Freshwater Fishes of Iran

Introduction - History of Research

Revised:  06 October 2010

Acknowledgements     Purpose     Materials and Methods     History of Research     Fisheries     Geography     Climate     Habitats     Environmental Change     Drainage Basins     Scientific Names     Fish Structure     Collecting Fishes     Preserving Fishes     Quotes

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 esocinus, B. grypus, B. kersin, B. luteus, B. sharpeyi, B. xanthopterus, Aspius vorax, Acanthobrama (presumably A. marmid), Cyprinion sp., Alburnus sp., Silurus triostegus, Mystus pelusius, Mastacembelus mastacembelus, Liza abu, Acanthopagrus sp., and Tenualosa ilisha.

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

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


Vessel, 5th millennium B.C., Susa, Khuzestan           

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



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


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 (, 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 Barbus sharpeyi) and shabbût (= probably Barbus grypus)(Perry, 1998). Probably the best example of an early "scientific" Islamic work on zoology is the fourteenth century "Nuzhatu-l-Qulub" or "Hearts Delight" by Hamdullah Al-Mustaufi Al-Qazwini (translated into English by Stephenson (1928)). Only the "tarikh" is identifiable as a freshwater fish - Alburnus tarichi from Lake Van in modern Turkey.

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

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

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

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

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

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

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

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

Scientific works relevant to Iran begin with the Systema Naturae, 10th edition, by Carolus Linnaeus (1701-1778) published in 1758 and in which scientific naming in zoology has its beginning. Linnaeus adopted many of the names from the system developed by Petrus Artedi (1705-1735) who, on a visit to Amsterdam to examine a collection of fishes from the East and West Indies, drowned in one of the canals. Genera subsequently found in Iran include Acipenser, Perca, Cobitis, Silurus, Salmo, Esox, Atherina, Mugil, Cyprinus, and Syngnathus and various species were described in these and other genera. After this date a variety of papers were published by authors in many countries describing fishes scientifically and some of these fishes were eventually found to occur in Iran, as with the Linnaean genera and species. Examples include Marc Elieser Bloch (1723-1799), a physician who began to devote himself to ichthyology at the age of 56, and Johann Gottlob Schneider (1750-1822) who collaborated with Bloch and published their "Systema Ichthyologiae" in 1801 after Bloch's death. This work contains all known species at that time (Bloch also wrote "Naturgeschichte der ausländischen Fische, 1785-1795) and in these works appear such Iranian species as diverse as the riffle minnow, Alburnoides bipunctatus, 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) and in a later work (1826) the pipefish (Syngnathus abaster) 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 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 and the minnow Rutilus frisii.

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

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

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

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

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

1. Barbus barbulus

2. Systomus albus var. alpina (= Barbus 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 (= Nemacheilus persa)

14. Acanthopsis linea (= Cobitis linea)

15. Lebias sophiae (= Aphanius sophiae)

16. Lebias punctata (= Aphanius sophiae)

17. Lebias crystallodon (= Aphanius sophiae)

In all, only 6 new species were discovered according to the modern interpretation of these taxa. In addition the following 22 described from Syria and modern Iraq by Heckel have since been found in Iran: Acanthobrama marmid, Aspius vorax, Barbus esocinus, B. grypus, B. kersin, B. lacerta, B. luteus, B. pectoralis, B. xanthopterus, Capoeta trutta, Alburnus mossulensis, Chondrostoma regium, Cyprinion kais, C. macrostomus, Garra rufa, G. variabilis, Squalius lepidus, Nemacheilus frenatus, N. tigris, Silurus triostegus, Aphanius mento and Liza abu. Heckel therefore described 28 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 travelled 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 travelled 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 subquincunciatus, Hemigarra elegans, and Nemacheilus griffithi. 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 councillor 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 sharpeyi and Macrones colvillii (= Mystus pelusius). Barbus faoensis (= B. 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 Nemacheilus 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 filippi, Barbus brachycephalus, Capoeta buhsei (from "Persia", apparently near Tehran (Berg, 1949)), Chondrostoma oxyrhynchum, Nemacheilus brandti, N. 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 (= B. mursa) and Labeobarbus euphrati from the Euphrates River (= B. 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 bampurensis, Nemacheilus sargadensis, Discognathus (= Garra) rossicus ?

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). 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. 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), by Banawari Lal Chaudhuri of the Indian Museum, Calcutta in 1909 who reported a new loach (Nemacheilus macmahoni (= N. rhadinaeus)) 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-Buchanan and Day, and Director of the Zoological Survey of India.

A. Ya. Nedoshivin and B. S. Il'in 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, Glyptothorax kurdistanicus, Anatirostrum profundorum, Knipowitschia caucasica and Knipowitschia iljini. His summary work "Freshwater Fishes of the U.S.S.R. and adjacent countries" was published in 1948-1949 and in English translation in 1962-1965 and has much of relevance to northern Iran, although the taxonomy is now dated. His 1940 work on the "Zoogeography of freshwater fish of the Near East" placed that fauna in context and included Iran but it was his 1949 work "Freshwater Fishes of Iran and adjacent countries" which has been the major modern work on Iranian fishes south of the Caspian Sea basin and the Lake Orumiyeh basin. This was based on collections deposited in the U.S.S.R. Academy of Sciences Zoological Institute in Leningrad (acronym ZIL, now St. Petersburg, Russia with the acronym ZISP). The collections had been made by two Russian biologists. The first of these was Nikolai Alekseevich Zarudnyi (1859-1919), a zoologist and ornithologist who made four journeys to Iran for which he was awarded medals and the Przheval'skii Prize by the Russian Geographical Society. His first journey in 1896 was to Kuchan, Sistan and Mashhad, his second in 1898 was to eastern Khorasan and Beluchistan, the third (1900-1901) was to Khorasan, Sistan and Beluchistan including the Bampur region and the Makran, and the last journey (1903-1904) was to Gorgan, western Khorasan, western Kuhistan, southern Irak-Ajemi and Khuzestan. Zarudnyi's material had previously been examined and described by Nikol'skii (see above). The second biologist was P. V. Nestorov who worked with the Turko-Persian Demarcation Commission in 1914 and collected fishes in the Tigris basin along the present Iran-Iraq frontier.

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

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

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

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


© Brian W. Coad (