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.
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.
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.
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 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.
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.
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.
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.
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.
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.
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.
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.
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.
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.
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.
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.
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.
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.
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.
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.
From the anteriormost part of the head to the tip of either
lobe of the caudal fin when that fin is normally splayed.
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.
From the anteriormost part of the snout to the bony margin
of the opercle (excluding the opercular membrane).
Maximum straight line depth excluding fins or fleshy and
scaly structures at fin bases
The distance between the opercles when in their normal,
closed position. The opercles are gently pressed into a closed position
if greatly dilated.
From the anteriormost part of the snout or upper lip at
the mid-line to the bony front margin of the orbit.
Greatest diameter between the bony rims of the orbit.
This distance is not always horizontal.
Greatest distance between the posterior bony orbit margin
and the bony opercular margin.
Least bony width between the orbits over the top of the
head in a straight line.
From the base of the anteriormost dorsal fin ray to the
tip of the snout or upper lip.
From the base of the anteriormost pelvic fin ray to the
anteriormost point on the head (snout or upper lip).
From the base of the anteriormost anal fin ray to the
anteriormost point on the head (snout or upper lip).
The oblique distance from the insertion of the anal fin
to the mid-point of the end of the hypural plate.
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).
From the extreme base of the uppermost, outermost or anteriormost
ray to the tip of the fin.
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.
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.
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:-
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:-
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.
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).
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).
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:-
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:-
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:-
This province demonstrates a great variation in mullet catch between years.
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:-
Abdolmalaki and Psuty (2007) give figures over a wide range of years for Iranian coastal catches in the southern Caspian Sea as follows:-
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.
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) :-
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:-
The Caspian Environment Programme (1998) gives annual production (in
thousands) of the main cultured fish species in government and private
hatcheries as follows:-
Hosseinzadeh (2003) gives the following figures in tonnes for total fisheries
production in Iran (note that southern waters are marine captures):-
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:-
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:-
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:-
Production by major fish-culturing provinces from Salehi (2004a) for carps is
as follows:-
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:-
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.
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.
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:-
Markazi (= Central or Tehran; sometimes split into Tehran and a southeast part called Markazi)
Hormozgan (or Hormozdgan or Saheli-ye Jazayer va Banader-e Khalij-e Fars va Darya-ye Oman)
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.
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.
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.
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).
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).
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.
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).
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.
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.
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.
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:-
Lake Kopibalbalch, Hassanlu Marsh, Yadergarlu Marsh and surrounding
marshes (37°00'N, 45°30'E)
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.
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:-
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.
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.
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.
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 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.
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).
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.
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).
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).
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:-
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 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.
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 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.
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.
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).
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.
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.
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.
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.
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.
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.
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 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.
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 (= 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.
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).
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.
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.
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.
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.
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:-
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.
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.
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):-
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.
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).
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.
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.
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.
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 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).
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
---------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------
6a. Pelvic fins absent; body very elongate ("eel-like") ---> 7
6b. Pelvic fins present; body not very elongate ---> 9
---------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------
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
---------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------
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)
---------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------
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
---------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------
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
---------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------
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
---------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------
12a. Blunt
grinding teeth in jaws = Sparidae (Acanthopgarus latus - Marine
basins)
12b.Teeth absent or, if present, sharp ---> 13
---------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------
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
---------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------
14a. Barbels absent ---> 15
14b. Barbels present ---> 18
Barbels in Paracobitis smithi (dorsal view) and in Barbus lacerta
---------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------
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
---------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------
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
---------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------
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)
---------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------
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
---------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------
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
---------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------
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
---------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------
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
---------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------
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
---------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------
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
---------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------
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
---------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------
26a.
No teeth in jaws; lateral line usually obvious ---> 27
26b. Teeth in jaws; no lateral line pores ---> 28
---------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------
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
---------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------
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
2b. = 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
male
female
5b. Females with flank spots ---> 6
6a. ?Females without lozenge-shaped spot at caudal fin base; flank spots large;
Tigris River basin =
Aphanius mesopotamicus
male
female
6b. Females with lozenge-shaped spot at caudal fin base; flank spots small; Kor River basin = Aphanius sophiae
male
female
--|----|----|--
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
2b. 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.
Lateral 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 AccountsThe 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
Back to Contents
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
Adult
Adult
Adult
Ammocoete
Disc
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
Back to
Contens
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
Back to
Contents
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.
.jpg)
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
Back to Contents
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
Back to Contents
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)
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.
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
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
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)
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)
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)
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
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)
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.
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
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 th
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