i. FRESHWATER FISHES
History of research. The most important early work on the fishes of Persia is that of Jakob Heckel, inspector at the Imperial Royal Court Collection of Natural History in Vienna. He described the collections sent by Theodor Kotschy to Vienna from around Shiraz, comprising twenty-two species. Other new species from Persia were collected by Pierre Aucher-Eloy, a French botanist, who traveled extensively in Persia from 1835-38, eventually dying at Eṣfahān from “an excess of zeal for natural sciences” (Cuvier and Valenciennes, XVII, p. 298). Albert Günther of the British Museum published a series of papers on fishes from Persia, or fishes from neighboring countries eventually to be found in Persia, e.g., the collections of the Afghan Delimitation Commission dispatched by the British government to mark the western borders of Afghanistan; collections made by Robert T. Günther in 1898 in the Lake Urmia basin comprising four new species. Eugen Keyserling joined a scientific expedition in 1858-59, sent by the Russian Imperial government to explore Khorasan. The difficulty of baggage transport limited the quantity of alcohol Keyserling could carry, and his early fish collections spoiled. However, he did draw cyprinid fishes from nature, and gave good descriptions of new species from Isfahan to Kermān. Filippo de Filippi, an Italian zoologist, professor at Turin and director of the museum, accompanied an Italian embassy to Persia in 1862, visiting Tabrīz, Qazvīn, Tehran, Rašt and the Caspian Sea. His companion Giacomo Doria collected fishes as far south as Shiraz. Seventeen species were described from the Caspian basin and inland waters of Persia, although locality data were poor in some instances. William Thomas Blanford accompanied the Persian Boundary Commission in 1872, publishing a two-volume account in 1876, but fish collections were minor and were not included. In association with this commission, Major St. John made collections from 1869-71 with a collector from the Indian Museum, Calcutta. Part of the collections was described by J. T. Jenkins in 1910 from material taken in Baluchistan and around Shiraz and deposited in Calcutta. Nelson Annandale, founder and director of the Zoological Survey of India, co-authored a review of the fishes of Sīstān with Sunder Lal Hora, based on specimens collected by the Seistan Arbitration Commission of 1901-4, and by officers of the Zoological Survey of India in the winter of 1918. Nine species were described. Alfons Gabriel and his wife collected fishes in the neighborhood of Bandar-e ʿAbbās, including specimens from the Genū hot spring and the Bašākerd mountains (q.v.). This material was described in 1929 by Maximilian Holly of the Naturhistorisches Staatsmuseum in Vienna, and contained two new species. Lev Semenovich Berg, a leading Soviet physical geographer and biologist, made many contributions to the ichthyology of Persia in a number of shorter articles, and in lengthy monographs from the late nineteenth century onwards. His three-volume summary work on the freshwater fishes of the U.S.S.R. and adjacent countrieswas published in Russian in 1948-49 and in English translation in 1962-65. It has much of relevance to northern Persia, 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 Persia, but it was his 1949 work on the freshwater fishes of Persia and adjacent countries which has been the major modern work on Persian fishes south of the Caspian Sea basin and the Lake Urmia basin. This was based on collections deposited in the Zoological Institute of the U.S.S.R. Academy of Sciences in Leningrad. The collections had been made by two Russian biologists. The first of these was Nikolaĭ Alekseevich Zarudnyĭ, a zoologist and ornithologist who made four journeys to Persia in the late nineteenth century. Zarudnyĭ’s material had previously been examined and described by Aleksandr Mikhailovich Nikolskiĭ .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 Persian-Iraq frontier. A recent thesis (Saadati) contains analyses of new collections and of works by a number of Persian authors (e.g. Rostami). Others listed in the bibliography have added new species and revised our understanding of the Persian ichthyofauna. The majority of these works are systematic and taxonomic. There have been relatively few studies of the ecology and behavior of Persian fishes, and those few are concentrated on the commercially important sturgeons, and on a few carp species in the Caspian Sea basin.
Fish species. The ichthyofauna of inland waters in Persia (rivers, lakes, large water reservoirs behind dams, aquaculture ponds, etc.) includes both native and exotic (acclimatized), resident and/or anadromous fishes (see Coad, 1995; 1998b). The systematic diversity of freshwater fishes in Persia is summarized in Table 1.
1. Natives comprise 169 species in 24 familes The Cyprinidae (carps and carp-like taxons), with 37 genera and 82 species, has the comparatively largest biodiversity, followed by the Balitoridae (hillstream loaches), with 1 genus and 21 species. Cyprinids and balitorids are found in all ten of the major watersheds of the country.
The Cyprinidae are the commonest and most individually numerous fishes in qanāts (Coad, 1996f), springs, and small streams in the vast desert basins. The species which have an underslung mouth and a fine cutting edge to the lower jaw thrive there, living off the small plants and other organisms adhering to the rocks. Many minnow species are small, but some attain a large size, up to 2.1 m and 136 kg for the “Tigris salmon” (Barbus esocinus; Figure 1). The smaller species are often a vital component of the ecosystem, and some make excellent aquarium fishes. The bitterling, Rhodeus amarus (Figure 2), is particularly colorful in the spawning season when males have a bright red iris and iridescent violet and steel-blue flanks, the throat and belly are orange to blood-red, the dorsal and anal fins are bright red, margined with black, and the caudal fin is green at the base and yellow at the edge. Reproduction in this species is also unusual. The female develops an ovipositor up to 6 cm long which deposits eggs inside freshwater clams where they develop protected from predators and even desiccation since the clam can move into deeper water in unfavorable conditions.
The endemics in Persia include two scientifically interesting tiny “blind” (i.e., eyeless) cave-fishes, both discovered in a small pool supplied by underground water at a locality near Dorūd in a valley of the Zagros mountains: Iranocypris typhlops (fam. Cyprinidae; Figure 3), “Asia’s first blind cave-fish,” a tiny cyprinid discovered in 1937 by two Danish scholars, A. F. Bruun and E. W. Kaiser (Smith, 1953, pp. 79, 80, 86, etc.); and N(o)emacheilus smithi Greenwood (now assigned to the fam. Balitoridae; Figure 4), “the first blind [cave-dwelling] loach” from Asia, discovered in 1976 by the British writer Anthony Smith, in deference to whom the species was named (vividly recounted in Smith, 1979, especially chaps. 5 and 6; see also idem, 1953).
Distribution. The distribution of fishes in Persian freshwater basins is summarized in Table 2. Distributions include species residing in the Caspian Sea, which has a salinity of about 12 parts per thousand, one third the salinity of seawater. The Caspian Sea has a mix of purely “marine” species, and others which also reside or spawn in fresh water. Members of 24 families are native to Persia, and another 16 families have been introduced with varying success. In regard to the native ichthyofauna, half the families are represented by only a single species, and only 3 families have more than 10 species. These 3 families comprise about 72 per cent of the species. Taxonomically, the fauna of Persia is dominated by the Ostariophysi (families Cyprinidae to Sisoridae in the table), which comprise 67 per cent of the native fishes.
Not only is species diversity concentrated in a few families, but also distribution is concentrated in a few basins (Table 2). Only the loaches, the carps, and the minnows are found in all Persian basins. Thirteen of the families are found in only one basin. The most speciose basins are the Tigris River basin, draining the Zagros Mountains to the head of the Persian Gulf, and the Caspian Sea basin, which includes both fresh and brackish water habitats. Both these basins have large rivers, small streams, and extensive marshes. Central Persia is poor in species, despite covering the largest area, and the waters there comprise small streams exposed to the sun, with little cover of habitat diversity. The Sīstān basin has large rivers and extensive marsh habitat, but is remote from the sea and from the centers of faunal diversity. Generally, the diversity of fishes declines with distance from the Caspian Sea and the Persian Gulf. Thus the Urmia and Tedzhen (Harīrūd) basins show strong affinities to the Caspian Sea basin, but have a less diverse fauna, while the internal basins of Baluchistan have similar faunas to coastal basins, without the marine elements. Some species are widespread and found in a number of basins, e.g. a minnow (Capoeta damascina; Figure 5) is found in six basins. Many other species are restricted to a single basin. In this respect, the Caspian Sea basin has 54 species found nowhere else in Persia. At 79 per cent of all the fish species represented in the country, it also accounts for a highly significant proportion of ichthyofauna diversity.
Relationships of the ichthyofauna. Persia lies at a faunal crossroads between Europe, Asia and Africa, and this is reflected in its fish fauna. The fishes of the Caspian Sea and its tributaries, along with neighboring basins such as the Tedzhen in the east and the Urmia in the west, have some degree of endemism, but most species are also found in the Mediterranean and Black Sea basins. There has been a long history of transgressions and isolation of the Caspian Sea, which can account for the similarity and differences between its fauna and that of Europe.
The origin of a number of southern species depends on marine migrations by freshwater fishes, which are known to have a wide tolerance of salinities. These include the herring (Tenualosa ilisha; Figure 6), the milkfish (Chanos chanos), the tooth-carp (Aphanius dispar; Figure 7), three goby species, and possibly the mullet (Liza abu; Figure 8). Just as the sea can be a highway facilitating dispersal of fishes, so can major rivers. The snow trout tribe (Schizocypris, Schizothorax and Schizopygopsis) of the Sīstān basin (altitude under 500 m) are apparently derived from the Hindu Kush mountains (altitudes over 5000 m) via the long Helmand River (q.v.) which crosses deserts to its terminal basin in Sīstān. These essentially montane fishes were probably driven down into lowlands by an ice age, and are now relict there. A number of basins have headwaters which rise close together on plains, and drainage capture has no doubt been a significant means of transfer and dispersal of fishes between the many currently isolated basins in Persia. A number of minnow species, such as Capoeta aculeata (Figure 9), are distributed in isolated basins in a manner which suggests headwater capture as the mechanism, perhaps aided by more extensive water courses under pluvial conditions in the past.
While the distribution of many fishes in Persia can be attributed to movements between watercourses, another explanation can be suggested for the distribution of tooth-carps in internal basins remote from the sea. These small fishes of low swimming ability are unlikely to have dispersed from the major lowland river basin of the Tigris onto the Persian plateau. Fossil evidence indicates that they are a relict of the Tethys Sea, which once covered Persia. This sea was lost through desiccation and continental uplift, and these fishes may have risen with the post-Pliocene uplift of the Zagros Mountains. Rivers of the Kor, and the Gulf and Hormoz basins, may have received some of their fishes by headwater capture, but another means suggests itself. Their faunas are derived from the separate Tigris River basin, many species being shared. As recently as 18,000 years ago, in the Pleistocene, the Tigris-Euphrates basin debouched into the mouth of the Persian Gulf near the Straits of Hormoz. Rivers of the Gulf and Hormoz basins would then have been tributaries to the expanded Tigris-Euphrates basin and could have derived their fishes directly from it. Since that time, sea level has fallen by as much as 120 m.
Certain distributions are not readily explainable. The minnow (Barilius mesopotamicus; Figure 10), the bagrid catfish (Figure 11), the sucker catfishes, and the spiny eel of the Tigris River and neighboring basins all have their nearest relatives in the Indus River basin, with a large distributional gap in between. A similar situation holds for a loach (Cobitis linea; Figure 12) and the cichlid (Iranocichla hormuzensis; Figure 13), both endemics of southern Persia, whose putative relatives reside far outside Persia’s borders, in Europe and Africa. These distributions require explanations involving selective loss of populations in intervening areas.
Dangerous fishes. A number of freshwater fishes are potentially dangerous to man if the eggs are consumed, although there are no documented fatalities in Persia. The eggs of the tench (Tinca tinca; Figure 14) of the Caspian Sea basin, and the snow trout (Schizothorax zarudnyi; Figure 15) of Sīstān, are particularly dangerous. Victims generally recover within three to five days, but severe cases can lead to paralysis, convulsions, coma, and death. The Indian stinging catfish (Heteropneustes fossilis; Figure 16), an exotic species in Ḵūzestān, has venomous pectoral fin spines and an aggressive stabbing behavior when disturbed. Anecdotal reports exist of fatalities from southern Persia (Coad, 1979a). The venom has both neurotoxic and haemotoxic components, and documented fatalities have occurred in India. Undoubtedly the most dangerous fish in Persia is the bull shark (Carcharhinus leucas; Figure 17), found in rivers of Ḵūzestān as far as 420 river kilometers from the sea. There are thirty-four documented unprovoked attacks with seventeen fatalities (Coad and Papahn); many other cases no doubt have gone unrecorded. The sharks seize people who are bathing, swimming, washing clothes or vehicles, or fishing.
Exotic fishes. At least twenty-nine species of fishes have been introduced to Persia from other countries, and perhaps twelve have become established and are now ineradicable (Coad, 1996e, 1996g; Coad and Abdoli, table 1). In addition, five native species have been transplanted to drainage basins where they do not occur naturally. Exotics have been imported for food, aquaculture, sport, research, ornamentation, weed control, health reasons, or accidentally. These exotics are often deleterious, introducing parasites and diseases, competing for food and habitat, swamping native genetic diversity, or consuming native fishes. The most widespread exotic is the mosquitofish (Gambusia holbrooki; Figure 18), introduced to eat the aquatic larvae of mosquitoes which carry malaria. This small fish is now found in all the water bodies of Persia, including many which were fishless.
Endangered fishes. A number of species found in Persia are quite rare, but this is a function of their natural distribution at the limit of the species range. For example the snakehead (Channa gachua; Figure 19) is known only from five specimens caught in Baluchistan (Coad, 1979b), but this is the westernmost distribution of the species. Fishes which are in particular danger are those of commercial importance in the Caspian Sea basin. These fishes depend on a migration from feeding grounds in the mouths of rivers or the sea itself, into freshwater for spawning. The construction of dams, pollutants from industry and farming, poaching, and excessive commercial catches, have all contributed to a decline in their numbers. Still other species are threatened because of their restricted habitat—a single event such as a chemical spill or a natural disaster could eliminate them. Among these are the cave fishes (Iranocypris typhlops of the carp family; Nemacheilus smithi of the hillstream loach family). These fishes are restricted to a cave system in the Zagros Mountains; the extent of this cave system is unknown. Holly’s pupfish (Aphanius ginaonis of the tooth-carp family; Figure 20) lives only in the Genū hot spring north of Bandar-e ʿAbbās. The temperature of the spring is 41° C; the fish live in the small stream flowing from it.
M. R. Ahmadi, “Aquaculture Industry in Iran,” in I Chiu Liao, Chung-Zen Shyu, and Nai-Hsien Chao, eds., Aquaculture in Asia: Proceedings of the 1990 Asian Productivity Organization Symposium on Aquaculture, Keelung, Taiwan, 5-13 September, 1990, Keelung, Taiwan, 1993, pp. 69-79.
M. R. Ahmadi and Gh. Wossughi, “Ein Beitrag zur fischereilichen Bedeutung des Hamun-Sees, Iran,” Österreichs Fischerei 41/10, 1988, pp. 209–16.
H. Ahnelt and J. Holcík, “Distribution of Two Species of the Genus Neogobius (Pisces: Gobiidae) in the Catchment Area of the Southern Caspian Sea,” Acta Universitatis Carolinae Biologica 40/1-2, 1996, pp. 99-114.
L. A. J. Al-Hassan, “Meristic Comparison of Liza abu from Basrah, Iraq and Karkhah River, Arabistan, Iran,” Cybium 8/3, 1984, pp. 107-8.
N. Annandale, “Notes on the Fish of the Genus Discognathus from India and Persia,” Records of the Indian Museum 18, 1919, pp. 65–78, pl. IX–XI.
N. Annandale and S. L. Hora, “The Fish of Seistan,” Records of the Indian Museum 18, 1920, pp. 151–203, pl. III.
G. Azari Takami, S. Etessami and A. Saremi “Régime alimentaire des esturgeons (Acipenseridés) sur les côtes sud de la mer Caspienne,” Cybium 3/10, 1980, pp. 65-72.
P. Banarescu and T. Nalbant, “The 3rd Danish Expedition to Central Asia, Zoological Results 34: Cobitidae (Pisces) from Afghanistan and Iran,” Videnskabelige Meddelelser fra Dansk naturhistorisk Forening 129, 1967, pp. 149-86, pl. XIX-XXI.
P. M. Banarescu and B. Herzig-Straschil, “A Revision of the Species of the Cyprinion macrostomus-group (Pisces: Cyprinidae),” Annalen des naturhistorischen Museums in Wien, 97B, 1995, pp. 411-20.
D. M. Bartley and K. Rana, “Stocking Inland waters of the Islamic Republic of Iran,” FAO Aquaculture Newsletter 18, 1998, pp. 16-19.
L. S. Berg, 1913. “Description of a New species of Garra (=Discognathus) from Eastern Persia,” Ezhegodnik Zoologicheskago Imperatorskoi Muzeya Akademii Nauk (St. Petersburg) 18, 1913, p. 61.
Idem, “Opisanie novogo vida Alburnus (Pisces) iz basseĭna oz. Urmii” (Description of a new species of Alburnus [Pisces] in the basin of the lake Urmia), Trudy Zoologicheskogo instituta Akademii nauk SSSR 26, 1925, pp. 213–14.
Idem, “Description of a New Siluroid Fish, Glyptosternum kurdistanicum from the Basin of the Tigris River,” Izvestiya Akademii Nauk SSSR 7, 1931, pp. 1267-70.
Idem, “Eine neue Barilius-Art (Pisces, Cyprinidae) aus Mesopotamien,” Zoologischer Anzeiger 100, 1932, pp. 332–34.
Idem, Acipenser güldenstädti persicus, a Sturgeon from the South Caspian Sea,” The Annals and Magazine of Natural History 10/13, 1934, pp. 317-18.
Idem, “Zoogeografiya presnovodnykh ryb Peredneĭ Azii” (Zoogeography of freshwater fish of the Near East), Uchenye zapiski Leningradskogo gosudarstvennogo universiteta, Seriya Geograficheskikh nauk 3, 1940, pp. 3-31.
Idem, Ryby presnykh vod SSSR i sopredel’nykh stran, 3 vols., Moscow 1948–49, tr. Israel Program for Scientific Translation as Freshwater Fishes of the USSR and Adjacent Countries, 3 vols., Jerusalem, 1962-65.
Idem, “Presnovodnye ryby Irana i sopredel’nykh stran"(Freshwater fishes in Iran and neighboring countries), Trudy Zoologicheskogo instituta Akademii nauk SSSR 8, 1949, 783–858.
P. G. Bianco, and P. Banarescu, “A Contribution to the Knowledge of the Cyprinidae of Iran (Pisces, Cypriniformes),” Cybium 6/2, 1982, pp. 75–96.
“The Blind White Fish of Persia,” Nature (London) 264, 1976, pp. 113-14.
N. Borodin, “Acipenser persicus, a Sturgeon from the Caspian Sea,” The Annals and Magazine of Natural History 20/9, 1926, pp. 26-28.
A. F. Bruun and E. W. Kaiser, “Iranocypris typhlops n.g., n.sp., the First True Cave Fish from Asia,” Danish Scientific Investigations in Iran (Copenhagen) 4, 1944, pp. 1-8.
B. W. Coad, “Poisonous and Venemous Freshwater Fishes of Iran,” Pahlavi Medical Journal 9/4, 1979a, pp. 388-407.
Idem, “Range Extension for the Snakehead Ophiocephalus gachua Hamilton-Buchanan (Osteichthyes: Channidae) in Iran,” Journal of the Bombay Natural History Society 75/2 (1978), 1979b, pp. 500-501.
Idem, “Environmental Change and Its Impact on the Freshwater Fishes of Iran,” Biological Conservation 19/1, 1980a, pp. 51-80.
Idem, “A Provisional, Annotated Check-list of the Freshwater Fishes of Iran,” Journal of the Bombay Natural History Society 76/1 (1979), 1980b, pp. 86-105.
Idem, “A Re-description of Aphanius ginaonis (Holly, 1929) from Southern Iran (Osteichthyes: Cyprinodontiformes),” Journal of Natural History 14/1, 1980c, pp. 33-40.
Idem, “First Record of the Milkfish, Chanos chanos (Forskal, 1775) from Iran and the Persian Gulf,” Journal of the Bombay Natural History Society 77/3, 1981a, pp. 522-24.
Idem, “Glyptothorax silviae, a New Species of Sisorid Catfish from Southwestern Iran,” Japanese Journal of Ichthyology 27/4, 1981b, pp. 291-95.
Idem, “Pseudophoxinus persidis, a New Cyprinid Fish from Fars, Southern Iran,” Canadian Journal of Zoology 59/11, 1981c, pp. 2058-63.
Idem, “Garra persica Berg, 1913, a Valid Species of Cyprinid Fish from Southern Iran,” Cybium 6/2, 1982a, pp. 97-100.
Idem, “The Identity of Alburnus maculatus Keyserling, a Cyprinid Fish from Esfahan Province, Iran,” Japanese Journal of Ichthyology 29/2, 1982b, pp. 227-28.
Idem, “A New Genus and Species of Cichlid Endemic to Southern Iran,” Copeia, 1982c, pp. 28–37.
Idem, “A Re-description and Generic Re-assignment of Kosswigobarbus kosswigi (Ladiges, 1960), a Cyprinid Fish from Turkey and Iran,” Mitteilungen aus dem hamburgischen Zoologischen Museum und Institut 79, 1982d, pp. 263-65.
Idem, “Alburnus doriae De Filippi, 1864, a Synonym of Leuciscus lepidus (Heckel, 1843), (Osteichthyes: Cyprinidae),” Matsya 9-10 ,1985, pp. 173-75.
Idem, “Zoogeography of the Freshwater Fishes of Iran,” in F. Krupp, W. Schneider and R. Kinzelbach, eds., Proceedings of the Symposium on the Fauna and Zoogeography of the Middle East, Mainz, 1985, TAVO, ser. A (Naturwissenschaften) 28, Wiesbaden, 1987, pp. 213-28.
Idem, “Aphanius vladykovi, a New Species of Tooth-Carp from the Zagros Mountains of Iran (Osteichthyes: Cyprinodontidae),” Environmental Biology of Fishes 23/1-2, 1988, pp. 115-25.
Idem, “Fishes of the Tigris-Euphrates Basin: A Critical Checklist,” Syllogeus (Ottawa), 68, 1991, pp. 1-49.
Idem, “Freshwater Fishes of Iran,” Acta Scientiarum Naturalium Academiae Scientiarum Bohemicae (Brno) 29/1, 1995, pp. 1-64.
Idem, 1996a. “Systematics of the Tooth-carp Genus Aphanius Nardo, 1827 (Actinopterygii: Cyprinodontidae) in Fars Province, Southern Iran,” Biologia (Bratislava) 51/2, 1996a, pp. 163-72.
Idem, 1996b. Systematics of the Shah Mahi, Chalcalburnus chalcoides (Güldenstädt, 1772), in the Southern Caspian Sea Basin (Actinopterygii: Cyprinidae),” Zoology in the Middle East 12, 1996b, pp. 65-70.
Idem, “Threatened Fishes of the World: Iranocypris typhlops Bruun and Kaiser, 1944 (Cyprinidae),” Environmental Biology of Fishes 46(4), 1996c, p. 374.
Idem, “Zoogeography of the Fishes of the Tigris-Euphrates Basin,” Zoology in the Middle East 13, 1996d, pp. 51-70.
Idem, 1996e. “Exotic Fish Species in the Tigris-Euphrates Basin,” Zoology in the Middle East 13, 1996e, pp. 71-83.
Idem, “Fishes from the Qanats of Iran,” Publicaciones Especiales Instituto Español de Oceanografía, 21, 1996f, pp. 63-79.
Idem, “Exotic and Transplanted Fishes in Southwest Asia,” Publicaciones Especiales Instituto Español de Oceanografía 21, 1996g, pp. 81-106.
Idem, “Freshwater Fishes of Iranian and Pakistani Baluchistan,” Biologia (Lahore) 42/1-2 (1996), 1997a, pp. 1-18.
Idem, “Shad in Iranian Waters,” The Shad Journal (Seattle) 2/4, 1997b, pp. 4-8.
Idem, “Threatened Fishes of the World: Lebias ginaonis (Holly, 1929)(Cyprinodontidae),” Environmental Biology of Fishes 51/3, 1998, p. 284.
B. W. Coad and A. Abdoli, “Exotic Fish Species in the Fresh Waters of Iran,” Zoology in the Middle East 9, 1993, pp. 65-80.
Idem, “Biodiversity of Iranian Freshwater Fishes,” Abzeeyan/Ābzīān (Tehran) 7/1, 1996, pp. iv, 4-10 (in Persian with English abstract).
B. W. Coad and L. A. J. Al-Hassan, A Bibliography of the Fishes of the Tigris-Euphrates Basin/Bibliographie der Fische des Euphrat-Tigris-Beckens, Heidelberg, 1989.
B. W. Coad and J. Holcík, “Systematics of the Cyprinid Fish Chalcalburnus atropatenae (Berg, 1925) from the Lake Orumiyeh Basin in Northwest Iran,” Biologia (Bratislava) 54/2, 1999, pp. 179-86.
B. W. Coad and F. Krupp, “Redescription of Barilius mesopotamicus Berg, 1932, a Poorly Known Cyprinid Fish from the Tigris-Euphrates Basin,” Cybium 7/1, 1983, pp. 47-56.
Idem, “Capoeta aculeata (Valenciennes in Cuv. and Val., 1844), a Valid Species of Cyprinid Fish from Iran (Teleostei: Cyprinidae),” Zoology in the Middle East 10, 1994, pp. 63-72.
B. W. Coad and N. Najafpour, “Barbus sublimus, a New species of Cyprinid fish from Khuzestan Province, Iran,” Ichthyological Exploration of Freshwaters 7/3, 1997, pp. 273-78.
B. W. Coad and F. Papahn, “Shark Attacks in the Rivers of Southern Iran,” Environmental Biology of Fishes 23/1-2, 1988, pp. 131-34.
G. Cuvier and A. Valenciennes, Histoire naturelle des poissons, 22 vols., Paris, 1828-49.
F. De Filippi, “Nuove o poco note specie di animali vertebrati raccolte in un viaggio in Persia nell’estate dell’anno 1862,” Archivio per la Zoologia, l’Anatomia e la Fisiologia 2, 1862, pp. 377–94.
Idem, “Riassunto del catalogo degli animali vertebrati delle provincie caucasiche e della Persia occidentale,” Atti della Società Italiana di Scienze Naturali 7, 1864, pp. 184-86.
F. De Filippi, Note de un viaggio in Persia nei 1862, Milan, 1865, “Pesci,” pp. 3, 63, 141, 357-60.
T. De Meulenaer and C. Raymakers, Sturgeons of the Caspian Sea and the International Trade in Caviar, Cambridge, U.K., 1996.
A. N. Derzhavin, “Iskusstvenoe razvedenie osetra na Sefid-Rude” (Artificial sturgeon-breeding on the Safīd Rūd), Russkiĭ gidro-biologischeskiĭ zhurnal 2/11-12, 1923, p. 250.
Idem, “Zametka o rybakh reki Keredzh, sev. Persiya” (A note on fishes of the river Karaj, northern Persia),” Izvestiya Bakinskoĭ ikhtiologicheskoĭ laboratorii 2/2, 1929, pp. 69-79.
Idem, “Presnovodnye ryby yuzhnogo poberezh’ya Kaspiya” (Freshwater fishes of the southern shore of the Caspian Sea), Trudy Azerbaidzhanskogo otdela Zakavkazskogo filiala Akademii nauk SSSR, Sektor zoologii 7, Baku, 1934, pp. 91–126.
N. A. Dmitriev, “Kefal’ v iranskikh vodakh Kaspiya” (Mullet in the Iranian waters of the Caspian), Priroda 12, 1946, pp. 74–75.
Idem, “Nekotorye dannye po biologii pekhleviĭskogo (enzeliĭskogo) puzanka Caspialosa caspia knipowitschi Iljin” (Some biological data on the Pahlavi [Enzeli] shad Caspialosa caspia knipowitschi Iljin), Zoologicheskiĭ zhurnal 26/6, 1947, pp. 559–60.
B. Elvira, “Taxonomic Revision of the Genus Chondrostoma Agassiz, 1835 (Pisces, Cyprinidae),” Cybium 11/2, 1988, pp. 111-40.
S. Etessami, “L’histologie des gonades chez deux cyprinides Alburnoides bipunctatus eichwaldi (Filipi, 1863) (sic) et Barbus mursa miliaris (Karaman, 1972) (sic) avec la description d’un cas d’hermaphrodisme chez ce dernier,” Cybium 6/2, 1982, pp. 5-13.
Food and Agriculture Organization of the United Nations, “Chinese Carps in Iran,” Aquaculture Bulletin 3/1, Rome, 1970a, p. 15.
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Table 1. Numbers of freshwater fishes.
Table 2. Distribution of freshwater fishes.
Figure 1. The Tigris salmon (Barbus esocinus). 2.1 m.
Figure 2. The bitterling (Rhodeus amarus). 18.0 cm., female.
Figure 3. Iranocypris typhlops. 5.3 cm.
Figure 4. Nemacheilus smithi. 6.5 cm.
Figure 5. Minnow (Capoeta damascina). 45.0 cm.
Figure 6. Herring (Tenualosa ilisha). 60.6 cm.
Figure 7. Tooth-carp (Aphanius dispar). 7.3 cm.
Figure 8. Mullet (Liza abu). 26.0 cm.
Figure 9. Capoeta aculeata. 20.0 cm.
Figure 10. Barilius mesopotamicus. 5.1 cm.
Figure 11. Bagrid catfish (Mystus pelusius). 22.9 cm.
Figure 12. Loach (Cobitis linea). 8.9 cm.
Figure 13. Iranocichla hormuzensis. 9.7 cm.
Figure 14. Tench ( Tinca tinca). 70.0 cm.
Figure 15. Snow trout (Schizochorax zarudnyi). 49.0 cm.
Figure 16. Indian stinging catfish (Heteropneustes fossilis). 30.5 cm.
Figure 17. Bull shark (Carcharhinus leucas). 3.24 m.
Figure 18. Mosquitofish (Gambusia holbrooki). 6.3 cm.
Figure 19. Snakehead (Channa gachua). 33.0 cm.
Figure 20. Holly’s pupfish (Aphanius ginaonis). 4.0 cm.
(Brian W. Coad)
Originally Published: December 15, 1999
Last Updated: January 26, 2012
This article is available in print.
Vol. IX, Fasc. 6, pp. 655-668
Brian W. Coad, “FISH i. FRESHWATER FISHES,” Encyclopædia Iranica, IX,6, pp. 655-668; available online at http://www.iranicaonline.org/articles/fish-i (accessed online at 17 August 2012).