up to 300 km have been recorded in the Volga River. Suf show strong homing tendencies. The upper lethal temperature is 35°C. High turbidity levels are preferred. During the day, suf shelter from strong light by descending in the water column (Collette et al., 1977; Marshall, 1977). Knipovich (1921) reports this species from depths of 11.0-11.9 m, possibly deeper, in the Iranian Caspian Sea. In Dagestan this species prefers areas where there is flowing water well-supplied with oxygen (Shikhshabekov, 1978) and avoids vegetation and therefore competition with Esox lucius. Suf will tolerate low salinities and can be found around river mouths in the Caspian Sea basin but the sea itself is too saline.

Age and growth

Males mature at 2-6 years (32 cm) and females at 3-6 years (42-44 cm). Life span may exceed 19 years, although in Lake Eğirdir, Turkey only 7 age groups were recorded (Becer and Ikiz, 1999a). Optimum growth occurs at 28-30°C (Marshall, 1977).

During 1932-1933 in the Anzali Mordab, 5-7 year old fish dominated in catches and weighed 2.6-7.4 kg but by the 1960s this had declined to 2-5 years and 1.6-2.7 kg (Hydrorybproject, 1965). Catches in 1971/72 in the commercial fishery of Iran were 3-7 years old, 33.0-55.0 cm long and weighed 370-2100 g (Razivi et al., 1972). Abdolmalaki (2005a) found age groups in the Iranian Caspian to be 2-5 years with 2-3 year olds forming 78.5% of the catch. The von Bertalanffy growth equation was Lt = 52.5*[1-exp-0.158*(t + 1.852)]. The instantaneous rate of total, natural and fishing mortality was 0.95 year-1, 0.31 year-1 and 0.64 year-1 respectively. The calculated exploitation ratio was 0.67, the estimated biomass was 31.56 tons, the minimum sustainable yield was 13.89 tons (lower than the total catch), and the fishery return coefficient was 2.87%. Abdolmalaki and Psuty (2007) report 6 age groups for coastal waters of the southern Caspian Sea, the length-weight relationship was w = -0.020606L2.85, and the von Bertalanffy parameters were L = 55.05 cm fork length (substantially less than in the Volga River delta at 79.0 cm and Aras Lake in Iran at 73.3 cm), K = 0.15, t0 = -2.59 and M = 0.31. More than 90% of the beach-seine caught fish were smaller than the minimum legal length. These authors also provide details of recruitment and fishing mortality for this population which is enhanced by introduction of fingerlings. Rahimibashar et al. (2008) found that the fish in the Aras Dam lake had allometric growth and age classes caught with cast nets and gill nets were 2+ to 5+ years.

Food

The suf is an ambush-pursuit predator. Feeding on fish begins at a length of 5-10 cm (2-3 months of age) depending on the relative abundance of zooplankton, invertebrates and forage fish. In the Volga Delta, spawning Rutilus rutilus  (presumably R. caspicus) in April-May is the most important food, up to 80% of the annual ration. In the 1960s and 1970s when the population of suf was 7 million fish, they ate 53,000 tonnes of Rutilus rutilus (presumably R. caspicus) (Caspian Sea Biodiversity Database, www.caspianenvironment.org). Adults are solitary but young fish feed in schools on nauplii, copepods and some rotifers. Some adults are cannibals (Collette et al., 1977; Marshall, 1977; Popova and Sytina, 1977) and Balik (1999) reports that in Lake Beyşehir, Turkey, a suf can eat its own species with a mean size of 35.9% of its length. Apparently many prey fish are seized and swallowed tail first. One Iranian specimen contained a Neogobius melanostomus and gobies are an important food item generally in the Caspian Sea, 17.8% of the diet compared to 59.9% for Rutilus rutilus (presumably R. caspicus).

Rahimibashar et al. (2008) found that the fish in the Aras Dam lake were carnivorous, almost gluttonous, feeding principally on bony fishes (92% food preference) with some crab larvae and aquatic insects.

Reproduction

The spawning migration begins in late March-early April in Dagestan with spawning in early to mid-April (Shikhshabekov, 1978). In Eğirdir Lake, Turkey spawning took place from April to June and Becer and Ikiz (1999b) give details of fecundity, egg diameters, and the relationships between length, weight and gonad weight and fecundity for fish that mature as young as ages 1-2. The spawning season over the range of this species is late February to late July, usually April-May at 12°C (range 6-22°C) as deep as 17 m. There are distinct spawning stocks.

In the Anzali Mordab, the main spawning area in the southern Caspian, the spawning run usually starts in the first 10 days of March at water temperatures of 8.0-9.5°C, ending at 12-14°C (Hydrorybproject, 1965; Razivi et al., 1972). Apparently, natural spawning has stopped completely in the mordab and this lagoon is stocked with fingerlings from spawners held at Aras, a border reservoir lying between Iran and Azerbaijan (Abdolmalaki and Psuty, 2007). Males build nests in depths of 30 cm to several metres on hard bottoms usually in turbid water. Each nest is a flat pit edged by gravel or shell. Plant roots are often exposed as a spawning substrate on which eggs are laid individually. The nest is guarded by the male and eggs are fanned. The female is driven away after spawning. Male suf are so devoted to protecting the nest that they will remain on site even if water levels fall and their backs stick out of the water. In addition they will try to bite humans if they approach the nest. Spawning is intermittent over several days and usually takes place at dawn. Maximum fecundity is 2.5 million eggs and egg diameters are up to 1.5 mm. Hatching occurs from 4 to 26.5 days, depending on temperature (Collette et al., 1977; Marshall, 1977). Females descend to the sea first from the Anzali Mordab after spawning and fry there are 19-33 mm long by the end of May (Hydrorybproject, 1965).

Parasites and predators

Eslami and Mokhayer (1977) examined 100 specimens of suf and found 20% to be infested with larvae of the nematode Anisakis. Ataee and Eslami (1999, www.mondialvet99.com, downloaded 31 May 2000) report the helminth Anisakis from the gastro-intestinal tract of fish from the Anzali wetland. This parasite can infest man if fish is eaten smoked, salted or fried at temperatures below 50°C. Mokhayer (1976b) records the acanthocephalan Corynosoma caspicum. Jalali and Molnár (1990a) record the monogenean Ancyrocephalus paradoxus from this species in the Safid River. Masoumian et al. (2005) recorded the protozoan parasite Trichodina perforata from this species in the Aras Dam in West Azarbayjan. Pazooki et al. (2007) recorded various parasites from localities in West Azarbayjan Province, including Diplostomum spathaceum and Argulus foliaceus from this species. Barzegar et al. (2008) record the digenean eye parasite Diplostomum spathaceum from this fish. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Achtheres percarum on this species.

The Caspian seal, Pusa caspica, is a significant predator on this species (Krylov, 1984) as are predatory fishes such as Esox lucius, Perca fluviatilis and Silurus glanis. Adult suf have few predators.

Economic importance

There is some opportunity for sport fishing for this species in the Anzali Mordab and potentially in various lakes around the country where it has been introduced (Anonymous, 1977). It is a very popular food fish in Iran (Razivi et al., 1972). It has also been studied in Iran as a control species for undesirable fishes (Annual Report, 1995-1996, Iranian Fisheries Research and Training Organization, Tehran, p. 80, 1997).

Nevraev (1929) reports on catches in various regions of Iran in the early years of the twentieth century. There were no evident trends of increase or decrease. In the Astara region from 1901-1902 to 1913-1914 the catch varied irregularly from 154 to 31,931 fish, in the Anzali region from 1901-1902 to 1918-1919 the catch varied from 608,300 to 3,367,000 fish, in the Safid River region from 1899-1900 to 1917-1918 the catch varied from 9983 to 125,182 fish, and in the Astrabad region from 1900-1901 to 1912-1913 the catch varied from 1400 to 22,900 fish.

Stocks of this species are known to fluctuate in Iran, as obviously do the catch statistics. Most fish are caught in beach seines although some are caught in gillnets, both legally and illegally (see below). The main fishing ground is coastal waters in the Anzali region. Catches in the 1920s were at 3000-4000 tonnes for the coastal zone of the southern Caspian Sea but declined drastically afterward (Razavi, 1999). The commercial catch in Iran from 1956/1957 to 1961/1962 varied from 206 kg to 20,945 kg (Vladykov, 1964; RaLonde and Walczak, 1972), from 1965/66 to 1968/69 it varied from 7 to 77 tonnes (Andersskog, 1970) and from 1963 to 1967 ranged from 0 to 14.6 t (RaLonde and Walczak, 1970b). In the 6 years from 1980 to 1985 catches were recorded by the Food and Agriculture Organization, Rome as respectively 0, 0, 0, 12, 13 and 10 t. Catches in 1990 were about 5-10 t and in 1996 about 35-40 t (Bartley and Rana, 1998b). In 2000-2001, the catch was 18 t or 11% of the total commercial catch in the Iranian Caspian Sea basin. 12 t were caught by beach seine along the coast, 3 t were taken in the Anzali Mordab and the rest was an estimated amount of unlicensed captures (Abdolmalaki, 2005a). In 2003-2004 the catch was 38 t, a decrease in comparison to the previous year, with 15 t of this from beach seine cooperatives. Most fish were immature and undersized and the catch was based on release of fingerlings (Abdolmalaki, 2006).

Summaries of catches of this species in the coastal southern Caspian Sea over 8 decadal periods is given in Fisheries. Recent catches from Abdolmalaki and Psuty (2007) are as follows:-

Year

Total catch (t)

Catch by beach seine (t)

Number of beach seine cooperatives

Beach seine efforts (hauls)

Number of illegal gillnets confiscated

1990

4.0

4.0

68

20.975

No data

1991

12.3

12.3

81

27,200

104,828

1992

10.0

8.7

88

30,239

109,446

1993

12.3

7.3

93

33,986

138,026

1994

40.2

22.6

91

27,868

215,381

1995

10.1

4.0

101

34,055

204,831

1996

8.0

2.8

109

42,847

270,727

1997

8.1

2.9

111

45,263

205,999

1998

95.0

54.8

125

52,574

222,897

1999

17.5

11.5

139

50,953

130,849

2000

18.0

12.0

147

56,913

82,678

2001

26.0

21.5

150

60,006

113,729

2002

30.0

20.3

150

57,310

141,506

2003

23.8

15.0

151

53,846

179,656

2004

22.5

14.4

151

49,809

261,875

In the period from 1933/34 to 1961/62 in the Bandar-e Anzali region catches varied from about 3483 t at the earlier date to 33 kg at the later one, with large variations between years. Holčík and Oláh (1992) report a catch of only 22 kg in the Anzali Mordab in 1990, and from 1932-1964 reported catches varied from 1 to 2581 t annually.

Hedayatifard and Jamali (2008) showed that this fish is a good source of polyunsaturated fatty acids and one of the best sources for omega-3 fatty acids, useful in preventing cardiovascular disease.

Khaval (2007) investigated polyculture of this species with silver, bighead, grass and common carp at the Safidrud Fisheries Research Station. Carp density was 3000 fish per hectare with 60% silver carp, 20% grass carp and 10% each for common and bighead carp. Suf weighed 2.1 g and were stocked at 250 fish per hectare. Survival rates were 93.33% for the suf and 83.77% for the Chinese carps. Over the period April to November, suf fingerlings reached 54.4 g on average and production was 4446.66 kg per hectare compared to 3212.8 kg without polyculture. Ghafouri Salah et al. (2008) studied physiological stress in this species and its effects on cortisol and muscle compounds.

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in aquaria and aquaculture, as food, in sport, in textbooks, and because it has been widely introduced outside its natural range. The suf or pike-perch is also fish-farmed in Europe for stocking purposes as eggs or young.

Conservation

The decline in catches from about 3000-4000 tonnes to about 30 t noted above has never been adequately explained. Overfishing, degradation of spawning grounds, fluctuating water levels and salinity variations in coastal waters, the volume of freshwater input, oxygen concentrations and input of organic material to estuaries and the sea, are all possible factors (Abdolmalaki and Psuty, 2007). The stocks of this species in Iranian waters also declined in the 1960s (Walczak and RaLonde, 1970; RaLonde and Walczak, 1972). Causes were reduction in estuarine habitat needed for spawning, man-made habitat changes and overfishing of the younger age classes and first year spawners. The catch declined from 125 tonnes annually in the 1940s to 14.6 t in 1967. Vladykov (1964) considered stocks all along the Iranian shore to be at dangerously low levels. Griffiths et al. (1972) suggested that stocks in Iran were on the verge of extinction and recommended a three-year ban on catching this species. Artificial raising of this species is difficult but more than 6 million fingerlings were raised and released into the Anzali Mordab in the Iranian year 1991-1992, a 100% increase over the previous year (Abzeeyan, Tehran, 4(2):VI, 1993). Fingerling production rose from 0.12 million in 1990, to 1.50 million in 1991 and 2.50 million in 1992 (Emadi, 1993a)(note that Matinfar and Nikouyan (1995) give 1.63 and 2.44 million fingerlings for 1991 and 1992). Fingerling production in 1995 was 2.269 million, in 1996 2.4 million and for 1996-1997 8 million (Bartley and Rana, 1998a; 1998b). The Sturgeon International Research Institute, which opened in 1994 near Rasht, released 5-8 million fry in 1996-1997 (Bartley and Rana, 1998b). The release in 1999 numbered 5 million "juveniles" (I.F.R.O. Newsletter, 23:4, 2000). Billard and Cosson (2002) cite an annual production of 5-10 million, mostly released in the Anzali lagoon and Moghaddam (2006) gives a figure of 5.13 million fingerlings for 2002. The highest number of fingerlings released in the Anzali Lagoon was 6,604,000 in 2003 with the lowest being 1,160,000 in 1993 for the period 1991-2003 according to Abdolmalaki and Psuty (2007). The latter authors also note that the beach seines used in Iran (see Fisheries in Introduction) do not protect young fish. There is a heavy mortality of discarded fish even when legal landing size is enforced and resources are inadequate to manage the fishery effectively. The minimum mesh size of the cod-end of the seines should be increased and its use monitored.

Ramin (1996) has studied semi-artificial propagation and rearing of fry of this species in Iran. Broodstock spawning occurs in March-April at 12-14°C on artificial nests of green wool bunches on wooden frames placed in ponds at 5 m intervals. Nests close to the bottom are preferred and eggs are dropped on them with an average fertilization of 30-90%. The nests with eggs on them are kept in a mist chamber and the eggs collected and placed in jars. Eyed eggs appear on day 3 or 4 of incubation. Yolk-sac absorption lasts 9-13 days and exogenous feeding fry measure 4-6 mm.

Lelek (1987) classifies this species as intermediate to vulnerable in Europe. Kiabi et al. (1999) consider this species to be vulnerable in the south Caspian Sea basin according to IUCN criteria. Criteria include commercial fishing, sport fishing, few in numbers, limited range (less than 25% of water bodies), absent in other water bodies in Iran, and present outside the Caspian Sea basin. Nezami et al. (2000) consider this species to be endangered because of overfishing, habitat destruction and spawning ground degradation.

Further work

The numbers of this species should be actively surveyed because of threats outlined above.

Sources

Deelder and Willemsen (1964) reviewed the biology of this species as did Craig (1987). Robins (1970) gave a bibliography of the genus Stizostedion (= Sander).

Iranian material: CMNFI 1970-0532, 1, 209.3 mm standard length, Gilan, Caspian Sea near Bandar Anzali (37º28'N, 49º27'E); CMNFI 1979-0431, 2, 211.5-241.0 mm standard length, Mazandaran, Now Shahr fish bazaar (no other locality data); CMNFI 1979-0455, 1, 68.5 mm standard length, Markazi, Manjil Dam (36º45'N, 49º17'E); CMNFI 1980-0127, 1, 266.1 mm standard length, Gilan, Caspian Sea near Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1980-0130, 1, 197.4 mm standard length, Mazandaran, river near Iz Deh (36º36'N, 52º07'E); CMNFI 1980-0150, 1, 280.8 mm standard length, Gilan, Caspian Sea at Safid River estuary (37º24'N, 49º58'E).

Sander marinus
(Cuvier, 1828)

Common names

suf-e daryai (= sea suf), souf-e-daryaee.

[daniz sifi in Azerbaijan; morskoi sudak or sea pike-perch, erroneously bërsh by fishermen in the Caspian Sea, both in Russian; sea zander, estuarine perch].

Systematics

Lucioperca marina was originally described from the Black Sea at Feodosiya. No types are known (Eschmeyer et al., 1996).

Key characters

This species is distinguished from S. lucioperca by the dorsal fins being well-separated, but by a distance less than eye diameter, anal fin spines are weak and closely joined to the soft rays, interorbital width in adults is much greater than eye diameter, the upper jaw extends back level with the posterior pupil edge or almost to the posterior eye edge in adults, the dorsal fin soft rays are 18 or less, and the spiny dorsal fin lacks large spots.

Morphology

First dorsal fin spines 12-14, second dorsal fin spines 1-4 followed by 12-18 soft rays, anal fin spines 2-4 followed by 9-12 soft rays, pectoral fin with 1 spine and usually 15 soft rays, and pelvic fin with 1 spine and 5 soft rays. Lateral line scales 75-88, scales above the lateral line 9-13 and caudal peduncle scales 34-36. Scales have few anterior radii, a crenulate anterior margin and a posterior focus. Gill rakers on the upper arch number 12-20, on the lower arch 11-17, reaching the second raker below when appressed, elongate and spinulose. Vertebrae number 38-44. There are 5-7 pyloric caeca. Cheeks are scaleless or almost scaleless. There are canines on the jaws and palatines. These counts are combined for several literature sources and include eastern and western populations of the southern Caspian Sea which show evident differences indicating distinct stocks. The gut is short and s-shaped.

Meristic values for Iranian specimens are:- first dorsal fin spines 13(2) or 14(1), second dorsal fin spines 2 (3), soft rays 16 (3); anal fin spines 2(3), soft rays 12(3); soft pectoral rays 15(3), pelvic fin with 1(3) spine and 5(3) soft rays, lateral line scales 79(2) or 82(1), scales above lateral line 10(1), 12(1) or 13(1), caudal peduncle scales 34(2) or 36(1); and total gill rakers 12(1), 13(1) or 15(1).

Sexual dimorphism

Unknown.

Colour

The back is light grey and the flanks have 12-15 dark bars, which are sometimes indistinct and may be absent, e.g. in females from the eastern Caspian which had small, irregular, dark-brownish speckles. Some fish are almost black and lack bars. The first dorsal fin lacks the strong spots seen in Sander lucioperca and is dark grey to black with patches of concentrated melanophores and clearer areas forming irregular and incomplete stripes, or darkly fringed and with a dark spot at the posterior base. The second dorsal fin and the caudal are finely spotted. Other fins are grey with some melanophores on rays. Eyes are silvery due to the tapetum lucidum. The peritoneum is brownish.

Size

Attains 62 cm (65 cm total length in Jolodar and Abdoli (2004)) and 2.2 kg.

Distribution

Found only in the northwestern Black Sea and the Caspian Sea. Reported from near Gasan-kuli in Turkmenistan (Berg, 1948-1949) and from the southeast Caspian Sea in Iran (Kiabi et al., 1999). Jolodar and Abdoli (2004)  and Abdoli and Naderi (2009) record it from the central, southwestern and southeastern regions of the Caspian Sea including in Astara. An old record is cited below under Habitat.

Zoogeography

The relationships of this species are discussed under the genus.

Habitat

This species lives in the Caspian Sea proper and rarely enters rivers. De Filippi (1865), however, did record "Un molto bello e grosso individuo....in un canale di Murdab, ove l'aqua era del pari sensibilmente dolce". It favours areas with rocky bottoms and does not migrate. In winter, part of the population moves into deeper water at depths of 30 m, rarely 100 m, while another part remains near the shore. The major concentration of this pike-perch is found near the shores of Turkmenistan, and secondly of Azerbaijan.

Age and growth

Sexual maturity is attained at 3-4 years for most fish with a few fish maturing at 2 years (Guseva, 1975). Life span is at least 12 years. Growth is slightly faster in females up to age 5, evens out later and males become larger (Kuliyev, 1981). Males on the spawning grounds average 41.2 cm and females 42.9 cm, with an average weight of 0.94 kg (Berg, 1948-1949).

Food

The principal foods are gobies (Gobiidae), young herring and tyulkas (Clupeidae), silversides (Atherinidae), and crayfish.

Reproduction

The male prepares a nest site and protects the eggs. Spawning takes place at 3-12 m on open, stony bottoms or in "nest-caves" and eggs are laid in a continuous layer. The male constructs and guards the nest. Spawning usually begins in the second half of April and ends in mid-May at temperatures of 10-17°C, and is most intense at 13-15°C (Gusev, 1974a). Up to 126,000 adhesive eggs are laid (Guseva, 1974a; 1975; Kuliyev, 1981) and are larger than in S. lucioperca. Fertilised eggs are 2.6-3.8 mm and at water temperatures of 14.7°C, incubation takes 12-17 days (Gusev, 1974a).

Parasites and predators

None reported from Iran.

Economic importance

The sea pike-perch was commercially fished off the Turkmenistan coast in the 1930s and 1940s with catches of 19 thousand centners (1 centner = 100 kg) or 2,271,000 fish. In 1927-1929 the annual average on the shores of Azerbaijan was 7000 centners and in 1930-1932 10,3000 centners. In 1930 the catch for the whole Caspian Sea was 909 thousand centners (Zenkevitch, 1963). The development of offshore oil deposits has drastically reduced stocks throughout the Caspian Sea (Guseva, 1974b; 1975; Kuliyev, 1981).

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in textbooks.

Conservation

This species has been proposed for inclusion in the "Red Book of the U.S.S.R." which forms the basis for measures to protect species (Pavlov et al., 1985). Kiabi et al. (1999) consider this species to be data deficient in the south Caspian Sea basin according to IUCN criteria. Criteria include possibly few in numbers, limited range (less than 25% of water bodies), absent in other water bodies in Iran, and present outside the Caspian Sea basin. The 2000 IUCN Red List lists this species as DD (Data Deficient).

Further work

The distribution and abundance of this species in Iranian waters needs to be examined to determine its conservation status.

Sources

Morphology based in part on Svetovidov and Dorofeeva (1963) and Kuliyev (1981).

Iranian material: CMNFI 1970-0532, 3, 156.3-165.5 mm standard length, Gilan, Caspian Sea near Bandar Anzali (37º28'N, 49º27'E).

Sparidae

The sea breams or porgies are found in the Atlantic, Indian and Pacific oceans and comprise 33 genera and about 115 species (Nelson, 2006). Some commonly enter estuaries and penetrate up rivers. Maximum length is about 1.2 m.

This family is characterised by a groove in the distal end of the premaxilla which accommodates the maxilla; the body is oblong to ovate and is compressed; the head is large with a steep upper profile; the preopercle margin is smooth; scales are weakly ctenoid, moderate in size and extend on to the cheeks and operculum; teeth are conical to incisiform and molar teeth are found in some at the rear of the jaw; there are no teeth on the vomer, palatines or tongue; the dorsal fin is continuous with an anterior spiny portion and a soft-rayed posterior portion about equal in size; with 10-13 spines and 10-15 soft rays respectively; spines fold into a groove; the anal fin has 3 spines (the second the largest) and 8-14 soft rays; branchiostegal rays 5-6; branchiostegal rays 5-6; and lateral line not continued onto the caudal fin but with enlarged scales near the head.

Many species in this family are hermaphrodites with male and female sex organs developing simultaneously, changing sex from male to female (protandry), or from female to male (protogyny). These fishes are often important as food or sought by anglers. Young fish may very different in colour to adults, usually being more vividly coloured with distinctive patterns. Most species are marine (see Marine List in Checklists in the Introduction) but a few enter fresh water and penetrate a considerable distance from the sea including one species in Iran.

Genus Acanthopagrus
Peters, 1855

Members of this genus have a compressed and moderately deep body, 4-6 enlarged incisiform teeth at the front of the jaws followed by 3-4 rows of molars, the second anal fin spine is longer than the third, there is a scaly sheath at the base of the dorsal and anal soft fins, and moderate-sized scales. Iwatsuki and Heemstra (2010) revised the genus in the western Indian Ocean.

Acanthopagrus latus
(Houttuyn, 1782)

Common names

شانك (= shanak), شانك زردباله (= shanak-e zardbaleh or yellowfin shanak).

[shanak, shaghoom, shaam, sha'm, shaem, sheim or sha-om in Arabic; yellow-finned porgy or seabream, yellow-finned black porgy, Japanese silver bream].

Systematics

Sparus latus was originally described from Japan.

Al-Hassan (1990) found differences in two meristic characters (pectoral and dorsal fin ray counts) but no differences in electrophoretic characters between populations from the Shatt al Arab and Khor al Zubair areas of southern Iraq. He concludes that there is only one stock of this species in southern Iraq as meristic variation may reflect environmental conditions.

Key characters

This species is the only sparid member recorded from Iranian freshwaters and is recognised by the dorsal fin spines alternately thick and thin and the colour pattern.

Morphology

Upper profile of head steep and convex back to above the posterior eye margin. The head bulges over the eye. Dorsal fin spines 11-13, soft rays 9-13. Anal fin with 3 spines, the second much stronger and wider than the third, and 8-9 soft rays. Pectoral fin branched rays 10-16. There is a strong spine in the pelvic fin and a well-developed axillary scale. Lateral line scales 41-46, or 48-50, or up to 55 depending probably on differing counting methods. The scales are vertical ovals with the anterior margin wavy where radii intersect. They have very fine circuli, moderate numbers of posterior radii, a subcentral posterior focus, and ctenii on the central part of the posterior margin extending inwards towards the focus. Four or five series of preopercular scales. The first pelvic fin ray is elongated as a small filament. There is a strong pelvic axillary scale. There are 3-4 scale rows sheathing the dorsal and anal fin bases. There are 4-6 compressed teeth in front of each jaw followed by 3-5 rows of molar teeth. The chromosome number is 2n=48 (Klinkhardt et al., 1995). The gut is an elongate s-shape.

Meristic values for Iranian specimens are:- dorsal fin spines 12, soft rays 10, anal fin spines 3, soft rays 8, pectoral fin branched rays 13, scales mostly lost.

Sexual dimorphism

This species is a protandrous hermaphrodite, being male early in its life and then becoming female later. Catches will include males, females and hermaphrodites, e.g. in Abu-Hakima's (1984a) study in Kuwait, there were 326 males, 343 females and 41 hermaphrodites.

Colour

Overall colour is a silvery-grey or silvery-white with the back darker and the belly yellowish. Scales each have a dark, brownish to golden spot at the base which line up to form apparent stripes along the flank. There is a dark blotch at the upper corner of the gill opening, on both the body and gill cover. There is a dark band over the head between the eyes and the edge of the operculum is dark. Dorsal fin spines are white and the membranes are grey, with dark margins between the spine tips. The soft dorsal fin is dark grey with a light orange tinge. There is a small back spot at the pectoral fin base and the fin is mostly hyaline with a light orange tinge. The anal and pelvic fins are a light yellowish-brown. The caudal fin is dark grey on the upper lobe and yellow on the lower with a black margin. The peritoneum is silvery brown in preserved fish with widely scattered melanophores.

Size

Reaches over 50 cm total length.

Distribution

Found from the Persian Gulf to Japan and north Australia. Recorded from the Helleh River in Bushehr Province, Iran (J. Holčík, pers comm., 1995) and from the Bahmanshir River (Marammazi, 1995; Eskandary et al., 1999). Also from the Shatt al Arab in Iraq on the border with Khuzestan in Iran (Al-Hassan et al., 1989; Hussain et al., 1989).

Zoogeography

This marine species enters rivers in southern Iran and presumably freshwater stocks are maintained from this marine gene pool.

Habitat

The usual habitat is over sand and rock bottoms in the sea down to about 50 m, but young fish may enter estuaries and may penetrate considerable distances inland, although some fish remain at sea permanently. The frequency of penetration into Iranian rivers along the Persian Gulf coast is not known. Larger specimens are known to penetrate the Shatt al Arab in autumn, October to December, and this water body is an important nursery for this species, found there year round as young. Adults emigrate from January to March (Al-Hassan, 1990; Hussain et al., 1987). At a freshwater station on the Shatt al Basrah Canal with salinities up to 3.5‰, Al-Daham and Yousif (1990) found this species to be the second most dominant after Liza subviridis, comprising 7.1% by number and 10.9% by weight. Al-Daham et al. (1993) found young fish in the Shatt al Basrah mostly from April to October. Cage-cultured fish are reared at 14-31°C in Kuwait Bay (Abou-Seedo et al., 2003). Temperatures in Kuwait Bay during spawning are 14-18ºC during the winter months, a situation mirrored by Liza klunzingeri but not other species which begin to spawn in April (Abou-Seedo and Dadzie, 2004).

Age and growth

A fast growing and hardy fish. Life in Iranian fresh waters has not been studied. A study of fish in the Shatt al Basrah canal, a man-made estuary of southern Iraq, was based on mostly small and immature fish (49-181 mm standard length) caught  mostly in April-October. The length-weight relationship was W = 0.0511 L2.893, the dominant age group was 1+ fish, the maximum age was 3+ years, fish grew to 95, 155 and 215 mm total length in their first three years of life, and mortality (Z) was 2.23 (Al-Daham et al., 1993).

In an aquaculture experiment in Kuwait, fish more than doubled in weight over a 6 week period (Jafri et al., 1981). Males mature at a smaller size (12.3-14.2 cm) than females (24.3-26.2 cm) in cage-culture in Kuwait Bay (Abou-Seedo et al., 2003). However fish in the Shatt al Arab are usually less than 20 cm long and most are immature, in age groups 0 and 1. The length-weight relationship for both sexes was W = 0.0916 L2.6601. The lowest condition factors were found in April and May, possibly because fish were spent after spawning or were in lower condition after the winter (Hussain et al., 1987). In Kuwait, ages up to 14 years have been reported with ranges in von Bertalanffy growth parameters for the years 1981-1985 of L (cm) = 38.3-52.29, K = 0.169-0.298, and t0 -0.213- to -2.237. Total mortality values (Z) were 0.432-0.709, this range suggesting that, even for relatively large samples (92-314 fish), may be too small to provide reliable estimates of Z in species with a large overlap in age groups and where old fish are sampled only with difficulty. The length-weight relationship was total weight = 0.02874 x total length2.79198. Growth and mortality estimates based on all data were L = 40.48 cm, K = 0.258, t0 = -0.965, tmax = 14 years and Z = 0.600 (Samuel and Mathews, 1987). Morgan (1985) gave values of L = 43, K = 0.20 and Z = 0.97 but his study excluded fish over 30 cm and below 19 cm which accounts for the difference in mortality; and Samuel and Mathews (1987) had a value about 2.0 for Z when only fish 20-30 cm long were analysed.

Heydarnejad (2009) gave the length-weight relationship for an Iranian sample as W = 0.0451TL3.091.

Food

Freshwater food habits not known for Iran in detail but the one specimen examined contained plant fragments and scales of a cyprinid.

In a study of the recovering Hawr al Hammar, diet was 60% shrimps and 40% insects (Hussain et al., 2006). Feeds on echinoderms, worms, crustaceans, insects, bivalve molluscs and plants in the sea (Nasir, 2000; Al-Daham et al., 1993). Hussain et al. (1987) found crabs and bivalves to be the most important items by percentage in the Shatt al Arab Fish, shrimps and aquatic insects were also taken and there was significant seasonal variation, with shrimps and aquatic insects more important in December and spring. Hussain et al (1994) found bivalves in 91% of fish by number and weight in the Khawr az Zubayr. Al-Daham et al. (1993) found fish in the Shatt al Basrah fed on, in order of occurrence, crustaceans (decapods, amphipods, isopods, mysids, cladocerans and cyclopoids), fishes (Liza spp., Barbus (= Carasobarbus) luteus, Thryssa purava, eggs and scales), molluscs (Corbicula, Lymnaea, Tryonia and Sphaeriidae), algae (Oscillatoria, Syndera, Fragillaria and Cladophora), higher plants (Vallisneria, Ceratophyllum, seeds and roots) and aquatic insects (Corixidae, Hemiptera, Odonata and Coleoptera). Crustaceans were most important  during July and November, molluscs in May and fishes during August. Hosseini (1998) examined food in coastal waters of Bushehr, Delvar and Rostami in the northern Persian Gulf of Iran and found 36.4% to contain crabs, 34.0% other fish and 13.4% shrimps. Snails and sea urchins were also eaten.

Reproduction

Al-Hassan et al. (1993) have shown for fish from the Shatt al Arab near Basrah, Iraq that haematological parameters vary with reproductive phase and between sexes. Cage-reared fish in Kuwait Bay have a prolonged spawning season from February to April. Fecundity there is up to 3,837,000 eggs. Spawning in the Shatt al Arab estuary is reported for April (Hussain and Ahmed, 1995) and Al-Daham et al. (1993) record a spawning season for the northwestern Arabian (= Persian) Gulf as January to April with a peak in February and March. Abu-Hakima (1984a) found the spawning period in Kuwait waters to be January to March with fecundity up to 2,152,993 eggs. This species is a protandrous hermaphrodite with males dominating in smaller size groups (22.3-24.2 cm) while females dominate in larger groups (24.3-26.2 cm) (Abou-Seedo et al., 2003). Samuel and Mathews (1987) give a spawning date of 1 December for their Kuwait sample. The gonadosomatic index was highest in February-March in Hosseini's (1998) study in coastal waters of the northern Persian Gulf.

The average absolute fecundity in coastal waters near Bushehr in Iran was 1,842,700, sex ratio was 1:1, and the spawning peak was January-February (Hossini and Savari, 2004).

Parasites and predators

None recorded.

Economic importance

A good food fish seen in bazaars along the Persian Gulf coast and in the Shatt al Arab. It was selling at U.S.$3.5-5.5 per kg in Kuwait about 1980, with 213 tons landed in 1995 for a value of U.S.$1,769,407 (Abou-Seedo et al., 2003). Experiments there indicate that this species can be farmed (Jafri et al., 1981; Abou-Seedo et al., 2003) and this has been proposed for Iranian waters in the Persian Gulf (Regunathan and Kitto, 2005). Experimental culture has been tried at Qeshm Island where a million larvae were produced in March with 100,000 larvae 2.0-3.5 cm long surviving in May (www.shilat.com, downloaded 7 June 2007). It is caught by trawls, handlines, in hadra (fixed stake nets) and gargoor (fish pots) and in the sport fishery in the Arabian (= Persian) Gulf (Samuel and Mathews, 1987; Carpenter et al., 1997).

Conservation

This marine species is fished commercially in the sea and populations there may be under some threat as a consequence. The status of freshwater populations is unclear as they appear quite rare and are presumably derived from marine populations at intervals.

Further work

The frequency of occurrence, detailed distribution and biology of this species in Iranian fresh waters needs study.

Sources

Iranian material: uncatalogued, 1, 62.9 mm standard length, Bushehr, about15 km above mouth in Helleh River (ca. 29º13'N, ca. 50º43'E).

Comparative material: BM(NH) 1974.2.22:1859, 1, 76.4 mm standard length, Iraq, Basrah (30º30'N, 47º47'E); BM(NH)1974.2.22:1858, 1, 76.3 mm standard length, Iraq, Beree (no other locality data).

Mullidae

Genus Mullus
Linnaeus, 1758

Mullus barbatus
Linnaeus, 1758

Introduced to the Soviet Caspian Sea basin in 1931-1934 but not subsequently observed (Baltz, 1991). No Iranian record.

Cichlidae

Cichlids are found in fresh and brackish waters of Central and South America, Africa, Madagascar, the Levant, southern India, Sri Lanka and southern Iran. There are over 1300 species (with several hundred more to be described) (Nelson, 2006) but only 1 is found in Iran. Maximum length is about 80 cm.

Cichlids are distinguished by having only a single nostril on each side; practically all other fishes have two nostrils. The lateral line is in 2 parts, an anterior and higher portion ending under the soft dorsal fin and a lower, mid-flank, posterior part beginning below where the first part ends and continuing to the tail base, usually 7-25 spines in the dorsal fin followed by 5-30 soft rays, anal fin usually with 3 spines, but some species have 4-9 or 12-15 spines, followed by usually 4-15 soft rays, scales ctenoid or cycloid, extending onto the head, a specialised pharyngeal bone in the throat breaks up food by pressing it against a hard pad on the skull base, mouth dentition highly specialised in relation to diet with scraping (for algae on rocks), pointed (to seize fish), crushing (for hard-shelled molluscs), winkling (for removing snails from their shell or picking eyes) or reduced and embedded (for egg eating). Body form varies greatly between species and many are colourful and highly prized as aquarium fishes. They are often highly territorial, defending a breeding area against all invaders. They are most common in still waters where there are branches, rocks or plants. Some cichlids are important food fishes but they have attracted scientific attention for their elaborate breeding behaviour and evolutionary history. Certain cichlids, for example, are mouthbrooders, carrying eggs and fry in their mouths to protect them, while others spawn on the substrate, build nests or nourish young from a skin secretion. African lakes contain rich species flocks of cichlids which show various feeding behaviours. How these species arose and adapted to different ways of life have been important to scientists in understanding the mechanisms of evolution and adaptation.

Murray (2001) reviews the fossil record and the biogeography of the family and suggests an origin less than 65 MYA in the Early Tertiary in contrast to other studies that give an origin over 130 million years ago. Their salinity tolerance has enabled them to cross marine barriers.

Genus Iranocichla
Coad, 1982

The genus is monotypic so its description is subsumed under the Species Account below.

Iranocichla hormuzensis
Coad, 1982

Colour photographs of males (females in background) courtesy of Thomas Schulz

Common names

mahi-e karoo, siklid Irani, siklid-e Hormuz, cichlid-e Hormuz.

[Hormuz cichlid, Iranian cichlid]

Systematics

Trewavas (1983) relates this species to Danakilia Thys van den Audenaerde, 1968, a genus with two species (Stiassny et al., 2010) from northeast Africa, on the basis of the low gill raker count, lower pharyngeal bone and teeth morphology, and morphometric characters such as a deep preorbital depth, long snout, head length and the small eye. Trewavas (1983) places Iranocichla in Danakilia but also agrees with Coad (1982a) that Iranocichla (and Danakilia) may be related to species in the genus Tristramella Trewavas, 1942 of the Jordan River basin and that this requires further investigation. Schwarzer et al. (2009) found molecular evidence for the relationship of Iranocichla and Tristramella (DNA material of Danakilia was lacking). I retain Iranocichla as a distinct genus until these relationships are examined more closely as some of the characters used to relate Danakilia and Iranocichla may be common responses to temperature and salinity extremes. In addition, Trewavas (1983) suggests a possible relationship of Danakilia with Oreochromis alcalicus (Hilgendorf, 1905) of the African Rift Valley. This species too is found in waters of high temperature and mineral content. Klett and Meyer (2002) group this genus with Sarotherodon, Oreochromis and Tristramella on mitochondrial NADH dehydrogenase subunit 2 gene sequences.

The type locality is the "Mehran River at 27°04'N, 54°35'E, Hormozdgan Province" (Coad, 1982a). The holotype, 94.2 mm standard length, is a female with eggs in the mouth held in the Canadian Museum of Nature, Ottawa under CMNFI 1979-0408A (see figure above). Paratypes are CMNFI 1979-0408B, 15, 24.3-86.5 mm standard length, same locality as the holotype and CMNFI 1979-0139, 35, 29.6-95.2 mm standard length, stream in Rasul River drainage between Chahar Berkeh and Tang-e Dalan, ca. 27°25.5'N, 54°59'E, Fars-Hormozgan border. Paratypes were deposited in the British Museum (Natural History), London under BM(NH) 1981.1.12:1-2 (2 specimens), Muséum national d'Histoire naturelle, Paris under MNHN 1981-107, 108 (2), California Academy of Sciences, San Francisco under CAS 47324 (2), the Royal Ontario Museum, Toronto under ROM 36389 (1) and the University of British Columbia, Vancouver under BC 81-1 (1).

Key characters

This is the only cichlid species in Iran, easily recognised by the single nostril opening on each side of the head.

Morphology

This cichlid is uniquely characterised by a nearly circular dental field on the lower pharyngeal bone, the teeth there being of uniform size and not enlarged medially and by cheek, operculum, belly, isthmus and area between the pectoral and pelvic fin bases naked or poorly scaled. Other significant characters are the posteriorly rounded dorsal and anal fins, short pectoral fins not reaching the vent, cycloid scales with granular posterior circuli bearing rounded or irregular protuberances, inferior apophyses for support of the swimbladder centred around the fourth vertebra (figured in Coad (1982a)), mesethmoid not meeting the vomer, modal vertebral count 29, median length of lower pharyngeal bone 31.8-40.9% (mean 35%) length of head, and pharyngeal blade/median length toothed area 0.6-1.0, mean 0.8.

Scales are regularly arranged on the flanks except that in some large specimens the regular scale rows are interspersed with irregularly distributed smaller scales, particularly on the upper flank. Scales may be absent entirely from the head, sparse above the lateral line anteriorly and on the belly posterior to the pelvic fins, absent from the dorsal and anal fin bases, absent from between the pectoral and pelvic fin bases and on the belly and isthmus anterior to the pelvic fins. However, in other specimens the head may be scaled dorsally to above the eyes, with scales variably imbricate, there may be 2-3 rows containing 4-7 minimally or non-imbricate scales on the cheek which is never completely scaled. The dorsal border of the opercle may have two large scales next to each other and a single scale may be present over the centre of the subopercular bone. Scales may be present on the whole belly, isthmus and between the pectoral and pelvic fin bases, but they are minute, embedded, and non-imbricate. Their extent and number varies between individuals. Small to minute scales, numbering up to about 20, are present on the caudal fin base, extending distally onto the fin membranes for more than half the fin ray length in some specimens.

Flank scales below the mid-point of the spiny dorsal fin and beneath the upper lateral line are cycloid or very weakly ctenoid. The focus is central and there are 9-14, mean 12.4, radii on the anterior field based on 5 scales from 7 adult specimens 59.2-87.1 mm standard length. Posterior circuli are granular so the exposed scale surface has rows of rounded or irregular protuberances.

The gut is a tightly coiled spiral with its apex ventral. Gut length in 5 specimens (59.2-90.5 mm standard length) is 6.8-8.3, mean 7.6, times the standard length. Gill rakers are short and rounded, reaching the adjacent raker or a little further when appressed.

The pharyngeal apophysis is of the Tilapia type (Greenwood, 1978). The mesethmoid does not meet the vomer, the intervening space being cartilaginous. Pores at the openings of the cephalic lateral line canals on the preorbital and preoperculum are single not multiple. The inferior apophyses for support of the anterior end of the swimbladder involve vertebrae 2 to 5, the fourth vertebra being involved in 8 out of 10 fish examined.

Teeth in the jaws are often irregularly arranged so that 4 rows are found in some places in both jaws. In some individual fish where teeth are regularly arranged there are 3 rows in the upper jaw and 4 rows in the lower jaw. Number of rows decreases laterally to one at the rictus. The outer row teeth are bicuspid with the lateral cusp the smaller, while inner row teeth are tricuspid, with the central cusp the most prominent. The upper jaw has more teeth than the lower jaw.

The diploid chromosome number is 2n=44, comprising 25 submetacentic, 18 subtelocentric and 1 metacentric chromosomes with an arm number of 70. The chromosome count may indicate a relationship to the Levantine Tristramella (Esmaeili et al., 2006).

Scales in upper lateral line 17(1), 18(1), 19(2), 20(8), 21(9), 22(10), 23(7), 24(4), 25(2), 26(1) or 29(1); scales in lower lateral line 9(6), 10(17), 11(14) or 12(9); total scales in lateral series 28(1), 29(2), 30(4), 31(9), 32(13), 33(5), 34(4), 35(4), 36(3) or 40(1); scales around caudal peduncle 16(15), 17(13), 18(15), 19(2) or 20(1); precaudal vertebrae 14(2), 15(53) or 16(11); caudal vertebrae 13(26), 14(35) or 15(5); total vertebrae 28(19), 29(40) or 30(7).

Dorsal fin spines 14(6), 15(46) or 16(14); dorsal fin branched rays 9(2), 10(36) or 11(28); anal fin branched rays 6(7), 7(20), 8(38) or 9(1); pectoral fin branched rays 11(42) or 12(24); and total gill rakers 14(6), 15(9), 16(24), 17(19), 18(6) or 19(1).

Sexual dimorphism

Head length is greater in females while pelvic fin length is smaller in females compared to males. Interorbital width is greater in males. Dorsal and anal fins are larger in males when expressed in terms of longest ray length in head length (Coad, 1982a). Colour differs as described below.

Colour

Live specimens are brightly coloured in spawning condition (based on aquarium photographs in Schulz (2004)). The male is brick-red on the lower sides and underside of the head with black on the dorsal head surface. The underside of the head may also be black. The belly anterior to the pelvics is black. The chin is white. The sides off the head have a few, scattered white spots but the body, dorsal and caudal fins are densely covered with white spots and blotches. Those on the dorsal fin are arranged in oblique rows and those on the caudal fin in bars. The anal fin has white spots also but these are not present distally. The pectoral fin has darkened rays but lacks spots. The pelvic fin has white spots proximally but less than the anal fin but is overall a dark black. Other reports and photographs (Svardal (2006) and Svardal and Svardal (2006)) show dominant spawning males to be black with brilliant turquoises blotches on the body but especially so on the fins. The female has an overall silvery colour with up to 9 faint to moderate flank bars. Fins are yellowish. The dorsal fin has a black tilapia-mark on the posterior dorsal fin.

Overall body colour outside the spawning season is a light lime green, with an iridescent tinge to the posterior edge of the operculum and on the back. The dorsal fin has light, lime-green, oblique bars, the last one or two black-edged and spot-like. The peritoneum is black.

Preserved specimens have the following pigmentation. Young fish have a distinct tilapia-mark, a spot on the rays of the soft dorsal fin typical of these cichlid fishes. The spot is black and is surrounded by a hyaline ring. Occasionally a second spot is found posterior to the first spot. The principal spot is often retained in adult fish. Young also have 7-11 bars along the flank which are also retained by adults but are then less distinct. In adults the dorsal fin rays and membranes are covered with melanophores interspersed with hyaline spots and irregular blotches. Wavy, oblique bars are found posteriorly on the soft dorsal fin in some specimens. The caudal fin has a series of about 7 narrow bars in some male specimens while females are uniformly grey. The anal fin is narrowly barred with up to 6 vertical to oblique bars in some specimens, in others uniformly pigmented grey proximally fading to hyaline distally. Pectoral and pelvic fins are not barred and are lightly pigmented, the pelvics being the darker. The head and body, including the belly, are more heavily pigmented to give an overall brown colour, lightest on the belly anterior to the pelvic fins in females. Scales are not pigmented on their free margins, which are pale.

Some specimens may be quite dark, particularly the back and fins and strikingly the lips.

Size

Attains 11.09 cm standard length or 12.95 cm total length (Esmaeili and Ebrahimi, 2006). Lamboj et al. (2006) give 13 cm, presumably total length.

Distribution

  

The cichlid is restricted to rivers draining to the Straits of Hormuz in southern Iran (Coad, 1982a; Abdoli, 2000). Svardal and Svardal (2006) also map this species at 27.770°N, 54.999°E, slightly to the north of samples mapped here. The distribution mapped by Stiassny in Keenleyside (1991) following Berra (1981) is too far north. The map in Berra (2001) is more accurate. Abdoli (2000) records this species from the lower Minab basin, lower Hasan Langi, middle to lower Kul, Gowdar and middle to lower Mehran rivers.

Specimens kindly sent to me by H. R. Esmaeili in 1997 are from the Dozdan River at 27°26'N, 57°10'E, an eastwards extension into the Minab River basin. The cichlid was not collected there in the 1970s. The new record may simply be filling in a collecting gap, a natural range extension or possibly the result of an introduction.

Bleher (2011) found only 2 of the 22 sites recorded by Coad (1982) to still have water, although I have observed river stretches drying up and re-connecting and not always the same stretches.

Zoogeography

Trewavas (1983) suggests that the ancestor of this cichlid was distributed across the Arabian Peninsula in the Late Pliocene/early Pleistocene when this area was more humid. Desiccation in the Pleistocene and Recent Periods then led to the extinction of the ancestor. A Miocene-Oligocene fossil "Tilapia" was reported from Jisan in southwest Saudi Arabia where cichlids are not now native (Brown, 1970) and would help support a southern distribution of cichlids as the origin of Iranocichla. However Trewavas (1983) reports that this fossil cannot be identified as a cichlid. Micklich and Roscher (1990) and Lippitsch and Micklich (1998) also report three species of what are presumably cichlids from southwest Saudi Arabia in the Baid Formation of Oligocene age at Ad Darb, Tihamat Asir. They belong to the basal grade of cichlids and to two different clades within the African assemblage. Whybrow and Clements (1999) record unidentified Cichlidae from the Early Oligocene from the coastal trip of Dhofar, Sultanate of Oman with a date of 33 MYA. Murray (2001) reviews these and other cichlid fossil material and the identity of Omani material as cichlids appears questionable. Southwest Saudi Arabian material is more clearly cichlid but does not point to a continuous distribution eastwards across the Arabian Peninsula. However, Bănărescu (1992b) considers that a common ancestor to Iranocichla and the Levantine Tristramella evolved in the Arabian Peninsula from African forebears in the Miocene, the latter lineage extending its range northwards to the Levant and the former eastwards to Oman and southern Iran, the Straits of Hormuz not then being in existence. Murray (2001) gives an earliest date for colonisation of Iranocichla ancestors to be the Middle Miocene when southern Iran rose above sea level. She does not consider a coastwise dispersal through brackish waters of Arabia to be a possible route as cichlids are not found there today but indicates a route through the Tethys Sea/Indian Ocean could be possible.

Coad (1982a) suggests another hypothesis. Iranocichla could be a relict of a once wider distribution across the Tigris-Euphrates basin in a northern arc rather than directly across the Arabian Peninsula. The absence of cichlids from southern Arabia today warrant this alternative hypothesis. Warm streams have probably been continually present in southern Arabia and support a limited fish fauna today. There is no apparent reason why cichlids should have become extinct there. Murray (2001) points out that Iranocichla lives at 40-400 m above sea level and is limited by mountains north of the present distribution, and so it must have arrived before the mountains attained their current height, to support Coad's hypothesis.

The headwaters of the Tigris-Euphrates basin are narrowly separated from the Levant Rift Valley today and at times in the past may have had direct exchanges of faunas (Kosswig, 1965; 1973; Krupp, 1987). The modern absence of cichlids from the Tigris-Euphrates basin may be explained by low temperatures. The effects of low temperature on Iranocichla have not been investigated but fingerlings of Tilapia aurea, which occurs naturally in the southern Levant Rift Valley, begin to die at 11°C and cease all motion at temperatures below 10°C (Chervinski and Lahav, 1976). Most of Syria, northern Iraq and the northern Arabian Peninsula have temperatures below 10°C in winter (Beaumont et al., 1976). Spot temperatures from southern Iran at the head of the Persian Gulf are about 12-13°C in January compared to 18-20°C around the Straits of Hormuz where Iranocichla occurs. Additionally the death rate of Tilapia aurea in fresh water is twice that in salt water at low temperatures: it may be pertinent that Iranocichla is found mainly in saline streams. This hypothesis can only be confirmed by fossil discoveries.

Habitat

The streams in which this species lives are subject to desiccation with continuous flow breaking up into isolated pools. The survival of cichlid populations in these pools varies between years and some pools may be fishless in one year and populated in another.

The area around the Straits of Hormuz is rich in salt domes and consequently most surface streams are saline, up to 80 mS. Cichlids are found in these streams but also in the Sar Khun oasis which is fresh with a conductivity of 1.6 mS. Apparently they can be transported at 10 mS as this is less stressful. Stream waters are cloudy to clear and colourless. Water temperatures in winter (November to March) range from 15 to 33°C and would be considerably higher in summer when air temperatures reach 45°C with no riparian shade and low water levels. Lamboj et al. (2006) and Svardal (2006) give water temperatures of 33-40°C and conductivity of 45-75 mS.

Streams are 1 to 50 m wide and consist of alternating riffles and pools with occasional backwaters. The bottom is pebbles, sand or mud. Aquatic vegetation is restricted to encrusting algae.

Kiabi and Abdoli (2000) found this species to be, with Cyprinion watsoni, the commonest in Hormozgan Province.

Age and growth

Unknown. Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 379 fish measuring 2.74-11.09 cm standard length. The a-value was 0.0349 and the b-value 3.047 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases).

Food

Gut contents of 5 specimens (41.2-90.5 mm standard length) included only algae and diatoms suggesting food is scraped from rocks and from bottom deposits. This is consistent with an elongate gut and black peritoneum. Aquarium specimens eat algal tabs but also appreciate insects and fish remains.

Reproduction

This species is a mouth brooder. A breeding female and a male were caught in a backwater on March 18 of the Mehran River (the type series). This backwater was 1-5 m wide, maximum depth was 40 cm over a mud bottom, the water was cloudy and highly saline (40mS) and temperature ranged from 26°C at the mouth of the backwater to 33°C at its head. Eggs in fish taken in November and January are small so the breeding season is deduced to be around March. Five eggs ranged in length from 3.2 to 3.8 mm, mean 3.6 mm and in width from 2.4 to 2.7, mean 2.5 mm. Total number of eggs from 2 females, 59.0-59.2 mm standard length was 36 and 38 respectively. Eggs are yellow-orange in preserved fish.

A female 116.9 mm standard length from the Mehran River had 153 larvae in her mouth, ranging in length from 9.6 to 10.9 mm (H. R. Esmaeili, pers. comm., 6 October 2005; Esmaeili et al., 2009). Esmaeili et al. (2009) found a sex ratio biased towards males in May and June, presumably because males were defending nests and easily caught, or possibly differential survival of the sexes. They suggest that the breeding season begins in March and lasts until the end of June with a peak in May. Eggs attained 3.76 mm and fecundity reached 151 eggs with a relative fecundity of 5.4 eggs per gram body weight. Esmaeili et al. (2010) detail gonad morphology and histology and confirmed peak spawning in May.

Schulz (2004) observed fish in the field and found each male occupying a territory defending a nest about 1 m from each neighbouring nest. The nests were made on light grey, fine sand and consisted of a pit approximately 15 cm in diameter. The pit was black because of anoxic conditions below the sand surface. The actual nest was about the same as the body length of the fish (8-10 cm) and lay at the centre of the pit. The pit was surrounded by a rim about 1.5 cm high with an internally indented margin. Simpler pits are built where building materials are unavailable. Females were present in schools in deeper water in the river centre. Individual females swam purposefully to the nest defended by the male. The male directed the female to the nest centre with folded up fins while the female spread her fins and showed radiating colour changes. Spawning occurred immediately and neighbouring males intervened continuously at a speed that did not allow full analysis of the movements. A defending male would chase away an intruding male allowing another male into the unprotected nest to mate with the female. A clutch of eggs was always inseminated by a whole group of males.

     

Spawning site photographs courtesy of Thomas Schulz

Parasites and predators

Unknown although piscivorous birds have been observed along the streams where the cichlid is found.

Economic importance

Saadati (1977) suggests that this salt-tolerant species could be a valuable resource if introduced into the saline and fishless waters of internal basins. However this is not advisable since the native fauna, evolved in a fishless environment, could be devastated before it has even been documented. Esmaeili et al. (2009) note that it is eaten by local people when available in large numbers in spring. It is now an aquarium fish in Germany (Schulz, 2002; 2004a; 2004b; Oliver Lucanus, pers. comm., 23 January 2004; Lamboj et al., 2006; Svardal, 2006; Svardal and Svardal, 2006). Articles in aquarium magazines give photographs in spawning condition, including mouth-brooding, and details for their maintenance, including water with a conductivity of 50-70mS/cm NaCl or sea salt mixture, tank water changed once a week, vegetarian food tabs (containing the blue-green alga Spirulina), and a temperature of 20-35°C, optimally 27°C. It has been noted that males, in continually defending a nest and courting, "wear out" earlier than females (Thomas Schulz, in litt., 12 October 2006).

Conservation

Axelrod (1993) states that "as pollution chokes the waters of southern Iran, we can expect this fish to disappear very quickly" and "Because of the restricted range of this fish and the continual warfare and oil pollution in the area, Iranocichla hormuzensis may well be on its way to extinction - if it is not gone already". Bailey (2006) apparently repeats this. However its habitat is mostly saline streams which cannot readily be used for agriculture or industry. The surrounding area is not industrialised, nor likely to be, and was never a war zone so pollution is not a problem for this species.

Flash floods are probably a significant problem as water drains rapidly off vegetation barren land. The scouring action may well displace or strand cichlids. Mouth brooding offers protection against floods and against associated fishes.

Svardal (2006) and Svardal and Svardal (2006) give details of capture, transport and aquarium care of this species.

Further work

Further investigations into the biology of this species are needed.

Sources

Rabbaniha (1993a, 1993b, 1994) gives Farsi accounts of this species and cichlids in general. The account is based principally on Coad (1982a).

Type material: See above, CMNFI 1979-0408A, CMNFI 1979-0408B, CMNFI 1979-0139, BM(NH) 1981.1.12:1-2,  MNHN 1981-107, 108, CAS 47324, ROM 36389 and BC 81-1.

Iranian material: CMNFI 1979-0138, 17, 25.6-97.3 mm standard length, Fars-Hormozgan border, stream in Rasul River drainage (ca. 27º32'N, ca. 54º58'30"E); CMNFI 1979-0140, 48, 24.2-59.4 mm standard length, Hormozgan, stream east of Kichal, Kul River drainage (27º14'N, 55º46'30"E); CMNFI 1979-0141, 5, 15.0-21.0 mm standard length, Hormozgan, Kul River (27º17'30"N, 56º03'30"E); CMNFI 1979-0142, 65, 21.3-82.8 mm standard length, Hormozgan, Baghu River (27º17'N, 56º28'E); CMNFI 1979-0143, 3, 25.0-26.7 mm standard length, Hormozgan, marsh in Hasan Langi River drainage (27º21'N, 56º50'30"E); CMNFI 1979-0148, 5, 33.2-92.7 mm standard length, Hormozgan, Sarzeh River (27º30'30"N, 56º15'30"E); CMNFI 1979-0181, Hormozgan, Kul River (27º17'30"N, 56º03'30"E); CMNFI 1979-0184, 12, 31.4-50.4 mm standard length, Hormozgan, effluent of Rasul River (27º11'N, 55º42'E); CMNFI 1979-0185, 7, 26.1-59.8 mm standard length, Hormozgan, stream in Rasul River drainage (27º06'N, 55º45'E); CMNFI 1979-0187, 2, 63.9-64.6 mm standard length, Hormozgan, stream at Sar Khun (27º23'30"N, 56º26'E); CMNFI 1979-0406, 5, 33.4-59.2 mm standard length, Hormozgan, stream north of Bandar-e Charak turnoff (26º48'N, 54º18'E);CMNFI 2007-0053, 17, 28.0-59.9 mm standard length, Hormozgan, Sarzeh River (ca. 27º36'N, 56º15'E); CMNFI 2007-0057, 14, 30.2-85.5 mm standard length, Hormozgan, Mehran River below Bastak (ca. 27º05'N, ca. 54º05'E); CMNFI 2007-0058, 2, 64.8-83.0 mm standard length, Fars, headwaters of Gowdar River (ca. 27º24'N, ca. 54º16'E)

Genus Oreochromis
Günther, 1889

Oreochromis niloticus
(Linnaeus, 1758)

Introduced to the Tigris River basin in Iraq but did not apparently survive winterkill (Herzog, 1969). Mutlak and Al-Faisal ( 2009), however, record O. aureus (possibly O. niloticus) from Basrah in southern Iraq and this species could easily become established in Iran. No Iranian record confirmed as yet. Red tilapias (Oreochromis sp.) have been studied in aquaponic systems in Iran so there is a potential for an exotic release (Rafiee and Saad, 2005). Fingerlings from Indonesia have been reared using saline waters at Bafgh, Yazd Province in 3 ton fibreglass tanks. Larvae were successfully grown to 2.0 kg at 28±1ºC (Iranian Fisheries Research Organization Newsletter, 57:2, 2009).

Genus Tilapia
Smith, 1840

Tilapia zilli
(Gervais, 1848)

Introduced to the Tigris River basin in Iraq but did not apparently survive (Job, 1967). Redbelly tilapias are established in the Syrian Euphrates (R. Beck, pers. comm., 2000) and a recent report by Beshar Abd Al-Hussain Al-Saadi (in litt., 10 October 2006) of a cichlid at Al Musayyib on the Euphrates River in Iraq may well be this species. Mutlak and Al-Faisal ( 2009), record this species from Basrah in southern Iraq and these could spread to Iranian waters. No Iranian record as yet. The Farsi name is تيلاپيا (= tilapia).

Gobiidae

The gobies are a world-wide family found mostly in warmer marine waters although some species enter fresh water and others live there permanently (see also Marine List in Checklists in the Introduction). The number of species is high and this may be the most speciose fish family in the world with about 210 genera and an estimated 2000 species, perhaps more. A diversity of gobies occurs in the Caspian Sea basin. Not all Caspian gobies have valid Iranian records but most will probably be found there. Several gobies penetrate southern waters of Iran from the Persian Gulf and Sea of Oman and are described here. Others will probably be discovered when more detailed surveys are made.

Gobies are easily distinguished by their pelvic fins being united as an adhesive or sucking disk or cup. Body form and coloration are diverse. The pattern of head canals, canal pores and neuromasts is distinctive and used in identifying and relating species (except in "Neogobius" (Pinchuk, 1991)). However the neuromasts may be sunken in narrow furrows or pits and completely covered by epithelium so they do not preserve well and this can lead to confusion in identifications (Zambriborshch, 1968). There is usually a short spiny dorsal fin (2-8 flexible spines) separated from, but close to, a soft dorsal fin. The soft dorsal fin and anal fin are longer than the caudal peduncle. Scales may be cycloid, ctenoid or rarely absent. No obvious lateral line. There are 5 branchiostegal rays. Gill membranes are connected to the isthmus and gill openings are moderate to wide, or very restricted in the mudskippers. The head is usually blunt and the mouth is usually large. Teeth are usually small and conical in one to several rows in both jaws. Miller in Miller (2003) gives a suite of osteological characters defining the family.

Most gobies are quite small (5-10 cm) and they are often very abundant. Maximum size is about 50 cm. Some of the world's smallest vertebrates are gobies from the Indian Ocean, mature at 8 mm. Others, however, are large and form part of fisheries in both the Caspian Sea and the Indian Ocean. They are not significant food fishes in Iran. Gobies tend to rest on the bottom and move in sudden, characteristic dashes. The male goby guards a nest. Food is crustaceans, worms, molluscs and small fishes. Many gobies are important in the aquarium trade since they are beautifully coloured, small and tough.

They are known generally as gav mahi (= cow fish) or sag mahi (= dog fish) or contain the word gel (= mud) in Iran. A general review in Farsi of the Caspian gobies is given by Aslaanparviz (1991).

The males of some Caspian species become black during the spawning season, their fins elongate, head shape alters and some even become naked. Loss of tubercles in adult male gobies of the genus Benthophilus makes it possible to identify only juveniles and females. The males build nests and guard the eggs. Life span of certain Caspian species is said to be as short as one year, e.g. some species of Benthophilus, Knipowitschia, and Proterorhinus. Neogobius pallasi, N. melanostomus and Ponticola gorlap consume the invasive ctenophore, Mnemiopsis leidyi, as much as 10-15% of total biomass in some areas (Mamedov, 2006).

The Black and Caspian Sea basins contain an endemic Sarmatian fauna of gobies. There are two main clades, the gobiine-benthophilines (or transverse gobiids) and the pomatoschistines (or sand gobies), that have probably been distinct for at least 40 million years. Miller (2001) and Miller in Miller (2003) reviews the evolutionary history of these two clades and their anatomical differences based on head papillae and osteology. The transverse gobiids include Mesogobius, Neogobius, Proterorhinus, Chasar, Anatirostrum, Benthophiloides, Benthophilus and Caspiosoma while the sand gobies include Knipowitschia and Hyrcanogobius. The Sarmatian fauna was separated from the Atlantic-Mediterranean fauna with the isolation of the Paratethys during the late Miocene Messinian salinity crisis as the Mediterranean dried. Partial flooding of the Mediterranean from the Paratethys in the early Pliocene allowed Sarmatian gobies to spread westwards. Within the Ponto-Caspian basin, evolution of species flocks was favoured by basin sub-divisions and rejoinings. The benthophilines may be a monophyletic group from these events.

Ahnelt and Duchkowitsch (2004) give information on the neogobiine stock. About 12-13 million years ago in the Middle Miocene, the Ponto-Caspian endemic and ancestral neogobiine stock may have differentiated from an Atlantic-Mediterranean gobiine stock. At this time the Paratethys was a sea with reduced salinity and a high level of endemism. The Neogobius-Proterorhinus stock developed independently from the recent Gobius stock that invaded the Mediterranean basin after that sea was restored about 5 million years ago in the Late Miocene.

Neilson and Stepien (2009a) using mitochondrial and nuclear genes detail the evolution and biogeography of the subfamily Benthophilinae restricting Neogbius to caspius, melanostomus and pallasi with other known Iranian species formerly in Neogobius placed in Ponticola. The Benthophilinae includes Babka, Mesogobius, Neogobius, Ponticola and Proterorhinus as well as the tadpole gobies (Anatirsotrum, Benthophiloides, Benthophilus and Caspiosoma). The origin of the genera Babka, Benthophilus, Caspiosoma, Mesogobius, Neogobius, Ponticola and Proterorhinus ranges from 4.29 to 6.25 MYA and the paper gives divergence times for major lineages in relation to geological events in the Ponto-Caspian. These events include connections with, and isolation from, the World Ocean and salinity changes in a range of 1-30 p.p.t. over the last 5 million years. Most genera diversified about 5 MYA when the Black and Caspian seas separated. More recent separation events occurred during the Pleistocene glaciations Proterorhinus diverged from Mesogobius about 6.18 MYA, the Ponticola clade diverged about 4.51-4.86 MYA, and freshwater Proterorhinus emerged 1.18 MYA.

The principal recent works on the systematics of Caspian gobies are by V. I. Pinchuk, D. B. Ragimov, Ye. D. Vasil'yeva, H. Ahnelt and P. J. Miller. Earlier works are by B. S. Iljin (also spelled Il'in or Ilyin). Later molecular studies are cited above.

Other gobies in Iran are the familiar tropical mudskippers which can move quickly over land, using the muscular-based paired fins to row across mud, and some can even clasp and climb mangroves. They can live out of water because the gill openings are small to prevent desiccation of the gills, oxygen can be taken into the chamber and absorbed through the gills and chamber wall, and they can also absorb oxygen through their skin. They often rest with the tail immersed in water for this purpose or roll around in shallow water to moisten themselves. They may live entirely in water, or will come onto land even when there is enough oxygen in the water. Their eyes are high on the head, protruding and able to revolve independently, and have a movable lower lid. The eyes are retracted periodically into small cups below the head to moisten them. Such eyes are very effective as a means to watch for potential enemies on land but their vision under water is blurred. Mudskippers have elaborate reproductive behaviour which involves tail standing, flip-flops, and fin displays. They are very territorial and defend their territory against other mudskippers and crabs. They can deliver a skin-breaking bite to humans even though they are only about 15 cm long!

Genus Anatirostrum
Iljin, 1930

This genus comprises only a single species and so its characters are those of the species. The snout is very distinctive and details of neuromasts are not given here as they are not needed in identification, although of importance in relating the genus. The genus is closely related to the tadpole goby clade comprising Benthophiloides-Caspiosoma-Benthophilus and details are give in Miller in Miller (2004). This author also gives an alternative terminology for the arrangement of neuromasts than that of Ahnelt et al. (2000).

Anatirostrum profundorum
(Berg, 1927)

Dorsal view

Common names

gavmahi nuk-ordaki (= duckbill goby), gavmahi-ye poozehderaz.

[duckbill tadpole goby].

Systematics

This species was originally described in the genus Benthophilus Eichwald, 1831 by Berg (1927) but later Iljin (1930) erected a new genus because of its unusual and distinctive morphology. The type locality is the Caspian Sea at 37°58'N, 52°22'E at a depth of 294 m (but see below).

There are 14 fish under ZISP 23134 recognised as syntypes in Ahnelt et al. (2000) although Berg (1927) mentions 15 fish in his description. Ragimov (1985) states that Berg described this species from a single young specimen and also visually observed 15 others for a total of 16 in the type series.

Key characters

The duckbill tadpole goby is characterised by the elongate and flattened head which is similar to a duck's bill. Unlike gobies of the genus Benthophilus the body has minute platelets and granules, there are 8 rather than 6, vertical, suborbital series of pit organs, no postorbital occipital series, and no chin barbel or cheek flaps behind the jaw angle (Ahnelt et al., 2000).

Morphology

First dorsal fin with 3-4 spines, usually 4, second dorsal fin with 1 spine followed by 8-11, usually 10, soft rays. Anal fin with 1 spine followed by 8-11 soft rays. Pectoral fin rays 14-16. Gill rakers on the posterior part of the arch are very short and anteriorly are minute. Pit organs on the side of the head are papilliform and clearly visible with the naked eye. Further details of anatomy are given by Ahnelt et al. (2000).

Iranian specimens had the following meristics:- first dorsal fin with 4(4) spines; second dorsal fin with 1(4) spine followed by 10(4) soft rays; pectoral fin rays 14(1), 15(2) or 16(1); anal fin with 1(4) spine followed by 11(4) soft rays; and total vertebrae 29(4).

Sexual dimorphism

Females may retain dermal granules when mature.

Colour

Overall, colour is a light grey or pale fawn fading to a whitish grey on the belly. Various speckles and melanophores are found on the back and upper flank. The dorsal, caudal and pectoral fins have dark grey speckles. The head sides from the snout to the cheek are dark with transversal suborbital papillae series whitish giving the impression of narrow light stripes below the eye and on the cheek. The peritoneum is black or densely covered in fine speckles.

Size

Reaches 11.2 cm, or 13 cm total length (Jolodar and Abdoli, 2004). Females may be larger than males (mean total length 84 mm versus 77 mm).

Distribution

Found in the southern Caspian Sea including Iranian waters (Berg, 1927; Ahnelt et al., 2000; Abdoli and Naderi, 2009).

Zoogeography

Known only from the Caspian Sea and one of the endemic Sarmatian fauna (see Family Account).

Habitat

Found to a depth of 294 m on white silt bottoms according to Berg (1927) but the data in ZISP states 244 sazhems (= 446.5 m). Recent Iranian material is from 45-80 m, at 9.7-16.4°C at 50 m (Ahnelt et al., 2000) and Jolodar and Abdoli (2004) state it lives mainly at 50-100 m depths in the south Caspian Sea.

Age and growth

Unknown.

Food

Unknown but the duck bill may be an adaptation for feeding on silt bottoms (Ragimov, 1986).

Reproduction

Apparently mature eggs reach 1.9 mm in diameter in the Iranian specimen.

Parasites and predators

Unknown.

Economic importance

This species is too rare to be of any economic importance.

Conservation

Conservation requirements are unknown. Kiabi et al. (1999) consider this species to be data deficient in the south Caspian Sea basin according to IUCN criteria.

Further work

More specimens need to be caught to assess its distribution, numbers, variation and biology.

Sources

Type material: See above (ZISP 23134).

Iranian material: CMNFI 1999-0023, 4, 76.1-79.1 mm standard length, Gilan, Caspian Sea off Astara (38º00'N, 49º30'E to 38º20'N, 50º00'E).

Genus Babka
Iljin, 1927

Babka gymnotrachelus
(Kessler, 1857)

Caspian Sea basin but no Iranian record although Kottelat and Freyhof (2007) map it from the Iranian shore. Gobius macropus De Filippi, 1863 described from Lake Palestrom near Poti, Georgia is a synonym.

Babka macrophthalma
(Kessler, 1877)
 

Reported from the south Caspian Sea by Naseka and Bogutskaya but no confirmed specimen from Iran.

Genus Benthophiloides
Beling and Iljin, 1927

Benthophiloides brauneri
Beling and Iljin, 1927

Originally described from the lower Dnieper River between Kherson and Kakhovka and the south Bug River between Novaya Odessa and Nikolayev, Ukraine and also recorded from the Caspian Sea basin but no Iranian record. Ragimov (1998c) and Miller (2004) give recent descriptions. Pinchuk and Miller in Miller (2004) question the validity of the Caspian Sea record for this species.

Benthophiloides turcomanus
(Iljin, 1941)

Caspian Sea basin, described from off Chikishlar at 37°45.5'N, 53°47'E and southwest of Ulsky Bank at 38°05'N, 52°34'E, Turkmenistan, but no Iranian record. Known only from the two type specimens, now lost (Reshetnikov et al., 1997). Miller in Miler (2004) places this species in Benthophiloides; formerly it was in the genus Asra Iljin, 1941. Miller in Miler (2004) gives a general map that encompasses the southeastern Caspian Sea including Iranian waters but the only records are not in Iran.

Genus Benthophilus
Eichwald, 1831

Recheck Benthophilus illustrations in this account for source (correct species)

The tadpole gobies are found in the Black and Caspian seas where there are about 20 species, 16 endemic to the Caspian (Pinchuk and Miller in Miller, 2004; Boldyrev and Bogutskaya, 2007). The general Farsi name for fishes in this genus is گاو ماهي (gav mahi) or سگ ماهي (sag mahi), not repeated under each species description.

Members of this genus are characterised by the broad and flattened head, dorsal muscles not extending to the eyes, no sensory canals or pores, first dorsal fin with 2-4 rays, well-separated form the second dorsal fin, the caudal fin has only 2 rows of papillae, head and body scaleless but with spinulose bony granules, scutes and platelets except in adult males which are naked (and cannot therefore be readily identified), anterior nostrils developed as small tubes overlying the upper lip, no swimbladder, a longitudinal dermal fold usually present on each side behind the mouth corner, and presence of a chin barbel. There is no pelagic larval stage and eggs are large and oligoplasmatic. Ahnelt (2003) discuss the unique specialisations in the postcranial skeleton of benthophiline gobies (which also includes Anatirostrum and Benthophiloides) including reductions in vertebral numbers and arrangement of pterygiophores. Pinchuk and Miller in Miller (2004) review other characters that show relationships of this specialised group within a larger tadpole-goby clade, the broad and flattened head being the most obvious.

Pinchuk and Miller in Miller (2004) give details of the head neuromast organs that characterise this genus, namely 6 transverse infraorbital rows, 1-4 before and 5s and 6s well above the level of the hyomandibular row b; row a is absent; row 5s is distant from row 4 which does not extend above the level of row b; rows 5i and 6i are arranged successively but row 5i is mostly below the level of row 6i separating the latter from the posterior end of row d; dorsal supraorbital rows o very short and well-separated in the dorsal midline; and supraorbital series p present in interorbit as 3 pairs of short transverse rows.

These fishes are found in brackish waters with a salinity up to about 20‰, in deeper waters, estuaries and coastal waters. Life span of these tadpole gobies is only a year, some individuals maturing at 6-7 months, called ephemery. Fish die after spawning, females earlier than males by 3-4 weeks (Boldyrev and Bogutskaya, 2004; 2007).

Pinchuk and Miller in Miller (2004) and Boldyrev and Bogutskaya (2004; 2007) give descriptions and keys for all Caspian Sea species. The 5 species cited here are the ones with records from Iran although it is likely other species will eventually be found along the southern Caspian Sea coast. Boldyrev and Bogutskaya (2007) gives keys and descriptions to all the species.

Benthophilus abdurahmanovi
Ragimov, 1978

Endemic to the Caspian Sea, found in the North Caspian and reported from the south Caspian Sea by Naseka and Bogutskaya (2009). No Iranian record.

Benthophilus baeri
Kessler, 1877

Common names

gavmahi-ye Baer, gavmahi-ye tokmehsar.

[Ber comcaxulu in Azerbaijan; Baer pugolovka or Baer tadpole goby in Russian].

Systematics

Benthophilus Baeri was described from the Caspian Sea at Mangyshlak Peninsula, Kazakhstan and in the South Caspian.

A specimen listed as a syntype is in the Natural History Museum, London under BM(NH) 1897.7.5:16 (32.5 mm standard length) from St. Petersburg University (sic, ZISP?) with others in the Zoological Institute, St. Petersburg (ZISP 2239, lectotype and ZISP 53665 (ex 2239) (4)) and SPU 395 (463/410) (1) (Eschmeyer et al., 1996; Pinchuk et al., 2004).

Lectotype: ZISP 2239. Paralectotypes: (11) BMNH 1897.7.5.16 [ex St. Petersberg Univ.] (1), SPU 395 [463/410] (1), ZIN 53665 [ex 2239] (4).

Key characters

This species has very large and high, conical tubercles arranged in rows, not all tubercles are spinous, dorsal row tubercles usually 11-17 (lower than other Benthophilus), ventral row tubercles 9-14 (lower than other Benthophilus), granules are mostly absent between body tubercles rows, interorbit with a shallow median groove, a chin barbel is present, there are 1-2 dermal barbels behind jaw angle, the first dorsal fin has 1-2 spines, and dark bands are absent.

Morphology

First dorsal fin with 0-2 spines, usually 1 (Ragimov (1985) and Boldyrev and Bogutskaya (2007) but 2 according to Pinchuk et al. in Miller (2004)), second dorsal fin with 1 spine followed by 6-9, usually 7, soft rays. Anal fin with 1 spine followed by 5-9, usually 6-7, soft rays. Dorsal row tubercles 11-17, usually 13-15, lateral row 1-12, usually 7-9 (but sometimes completely absent), and abdominal row 9-14, usually 11-12. Total vertebrae 24-27. Tubercles on the head and flanks are large and can bear 1-4 sharp spines but only some tubercles retain spines. The dorsal tubercle row continues onto the head ending near the eyes while the lateral row is short, extending from a level between the dorsal fins and ending at the rear of the second dorsal fin. There may be a few small granules scattered between the major row tubercles and on the head. There is a long chin barbel, about equal to eye diameter. There is one neuromast in row 7 behind the eye.

A single Iranian specimen had ?

Sexual dimorphism

The second dorsal and anal fins of large males are enlarged.

Colour

The back, upper flank and upper head are pale grey to pale brown with dark grey speckles, and often minute dark dots form dark areas. Some bands may be weakly expressed on the back and flank, more obvious in immature specimens. The lower body and belly are whitish. The pectoral, second dorsal and caudal fins have dark grey speckles and bars while the other fins are pale.

Size

Reaches 8.6 cm for males and 6.9 cm for females.

Distribution

Found in the central and southern Caspian Sea, rarely in the northwest Caspian. Reported from Gorgan Bay, the southeast Caspian Sea, southwest Caspian Sea and south-central Caspian Sea in Iran (Kiabi et al., 1999; map in Reshetnikov, 2002; Pinchuk et al. in Miller, 2004; Abdoli and Naderi, 2009).

Zoogeography

A Caspian Sea endemic, part of a unique fauna poorly studied and collected in Iranian waters, as are all its relatives below.

Habitat

Found in depths of 5 to 81 m in littoral areas and also known to enter rivers in Russia (Reshetnikov et al., 1997). Found at 0.5-30.0 m in warm seasons and migrates to deeper water at 100 m in winter (Boldyrev and Bogutskaya ,2007). During the spawning season they are absent from shallow waters (5-10 m) and deeper waters (30-50 m)(Pinchuk et al. in Miller, 2004).

Age and growth

Up to 4 age groups are known (Reshetnikov, 2002) although Pinchuk et al. in Miller (2004) indicate life-span may be only 18 months.

Food

Corophiid crustaceans and some molluscs such as gastropods are principal diet items. Cumaceans, amphipods and Nereis worms are also important food items (Pinchuk et al. in Miller, 2004).

Reproduction

Mature males measure 8 cm while a mature female from the same collection was 6 cm (Berg, 1948-1949) while Pinchuk et al. in Miller (2004) state that both sexes are mature by 4.0 cm, in the first year of life. Eggs have a maximum diameter of 1.2-1.8 mm and a minimum diameter of 0.3-0.5 mm and fish up to 5.2 cm long produce up to 1860 eggs (Reshetnikov, 2002). Spawning takes place from the second half of May to August and rarely into September with a second batch of eggs laid in July (Pinchuk et al. in Miller, 2004). Eggs are laid on silt and silt-sand bottoms, probably using shells, at 10-25 m (Pinchuk et al. in Miller, 2004).

Parasites and predators

None recorded from Iran.

Economic importance

This species is food for sturgeon and Sander species (Pinchuk et al. in Miller, 2004).

Conservation

Kiabi et al. (1999) consider this species to be data deficient in the south Caspian Sea basin according to IUCN criteria. Criteria include possibly few in numbers, limited range (less than 25% of water bodies), absent in other water bodies in Iran, and absent outside the Caspian Sea basin.

Further work

The biology and distribution of this species in Iranian waters needs to be studied.

Sources

Type material: See above (BM(NH) 1897.7.5:16).

Iranian material:

Benthophilus casachicus
Ragimov, 1978

Endemic to the Caspian Sea and found on the eastern Caspian coast, the Volga delta area, and from the south Caspian Sea by Naseka and Bogutskaya (2009). No Iranian record.

Benthophilus ctenolepidus
Kessler, 1877

Common names

gavmahi-ye shafaf or shaffaf (= transparent or clear goby); gavmahi-ye Lankaran or gavmahi-ye Lenkoran.

[prozrachnaya pugolovka or transparent tadpole goby, shipogolovaya pugolovka or spiny-headed (?) tadpole goby in Russian; spiny-scaled tadpole goby; Lenkoranskaya pugolovka or Lenkoran tadpole goby in Russian for the subspecies in the southern Caspian Sea].

Systematics

A subspecies reported from off the Iranian coast was Benthophilus ctenolepidus pinchuki Ragimov, 1982 but this is now recognised as a distinct species by Pinchuk and Miller in Miller (2004) (see below). The lectotype of Benthophilus ctenolepidus as designated by Ragimov (1982) is in the Zoological Institute, St. Petersburg under ZISP 10897 with 3 paralectotypes lost and 1 paralectotype from ZISP in the Natural History Museum, London under BM(NH) 1897.7.5:13 (66.9 mm standard length) (Eschmeyer et al., 1996). The type locality is "yuzhnoi i srednei chastyakh' Kaspiiskago morya" (= southern and middle Caspian Sea), or Caspian Sea, 40°08'N, 0°26'E off Baku, Azerbaijan (in Eschmeyer's "Catalog of Fishes" online, downloaded 22 August 2007). Records of B. magistri Iljin, 1927 in the Caspian Sea, as B. m. lencoranicus Ragimov, 1982, are this species (Boldyrev and Bogutskaya, 2007). The taxon Benthophilus magistri lencoranicus Ragimov, 1982 was described from "raione protib Zelenogo bugra (38°10' s. sh.)" (= area opposite Green Hill, Turkmenistan). The holotype of lencoranicus is in the Caspian Biological Station, Institute of Zoology, Azerbaijan under CBSIZA 546 (no. 2), paratypes under 546 (20) and additional non-type material in the Zoological Institute, St. Petersburg under ZISP 23131 and 41912 (9) (Eschmeyer et al., 1996). Boldyrev and Bogutskaya (2007) were unable to locate IZA 546.

Key characters

The narrow head has a trough-shaped depression or groove on the occiput, the large tubercles are vertically elongate, curved and crest-like and have their rear edges fringed with a comb of spines, dorsal row tubercles usually 30-33, abdominal row tubercles usually 24-26, temporal and occipital region without large tubercles but having granules, the body has tiny granules anteriorly interspersed among the large tubercles, dermal filaments present or absent, a small but well-developed chin barbel is present, the dermal process behind jaw angle is lobed, narrow, with an acute protuberance, first dorsal fin spines 3-4 (rarely 2), and back without brown bands.

Morphology

First dorsal fin with 3-4 spines, second dorsal fin with 1 spine followed by 8-11, usually 9-10, soft rays. Anal fin with 1 spine followed by 7-11, usually 8, soft rays. Total vertebrae 29-230. Dorsal row tubercles 27-33, usually 30-33 (?) or 22-24 (Boldyrev and Bogutskaya, 2007), lateral row 11-22, usually 17-19, abdominal 20-26, usually 24-26 (22-24 in Boldyrev and Bogutskaya (2007)). Tubercles on the head (when present) and flanks are large and bear sharp spines, the flank tubercles being angular in shape. The dorsal row of tubercles is incomplete, usually beginning below the first dorsal fin. The first 3-4, and as many as 7, dorsal row tubercles are markedly smaller than the succeeding ones. The dermal plate or fold behind the mouth corner is large (1.2 times eye diameter), oval, and without a wavy margin.

Sexual dimorphism

Undocumented.

Colour

Mostly transparent as the name suggests, a light to ash grey. The body bears dark lines and speckles but no strong dark bands, spots or blotches. The belly is yellowish-white. The dorsal and pectoral fins have thin banding and the pelvic and and anal fins are pale.

Size

Reaches 10.7 cm.

Distribution

Found in the central, southwestern and southeastern Caspian Sea and in adjacent Iranian waters (Ragimov, 1965; Abdoli and Naderi, 2009). Records are from Astara on the Azerbaijan-Iran border to Hasan Kuli in Turkmenistan and adjacent waters.

Zoogeography

A Caspian Sea endemic.

Habitat

Found in depths of 0.5-74 m in littoral areas but deeper than B. magistri and B. macrocephalus. It feeds and spawns in 0.5-10.0 m and migrates to deeper water in winter. Occurs in higher salinity areas at 12.4-13.0‰. Jolodar and Abdoli (2004) place it at 30-200 m depths in the south Caspian Sea basin. The pelvic fin is less developed, shorter and not as dilated than in these shallow water species.

Age and growth

Life span is probably not more than 18 months with maturity at one year (Pinchuk and Miller in Miller, 2004).

Food

Food in northern Caspian specimens was found to include mostly corophiid amphipods, with some gammarids, other crustaceans, and gastropods (Pinchuk and Miller in Miller, 2004).

Reproduction

Spawning is thought to occur from spring to autumn and into winter (Pinchuk and Miller in Miller, 2004).

Parasites and predators

None reported from Iran.

Economic importance

None.

Conservation

Kiabi et al. (1999) consider this species to be data deficient in the south Caspian Sea basin according to IUCN criteria. Criteria include possibly few in numbers, limited range (less than 25% of water bodies), absent in other water bodies in Iran, and absent outside the Caspian Sea basin.

Further work

The biology and distribution of this species in Iranian waters needs to be studied.

Sources

Based on Pinchuk and Miller in Miller (2004).

Type material: See above under B. ctenolepidus (BM(NH) 1897.7.5:13).

Benthophilus granulosus
Kessler, 1877

Endemic to the Caspian Sea and found in coastal areas of the North and eastern and western Middle Caspian. No Iranian record although Kottelat and Freyhof (2007) map this species from the Iranian shore.

Benthophilus grimmi
Kessler, 1877

A Caspian Sea endemic found in the western part of the Middle Caspian and the northern part of the South Caspian south to about 39º40'N, and from the south Caspian Sea by Naseka and Bogutskaya (2009). No Iranian record.

Benthophilus kessleri
Berg, 1927

A Caspian Sea endemic found in coastal waters of the eastern Middle Caspian, and from the south Caspian Sea by Naseka and Bogutskaya (2009). No Iranian record.

Benthophilus leobergius
Berg, 1949

Common names

gavmahi-ye akhtari (= stellate or starry goby), gav mahi bacheh qurbaqei, gavmahi-ye bacheghoorbagheie, sebele.

[ulduzlu comcaxul or khazar julduzlu chomchakhulu in Azerbaijan; zvezdchataya pugolovka or stellate tadpole goby in Russian; Caspian starry goby, Caspian stellate goby].

Systematics

This taxon was originally described as a subspecies of Benthophilus stellatus (Sauvage, 1874). B. stellatus is now restricted to fish from the Black Sea. Note that views do conflict on the status of this taxon and more studies remain to be done (Pinchuk et al. in Miller, 2004).

The subspecies leobergius (sic; leobergi has been used, see below) was first proposed by Iljin (1949) and was thought not be available as no distinguishing features were given in the description (Eschmeyer et al., 1996) although Berg (1948-1949) gives a description referring to "higher development of spiny tubercles" and colouration "less bright" than then Black Sea form in his volume III dated 1949 (not 1948 as given in Eschmeyer et al. (1996)). Eschmeyer et al. (1996) consider that Berg may be the author of this taxon and Pinchuk et al. in Miller (2004) concur. The holotype is probably in the Zoological Institute, St. Petersburg under ZISP 10891 (Eschmeyer et al., 1996). Berg (1948-1949) notes that the taxon is found "in the freshened regions throughout the Caspian Sea, south to Astrabad Bay (No. 10891, 5 August 1874, 62 mm)". This may be the type locality, i.e. Gorgan (= Astrabad) Bay, Iran. However this specimen may be merely recording the southernmost distribution of the taxon and a larger fish at 103 mm under ZISP 23128 labeled as from the Caspian Sea could be the holotype as it is depicted in two views (Figure 858, p. 1115 in the Russian text and p. 196 in the English text, and Figure 859, p. 1116 Russian and p. 197 English). Both these fish may be syntypes. Iljin (1949) simply referred to "Kaspii", i.e. the Caspian Sea, as the type locality. Boldyrev and Bogutskaya (2007) selected ZISP 23128 as the lectotype.

Benthophilus macrocephalus variety b (large-scaled) Kessler, 1877 is a synonym of B. s. leobergius according to Berg (1948-1949). Benthophilus aculeatus Baer in Lukina, 1984 is possibly a synonym, with no types known (Boldyrev and Bogutskaya, 2007).

Kottelat (1997) considers, tentatively, that Benthophilus stellatus is restricted to the Black Sea basin. Furthermore, Kottelat points out that perhaps the Caspian Sea species is Benthophilus leobergi Ragimov, 1978, leobergi since the species is founded on a personal name and should be emended to this spelling and Ragimov, 1978 since the name is a nomen nudum in Iljin (1949) (no description or diagnostic characters (but see above)) until Ragimov (1978) makes the name available by listing characters distinguishing it from B. s. casachicus. Benthophilus casachicus Ragimov, 1978 is then another Caspian Sea species.

Key characters

Distinguished by stellate tubercles, mostly spinous at the tip, tubercles rows on body well-defined but scattered on head, dorsal tubercles 25-30, abdominal tubercles 21-25, granules present predorsally but mostly absent from head and body, interorbit without a median groove, chin barbel present but small (eye diameter or less), dermal process behind jaw angle obvious, 3-4 first dorsal fin spines, and first dorsal fin patterned. and dark bands present.

Morphology

The snout is very long, 30-33% of head length, and the upper jaw projects. The head is wider than long. There are 3-4 neuromasts in row 7 (also in B. macrocephalus, usually 1 in other species). First dorsal fin spines 2-6, usually 3-4, second dorsal fin with 1 spine and 6-11 soft rays, usually 8-10, anal fin with 1 spine and 7-9 soft rays. Pectoral fin rays 15-18. The gill arch is without evident rakers and is irregularly tuberculate. Vertebrae number 28-31. There are 10-11, often 11, precaudal vertebrae (commonly 9 in other Benthophilus). There is a flap behind the jaw angle. The gut is moderately long with several convolutions. The body bears 3 rows of very spinulose tubercles; the upper row numbering 25-30, usually 26-28 or 27-29 (authors vary), the lower 21-25, usually 23-25 or 22-24 (authors vary), and the dorso-lateral row 15-24, usually 18-20. The ventro-lateral row is usually absent.There are numerous small tubercles between the main rows on the flanks, particularly the dorsal and lateral rows. The head surface has numerous small and large tubercles, arranged irregularly. Small tubercles or granules are even found on the medial half of the eyeball. There are 2 well-developed tubercles on the midline between the eyes (in 80% of specimens) as opposed to one or none in other Benthophilus.

The chromosome number of B. leobergi is 2n=44 with 46-48 chromosomal arms (Grigoryan and Vasil'ev, 1993; Vasil'yev and Grigoryan, 1993) or 2n=46 (Klinkhardt et al., 1995).

Iranian specimens had the following meristics: first dorsal fin spines 3(1) or 5(1), second dorsal fin with 1 spine and 9 soft rays (2), anal fin with 1 spine and 8 soft rays (2), and pectoral fin with 17(2) branched rays. The extent of tubercle rows is difficult to distinguish since their size gradually decreases posteriorly and their arrangement is irregular anteriorly. Dorsal row tubercles 31(2), lateral row tubercles ca. 19(2), and lower row tubercles 23(1) or 24(1).

Sexual dimorphism

The pectoral fin is longer than the pelvic fins in males when spawning. Females have a very small chin barbel.

Colour

Overall colour is brownish to whitish-grey. The head surface and upper flank have large, dark brown spots and small dark speckles. The body has about 5 transverse bands which circle the first dorsal fin, rear of the second dorsal fin and caudal fin base when viewed from above. Rows of speckles are found on the second dorsal, pectoral and caudal fins.

Size

Reaches 10.7 cm.

Distribution

Found in the Caspian Sea. In Iran, it is reported from Gorgan Bay (Abdoli, 1994a). the Safid River (Abbasi et al., 1999), the southeast Caspian Sea, southwest Caspian Sea and south-central Caspian Sea (Kiabi et al., 1999; Abdoli and Naderi, 2009). Jolodar and Abdoli (2004) consider it to be widely distributed along the Iranian coast but more abundant in Gomishan Wetland and Gorgan Bay. It is also recorded between Kultuk and Astara in Azerbaijan, near the Iranian border (Ragimov, 1965).

Zoogeography

Endemic to the Caspian Sea.

Habitat

This species is found in the Caspian Sea and in brackish and fresh waters, and is widespread in coastal waters over sand and shell debris bottoms down to at least 100 m. Said to prefer salinities below 9‰. Feeding and spawning occur in shallow water at 0.5-10.0 m, retreating to deeper water in winter (Boldyrev and Bogutskaya, 2007). It is said not to enter fresh waters unlike its relative B. stellatus but is known from fresh water in the Volga delta. In spring and autumn it can be found in coastal zone at 1-20 m. It prefers warmer waters at 18-26ºC (Caspian Sea Biodiversity Database, www.caspianenvironment.org). e

Age and growth

Young fish reach 1.83 cm in January and early February after spawning in autumn and reach their maximum size in July-August of the following year at 8.5-9.4 cm and 21.6-24.7 g. The spring spawners reach 1.9-2.0 cm and 0.17-0.24 g in May-June and 3.5-5.9 cm and 1.1-5.4 g in July-August. With two spawning seasons, there are always fish of markedly different size and maturation stage in any population. Life span is about 18 months (Ragimov, 1985a). Abdoli et al. (2006) examined fish from the Gomishan and Miankaleh wetlands and found a length (L)-weight (W) relationship of W = 0.0499L2.5 for males.

Food

Diet is dominated by molluscs (bivalves and gastropods) but includes crustaceans, worms, insect larvae and small fishes. An Iranian specimen contained the remains of a crab.

Reproduction

In the southern Caspian spawning occurs at depths of 30-35 m and in October-December and March-April (in the northern Caspian it is in shallow water (1-5 m) and during summer months). Batch spawning occurs and eggs number up to 2136 large and 1284 small with diameters 1.5-2.1 mm and 0.3-0.7 respectively (Ragimov, 1985a). Fecundity may reach 3500 eggs and maximum diameter 4.5 mm (Reshetnikov, 2002). Males protect the nests; unprotected nests have their eggs eaten by crustaceans within 2-3 hours (Caspian Sea Biodiversity Database, www.caspianenvironment.org).

Parasites and predators

Sattari et al. (2004) record the nematode Anisakis sp. from this species in Gilan. This fish is eaten by other fishes and by seals.

Economic importance

None.

Conservation

Kiabi et al. (1999) consider this species to be of least concern in the south Caspian Sea basin according to IUCN criteria. Criteria include medium numbers, medium range (25-75% of water bodies), absent in other water bodies in Iran, and present outside the Caspian Sea basin.

Further work

The biology and distribution of this species in Iranian waters needs to be studied.

Sources

Iranian material: CMNFI 1971-0326A, mm standard length, (): CMNFI 1970-0543, (); 3, 90.4-90.9 mm standard length, Mazandaran, Gorgan Bay and the Caspian Sea.

Benthophilus leptocephalus
Kessler, 1877

A Caspian Sea, deep-water endemic found in the western Middle Caspian south to 39º41'N and the eastern South Caspian south to Hasan Kuli in Turkmenistan. No Iranian record.

Benthophilus leptorhynchus
Kessler, 1877

A Caspian Sea, deep-water endemic known from the western coast of the Middle Caspian and the north of the South Caspian south to Baku. No Iranian record.

Benthophilus macrocephalus
(Pallas, 1788)

Common names

gavmahi-ye vazaghi (= tadpole goby).

[iribas comcaxul in Azerbaijan; Kaspiiskaya pugolovka or Caspian tadpole goby in Russian; bighead goby].

Systematics

Eschmeyer et al. (1996) gave the publication date as 1788 (later 1787), Berg (1948-1948) and Pinchuk and Miller in Miller (2004) as 1787. The original description is dated 15. Mars. 1787 but was probably published in the following year (not verified). The type locality is the Caspian Sea near the mouths of rivers and streams, and in small inlets, although the type of Gobius macrocephalus was probably from the Volga delta. No type specimens are known.

Key characters

Distinguished by having moderate, stellate tubercles, spinous at their apex with usually 22-23 (but see below) in the dorsal row and 19-24 in the abdominal row, numerous granules over the head and body, head wide, width less than to slightly more than length, large snout tubercle, two rows of tubercles from dorsal rows extend onto temporal and occipital areas, no occipital groove, chin barbel present, dermal process behind jaw angle very large (base length 20-26% head length), rounded or scalloped in larger fish, first dorsal fin spines 3-4, no dark annular bands.

Morphology

There are 3-4 neuromasts in row 7 (shared with B. leobergius, usually one in other Benthophilus). The head is very wide , 103-111% head length (only B. leobergius has similar measurements). First dorsal fin with 2-4 spines, usually 3, second dorsal fin with 1-2 spines, usually 1, followed by 6-10 soft rays, mostly 8-9. Anal fin with 1 spine followed by 6-9, usually 7, soft rays. There are no evident gill rakers on the arch but rounded tubercles are present. Total vertebrae number 27-28, with 17-19 caudal vertebrae, usually 18. The head and body are covered with minute granules with larger stellate scutes or tubercles bearing sharp spines on the body and on the dorsal and lateral head surfaces. Granules extend onto the medial half of the eyeball and onto the pectoral fin base. Scutes are few and scattered on top of the head, strongly developed and close together on the side of the head. There are three longitudinal rows of tubercles on the body with 19-27, usually 22-23 (or 23-27, modally 25, authors differ) tubercles in the dorsal row, 8-16, usually 10-13, laterally, and 19-24 abdominally, usually 21-22. The ventro-lateral row is usually absent. The belly is naked. The gut is moderately elongate with several convolutions.

Iranian specimens had the following meristics: first dorsal fin spines 3(3), second dorsal fin with 1 spine and 8 soft rays (4), anal fin with 1 spine and 7 soft rays (4), and pectoral fin with 16(1) or 17(3) branched rays. The extent of tubercle rows is difficult to distinguish since their size gradually decreases posteriorly and their arrangement is irregular anteriorly. Dorsal row tubercles 25(4), lateral row tubercles 9(1), 13(2) or 14(1), and lower row tubercles 21(1), 22(2) or 23(1).

Sexual dimorphism

Mature males lose the granules, scutes and tubercles, their dorsal and anal fins enlarge and their cheeks become more muscular or inflated.

Colour

Overall colour is ash-grey with scattered melanophores on the upper flank and dorsal surface. Melanophores line the pectoral and caudal fin rays being best developed on the more dorsal rays.

Size

Reaches 13 cm.

Distribution

This species is reported from the Caspian Sea including the south from the Anzali Mordab and Gorgan Bay in Iran and between Kultuk and Astara in Azerbaijan (Ragimov, 1965).

Zoogeography

A Caspian Sea endemic.

Habitat

Found mostly in the sea over muddy bottoms but may approach river mouths. Commonly at 0.5-10.0 m, migrating to deeper areas at 20-25 m in winter. De Filipii (1865) reports this species from the Murdab (= Anzali Mordab presumably) in relatively fresh water.

Age and growth

Young fish taken in early June are 2.4-3.5 cm and 0.35-1.00 g and by the end of November they are 7.0-8.0 cm and 10.15 g, the maximum values for the species. By February-March they are nearly mature (Ragimov, 1985a). There are up to 4 age groups (Reshetnikov, 2002) although Pinchuk et al. in Miller (2004) suggest 18 months.

Food

Food includes molluscs (80%) crustaceans, fish, worms and chironomids (Zenkevitch, 1963). Iranian fish contained clams, crustaceans and a polychaete in gut contents.

Reproduction

Spawning takes place in April-May and rarely up to mid-June near the shore on silt or silt-sand bottoms with a mixture of shells. The eggs are laid in large, empty mollusc shells. Egg laying occurs in two batches as females carry eggs of two sizes. Fecundity of the larger eggs is up to 3180 and the smaller 1403. Their diameters are 2.0-3.0 mm and 0.3-0.6 mm respectively. Young are caught in early June (Ragimov, 1985a). Males probably die after spawning. An Iranian fish, 77.9 mm standard length, caught on 13 March had minute but developing eggs.

Parasites and predators

None reported from Iran.

Economic importance

None.

Conservation

Numbers in Iranian waters and threats to this species need to be assessed.

Further work

The biology and distribution of this species in Iranian waters needs to be studied.

Sources

Iranian material: CMNFI 1970-0543, CMNFI 1970-0544, mmm standard length, (); CMNFI 4, 55.3-82.8 mm standard length, Gilan, Caspian Sea near Bandar Anzali ().

Benthophilus mahmudbejovi
Ragimov, 1976

Endemic to the Caspian Sea and found on the western coasts of the Middle and South Caspian and the Volga delta region. No Iranian record.

Benthophilus pinchuki
Ragimov, 1982

Common names

None.

[Pugolovka Pinchuka or Pinchuk's tadpole goby in Russian]

Systematics

Originally described as a subspecies of B. ctenolepidus, Pinchuk and Miller in Miller (2004) recognise it provisionally as a distinct species. This taxon was described from "raione Belogo bugra y vostochnogo poberezh'ya Yuzhnogo Kaspiya" (= area of the White Hill (? locality not ascertained further by me) near the eastern coast of the southern Caspian Sea). A specimen listed as a syntype is in the Natural History Museum, London (no number, noted in October 1982). The holotype of pinchuki is in the Caspian Biological Station, Institute of Zoology, Baku, Azerbaijan under CBSIZA 52 (later in ZISP 53569, ex IZA), paratypes are under CBSIZA 304 (24, later 8 fish and ZISP 53660 9ex IZA 304) (6)). Additional material is in the Zoological Institute, St. Petersburg under ZISP 33143 (6) and 44346 (1) (Eschmeyer et al., 1996; on-line version downloaded 22 August 2007).

Key characters

Tubercles are vertically elongated, curved and crest-like, with a spiny fringe on their rear edge, dorsal row tubercles complete and 30-33, ventral row tubercles 23-27, granules are absent postorbitally on the head and on the body, temporal and occipital region without large tubercles, head narrow, the interorbit has a shallow median groove, dermal filaments present or absent, chin barbel usually present, dermal process behind jaw angle narrow (less than eye diameter) and barbel-like, first dorsal fin spines 3-4, and colour without dark bands.

Morphology

The first dorsal fin has 3-4 spines, usually 4, the second dorsal fin 1 spine and 9-11 soft rays, and the anal fin 1 spine and 8-11 soft rays (Ragimov, 1982; note the holotype has a much lower count of anal fin rays than figure in table). Dorsal row tubercles 30-33, usually 31-32 (other species having 30 or less usually except B. ragimovi), ventral row tubercles 23-27, and dorso-lateral row tubercles 10-23. Total vertebrae 30-32 and caudal vertebrae 21-23 (26-31 and and 17-21 in most other Benthophilus). The pelvic fin extends to the anus or beyond. Chin barbel less than half eye diameter. One neuromast in row 7.

Sexual dimorphism

Males may have longer pectoral fins than females and grow larger than females.

Colour

Not described except for the absence of dark bands, blotches and spots. Probably an overall greyish.

Size

Reaches 8.9 cm in males.

Distribution

Found throughout the Caspian Sea.

Zoogeography

Endemic to the Caspian Sea.

Habitat

Known down to 282-294 m, usually 50-100 m but as shallow as 20 m, and a salinity of 12.4-13.2‰.

Age and growth

Life span is probably 18 months with maturity at 1 year.

Food

Unknown.

Reproduction

Spawning probably occurs in spring as fry have been caught in late April.

Parasites and predators

Unknown.

Economic importance

None.

Conservation

Numbers in Iranian waters and threats to this species need to be assessed. Boldyrev and Bogutskaya (2007) call it a rare deepwater species.

Further work

The biology and distribution of this species in Iranian waters needs to be studied.

Sources

No specimens examined, based on Pinchuk and Miller in Miller (2004).

Benthophilus ragimovi
Boldyrev and Bogutskaya, 2004

Endemic to the Caspian Sea from the western coast of the Middle and South Caspian, south to Astara. No Iranian record.

Benthophilus spinosus
Kessler, 1877

Endemic to the Caspian Sea from the eastern and western coast of the Middle Caspian , the southeastern North Caspian and southeastern South Caspian. No Iranian record.

Benthophilus svetovidovi
Pinchuk and Ragimov, 1979

A Caspian Sea endemic found on the eastern coast of the Middle Caspian, and from the south Caspian Sea by Naseka and Bogutskaya (2009). No Iranian record.

Genus Boleophthalmus
Valenciennes, 1837

This genus of gobies comprises several species in the Indian Ocean including one reported from Iran.

The genus is characterised by a very elongate, compressed body, a rounded head, greatly thickened head and nape epidermis (as protection against drying when out of water), scales cycloid, moderate to minute in size, eyes high on the head and close together, lower eyelid or dermal cup well-developed, mouth slightly oblique, teeth in a single row in each jaw, upper jaw teeth conical with some canines, lower jaw teeth flattened, obliquely notched and almost horizontal, tongue truncate and nearly all adnate to mouth floor, gill opening small and oblique, and second dorsal and anal fins long (23 or more rays).

Boleophthalmus dussumieri
Valenciennes, 1837

Common names

eshlambo, neeshlambo, gel khorak.

[kelb-el-daw, abou-shlembo, abu-shelamboo or triton in Arabic; gullo in Pakistan; Dussumier's mudskipper].

Systematics

Boleophthalmus Dussumieri was originally described from Bombay, India and the holotype is in the Muséum national d'Histoire naturelle, Paris under MNHN A.1468 (Bauchot et al., 1991).

Synonyms are Boleophthalmus dentatus Valenciennes in Cuvier and Valenciennes, 1837, described from Bombay and reported from the Shatt al Arab, Iraq by Iraqi authors, and Boleophthalmus chamiri Holly, 1929 from "Chamir an der Straße von Clarence gegenüber der Insel Tawilah in der Meerenge von Hormus, südöstliches Persien" (Holly, 1929a; Berg, 1949) (Khamir is at 26°57'N, 55°36'E in Khowran Strait (= Clarence Strait) opposite Qeshm Island (= Tawilah or Tavileh Island). Boleophthalmus boddarti (Pallas, 1770) as reported by Relyea (1981) does not occur in the Persian Gulf region (Murdy, 1989).

Mirza et al. (1996) separate B. dentatus and B. dussumieri on the basis of first dorsal fin height, body depth, number of teeth on the jaws and the membrane between the two dorsal fins. Murdy (1989) defines the genus Boleophthalmus in part by the dorsal fins being separate, but does not give first dorsal fin height, body depth and tooth counts for B. dussumieri or its synonyms.

A syntype of Boleophthalmus chamiri is in the Naturhistorisches Museum Wien (NMW 13804).

Key characters

The anal fin base and second dorsal fin base are 34% or more of standard length and there are 2 canine teeth internal to the lower jaw symphysis in contrast to Periophthalmus, the other mudskipper of southern Iran.

Morphology

First dorsal fin spines 4-5, usually 5, second dorsal fin with 1 spine and 24-28 soft rays, anal fin with 1 spine and 23-27 soft rays, and pectoral fin branched rays 16-20. Scales are minute and embedded, 103-185 in longitudinal series; predorsal scales 48-56. Each scale is rounded with radii around a posterior focus. Circuli are few. There are 3 canine teeth on each side of the upper jaw (1-2 may be lost), overlapping it when the mouth is closed and obviously protruding in some preserved fish, and 39-76 upper jaw teeth and 43-71 notched lower jaw teeth, the latter projecting almost horizontally from the jaw. The number of teeth increases with age and B. chamiri was founded on juveniles with fewer teeth (Berg, 1949). Gill rakers are partly embedded in a membrane and not always easy to count; the range is 12-14 for Iranian specimens. Upper arm rakers are long and pointed, less developed on the lower arm. When appressed, each raker touches the adjacent one. Gut very elongate and coiled. Muscle blocks may be evident in preserved fish.

Meristic values for Iranian specimens are:- first dorsal fin with 4(2) or 5(20) spines, second dorsal fin soft rays 24(3), 25(5), 26(9), 27(3), or 28(2), anal fin soft rays 23(3), 24(4), 25(13), 26(1) or 27(1), and pectoral fin rays 17(3), 18(8) or 19(11). Total vertebrae 26(10).

Sexual dimorphism

The male has a higher dorsal fin and larger canine teeth than the female. Females were described as B. dussumieri while males were described as B. dentatus (Berg, 1949).

Colour

Overall colour a greyish violet or light blue. The eye is light yellow to brown around the rim. The opercle, pectoral fin base and area just above the pectoral base have small green spots. There are 3 greenish-brown blotches on the nape, 3 more at the first dorsal fin base and 2 others at the anterior second dorsal fin base. The first dorsal fin is purple to light blue with small black or brown spots, or with greenish spots at the base and wavy white lines and bluish spots over the whole fin. The second dorsal fin has 2-6 horizontal rows of oblong blue (or white in preservative) spots on the fin membranes on a purplish background. Large white spots are found on the base of interradial membranes 7, 10, 14, 17, and 22 on a specimen examined by Murdy (1989) but this varies markedly between individuals seen by me. The margin of this fin is black, edged with orange (white in preservative). The caudal fin is black distally and bluish near the base. The anal fin is transparent, the pectoral fin bluish and the pelvic fin whitish.

In preservative, larger fish are darker than small ones. The body is speckled darker above than below but this is still much lighter than the dark dorsal fins. The first dorsal fin is very dark, almost black, because the small pigment spots are so dense. The second dorsal fin is lighter with alternate horizontal light and dark rows on the membranes. Rays are somewhat lighter or equally dark. The second dorsal fin margin is transparent but some fish may lack this feature and be dark interradially (because of a dark row) and light at ray tips. The caudal fin rays are dark and the membranes are dark next to the rays but light in the centre. The anal fin is immaculate and the pectoral and pelvic fins are very light with some pigment lining the rays. Some pectoral fin pigment appears to be extensive vascularisation. Peritoneum dark brown to black.

Young fish have several vague bars posteriorly on the upper flank above the anal fin level. The anterior flank and side of the head is covered with small spots.

Size

Attains 24 cm and 35.4 g.

Distribution

Found from Iraq to Pakistan and the west coast of India. Reported from the Shatt al Arab, Iraq near Iranian Khuzestan by Menon (1956), Khalaf (1961), and Al-Nasiri and Hoda (1976). In Iran it is recorded from the Sarbaz River at Bahu Kalat in the Makran, and the Karun River near Mohammerah (Berg, 1949) and in the Gulf, Hormuz and Makran basins.

Abdoli (2000) maps this species from the lower Karun and Arvand rivers in the Tigris River basin, the lower Hilleh and Mand rivers in the Gulf basin, the Mehran, Kul and Minab rivers in the Hormuz basin and lower reaches of Makran rivers from the Jagin to Bahu Kalat.

Zoogeography

This species has ready access to marine waters and can easily move between rivers via the sea.

Habitat

This species is active on mud flats in streams under and above tidal influence but most information on behaviour and biology is based on studies of fish on marine tidal flats (as recounted below). In southern Iranian rivers, this species can be seen swimming with its head out of the water and pectoral fins splayed, presumably on the look-out for danger. Populations are very numerous on mudflats on lower reaches of rivers in Khuzestan. It hides in mud burrows when approached and a plume of dark muddy water can be seen emerging from the burrow, or it may bounce over the water surface. They are difficult to catch with a seine when approached from the shore because of this behaviour, scurrying rapidly from the land and shallows into deeper water, muddying the water and obscuring vision, and thus disappearing from view. However they can be caught in shallower rivers by using a seine net from mid-river to shore, after waiting for them to re-emerge and enter the shallows. Their reaction is still flight to water but the movement from water to land confuses them and they can be scooped up in their panic. In Khuzestan, local people whip them out of the water with palm fronds.

Both sexes of this species construct U-shaped, vertical burrows in sand to a depth of about 1 m in marine situations. Clayton and Vaughan (1986) illustrate burrow shapes, outline construction methods and experimented on territoriality by removal of fish. A polygonal, mostly pentagonal but significant numbers of hexagonal and quadrangular, dam of mud is built up around the burrow hole using the mouth, again by both sexes, and is used to reduce aggression between neighbours (Clayton and Vaughan, 1982; Clayton and Wright, 1989). This dam serves to keep water in the burrow at low tide although not all populations maintain this dam as completely. The burrows are used as shelter from low temperatures during winter and high temperatures in summer, at high tides, and when danger threatens. Air temperatures in Kuwait reach 51.1°C in summer and fall to -4°C in winter, conditions mirrored in Iraq. Polygonal dams were not seen in rivers, perhaps because of river flow and tidal fluctuations. Fish are active in the 13-40°C range and daily variations may be as much as 20C°. Fish tend to be inactive in winter in the northern Gulf, with feeding the main activity, and are easy prey for migratory birds feeding on the mudflats.

At low tide the fish emerges and raises its head, scanning the surrounding surface and recognising humans as threats at 5-7 m. Movement is with the aid of the muscular pectoral fins and the gill covers are puffed out, presumably to obtain water or to maintain a moist atmosphere for the gills. It does not usually travel further than the mud walls of its territory in contrast to Periophthalmus waltoni and is exposed to air for much shorter periods being submerged in water or watery mud most of the time. The goby basks when recently emerged from its burrow in order to raise body temperature to 14°C before moving out onto the mud. The minimum temperature for emergence is 10°C. Some larger fish will make rapid and short sorties from the burrow to test air temperatures. Evaporative loss of body fluids is used to cool the fish in summer and the body is kept wet by immersion in pools and burrows. Body temperatures do not exceed 33°C even when air temperatures were over 40°C. There can be up to 52 fish in 100 sq m in Kuwait (Tytler and Vaughan (1983) and Clayton and Vaughan (1988) on B. boddarti in Kuwait marine waters, identified as this species by Murdy (1989)). At the Khowr-e Mussa in Iran, each fish defends a dammed territory of about 1 sq m, this area containing the epiphytobenthic food it requires (Höpner and Kazem Maraschi, 1999).

The skin has large numbers of mucous cells which prevent desiccation on the mud flat and lubricate the fish during burrowing (Salih and Al-Jaffery (1980) on B. dentatus from Al Faw (= Fao), assumed to be this species). Gas exchange in air and water takes place through the gill, inner operculum, nasal, body and outer operculum skin in that order, as adaptations to low oxygen conditions in estuarine areas and to life out of water (Al-Kodhomiy and Hughes (1988) on B. boddarti but presumed to be this species). In addition to the territorial individuals, there are also smaller, non-territorial fish which roam more freely and are called "errant" (Clayton and Vaughan, 1988). Clayton and Vaughan (1988) describe in detail a series of body postures and movements associated with being stationary, moving, feeding, maintaining moisture levels, construction, interacting with other individuals, avoiding predators, and courtship. Males raise their dorsal fins when they meet, gape widely and fight mouth to mouth.

Age and growth

On the Karachi coast of Pakistan, juveniles and young of the year are found from September to December, growing from 45 mm standard length to 60 mm over three months (Hoda and Akhtar, 1985). In Gujarat, India three age groups are reported (Soni and George, 1986).

Food

Diet includes diatoms (Pankow and Huq (1979) recorded 114 species at Al Faw (= Fao)), blue-green algae, filamentous algae, insects, crustaceans, nematodes and teleost fish eggs in marine situations, the polygonal enclosure acting as a "farm" promoting growth of diatoms (Sarker et al., 1980; Clayton and Vaughan, 1988). It grazes on surface films of yellow-green algae at Fao (Pankow and Huq, 1979) and brown algae in Kuwait. The grazing action, with side-to-side head movements, is distinctive for this species among other mudskippers in the Gulf (Clayton and Vaughan, 1988). The fish leave their burrows on feeding excursions. Young are found in non-territorial groups in shallow pools, where in winter they take some time to warm up before leaving to feed. The pools offer a protective environment for the young.

These mudskippers are fed on by gulls, terns, bitterns, egrets and herons as well as a wide variety of other fishes and sea snakes (Clayton and Vaughan, 1988).

Reproduction

Spawning in the marine environment of the Khawr az Zubayr, Iraq takes place in June, the release of larvae being timed to take advantage of phytoplankton and zooplankton blooms (Hussain and Ahmed, 1999).

On the Karachi coast of Pakistan, 50% maturity (= first maturity) is given as 69 mm standard length in males and 72 mm in females (Hoda and Akhtar, 1985) while maturation starts at 60-65 mm standard length in males and 70-75 mm in females according to Hoda (1987). Spawning there occurs twice each year during April-May and July-September (Hoda, 1986) or April-May and July-August (Hoda and Akhtar, 1985). Fecundity range is 986-4912 eggs for fish 74-110 mm long and maximum egg diameter is 1.05 mm (Hoda, 1986). Hoda and Akhtar (1985) give a range of 970 eggs at 75 mm standard length to 4113 eggs at 110 mm, mean 2371 eggs.

Clayton and Vaughan (1988) give the start of the breeding season in Kuwait as March but the majority of males are only displaying by mid-April. Courtship continues until August and most territories have breached walls where fish have crossed over into other territories to find mates. Larvae appear as early as mid-July although most appear later than this. Territories are repaired in October and November.

Courtship involves competition between males for neighbouring females. Each male has on average 5 neighbours but only half will be females. Males identify themselves and try to attract females by elevating their fins and leaping from a lateral prone position with about two-thirds of the fish clearing the ground. Other signaling devices are pectoral fin waves, lateral tail-beats, and quivers. The female advances and the male leads her into his burrow where spawning presumably occurs. While errant females may be mated with territorial males, errant males having no territory do not show the leaping behaviour.

Parasites and predators

Migrating birds are heavy predators of this fish on mud flats (Tytler and Vaughan, 1983). Indian pond herons are known to eat this fish in Iran. E. Kahrom in the Crocodile Specialist Group Newsletter reports that muggers (marsh crocodile, Crocodylus palustris) eats a Periophthalmus in the Sarbaz River of Iranian Baluchestan (www.flmnh.ufl.edu/natsci/herpetology/newsletter/news173b.htm, downloaded 15 February 2002). However this may well be Boleophthalmus dussumieri.

Economic importance

None, although children play with them in the Karun River (www.abadan.net/abadanidictionary.html, downloaded 4 December 2003).

Conservation

There has been no assessment of the conservation status of this species but it is common on mudflats in the lower reaches of rivers Iran.

Further work

The biology of this species in Iranian fresh waters has not been investigated.

Sources

Iranian material: CMNFI 1979-0141, 1, 86.2 mm standard length, Hormozgan, Rud-e Kul (27º17'30"N, 56º03'30"E); CMNFI 1979-0142, 5, 74.6-137.5 mm standard length, Hormozgan, Baghu River (27º17'N, 56º28'E); CMNFI 1979-0145, 10, 42.6-136.6 mm standard length, Hormozgan, Geru River (26º55'N, 57º01'30"E); CMNFI 1979-0322, 5, 57.1-111.8 mm standard length, Baluchestan, Bahu Kalat River (ca. 25º45'N, ca. 61º26'E); OSU 8121, 1, 61.4 mm standard length, Baluchestan, Bahu Kalat (no other locality data).

Comparative material:- CMNFI 1985-0182, 3, 44.7-68.1 mm standard length, Iraq, Shatt al Arab (no other locality data); BM(NH) 1976.12.8:4-7, 4, 103.9-128.2 mm standard length, Iraq, Basrah (30º30'N, 47º47'E); BM(NH) 1954.11.10:3-7, 6, 79.9-147.5 mm standard length, Iraq, Fao (29º58'N, 48º29'E).

Genus Caspiosoma
Iljin, 1927

Caspiosoma caspium
(Kessler, 1877)

Caspian Sea basin, described and recorded from the northern and middle Caspian Sea, but no Iranian record. Ragimov (1998c), Miller (2004) and Reshetnikov (2002) give recent descriptions.

Genus Chasar
Vasil'eva, 1996

This genus has a single species in the Caspian Sea. The characters of the genus are covered in more detail by Miller in Miller (2004) and are the characters of the species below. It is recognised as distinct from the related Neogobius by head sensory papillae patterns (especially an additional transverse row before row b) and modally 7 first dorsal fin rays rather than 6. The number of the transverse suborbital neuromast rows of the cephalic lateral line system is an important diagnostic feature for the monotypic genus Chasar according to Miller (2004). Pinchuk and Ragimov (1985) count these rows consecutively from anterior to posterior as 1 – 8 while Miller (2004) postulates an additional transverse row anterior to the longitudinal suborbital row b labelling the posteriormost row as row 7 (Pinchuk et al. in Miller, 2004). A Farsi common name for Neogobius and related gobies is گاو ماهي (gav mahi).

Chasar bathybius
(Kessler, 1877)

Common names

[bychok-glubokovodny or deepwater goby]

Systematics

Described from Svinoi Island south of Baku, Azerbaijan. No types known.

Key characters

Nape with cycloid scales anteriorly, not quite reaching orbit level, head depth at eyes about equal or slightly less than width between upper origins of opercles, interorbital distance equal to or slightly less than orbit, angle of jaw below pupil of eye, snout longer than eye, up to 1.8 times orbit diameter, upper lip not swollen at angle, pelvic fin not reaching anal fin, except in young, anterior pelvic fin membrane without well-developed lateral lobes, caudal peduncle depth 0.3-0.5 length, first dorsal fin with 7 rays and little higher than second dorsal fin (which is lower posteriorly), overall colour semi-transparent, and lateral series scales mostly 55-60.

Morphology

First dorsal fin with 6-8 spines, usually 7, second dorsal fin with 1 spine and 14-16 soft rays, and anal fin with 1 spine and 11-14 soft rays, lateral series scales 55-65. Scales are overall ctenoid but cycloid on the nape, breast, opercle, abdomen and and lobe of the pectoral fin. The pectoral fin is long reaching back well beyond the level of the second dorsal fin origin. Lateral line system usually with eight or nine transversal suborbital neuromast rows, two rows ventral to the longitudinal suborbital row b. Four neuromast rows on the opercle. Lateral line system with eight suborbital neuromast rows arranged transversally, five before, three above and two below the longitudinal row b. Four rows of free neuromasts on the opercle instead of the characteristic three rows of all other neogobiines.

Meristics for Iranian specimens: first dorsal fin with 6(1), 7(14) or 8(1) spines, second dorsal fin with 14(4), 15(9) or 16(3) soft rays, anal fin with 11(3), 12(9) or 13(4) soft rays, pectoral fin with 18(10) or 19(6) rays, scales in lateral series 55(1), 56(3), 57(6), 59(1) or 60(1), and transverse rows of scales 16(8) or 17(4).

Sexual dimorphism

Males grow larger than females.

Colour

The body is semi-transparent with reddish or yellowish-brown colour. The myotomes are visible but scales are difficult to see. The back is darker than the flank and belly. Preserved fish have 4 faint saddles and faint spots along the lateral midline, the first dorsal fin has two brownish bands, and young have a stripe-like spot posteriorly on the first dorsal fin.

Size

Attains 29.0 cm.

Distribution

Caspian Sea basin, recorded between Kultuk and Astara in Azerbaijan (Ragimov, 1965). Kiabi et al. (1999) report this species from the southeast Caspian Sea, southwest Caspian Sea and south-central Caspian Sea in Iran. Miller in Miller (2004) maps this species from the Safid River westwards and from Gorgan Bay but there are no text records to go with the map. Recently recorded from Iran (Ahnelt, et al., 2007). South coast of the Caspian Sea especially in Gorgan Bay (Jolodar and Abdoli, 2004).

Zoogeography

Endemic to the Caspian Sea but related to the speciose genera Neogobius and Ponticola.

Habitat

A marine species, not entering fresh waters, found down to 75 m and perhaps deeper, especially in colder conditions. It is found on sandy and shelly bottoms and in smaller numbers on firm silt.

Age and growth

Life span is estimated to be 4 years, with maturity in the second year and spawning every year. Abdoli et al. (2006) examined fish from the Gomishan and Miankaleh wetlands and found a length (L)-weight (W) relationship of W = 0.0777L2.44 for sexes combined.

Food

Fish are an important food for larger specimens, smaller fish take crustaceans. Fish food includes Knipowitschia iljini and crustaceans include the decapod Palaemon, corophiids, mysids and some gammarids. A small number of worms (Nereis) are also eaten.

Reproduction

Eggs of only one size have been found in this species, suggesting a single spawning each year. Up to 2979 eggs have been recorded with a maximum diameter of 2.6 mm. Spawning in the southern part of the Middle Caspian Sea takes place from the second half of June until July. Males approach the coast in the first half of March or April while females remain at 20-30 m. Females enter shallow water in late April and in greater numbers in May and June. After spawning, fish retreat to areas deeper than 10-15 m.

Parasites and predators

Eaten by sturgeons and the Caspian seal. Sattari et al. (2004; 2005) surveyed this species in the inshore area of the Caspian Sea, recording Anisakis sp. and Dichelyne minutus.

Economic importance

None in Iran but caught in Lenkoran, Azerbaijan with fixed nets and in northern Azerbaijan with cast nets. The fish are oily but pleasant tasting (Pinchuk et al. in Miller, 2004).

Conservation

Kiabi et al. (1999) consider this species to be data deficient in the south Caspian Sea basin according to IUCN criteria. Criteria include medium numbers, medium range (25-75% of water bodies), absent in other water bodies in Iran, and absent outside the Caspian Sea basin.

Further work

The distribution of this species in Iranian waters needs clarifictaion.

Sources

Based on Pinchuk et al. in Miller (2004) and Ahnelt et al. (2007).

Iranian material: CMNFI 1971-0352, 8, 141.7-195.8 mm standard length, Gilan, Caspian Sea + Shalman River (37°28’N, 49°27'E, 37°08’N, 50°15’E); CMNFI 2006-0022, 3, 142.9-195.4 mm standard length, Gilan, southwest Caspian Sea off Astara (38°00’N, 49°30’E to 38°20’N, 50°00'E); CMNFI 2006-0023, 1, 99. 4 mm standard length, Gilan, Chaboksar (36°58’N, 50°34’E); CMNFI 2006-0024, 2, 160.4-183.1 mm standard length, Gilan, Talesh (37°48’N, 48°55’E); CMNFI 2006-0025, 2, 174.7-187.9 mm standard length, Gilan, Talesh (37°48’N, 48°55’E); CMNFI 2006-0026, 2, 76.7-81.3 mm standard length, Gilan, southwest Caspian Sea off Bandar Kiashahr (37°25’N, 49°57’E).

Genus Glossogobius
Gill, 1859

The gobies of this genus are found in fresh, brackish and marine waters in the Indo-West Pacific area. Most species are freshwater residents and there are about 24 species with one in Iran. They are characterised by an elongate body, anteriorly rounded and posteriorly compressed, head depressed, snout elongate, scales on the body ctenoid and on the head cycloid, head scaled behind eyes, cheeks and operculum scaleless, anterior nostril a short tube, lower jaw prominent, tongue bilobed, teeth in several rows in each jaw, gill openings wide, isthmus narrow, no flaps on the inner edge of the shoulder girdle, and pectoral fin without free rays and base scaled.

Glossogobius giuris
(Hamilton, 1822)

Sarbaz River, courtesy of Asghar Mobaraki.

Common names

gel-ye mahi cheshm navari (= band- or bar-eyed goby) or gavmahi-ye cheshmnavari.

[guloo in Pakistan; bar-eyed goby, tank goby, white goby, flathead goby, crocodile goby].

Systematics

Gobius giuris was originally described from the Ganges River, India.

Key characters

As the sole representative of the genus in Iran, the characters of the genus serve to identify it.

Morphology

The first dorsal fin has 6 spines, the second dorsal fin has 1 spine followed by 7-9 soft rays, the anal fin has 1 spine and 7-9 soft rays and the pectoral fin has 16-21 branched rays. Lateral series scales 28-36. Scales on the head are mostly cycloid while body scales are ctenoid. The exposed portion of flank scales are diamond-shaped. Scales are rectangular in overall shape with a rounded anterior margin which may be slightly indented above and below the mid-point, parallel dorsal and ventral margins, and a posterior margin with two straight edges meeting at a rounded central tip. The anterior dorsal and ventral corners are sharp. There are fine ctenii on the posterior margin. Numerous radii radiate anteriorly from the rounded central tip of the posterior margin. Circuli are fine. Esmaeili et al. (2009) give details of scale morphology using scanning electron microscopy. Gill rakers are short and reach the one below when appressed. The cheek has 3-5 longitudinal mucus canals. The gut is short and s-shaped.

The chromosome number is 2n=46, with 46 acrocentric chromosomes and 46 chromosomal arms (Subrahmanyam, 1969; Ráb, 1985; Vasil'ev and Grigoryan, 1993; Klinkhardt et al., 1995).

Meristics on Iranian specimens: first dorsal fin with 6(3) spines, second dorsal fin soft rays 9(3), anal fin soft rays 8(3), and pectoral fin rays 19(1) or 20(2). Scales in lateral series 32(1) or 36(1), total gill rakers 12(1) or 14(1) and total vertebrae 27(1).

Sexual dimorphism

Males are larger than females. Male dorsal fins are longer and their overall colour is brighter (Pethiyagoda, 1991).

Colour

Olive-green to blackish-green or yellowish to sandy brown on the back with flanks similar but lighter, often a greenish-yellow. The belly is cream coloured. The head and body are darkly blotched and spotted. The flank may have 5 regularly spaced blotches. The sides of the head have irregular dark to violet spots. The fins have thin bars appearing as rows of spots on each fin ray, in some concentrated at the base of the second spine of the first dorsal fin. Fins are a yellowish-green. The first dorsal fin may have a dark spot posteriorly and the anal fin a black margin. The pectoral fin has a dark spot above the base. Small preserved fish are immaculate. The peritoneum is silvery with a few small and scattered melanophores.

Size

Attains about 50 cm.

Distribution

Found in the Indo-West Pacific entering rivers. Abdoli (2000) maps this species as questionably present in the lower Karun and Arvand rivers of the Tigris River basin, questionably in the Gulf and Hormuz basins; and in lower reaches of Makran rivers from the Jagin to the Bahu Kalat or Sarbaz. It is recorded from the Hormuz and Makran basins by specimens.

Zoogeography

The marine abilities of some members of this family would aid in dispersal between river systems connected to the sea.

Habitat

Sand and mud substrates are preferred over rock in Sri Lanka (Pethiyagoda, 1991). The fish lies embedded in the sand with only the eyes protruding and rarely swim freely. Young fish may form schools or may rest close to shore in shallow water where they can be caught by hand. Tekrival and Rao (1999) report its aquarium preferences and habits as 22-25°C, pH 6.5-7.2, predator, almost dark lighting, bottom dwelling with stones preferred and a pair or more can be kept.

Age and Growth

Hossain et al. (2009) found a value close to isometric for this fish in the Ganges River of northwestern Bangladesh (b = 3.068 for total length and combined sexes, 2.954 for males and 3.293 for females)

Food

Diet includes small fishes seized from its concealment in the sand by a sudden lunge; this species being a visual ambush predator. It may browse on the substrate and take living food. An Iranian specimen had crustacean remains.

Reproduction

Eggs are green and when laid become strongly attached to any submerged structure by a tube 3-8 mm long. The eggs are 0.5 mm in diameter. Freshwater populations may migrate to the sea to breed as there is a marine larval stage although some are reported to be land-locked in South Asia and hence freshwater breeders (Pethiyagoda, 1991). The spawning period is June to September, peaking in July to August (Saksena, 1980).

Parasites and predators

None reported from Iran.

Economic importance

This goby is a food fish in South Asia and can be caught on light angling gear. It has been reported as being ciguatoxic (intermittently poisonous through feeding on toxic food) (Bagnis et al., 1970).

Conservation

The distribution and abundance of this species in Iranian waters has not been examined and no conservation assessment can be made.

Further work

This species seems to have a fairly restricted distribution in Iran and this should be investigated further.

Sources

Iranian material: CMNFI 1979-0322, 2, 20.7-53.6 mm standard length, Baluchestan, Sarbaz River (ca. 25º45'N, ca. 61º26'E); OSU 8124, 1, 162.0 mm standard length, Baluchestan, Bahu Kalat, Sarbaz River (25º43'N, 61º25'E).

Genus Hyrcanogobius
Iljin, 1928

Hyrcanogobius bergi
Iljin, 1928

This species was described from the north Caspian Sea near the mouths of the rivers Volga, Ural and Emba. Ragimov (1977) reports this species from off Gasan Kuli at 37°30'N in Turkmenistan near Iranian waters. Shakirova and Sukhanova (1994) record this species from the Atrek lakes of Turkmenistan, the Atrek River flowing into Iran for part of its course. Gobius longecaudatus var. c Kessler, 1877 described from Baku Bay is a synonym. Miller and Pinchuk in Miller (2004) map this species from Iranian waters (Gorgan Bay) but this is not confirmed by specimens. Miller and Pinchuk in Miller (2004) give a recent description of this species.

Genus Knipowitschia
Iljin, 1927

This genus of sand gobies has about 13 species in the Caspian, Black, Aegean and Adriatic seas and adjacent fresh waters with 2 recorded from Iran. Later spelt Knipovitschia, incorrectly, in the English summary of Iljin's 1928 description. The original description is apparently in Iljin (1927d) (as is that of Caspiosoma) but was repeated in Iljin (1928). Hyrcanogobius Iljin, 1928 and Bubyr Iljin, 1930 are synonyms (Economidis and Miller, 1990) although Miller in Miller (2004) separates Hyrcanogobius.

The body is fusiform without a strongly depressed head, the dorsal profile anteriorly being straight. The mouth angle does not reach past the anterior part of the eye. The mouth is oblique. The eyes are moderate in size and may be lateral or directed upwards. The body usually bears imbricate ctenoid scales but the head is naked as can be the anterior part of the back as far as the second dorsal fin and the middle of the abdomen to the anal fin. There may be axillary scales and scales on the belly midline. Some members lack scales except for the axillary region and the caudal peduncle. Lateral series scales less than 40. There are no barbels and the anterior nostrils are not an extended tube-like structure. The posterior nostrils are pore-like. The nape lacks a bony crown and only the posterior part of the cranium is covered with dorsal muscles. The preoperculum lacks teeth or projections. The isthmus is broad and the branchiostegal membrane is attached to its entire lateral margin. Jaw teeth are conical, caniniform and in several rows forming a band. The outer row is slightly enlarged but there are no strongly enlarged teeth. The vomer lacks teeth. The tongue is truncated or slightly rounded. The gill rakers are small and broadly spaced apart. The pectoral fins lack fine, silk-like rays. The anterior edge of the pectoral girdle lacks dermal flaps. The pelvic fin disc reaches to, or near to, the anus. The pelvic fin anterior membrane is well-developed but without obvious lateral lobes. The caudal peduncle is slender and longer than the second dorsal fin base. The upper angle of the caudal fin is nearly rectangular and the caudal fin is rounded and shorter than the head. The head canals are variably developed or absent, and may be damaged by abrasion in poorly preserved material. The first dorsal fin modally has 6 or 7 rays and vertebral modes are 30-33. The swimbladder is present or reduced. There is no perianal organ. Colour is light. Postlarvae are planktonic.

Miller in Miller (2004) also defines this genus by characters of the head lateral line system, namely having the infraorbital longitudinal row a present with typically at least 2 transverse rows from this row; transverse infraorbital rows between levels of rows b and d along cheek, with rearmost descending behind posterior end of row d; transverse row behind jaw angle not descending below level of row d; row b short, to below rear eye; anterior row tra distant from row b; anterior dorsal row g distant from row n; snout rows r and s linear; uniserial row of papillae along upper edge of each orbit when canals are absent but never with transverse rows; anterior oculoscapular canal (if present) ending anteriorly in interorbit, dividing from pore κ to end at double pores β; posterior oculoscapular present or absent; and preopercular canal when present always lacks middl;e pore δ.

Vasil'eva and Kuga (2001) propose that the separation of Knipowitschia from Pomatoschistus is doubtful based on similar cranial osteology and the variation in the degree of development of head canals seen in Knipowitschia. However Miller in Miller (2004) disagrees and presents arguments for a "sand-goby" group of 5 distinct genera, of which only Knipowitschia and Hyrcanogbius are known from Iran (Pomatoschistus, Gobiusculus and Economidichthys being the others).

Miller (1990) states that the majority of the members of this genus evolved after the draining of the Western Paratethys basin into the Adriatic Sea basin in the late Miocene about 6.0-5.5MYA.

.Members of this genus are called gav mahi and سگ ماهي (sag mahi) in Farsi and this not repeated under each Species Account.

Knipowitschia caucasica
(Berg, 1916)

Common names

gel-ye mahi qafqazi (= Caucasian goby) or gavmahi-ye qafqazi.

[gafgaz xulu in Azerbaijan; bubyr kavkazskii in Russian; Caucasus bald goby; Caucasian dwarf goby].

Systematics

Pomatoschistus caucasicus was originally described from Batumi/Temirgoe, a station south of the Sulak mouth on the Black Sea.

The name of this species first appeared in Kavraiskii in Radde (1899) as a museum name without a description ( a nomen nudum) and was subsequently made available by L. S. Berg in 1916 with the type locality being a swamp near Batum and Lake Inkit near Pitzunda, Georgia (see Eschmeyer (1990) and Eschmeyer et al. (1996) for details).

Gobius lenkoranicus Kessler, 1877 described from "pribrezhnom' bolote, po blizosti Lenkorana" (= a coastal marsh near Lenkoran, Azerbaijan) is a synonym, tentatively so according to Economidis and Miller (1990). This name was suppressed by the International Commission on Zoological Nomenclature (see Kottelat, 1997; Miller et al. in Miller, 2004)). This earlier name for what could be caucasica was suppressed as a nomen dubium because of an inadequate original description in its details and the small and poorly-preserved holotype which could not be readily compared with caucasica. The name could also have been suppressed as a nomen oblitum as it had not been used for 50 years (Economidis and Miller, 1990).

Formerly placed in the genus Pomatoschistus Gill, 1863 and in the genus Bubyr Iljin, 1930.

Three syntypes from Lake Temirgorje, Georgia (formerly in the Tiflis Museum) are in the Natural History Museum, London under BM(NH) 1896.3.28:26-28 (22.1-25.2 mm standard length) (Eschmeyer et al., 1996; personal observations). Economidis and Miller (1990) list these three specimens (as evidenced by the catalogue numbers 26-28) as two males (22.8-24.0 mm standard length) and two females but give the length of only one female (24.5 mm). Miller et al. in Miller (2004) consider the material used by L. S. Berg (see above) to be the type series (ZISP 15343 from Lake Inkit and ZISP 15321 a swamp near Batum).

Key characters

The anterior oculoscapular lateral line head canals unite in the posterior interorbit, with a single median pore κ, and canals extend anteriorly to pores λ. The preopercular canal is present (Miller in Miller, 2004).

Morphology

First dorsal fin with 5-8, usually 6, spines (rarely 4 as an anomaly, e.g. in Berg (1948-1949, Fig. 784 (Iljin, 1956)), second dorsal fin with 1 spine and 6-10 branched rays, usually 7-8, anal fin with 1 spine and 5-10 branched rays, usually 7-8, and pectoral fin with 13-18 rays. Lateral scales 29-38. Vertebrae 30-33. The back in front of the first dorsal fin and the belly in front of the anal fin are usually scaleless. The caudal peduncle is completely scaled and there is a complete row of scales anteriorly along the lateral midline, although other flank scales may be missing. The anterior oculoscapular head lateral line canals join in the posterior interorbit region, with one pore κ, and paired canals in the interorbit each to a pore λ and single pore α. The posterior oculoscapular canal is typically present, the preopercular canal is present and there are no additional pores on the horizontal anterior oculoscapular canal behind the eyes (Miller et al. in Miller, 2004). The subopercular canal is variably present which has led to different generic assignments for this species. The larva has been described by Daoulas et al. (1993).

Meristic values for Iranian specimens are:- first dorsal fin spines 6(4), second dorsal fin rays after a spine 6(1) or 8(3), anal fin rays after a spine 8(2) or 9(2), lateral scales 33(1), 34(1), 36(1), 39(1), and total gill rakers 9(3) or 10(1). Rakers are very short, not reaching half way to the adjacent one when appressed.

Sexual dimorphism

See under colour. Males are larger than females. There are various morphometric differences (Miller et al. in Miller, 2004).

Colour

Overall colour is grey to fawn with olive-green tints. Fish in hypersaline habitats are darker. Males are generally darker than females and have distinct dark bars on the flanks below the two dorsal fins and on the caudal peduncle while females have irregular spots or blotches along the flank mid-line. Nuptial males have 4 prominent bars. However some males may lack bars and have spots and some females lack spots (Ahnelt et al., 1995). The head in males is densely pigmented, including ventrally. The caudal peduncle is completely pigmented, usually including the ventral surface. The flank bar under the first dorsal fin in males is long and usually runs from near the dorsal mid-line to the belly. The male first dorsal fin has three oblique dark bands and a dark spot medially on rays 5 and 6 extending posteriorly on the fin membrane. The distal edge of this dorsal fin is dark. The second dorsal fin has three bands also and a dark edge. Males have dark pigment on the pelvic fins, head and breast, and a silver hue on the anal fin during the breeding season. The male pectoral fin has a short and oblique dark band on the upper rays. The anal fin in males is normally dark with a pale margin while in females it is pale overall. The caudal fin has thin dark bands.

Females are a pale fawn overall. The female has a densely spotted first dorsal fin while the male has a blue blotch at the fin rear when in breeding condition. The first dorsal fin in females has a distinctive middle band but the others are faint. The second dorsal fin in females is only faintly banded. Breeding females have 2 intensely black areas on the first dorsal fin, a black spot on the lower jaw, 2 rows of melanophores running obliquely from the eyes to the upper jaw, and a yellow abdominal region (Economou et al., 1994). The back has fine melanophores extending to the rear of the second dorsal fin. There are light saddles at the origins of the first and second dorsal fins, the end of the second dorsal fin and on the caudal peduncle. The mid-flank has several blotches, some slightly deeper than wide. The rest of the body below these blotches is pale except for some short vertical groups of melanophores below the lateral midline and melanophores along the base of the anal fin. The breast is pale as opposed to dark in males.

Large and minute pigment blotches and spots variably developed are scattered over the whole head and body in both sexes of an Iranian sample. The peritoneum has large pigment spots ventrally or is almost immaculate.

Size

Reaches 48.6 mm, perhaps 69 mm, total length. Fish in Gorgan Bay reached 46.0 mm (Afraei and Hassannia, 1999).

Distribution

Found in the Aegean, Black and Caspian Sea basins. Introduced to the Aral Sea. Caspian localities include Imeni Kirova or Kyzylagach Bay near the northwestern Iranian border, the lower Safid River and lower Babol River, near Anzali and Ashuradeh (Derzhavin, 1934). It is reported from Gorgan Bay, the delta of the Karasu, the Atrak River, the southeast Caspian Sea, southwest Caspian Sea and south-central Caspian Sea (Kiabi et al., 1999; Jolodar and Abdoli, 2004; Miller et al. in Miller, 2004; Abdoli and Naderi, 2009).

Zoogeography

This species is part of  marine fauna in the eastern Mediterranean and the Black-Caspian-Aral seas basins and its relationships are outlined under the genus description above.

Habitat

This species lives and reproduces in salinities as high as 59.5‰ and up to 83‰ (Zenkevitch, 1963; Miller et al. in Miller, 2004) but also enters fresh water. Preferred conditions in the Aegean area are 15-20‰ (Ahnelt et al., 1995; Miller et al. in Miller, 2004). Juveniles prefer sand, mud or gravel bottoms near shore in Greece (Daoulas et al., 1993) but weedy shallows are also favoured (Economidis and Miller, 1990). They may overwinter in deeper water at 1-2 m in the Volga delta (Economidis and Miller, 1990) and migrate into shallow water in April-May prior to spawning (Miller et al. in Miller, 2004). The Atrek River population moves downstream (Miller et al. in Miller, 2004). Afraei et al. (2001) found this species to be most frequent in Gorgan Bay in January and least frequent in March.

Age and growth

Life span may be only 1 year and as a consequence numbers can fluctuate widely in any given year depending on environmental conditions. In the Evros Delta of the north Aegean Sea, fish grow rapidly during the summer after hatching, breed from the end of April to the end of July in the following summer and grow rapidly again from July to September. Older males die the following year after February but some females will survive to spawn a second time at the end of April and the beginning of May and then die. Fish may be mature at 14.5 mm total length (Kevrekedis et al., 1990). In the spawning season, females comprise as much as 85.5% of the population (Miller et al. in Miller, 2004).

A study of this species in Gorgan Bay, Golestan (Iranian Fisheries Research Organization Newsletter, 23:2, 2000) showed that the maximum size obtained was 46 mm with average lengths of 33 and 37 mm for males and females respectively. The sex ratio was 1:1.1 for males to females and growth is positive allometric (see also Afraei and Hassannia (1999) and Afraei et al. (2001)). In the Volga delta spawning males were 33.3 mm and females 30.2 mm on average (Miller et al. in Miller, 2004). Abdoli et al. (2006) examined fish from the Gomishan and Miankaleh wetlands and found a length (L)-weight (W) relationship of W = 0.135L2.72 for males, 0.0377L1.97 for females and 0.0182L2.51 for sexes combined.

Food

Diet was dominated by polychaete worms and chironomids (88%) in one study but crustaceans such as amphipods, copepods, ostracods, cladocerans, insects such as choronomids, bivalve mollusc larvae, and fish are also taken and items may be both benthic and planktonic.

Reproduction

Females may be carrying eggs at 20-24 mm and 8-10 months of age. Fecundity reaches 1389 eggs but can be as low as 60 eggs. Fecundity in the Caspian Sea ranges from 209 to 786 eggs with a mean of 423, from 209 to 382, mean 285 and from 527 to 863, mean 715.6 for various reports (Kevrekedis et al., 1990). Batches consist of 80-100 eggs. Eggs are cylindrical, 2.6 x 0.97 mm. Larvae are pelagic and as small as 4.1 mm standard length. Relative fecundity in the Gorgan Bay study in Iran was 290-550, mean 395.5 eggs (Afraei et al., 2001).

There is a pre-spawning migration into shallow water in April in the Caspian Sea. The male approaches the female from below, touching her lower jaw with his snout and leading her to a reed nest. Such behaviour occupies several hours to more than a day. The female enters the reed nest, inverts and deposits eggs on the roof of the nest. The underside of bivalve shells or gravel may also be used. The male then fertilises the eggs, inverting to do so. The eggs are laid in a line and the pair will form other lines. Egg patches may eventually contain as many as 3000 eggs at different developmental stages, which indicates males may spawn with several females. Spawning takes about 1.5 hours and the female gradually loses the brightness to her breeding colours and almost all breeding pigment is lost within a day. The pair mated 4 times over a period of 35 days (28 March, 6 April, 14 April and 2 May in an aquarium with fish from Lake Trichonis, Greece). The male remains mostly inside the reed nest, aerating the eggs and defending them against other gobies. At irregular intervals (8-60 minutes), the male would leave the nest for 1-5 minutes, occasionally feeding but always remaining near the nest.

Spawning in the pre-estuary area of the Volga River in the Caspian Sea takes place from the middle of April to the end of May, rarely to June. Spawning in the Caspian takes place in shallow water with some current at a depth of 0.15 to 1.5 m, a temperature of 15-27°C and over a wide range of salinities (Kevrekedis et al., 1990).

Parasites and predators

None reported for Iran.

Economic importance

None.

Conservation

This species seems to be rare in Iran with only Derzhavin's (1934) report and one other record. Its status cannot be assessed until more is known about its distribution. Kiabi et al. (1999) consider this species to be data deficient in the south Caspian Sea basin according to IUCN criteria. Criteria include abundant in numbers, habitat destruction, limited range (less than 25% of water bodies), absent in other water bodies in Iran, absent outside the Caspian Sea basin (sic).  Vulnerable in Turkey (Fricke et al., 2007).

Further work

Further collecting will need to be carried out to determine its abundance and distribution in Iranian waters. Its small size may have led to it being ignored or confused with other gobies as young and it may be more widely distributed than current knowledge indicates.

Sources

Iranian material: Uncatalogued, 4, 22.2-35.3 mm standard length, Mazandaran, Gorgan Bay (no other locality data).

Knipowitschia iljini
Berg, 1931

Common names

gavmahi-ye Iljin, gel-ye mahi.

[Iljin's bychok in Russian, Iljin xulu in Azerbaijan].

Systematics

At least 4 syntypes are in the Zoological Institute, St. Petersburg (ZISP 22052) along with ZISP 24370 and possibly ZISP 24424 (Miller and Pinchuk in Miller, 2004). Although Berg (1931b) states "many specimens caught" this may refer to abundance while only 4 fish are illustrated and may be the type series. This species was described from "in the middle part of the Caspian Sea".

Key characters

The anterior oculoscapular lateral line head canals are more or less separate in the midline of the posterior interorbit, with double or separate pores κ, and canals extend anteriorly through interorbit of variable extent. The preopercular canal is absent or present (Miller in Miller, 2004).

Morphology

The first dorsal fin has 6-8 spines, usually 7, the second dorsal fin has 1 spine followed by 8-10 soft rays, the anal fin has 1 spine and 7-10, usually 9, soft rays, the pectoral fin has 15-18 rays. Lateral series scales 28-38. Vertebrae 31-33. Scales are ctenoid. The belly, head and the dorsal surface back to the second dorsal fin are all naked. There is no small, straight canal extending posteriorly from the junction of the orbital canals. The caudal fin is rounded and symmetrical. The anterior oculoscapular lateral line head canals do not unite in the posterior interorbit. Canals anterior to pores κ are typically absent and pore κ is double. The posterior oculoscapular canal is absent, the preopercular canal absent or present. Pore "a" is single or double with sometimes an additional pore or two on horizontal anterior oculoscapular canal behind eye.

Sexual dimorphism

Males have darker overall colouration and darker fins than the females (see below). The first dorsal fin in males has free tips and usually a dark spot at the rear margin. Males have a pointed genital papilla while females have a distally bifurcated papilla. The pelvic fins reach the genital papilla in males while in females they do not reach the vent. Females have a more slender caudal peduncle and the snout is more produced than in males.

Colour

The body overall has a brownish reticulate pattern. The back and flanks bear about 6-10 dark bands in males while in females bands are faint and restricted to the posterior flank. The male has a darkly edged caudal fin and the pectoral fin is dark ventrally. The dorsal fins have dark stripes and the anal fin has a dark stripe along its margin in males while females lack stripes. Females bear small dark spots irregularly distributed on the back and anterior flanks.

Size

Attains about 4.7 cm for males and 5.0 cm for females. Ragimov (2003) gives a maximum length of 5.2 cm.

Distribution

Ragimov (1965) reports this species between Kultuk and Astara in Azerbaijan and generally in the central and southern Caspian Sea, and mapped around the coast of the whole Caspian by Miller and Pinchuk in Miller (2004).

Zoogeography

This species is part of a marine fauna in the eastern Mediterranean and the Black-Caspian-Aral seas basins and its relationships are outlined under the genus description above.

Habitat

Found benthically and in the water column with adults in shallow coastal areas at 20-70 m with juveniles offshore as deep as 500 m Not recorded from fresh water.

Age and growth

Life span is estimated at one year (Miller and Pinchuk in Miller, 2004).

Food

Only mysids have been recorded from 8 fish examined (Miller and Pinchuk in Miller, 2004).

Reproduction

Gonad maturity begins in late August to early September, many fish are ripe by the end of January and most by the end of February. Spawning occurs from early April to possibly as late as September with a peak in May-June (Miller and Pinchuk in Miller, 2004). Fecundity reaches 2240 eggs and egg diameter 0.8 mm (Miller and Pinchuk in Miller, 2004).

Parasites and predators

None reported from Iran but sturgeon and Sander species eat this fish (Miller and Pinchuk in Miller, 2004).

Economic importance

None.

Conservation

Apparently rare in Iran but this may reflect collecting effort for this small species. Insufficient work has been devoted to this species to assess its status.

Further work

Further collecting will need to be carried out to determine its abundance and distribution in Iranian waters. Its small size may have led to it being ignored or confused with other gobies as young and it may be more widely distributed than current knowledge indicates.

Sources

Iranian material: None.

Knipowitschia longecaudata
(Kessler, 1877)

Black and Caspian seas, described from the southern and middle Caspian Sea, but no Iranian record. Ragimov (1965) reports this species between Kultuk and Astara in Azerbaijan. Sometimes spelt longicaudata erroneously in the literature.

Genus Mesogobius
Bleeker, 1874

This is an endemic Ponto-Caspian genus with 3-4 species, with one species being of uncertain status (see below). It is distinguished by having three (rather than the more usual two) transverse head papillae rows between rows b and d. Other characters include suborbital papillae in transverse rows, no row a, snout with longitudinal rows s1 and s2, no perianal organ; row 5i not below level of row 6i, 6i at or opposite end of row d, scales normal, anterior nostril elongate but not overhanging lip, and pelvic fins without lobules. A detailed, recent description is given by Miller in Miller (2004).

Mesogobius nigronotatus
(Kessler, 1877)

Caspian Sea basin, described from Fort Aleksandrov (= Shevchenko) on the Caspian Sea coast of Kazakhstan, and from the south Caspian Sea by Naseka and Bogutskaya (2009), but no Iranian record. The only specimen was lost (Reshetnikov et al., 1997). M. nonultimus may be a synonym (Pinchuk and Miller in Miller, 2004).

Mesogobius nonultimus
(Iljin, 1936)

Common names

[bychok neposledniy or seriy bychok-martovic in Russian; Caspian toad goby]

Systematics

Described from 24 miles southwest of Ulsky Bank, Caspian Sea, Turkmenistan. The type is lost. May be a subspecies of Mesogobius batrachocephalus (Pallas, 1814) although Miller revises his opinion in Pinchuk and Miller in Miller (2004) and recognises it as a distinct species. See also under M. nigronotatus above.

Key characters

It is distinguished by having three (rather than the more usual two) transverse head papillae rows between rows b and d. Nape scales weakly ctenoid, almost reaching median pore κ, snout less than twice eye diameter, lower jaw not protruding, and males with distinct breeding colours.

Morphology

First dorsal fin rays 6-7, second dorsal fin with 1 spine and 16-19 (usually 17-18) soft rays, and anal fin with 1 spine and 15-18 soft rays. Scales ctenoid except for a few, small cycloid scales on the opercle. The breast is naked. Lateral line scales 73-83. Lateral line system with 8 transverse suborbital neuromast rows. Three rows of free neuromasts on the opercle. The height of the second dorsal fin diminishes posteriorly. Head width 1.25 depth.

Meristics for an Iranian specimen: first dorsal fin with 6 spines, second dorsal fin 1 spine and 19 soft rays, anal fin with 17 soft rays and pectoral fin with 16 branched rays. Lateral series scales 80 and transverse scale rows 20.

Sexual dimorphism

Male spawning colour is distinctive and males are larger than females.

Colour

Overall colour is pale grey with dark bands, up to 6 on the body. The first band is in front of or below the first dorsal fin and is visible from above. Intensity of bands varies with ones below the middle of the second dorsal fin and before the caudal fin origin being the most intense. The first dorsal fin has two bands, the dorsal, pectoral and caudal fins are dark while the pelvic and anal fins are pale. Males have a dark, bluish-black spawning colouration with spots and bands on the upper body becoming indistinct.

Size

Reaches 17.4 cm.

Distribution

Probably the whole Caspian Sea basin. Known from the Turkmenistan and Daghestan coasts and from recorded between Kultuk and Astara in Azerbaijan (Ragimov, 1965). Pinchuk and Miller in Miller (2004) map this species from the entire Caspian Sea including the Iranian shore but only recently has it been recorded from Iran (H. Ahnelt, et al., 2007).

Zoogeography

This species is related to Neogobius and Ponticola, more speciose genera of Ponto-Caspian gobies.

Habitat

This species has been collected down to 50 m, perhaps deeper, but little else is known of its habits. It is suspected to feed, winter and spawn in deeper waters without an onshore migration.

Age and growth

Maturity is attained in the second year of life.

Food

Unknown.

Reproduction

Fecundity reaches 1544 eggs, probably laid in a single batch. The spawning season off the Lenkoran coast is late March and early April, the earliest of all Caspian Sea gobies.

Parasites and predators

Unknown.

Economic importance

None.

Conservation

Numbers in Iranian waters and threats to this species need to be assessed.

Further work

The biology and distribution of this species in Iranian waters needs to be studied.

Sources

Based in part on Pinchuk and Miller in Miller (2004).

Iranian material: CMNFI 2006-0022, 1, 114.2 mm standard length, Gilan, southwest Caspian Sea (38º00'N, 49º30'E to 38º20'N, 50º00'E) (Ahnelt et al., 2007).

Genus Neogobius
Iljin, 1927

This genus of gobies is found in the Black and Caspian seas where there are about 14 species, some large enough to be the object of commercial fisheries. The general Farsi name for fishes in this genus is گاوماهي (gav mahi), سگ ماهي (sag mahi) or gel-ye mahi, not repeated under each species.

Berg (1948-1949) and various papers by Pinchuk (1976, 1977, 1991) recognise Neogobius as a genus but Vasil'yeva (1989) considers that the separation of Neogobius, even as a subgenus, is doubtful and refers the species to Gobius. Simonovic (1999) briefly reviews the literature on Neogobius and its distinction from Gobius. Presence or absence of a swimbladder, egg size, and presence or absence of pelagic larvae are characters which have been used to recognise genera or subgenera but these may be phylogenetically independent (Svetovidov, 1964b). Various subgenera may be recognised within Gobius such as Gobius Linnaeus, 1758, Neogobius Iljin, 1927 (type species N. fluviatilis), Apollonia Iljin, 1927 (monotypic for N. melanostomus according to Pinchuk (1991) although Vasil'yeva (1989) and Dobrovolov et al. in Charlebois et al. (1997) add the species fluviatilis on the basis of osteology and electrophoretic studies; and then later she (Vasil'eva, 1996a) places both these species in the subgenus Neogobius, again on osteological grounds), Ponticola Iljin, 1927 including such species as ratan (type species), and Eichwaldia for caspius (type species)(Eichwaldia is preoccupied by Eichwaldia Billings 1858 in fossil brachiopods, replaced by and objective synonym of Eichwaldiella Whitley, 1930). Miller and Vasil'eva in Miller (2003 - Vasil'eva's assignments) give the following subgenera, with definitions (these subgenera may eventually be elevated to genus once a thorough analysis is complete (see below); note assignments conflict in part with the above:- Neogobius (with one species, caspius), Apollonia (fluviatilis and melanostomus), Babka (gymnotrachelus, not in the Caspian Sea), Ponticola (the remainder - including species not in the Caspian Sea). Miller in Miller (2004) place Neogobius bathybius in the genus Chasar and this is followed here. Neilsen and Stepien (2006) and Stepien and Tumeo (2006) used mtDNA cytochrome b sequence data and consider Apollonia (N. melanostomus and N. fluviatilis) to be highly divergent from all other neogobiins, Mesogobius is the sister genus of Proterorhinus and a clade of these two genera is sister to Neogobius. Neilsen and Stepien (2009a) using two mitochondrial and two nuclear genes conclude that Neogobius, or in a restricted form Apollonia, consists of fluviatilis, melanostomus and caspius, Babka is monotypic (species gymnotrachelus), and Ponticola includes cyrius, gorlap, ratan and syrman among others. Mesogobius and Proterorhinus are sister groups. Tribes are also delineated. Apollonia is synonymised with Neogobius.

Pinchuk (1991) states that many of the characters used to define genera are composed for convenience in identification rather than indicating natural relationships, i.e. the characters are simply those used in keys. Ahnelt and Holčík (1996) call the taxonomy of this genus "variable, unstable and frequently uncertain" since variation between Black and Caspian Sea populations is usually very small and since there are migrating and resident marine and freshwater populations of the same species. Miller and Vasil'eva in Miller (2003) state that the genus is paraphyletic (included species do not share an autapomorphy) but retain it for convenience on morphological grounds, a practice followed here until a cladistic revision is performed.

These gobies are characterised by having an elongate body, compressed posteriorly, mouth of moderate size, 6 first dorsal fin rays, more than 10 rays in the second dorsal fin (large number of rays in the unpaired fins according to Pinchuk (1991) compared to Gobius sensu stricto), anterior nostrils tubular but not very elongate, posterior nostrils near eye, ctenoid scales of moderate size, a scaled nape and predorsal area, cheek naked, the pelvic fin anterior membrane may have obvious lateral lobes, no barbels, upper rays of pectoral fin may be free and filament-like (Pinchuk (1991) states that they are always within the fin membrane or joined), teeth conical and in a few series, back muscles extend forward almost to the eyes, tongue not notched or only slightly notched, no swimbladder in adults, 32-35 vertebrae modally (Gobius has only 28 and Proterorhinus 30-32 (Simonovič et al. (1996)), dorsal pterygiophore formula (3)22110, anal pterygiophores before first caudal haemal spine 2 or 3, large, oligoplasmatic roe, and no pelagic larvae. Miller and Vasil'eva in Miller (2003) give details of neuromast organs as follows:- infraorbital neuromasts typically in 7 transverse rows, 4 before and 3 above hyomandibular row b, and no row a. Row 7 is made up of a few to several papillae in a row descending postero-ventrally from just in front of the anterior oculoscapular pore α. Rows 5i and 6i are separated with 5i well behind the anterior end of hyomandibular row b and with 6i reaching or falling short of row b. Dorsal supraorbital rows o are separate in the dorsal midline. Hyomandibular row z ends near pore χ. Supratemporal accessory line x1 ending anteriorly behind pore β. Anterior oculoscapuler, posterior oculoscapular, and preopercular canals present. These have the pores σ,λ,κ,ω,α,β and ρ; θ and τ; and χ, δ and ε. Anterior oculoscapular pore ρ and posterior oculoscapular pore θ well separated.

Neogobius is distinguished from the closely related genus Mesogobius Bleeker, 1874 by having a smaller number of infraorbital transverse rows of pit organs and the transverse parietal posterior rows of pit organs are separated by a wide interspace (Pinchuk, 1991).

Simonovič et al. (1996) consider this genus of gobies to be a young one as evidenced by the distribution of species in the Caspian and Aral seas which closed recently. Simonovic (1999) concludes that "Neogobius" (and Proterorhinus) evolved from a common ancestor in the Dacian Basin of the eastern Paratethys (now the Black Sea) or its tributaries during the early to mid-Pliocene, as evidenced by their absence from Adriatic, Ionian and Aegean basins. Divergence probably occurred in freshwater, riverine habitats as development is direct without a pelagic larval stage as found in marine gobiids. The hypersalinity of the isolated Black Sea in the late Miocene to early Pliocene was the causal event that led to rapid speciation of "Neogobius" and the genus retained direct development when it returned to the Black and Caspian seas. Dillon and Stepien (2001) using mitochondrial DNA suggest that the genera "Neogobius" and Proterorhinus diverged about 5.2±1.0 MYA in the late Miocene/early Pliocene from a common ancestor shared with Gobius during the isolation of the Paratethys Basin from the Mediterranean Tethys Sea. This contradicts suggestions that the neogobiins diverged from Gobius during the Quaternary interglacial connections of the Black and Mediterranean seas.

Masoumian and Aghl Mandi (2009) record Neogobius spp (presumably including Ponticola spp.) infected with Ceratomyxa caspia, a myxosporean, from the Iranian shore of the Caspian Sea.

Neogobius caspius
(Eichwald, 1831)

Common names

gel-ye mahi Khazari (= Caspian goby), gavmahi-ye Khazari, gavmahi-ye Daryaye-e Khazar.

[xval xulu in Azerbaijan; Khvalynskii bychok or Hyrcanian goby in Russian; Caspian goby].

Systematics

No major synonyms and no types known. Described in Latin from "Hab. in Caspio mari, in sinu Bacuensi" (in the Caspian Sea, in Baku Bay, Azerbaijan).

Key characters

This species is separated from other Caspian gobies in Iran by the rear nostril being slit-like and far from the eye margin. It has a comparatively small mouth.

Morphology

Head canals and free neuromasts on the body are typical of a Neogobius. First dorsal fin spines 5-7, usually 6, second dorsal fin with 1 spine followed by 14-17 soft rays. The first ray of the second dorsal fin may be elongated and branched at its tip and in such cases was counted as a "spine". Anal fin with 1-2 spines, usually 1, and 10-14 soft rays. Lateral series scales 57-71. Predorsal area and nape scaled, anteriorly reaching to the orbit (postorbital section of the supraorbital canal of the cephalic lateral line system). Anteriormost scales cycloid. Scales have a diamond-shaped pattern on the flank. Individual scales are squarish with the upper and lower margins straight to slightly rounded or, more rarely, each scale has an overall rounded shape. The posterior margin has two straight edges meeting at the posterior centre. The posterior margin bears ctenii. The anterior margin is wavy where the radii terminate at indentations and there is a central protuberance. Moderately numerous radii radiate anteriorly from the central point of the posterior margin, occupying the entire anterior field and spilling into the lateral fields. Circuli are fine. There is a dermal papilla on the posterior edge of the gill chamber, just under the gill cover. The gill rakers are short anteriorly and posteriorly but vary in length and the longer central rakers reach the second adjacent raker when appressed. Rakers are commonly forked or even antler-like. Dorsally the smallest rakers are difficult to distinguish from mouth cavity tubercles and only definite rakers were included in the count. The pelvic fin anterior membrane has pointed lobes. The gut is an elongate s-shape with convolutions or twists along its length. The genital papilla is bifurcated at the tip.

Meristics for Iranian specimens: first dorsal fin spines 5(1), 6(37) or 7(2); second dorsal fin soft rays 14(2), 15(10), 16(27) or 17(1); anal fin soft rays 11(2), 12(25) or 13(13); pectoral fin rays 17(6), 18(29) or 19(5); lateral series scales 60(2), 61(4), 62(6), 63(8), 64(10), 65(6), 66(1), 67(1), 68(1) or 70(1); total gill rakers 8(1), 9(3), 10(17), 11(14), 12(4) or 13(1); and total vertebrae ?.

Sexual dimorphism

Colour differences are given below.

Colour

Overall colour is a dark grey or grey-brown with a clearly contrasting white to pearly white belly and lower head surface. The back may have 7-8 dark brown or grey saddles. The flanks have 6-10 obscure dark spots. The first dorsal fin has 2-4 dark brown stripes and a black spot over the last three rays. The black spot may not be very evident and merges with the general background pigmentation. There are no clearly defined spots on the fins although the rays are brown in all but the pelvic and anal fins. The anal and pelvic fins are pearly-white. The caudal fin has a narrow whitish transparent edge. The iris has a narrow golden ring. The genital papilla is white or grey. The peritoneum is silvery-brown in preserved specimens.

Breeding males are black all over, often including the margin of the first dorsal fin. First dorsal fin ray tips may be yellowish. The belly is whitish to pale grey in front of the pelvic fins. There is a dark spot at the end of the first dorsal fin. The second dorsal fin has a narrow colourless margin but fin tips are not extended into free ends as obviously as in some Neogobius species. All fins have a fringe effect although this is not well developed in the first dorsal fin. The caudal fin has a colourless or whitish margin. The dorsal fin spot is lost temporarily after spawning.

Size

Reaches 20.2 cm.

Distribution

This species is found in the Caspian Sea and is recorded between Kultuk and Astara in Azerbaijan (Ragimov, 1965). In Iran records are from both the southwestern, south-central and southeastern Caspian Sea, and Gorgan Bay (Abdoli and Naderi, 2009).

Zoogeography

Endemic to the Caspian Sea.

Habitat

Found throughout the Caspian Sea but does not enter fresh water. It is found on sand or pebble bottoms, less often on stones and rarely on soft mud. They tend to remain inshore until the winter when temperatures fall although high summer temperatures may drive them into deeper and cooler water for a short period (Pinchuk and Miller in Miller, 2003).

Age and growth

Sarpanah Sarkohi et al. (2010) found fish to reach age 5 years in the Iranian Caspian Sea sample sites listed below.

Food

Gut contents include polychaetes, crustaceans, small bivalve molluscs and fish. In some areas molluscs predominate, in others crustaceans. The study by Sarpanah Sarkohi et al. (2010) of fish taken in bottom trawlers from three stations at Astara, Anzali and Chaboksar found a relative gut length of less than one suggesting this fish is carnivorous. Intensity of fullness of guts was highest from April to October. The main food items were molluscs (principally Cardium, Didacna, Mytilaster, Abra), worms (principally Nereis, oligochaetes) and crustaceans (principally Niphargoides, Petrcuma (sic), Paramysis) with food preferences of 100%, 89% and 74% respectively. Some chironomids and gobiids were also eaten.

Reproduction

Egg sizes in Iranian specimens reach 2.1 mm and fish taken on 14 May have eggs 1.9 mm in diameter suggesting a spring to summer spawning season. This species approaches coastal areas in the southern Caspian Sea at the end of March and the beginning of April, later than P. syrman and P. gorlap. Females may have up to 1007 large eggs and up to 1985 small eggs, indicating repeat spawning (Pinchuk and Miller in Miller, 2003). Sarpanah Sourkouhi et al. (2008) examined fish form the Gilan coast and found the highest gonadosomatic indices in June, suggested spawning occurs several times, found a sex ratio of 1.47:1 in favour of females, an absolute fecundity of 212-1234 eggs, a relative fecundity of 18.73-29.89 eggs/g body weight and an egg diameter range of 0.06-0.23 mm.

Parasites and predators

Sattari et al. (2002) and Sattari (2004) records the presence of the nematode, Eustrongylides excisus, in this species. This parasite can damage muscles in commercial species and render them unsuitable for sale. This species is eaten by the Caspian seal and by sturgeons. Daghigh Roohi and Sattari (2004) record Eustrongylides excisus, and Dichelyne minutus from this species in the southwestern Caspian Sea of Iran. Sattari et al. (2004; 2005) surveyed this species in the inshore area of the Caspian Sea, recording Eustrongyloides excisus and Dichelyne minutus.

Economic importance

Not reported as of economic importance in Iran, it has been caught commercially in the former Soviet Union and on rod and line. It does form part of the diet of sturgeons (Pinchuk and Miller in Miller, 2003).

Conservation

Trends in numbers of individuals of this species and threats to it have not been examined so its conservation status is unknown.

Further work

The biology of this species needs investigation.

Sources

Pinchuk and Miller in Miller (2003) is the most recent summary on this species.

Iranian material: CMNFI 2006-0022, 1,124.8 mm standard length, Gilan, south-west Caspian Sea off Astara (38°00’N, 49°30’E to 38°20’N, 50°00'E); CMNFI 2006-0027, 2, 113.4-121.1 mm standard length, Gilan, Talesh (37°48’N, 48°55’E); NMW 80605, 1, 116.1 mm standard length, Mazandaran, Now Shahr beach (36°39’N, 51°31’E); NMW 95073, 1, 92.8 mm standard length, Mazandaran, Gomishan lagoon near estuary of Gorgan River (36°59’N, 54°00’E).

Neogobius melanostomus
(Pallas, 1814)

Neogobius melanostomus from Wikimedia Commons.

Neogobius melanostomus from Wikimedia Commons.

Common names

gavmahi-ye gerd (= round goby), gel-ye mahi gerd (= round goby), gavmahi domgerd, gavmahi-e-domguerd, sebele.

[xazar kirda xulu in Azerbaijan; Kaspiiskii bychok-kruglyak or Caspian round goby in Russian; black spotted goby].

Systematics

Gobius melanostomus was originally described from Sevastopol and Balaklava, Ukraine, on the Black Sea. No types known.

Gobius affinis Eichwald, 1831 described in Latin from "Hab. in Caspii maris sinu bacuensi, balchanensi" (Baku and Balkhan bays in the Caspian Sea, no types known) and Gobius sulcatus Eichwald, 1831 described in Latin from "Hab. in Caspii maris sinu balchanensi" (Balkhan Bay in the Caspian Sea, no types known), are synonyms. The Caspian Sea population was referred to Neogobius melanostomus affinis (Eichwald, 1831) but subspecies are no longer recognised (Iljin, 1956; Pinchuk, 1976). However, Brown and Stepien (2008) conclude on DNA evidence that Black and Caspian Sea populations diverged ca. 350,000 years ago and this supports subspecies separation.

Key characters

Pelvic fin anterior membrane with rounded and shallow lateral lobes; lobes not more than one-sixth width of anterior edge of membrane, or lacking entirely; nape scaled and at least anterior nape scales cycloid; first dorsal fin with large dark spot at rear; and lateral series scales usually 49-55.

Morphology

This species is separated from other Caspian gobies in Iran by having a small mouth, rounded, shallow, poorly-expressed lobes to the pelvic anterior membrane, scales in lateral series 41-59 (usually 47-55), generally at the low end of this range, mid- and anterior nape scales are cycloid as are most of those on the gill cover, throat, pectoral in bases and part of the abdomen, the pelvic disc is 0.6-0.8 pelvic base to anal fin length, the jaw angle is below the level of the anterior part of the eye, the posterior nostril is close to the eye margin, and the first dorsal fin has an obvious dark spot posteriorly.

First dorsal fin spines 5-8, usually 6, second dorsal fin with 1 spine and 11-18 soft rays, anal fin with 1 spine and 9-14 soft rays, and pectoral fin with 16-20 branched rays. Scales appear as diamond-shaped on the flank since the posterior margin has two straight edges coming to a central point. However scales can be quite rounded even adjacent to those forming a diamond pattern. The posterior margin bears ctenii. Dorsal and ventral margins are rounded and the anterior margin is indented above and below a central protuberance. Radii radiate anteriorly from the centre of the posterior margin. Circuli are fine. Gill rakers 9-13, presumably on lower arch only, see below. Gill rakers may reach the second adjacent raker when appressed and although usually regular can be branched and variable in size along the arch. Vertebrae 31-34.

Head depth at the eyes is 0.9-1.2 head width between the upper origins of the opercles. The interorbital width is 0.8 to slightly less than eye diameter. The snout is longer than the eye, 1.1-1.4 times. The upper lip narrow slightly rearward and is about half the lateral preorbital area (between lip and eye). Caudal peduncle depth is two-thirds its length. The tongue is truncate or slightly notched. The posterior teeth on the dentary bone are smaller than the anterior teeth and all are blunt. Pharyngeal teeth are molariform, adapted to crushing molluscs. The anal papilla is bifurcated at the tip. The gut is short and roughly s-shaped with some convolutions, particularly on the posterior section.

The chromosome number for "N. m. affinis" is 2n=46 with 46 chromosomal arms (Ráb, 1985; Vasil'ev and Grigoryan, 1993; Simonovic, 1999).

Meristics for Iranian fish:- first dorsal fin spines 5(6) or 6(34); second dorsal fin soft rays 12(1), 13(2), 14(18), 15(18) or 16(1); anal fin soft rays 10(3), 11(17), 12(19) or 13(1); total pectoral fin rays 17(11), 18(28) or 19(1); lateral series scales 48(1), 49(3), 50(8), 51(4), 52(8), 53(4), 54(5), 55(4), 56(2) or 57(1); total gill rakers 11(5), 12(15), 13(11), 14(7) or 15(2); total vertebrae ?.

Sexual dimorphism

Females are smaller than the male. Mature males have larger dorsal, pectoral and anal fins, swollen cheeks, and colour is different (see below).

Colour

Overall colour is yellowish-grey or brownish-grey (fawn) with 8-9 large, dark brown, blue-grey to black flank blotches. There are 4 saddles across the back. The first dorsal fin has a black blotch posteriorly over the last two rays (and sometimes on the interradial membrane between rays 4 and 5), and the blotch or spot may have a white border in juveniles. The spot can be absent as reported by Jude (1997) for fish introduced to the North American Lake Erie. The sides of the head have oblique, curved bands or a reticulate pattern of a rusty to gingery tint on a bluish-grey background. The pattern is darker on the opercle and snout. The first dorsal fin may be tinged green and has 3 rusty-brown bands and a narrow margin may be a rusty or yellowish colour. The second dorsal and caudal fins may have a light margin even in non-spawning individuals. The second dorsal fin has 3-4 rows of small rusty spots. The caudal fin also bears rusty spots anteriorly but not in very definite rows. The anal fin is grey to dark brown. The pectoral fin lacks spots and is almost colourless but may be tinged with yellow. The pelvic disc is grey with a rusty colour in the centre. The genital papilla is grey or unpigmented. The iris is yellow to orange. The peritoneum is a silvery brown.

Colour varies with habitat. In bays with algal growth fish are darker, blue-grey tints are lost, there are more yellowish-brown colours and there is an increase in number and size of dark-brown spots. Others may have an olive-green tint.

When spawning, males are charcoal black overall, sometimes with a bluish tint, and with indistinct yellow spots. The dorsal, caudal and anal fins are white-edged or yellow, although the first dorsal margin can be black and there may be practically no margin to the anal fin. The black spot on the first dorsal fin can still be seen against the light. There are no free tips to fin rays according to literature reports but Iranian specimens have all fin tips separated to form a fringe.

The young are a slate grey overall with a mottled and blotched flank. The mid-flank blotches may form a series of about 6. There is a spot at the upper base of the pectoral fin. The dorsal fin spot is centred on the membrane between the last two rays but spills over onto the adjacent membranes, particularly the posterior one. Fins have pigment on the rays and membranes with no clear pattern. The dorsal fins are darker than the anal which is darker than the pectoral or pelvic. There is a bar from the lower eye margin to the upper lip just anterior to the mouth corner.

Size

Reaches 29.0 cm total length. Freshwater fish are smaller than marine ones.

Distribution

Found from the Aegean to the Black and Caspian seas. Introduced to the Aral Sea in the 1950s but increasing salinity in the 1980s eliminated the population. Now in the Baltic Sea via rivers and canals or in ship ballast. Also present by introduction in the Great Lakes of North America where first reported in 1990, apparently from several sources (Brown and Stepien, 2006).

Reported from a wide range of rivers along the Caspian coast of Iran including the Gorgan, Tonekabon, Safid, Dyusan, Shimrud, Chemkhaleh, Rud-e Sere, Kargan, Navarus, Chalus, Babol and Haraz rivers, the Anzali Mordab, Gorgan Bay, the southeast Caspian Sea, southwest Caspian Sea and south-central Caspian Sea (Holčík and Oláh, 1992; Abbasi et al., 1999; Kiabi et al., 1999; Abdoli, 2000; Jolodar and Abdoli, 2004; Abdoli and Naderi, 2009). Apparently absent from the Atrak River in Iran and of uncertain presence in the Aras River (Abdoli and Naderi, 2009).

Zoogeography

See genus account above.

Habitat

This species is found in inshore waters at depths to about 20 m, sometimes to 70 m or even 200 m in deeper sea areas in winter, on rock, gravel, shell, sand or silt bottoms. Aquarium specimens are seen to hide in crevices and under rocks. In the Caspian Sea, it may be found among eelgrass, Zostera. It also enters rivers. Larvae live near the bottom. The species prefers littoral areas where wave action keeps oxygen levels high and decaying material is reduced. It leads a sedentary life (Moskal'kova, 1996). It may be found from fresh waters to salinities of 40.5‰ from -1 to 32°C, and respiring through the skin can tolerate low oxygen conditions from 0.3-0.9 ml/l (Moskal'kova, 1996; Jude, 1997).

In the North American Great Lakes basin, where it is an exotic, this species prefers rocky habitats and is more active during the day than the night. The mean density in one river was approximately 7 gobies/sq m and can reach as high as 90 gobies/sq m since they aggregate. This goby has a high site fidelity in mark-recapture studies (Ray, 1997; Ray and Corkum, 1997a).

The gobies can tolerate a flow of 0.34 m/s for 3-4 minutes but at higher levels they retreat to the bottom and brace themselves against the current using the pectoral fins.

The onshore spawning migration in the southern Caspian Sea occurs in spring, later than P. syrman and P. gorlap. A spawning area is chosen on the basis of stones being present rather than depth. Larger males approach the shore first. There is an offshore movement as winter begins and in the hottest part of summer.

Age and growth

Maturity is attained in the second year by females and in the third year by males but may be as early as 1 year. Life span can exceed 5 years for females. Males die after spawning. Faster growth results in a shorter life span. At the end of the first year of life they can reach 5.5-6.0 cm (Moskal'kova, 1996). In North America, populations of this exotic are dominated by age one fish (MacInnis and Corkum, 1997a). Growth is slower in North America and maximum size is less than in its native habitat, probably because temperatures are lower than in Europe (Jude, 1997). Abdoli et al. (2006) examined fish from the Gomishan and Miankaleh wetlands and found a length (L)-weight (W) relationship of W = 0.0091L3.17 for males, 0.0249L2.7 for females and 0.0112L3.08 for sexes combined.

Food

Molluscs predominate with significant amounts of crustaceans in the north Caspian Sea but this can vary annually with crustaceans becoming predominant. They feed most heavily in the post-spawning period in July and August (Opalatenko, 1979). Other important foods are bivalves, such as zebra mussels, snails, polychaetes, chironomids, other aquatic insects, goby eggs and small fishes. Attached clams and mussels are bitten off the substrate by the large jaw teeth and crushed by the pharyngeal teeth (Moskal'ova, 1996). Molluscivory is also documented for Danube River populations (Simonovič et al., 2001) and for Baltic Sea populations (Skora and Rzeznik, 2001) and molluscs appear to be the preferred diet wherever this species is found. An exception may be the southern Caspian Sea where one study found crustaceans to predominate (Pinchuk et al. in Miller, 2003).

In the Caspian Sea, introduced species, such as crabs, the worm Nereis diversicolor and the mollusc Abra ovata, are also eaten (Kosarev and Yablonskaya, 1994). Iranian specimens contain plant fragments, mollusc shells, shrimps, fish remains, aquatic insects and polychaetes.

In North America, immature round gobies are the most successful predators on round goby eggs. Round gobies are said to be aggressive and will take a baited hook. However this goby may need to be stationary to detect prey. Nocturnal feeding occurs. The blunt teeth are indicative of a primarily mollusc diet.

Experimental studies in North America have shown this species to prefer individual and clumped zebra mussels to other foods such as sphaeriid clams. Feeding rates vary between 36 and 47 mussels (4.5-12.5 mm long) per day for fish 6-10 cm long, over 100 mussels per day when the mussels are smaller than 4 mm (Ghedotti et al., 1995). Mussels can remain in the mouth and pharynx for less than one hour to over 12 hours; shells are then split and ejected. However some are crushed and passed through the digestive tract. In the upper Detroit River, Canada, round gobies eat zebra mussels (58%), snails (6%) and other invertebrates (36%) such as aquatic insects, softshelled crayfish and zooplankton. In laboratory studies, round gobies ate an average of 1 g of mussels in 24 hours, smaller mussels being preferred (Ray and Corkum, 1997b). However, field studies in Lake Michigan show that round gobies are unlikely to remove zebra mussels (also a pest in North America) from a habitat (Djuricich and Janssen, 2001). These gobies are also predators on lake trout eggs under laboratory conditions in North America (Chotkowski and Marsden, 1999). They are also reported to eat sturgeon eggs in North America.

Reproduction

There is a movement inshore to waters about 0.2-1.5 m deep for spawning, males preceding females. Spawning takes place at the end of April to September in near shore areas of the north Caspian Sea and over a similar period in the Baltic Sea. Iranian specimens have well-developed testes on 10 April and spawning may occur earlier than in the North Caspian. There may be repeat spawning, up to 6 times every 18-20 days, as indicated by captive specimens. Peak spawning occurs at 15°C (range 10-30°C). Eggs are attached to or under rocks, in cavities, under logs or in such objects as cans and are aggressively guarded by the male. Early maturity, a very long spawning season and aggressive defense contribute to high egg, larvae and fry survival. The male prepares the nest site with a secretion from a cement gland which is coated over the site. As the female lays each egg, it is glued to the underside of the nest cavity roof in single layer rows. Egg clutches can be transported on the hulls of ships (Ahnelt et al., 1998). Fecundity reaches 6,177 eggs. Eggs are pale yellow, orange or pink, with red pigmentation. the eggs are ovoid with a sharp apex and measure 3.9 by 2.2 mm. Details of development are given by Moskal'kova (1996).

In North America, the species has been shown to be a multiple spawner with an extended reproductive season and male guarding of eggs (MacInnes, 1997). Here maturity occurs as early as year one and 43 mm standard length, a year earlier than in their natural habitat. Spawning extends from May to early August. Mean fecundity is 198 eggs. Each nest is utilised by large numbers of females since as many as an estimated 9462 eggs were found in each artificial nest site (MacInnis and Corkum, 1997b). Such a nest site can lose 50-70% of the eggs to predators while smaller clutches lose few eggs.

Jude (1997) notes that in North America some males do not defend a nest, the reason being unknown. This may well be due to such factors as availability of suitable spawning sites, large size or aggressiveness. He reports that after reaching a large size, males spawn once and then die, although females spawn up to six times through spring and summer, about every 20 days. Elsewhere males guard the nest from predators and fan the eggs to oxygenate them and to reduce siltation and fungal infection. Males will eat infected and unfertilised eggs. Males defend the nest by flaring the gills, spitting sand, darting at nearby intruders, biting, and growling for 1.0-1.5 seconds loud enough to be heard 10 m away. Nest sites in North America may be as deep as 7 m (Corkum and Wickett, 1998).

Males make several sounds including one attracting females to nest sites and another to intimidate other males. Females respond to male calls by a quieter sound (Jude, 1997).

Parasites and predators

Fil mahi (Huso huso), chalbash (Acipenser gueldenstaedtii), uzun burun (Acipenser stellatus), mash mahi (Aspius aspius), ordak mahi (Esox lucius), suf (Sander lucioperca) and seals are predators. Daghigh Roohi and Sattari (2004) record Dichelyne minutus and Corynosoma strumosum from this species in the southwestern Caspian Sea of Iran. Sattari et al. (2004; 2005) surveyed this species in the inshore area of the Caspian Sea, recording Dichelyne minutus. Sattari et al. (2007) report the nematode Eustrongylides excisus for this species in Iranian waters.

Economic importance

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in aquaria, as food and because it has been introduced outside its natural range. It is caught by anglers in some parts of the Caspian Sea basin.

It is present in the North American Great Lakes and is expanding (Ray and Corkum, 2001; Clapp et al., 2001; Jude et al., 1992; Jude, 2001). It is an introduction from the ballast waters of ships from multiple sources, perhaps including the Black Sea (Dillon and Stepien, 2001). Round gobies may displace native fish, compete for food and eat their eggs and young, and are the target of major efforts to study and contain them (Arrigoni and Berg, 2001; Charlebois et al., 2001; Diers et al., 2001; French and Jude, 2001; Janssen and Jude, 2001; Kaur et al., 2001; Schreier et al., 2001; Wolfe, 2001). They may also accumulate and pass on large concentrations of such chemicals as PCBs to sport fishes in North America (Jude, 1997). An extensive literature is now available on the Great Lakes populations.

It is an important food species in the Black Sea marketed fresh and formerly canned in tomato sauce. In the Sea of Azov in 1950-1960, gobies made up the bulk in local fisheries and round gobies comprised almost 90% of a catch exceeding 90,000 t (Moskal'kova, 1996). Commercial catches in the Caspian Sea have comprised up to 88.2% of goby catches but pollution has severely depleted stocks (Pinchuk et al., 2003).

Conservation

Kiabi et al. (1999) consider this species to be of least concern in the south Caspian Sea basin according to IUCN criteria. Criteria include medium numbers, habitat destruction, widespread range (75% of water bodies), absent in other water bodies in Iran, and absent outside the Caspian Sea basin (as N. m. affinis).

Further work

Biology in Iran has not been studied as well as elsewhere.

Sources

Charlebois et al. (1997) and Pinchuk et al. (2003) give general reviews of biology and are excerpted briefly above.

Iranian material:

Neogobius pallasi
(Berg, 1916)

Common names

gavmahi-ye tireh (= dark goby), gel-ye mahi rudkhanehai (= river goby), gavmahi-ye sheni, sebele.

[gumlur xulu in Azerbaijan; Kaspiiskii bychok-pesochnik or Caspian sand goby, rastrepka (= a person with untidy hair in reference to the fringed ends of the fin rays), chernysh (= darkie), khylak (when emaciated), all in Russian; Caspian monkey goby].

Systematics

Gobius fluviatilis was originally described in part from near the mouths of rivers falling into the Black Sea and similarly the Caspian Sea. No types are known. Neogobius fluviatilis pallasi (Berg, 1916) was the subspecies of the Caspian Sea basin. Kottelat and Freyhof (2007) recognise pallasi as the Caspian Sea species and restrict fluviatilis to the Black Sea basin. The type locality is the Caspian Sea, mouths of the Ural and Volga and syntypes are in the Zoological Institute, St. Petersburg under ZISP 2195, 2204, 23137, 30729, 30736, 30738, 30919, 30920, 30924-26. 33182, 34276, 34277 (Eschmeyer's "Catalog of Fishes", downloaded 23 August 2007).

Abd Elahpour Biria et al. (2009) examined fish from 4 localities on the Gilan coast using morphometric characters and found 3 different populations in Astara, Anzali and Chamkhaleh and Chaboksar.

Key characters

Pelvic fin anterior membrane with rounded and shallow lateral lobes. less than one-fifth rear edge width; pelvic fin disc 0.9 to more than distance to anal fin origin; nape with ctenoid scales; first dorsal fin without large dark spot (except in juveniles); lateral series scales usually 55-70; pelvic fin membrane anterior lobes small, rounded; angle of jaw anterior to orbit; and two transverse infraorbital papillae rows below longitudinal hyomandibular row b.

Morphology

The head width (between upper origins of opercles) is about equal to head depth (at eyes). The interorbital distance is not more than 0.75 eye diameter. The top of the head and the occiput are scaled with ctenoid scales. The upper lip is expanded towards the mouth corners, sometimes narrowing at the very end. The angle of the jaws reaches back at a level between the eye and the posterior nostril. The snout is longer than the eye diameter (1.6 times). The upper lip is not swollen at the angle, and measures 0.4-0.6 lateral preorbital area (between lip and eye). The first dorsal fin is high with an acute anterior profile, the first branched ray about twice as long as the penultimate ray. The pelvic fin has small, rounded lateral lobes (< 0.2 rear edge width) and almost reaches the anal fin (0.9 distance) or extends beyond the anal fin origin. The caudal peduncle depth is 0.5-0.6 length.

First dorsal fin spines 5-7, usually 6, second dorsal fin with 1 spine followed by 11-18, usually 15-17, soft rays, anal fin with 1 spine and 11-17 soft rays, usually 13-15, and pectoral fin rays 16-19. Lateral line scales 49-69, mostly 52-65. Gill rakers 6-10. Vertebrae 31-35. The chromosome number is 2n=46, with 46 acrocentric chromosomes and 46 chromosomal arms (Grigoryan and Vasil'ev, 1993; Klinkhardt et al., 1995; Simonovic, 1999).

N. pallasii is distinguished by the dorsal fin spot in young fish and an average number of lateral series scales being 53 as opposed to 62 for Black Sea fish.

Sexual dimorphism

See under colour. Males become slimmer at spawning and their are morphometric differences between the sexes (Pinchuk et al. in Miller, 2003).

Colour

Overall colour is pale and sandy, in cantrast to other gobies. On the flanks and over the back there is a series of 8-9 brownish elongate spots or blotches. The belly, lower head and genital papilla are pearly white. A dark bar runs from the eye antero-ventrally to the lip. The first dorsal fin has a dark spot posteriorly, although it may be absent in some adults, and is characteristic of the former subspecies pallasi, and found in small fish in the southern Caspian, persisting in adults in the northern Caspian (Pinchuk et al. in Miller, 2003). The first dorsal fin has 1-2 rows of grey to black spots and the second dorsal fin has 1-3 rows of pale grey spots. The caudal, anal, pectoral and pelvic fins are mostly grey on the rays and immaculate.

Breeding males become black or blue-black with the dorsal, caudal and anal fins bearing a yellow-orange margin. Some fish have a very narrow colourless margin to the pectoral and pelvic fins. The second dorsal fin may only be yellowish on its first ray, the remainder being white and the caudal margin may be colourless. Rays in the second dorsal fin of breeding males become extended into free tips.

Size

Reaches 24 cm total length although in the Caspian the largest fish are 16.0 cm males (Pinchuk et al. in Miller, 2003).

Distribution

Found in the Caspian Sea and its basin. In Iran, it is found in rivers along the Caspian coast including some upper reaches, the Aras River middle reaches and its tributary the Qareh Chai, the Anzali Mordab, Gorgan Bay, at Bandar Gaz, in the Gorgan, Madarso (or Madar Su), Tajan, Babol, Haraz, Sardab, Tonekabon, Pol-e Rud and Safid rivers (including their middle to upper reaches such as the lower Qezel Owzan and Shahrud of the Safid River basin), the southeast Caspian Sea, the southwest Caspian Sea and the south-central Caspian Sea (Shamsi et al., 1997; Abbasi et al., 1999; Kiabi et al., 1994; 1999; Abdoli, 2000; Abdoli and Naderi, 2009). Note that Abdoli (2000) records it from the upper Atrak River in contrast to earlier authors.

Zoogeography

The southern Caspian Sea and its tributary rivers is inhabited by a distinct subspecies, presumably on account of isolation from populations in the Black Sea. See genus account above.

Habitat

This species is found in inshore brackish waters and enters rivers. Salinities up to 46.9‰ are tolerated in marine bays. It is found down to 25 m although depths of 0.5-10 m are preferred, shallower in spring (0.5-5.0 m), slightly deeper in summer and deeper still in fall (5-10 m)(Pinchuk et al. in Miller, 2003). It is found over sand, mud and sandy-shelly bottoms. In the Madarsoo stream of Golestan National Park, abundance ranged from 0.1 to 0.75 fish per sq m and biomass from 0.21 to 4.88 g/m2 (Abdoli et al., 2002). Floods in this stream reduced the population, presumably through destruction of the substratum and its food supply. Smaller fish were caught in riffles while larger fish lived close to to the river bank in depths up to 0.5 m. Abdoli and Rahmani (2001) found this species to represent 95% of the goby catch in the Madarsoo stream, with N. melanostomus occupying the remainder.

Age and growth

Sexual maturity is attained at 2 years of age (9-12 cm for males, 8.5-10.5 cm for females) but may be as early as 1+ year (5.5 cm), particularly in the Caspian. Spawning males near Bandar-e Anzali were 12.4-15.2 cm long. Life span is 6 years but perhaps only 2+ years in the Caspian (Pinchuk et al. in Miller, 2003). Abdoli et al. (2002) found a sex ratio of 1 male to 3 females in the Madarsoo stream. Patimar et al. (2008) reported 5 age groups in fish from the Zarrin-Gol River in the eastern Alborz Mountains. Maximum total lengths were 12.1 cm in males and 10.8 cm in females, both 4+ years old. Fish aged 2+ and 3+ years were the most frequent in the population. sex ratio was equal and growth was negative allometric. Condition factor was highest in early September. Males showed a higher growth rate than females after age 1+ years. Abdoli et al. (2006) examined fish from the Gomishan and Miankaleh wetlands and found a length (L)-weight (W) relationship of W = 0.0059L3.291 for males, 0.0128L2.92 for females and 0.0057L3.29 for sexes combined.

Food

Diet items include crustaceans with lesser amounts of molluscs, polychaetes, chironomids and small gobies in the north Caspian Sea. They feed most heavily in the post-spawning period in July and August (Opalatenko, 1979). The quantity of amphipods eaten increases with age while the amount of mysids decreases. Introduced species, such as crabs, the worm Nereis and the mollusc Abra, are also eaten (Kosarev and Yablonskaya, 1994). In the Madarsoo stream of Golestan National Park in Iran, Ephemeroptera and Chironomidae are the most important food items but also included Trichoptera, other aquatic and terrestrial insects, crustaceans (including freshwater crabs), arachnids, worms and snails (Abdoli and Rahmani, 1999; Abdoli and Rahmani, 2001; Abdoli et al., 2002). Alavi Yeganeh and Kalbasi (2006) found fish in shallow water at Noor fed on amphipods, fish larvae, barnacles, palaemonid shrimps, mytilid bivalves, xanthid crabs, cardid bivalves, fish eggs and oligochaete worms, in order of preference. Diet varied with season, males feeding less during spawning for example Patimar et al. (2008) found the most frequent food items of fish in the Zarrin-Gol River were trichopterans, molluscs, chironomids and gammarids.

Reproduction

Spawning can take place from late April to September, varying with locality. Peak spawning in the Volga River delta is in May. Spawning is recorded at 18-26ºC, 20-28‰ and 0.5-1.3 m (Pinchuk et al. in Miller, 2003). In the Madarsoo stream of Iran, spawning peaked in March with an absolute fecundity of up to 532 eggs (Abdoli et al., 2002). Fecundity reaches 1025 eggs in the southern Caspian (perhaps 12,800 eggs elsewhere) and egg diameter 2.1 mm. Yellow-amber or pale pink eggs are laid on or under stones and plants. Nests are excavated by the male under natural or artificial objects. The male defends the nest and larger males are more successful. However, the stress associated with constant defense against predators and competing males leads to extreme emaciation. Nests contain more eggs than fecundity would indicate suggesting that more than one female lays eggs in a nest (Pinchuk et al. in Miller, 2003). Patimar et al. (2008) found the Zarrin-Gol River population to have an average absolute and relative fecundity of 508.47 eggs and 61.27 eggs/g. The reproductive season was March to April.

Parasites and predators

Pazooki and Aghlmandi (1998) found the nematode Dichelyne minutus infecting the intestine of 6.6% of fish examined from the Tajan River in Mazandaran. and Sattari et al. (2002) and Sattari (2004) records the presence of the nematode, Eustrongylides excisus, in this species. This parasite can damage muscles in commercial species and render them unsuitable for sale. Fil mahi (Huso huso), chalbash (Acipenser gueldenstaedtii) and suf (Sander lucioperca) are predators. Sattari et al. (2004; 2005) surveyed this species in the inshore area of the Caspian Sea, recording Eustrongyloides excisus and Dichelyne minutus. Daghigh Roohi and Sattari (2004) record Eustrongylides excisus, Dichelyne minutus and Corynosoma strumosum from this species in the southwestern Caspian Sea of Iran.

Economic importance

This is the most economically important goby in the north Caspian Sea according to Berg (1948-1949). Iljin (1956) considered this an error; it is the most numerous species but commercial fishing does not occur there. Pinchuk et al. in Miller (2003) state however that it comprised 54% of the entire goby catch, although only 7.7% in southern Dagestan. It is food for sturgeons and Sander lucioperca.

Conservation

Kiabi et al. (1999) consider this species to be of least concern in the south Caspian Sea basin according to IUCN criteria. Criteria include abundant in numbers, habitat destruction, widespread range (75% of water bodies), absent in other water bodies in Iran, and absent outside the Caspian Sea basin (as N. f. pallasi).

Further work

On of the better known gobies in Iran but could still use work on its biology.

Sources

Genus Periophthalmus
Bloch and Schneider, 1801

This mudskipper genus is found in the eastern Atlantic, Indian and western Pacific oceans. There are 12 species in the genus as yet undefined by a synapomorphy (Murdy, 1989). The pelvic fins are only partially united or may be totally separate. There are usually 12-14 pectoral fin rays, 4-17 first dorsal fin spines, and 8-13 second dorsal and anal fin rays. Scales are cycloid and cover the whole body and head except for the snout, isthmus and interorbital region in most species. Caninoid teeth are present in both jaws as a single row, with the anterior ones larger and pointed. The eyes are erectile and have a dermal cup covering the lower portion. There is a median fleshy ridge anterior to the eyes.

Mudskippers are found on muddy shores associated with mangroves but also venture into rivers. They spend much time out of the water.

Periophthalmus waltoni
Koumans, 1955

 

Common names

gel-cheragh (= mud-eater, mud-grazer), mahi-sag (= dog fish), gel khorak.

[shelambo, abou-shlembo or abu-shelamboo in Arabic; gullo in Pakistan; dark-blotched mudskipper, Walton's mudskipper, spotted mudskipper].

Systematics

This species was originally described from Iraq and Pakistan. The type locality is unclear as the information given by Koumans (1955) gives more than one locality, as follows:-

"Habitat.-Mesopotamia, Karachi.
I have seen 2♂ and 2♀. These were from the following localities:-
                 Indian Museum Collection.
Fao estuary of the Shatt-el-Arab, Mesopotamia, ♀        H. J. Walton.
Karachi ♂ and ♀                                                          W. D. Cumming.
Mouth of Shatt-el-Arab ♂                                             W. T. Blanford.".

The type locality could be Fao and the holotype is the female specimen caught by H. J. Walton and named for him, the remainder being paratypes - by implication. Alternatively and probably correctly, these are all syntypes as no holotype was formally designated. The date of publication of Koumans work is 1955, not 1941 as cited by Murdy (1989).

This species has been confused with Periophthalmus koelreuteri (Pallas, 1770), itself a synonym of Periophthalmus barbarus (Linnaeus, 1766) according to Murdy (1989). P. barbarus does not occur in the Persian Gulf region. P. waltoni has been placed in the synonymy of P. koelreuteri in works on Persian Gulf fishes, e.g. by Relyea (1981), but the revision by Murdy (1989) is followed here. Another mudskipper species listed in Coad (1991b), Periophthalmus weberi Eggert, 1935, should be deleted from the list as Murdy (1989) does not report it from the Persian Gulf.

Two syntypes of Periophthalmus waltoni are in the Rijksmuseum van Natuurlijke Historie, Leiden (RMNH 17004) and 4 syntypes are in the Zoological Survey of India, Calcutta (possibly lost) (Murdy, 1989). However, this is more fish than listed in Koumans (1955).

Key characters

The pelvic fins are totally united into a sucker with a well-developed basal membrane or frenum (absent in P. barbarus). The anal fin base and second dorsal fin base are 27% or less of standard length and there are no canine teeth internal to the lower jaw symphysis in contrast to Boleophthalmus, the other mudskipper of southern Iran.

Morphology

First dorsal fin spines 10-14, second dorsal fin with 1 spine and 12-14 soft rays. Anal fin with 10-13 soft rays. Pectoral fin rays 13-15. Scales in lateral series 90-121, predorsal rows 27-37. The upper jaw has 4-5 anterior teeth on each side much larger than lateral teeth. Upper jaw teeth 19-28, lower jaw teeth 17-23. Gill rakers 9-16, mean 11, on each arch, triangular in shape with 3-4 spines at the top of each raker (Barak et al., 1994). The first dorsal fin lacks elongate spines and is slightly rounded. The dorsal fins are not connected by a membrane. The gut shortens with growth in terms of body length indicative of a carnivorous habit (Barak et al., 1994).

Sexual dimorphism

Females have a lozenge-shaped first dorsal fin (i.e. anterior 6 spines equal in height followed by abruptly shortened ones) with a height less than body depth; in males this fin equals body depth.

Colour

The overall colour is a light grey-brown. The flank bears blotches which, below the soft dorsal fin, take on the form of 8 short bars and there is a scattering of small silvery or white dots. Ventrally the spots form roughly transverse silvery stripes. The head top is also blotched and the sides bear a few small white dots. There are 7 irregular black blotches along the back. The first dorsal fin is dusky brown to light grey and its margin is yellowish between spines 1 to 8 and dark just below this except for the posteriormost 3-4 spines. There is a series of light spots on each interray membrane on the basal half of the fin. The second dorsal fin is dusky brown, with a pale margin, followed by a black stripe, and then with a white spot at the middle of each membrane thus forming a horizontal dotted line along the mid-fin. The caudal fin is a dusky brown, darker basally. There are 3 black blotches on the caudal peduncle. The pectoral fins are a light dusky brown on the rays with the membranes on the upper half pale with dark, brownish streaks in mid-fin, and a whitish margin is present. The pelvic fins are blackish with a reddish-brown or pale margin.

Size

Reaches 20.0 cm total length (Mhaisen and Al-Maliki, 1996).

Distribution

Found from the Persian Gulf to Pakistan. Reported from the Shatt al Arab, Iraq near Iranian Khuzestan by Khalaf (1961), Al-Nasiri and Hoda (1975b, 1976), Al-Hassan et al. (1989), and Hussain et al. (1989; 1997). Specimens believed to be this species have been reported from Iranian fresh waters but no specimens have been examined. Holly (1929b) records this species from the type locality of Barbus baschakirdi in the Bashakird Mountains (see Cyprinion watsoni). The Museum of Comparative Zoology, Harvard has a Periophthalmus species recorded from Manyuhi on the Shatt al Arab in irrigation ditches (MCZ 149602) but the identity of this specimen needs to be confirmed.

Abdoli (2000) maps this species from the Arvand River in the Tigris River basin, the lower Zohreh (= Hendijan), lower Helleh and Mand rivers in the Gulf basin, the Kul and Mehran rivers of the Hormuz basin, and the lower reaches of rivers of the Makran from the Jagin to the Bahu Kalat.

Zoogeography

This species has ready access to the sea and can colonise rivers along the Iranian coast.

Habitat

The habitat and behaviour of this species is similar to Boleophthalmus dussumieri (q.v.; Tytler and Vaughan (1983) review behaviour of this species in Kuwait, their Periophthalmus koelreuteri being this species according to Murdy (1989)). However it favours areas of mud which are drier on the upper and middle regions of the shore. The burrow is Y-shaped with two entrances and a depth of 30 cm in mud. Fish occupy several burrow systems within a 2-3 m home range (Clayton and Snowden, 2000). Young first appear on mud flats in April but stay in wet mud or shallow pools. They move onto drier areas by May after some growth (1.5-2.5 cm). There are up to 30 fish per 100 sq m on Kuwait mud flats. The activity range of body temperatures is 14-20°C in January and 26-34.5°C in June. Body temperatures do not exceed 35°C even when air temperatures exceed 40°C because of evaporative cooling. In tidal pools of a few centimetres depth in the Khawr az Zubayr, water temperatures ranged from 16 to 44ºC, pH from 7.6 to 8.7, dissolved oxygen from 5.2 to 8.8 mg/l and salinity from 10.2 to 20.2‰ (Al-Daham and Al-Noor, 2000). Holes from 50 to 80 cm depth, with two openings separated by not more than 50 cm, are constructed as refuge from extreme temperatures, high winds and predators. The mud is carried to the outside of the hole in the fish's mouth, and forms a dam-like structure around the shelter not exceeding 8 cm in height. The fish spends most of the ebb tide outside the shelter.

As well as being found on mudflats of the sea shore, this mudskipper is found in the Shatt al Arab and other rivers of the Persian Gulf.

Age and growth

Barak et al. (1994) studied this species in the Khawr az Zubayr, Iraq and found growth to be allometric with the regression coefficient significantly (p<0.05) greater than 3. (W = 1.74x10-4 L2.35 (r = 0.99)). Al-Daham and Al-Noor (2000) found the smallest mature males were 7.3 cm and females 7.4 cm in the Khawr az Zubayr with maturity at age one year.

Food

Fiddler crabs are stalked as the main prey of this mudskipper in Kuwait (Tytler and Vaughan, 1983) but shrimps and other invertebrates are also taken. High wind speeds and surface temperatures below 15ºC inhibited hunting. Hunts were spatially well separated because of the long time period before crabs returned to the surface after a hunt in their vicinity (Clayton and Snowden, 2000). On the intertidal mudflats of Iraq at the head of the  Persian Gulf, this mudskipper feeds predominately on fish when larger than 90 mm (frequency 78.9%, occurrence 80.0%) with shrimps (14.1%, 20.0%), insects (3.2%, 10.0%), copepods 2.1%, 5.0%) and Daphnia sp. (1.7%, 5.0%). Fish smaller than 80 mm had values much lower for fish (6.7%, 8.3%) and higher for copepods (75.1%, 83.3%) with Daphnia sp. at 13.3% and 16.6%, insects at 2.7% and 4.1% and shrimps at 2.2% and 4.1%. Barak et al. (1994) found this species to be cannibalistic and piscivorous in the Khawr az Zubayr based on gill raker gap, gut-length/body length ratios decreasing with size, and stomach contents. Copepods were the major food item of smaller mudskippers (<70 mm) while mudskippers larger than this took fish. Mhaisen and Al-Maliki (1996) report food items as percentage of occurrence from fish caught in the Khawr az Zubayr estuary of Iraq to be 67.2% crustaceans (shrimps, crabs, barnacles and others), 34.5% snails, 3.4% insects and 1.7% fishes. Some fish consume plant material in winter months only, presumably in the absence of animal food.

Reproduction

Spawning in the marine environment of the Khawr az Zubayr takes place in March, the release of larvae being timed to take advantage of phytoplankton and zooplankton blooms (Hussain and Ahmed, 1999). Al-Daham and Al-Noor (2000) give the spawning period as March to April in the Khawr az Zubayr with a fecundity of 11,560-35,005 eggs for fish 7.4-12.9 cm long. A water temperature over 18ºC was the primary factor for spawning stimulation.

Parasites and predators

Mhaisen and Al-Maliki (1996) give information for Iraqi fish. E. Kahrom in the Crocodile Specialist Group Newsletter reports that muggers (marsh crocodile, Crocodylus palustris) eats a Periophthalmus in the Sarbaz River of Iranian Baluchestan (www.flmnh.ufl.edu/natsci/herpetology/newsletter/news173b.htm, downloaded 15 February 2002). However this may well be Boleophthalmus dussumieri.

Economic importance

None.

Conservation

No study of the numbers of this species in Iranian waters has been carried out so its status cannot be assessed.

Further work

The biology and population numbers of this species in Iran needs study.

Sources

The colour description is based on Kuronuma and Abe (1986).

Iranian material: None seen.

Comparative material: CMNFI 1985-0183, 2, 35.1-40.7 mm standard length, Iraq, Shatt al Arab (no other locality data); BM(NH) 1954.11.10:1-2, 2, 112.6-119.0 mm standard length, Iraq, Fao (29º58'N, 48º29'E); BM(NH) 1976.12.8:1-2, 76.1-84.1 mm standard length, Iraq, Basrah (30º30'N, 47"47'E); BM(NH) 1981.3.19:7-10, 4, 91.3-104.3 mm standard length, Kuwait mudflats (no other locality data).

Ponticola
Iljin, 1927

Ponticola cyrius
(Kessler, 1874)

Common names

gav mahi rud-e Kura (= Kura River goby) or gavmahi-ye Rud-e Kurosh.

[Kura goby, Kura River goby].

Systematics

Studies by Vasil'eva (1995a) and Vasil'yeva and Vasil'yev (1995) indicate that gobies from the upper and middle Kura River described as Gobius cyrius Kessler, 1874 and Gobius Weidemanni Kessler, 1874, and identified by various authors as Gobius platyrostris, Gobius constructor, Gobius platyrostris cyrius, Gobius cephalarges constructor, Neogobius cephalarges constructor and Neogobius platyrostris constructor should all be referred to Neogobius (now Ponticola) cyrius.

Gobius cyrius was described from the Kura River near Borzhom, Georgia and 3 syntypes are in the Zoological Institute, St. Petersburg under ZISP 2235 (Eschmeyer et al., 1996). One of these, a male 103 mm total length and 84 mm standard length, is designated as the lectotype by Vasil'yeva and Vasil'ev (1995). Two syntypes of Gobius Weidemanni described from Transcaucasia are under ZISP 2224.

Key characters

The pelvic fin anterior membrane has large, angular lateral lobes, at least one-fifth width of rear edge and pelvic fin less than nine-tenths distance to anal fin; the posterior nostril is near the edge of the orbit; lateral series scales usually 54-76; upper lip moderately swollen, width about 0.75 to more than length lateral preorbit; and interorbital distance 0.4-0.8 eye diameter.

Morphology

The head width is much greater than its depth. The interorbital distance is not more than half the eye diameter. The top of the head and the occiput are scaled with cycloid scales. The upper lip is expanded towards the mouth corners, sometimes narrowing at the very end. The lower jaw protrudes and the angle of the jaws reaches back below the anterior half of the eye. The snout is longer than the eye diameter (1.3-14 times). The upper lip is swollen at the angle. The second dorsal fin is of equal height along its length or rising slightly higher at its middle or end. The pelvic fin has a well-developed membrane with pointed lobes and does not extend to the anal opening. The caudal peduncle is compressed, depth about 0.30-0.35 length. The caudal fin is usually rounded but may be pointed, truncated or scalloped. The predorsal area is marbled and the first dorsal fin has a distal dark band.

First dorsal fin with 6-8, usually 6, spines, second dorsal fin with 1 spine and 15-20, usually 18-19, soft rays, and anal fin with 10-16, usually 14, soft rays (but see below), pectoral fin rays 17-19, and lateral line scales 53-78. Vertebrae 33-34. The chromosome number is 2n=36-42 (Klinkhardt et al., 1995).

Meristics in Iranian material: first dorsal fin spines 6(38), second dorsal fin with 15(7), 16(28) or 17(3) soft rays, anal fin with 10(1), 11(8), 12(26) or 13(3) soft rays, and pectoral fin rays 17(3), 18(28) or 19(7). Lateral line scales 54(2), 55(5), 56(7), 57(11), 58(6), or 59(4).

Diploid chromosome number 36-42, chromosome arms 46, polymorphism seen in a pair of subteleocentric chromosomes, the number of which varies from 0 to 2.

Sexual dimorphism

Males may not become black during the breeding season (Vasil'yeva and Vasil'yev, 1995). There are various differences in morphometric characters, summarised in Vasil'eva and Vasil'ev in Miller (2003).

Colour

The body is marked by a reticulate or marbled pattern in dark fawn, by scattered light spots and by 8-10 elongate dark spots along the lateral line. The back anterior to the dorsal fin is marbled. The belly is paler than the flank. The first dorsal fin is dark, particularly in the upper part where the darkening may form a band, with a transparent margin and a conspicuous black spot anteriorly below the tip of the fin. The first dorsal, caudal and anal fins have light margins but the fins are mostly dark. The light edge is obvious in the anterior parts of the first dorsal and anal fins. Only the second dorsal fin ray tips are light. The pectoral and pelvic fins are both uniformly dark.

Size

Attains 101.3 mm standard length, 13.0 cm total length.

Distribution

This species is found in the Caspian Sea basin including the Anzali Mordab and tributary rivers, the Massuleh River at Siahbar, and the Pasikhan River just above confluence with Pirbazar River and a tributary near Islamabad (Ahnelt and Holčík, 1996; Abdoli and Naderi, 2009) and the upper and Middle Kura River and upper Aras River of the former U.S.S.R. (Berg, 1948-1949; Vasil'eva and Vasil'ev in Miller, 2003).

Zoogeography

This species is distributed in rivers of the Caspian Sea basin in Azerbaijan and Iran while Neogobius constructor, with which it has been confused, is found in the Black Sea basin. The absence of P. cyrius from the lower Kura River could have been prevented by the prior penetration of P. gorlap according to Vasil'eva and Vasilev (1995).

Vasil'eva and Vasilev (1995; 1996a) consider that Sarmatian marine ancestors of modern P. cyrius and N. constructor most probably moved into fresh water during the Miocene, about 5 MYA. Mountain building in the Caucasus isolated the Kura and other rivers which once flowed to the Black Sea and their course became oriented to the Caspian Sea. Isolation of the gobies resulted in speciation.

Ahnelt and Holčík (1996) present an alternative view. They place the origin of this species in the Quaternary when fresh water from melting glaciers connected the Black and Caspian seas via the Manych Channel allowing freshwater fishes to disperse. This occurred four times during the Pleistocene, the last time about 14-12,000 years BP. A goby species was able to colonise both fresh and brackish water and particular stocks began to favour certain habitats. Those stocks which favoured headwaters evolved into P. cyrius while those which favoured lower regions and coastal areas evolved into P. gorlap. Ahnelt and Holčík (1996) point out that there is an analogous situation in the Black Sea with N. constructor and the species N. rhodioni Vasil'yeva and Vasil'yev, 1995. Additionally all west Caucasian and Caspian riverine gobies are very similar morphologically as evidenced by the difficulty in constructing reliable keys, suggestive of recent differentiation about 11,000-9,000 years BP after the isolation of the Black and Caspian seas.

Habitat

This small goby is found only in freshwater in mountain and foothill zones in contrast to P. gorlap. It is found in shallow (3 m or less) streams with a high gradient and current speed (1.6-4.9 m/sec on the bottom) in Iran (Ahnelt and Holčík, 1996). However, Vasil'yeva and Vasil'yev (1995) record it as absent in areas of strong current in the Kura River, preferring slow current and muddy bottoms. This type of habitat is limited in Iranian rivers which are much shorter and smaller than those of the Caucasus where an extensive lowland course is found. In Iran, the stream bed is composed of stones and boulders with some pebbles in the upper reaches and pebbles and gravel in the lower reaches. Small body size is an advantage for hiding under rocks to avoid predators and to escape the rapid current.

Age and growth

Life span is up to 3 years with maturity attained in the second or third year (Vasil'eva and Vasil'ev in Miller, 2003).

Food

Vasil'eva and Vasil'ev in Miller (2003) report feeding on insect larvae and parts of plants.

Reproduction

This species may contain eggs in three size groups, the largest being 2.81 mm in diameter, indicative of batch spawning. Total fecundity reached 1060 eggs. Spawning occurs from May to August at 11-20ºC, in some localities reported only for April-May (Vasil'eva and Vasil'ev in Miller, 2003).

Parasites and predators

Unknown.

Economic importance

None reported although in Azerbaijan some are caught by anglers.

Conservation

Further field work is necessary to assess numbers and status of this species.

Further work

The biology, distribution and conservation status of this species, recently recorded from Iran, remain to be examined in detail.

Sources

Counts and data are based on Ahnelt and Holčík (1996).

Ponticola goebelii
(Kessler, 1874)

Common names

gavmahi-ye ratan, gel-ye mahi ratan. ?name cahnge

[Kaspiiskii bychok-rotan or Caspian ratan or rotan goby in Russian].

Systematics

The Caspian Sea basin subspecies was usually referred to Neogobius ratan goebelii (Kessler, 1874). Gobius Goebelii Kessler, 1874 was described from Baku, Azerbaijan and Gobius ratan was originally described from Odessa on the Black Sea.

Gobius Bogdanowi Kessler, 1874 described from the Caspian Sea at Makhachkala (Petrovsk) is a synonym (the male of N. ratan goebeli according to Iljin (1956) although Pinchuk (1991) and Ragimov (1998a) consider it to be a subspecies of N. ratan). Pinchuk (1976) notes that fish from the Azerbaijan and Iranian coast differ from those from Dagestan and Turkmenistan, the first having fewer scales, but the number of specimens were insufficient to define taxa.

Gobius Goebelii has a syntype in the Naturhistorisches Museum Wien under NMW 33910 and others possibly in the Zoological Institute, St. Petersburg (ZISP 2229-30) (Eschmeyer et al., 1996; Pinchuk et al., 2003). Two probable syntypes of Gobius ratan are in the Muséum national d'Histoire naturelle, Paris under MNHN A.1125 and a syntype is in the Zoologisches Museum Berlin (Museum für Naturkunde, Universität Humboldt, Berlin) under ZMB 2098 (Bauchot et al., 1991). Two syntypes of Gobius Bogdanowi are possibly in the Zoological Institute, St. Petersburg (ZISP 10902) (Eschmeyer et al., 1996; Pinchuk et al., 2003).

Key characters

Pelvic fin anterior membrane with angular lateral lobes, not more than one-fifth width of rear edge; pelvic fin almost reaches the anal fin (0.9 distance) or extends beyond the anal fin origin; posterior nostril near edge of orbit; anterior nape with cycloid scales; rear of first dorsal fin without a dark spot; lateral series scales usually 49-54; and upper lip width 0.4-0.67 lateral preorbital width (lip to orbit).

Morphology

Pelvic fin anterior membrane with angular lateral lobes, rather than rounded; lobes small, not more than 0.2 width of rear edge; posterior nostril near edge of orbit; angle of jaw below pupil of eye; upper lip slightly swollen posteriorly, 0.5-0.67 lateral preorbital area (between lip and eye); first dorsal fin with sub-horizontal upper profile; rear of first dorsal fin without a dark spot; head depth at eyes slightly less to somewhat greater than width between upper origins of opercles; snout length about 0.9-1.2 orbit; lateral series scales usually 52-61; interorbital width 0.4-0.6 eye diameter; nape scales cycloid; pelvic fin almost reaches the anal fin (0.9 distance) or extends beyond the anal fin origin; the caudal peduncle depth almost equals its length, the posterior nostril is close to the eye margin, and colour dark brown with few spots (spawning males black). The two subspecies listed above are distinguished partly by scale counts, 63-66 in bogdanowi and 63 or less in goebeli.

First dorsal fin with 5-7, usually 6, spines, second dorsal fin with 1 spine and 15-19 soft rays, anal fin with 1 spine and 11-15 soft rays, and pectoral rays 17-21. Range in lateral series scales 49-70. Vertebrae 32-34.

Sexual dimorphism

Male colouration is distinctive and they have a higher dorsal fin.

Colour

Overall colour is dark brown with a few, discrete, rounded pale spots and, along the mid-flank, a series of small horizontal bars. Pale saddles cross the back. The first dorsal fin has a yellow or intense orange margin above a black bar (white in preserved fish). This fin also has a dark blotch, often with a bluish tint, on the first and second and second and third interradial membranes, and sometime son the third and fourth interradial membrane. The second dorsal fin is greyish with 3-4 rows of brown spots proximally, and it may have a pale margin. The pectoral and caudal fins have rows of rusty-brown spots proximally. The pelvic fin disc and the anal fin are grey with a broad pale margin. The belly is grey. Breeding males become black with a bluish tint.

Size

Reaches 23 cm.

Distribution

Found in the Black and Caspian seas including Iranian waters of the latter.

Zoogeography

See genus account above.

Habitat

This species lives in inshore waters over rocks, gravel and stones, although a spawning male was caught over silt near the Kura River, and a female was caught at 11 m at Krasnovodsk (Pinchuk et al. in Miller, 2003). It rarely enters fresh waters, at least in the Caspian.

Age and growth

Sexual maturity is attained at 2 years.

Food

The main food items are crustaceans, with smaller quantities of worms, molluscs and fish.

Reproduction

Spawning takes place from the end of March to the end of May, with a single repeat spawning. Eggs are deposited on and between stones. Females contain 22-360 ripened eggs (Pinchuk et al. in Miller, 2003).

Parasites and predators

None reported from Iran.

Economic importance

None as it is disliked by commercial and sport fishermen and is not common in the Caspian Sea (Pinchuk et al. in Miller, 2003).

Conservation

No data for Iran. Endangered in Turkey (Fricke et al., 2007).

Further work

The biology of this species in Iranian waters requires study.

Sources

Iranian material:

Ponticola gorlap
(Iljin, 1949)

Common names

mahi kafzi (= bottom-dwelling or benthic fish), gel-ye mahi sar bozorg (= bighead goby), gavmahi-ye sarbozorg, gav mahi sar gondeh, sebele, gorzak.

[Names once used for Neogobius kessleri gorlap may be applied to this species: iribogaz xul for Azerbaijan; bychok-golovach or bighead goby in Russian; gorlap goby, Caspian bighead goby]

Systematics

The Caspian Sea basin was thought to contain the subspecies Neogobius (now Ponticola) kessleri gorlap (Iljin, 1949). Iljin (1956) and Vasil'yeva (1991) considered P. k. gorlap to be a distinct species while Berg (1948-1949) and Pinchuk (1977) on external morphological grounds considered it to be a subspecies. Karyotype data (Vasil'ev and Grigoryan, 1993) shows P. k. kessleri from the Black Sea basin to have 29 chromosomes in males and 30 in females and the number of biarmed chromosomes in 2n=29 individuals is 17 and in 2n=30 individuals it is 16. Nominal P. k. gorlap from the Caspian Sea have 2n=46 in both sexes (Vasil'yev and Vasil'yeva, 1992). Vasil'ev and Grigoryan (1993), Vasil'yev and Vasil'yeva (1992), Vasil'yeva et al. (1994) and Vasil'yeva and Vasil'yev (1994) referred the Caspian Sea basin populations to Neogobius gorlap, but indicated that this taxon required redescription and this was carried out in 1996. Vasil'eva and Vasil'ev (1996) consider that Gobius gorlap Iljin is unpublished and nonvalid, i.e. a nomen nudum (Iljin's 1941 manuscript is lost) and give the taxon the name Neogobius iljini with a type locality in the Mangyshlak region of the Caspian Sea (holotype in the Zoological Museum of Moscow State University under ZMMGU P-19726 with paratypes ZMMGU P-46453 (3 specimens); note that the Zoological Museum of Moscow University (ZMMU; their acronym) has P-19726 and P-4643 (sic)(Pavlinov and Borissenko, 2001)). Kottelat (1997) points out that Berg (1948-1949; volume 3 including gobies dated 1949) had access to and used some of Iljin's data, thus making the name gorlap available. Indeed, any of the subsequent papers by Vasil'eva and Vasil'ev and others using the name with descriptive details would have made the name available. Neogobius iljini is therefore a junior synonym of Ponticola gorlap (Iljin in Berg, 1949). There is no type series for P. gorlap as it is based on Iljin's data as outlined above.

Key characters

Pelvic fin anterior membrane with angular lateral lobes, lobes about one-sixth to almost one-half width of anterior edge of membrane; pelvic fin less than nine-tenths distance to anal fin; posterior nostril near edge of orbit; anterior nape with ctenoid or cycloid scales; rear of first dorsal fin without a dark spot; lateral series scales usually 54-76; lateral lobes of pelvic fin anterior membrane large, at least one-fifth width of rear edge; upper lip not markedly swollen, width at least 0.6 lateral preorbital width (lip to orbit), if less than 0.75, then nape scales ctenoid; interorbital distance 0.8-0.9 eye diameter; and caudal peduncle depth 0.67-0.75 length.

Morphology

This species is separated from other Caspian gobies in Iran by having a completely scaled nape, scales usually cycloid on the sinciput (anteriorly on the head just behind the eyes) (70-100% cycloid; 0-30% uniseriate ctenoid in Kura River fish) and cycloid or ctenoid posteriorly over the operculum (60-100% ctenoid in Kura River fish, 0-50% ctenoid in others); scales on the middle of the upper third of the operculum usually cycloid (70-100%); head depth at the eyes is slightly less or about equal to head width between the upper origins of the opercles; a large mouth with the jaw angle below the anterior part of the eye; the snout is much longer than the eye, 1.5-1.6 times; upper lip not expanded, or only slightly, less than lateral preorbital area (between lip and eye); interorbital distance less than 0.85 eye diameter; first dorsal fin not higher than second dorsal fin; moderately large, angular or pointed rather than rounded lateral lobes to the anterior pelvic membrane (rather small and obtuse lobes according to Ahnelt and Holčík (1996)), the membrane is 0.2-0.3 the width of the rear edge as opposed to not more than one-sixth; pelvic disc length is about 0.8 length of pelvic origin to anus length; scales in lateral series 53-79; caudal peduncle depth is 0.6-0.75 its length; the posterior nostril is close to the eye margin; and overall colour is yellowish and marbled.

The head in fish from the Kura River and possibly Iran have a rather subcylindrical shape (43.8-60.0%), while marine and north and middle Caspian Sea basin fish have a depressed and wide head, the width being markedly greater than the depth. The caudal fin is rounded, oval, or rarely elongated.

First dorsal fin spines 5-7, usually 6, second dorsal fin with 1 spine followed by 15-20, usually 18, soft rays, anal fin with 1 spine and 11-16, usually 13-15 (but see below), soft rays, pectoral fin rays 17-22, and pelvic fin rays 8-9. Marine specimens and those from the lower Volga River and Yashan Lake in the Uzboi River system of Turkmenistan have less than 15 anal fin branched rays while freshwater populations (in the Kura River at least, perhaps Iran) have 30-60% of all fish with 15 rays. Scale counts in the southern Caspian Sea are less (53-74) on average than for other populations (61-79) (Vasil'eva and Vasil'ev, 1996). Gill rakers 5-8. Vertebrae 33-35. The chromosome number for P. gorlap is distinctive with 2n=46 with 46 chromosomal arms (Vasil'ev and Grigoryan, 1993; Esmaily and Kalbassi, 2008).

Meristic values for Iranian specimens are: first dorsal fin spines 6(16), second dorsal fin soft rays 16(4), 17(11) or 18(1), anal fin soft rays 12(13) or 13(3), pectoral fin rays 18(8) or 19(8) and lateral line scales 63(5), 64(2), 65(2), 66(2), 67(2) or 68(2).

Sexual dimorphism

Abdurakhmanov (1962) reports on fish from Azerbaijan that head width, predorsal distance, second dorsal fin and anal fin bases and length of third branched anal fin ray are all greater in males while pelvic fin length and eye diameter are greater in females. In sea populations, females are larger than males.

Colour

A strongly yellow to reddish-yellow coloration distinguishes this species (Berg, 1948-1949; Vasil'eva and Vasil'ev, 1996) although Gavlena (1977) describes Volgograd Reservoir specimens as brown and others agree fish may be dark brown to dark grey or greenish. There are 4-8 elongated dark brown spots along the mid-flank in freshwater specimens, the last forming a horizontal "T" with the cross-bar on the caudal fin base. The flanks are usually marbled dark brown with light brown and yellowish strips, sometimes in a cellular design. There are light spots, darkly rimmed, along the sides of the head and near the pectoral fin bases, the latter also being likened to a network, reticulate pattern or cellular design. Cheeks with reticulate or roundish-cellular design. Short, longitudinal stripes often present below the eyes. The belly is light grey or light yellowish.

Dorsal, caudal and pectoral fins yellowish to light brownish. Second dorsal fin branched ray tips white. The fins typically have rows of small dark, orange-grey spots, although the pelvic fin has only a little dark grey or yellowish pigment with some brownish strips or many minute spots. The first dorsal fin lacks a large spot but has 3-4 series of spots; the second dorsal fin has 2-6, usually 3-4, series of spots; the anal fin is yellow to grey with occasional spots; the pectoral fin has 8-10 series of spots; and the caudal fin has 7-8 series of spots. The ends of the second dorsal fin branched rays are white. The anal fin has a light border as does the pelvic disc, although the latter sometimes has a black centre. The caudal fin has many minute spots. The genital papilla is dark grey. Young fish may appear spotted but have a more reticulate pattern on the flank than N. goebelii and are generally darker.

The dorsal fins in breeding males have a wide yellowish or white margin and the second dorsal fin rays have free tips. The margin is rusty in a female. The head can be a dark bluish-grey and the flanks are dark with some paler markings. Fin membranes become dark.

Size

Reaches 22.5 cm. Freshwater populations have fish much smaller than those in the sea (Vasil'eva and Vasil'ev in Miller, 2003).

Distribution

Found in the inshore Caspian Sea and tributary rivers. It is reported from between Kultuk and Astara in Azerbaijan (Ragimov, 1965) and from the Uzboi Lakes in Turkmenistan. In Iran, it is reported from a wide range of rivers from Astara to the Gorgan and probably the Atrak, including the Siazan, Safid, Duyusan, Shimrud, Khoshkerud, Palarud, Naviq, Shasdehrud, Shafarud, Kargan, Gharasu, Tajan, Babol, Haraz, Pol-e Rud and Safid, the Aras River, the Anzali Mordab and Bahambar, Siah Darvishan, Pasikhan, Massouleh and Pir Bazar river, Gorgan Bay, the southeast Caspian Sea, southwest Caspian Sea and south-central Caspian Sea (Holčík and Oláh, 1992; Ahnelt and Holčík, 1996; Abbasi et al., 1999; Kiabi et al., 1999; Abdoli and Naderi, 2009). Abdoli and Naderi (2009) consider it to be absent from the Atrak, Sardab, Tonekabon and Aras rivers.

Zoogeography

This species is thought to have evolved from populations which settled in the lower reaches of rivers and in coastal areas while those populations which penetrated headwaters evolved into P. cyrius (q.v.) (Ahnelt and Holčík, 1996). See also genus account above.

Habitat

This species is found in fresh and brackish waters in rivers and lagoons and in inshore waters of the Caspian Sea. In the sea, it is found down to 10 m, rarely to 20 m, on rocky and dense sand bottoms. It moves inshore in the sea for spawning and retreats to deeper water (6-12 m) to overwinter (Vasil'eva and Vasil'ev in Miller, 2003). It may be found among reeds. It can live at river sites with a current velocity at 4.5 m/sec on the bottom (Holčík and Oláh, 1992). This species is allopatrically distributed with Ponticola cyrius (q.v.). Ahnelt and Holčík (1996) rate this species as a eurytope, inhabiting both fresh and brackish waters, slow and rapid currents, and clay and pebble and gravel bottoms. Kirilenko and Shemonaev (2010) found fish in the Kuibyshev Reservoir in Russia to prefer rocky, sandy-slimy grounds rich in higher aquatic vegetation and are sometimes found among cane roots.

Age and growth

Brackish water populations grow larger than freshwater ones (Ahnelt and Holčík, 1996). Males attain 3 years, females 2 years, and males grow faster than females. Maturity is reached at 1+ for most females and 2+ years for males (Vasil'eva and Vasil'ev in Miller, 2003). Kirilenko and Shemonaev (2010) found a female in the Kuibyshev Reservoir that was 4 years old while the oldest fish in the 3+ class were all males, fish matured at 1+ years for both sexes. Abdoli et al. (2006) examined fish from the Gomishan and Miankaleh wetlands and found a length (L)-weight (W) relationship of W = 0.1L2.39 for males, 0.0595L2.588 for females and 0.1L2.4 for sexes combined.

Food

The principal food in the sea is fishes, mostly gobies and Atherina, but also includes crustaceans. Benthic food means this species competes with commercial fishes. In fresh water, fishes eaten include Nemacheilidae and the cyprinid Alburnoides bipunctatus (= eichwaldii), as well as crustaceans such as gammarids (e.g. Dikerogammarus haemobaphes), gobies such as its own species and N. melanostomus, insects and clams (Gavlena, 1977). Kirilenko and Shemonaev (2010) found food in the Kuibyshev Reservoir to be mainly  crustaceans and chironomids.

Reproduction

Spawning takes place in June in the Volgograd Reservoir and each female may contain up to 576 eggs (Gavlena, 1977). In the Caspian Sea, fish have as many as 3506 large eggs and 3399 smaller ones. Spawning takes place on rock and shingle bottoms with eggs laid on various objects. The males of this species are nest guarders. Spawning in the southern Caspian is in April and May with two batches of eggs deposited. In the lower Volga River, spawning is from the end of April to July (Vavsil'eva and Vasil'ev in Miller, 2003).

Parasites and predators

The Caspian seal, Pusa caspica, is a predator on this species (Krylov, 1984). Pazooki and Aghlmandi (1998) found the nematode Dichelyne minutus infecting the intestine of 100% of fish examined from the south Caspian Sea in Iran. and Sattari et al. (2002; 2004) and Sattari (2004) record the presence of the nematode, Eustrongylides excisus, in this species. This parasite can damage muscles in commercial species and render them unsuitable for sale. Dichelyne minutus was also recorded. Daghigh Roohi and Sattari (2004) record Eustrongylides excisus, Dichelyne minutus and Corynosoma strumosum from this species in the southwestern Caspian Sea of Iran. Sattari et al. (2005) surveyed this species in the inshore area of the Caspian Sea, recording Eustrongyloides excisus and Dichelyne minutus.

Economic importance

None in Iran although it has formed a small part of the goby catches in the former Soviet Union. It is also caught by anglers (Vasil'eva and Vasil'ev in Miller, 2003).

Conservation

Kiabi et al. (1999) consider this species to be of least concern in the south Caspian Sea basin according to IUCN criteria. Criteria include abundant in numbers, widespread range (75% of water bodies), absent in other water bodies in Iran, and present outside the Caspian Sea basin.

Further work

The biology of this specie sin Iran needs work.

Sources

Meristics and colour description based in part on Gavlena (1977) describing fish from the Volgograd Reservoir, identified as N. kessleri. Counts and data for Iranian fish are based on Ahnelt and Holčík (1996).

Iranian material:

Ponticola syrman
(Nordmann, 1840)

Common names

None (other than general names listed under genus above).

[shirman in the Ukraine, syrman goby, Kaspiiskii shirman or Caspian syrman goby]

Systematics

Gobius syrman is described from Odessa, Dniester estuary, in the former U.S.S.R., now Ukraine). Gobius Trautvetteri Kessler, 1859 described from the lower Bug and Dniester rivers in the former U.S.S.R. and Gobius eurystomus Kessler, 1877 described from the southern Caspian Sea are synonyms. Berg (1948-1949) has eurystomus as a valid subspecies of Neogobius syrman but this is not accepted now (Reshetnikov et al., 1997). Presumed syntypes of G. syrman are in the Natural History Museum, London (BM(NH) 1872.5.30:35, 105.0 mm standard length), Muséum national d'Histoire naturelle, Paris (MNHN A. 1126) and in the Naturhistorisches Museum Wien (MNMW 30099. Syntypes of G. eurystomus are under BM(NH) 1897.7.5.8-9 (2, 73.6-88.6 mm standard length), in the Zoological Institute, St. Petersburg (ZISP 10904-05), and perhaps NMW 29176. (Pinchuk et al. in Miller, 2003).

Key characters

Three transverse suborbital neuromasts rows occur ventral to the longitudinal suborbital row b while all other Neogobius have typically two rows. Additionally, pelvic fin anterior membrane with rounded and shallow lateral lobes, lobes not more than one-sixth width of anterior edge of membrane, or lacking entirely; pelvic fin disc two-thirds distance to anal fin origin; nape with ctenoid scales; first dorsal fin without large dark spot; lateral series scales usually 55-70; and angle of jaw below pupil of eye.

Morphology

First dorsal fin with 5-7, usually 6, spines, second dorsal fin with 1 spine and 15-19, usually16-18 soft rays, anal fin with 1 spine and 10-15, usually 12-14, soft rays, and pectoral rays 17-21. Range in lateral series scales 56-79. Vertebrae 34-35. Pelvic fin anterior membrane with very shallow, rounded lateral lobes. Posterior nostril near edge of orbit. Angle of jaw below anterior part of pupil, upper lip with uniform width, about half lateral preorbital area (between lip and eye). First dorsal fin relatively high, with rounded upper profile, without an upper anterior dark blotch. Second dorsal fin highest in middle. Head depth at eyes 1.1-1.2 width between upper origins of opercles. Interorbital width less than or equal to eye diameter. Snout length 1.3-1.4 orbit. Lateral series scales usually 59-67. Nape and predorsal area scaled completely. Breast, base of pectoral fin and upper part of opercle scaled, cycloid scales on ventral part of abdomen, opercle and anterior part of nape. Pelvic fin about two-thirds distance to anal fin origin. Caudal peduncle depth about half length. Posterior nostril close to the eye margin. The head canals and the free neuromasts on the body are typically arranged for a neogobiine. Three transverse suborbital neuromasts rows occur ventral to the longitudinal suborbital row b.

Sexual dimorphism

Males may have a more intensely dark distal band and body spots during spawning. Male median fin rays are slightly elongated during spawning. Females are usually smaller than males.

Colour

Overall colour pale grey, brownish-grey or more rarely yellowish-brown (perhaps fish parasitised with the strigeid nematode Neascus), with pale saddles across the back extending onto the flank and sometimes meeting dark blotches along the mid-flank. The body has large grey-brown markings arranged in a chequer-like pattern. The head has brown markings laterally and a dark band antero-ventrally from the eye to the upper lip. The first dorsal fin has a pale yellowish margin set off by a dark stripe above two brown stripes proximally. The second dorsal fin bands are less distinct and the anal fin has an indistinct, bluish band and a white margin. The pectoral fins are yellowish and the pelvic disc is pale. Belly pale. Males do not become dark during spawning (Pinchuk et al. in Miller, 2003).

Size

Attains 29.2 cm.

Distribution

Black and Caspian Sea basins including the Imeni Kirova or Kyzylagach Bay near the northwestern Iranian border (Kuliev, 1989) and between Kultuk and Astara in Azerbaijan (Ragimov, 1965) and recently recorded from Iran (Ahnelt et al., 2007). Reported from the Anzali Mordab and Gomishan Lagoon, Gorgan Bay, and the southwestern, southeastern and south-central Caspian Sea (Jolodar and Abdoli, 2004; Abdoli and Naderi, 2009). Introduced to the Aral Sea.

Zoogeography

See genus account above.

Habitat

This goby is found principally in the sea, with one report from fresh water in the Caspian Sea basin (Emba River, Kazakhstan). It is unusual in being found on mud and silt bottoms and can tolerate low oxygen concentrations (to 20% saturation). It is rare offshore at 6-20 m in the Caspian Sea. It is found in the Caspian at 1-10 m, retreating in summer to slightly deeper water, reappearing as autumn cooling occurs, and then spending the winter at 30-50 m. It approaches shores earlier than other gobies for spawning, as early as the latter half of February for males with females in the second half of March (Pinchuk et al. in Miller, 2003).

Age and growth

Maximum age may be 6 years although most fish probably live only half this time. Maturity may be attained shortly before age 1 year and by 2 years for most fish. Abdoli et al. (2006) examined fish from the Gomishan and Miankaleh wetlands and found a length (L)-weight (W) relationship of W = 0.0033L3.41 for males, 0.0033L3.5 for females and 0.003L3.5 for sexes combined.

Food

Fish, crustaceans and Nereis worms are the main diet items in Caspian Sea fish examined. Fish included Knipowitschia and fry of Neogobius pallasi and N. melanostomus as well as clupeids and its own species. Smaller individuals take crustaceans and larger ones favour fish (Pinchuk et al. in Miller, 2003).

Reproduction

The spawning season is from the beginning of April into May in the southern Caspian Sea. The eggs are found in two sizes, indicative of batch spawning. Up to 8474 ripe eggs are found in each female. Eggs are 4.2 by 1.9 mm and are laid under and between stones, on any available object or even muddy bottoms.

Parasites and predators

This species is eaten by Sander lucioperca and fry are eaten by sturgeons.

Economic importance

This species has been fished in former Soviet waters of the Caspian Sea because of its large size but the proportion in catches varies from 1-50%. It is important as food for commercial species.

Conservation

Only recently recorded from Iran, its conservation status in unknown.

Further work

The distribution and abundance of this species in Iranian waters needs to be surveyed as it is an important food for commercial fishes.

Sources

Pinchuk et al. in Miller (2003) is a recent summary of data on this species.

Iranian material: CMNFI 1970-0544, 1, 141.7 mm standard length, Gilan, near Bandar-e Anzali, (37°28’N, 49°27’E); CMNFI 1979-0689, 9, 87.1-124.0 mm standard length, Gilan, Safid River at Hasan Kiadeh (37°24’N, 49°58’E); CMNFI 1970-0586, 1, 100.1 mm, Mazandaran, Gorgan Mordab at Ashuradeh-ye Kuchak (36°50’N, 53°56’E); NMW 95074, 2, 100.1–130.2 mm, Mazandaran, south-east Caspian Sea near estuary of Gorgan River (36°59’N, 53°59’E); CMNFI 1979-0788, 2, 135.3–146.2 mm standard length, Mazandaran, Gorgan River (37°00’N, 54°07’E); NMW 95075, 1, 186.6 mm standard length, Mazandaran, south-east Caspian Sea, Mian Kaleh near Behshahr (36°53’N, 53°32’E).

Genus Proterorhinus
Smitt, 1900

This genus is characterised by having 6 first dorsal fin rays, a pelvic disc without obvious lateral lobes and, uniquely in Ponto-Caspian gobies, a very elongate anterior nostril which hangs over the lip. Miller in Miller (2004) give additional characters, in particular the distribution of neuromasts on the head. It contains only a single species found in the Black and Caspian seas and their tributary rivers and in the northern Aegean Sea in Greek fresh waters. There may be two species if P. semilunaris is recognised (see below). Simonovič (1999) considered this genus to be a subgenus of Neogobius but Ahnelt and Duchkowitsch (2001) provisionally consider Proterorhinus to be distinct based on their study of the lateral line system and consider it distinct based on osteology (Ahnelt and Duchkowitsch, 2004). Miller in Miller (2004) recognises the genus as distinct as do Stepien and Tumeo (2000) based on mtDNA. Simonovič et al. (1996) consider this genus of gobies to be a young one as evidenced by the distribution of the species in the Caspian Sea which closed recently.

Reshetnikov et al. (1997) and Miller in Miller (2004) have the date for this genus as 1899 but Eschmeyer's "Catalog of Fishes" states that it was published in 1900 although dated 1899.

Proterorhinus nasalis
(De Filippi, 1863)

Common names

gav mahi, sag mahi, gel-ye mahi marmari, gavmahi-ye marmari, sebele.

[marmar xul in Azerbaijan; bychok-tsutsik or tubenose goby, marmornyi bychok or marbled goby, both in Russian; mottled goby].

Systematics

The Caspian Sea tubenose goby has generally been referred to Proterorhinus marmoratus (Pallas, 18l4). Gobius marmoratus was originally described from Sevastopol, Ukraine, on the Black Sea. Types are lost (Miller in Miller, 2004). Gobius nasalis De Filippi, 1863 described in Latin from the "Mar Caspio presso Baku" and Gobius blennioides Kessler, 1877 described from "Bakinskoi bukhte" (= Baku Bay), were regarded as synonyms. The meristic characters reported by De Filippi (1863) for nasalis are probably in error according to Berg (1948-1949) but the only character falling outside the ranges compiled below is for soft dorsal rays given as 21.

Stepien and Tumeo (2006) used mtDNA cytochrome b sequence data and consider there are two species in this genus. P. marmoratus is a marine species in the Black Sea and P. semilunaris (Heckel, 1837) is a freshwater species in other Eurasian habitats. The type locality of the latter is "Maritza [Marizza] R., near Plovdiv, e. Rumelia, Balkan region of Bulgaria". Caspian Sea basin Proterorhinus were not investigated. Kottelat and Freyhof (2007), Freyhof and Naseka (2007) and Sorokin et al. (2011) restrict marmoratus to brackish waters in Sevastopol and consider nasalis to be the name for Iranian Caspian Sea populations.

Neilson and Stepien (2009b) examined mtDNA and nuclear DNA and confirmed the presence of a distinct taxon in the Caspian Sea basin. They advocated P. semipellucidus as the name for the Caspian Sea basin taxon. Gobius semipellucidus Kessler, 1877 was described from a single specimen taken at the "ust'e rechki Kara-su, vpadayushchei vb Astrabadskii zalivb" (= mouth of the Karasu River falling into Astrabad Bay (Qareh Su, Gorgan Bay) and the holotype is in the Zoological Institute, St. Petersburg (ZISP) (Eschmeyer et al., 1996). P. semipellucidus is given as a valid species in Eschmeyer's "Catalog of Fishes" but is presumably a synonym of P. nasalis. The relationship between marine Caspian Sea (brackish) populations (nasalis) and true freshwater populations (possibly semipellucidus) remains to be resolved. Neilson and Stepien (2009b) examined only freshwater and brackish (0-6 p.p.t.) specimens from the Caspian Sea basin. Sorokin et al. (2011) consider nasalis to be the taxon widely distributed in the Caspian and Sea of Azov basins with pellucidus a synonym, although further work is needed to determine if pellucidus is the freshwater taxon.

The syntypes of Gobius nasalis are in the Istituto e Museo di Zoologia della R. Università di Torino under MZUT N.672 (7 specimens), the Natural History Museum, London under BM(NH) 1869.3.4:34 (1, 43.4 mm standard length), in the Museo Civico di Storia Naturale di Genova under MSNG 12655 (2) and MSNG 36228 (3), in the Naturhistorisches Museum Wien under NMW 33894 (1), NMW 33895 (1) and NMW 33896 (1), and in the Museum für Naturkunde, Universität Humboldt, Berlin under ZMB 5015 (3) (Eschmeyer et al., 1996; Miller in Miller, 2004; 43.3-48.4 mm standard length, measured in February 2006). The holotype of Gobius semipellucidus is lost. Ahnelt and Mikschi (2008) give details on the types of Gobius semilunaris.

Key characters

The combination of the pelvic fins forming a disc, the head not being flattened, and the anterior nostrils forming a tube overhanging the upper lip are distinctive. The origin of the first dorsal fin is displaced anteriorly in comparison to other Ponto-Caspian and Atlantic-Mediterranean gobioids, and this reflects the differing postcranial osteology detailed in Ahnelt and Duchkowitsch (2004).

Morphology

First dorsal fin spines 5-7, usually 6, second dorsal fin with 1 spine and 12-20, usually 16-17, soft rays, anal fin with 1-2 spines and 11-17, usually 13-14, soft rays, and pectoral fin with 14-16 branched rays. Scales in lateral series 34-53. Scales are a vertical oval with the anterior margin rounded to slightly wavy, the upper and lower margins rounded and the posterior margin with almost straight upper and lower elements coming almost to a central point. This causes the exposed portion of the scales to be delimited by almost straight edges and flank scales have a diamond pattern. Circuli are numerous and fine. Radii are numerous and radiate back from a focus almost at the posterior margin. The posterior scale margin bears fine ctenii. Gill rakers 3-6, short and concentrated around the angle of the gill arch. A membrane partly closes off and narrows the gill opening at the gill raker level and may partially encompass a raker making counts difficult. Vertebrae 30-33. The chromosome number is 2n=46, with 46 acrocentric chromosomes and 46 chromosomal arms (Ráb, 1985; Grigoryan and Vasil'ev, 1993; Simonovič, 1999). The predorsal area on the back and the nape are scaled. The cheek has infraorbital neuromast organs in 7 transverse rows from the lower orbit border without uniserial papillae between the rows. The preopercular canal has three pores, the anterior oculoscapular canal has 7 pores and the posterior oculoscapular canal has 2 pores. Ahnelt and Duchkowitsch (2001) give a detailed description of the lateral line system on the head. The gut is short and s-shaped. The genital papilla has a two-lobed tip.

Freyhof and Naseka (2007) note that Caspian tubenose gobies are most similar to P. semilunaris (Heckel, 1837) of the Aegean and Black seas basins and are distinguished by having a shorter posterior membrane to the first dorsal fin, not reaching the origin of the second dorsal fin (versus reaching), a longer anterior naris reaching to the middle or to the lower margin of the lower lip when folded down (versus to the upper lip or to the upppermost margin of the lower lip), and a smaller eye (diameter 16-21% head length versus 20-28%).

Meristic values for Iranian specimens are:- first dorsal fin spines 5(1), 6(28) or 7(1); second dorsal fin soft rays 13(1), 14(6), 15(17) or 16(6); anal fin soft rays 12(13), 13(14), 14(2) or 15(1); total pectoral fin rays 14(18) or 15(12); lateral series scales 36(1), 38(4), 39(7), 40(7), 41(4), 42(5), or 44(2); total gill rakers 3(18), 4(9) or 5(3), although accuracy is doubtful because of the membrane; and total vertebrae 33(10), 34(16) or 35(3).

Sexual dimorphism

Males are overall darker than females, e.g. one at 60.1 mm standard length had pigment on both rays and membranes of fins, and all fin margins frilly and pale or orange-yellow. Cheeks are swollen in spawning males and fins are longer than in females.

Colour

Overall colour is brownish or yellowish-brown to olive or yellowish-green with about 5 oblique and irregular bars or blotches across the flank. Colour varies with the background, being darker on mud bottoms and a bright reddish-brown on sand. The head has brown-grey bands over the top and sides on a pale background. The first dorsal fin may have a pinkish tinge and irregular grey blotches. The second dorsal fin has several series of brown spots with the upper 2-3 greenish-yellow. The caudal fin is grey interrupted by up to 9 narrow bands of greenish-yellow. There is a triangular black spot at the caudal fin base, flanked by two white spots. The pectoral fin is similarly greyish with 5-7 narrow bands of pale green or yellow. The pelvic fin has 3-4 yellow bands and the anal fin 6-7 yellowish and oblique bands over a grey background. The iris has a narrow golden ring. Spawning males have all fins dark, although the pectoral fin is lighter than the rest and the caudal fin has an obvious white margin. The genital papilla is grey. Breeding females have a colour similar to that of both sexes in winter. Young (16-22 mm) have an unpigmented median band, wider than the eye, from the head to the first dorsal fin and even to the caudal fin. In front of the first dorsal fin is a dark saddle. The peritoneum is light brown to silvery with varying amounts of melanophores.

Size

Attains 15.0 cm although Caspian specimens reputedly reach only 7.6 cm.

Distribution

Found in the Caspian Sea basin and the Uzboi Valley of Turkmenistan. It is found in a wide range of rivers on the Caspian coast of Iran including the Qareh Su falling into Gorgan Bay, Kargan River estuary near Hashtpar, lower Safid,  Rud-e Sera and Babol, in the Anzali Mordab and Kolesar River, the southeast Caspian Sea, the southwest Caspian Sea and south-central Caspian Sea (Derzhavin, 1934; Holčík and Oláh, 1992; Abbasi et al., 1999; Kiabi et al., 1999; Abdoli and Naderi, 2009).

Zoogeography

The genus Proterorhinus diverged from the related genus Neogobius about 5.2MYA during the late Miocene/early Pliocene according to mitochondrial DNA evidence (Dillon and Stepien, 2001). Freshwater and marine/brackish populations apparently diverged 3.82-4.30 MYA and freshwater lineages in the Black and Caspian seas diverged 0.92-1.03 MYA (Neilson and Stepien, 2009b).

Habitat

This species occurs in both salt and brackish waters and enters the fresh water of rivers. There are some permanent freshwater populations in Europe. The usual habitat is shallow sea bays, offshore banks or flowing water of streams but it may also be found in ponds and canals overgrown with vegetation. Where current is strong it hides under boulders. Some fish are found below 3 m depths in the sea. It is often found under stones or among weed, to which it can retreat rapidly if disturbed. Most activity takes place at night.

Age and growth

Life span is variably reported as 2 or about 5 years with maturity attained at 1-3 years. Young-of-the-year can mature by autumn at 5.5 cm in the Caspian Sea (Miller in Miller, 2004).

Food

Kuliev (1989) records insect remains, gammarids, daphnia, chironomids, molluscs and fish eggs in the guts of this species in Kirova Bay of Azerbaijan. For the Caspian Sea as a whole, amphipods predominate at 45.7% followed by mysids at 21.8%, cumaceans at 16.4% and decapods at 7.2%. Molluscs make up 5.6% (Kosarev and Yablonskaya, 1994). Iranian specimens contained crustacean and insect remains. Elsewhere polychaete worms and small fish have been found in stomachs of this species (Miller in Miller, 2004).

Reproduction

Both sexes produce sounds during the breeding season. Eggs are laid from shallow water (0.5 m) down to 20 m in bowl-shaped nests, under rocks, in empty mollusc shells, and on roots and stems of plants. Empty cans may be used. Two females may lay eggs in the same nest. Spawning begins in mid-April in the southern Caspian and may continue as late as the first half of August. Some fish may be mature in the middle of March in Iranian waters. Fish presumed to be young-of-the-year at 9.8 mm standard length have been collected on 26 April, by 10-11 June fish are 11.2-11.6 mm (mean 11.4 mm), on 20 June the 5 smallest fish of a sample were 12.5-14.1 mm, mean 13.6 mm while the 5 largest were 21.0-25.2 mm, mean 23.4 mm, and on 12 October the 5 smallest were 16.6-19.9 mm, mean 17.7 mm while the 5 largest were 26.4-33.3 mm, mean 29.6 mm. By 15 March, eggs in a 49.7 mm standard length fish are 1.3 mm in diameter and well-developed. This data suggests an extended spawning season in Iran. However young are first found in mid-June according to Ragimov (1987). Eggs are deposited in 2-3 batches and are guarded. Females contain up to 1335 eggs of diameter 3.3-3.6 by 1.4-1.5 mm. Elsewhere up to 2500 eggs may be laid. Egg clutches can be transported on the hulls of ships (Ahnelt et al., 1998). Males are nest guarders.

Parasites and predators

Unknown for Iran.

Economic importance

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in aquaria and because it has been introduced outside its natural range (Jude, 2001).

Conservation

Lelek (1987) classifies this species as vulnerable in Europe on account of water level changes. Kiabi et al. (1999) consider this species to be data deficient in the south Caspian Sea basin according to IUCN criteria. Criteria include few in numbers, medium range (25-75% of water bodies), absent in other water bodies in Iran, and present outside the Caspian Sea basin.

Further work

The biology of this species in Iran is not well known.

Sources

Type material: See above for Gobius nasalis (ZMB 5015).

Iranian material: CMNFI 1970-0530, 66, 41.6-64.5 mm standard length, Gilan, Nahang Roga River (37º28'N, 49º28'E); CMNFI 1970-0542, 5, 48.0-59.6 mm standard length, Gilan, Old Safid River estuary (37º23'N, 50º11'E); CMNFI 1970-0585, 2, 52.6-55.9 mm standard length, Gilan, Nahang Roga River (37º28'N, 49º28'E); CMNFI 1970-0590, 5, 39.5-47.7 mm standard length, Mazandaran, Shesh Deh River near Babol Sar (no other locality data); CMNFI 1979-0787, 3, 52.7-59.1 mm standard length, Gilan, Nahang Roga River (37º28'N, 49º28'E).

Genus Rhinogobius
Gill, 1859

Members of this genus are characterised by an elongate body which is compressed posteriorly, a depressed head, a long snout, usually small scales on the occiput, short soft dorsal and anal fins both with 9-10 rays, the first dorsal fin with 6 spines, scales are ctenoid and number 23-40 in lateral series, anterior nostrils are tubular, the tongue is not notched, and teeth are simple.

There are about 48 species in Japan, Korea, Taiwan, China and the Amur River basin of eastern Asia. One species, presumably from the Amur River basin in the former Soviet Union, has been introduced to Turkmenistan and has reached Iranian waters.

Rhinogobius similis
Gill, 1859

Common names

gel-ye mahi, gavmahi-ye talabi (= pond or lake goby).

[lake goby]

Systematics

Rhinogobius similis was originally described from Shimoda, Japan

Rhinogobius similis lindbergi Berg, 1933 described from the Amur and Ussuri rivers, Russia is the subspecies of the Amur River basin according to Pinchuk (1978) but Eschmeyer (1998) records it as a synonym of the type subspecies. The Iranian specimens probably had their ultimate origin in the Amur River basin. Vasil'eva (2007) and Vasil'eva and Kuga (2008) consider that the Rhinogobius introduced to Central Asia are R. cheni (Nichols, 1931), a Chinese species of the Yangtze River, and this could be the species found in Iran. However, the large number of species in the genus, the poor state of knowledge on these fishes and the source presumably being the Amur River, leaves this identification as open.

Key characters

The only member of its genus in Iran, its key characters are those outlined in the genus.

Morphology

Dorsal fin spines 5-8, dorsal soft rays 7-10 after an initial unbranched ray, anal fin soft rays 6-10 after an initial unbranched ray, lateral series scales 28-36. Scales can be absent from the midline of the nape and the parietal and occiptal regions but this is variable (the nominal subspecies lindbergi lacks scales). Adult fish have the dorsal spines elongated (but not in lindbergi, not even in males). Scales in small Iranian specimens are a vertical oval, the posterior margin is fringed with ctenii, circuli in these small specimens are moderately devloped, radii radiate from the central point of the posterior margin to the anterior margin, the posterior margin has almost straight upper and lower margins coming to a central point so flank scales form a diamond pattern. The gut is a short s-shape. Gill rakers number 8-12 and are short and touch the adjacent one when appressed. Total vertebrae number 24-28.

Meristic values for Iranian specimens are:- first dorsal fin spines 6(3); second dorsal fin soft rays 9(3); pectoral fin rays 18(2) or 19(1); lateral series scales 31(3); total gill rakers 10(1) or 11(2); and total vertebrae 27(2).

Sexual dimorphism

Unknown.

Colour

Overall colour is a pale yellow or light brown with 6 brownish blotches on the flank and 6 blotches on the back, their position alternating. A dark band runs through the eye to the posterior edge of the mouth. The upper head is speckled brown. The belly is white. There is a dark spot between the first and second dorsal fin spines in the nominal lindbergi subspecies. There are 2 rows of spots on the first dorsal fin and 3-4 more distinctive rows on the second dorsal fin. The caudal fin has 2-5 rows of spots. The anal fin bears pigment distally on the membranes. The pectoral and pelvic fins are greyish. The pigmentation of the type subspecies is given by Pinchuk (1974) but may differ in detail from the nominal lindbergi subspecies so is not given here.

Young fish have 6 blotches along the mid-flank and membranes of fins are either strongly pigmented overall or bear up to 6 bars on the first dorsal fin, second dorsal fin and caudal fin with pectoral, pelvic and anal fins much lighter, almost immaculate. There is a small blotch between the first and second first dorsal fin rays near the fin base. The distal margin of the anal fin is yellowish-orange. The peritoneum is silvery but has a strong development of melanophores dorsally.

Size

Attains 100 mm.

Distribution

Aliev et al. (1988) refer to an exotic species of goby occurring in the Tedzhen River of Turkmenistan in the text of their work but not in the table of distributions. A goby is also reported by Shakirova and Sukhanova (1994) and Sal'nikov (1995) from the Karakum Canal and the Kopetdag Reservoir in Turkmenistan on the northern borders of Iran. Specimens have been collected from Sarakhs, Iran in the Tedzhen (= Hari) River (Coad and Abdoli, 2000a; Abdoli et al., 2000) and Abdoli (2000) records it from the Kashaf and Hari rivers in the Tedzhen River basin.

Zoogeography

An exotic or alien species from eastern Asia, introduced to Turkmenistan from where it presumably entered Iranian waters. The genus Rhinogobius is in need of revision and the correct name for the taxon in Iran may change.

Habitat

The Iranian specimens were caught in the Hari River which had a depth of 35-120 cm , a width of 12-27 m, a turbidity reading of 15 cm, a water temperature of 12°C and a coarse gravel bed. It eastern Asia it is found in both rivers and lakes (hence the common name of lake goby as used in Japan). It is found on shallow sandy bottoms and on the upper surface of large stones in shallow water. Water temperatures reach 28°C (Kopylets and Dukravets, 1981).

Age and growth

Life span exceeds 4 years (Kopylets and Dukravets, 1981).

Food

Unknown.

Reproduction

Egg production reaches 1862 eggs in total (Kopylets and Dukravets, 1981).

Parasites and predators

Unknown for Iran.

Economic importance

Welcomme in Courtenay and Stauffer (1984) reports this species to be a pest when introduced.

Conservation

As an introduced species this fish requires no conservation.

Further work

The spread of this species in Iranian waters and its effects on native fauna should be documented.

Sources

Iranian material: Uncatalogued material, 5, 42.4-47.6 mm standard length, Khorasan, Hari River near Sarakhs (36º30'N, 61º10'E).

Channidae

This family is known as the snakeheads or serpent-heads because of the characteristic broad head with large scales and a large oblique mouth. They are found from Africa to eastern Siberia and Southeast Asia and comprise about 29 species but the family is need of a revision. Maximum size is about 1.2 m.

Snakeheads are characterised by paired accessory organs or suprabranchial organs in the upper gill chamber (above and behind the gills) which enable these fishes to breathe air, survive low oxygen conditions and even reputedly travel overland; an elongate rounded body becoming compressed posteriorly; the dorsal and anal fins are long and of even height, and spineless; pelvic fins are present or absent; the mouth is large and the lower jaw protrudes; there are teeth on the jaws, vomer and palatines; gill openings are wide and the gill membranes are united but free from the isthmus; branchiostegal rays number 5; the caudal fin is rounded; scales are small and cycloid or ctenoid; and colour is highly variable, rapidly changing to suit the surroundings.

Snakeheads are ambush predators, living in still waters although some inhabit the larger rivers. Most species build a bubble nest in vegetation, laying and fertilising the eggs below it so that they float up into the bubbles. Others are mouth brooders. One or both adults guard the nest and young and will attack intruders savagely, including humans according to folklore. Many species are known to aestivate in summer when the habitat dries. They can be carried alive wrapped in wet cloths or vegetation and may be introduced into areas outside their natural distribution.

A number of species grow large enough to be an important food in Southeast Asia but are also pests, eating more valuable species. Some species are popular in the aquarium trade. Juveniles may be called, incorrectly, "larvae" in the aquarium trade and are brighter in colour than adults.

Genus Channa
Scopoli, 1777

Channa Scopoli, 1777 has priority over Ophicephalus Bloch, 1793 (Eschmeyer, 1990). Ophiocephalus is an incorrect emendation. See also Myers and Shapovalov (1931-1932), DeWitt (1960), Ettrich and Schmidt (1989) and Pethiyagoda (1991) for discussions on the taxonomy of this genus and species.

The characters listed under the family above obtain for the genus which has only one confirmed Iranian species.

Channa argus
(Cantor, 1842)

Reported as Channa argus warpachowskii (Berg, 1909) from the Karakum Canal and Kopetdag Reservoir in Turkmenistan as an exotic from China (Shakirova and Sukhanova, 1994; Sal'nikov, 1995). This canal connects with the Tedzhen River basin shared with Iran and ultimately may connect with the Caspian Sea basin. No Iranian record.

Channa gachua
(Hamilton, 1822)

Common names

mahi-ye sarmari (= snakeheaded fish).

[tond, dolli or dauli in Pakistan; dwarf snakehead, frog snakehead, brown snakehead, oriental snakehead, smooth-breasted snakehead].

Systematics

Ophicephalus gachua was originally described from Bengal, India. Possible syntypes are in the Natural History Museum, London under BM(NH) 1858.8.15.54 (1) and BM(NH) 1858.8.15.144 (1) (Eschmeyer's "Catalog of Fishes", downloaded 29 August 2007).

Key characters

The large head scales and elongate dorsal and anal fins are distinctive.

Morphology

Dorsal fin rays 30-37 (the last 2 rays counted as 1 where close together at base), anal fin rays 20-24, pectoral rays 13-16, and lateral line scales 39-47. Pelvic fins may be present or absent. The lateral line is displaced down one row under, or just beyond, the posterior end of the pectoral fin. Scales have a vertical anterior margin, parallel dorsal and ventral margins and a rounded posterior margin. The anterior dorsal and ventral corners are square cut, rounded to sharp. Circuli are fine and numerous on the anterior field but on the posterior field become coarser and are parallel to the horizontal axis. Radii are numerous in the anterior field and radiate from a central focus. Gill rakers are minute. The gut is short and s-shaped. The anterior nostril is tube-shaped and hangs over the upper lip to the mouth. The chromosome number is 2n=78 (Banerjee et al., 1988; Klinkhardt et al., 1995).

Meristic values for Iranian specimens are:- dorsal fin rays 33(1), 34(3) or 35(1) (the last 2 rays counted as 1 where close together at base); anal fin rays 22(5); pectoral rays 14(3), 15(1) or 16(1); and lateral line scales 41(2), 43(2) and 44(1).

Sexual dimorphism

Females have a dark eye-spot at the posterior part of the dorsal fin (Pethiyagoda, 1991).

Colour

Colour varies with the habitat. The back is usually greenish-grey to brownish with bluish tints and the flank is crossed by irregular oblique bars, less obvious in adults than young. The background colour of the flank is dove grey with a violet sheen in males. The dorsal, anal and caudal fins are slate-coloured and have characteristic, narrow and strong orange margins (white in preservative). The membranes of these fins may be an iridescent green. The dorsal fin may have a blue or bluish-green stripe with a vertical extent from the fin mid-point distally about half-way to the fin margin on the membranes. The caudal fin has blue and green rays. Females have a dark eye-spot at the end of the dorsal fin. The pectoral fin base is dark blue and there are 4-5 orange and blue bands on the fin, with the margin orange. Bars on the pectoral fin are very distinctive in preservative, alternating dark and light. The iris is reddish. The peritoneum is silvery.

Size

Attains 33 cm (Day, 1875-1878) although Courtenay and Williams (2004) give 17 cm, probably more accurate given taxonomic confusion in the past.

Distribution

Reported from the Dasht and Rakshan rivers, the Makran coast and the Mashkel (= Mashkid) River basin in Pakistan (Zugmayer, 1913). Iranian records were limited to 4 specimens collected by N. A. Zarudnyi in the Bampur river (upper or middle course) during 15-23 July 1898 (27 June-4 July in Berg (1949)) until 1 specimen was found in the Halil River basin near Sabzeveran at 28°39'N, 57°45'E, over 300 km northwest of the Bampur River at Bampur (Coad, 1979a). Abdoli (2000) reports this species from the Halil and questionably from the Iranian Makran. Ebrahimi (2001) maps two localities in the Halil River. The map in Berra (2001) has this species too close to the Straits of Hormuz and the map in Courtenay and Williams (2004) does not extend far enough into Iran.

Zoogeography

The distribution of this species in southeastern Iran is confirmed in Coad (1979a), a record not noted by Bănărescu (1992b). It is the westernmost occurrence of the species in Asia. It is found eastwards to Indonesia (Ettrich and Schmidt, 1989).

Habitat

This species can survive in turbid and poorly oxygenated water because of its ability to breathe air. Air breathing is so well developed that this snakehead can travel overland between water bodies, using a hopping motion. Two pharyngeal suprabranchial cavities extend along the body to the caudal peduncle and are lined with vascularized mucous membranes. Mountain streams, rivers, lakes, reservoirs, ponds, rice paddies and even hot springs are recorded as habitats for this species in South Asia. Clear water in shallow streams and swamps in forested areas are preferred (Courtenay and Willimas, 2004) while Pethiyagoda (1991) states that flowing water is preferred. It is found in mud among emergent vegetation in Sri Lanka (De Silva, 1991). It may be largely nocturnal. This species can survive temperatures as low as 13°C, a pH range of 3.1-9.6, brackish water, and waters low in calcium, low in mineral content and pronounced anion excess (Lee and Ng, 1994). Low temperatures may be a limiting factor in its distribution in Iran.

Age and growth

Some populations in mountain streams are mature at about 13 cm while in lower areas maturity is reached at about 10.2 cm at 20 months. The population examined in Sri Lanka has 90% of the individuals less than 24 months old and 99% less than 38 months. Longevity is about 6 years (De Silva, 1991).

Food

This species is recorded as a nocturnal predator on other fishes and on frogs but most diet studies indicate that insects and crustaceans are the main foods (De Silva, 1991; Pethiyagoda, 1991). Young fish feed on unfertilised eggs from the mother for about 4 weeks. These eggs are released and fall in the water (while those which hatch float). The young stimulate egg release through close body contact with the mother, who swims in a circle while releasing the eggs. In aquaria, a female will take Artemia nauplii into its mouth and swim over to the male which releases the young to feed on the nauplii as they are emitted from behind the gill cover (Ettrich and Schmidt, 1989). An Iranian specimen had only sand grains in its gut.

Reproduction

Spawning takes place over silt or gravel bottoms or in areas of cleared vegetation forming a "nest". Vegetation is cleared by fin movements and can be 15 cm across (Ettrich and Schmidt, 1989). Some reports have the female swimming belly up and the male then fertilises the eggs as they are released by swimming diagonally over the female's vent. Ettrich and Schmidt (1989) state that the male forms a loop around the belly region of the female, an intensive and long-lasting process. This occurs after pair-bonding lasting several weeks which serves to synchronise reproduction, necessary since all the eggs are released at once. The male picks up the eggs in his mouth and keeps them there for 4-5 days until hatching (Ettrich, 1989; Pethiyagoda, 1991), as the eggs are oily and slowly float to the surface. The fry may also be protected in the male's mouth for up to three days before releasing them, but retain the fry behind the gill cover when danger threatens or night descends (Ettrich and Schmidt, 1989). Egg numbers vary between 20 and 200 per spawning (Lee and Ng, 1994). Fecundity ranges from 389 and 7194 ((De Silva, 1991; Courtenay and Williams, 2004). Brood size (97-343 larvae) is smaller than the number of mature eggs reported for females according to De Silva (1991) and many eggs either fail to develop or are lost to predators despite parental care. An Iranian fish caught on 6 May contained small and possibly atretic eggs. Egg diameter is 2.6 mm and the eggs are a golden yellow.

Ettrich and Schmidt (1989) report that 6 days after being released from the mouth, the fry ascend to the water surface and draw air. A foam nest is produced under which the young fish hide.

De Silva (1991) found breeding to take place throughout the year in Sri Lanka, with enhanced breeding in May to July and October to December. In the Karnataka State, India this species breeds from May to August. Individuals appear to spawn once in each rainy season in Sri Lanka.

Parasites and predators

None reported from Iran.

Economic importance

This species is too rare in Iran to be of any economic importance but in Sri Lanka it consumes pests in rice paddies (De Silva, 1991) and in Singapore in 1990 sold for up to Singaporean $60 (Courtenay and Willimas, 2004).

Conservation

This species would be difficult to conserve in Iran as it is rare and at its extreme westernmost distribution. Survivability may be marginal and numbers low.

Further work

Surveys to determine the distribution of this species, its numbers and its environmental requirements may enable conservation to be effective.

Sources

Iranian material: ZISP 11714, 4, 42.9-70.2 mm standard length, Baluchestan, Bampur River (no other locality data); CMNFI 1979-0220, 1, 110.9 mm standard length, Kerman, irrigation ditch, 2 km south of Jiroft (= Sabzeveran) (28°39'N, 57°45'E).

Comparative material: BC 66-32, 2, 63.8-71.5 mm standard length, East Pakistan, Chittagong Hill Tracts, Karnaphuli Reservoir tributary (no other locality data); BC 66-38, 23, 79.5-124.9 mm standard length, East Pakistan, Dacca, Bellabor Fish Market (no other locality data); BC 66-44, 2, 109-6-122.3 mm standard length, East Pakistan, Tipperali, Chandpur (no other locality data).

Scophthalmidae

Genus Psetta
Swainson, 1839

Psetta maxima
(Linnaeus, 1758)

Introduced to the Soviet Caspian Sea basin as the subspecies Psetta maxima maeotica (Pallas, 1811) in 1930-1931 but not subsequently observed. The introduction probably failed because the pelagic eggs were not buoyant at the low Caspian salinities (Baltz, 1991). No Iranian record.

Pleuronectidae

Genus Platichthys
Girard, 1854

Platichthys flesus
(Linnaeus, 1758)

Introduced to the Soviet Caspian Sea basin in 1902 and 1930-1931 but not subsequently observed (Shukolyukov, 1937a). The introduction probably failed because the pelagic eggs were not buoyant at the low Caspian salinities (Baltz, 1991). Voronina (1999) is a recent systematic review of this species. No modern Iranian record although some were caught in the Anzali Mordab outlets in the 1930s (Holčík and Oláh, 1992). The Farsi name is pahn mahi.

 


Bibliography
1711-2010

This section contains all the papers and books referred to in the text of the web site on Freshwater Fishes of Iran but also attempts to be a bibliography on these fishes and their environment. Entries ceased at the end of 2010 except for taxonomic and systematic works. The volume of literature on Iranian fishes, and the ease of search and access through the internet, render additions tedious and nugatory.

A partial version of this bibliography was first posted on 10 September 1996 and maintained by Yazdan Keivany, then at the University of Alberta, Edmonton. There has been a rapid increase in studies on fishes of Iran, starting in the 1990s. Prior to 1900, this bibliography lists less than 100 publications relevant to this work, many not strictly on Iranian fishes. On a decadal basis, it is only in the 1960s that publications exceed 100 and by the 1990s are an order of magnitude larger.

The large number of articles listed here, the length of many, their limited availability, and costs involved make it impossible to loan xeroxes except in restricted cases. Please use your interlibrary loan service.

The website http://www.sid.ir (Scientific Information Database or SID, Tehran) appears in Farsi and English and lists publications in Iranian journals, with abstracts in English and Farsi. One journal given is listed as  "Virtual" sometimes without date or pagination and is presumably some form of preliminary listing.

Students of Iranian fishes should also consult the Freshwater Fishes of Iraq website for its Bibliography as this includes additional and complementary studies on species shared with Iran.

There is a moderately extensive grey literature on Iran, mostly manuscript reports of the Department of the Environment, Tehran. Some of these are in English and were written by Peace Corps workers in the 1960s and 1970s. I have cited all those I came across but the listings are by no means exhaustive. Additionally, magazines and newspapers in Iran carry articles on fishes and numerous conference presentations appear as titles, abstracts and even lengthy articles. Regretfully, a complete survey of these was not possible although all those found are included.

The internet now carries English translations of Iranian newspapers and press releases. The URL (Universal Resource Locator) for these and for websites having information on Iran are cited directly in the text and not in this bibliography. Note that links often change or are no longer functional and these textual references serve only to document a source. Generally the date when they were downloaded is given. They may still be accessible but are not hot linked in the text. Some papers cited in this Bibliography have only been seen in their online version and pagination of the original, printed version is unknown. Others are known only from abstracts as costs prevent obtaining all papers when these are marginal to the main purpose of this website - describing the fishes and their general biology. Wherever possible the language of the article is indicated by the title (for European languages including Turkish) or by adding the language at the end. In some cases, particularly for articles published in Iran and seen only in English abstract, the language could be Farsi or English as journals may have articles in both languages.

Transliteration of personal names into English from other languages, notably Farsi, often gives variant spellings. Iranians may spell their name differently in different papers (you know who you are). When I am aware that the same person is meant, the original journal spelling is given followed by (sic) to indicate that the author may appear under another spelling. The same (sic) is also used to denote misspellings in titles (if these are egregious). Note that family names in Farsi may appear in two forms, a single name or a double-barrelled name (separated or hyphenated), and these are variously used in papers. Personal names may be abbreviated to a single letter when appearing in English but sometimes appears as two letters to accord with the Persian alphabet (e.g. there are single letters s and sh).

The "Iranian Journal of Fisheries Sciences" (from 7(2) 1998 onwards) also appeared earlier as the "Iranian Fisheries Scientific Journal" (3(2) 1994 to 7(1) 1998), and earlier still as "Iranian Fisheries Journal", "Iranian Fisheries Scientific Bulletin" and "Iranian Fisheries Bulletin", is published in Farsi with English abstracts. It now appears as the "Iranian Scientific Fisheries Journal" in Iranian abstracting journals and websites. The "Iranian Journal of Fisheries Sciences" (starting in January 1999 with volume 1(1)) is published in English with Farsi abstracts. The Iranian Fisheries Research Organization (formerly Iranian Fisheries Research and Training Organization) publishes these journals and has a website with lists of publications.

Note that articles published in the "Journal of Ichthyology" are translations from the original Russian journal "Voprosy Ikhtiologii" although pagination (and sometimes issue number) are not the same. Only "Journal of Ichthyology" pagination is cited here but readers may wish to avail themselves of the Russian versions of these articles (however some earlier issues of "Voprosy Ikhtiologii" are only available in Russian). Prior to 1970 the English translation was called "Problems of Ichthyology". Year of publication as marked on the cover of each issue was the one cited here although internal evidence indicates translated issues were often published in the following year, a delay presumably because of translation time required. The date thus used is the Russian date of publication which is the relevant one for the few taxon descriptions appearing in this journal.

The Journal of the Faculty of Veterinary Medicine, University of Tehran contains many articles on aquaculture in Iran and other articles on Iranian fishes generally. It sometimes appears in abstracting services, and in this bibliography, as Journal of Veterinary Research.

Some articles may be identified by journal name and a DOI but not pagination. DOI is the Digital Object Identifier and is found, in these cases, on articles that appear on-line but are not yet in print. Eventually volume, issue and pages replace the DOI herein. Occasionally, some articles seem to languish as a DOI but for most the journal pagination follows within a few months.

A

Abaee, S. 2001. Rapid assessment of point sources pollution in Iranian part of Caspian Sea area using GIWA methodology. Caspian Environment Programme, Baku, Azerbaijan and Tehran. 8 pp., 3 figs., appendices A-B.

Abakumov, V. A. 1960. The damage done by lampreys to fish stocks. Fisheries Research Board of Canada Translation Series, 274:1-2.

Abassali Zadeh, A. 2003. Effects of ablading of sensory barbels on growth of Huso huso (young stage). Iranian Journal of Fisheries Sciences, 12(3):193-200. In Farsi.

Abaychi, J. and Al-Saad, H. T. 1988. Trace elements in fish from the Arabian Gulf and the Shatt al-Arab River, Iraq. Bulletin of Environmental Contamination and Toxicology, 40:226-232.

Abaychi, J. K., Darmoian, S. A. and DouAbul, A. A. Z. 1988. The Shatt al-Arab River: A nutrient salt and organic matter source to the Arabian Gulf. Hydrobiologia, 166:217-224

Abbasi, H. 1974. Neur Lake experimental commercial fishery, 27 Mehr - 16 Aban. L-6-53, Job Progress Report, Department of the Environment, Tehran. 9 pp. MS.

Abbasi, H. 1975. Neur Lake commercial fishery, 29 Mehr - 19 Aban 1354. Job Progress Report, Fisheries and Aquatic Ecology Section, Department of the Environment, Tehran. MS (Abstract).

Abbasi, K. 2003. First record of black carp, Mylopharyngodon piceus from southern Caspian Sea (Iran). Iranian Journal of Fisheries Sciences, 12(2):139-146. In Farsi.

Abbasi, K. 2005. Studying alien fishes and macrocrustaceans distribution and their effects on rivers and wetlands of the Iranian basin of Caspian Sea. Borok II, International Workshop, Invasions of Alien Species in Holarctic, 27 September-1 October 2005, Borok (abstract).

Abbasi, K. 2006a. Identification and distribution of fish fauna in Shafarud River, Guilan Province. Iranian Journal of Fisheries Sciences, 15(2):73-86. In Farsi.

Abbasi, K. 2006b. Identification and distribution of fish fauna in Hevigh River (Guilan Province). Iranian Journal of Biology, 18(4):370-382. In Farsi.

Abbasi, K., Keyvan, A. and Ahmadi, M. R. 2004. Morphometric and meristic characteristics of Vimba vimba persa in Sefidrud River. Iranian Journal of Fisheries Sciences, 13(1):61-76. In Farsi.

Abbasi, K., Keyvan, A. and Ahmadi, M. R. 2005. Studying natural reproduction, spawning grounds and spawning period of Vimba vimba persa population in Sefid-Roud River, Guilan Province in north Iran. Iranian Journal of Fisheries Sciences, 14(3):113-126. In Farsi.

Abbasi, K., Moradkhah, S. and Sarpanah, A. N. 2007. Identification and distribution of fish fauna in Siahdarvishan River (Anzali Wetland basin). Pajouhesh va Sazandegi, 19(1)(74):27-39. In Farsi.

Abbasi, K. and Sabkara, J. 2004a. Studying food items of Leuciscus ulanus, an endemic fish species in Iran. The 1st Congress on Animal and Aquatic Sciences, University of Tehran, 31 August - 2 September 2004, p. 492-495.

Abbasi, K. and Sabkara, J. 2004b. Studying some biological characteristics of Leuciscus ulanus (Cyprindae), an endemic fish and little-known in Iran. 2nd International Conference on Applied Biology, Mashad, 29-30 September 2004, p. 29.

Abbasi, K. and Sabkara, J. 2004c. Studying Alosa caspia caspia (Caspian shad) diet in southeastern Caspian Sea (Mazandaran and Golestan provinces). Iranian Journal of Biology, 17(3):272-290. In Farsi.

Abbasi, K., Salavatian, S. M. and Abdollapoor, H. 2005. Investigating fish diversity and distribution in the Mahabad-Chai River of the Lake Urmia basin, north-western Iran. Iranian Journal of Fisheries Sciences, 13(4):75-94. In Farsi.

Abbasi, K. and Sarpanah, A. 2001. Fish fauna investigation in Aras Reservoir and its Iranian tributaries. Iranian Journal of Fisheries Sciences, 10(2):41-62. In Farsi.

Abbasi, K., Valipour, A., Talebi Haghighi, D., Sarpanah, A. and Nezami, Sh. 1999. Atlas of Iranian Fishes. Gilan Inland Waters. Gilan Fisheries Research Centre, Rasht. vi + 113 pp. In Farsi.

Abbasi, K. and Valipour, A. R. 2005. Studying the Silurus glanis Linnaeus, 1758 food items in Anzali lagoon. Pajouhesh va Sazandegi, 17(1)(66):14-24. In Farsi.

Abbasi, M. and Gharezi, A. 2008. Morphology and histology of the digestive tract of Iranian blind cave fish (Iranocypris typhlops). Iranian Veterinary Journal, 4(2)(19):60-69. In Farsi.

Abbasian, G-R. 1997. Fisheries of Iran: Development possibilities. INFOFISH International, Kuala Lumpur, 2:14-18.

Abbasov, G. S. 1980. The ichthyofauna of the main freshwater bodies of Azerbaidzhan. Journal of Ichthyology, 20(4):140-142.

Abbaspour, M. and Sabetraftar, A. 2005. Review of cycles of drought and indices of drought and their effect on water resources, ecological, biological, agricultural, social and economical issues in Iran. International Journal of Environmental Studies, 62(6):709-724.

Abd, I. M., Rubec, C. and Coad, B. W. 2009. Key biodiversity areas - Rapid assessment of fish fauna in southern Iraq, p. 161-171. In: Krupp, F., Musselman, J. L., Kotb, M. M. A. and Weidig, I. (Eds.). Environment, Biodiversity and Conservation in the Middle East, Proceedings of the First Middle Eastern Biodiversity Congress, Aqaba, Jordan, 20-23 October 2008. BioRisk 3(Special Issue):219 pp. ISSN 1313-2652 (online), ISSN 1313-2644 (print). Pensoft Publishers, Sofia-Moscow (http://pensoftonline.net/biorisk/index.php/journal/article/view/15/27).

Abd Almaleki, Sh and Ghaninezhad, D. 2007. Stock assessment of the Caspian Sea kutum (Rutilus frisii kutum) in Iranian coastal waters of the Caspian Sea. Iranian Scientific Fisheries Journal, 16(1):103-114. In Farsi.

Abd Elahpour Biria, H., Abbasi, K., Sarpanah, A. N. and Pourgholami Moghadam, A. 2009. Population of sand goby (Neogobius fluviatilis pallasi) in southwest coasts of the Caspian Sea using morphological characteristics. Iranian Scientific Fisheries Journal, 18(2):81-90. In Farsi.

Abdi, K. 1999a. Survey on the four new species of parasitic leeches of Mahabad's Lake Dam in Iran. In: 26th World Veterinary Congress, Lyon, France, 23-26 September 1999 (abstract).

Abdi, K. 1999b. Survey on the crustacean parasites of fishes of Mahabad's reservoir dam with new records of Tracheliastes polycolpus in Iran. In: 26th World Veterinary Congress, Lyon, France, 23-26 September 1999 (abstract).

Abdi, K., Jalalai, B. Moobedi, I. and Naem, S. 1995. Identification of crustacean parasites in Mahabad Reservoir. Pajouhesh va Sazandegi, 36:128-132. In Farsi.

Abdi, K., Mobedi, I. and Rostamzad, R. 2006. The first outbreak of acanthocephalosis in rainbow trout fish farms of Iran. Pajouhesh va Sazandegi, 18(2)(71):9-12. In Farsi.

Abdolhay, H. 1996. Aquaculture status and development in the Islamic Republic of Iran. Paper presented to the Working Group on Aquaculture, Indian Ocean Fishery Commission Committee for the Development and Management of the Fishery Resources of the Gulf, Cairo, Egypt, 1-3 October, 1996.

Abdolhay, H. 1997a. Artificial reproduction of fish for stock enhancement in the Caspian Sea. Seventh Conference of Shilat, Responsible Fisheries, 17-18 February 1997, Tehran. p. 187-207. In Farsi.

Abdolhay, H. 1997b. Fingerling production and stock enhancement of sturgeon in south of Caspian Sea. Third International Symposium on Sturgeon, Piacenza, Italy, 8-11 July 1997, Book of Abstracts (poster).

Abdolhay, H. and Baradaran Tahori, H. 1998. Review of reproduction of sturgeon from 1991-1997 in Iranian hatchery. First National Symposium on Sturgeon, Rasht, 15-16 December 1998, p. 107-124. In Farsi.

Abdolhay, H. and Baradaran Tahori, H. 1999. Fingerling production and stock enhancement of sturgeon in south of Caspian Sea. Journal of Applied Ichthyology, 15(4-5):298.

Abdolhay, H. and Baradaran Tahori, H. 2006. Fingerling production and release for stock enhancement of sturgeon in the southern Caspian Sea: an overview. Journal of Applied Ichthyology, 22(s1):125-131.

Abdolhay, H., Baradaran Tahouri, H. and Amini, R. 2006. An audit of sturgeon reproduction in Iran over the years 1998-2002. Iranian Scientific Fisheries Journal, 14(4):97-112. In Farsi.

Abdolhay, H. A., Daud, S. K., Pourkazemi, M., Siraj, S. S., Rezvani, S., Mostafa, K. A. S. and Hosseinzadeh Sahafi, H. 2010. Morphometric studies of mahisefid (Rutilus frisii kutum, Kamensky, 1901) from selected rivers in the southern Caspian Sea. Iranian Journal of Fisheries Sciences, 9(1):1-18.

Abdoli, A. 1992. A new fish species in Mazandaran ab bandans. Abzeeyan, Tehran, 2(8):54-55. In Farsi.

Abdoli, A. 1993a. Threespine stickleback in the Caspian Sea. Abzeeyan, Tehran, 3(10 & 11):45. In Farsi.

Abdoli, A. 1993b. Fishes of Golestan National Park's river. Abzeeyan, Tehran, 4(3):32-33, XIII. In Farsi.

Abdoli, A. 1993c. Three-spined stickleback in southern basins of the Caspian Sea. Abzeeyan, Tehran, 4(5):10-13, III. In Farsi.

Abdoli, A. 1994a. Tadpole goby Benthophilus stellatus leobergius in Gorgan Bay. Abzeeyan, Tehran, 4(12):58-59. In Farsi.

Abdoli, A. 1994b. Ecological study of fish populations in Sardabrud River and Chalus River, Mazandaran, Iran. M.Sc. Thesis, University of Tehran. iv + 94 pp, 28 figs, 20 tables.

Abdoli, A. 1998a. An investigation of some biological characteristics and distribution of Pseudorasbora parva in freshwaters of Iran. Journal of Agricultural Sciences and Natural Resources, 8(1):33-40. In Farsi.

Abdoli, A. 1998b. Fish species status in southern Caspian Sea. First International Student Seminar of Coastal University of the Caspian Sea, Astrakhan, Russia (title).

Abdoli, A. 2000. The Inland Water Fishes of Iran. Iranian Museum of Nature and Wildlife, Tehran. 378 pp. In Farsi.

Abdoli, A., Allahyari, S., Kiabi, B. H., Patimar, R., Ghelichi, A., Mostafavi, H., Aghili, S. M. and Rasooli, P. 2009. Length-weight relationships for seven Gobiid fish species in the southeastern Caspian Sea basin, Iran. Journal of Applied Ichthyology 25(6):785-786.

Abdoli, A., Coad, B. and Naderi, M. 2000. First record of Rhinogobius similis, Gill 1859 in Iran. Iranian Journal of Fisheries Sciences, 9(1):73-76, 9. In Farsi.

Abdoli, A. Ghorbani, S., Niksirat, H., Mashhadi Ahmadi, A. and Golzarianpour, K. 2010. Iranian Wild Nature, Brown Trout. Plan for the Land, Tehran (pamphlet) In Farsi.

Abdoli, A., Golzarianpoor, K., Kiabi, K. H. and Patimar, R. 2010. Status of the endemic loaches of Iran. International Loach Conference 2010, 31 August to 3 September 2010, Prague, Czech Republic (abstract).

Abdoli, A., Kiabi, B., Patimar, R. and Allahyari, S. 2008. Study of species diversity of fishes of Gomishan wetland in south east of Caspian Sea. International Conference on Biodiversity Conservation and Management, University of Science and Technology, Cochin, Kerala, India, 3-6 February 2008 (abstract).

Abdoli, A. and Naderi, M. 2009. Biodiversity of Fishes of the Southern Basin of the Caspian Sea. Abzian Scientific Publications, Tehran. 243 pp. In Farsi.

Abdoli, A., Naderi, M., Aboo, M., Fazli, H. and Afraee, M. A. 1996. Spawning time and fecundity of the mullet, Liza auratus (sic) in south eastern part of the Caspian Sea. Abzeeyan, Tehran, 7(2):24-26, III. In Farsi.

Abdoli, A., Niksirat, H. and Patimar, R. 2011. Variation of some biological characteristics of Caspian brown trout in southern Caspian Sea. The 4th International Symposium on Stock Enhancement and Sea Ranching, 9th Asian Fisheries and Aquaculture Forum, Shanghai Ocean University, April 21 to 23, 2011 (abstract).

Abdoli, A., Patimar, R. Mostafavi, H., Kiabi, B. H. and Rasooli, P. 2007. Feasibility of Le Cren Method for population estimation of freshwater fish in Iran. Fish Stock Assessment Methods for Lakes and Reservoirs: Towards the True Picture of Fish Stock, 11-15 September 2007, Ceske Budejovice, Czech Republic (poster).

Abdoli, A and Rahmani, H. 1999. The populations, food consumption and reproduction of Neogobius fluviatilis in a small stream of Madarso, Golestan National Park, North East of Iran. Seventh International Symposium on the Ecology of Fluvial Fishes, 10-13 May 1999, Lodz, Poland (poster).

Abdoli, A. and Rahmani, H. 2001. Food habits of two species of Gobiidae Neogobius fluviatilis, Neogobius melanostomus in the Madarsoo stream, Golestan National Park. Journal of Agricultural Sciences and Natural Resources, 8(1):3-15, 191. In Farsi.

Abdoli, A., Rahmani, H. and Rasooli, P. 2002. On the occurrence, diet and reproduction of Neogobius fluviatilis in Madarsoo stream, Golestan National Park, (north eastern Iran). Zoology in the Middle East, 26:123-128.

Abdoli, A., Rasooli, P. and Mostafavi, H. 2008. Length-weight relationships of Capoeta capoeta capoeta (Gueldenstaedt, 1772) in the Gorganrud River, south Caspian basin. Journal of Applied Ichthyology, 24(1):96-98.

Abdoli, A., Rasooli, P. and Soltaninasab, S. 2008. A contribution to the biology of Acanthalburnus urmianus (Günther, 1899) (Osteichthyes: Cyprinidae): an endemic fish of Iran. Zoology in the Middle East, 43:111-112.

Abdoli, A., Rasooli, P., Yazdandad Bibalan, H. and Abdoli, L. 2007. A study on some ecological aspects of snow trout (Schizothorax pelzami) from Laiinsoo River in northeastern Iran. Environmental Sciences, 4(3):69-76.

Abdoli, A. and Skandari, S. K. 1999. Reproductive biology of khramulya (Capoeta capoeta gracilis) in Madarso stream in Golestan National Park. Journal of Agricultural Sciences and Natural Resources, 6(3):38-51. In Farsi.

Abdollapour Bereya, H., Abbasi, K., Sabkara, J. and Keyvan, A. 2007. Studying Caspian shad (Alosa caspia caspia) diet in southwest coastal area of the Caspian Sea, Guilan Province waters. Iranian Scientific Fisheries Journal, 16(1):115-128. In Farsi.

Abdolmalaki (= Abd Almaleki), Sh. 2000. Survey of parasites of fishes of Makoo Reservoir. Department of Fisheries and Sciences, Iran. 21 pp. In Farsi.

Abdolmalaki, Sh. 2001. Catch statistic review of mullet fishery in Iranian coastal waters of the Caspian Sea. Iranian Journal of Fisheries Sciences, 9(4):39-52. In Farsi.

Abdolmalaki, Sh. 2004. Fishery status and stock assessment of silver carp (Hypophthalmichthys molitrix) in Mahabad Reservoir in fishing season of 1998-1999. Iranian Journal of Fisheries Sciences, 13(1):77-96. In Farsi.

Abdolmalaki, Sh. 2005a. Stock assessment of the pick (sic) perch (Stizostedion lucioperca) in the southern coastal waters of the Caspian Sea. Iranian Journal of Fisheries Sciences, 13(4):95-110. In Farsi.

Abdolmalaki, Sh. 2005b. Population dynamics of Caspian sea bream (Abramis brama orientalis) in Iranian coast of the Caspian Sea in 2000-2001. Pajouhesh va Sazandegi, 18(3)(68):85-92. In Farsi.

Abdolmalaki, Sh. 2006a. Stock assessment of bonyfishes in the Iranian coastal water of the Caspian Sea. Iranian Fisheries Research Organization, Agriculture Research and Education Organization, Ministry of Jihad-e-Agriculture, Bandar Anzali. http://en.ifro.ir (abstract).

Abdolmalaki, Sh. 2006b. Trends in stock fluctuations of Rutilus frisii kutum in the Caspian Sea. Iranian Journal of Fisheries Sciences, 15(2):87-100. In Farsi.

Abdolmalaki, Sh., Amirkhani, A., Borani, M., Ghaninejad, D., Rastin, R., Porgholami, A. and Moradkhah, S. 1998. A survey on catch status and sexual maturity of mullets in Iranian coastal water of the Caspian Sea (Gilan Province) in October 1998. Research Report, Fisheries Research Center of Gilan Province, Bandar Anzali. 13 pp. In Farsi.

Abdolmalaki, Sh. and Ghaninezhad, D. 1999. A survey on releasing fingerlings and catching pikeperch (Stizostedion lucioperca) in Iranian coastal waters of the Caspian Sea. Proceedings of the First National Congress on Caspian Sea Bony Fish, 21-23 January 1999, Bandar Anzali, Gilan. In Farsi.

Abdolmalaki, Sh. and Ghaninezhad, D. 2007a. Stock assessment of the Caspian kutum Rutilus frisii kutum in the Iranian coastal waters of the Caspian. Sea. Iranian Journal of Fisheries Sciences, 16(1):103-114.

Abdolmalaki, Sh. and Ghaninezhad, D. 2007b. Releasing fry of kutum (Rutilus frisii kutum) for restocking in Iranian waters of the Caspian Sea. Abzeeyan, 36:8-13. In Farsi.

Abdolmalaki, Sh. and Psuty, I. 2007. The effects of stock enhancement of pikeperch (Sander lucioperca) in Iranian coastal waters of the Caspian Sea. ICES Journal of Marine Sciences, 64:8 pp. (advance copy).

Abdolmalaki, Sh., Sadat, H. A. and Nahrevar, R. 2007. Catch status and population structure of kutum (Rutilus frisii kutum) in Iranian coastal water of the Caspian Sea. Journal of Marine Sciences and Technology, 6(3-4):51-62.

Abdolmaleki, S. 1994. An investigation on the bentic (sic) macrofauna of the Anzali Lagoon. Iranian Fisheries Scientific Bulletin, (5):27-38, 4. In Farsi.

Abdurakhmanov, Yu. A. 1962. Ryby Presnykh vod Azerbaidzhana [Freshwater Fishes of Azerbaidzhan]. Akademii Nauk Azerbaidzhanskoi SSR, Institut Zoologii, Baku. 407 pp.

Abdurrahmanov, Y. A. 1966. Azarbaycanin Faunasi. VII Cild. Baliglar (Pisces). [The Fauna of Azerbaidzhan. VII. Fishes (Pisces)]. Zoologiya Institutu, Azarbaycan SSR Elmlar Akademiyasi, Baki. 224 pp. In Azerbaijani.

Abdurakhmanov, Yu. A. 1975. Transformation of the diadromous Kura shemaya Chalcalburnus chalcoides into a land-locked population in the Mingechaur Reservoir. Journal of Ichthyology, 15(2):189-196.

Abdurakhmanov, Yu. A., Kuliev, Z. M. and Agayarova, A. E. 1968. Materialy po Biologii i Raspredeleniyu Ryb u Azerbaidzhanskogo Poberezh'ya Srednego i Yuzhnogo Kaspiya [Materials on the biology and distribution of fishes at the Azerbaidzhan coast of the Central and Southern Caspian]. In: The Biology of the Central and Southern Caspian. Moscow. pp. 113-146.

Abdurakhmanov, Yu. I. and Kuliyev, Z. M. 1968. European eel in Azerbaidzhan waters. Problems of Ichthyology, 8:592-594. In English.

Abdusamadov, A. S. 1986. Biology of white amur, Ctenopharyngodon idella, silver carp, Hypophthalmichthys molitrix, and bighead, Aristichthys nobilis, acclimatized in the Terek region of the Caspian basin. Journal of Ichthyology, 26(4):41-49.

Abedi, M., Shiva, A. H., Mohammadi, H. and Malekpour, R. 2010. Reproductive biology and age determination of Garra rufa Heckel, 1843 (Actinopterygii: Cyprinidae) in central Iran. Turkish Journal of Zoology, 35(3):317-323.

Abedian Kennari, A. M., Oveisipour, M. R. and Nazari, R. M. 2007. Effects of n3-HUFA enriched Daphnia magna on growth, survival, stress resistance, and fatty acid composition of larvae of Persian sturgeon (Acipenser persicus). Iranian Journal of Fisheries Sciences, 7(1):1-14.

Abell, R., Thieme, M. L., Revenga, C., Bryer, M., Kottelat, M., Bogutskaya, N., Coad, B., Mandrak, N., Contreras Balderas, S., Bussing, W., Stiassny, M. L. J., Skelton, P., Allen, G. R., Unmack, P., Naseka, A., Ng, R., Sindorf, N., Robertson, J., Armijo, E., Higgins, J. V., Heibel, T. J., Wikramanayake, E., Olson, D., López, H. L., Reis, R. E., Lundberg, J. G., Sabaj Pérez, M. H. and Petry, P. 2008. Freshwater Ecoregions of the World: A new map of biogeographic units for freshwater biodiversity conservation. BioScience, 58(5):403-413.

Abich, H. 1856. Vergleichende Chemische Untersuchungen der Wasser des Caspischen Meeres, Urmia- und Van-See's. Mémoires de l'Académie Impériale des sciences de Saint-Pétersbourg, sixième série, Sciences mathématiques et physiques, 7:57 pp., 2 plates.

Abou-Seedo, F., Dadzie, S., Al-Kanaan, K. and Sukumaran, J. V. 2003. Aspects of reproductive biology of the hermaphroditic yellowfin seabream, Acanthopagrus latus (Hottuyn (sic), 1782), in cages in Kuwait Bay. Zoology in the Middle East, 29:51-58.

Abtahi B., Esmaili Sari, A., Khodabandeh, S. and Seifabadi J. 2001. Ctenophores of the Caspian Sea and their socio-economic impacts. The Second International Iran and Russia Conference (Agriculture & Natural Resources), Moscow. Abstract, p. 164.

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Coad, B. W. 1979b. Poisonous and venomous freshwater fishes of Iran. Pahlavi Medical Journal, 9(4):388-407.

Coad, B. W. 1980a. A provisional, annotated check-list of the freshwater fishes of Iran. Journal of the Bombay Natural History Society, 76(1)(1979):86-105.

Coad, B. W. 1980b. A re-description of Aphanius ginaonis (Holly, 1929) from southern Iran (Osteichthyes: Cyprinodontiformes). Journal of Natural History, 14(1):33-40.

Coad, B. W. 1980c. Environmental change and its impact on the freshwater fishes of Iran. Biological Conservation, 19(1):51-80.

Coad, B. W. 1981a. Glyptothorax silviae, a new species of sisorid catfish from southwestern Iran. Japanese Journal of Ichthyology, 27(4):291-295.

Coad, B. W. 1981b. First record of the milkfish, Chanos chanos (Forskal, 1775) from Iran and the Persian Gulf. Journal of the Bombay Natural History Society, 77(3)(1980):522-524.

Coad, B. W. 1981c. A new genus and species of cichlid endemic to southern Iran. Program of the American Society of Ichthyologists and Herpetologists, 61st Annual Meeting, 21-26 June 1981, Corvallis, Oregon. 1 p. (Abstract).

Coad, B. W. 1981d. Fishes of Afghanistan, an annotated checklist. Publications in Zoology, National Museums of Canada, 14:v + 26 pp.

Coad, B. W. 1981e. Pseudophoxinus persidis, a new cyprinid fish from Fars, southern Iran. Canadian Journal of Zoology, 59(11):2058-2063.

Coad, B. W. 1981f. A bibliography of the sticklebacks (Gasterosteidae: Osteichthyes). Syllogeus, Ottawa, 35:1-142.

Coad, B. W. 1982a. A new genus and species of cichlid endemic to southern Iran. Copeia, 1982(1):28-37.

Coad, B. W. 1982b. Review of "Euphrates and Tigris, Mesopotamian Ecology and Destiny" by Julian Rozska (sic) with contributions by J. F. Talling, F. R. S. and Dr. K. E. Bainster (sic). Monographiae Biologicae, Volume 38, Dr. W. Junk by (sic) Publishers, The Hague, 1980. x + 122 pp., 36 figs., map.-U.S. $33.31. Matsya, Bulletin of the Indian Society of Ichthyologists, 7(1981):102-104.

Coad, B. W. 1982c. Garra persica Berg, 1913, a valid species of cyprinid fish from southern Iran. Cybium, 6(2):97-100.

Coad, B. W. 1982d. The identity of Alburnus maculatus Keyserling, a cyprinid fish from Esfahan Province, Iran. Japanese Journal of Ichthyology, 29(2):227-228.

Coad, B. W. 1982e. Studies on the systematics and zoogeography of the freshwater fishes of Iran. Programme of the Fourth Congress of European Ichthyologists, 20.-24. 9. 1982, Hamburg, West Germany. 1 p. (Abstract).

Coad, B. W. 1982f. A re-description and generic re-assignment of Kosswigobarbus kosswigi (Ladiges, 1960), a cyprinid fish from Turkey and Iran. Mitteilungen aus dem hamburgischen Zoologischen Museum und Institut, 79:263-265.

Coad, B. W. 1983. Review of "The Freshwater Fishes of India, Pakistan, Bangladesh, Burma and Sri Lanka - A Handbook". By K. C. Jayaram. 1981. Zoological Survey of India, Calcutta. xxii + 475 pp., 208 figs., 13 plates (2 color). Rs 100.00 or $22.50. Copeia, 1983(1):280-282.

Coad, B. W. 1984. Acanthobrama centisquama Heckel and the validity of the genus Mirogrex Goren, Fishelson and Trewavas (Osteichthyes: Cyprinidae). Hydrobiologia, 109(3):275-278.

Coad, B. W. 1985. Alburnus doriae De Filippi, 1864 a synonym of Leuciscus lepidus (Heckel, 1843)(Osteichthyes: Cyprinidae). Matsya, 9-10(1983-1984):173-175.

Coad, B. W. 1987a. The tooth-carps of Iran. Program and Abstracts, American Society of Ichthyologists and Herpetologists, 67th Annual Meeting, 21-26 June 1987, Albany, New York. p. 39 (Abstract).

Coad, B. W. 1987b. Zoogeography of the Freshwater Fishes of Iran, pp. 213-228, 1 figure, 2 tables. In: Krupp, F., Schneider, W. and Kinzelbach, R. (Eds.). Proceedings of the Symposium on the Fauna and Zoogeography of the Middle East, Mainz, 1985. Beihefte zum Tübinger Atlas des Vorderen Orients, Reihe A (Naturwissenschaften), 28, Dr. Ludwig Reichert Verlag, Wiesbaden, 338 pp.

Coad, B. W. 1988. Aphanius vladykovi, a new species of tooth-carp from the Zagros Mountains of Iran (Osteichthyes: Cyprinodontidae). Environmental Biology of Fishes, 23(1-2):115-125. (Reprinted with same pagination in McAllister, D. E. and Kott, E. (Eds.). On lampreys and fishes: a memorial anthology in honor of Vadim D. Vladykov. Developments in Environmental Biology of Fishes, 8:1-162, Kluwer Academic Publishers, Dordrecht/Boston/London).

Coad, B. 1989. Tooth-carps and Tangs or Orogeny and Speciation in the Zagros Mountains of Iran. Research Results Conference, National Museum of Natural Sciences, Ottawa, 2-3 October 1989. 2 pp. (Abstract).

Coad, B. W. 1991a. The qanat ichthyofauna of Iran. Research Results CONFERENCE des résultats de recherche 1991, Canadian Museum of Nature / Musée canadien de la nature, Ottawa, March 21, 1991 / le 21 mars 1991. p. 27 (Abstract).

Coad, B. W. 1991b. Fishes of the Tigris-Euphrates Basin: A Critical Checklist. Syllogeus, Ottawa, 68:1-49.

Coad, B. W. 1992a. Marine Fishes of the Persian Gulf and Sea of Oman. Checklist and Bibliography. Syllogeus, Ottawa, MS. 214 pp.

Coad, B. W. 1992b. Out of Africa? The large-scale barbs of Southwest Asia. Collections and Research CONFERENCE des collections et recherche 1992, Canadian Museum of Nature / Musée canadien de la nature, Ottawa, February 18, 1992 / le 18 février 1992. p. 28. (Abstract).

Coad, B. W. 1992c. Zoogeography of the fishes of Mesopotamia. Collections and Research CONFERENCE des collections et recherche 1992, Canadian Museum of Nature / Musée canadien de la nature, Ottawa, February 18, 1992 / le 18 février 1992. p. 30. (Abstract).

Coad, B. W. 1992d. Environmental change and its impact on the freshwater fishes of Iran, pp. 103-104. In: Trussell, D. (Ed.). The Social and Environmental Effects of Large Dams. Volume III. A Review of Literature. Wadebridge Ecological Centre, Camelford, U.K. xiv + 243 pp. (abstract of Coad (1980c)).

Coad, B. W. 1993. Expedition Field Techniques. Fishes. Expedition Advisory Centre, Royal Geographical Society, London. iv + 82 pp., 25 illustrations.

Coad, B. W. 1994a. Exotic and transplanted fishes in Southwest Asia. VIII Congress Societas Europaea Ichthyologorum, Oviedo, Spain, September 26 to October 2, 1994, p. 20 (abstract).

Coad, B. W. 1994b. Fishes from the qanats of Iran. VIII Congress Societas Europaea Ichthyologorum, Oviedo, Spain, September 26 to October 2, 1994, p. 75 (abstract).

Coad, B. W. 1994c. Criteria for assessing the conservation status of taxa. Science Forum Scientifique, Canadian Museum of Nature / Musée canadien de la nature, Ottawa, November 21 Novembre 1994, p. 35 (abstract).

Coad, B. W. 1994d. Exotic and transplanted fishes in Southwest Asia. Science Forum Scientifique, Canadian Museum of Nature / Musée canadien de la nature, Ottawa, November 21 Novembre 1994, p. 36 (abstract).

Coad, B. W. 1995a. Freshwater Fishes of Iran. Acta Scientiarum Naturalium Academiae Scientiarum Bohemicae, Brno, 29(1):1-64.

Coad, B. W. 1995b. Expedition Field Techniques: Fishes. Expedition Advisory Centre, Royal Geographical Society, London. 2nd Edition. v + 97 pp., 27 illustrations.

Coad, B. W. 1996a. Systematics of the tooth-carp genus Aphanius Nardo, 1827 (Actinopterygii: Cyprinodontidae) in Fars Province, southern Iran. Biologia, Bratislava, 51(2):163-172.

Coad, B. W. 1996b. Systematics of the shah mahi, Chalcalburnus chalcoides (Güldenstädt, 1772), in the southern Caspian Sea basin (Actinopterygii: Cyprinidae). Zoology in the Middle East, 12:65-70.

Coad, B. W. 1996c. Threatened fishes of the world: Iranocypris typhlops Bruun and Kaiser, 1944 (Cyprinidae). Environmental Biology of Fishes, 46(4):374.

Coad, B. W. 1996d. Freshwater fishes of Iranian and Pakistani Baluchistan. Second Symposium on Fish and Fisheries of Pakistan, November, 25, 26, 1996, Department of Zoology, Government College, Lahore, p. 25-26 (abstract).

Coad, B. W. 1996e. Zoogeography of the fishes of the Tigris-Euphrates Basin. Zoology in the Middle East, 13:51-70.

Coad, B. W. 1996f. Exotic fish species in the Tigris-Euphrates basin. Zoology in the Middle East, 13:71-83.

Coad, B. W. 1996g. Fishes from the qanats of Iran. Publicaciones Especiales Instituto Español de Oceanografía, 21:63-79.

Coad, B. W. 1996h. Exotic and transplanted fishes in Southwest Asia. Publicaciones Especiales Instituto Español de Oceanografía, 21:81-106.

Coad, B. W. 1997a. Status of shad in Iranian waters? Shad Journal, 2(2):3.

Coad, B. W. 1997b. Systematic biodiversity in the freshwater fishes of Iran. Ninth International Congress of European Ichthyologists (CEI9) "Fish Biodiversity". Italy 1997 (Napoli-Trieste). Book of Abstracts, p. 25.

Coad, B. W. 1997c. Freshwater fishes of Iranian and Pakistani Baluchistan. Biologia, Lahore, 42(1 & 2)(1996):1-18.

Coad, B. W. 1997d. Systematic biodiversity in the freshwater fishes of Iran. Science FORUM scientifique, Canadian Museum of Nature / Musée canadien de la nature, Ottawa, November 20 novembre, 1997, p. 27 (abstract).

Coad, B. 1997e. Shad in Iranian waters. The Shad Journal, Seattle, 2(4):4-8.

Coad, B. W. 1998a. Threatened fishes of the world: Lebias ginaonis (Holly, 1929)(Cyprinodontidae). Environmental Biology of Fishes, 51(3):284.

Coad, B. W. 1998b. Lebias ginaonis (Holly, 1929). Holly's pupfish. Killi Kontakt, Wommelgem, 4 pp., 1 figure.

Coad, B. W. 1998c. Description of Barbus sublimus Coad and Najafpour, 1997, 2 pp., 2 figures. In: Berrebi, P. (Ed.). Atlas Barbus on the Internet, Université Montpellier, France (http://162.38.181.13/webbarbus/).

Coad, B. W. 1998d. Description of Barbus grypus Heckel, 1843, 2 pp., 2 figures. In: Berrebi, P. (Ed.). Atlas Barbus on the Internet, Université Montpellier, France (http://162.38.181.13/webbarbus/).

Coad, B. W. 1998e. Systematic biodiversity in the freshwater fishes of Iran. Italian Journal of Zoology, 65 (Supplement):101-108. (Proceedings of the Ninth Congress of European Ichthyologists (CEI-9) "Fish Biodiversity" organised in Naples at the University Federico II and held in Trieste - Italy, 24-30 August 1997).

Coad, B. W. 1998f. Expedition Field Techniques: Fishes. Expedition Advisory Centre, Royal Geographical Society, London. 2nd Revised Edition. v + 97 pp., 27 illustrations. ISBN 0-907649-71-8.

Coad, B. W. 1999a. Researchs and scientists. Brian W. Coad. Iran Nature and Wildlife Magazine, Farvadin 1378(3):2 pp., 2 figures. In Farsi and English.

Coad, B. W. 1999b. (Fish). i. Freshwater fishes, p. 655-669. In: Yarshater, E. (Ed.). Encyclopædia Iranica. Bibliotheca Persica Press, New York. Volume IX, Fascicle 6. Festivals VIII-Fish.

Coad, B. W. 1999c. Verspreiding van Aphanius-soorten in Iran. Killi-Kontakt, Wommelgem, 27(4):102-110. In Flemish.

Coad, B. W. 2000a. Criteria for assessing the conservation status of taxa (as applied to Iranian freshwater fishes). Biologia, Bratislava, 55(5):539-557.

Coad, B. W. 2000b. Distribution of Aphanius species in Iran. Journal of the American Killifish Association, 33(6):183-191.

Coad, B. W. 2000c. Aphanius ginaonis (Holly, 1929). Holly's pupfish. Journal of the American Killifish Association, 33(6):192-194.

Coad, B. W. 2003. Freshwater fishes, 15 pp., 20 figures, 2 tables. In: Yarshater E. (Ed.). Encyclopædia Iranica (Daneshnameh-ye Iranika). Volume IX, Fascicule 6. Festivals VIII - Fish. Bibliotheca Persica Press, New York. (on-line version of Coad (1999b) at http://www.iranica.com/articles/v9f6/v9f641a.html, downloaded 30 July 2003).

Coad, B. W. 2004. Review of "Caviar: The Strange History and Uncertain Future of the World’s Most Coveted Delicacy” by Inga Saffron. 2002. Broadway Books, New York. xv + 270 pp. U.S. $35.95. Canadian Field-Naturalist, 117(2)(2003):324-325.

Coad, B. W. 2006. Endemicity in the freshwater fishes of Iran. Iranian Journal of Animal Biosystematics, 1(1)(2005):1-13.

Coad, B. W. 2007. Review of “The Complete Fauna of Iran” by Eskandar Firouz. 2005. I. B. Tauris. London, New York. xiv + 322 pp. US$90.00. Canadian Field-Naturalist, 120(1)(2006):114-116.

Coad, B. W. 2008a. Fishes of Tehran Province and adjacent areas. Shabpareh Publications, Tehran. 244 pp., 38 figures. ISBN 978-600-5038-02-6. (40,000 rials).

Coad, B. W. 2008b. Biodiversity in Iranian tooth-carps (Cyprinodontidae). International Congress on Documenting, Analysing and Managing Biodiversity in the Middle East 20-23 October 2008, Intercontinental Hotel, Aqaba, Jordan (abstract).

Coad, B. W. 2009a. Threatened fishes of the world: Luciobarbus subquincunciatus (Günther, 1868) (Cyprinidae). Environmental Biology of Fishes, 86(2):323.

Coad, B. W. 2009b. A new species of tooth-carp, Aphanius mesopotamicus, from Iran and Iraq (Actinopterygii, Cyprinodontidae), p. 149-163., In: Neubert, E., Amr, Z., Taiti, S. and Gümüs, B. (Eds.).Animal Biodiversity in the Middle East. Proceedings of the First Middle Eastern International Congress, Aqaba, Jordan, 20-23 October 2008, Aqaba, Jordan. ZooKeys, 31 (Special Issue):252 pp. Pensoft Publishers, Sofia-Moscow.

Coad, B. W. 2009c. Alburnus zagrosensis n. sp., a new species of fish from the Zagros Mountains of Iran (Actinopterygii: Cyprinidae). Zoology in the Middle East, 48:63-70.

Coad, B. W. 2010. Freshwater Fishes of Iraq. Pensoft Publishers, Sofia-Moscow. 294 pp.,16 colour plates. ISBN 978-954-642-530-0, Pensoft Series Faunistica, 93, ISSN 1312-0174.

Coad, B. W. (Continuing). Bibliography on the Freshwater Fishes of Iran. http://gause.biology.ualberta.ca/Keivany/irbiblcoad.html. First posted 10 September 1996. Page originally maintained by Yazdan Keivany at the University of Alberta, Edmonton and now replaced by this website at www.briancoad.com, maintained by Brian W. Coad and Nicholas P. Coad, Ottawa, Ontario, Canada.

Coad, B. W. (Continuing). Freshwater Fishes of Iran. First posted 26 February 2002 at www.briancoad.com, maintained by Brian W. Coad and Nicholas P. Coad, Ottawa, Ontario, Canada.

Coad, B. W. (Continuing). Freshwater Fishes of Iraq. First posted 28 September 2005 at www.briancoad.com, maintained by Brian W. Coad and Nicholas P. Coad, Ottawa, Ontario, Canada.

Coad, B. W. and Abdoli, A. 1993a. Exotic fish species in the fresh waters of Iran. Science Forum scientifique, Canadian Museum of Nature / Musée canadien de la nature, Ottawa, November 17 novembre 1993. p. 40. (Abstract).

Coad, B. W. and Abdoli, A. 1993b. Exotic fish species in the fresh waters of Iran. Zoology in the Middle East, 9:65-80.

Coad, B. W. and Abdoli, A. 1996. Biodiversity of Iranian freshwater fishes. Abzeeyan, Tehran, 7(1):4-10, IV. In Farsi, English abstract.

Coad, B. W. and Abdoli, A. 2000a. Rhinogobius cf. similis Gill, 1859, a goby new to the fish fauna of Iran and the problem of alien invasions. Zoology in the Middle East, 20:55-59.

Coad, B. W. and Abdoli, A. 2000b. Systematics of an isolated population of tooth-carp from northern Iran (Actinopterygii: Cyprinodontidae). Zoology in the Middle East, 21:87-102.

Coad, B. W. and Al-Hassan, L. A. J. 1989a. A Bibliography of the Fishes of the Tigris-Euphrates Basin / Bibliographie der Fische des Euphrat-Tigris-Beckens. Max Kasparek Verlag, Heidelberg. 56 pp. ISBN 3-925064-05-2. DM 19.80. (dated 1988).

Coad, B. W. and Al-Hassan, L. A. J. 1989b. Freshwater shark attacks at Basrah, Iraq. Zoology in the Middle East, 3:49-53.

Coad, B. W., Alkahem, H. F. and Behnke, R. J. 1983. Acanthobrama hadiyahensis, a new species of cyprinid fish from Saudi Arabia. Publications in Natural Sciences, National Museums of Canada, 2:v + 6 pp.

Coad, B. W., Atz, J. W. and Keivany, Y. 2000. Fish imagery in art 82: Jonah and the fish. Environmental Biology of Fishes, 57(1):9.

Coad, B. W. and Bogutskaya, N. G. 2009. Alburnoides qanati, a new species of cyprinid fish from southern Iran (Actinopterygii, Cyprinidae). ZooKeys, 13:67-77.

Coad, B. W. and Bogutskaya, N. G. 2010. Petroleuciscus esfahani, a new species of fish from central Iran (Actinopterygii: Cyprinidae). Zootaxa, 2534:37-47.

Coad, B. W. and Delmastro, G. B. 1985. Notes on a sisorid catfish from the Black Sea drainage of Turkey. Cybium, 9(3):221-224.

Coad, B. W. and Esmaeili, H. R. 2009a. Desert Fishes of Iran. Desert Fishes Council 41st Annual Meeting, Furnace Creek Ranch, Death Valley National Park, California, 19 to 22 November, 2009 (abstract).

Coad, B. W. and Esmaeili, H. R. 2009b. Desert Fishes of Iran. Videotape of Coad and Esmaeili (2009a) presented at the Native Aquatic Species Restoration Webinar CSP3901, National Conservation Training Center, Shepherdstown, West Virginia, 10 December 2009.

Coad, B. W. and Holčík, J. 1999. Systematics of the cyprinid fish Chalcalburnus atropatenae (Berg, 1925) from the Lake Orumiyeh basin in northwest Iran. Biologia, Bratislava, 54(2):179-186.

Coad, B. W. and Holčík, J. 2000. On Silurus species from Iran (Actinopterygii: Siluridae). Folia Zoologica, Prague, 49(2):139-148.

Coad, B. W. and Hussain, N. A. 2007. First record of the exotic species Hemiculter leucisculus (Actinopterygii: Cyprinidae) in Iraq. Zoology in the Middle East, 40:107-109.

Coad, B. W., Hussain, N. A., Ali, T. S. and Limburg, K. E. 2003. Middle Eastern shads. In: Limburg, K. E. and Waldman, J. R. (Eds.). Biodiversity, Status, and Conservation of the World’s Shads. American Fisheries Society Symposium, 35:59-67 (ISBN 1-888569-51-4, ISSN 0892-2284).

Coad, B. W. and Keivany, Y. 1998b. Lebias vladykovi (Coad, 1988). Killi-Kontakt, Wommelgem, 26(3):43-48. In Flemish.

Coad, B. W. and Keivany, Y. 2000. Aphanius vladykovi Coad, 1988. Zagros pupfish, mahi-e gour-e khari. Journal of the American Killifish Association, 33(6):195-198.

Coad, B. W. and Keivany, Y. 2002a. Aphanius vladykovi Coad, 1988 "Zagros pubfish" (sic) mahi-e gour-e khari. Skandinaviska Killi Sällskapet Killibladet, 2002(2):3-6, 1 figure. In Swedish (translation of Coad and Keivany, 2000).

Coad, B. W. and Keivany, Y. 2002b. Reviews of "Atlas of Iranian Fishes: Gilan Inland Waters. K. Abbasi, A. Valipour, D. Talebi Haghighi, A. Sarpanah, and Sh. Nezami. 1999. Gilan Fisheries Research Centre, Rasht. vi + 113 p. 35,000 Rls ($4.38)(paperback). The Inland Water Fishes of Iran. A. Abdoli. 2000. Iranian Museum of Nature and Wildlife, Tehran. ISBN: 964-6902-01-4. 378 p. 75,000 Rls ($9.38)(hardbound). A Guide to the Fauna of Iran. E. Firouz. 2000. Iran University Press (University Publication Centre), Tehran. ISBN: 964-01-0956-8. vi + 491 p. 45,000 Rls ($5.63)(hardbound). Freshwater Fishes of Iran. H. Mohammadian. 1999. Sepehr Publications Centre, Tehran. ISBN: 946-6123-18.X. vii + 178 p. 12,000 Rls ($1.50)(paperback)". Copeia, 2002(4):1164-1166.

Coad, B. W. and Keyzer-de Ville, N. 2004. On the systematics and distribution of the snow trout, Schizothorax pelzami Kessler, 1870, in Iran (Actinopterygii: Cyprinidae). Zoology in the Middle East, 32:57-62.

Coad, B. W. and Keyzer-de Ville, N. 2005. On the validity of the species in the snow-trout genus Schizocypris Regan, 1914 (Cyprinidae: Actinopterygii). Zoology in the Middle East, 35:35-42.

Coad, B. W. and Krupp, F. 1983. Redescription of Barilius mesopotamicus Berg, 1932 a poorly known cyprinid fish from the Tigris-Euphrates basin. Cybium, 7(1):47-56.

Coad, B. W. and Krupp, F. 1994. Capoeta aculeata (Valenciennes in Cuv. & Val., 1844), a valid species of cyprinid fish from Iran (Teleostei: Cyprinidae). Zoology in the Middle East, 10:63-72.

Coad, B. W. und Kuru, M. 1986. Bibliographie der Fische der Türkei/A Bibliography of the Fishes of Turkey, pp. 15-77. In: Kasparek, M. (Ed.). Zoologische Bibliographie der Türkei. Zoological Bibliography of Turkey. Pisces, Amphibia, Reptilia. Max Kasparek Verlag, Heidelberg. 118 pp. ISBN 3-925064-01-X. DM 24.50.

Coad, B. W. and McAllister, D. E. (Continuing). Dictionary of Ichthyology. First posted 4 September 2002 at www.briancoad.com, maintained by Brian W. Coad and Nicholas P. Coad, Pure Throttle Technologies Inc., Ottawa, Ontario.

Coad, B. W., Mehrani, R. and Najafpour, N. 2009. Threatened fishes of the world: Paracobitis smithi (Greenwood, 1976) (Balitoridae). Environmental Biology of Fishes, 84(3):323.

Coad, B. W. and Najafpour, N. 1997. Barbus sublimus, a new species of cyprinid fish from Khuzestan Province, Iran. Ichthyological Exploration of Freshwaters, 7(3):273-278.

Coad, B. W. and Nalbant, T. T. 2005. A new genus and a new species of a remarkable nemacheilid fish from Iran (Pisces: Ostariophysi: Nemacheilidae). Travaux du Muséum National d’Histoire Naturelle "Grigore Antipa", Bucharest, 48:303-308.

Coad B. W. and Nalbant T. T. 2006. On the systematic position of the recently described genus and species Ilamnemacheilus longipinnis from Iran (Pisces: Ostariophysi: Nemacheilidae). 3rd International Conference Loaches of the Genus Cobitis and Related Genera. Biology, Systematics, Genetics, Distribution, Ecology, Conservation. Sibenik, Croatia, 24-29 September 2006 (title).

Coad, B. W. and Papahn, F. 1988. Shark attacks in the rivers of southern Iran. Environmental Biology of Fishes, 23(1-2):131-134. (Reprinted with same pagination in McAllister, D. E. and Kott, E. (Eds.). On lampreys and fishes: a memorial anthology in honor of Vadim D. Vladykov. Developments in Environmental Biology of Fishes, 8:1-162, Kluwer Academic Publishers, Dordrecht/Boston/London).

Coad, B. W. and Sarieyyüpoğlu, M. 1988. Cobitis elazigensis, a new species of cobitidid fish from Anatolia, Turkey. Japanese Journal of Ichthyology, 34(4):426-430.

Coad, B. W. and Vilenkin, B. Ya. 2004. Co-occurrence and zoogeography of the freshwater fishes of Iran. Zoology in the Middle East, 31:53-61.

Coad, B. W. with Waszczuk, H. and Labignan, I. 1995. Encyclopedia of Canadian Fishes. Canadian Museum of Nature, Ottawa and Canadian Sportfishing Productions, Waterdown. viii + 928 pp.

COFREPECHE. 1997. Iranian Fisheries Sector Study. Parts 1-4. Report prepared for Shilat (Fisheries Department), Jihad-e Sazandagi, Islamic Republic of Iran.

Collares-Pereira, M. J. 1994. The karyology of barbins and the possible plesiomorphic condition of polyploïdy in Cyprinidae. Bulletin français de la pêche et de la pisciculture, 334:191-199.

Collette, B. B. 1982. Rediscovery of Hyporhamphus xanthopterus, a half beak endemic to Vembanad Lake, Kerala, southern India. Matsya, 7(1981):29-40.

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Collette, B. B. and Bănărescu, P. 1977. Systematics and zoogeography of the fishes of the Family Percidae. Journal of the Fisheries Research Board of Canada, 34:1450-1463.

Compagno, L. J. V. 1984. FAO Species Catalogue. Volume 4. Sharks of the World. An Annotated and Illustrated Catalogue of Shark Species Known to Date. Part 1 - Hexanchiformes to Lamniformes, Part 2 - Carcharhiniformes. Food and Agriculture Organization, Rome, Fisheries Synopsis, 125(4), part 1:viii + 1-249; part 2:x + 251-655.

Compagno, L. J. V. 1988. Sharks of the Order Carcharhiniformes. Princeton University Press, Princeton, New Jersey. xxii + 486 pp., 21 figs., 35 plates.

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Cornwallis, L. 1976. The impact of the Mahabad Project on Lake Rezaiyeh and its southern satellite wetlands together with proposals for the conservation of the latter. Division of Nature Conservation, Department of the Environment, Tehran, Report Number W-1-35-2, 11 pp.

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Cressey, G. B. 1958b. Qanats, karez, and foggaras. Geographical Review, 48: 27-44.

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Dadikyan, M. G. 1973. Variability of the Armenian riffle minnow (Alburnoides bipunctatus eichwaldi (Filippi)) in relation to the altitude at which it occurs. Journal of Ichthyology, 13(1):68-78.

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Danesh-e-Khoshasi (sic), A. 1998. Semi-natural breeding of Chinese carps. Iranian Fisheries Scientific Journal, 6(4):19-34, 3. In Farsi.

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Esmaeili, H. R. 2006. Freshwater fish diversity in the Gulf drainage basins of Iran. The 1st International Conference on the State of the Gulf Ecosystem: Future and Trends. 5-7 March 2006, Rotana Hotel, Al Ain, United Arab Emirates University (abstract).

Esmaeili, H. R., Baghbani, S., Zareian, H. and Shahryari, F. 2009. Scale morphology of tank goby Glossogobius giuris (Hamilton-Buchanan, 1822) (Perciformes: Gobiidae) using scanning electron microscope. Journal of Biological Sciences, 9(8):899-903.

Esmaeili, H. R. and Coad, B. W. 2005. Range extension for Mystus pelusius (Solander in Russell, 1794) (Actinopterygii: Bagridae) in southern Iran. Zoology in the Middle East, 34:112-114.

Esmaeili, H. R., Coad, B. W., Gholamifard, A., Nazari, N. and Teimory, A. 2011. Annotated checklist of the freshwater fishes of Iran. Zoosystematica Rossica, 19(2)(2010):361-386.

Esmaeili, H. R. and Ebrahimi, M. 2006. Length-weight relationships of some freshwater fishes of Iran. Journal of Applied Ichthyology, 22:328-329.

Esmaeili, H. R., Ebrahimi, M., Ansari, T. H., Teimory, A. and Gholamhosseini, G. 2009. Karyotype analysis of Persian stone lapper, Garra persica Berg, 1913 (Actinopterygii: Cyprinidae) from Iran. Current Science, 96(7):959-962.

Esmaeili, H. R., Ebrahimi, M. and Rahmati, M. 2002. Loaches of Iran. II International Conference, Loaches of the Genus Cobitis and Related Genera. Biology, Systematics, Genetics, Distribution, Ecology, Conservation. September 09-13, 2002, Olsztyn, Poland (title).

Esmaeili, H. R., Ebrahmi, M., Piaravar, Z. and Teimory, A. 2006. First report on karyotype of spiny eel Mastacembelus mastacembelus (Banks and Solander in Russel, 1794) from south of Iran. Journal of Science, Al-Zahra University, Tehran, 18(3):48-54. In Farsi.

Esmaeili, H. R., Ebrahimi, M. and Saifali, M. 2008. Karyological analysis of five tooth-carps (Actinopterygii: Cyprinodontiae) from Iran. Micron, 39:95-100.

Esmaeili, H. R., Ebrahimi, M., Teimory, A. and Ansary, T. H. 2008. First karyological analysis of an endemic fish, Isfahan tooth-carp, Aphanius isfahanensis (Actinopterygii: Cyprinodntidae) from Iran. Journal of Applied Animal Research, 33:73-76.

Esmaeili, H. R., Ebrahimi, M., Teimori, (sic) A. and Ansari (sic), T. H. 2009. First karyological analysis of an endemic fish, Zagros tooth-carp, Aphanius vladykovi Coad, 1988 (Actinopterygii: Cyprinodntidae) from Iran. Iranian Journal of Science and Technology, Transaction A, 33(A4):349-354.

Esmaeili, H. R., Ganjali, Z. and Monsefi, M. 2009. Reproductive biology of the endemic Iranian cichlid, Iranocichla hormuzensis Coad, 1982 from Mehran River, southern Iran. Environmental Biology of Fishes, 84(1):141-145.

Esmaeili, H. R., Ganjali, Z. and Monsefi, M. 2010. Gonad morphology and histology of the endemic Hormuz cichlid, Iranocichla hormuzensis Coad, 1982 from Mehran River, southern Iran. IUFS Journal of Biology, 69(1):1-12.

Esmaeili, H. R. and Gholami, Z. 2007. Investigations on the surface ultrastructure of scale of Geno tooth-carp, Aphanius ginaonis (Holly, 1929) (Actinopterygii: Cyprinodontidae) using scanning electron microscope. Iranian Journal of Biology, 20(2):307-314. In Farsi.

Esmaeili, H. R., Gholami, Z., Nazari, N., Gholamifard, A., Shahryari, F., Baghbani, S. and Ebrahimi, M. 2009. Karyotype analysis of an endemic sucker catfish, Glyptothorax silviae Coad, 1981 (Actinopterygii: Sisoridae), from Iran. Turkish Journal of Zoology, 33(4):409-412.

Esmaeili, H. R., Gholamifard, A. and Freyhof, J. 2011. Ichthyofauna of Zarivar Lake (Iran) with the first records of Hemiculter leucisculus and Alburnus hohenackeri in the Tigris drainage. Electronic Journal of Ichthyology, 7(1):1-6.

Esmaeili, H. R., Gholamifard, A., Teimori, A., Baghbani, S. and Coad, B. W. 2010. Xiphophorus hellerii Heckel, 1848 (Cyprinodontiformes, Poeciliidae), a newly introduced fish recorded from natural freshwaters of Iran. Journal of Applied Ichthyology, 26(6):937-938.

Esmaeili, H. R., Hagmoradlo, A. and Niksirat, H. 2003. Exotic fishes of Iran with special reference to Gasterosteus aculeatus. World Aquaculture 2003, 20-23 May, Salvador, Brazil (title).

Esmaeili H. R. and Niknejad, V. 2006. Preliminary study of southern spined loach Cobitis linea (Heckel, 1849) (Pisces; Cobitidae) in Fars province, Southwest Iran. 3rd International Conference Loaches of the Genus Cobitis and Related Genera. Biology, Systematics, Genetics, Distribution, Ecology, Conservation. Sibenik, Croatia, 24-29 September 2006 (title).

Esmaeili H. R. and Niknejad, V. 2006-2007. Scale morphological studies of 4 loaches (Actinopterygii: Cypriniformes) in Fars Province. Journal of Science, Al-Zahra University, 19(2):1-10, 67. In Farsi.

Esmaeili, H. R. and Piravar, Z. 2006a. Karyotype of Persian chub, Petroleuciscus persidis (Coad, 1981) (Actinopterygii: Cyprinidae) from southern Iran. Turkish Journal of Zoology, 30:1-3.

Esmaeili, H. R. and Piravar, Z. 2006b. On the karyotype of Cyprinion tenuiradius Heckel, 1849 (Pisces: Cyprinidae) from the southwest of Iran. Zoology in the Middle East, 39:75-80.

Esmaeili, H. R. and Piravar, Z. 2007. Karyotype analysis of Garra rufa (Heckel, 1843) (Actionopterygii: Cyprinidae) in Fars Province. Iranian Scientific Fisheries Journal, 16(3):11-18. In Farsi.

Esmaeili, H. R., Piravar, Z. and Ebrahimi, M. 2006. Karyological analysis of Iranian cichlid fish, Iranocichla hormuzensis Coad, 1982 (Perciformes, Cichlidae) from southern Iran. Journal of Applied Animal Research, 30:77-79.

Esmaeili, H. R., Piravar, Z. and Shiva, A. H. 2007. Karyological analysis of two endemic tooth-carps, Aphanius persicus and Aphanius sophiae (Pisces: Cyprinodontidae) from southwest Iran. Turkish Journal of Zoology, 31:69-74.

Esmaeili, H. R. and Shiva, A. H. 2006. Reproductive biology of the Persian tooth-carp, Aphanius persicus (Jenkins, 1910)(Cyprinodontidae), in southern Iran. Zoology in the Middle East, 37:39-46.

Esmaeili, H. R. and Teimori, A. 2009. Exotic and introduced fish species of Iran and their impact on native fishes. 13th European Congress of Ichthyology, 6-12 September 2009, Klaipeda, Lithuania (abstract).

Esmaeili, H. R., Teimory, A. and Coad, B. W. 2008. Qanat systems as fish refuge. International Congress on Documenting, Analysing and Managing Biodiversity in the Middle East 20-23 October 2008, Intercontinental Hotel, Aqaba, Jordan (title).

Esmaeili, H. R., Teimory, A., Coad, B. W. and Gholami, Z. 2008. Threatened fishes of the world: Cobitis linea (Heckel, 1849). Environmental Biology of Fishes, 83(4):407-408.

Esmaeili, H. R., Teimory, A., Coad, B. W. and Gholami, Z. 2009. Threatened fishes of the world: Seminemacheilus tongiorgii Nalbant and Bianco, 1998 (Balitoridae). Environmental Biology of Fishes, 84(4):375.

Esmaeili, H. R., Teimory, A., Gholami, Z. and Hosseinie, F. 2006. Range extension of Barbus sublimus Coad and Najafpour, 1997 (Actinopterygii: Cyprinidae) and its sympatric species in southwest of Iran. Iranian Journal of Animal Biosystematics, 2(1):19-24.

Esmaeili, H. R., Teimory, A. and Khosravi, A. R. 2007. A note on biodiversity of Ghadamghah spring-stream system in Fars province, southwest Iran. Iranian Journal of Animal Biosystematics, 3(1):15-23.

Esmaeili, H. R., Teimory, A. and Piravar, Z. 2009. Otolith morphology of some freshwater fishes of Iran. Iranian Scientific Fisheries Journal, 17(4):163-168. In Farsi.

Esmaeili, H. R. and Timoorei, A. 2006. Morphology of urohyal bone and its importance in taxonomy of some freshwater fishes of Iran. Iranian Scientific Fisheries Journal, 15(3):1-8. In Farsi.

Esmaeili, H. R., Yazdanpanah, M. and Monsefi, M. 2005. Reproductive biology of doctor fish, Garra rufa (Cyprinidae: Garrinae), in southwest of Iran. Journal of Fish Biology, 67(Supplement B):282.

Esmaeili, H. R., Zareian, H., Gholamhosseini, A., Ebrahimi, M., Gholami, Z. Teimori, A. and Ansari, T. A. 2010. Karyotype analysis of the king nase fish, Chondrostoma regium (Heckel, 1843) (Actinopterygii: Cyprinidae) from Iran. Turkish Journal of Fisheries and Aquatic Sciences, 10(4):477-481.

Esmaeilzadeh, K. R., Sahari, M. A. and Hamidi, E. Z. 2004. Comparative study on nutrient compositions of kutum (Rutilus frisii kutum) and grass carp (Ctenopharyngodon idella) and their marinade qualities. Iranian Journal of Fisheries Sciences, 12(4):13-28. In Farsi.

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Esmaili Sari A., Abtahi, B., Seyfabadi, J. and Khodabandeh, S. 2001. Invasive comb-jelly and future of the Caspian Sea. Naghshe Mehr Publication, Tehran, p. 144. In Farsi.

Esmaili Sari A., Abtahi, B., Seifabadi, J. Khodabandeh, S. Ershad, H. 2001. A study on the density and distribution of comb-jelly, Mnemiopsis leidyi, in the Caspian Sea. Daneshvar, 31:145-148. In Farsi.

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Esmaili Sari, A., Farshchi, P and Darvishi, F. 2002. Feeding competition between invasive comb-jelly (M. leidyi) and anchovy kilka (C. engrauliformis) in south coast of the Caspian Sea. Iranian Journal of Marine Sciences, 1(4):25-42, 143. In Farsi.

Esmaili Sari, A., Imandel, K. A. and Georgez, Sh. 2001. The valuation of the damages resulting from oil material on Shadegan Wetland environment. Journal of Environmental Science and Technology, 2001(9):21-30. In Farsi.

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