Freshwater Fishes of Iran

Species Accounts

Cobitidae to Cyprinodontidae

Revised:  06 August 2013

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Cobitidae

This family of loaches, sometimes called sting-loaches, is found in Eurasia and Morocco and has about 28 genera with about 236 species (Berra, 2001; Nelson, 2006; Eschmeyer and Fong, 2011). Berra (2001) does not indicate the more southern distribution of this genus in Khuzestan and Fars provinces of Iran. Five to six species are recorded from Iran.

Anonymous (1988a) places Cobitidae on the Official List of Family-Group Names in Zoology (rather than the grammatically correct but unused Cobitididae) and Cobitis taenia is designated as the type species for the genus Cobitis (see also Kottelat (1986) for further information).

The body form is fusiform to rounded or elongate; the mouth is subterminal and has 3-6 pairs of barbels; the mental lobes of the lower lip have two parts: the anterior which is usually short and sometimes divided into lobules, and the posterior which is flap-like and longer and sometimes divided into 2 or more barbel-like extensions; there is 1 row of pharyngeal teeth; and there is an erectile spine in a groove below the eye (anterior in a non-Iranian genus). Iranian species have one pair of rostral barbels and a rounded or slightly emarginate caudal fin and belong to the subfamily Cobitinae. Menon (1992) considers that structural details of the bony covering of the swimbladder and the nature of the scales are only of use at the generic level. Lip structures, fin positions relative to one another and secondary sexual characteristics in males are important characters in differentiating species in India. Economidis and Nalbant (1996) discuss characters used in the study of these fishes and consider scales to be characteristic of each species along with colour pattern, sexual dimorphism, suborbital spine morphology, barbel and mental lobe morphology, and others. Black spots at the caudal fin base and four longitudinal pigment zones on the flank (Z1-4 or Gambetta's zones 1-4) are important in distinguishing and describing species. Males have 1-2 laminar projections on the dorsal surface of the anterior pectoral fin rays known as the laminae circularis or Canestrini's scales. Hybrid lineages are known, produced by gynogenesis and are nearly all-female. Males of bisexual lineages are sperm donors but the sperm only induces egg development and contributes no genetic material. The all-female lineages are therefore sperm parasites and have to occur in sympatry with one of the parental species in a hybrid complex. As a result, hybridogenous individuals in the complex are difficult to distinguish from the bisexual parent species on external characters. An unbalanced sex ratio in a population with more females than males is usually evidence that hybridogenous lineages are present (Kottelat and Freyhof, 2007). Maximum size is about 40 cm but most are much smaller.

The origins of this group of loaches may well lie at the end of the Eocene or in the early Oligocene in South China, spreading along a northern route through Europe and Siberia during the Oligocene-Miocene-Pliocene period and then later southwards into Southwest Asia (Sawada, 1982; Menon, 1987; 1992; Bănărescu and Nalbant, 1998; Šlechtová et al., 2008; Tang et al., 2008). An early Oligocene route also existed between the Anatolian landmass and Central Asia (Tang et al., 2008) and cobitids may have invaded the Euro-Mediterranean zoogeographic subregion at least five times independently based on cytochrome b data.

Some members of this family can live in oxygen-poor waters. They take in air at the surface, and pass it through the intestine where the mucosa absorbs the oxygen and carbon dioxide waste is released through the vent. As a consequence, they may be very sensitive to air pressure changes and become restless when it falls and can be used to predict the weather. Foods are mostly small insects, worms and crustaceans detected by the aid of the barbels on the habitat bottom. Some eat algal films or mats. Most species bury themselves in sand or mud during the day, emerging to feed at night. Movement is by undulations of the body, particularly marked in the more elongate species. A consequence of this form of movement is a reduction in fin size and variation. Reproduction involves the male chasing the female, entering vegetation and wrapping around the female as eggs are released and fertilised. Eggs swell and reach as large as 3.5 mm in diameter and as a result are retained in the vegetation although they are not adhesive.

A number of species are popular aquarium fishes, including the coolie or kuhli loaches and the weatherfish. None of the Iranian fishes are used in this fashion but they can be quite colourful. Cobitis cf. taenia is a potential fishing bait for predatory fishes such as Sander lucioperca and has been examined experimentally for this purpose in Turkey (Kuşat et al., 1995).

Genus Cobitis
Linnaeus, 1758

These fishes are found in Europe, North Africa and Asia. There are 3-4 species in Iran. They are known generally as سگ ماهي (sag mahi meaning dog fish) in Farsi, the equivalent of loach in English. This name is not repeated under each Species Account. Also called لوچ (= louch in Farsi, from loach).

This genus is characterised by an elongate and compressed body, a usually bifid, erectile spine below the eye (sometimes hidden under the skin), 3 pairs of short barbels (4 at the snout tip and 2 at the mouth corners), minute scales cover the body (as many as 200 but they are seldom counted accurately), lateral line faint or indistinct, dorsal and anal fins small, caudal fin rounded or truncate, and swimbladder in a bony capsule with a free portion visible. Males have bony extensions of their pectoral fin rays, known as lamina circularis or scale of Canestrini, and no swellings of their body sides.

Polyploid unisexual, bisexual-unisexual complexes and gynogenetic forms of Cobitis exist in the basins of the Baltic, Black, Caspian and Mediterranean seas (Vasil'ev and Vasil'eva, 1996; Kottelat, 1997; Vasil'eva and Vasil'ev, 1998; Vasil'ev et al., 1999; Bohlen, 2001; Vasil'ev et al., 2011). The species are morpohologically undifferentiated and therefore require detailed study to resolve taxonomic and systematic problems. The composition of Iranian species has not been investigated throroughly.

The earliest fossil record is from the middle Miocene about 15MYA with divergence of Sabanejewia and Cobitis at 12-13MYA (Ludwig et al., 2001).

These loaches often bury themselves in mud to overwinter or escape predators. The spine under the eye when erected is an anti-predator device, discouraging swallowing by other fishes and birds. The fish is said to actively swing the head from side to side to prick predators. Vasil'ev et al. (2011) review evolutionary ecology of clonal-bisexual complexes. Clonal forms, for example, are polyploids and have larger eggs and therefore reduced individual fecundity, and hatchlings are larger, perhaps with higher fitness initially in a local environment. The usual population fecundity of clonal forms being twice that of a bisexual species does not apply here because of the larger eggs.

Cobitis amphilekta
Vasil'eva and Vasil'ev, 2012

Described from the Kyzylagach Bay in Azerbaijan and the northeastern Caspian Sea, this taxon has not been recorded from Iran.

Cobitis faridpaki
Mousavi-Sabet, Vasil'eva, Vatandoust and Vasil'ev, 2011

Common names

Faridpak's spine loach. Some Farsi names formerly under C. taenia (see C. keyvani) may apply here as some Caspian Iranian Cobitis may be this species.

Systematics

The holotype is a male, 51.3  mm standard length, Mazandaran, Siahrud River (sic), 36º26'85.05"N, 52º56'70.08"E under GUIC (Ichthyological Museum, Department of Fisheries, Natural Resources Faculty, University of Gilan, Rasht) CC1403MA. Paratypes are GUIC CC1403M, 3 specimens, 54.7-62.3 mm standard length, same locality as holotype, C1404M, 9, 48.5-61.4 mm standard length, same locality as holotype, GUIC CC1405, 10, 49.2-67.4 mm standard length, same locality as holotype, ZMMU (Zoological Museum of Moscow State University) P-22694, 2, 50.6-63.2 mm standard length, same locality as holotype.

Cobitis taenia Linnaeus, 1758 was formerly the species for Iranian Caspian Cobitis (see below under C. keyvani) and would have been used for this taxon in part.

The new species is named for Farhad Faridpak (1911-1996) an Iranian ichthyologist working on Caspian Sea fishes.

Key characters

This species is distinguished from the other members of the genus in Iran by lacking a second lamina Canestrini at the base of the first pectoral fin ray (C. linea) and by lacking large flank spots (C. keyvani).

Morphology

The body is compressed and relatively deep. Subdorsal scales are rounded with a reduced eccentric focus. The single lamina Canestrini is axe-shaped and relatively broad. Dorsal fin with 2-3 unbranched and 6-7, usually 7, branched rays, anal fin with 2-3 unbranched and 5-7, usually 7, branched rays, pectoral fin with 7-9 branched rays, pelvic fin with 6-7 branched rays, and caudal fin with 13-16, usually 14(64%) or 16(32%), branched rays. Total vertebrae 40-41, usually 40 and gill rakers 11-12, usually 11.

Sexual dimorphism

Females are more stout than males and reach a longer body length. Male barbels and pectoral fins are longer and males have lamina Canestrini.

Colour

Flanks are pale yellow in life with 25-30 dark brown spots or blotches, smaller than eye diameter and usually merged. The Gambetta's zones of pigment are nearly complete, the fourth zone composed of the 25-30, small spots. The second Gambetta's zone on the upper flank is a narrow zone of brown spots between the first and third zones both of which are wider and composed of small dots. The lower head and abdomen are yellowish-white. The head is darkly mottled with dots and there is usually a dark stripe from the eye to the snout tip. Males have almost orange dorsal and caudal fins with dark spots. Females have yellow-orange dorsal and caudal fins with 3-4 rows of dark spots. Other fins are whitish without pigmentation. The caudal fin base has a black spot smaller than eye diameter on its upper part (sometimes absent),

Size

Attains 78.2 mm total length.

Distribution

Described from the Siahrud or Siah River in Mazandaran and presumably present also in the Babol River (Mousavi Sabet et al., 2011), if not all the rivers of the Iranian Caspian coast.

Zoogeography

See genus and family account.

Habitat

Unknown.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

None.

Conservation

Known only from a restricted area with no data on numbers and threats.

Further work

The biology of this species and its conservation status need study.

Sources

The type description.

Cobitis keyvani
Mousavi-Sabet, Yerli, Vatandoust, Özeren and Moradkhani, 2012

Anzali Wetland, June 2009, courtesy of K. Abbasi
Anzali Wetland, June 2009, courtesy of K. Abbasi

 CMNFI 1979-0435, 1, 62.0 mm standard length, Gilan, stream 10 km west of Ramsar (36°57'N, 50°37'E), B. W. Coad
CMNFI 1979-0435, 1, 62.0 mm standard length, Gilan, stream 10 km west of Ramsar (36°57'N, 50°37'E), B. W. Coad

Common names

mahi roshtegar or roftegar (= dustman, cleaner or sweeper fish, presumably from the bottom-dwelling habit); sagmahi-ye sangi (= stone loach), sagmahi-ye juibari or sagmahi-e-jooibari, mar mahi (= snake fish); loch or louch in Khuzestan (presumably from the English name, but also louch means a person with a squint); gel khorak in Khuzestan (latter two not necessarily this taxon, see below).

[lakh mukhattat in Arabic; zagafgaziya iliskani in Azerbaijan; shchipovka or shchipovka zakavkazskaya or Transcaucasian spiny loach in Russian; spined or spiny loach, stone loach, spiny stone loach, weatherfish, spotted weatherfish, Siberian loach; all these being for C. taenia but the Azerbaijani and Russian names presumably applicable to this species].

Systematics

Kottelat and Freyhof (2007) considered that the taxon in the southern Caspian Sea basin was not Cobitis taenia Linnaeus, 1758 as long used. C. taenia is restricted to northern and eastern Europe. Cobitis Taenia was originally described from Europe although Kottelat (1997) gives Lake Mälaren, Sweden.

Cobitis taenia satunini Gladkov, 1935 has been considered the subspecies or species of some southern Caspian Sea basin Cobitis, distinguished by a truncate snout rather than gradually tapering to the tip as in European spined loaches as well as other characters listed by Gladkov (1935) including head depth comprising 63.5% of head length on average and 12.27% of standard length, and caudal peduncle shorter than the head (on average 16.27% of standard (?) length). However, it was described from western Transcaucasia, outside the Caspian Sea basin, and material from the Caspian coast of Iran has not been examined and compared with it in detail. Berg (1948-1949) referred specimens from the upper Kura River basin to this subspecies but lacked material from the lower Kura, Lenkoran and the southern Caspian Sea coast.

The holotype of Cobitis taenia satunini described from "Kavkaz" and "nizov'ya r. Kintrish" (= lower Kintrish stream in the Caucasus; the Kintrish stream mouth is at 41°48'N, 41°46'E) is in the Zoological Museum of Moscow State University under MMSU P.2852 with a cotype (= paratype) under MMSU P.2317 (Eschmeyer et al. (1996) has MMSU P.2251, quoting Svetovidova (1978), which disagrees with Gladkov (1935)). The Zoological Museum of Moscow University (ZMMU; their acronym) has P-2852 as the holotype plus P-2313 as 2 paratypes (Pavlinov and Borissenko, 2001).

The holotype of Cobitis keyvani is a female, 62.9 mm standard length, "from the Keselian stream, Talar River, southeast of the Caspian Sea basin, Mazandaran province, north of Iran, 36°11’74.09 "N, 53°00’92.01"E", catalogued in the Ichthyology Museum, Department of Fisheries, Faculty of Natural Resources, the University of Gilan as GUIC CC1389MA. Paratypes are GUIC CC1389M, 8,, 37.4-80.3 mm standard length, GUIC CC1390M, 6, 44.4-67.4 mm standard length, and GUIC CC1391, 5, 48.3 and 68.9 mm standard length (sic), all same locality as holotype. The species is named for Prof. Dr. Amin Keyvan (1930-2007), Iranian ichthyologist.

Key characters

This species is distinguished from the C. linea by lacking a second lamina Canestrini at the base of the first pectoral fin ray, and from C. faridpaki by large, dark and obvious spots along the mid-flank.

Morphology

Some specimens are elongate while others are deeper bodied. Dorsal fin unbranched rays 2-3, branched rays 6-7, anal fin unbranched rays 2-3, branched rays 4-7, usually 5, pectoral fin branched rays 6-9, usually 7-8, pelvic fin branched rays 5-7, usually 6, caudal fin branched rays 10-16, usually 14, gill rakers 9-10, and total vertebrae 39-40.

Meristics for Iranian specimens:- dorsal fin branched rays 6(29) or 7(51); anal fin branched rays 4(1), 5(78) or 7(1); pectoral fin branched rays 6(4), 7(35), 8(38) or 9(3); pelvic fin branched rays 5(6), 6(70) or 7(4); caudal fin branched rays 10(1, but deformed), 12(2), 13(7), 14(68), 15(1) or 16(1); total vertebrae ?. Flank spots number 9(1), 10(2), 11(9), 12(17), 13(14), 14(18), 15(16), 16(14), 17(8), 18(11), 19(5), or 20(3) (excluding fish with bars, see below). The type description gives 14-20 (and later 17-24) flank spots. The type material has counts as follows:- dorsal fin branched rays 6(10) or 7(10); anal fin branched rays 5(17) or 7(3); pectoral fin branched rays 6(2), 7(12) or 8(6); pelvic fin branched rays 5(2), 6(13) or 7(5); caudal fin branched rays 13(1), 14(12), 15(1) or 16(6); total vertebrae 39(1) or 40(19); and gill rakers 9(1) or 10(19).

Sexual dimorphism

The second ray of the male pectoral fin is thickened and there is an enlarged scale at the base (Canestrini scale). The body in front of the dorsal fin of males is not distended as in S. aurata. Females are longer than males, and pectoral fins and barbels are longer in males (Mousavi-Sabet et al., 2012).

Colour

The back is light brown, the flanks pale yellow and the belly and lower head are yellowish-white. Five lateral pigmentation zones exist with L1 narrower than L2, L3 with 20-28 obvious dark speckles, L4 with a few small spots and reduced in the postdorsal fin part, L2 and L4 zones with irregular spots, and L5 zone wider than L1 and L3 (Mousavi-Sabet et al., 2012). The flank spots of L5 are usually clear and distinct, usually round but sometimes squarish, and some can be poorly defined, especially on the anterior flank. Anterior flank spots may merge to form short bars although their origin from spots can still be discerned. Rarely a long bar forms extending anteriorly from the mid-dorsal fin level. All fish have several large and distinct spots on the posterior flank. The number of spots can vary between high and low counts within the same sample, and are readily visible on even the smallest fish. The base of the caudal fin rays has a dark spot dorsally, much denser in pigmentation than flank spots. Some fish have a ventral caudal fin spot as well. Large spots along the mid-line of the back may be present or absent. The dorsal and caudal fins have spots or bars of pigment arranged in 3-5 rows. Other fins are whitish. The head is mottled with brown dots. A bar extends from the eye antero-ventrally. A postero-dorsal bar is present in some fish, meeting the one from the opposite side at the midline of the back of the head. This bar can be a row of spots or there may be no discernible pattern in this area. The iris is silvery, slightly golden or orange.

Size

Attains 80.3 mm standard length (Mousavi-Sabet et al., 2012).

Distribution

Found from Astara to Gorgan Bay including the Anzali Mordab, the Aras, Safid, Pol-e Rud, Sardab, Takar, Nessa, Haraz, Talar, Babol. Chowbar, Qasemabad, Qa'emshahr, Tonekabon, Shirud, Tajan and Gharasu rivers, and from the upper Karkheh.

Reports of Cobitis taenia from the middle Dez, Kashkan, Qareh Chai, Simarreh and lower Gav Masiab rivers in the Tigris basin may represent another new species (Saadati, 1977; Roshan Tabari, 1997; Abbasi et al., 1999; Kiabi et al., 1999; Abdoli, 2000; Abdoli and Naderi, 2009). An anecdotal report of a Cobitis from the Qareh Chai near Hamadan in the Namak Lake basin needs specimens for confirmation.

Zoogeography

Records from the Tigris River basin of Iran are based on literature (Saadati, 1977) and a ???single specimen (CMNFI 1979-0285). Mid-flank spots are less distinct than in Cobitis keyvani from the Caspian Sea basin of Iran of similar size being smaller and more numerous and the stripe on the centre of the back is continuous rather than spots as in most Caspian fish. Identification is tentative.

Habitat

This species remains buried in sand, mud which is not too thick, or dense weed growths during the day, being active at night, and is mostly solitary. Swimming is by undulating motions over short distances. When concealed, the body is bent into an arch so only the head and tail protrude. It prefers cool, clear running waters. Along the Caspian shore it is found in the lower reaches of rivers (Jolodar and Abdoli, 2004).

Age and growth

Females dominated the population in the Safid River and growth was positively allometric (Patimar et al., 2010). ??Mousavi Sabet et al. (2011) examined fish from the Babol River and found the percentage of females was significantly higher than males, and mature females were longer than 45 mm total length and 2+ years old and mature males were longer than 35 mm and 1+ years old. Life span was 5+ years for females and 3+ years for males. Mousavi-Sabet et al. (2012) found the population in the Talar River had significantly more females than males, females were mature at lengths longer than 49 mm and were at age 3+ while males matured at lengths longer than 45 mm and were at age 1+, and maximum age was 5+ years.

Food

Diet for related C. taenia is small crustaceans such as ostracods, copepods and rotifers in the bottom mud or sand. A mouthful of mud or sand is taken in, chewed, food items extracted, and the residue expelled convulsively through the gill openings.

Reproduction

Spawning of related C. taenia takes place from April to June in slow to still water. Eggs are laid on sand, stones and vegetation in several batches. Eggs may be deposited on the roots of water plants cleared of debris by males rooting among them. Males use their enlarged pectoral fins to turn the female during spawning.

The population in the Safid River had egg diameters up to 1.02 mm, absolute fecundity reached 8111 eggs and relative fecundity up to 383 eggs/g (Patimar et al., 2010). ??Mousavi Sabet et al. (2011) found Babol River fish spawning from the beginning of May to late July at water temperatures of 19.1-24.6ºC. Average absolute and relative fecundities were2172 and 590 eggs with ranges of 734-3562 and 347 to 945 eggs, respectively. Egg diameters were up to 1.4 mm with an average of 0.58 mm. Mousavi-Sabet et al. (2012) found the population in the Talar River had an average egg diameter of 0.56 mm, a maximum egg diameter of 1.4 mm, spawning took place from May to late July at 18.7-24.0°C, and mean (and maximum) absolute and relative fecundities were 2211 (3319) and 586 (902) eggs respectively.

Parasites and predators

This species (identified as C. taenia) is infected with Clinostomum complanatum, a parasite that can cause laryngo-pharyngitis in humans, in the Shirud of western Mazandaran Province (Shamsi et al., 1997).

Economic importance

None. Palicka (1996) gives a short account of aquarium care for C. taenia which may prove applicable to this species.

Conservation

This species was commonly caught in Iranian streams along the Caspian coast. Kiabi et al. (1999) consider it (as C. taenia) 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 (the latter two criteria no longer applicable).

Further work

The biology of this species needs study.

Sources

Iranian material: CMNFI 1970-0511, 35, 32.0-74.1 mm standard length, Gilan, Shafa River estuary (37°35'N, 49°09'E); CMNFI 1970-0514, 39, 37.6-79.4 mm standard length, Gilan, Shafa River estuary (37°35'N, 49°09'E); CMNFI 1970-515, 22, 38.8-73.7 mm standard length, Gilan, Shafa River estuary (37°35'N, 49°09'E); CMNFI 1970-0516, 2, 44.2-52.1 mm standard length, Gilan, Limir River at Lomir (38°14'N, 48°52'30"E); CMNFI 1970-0518, 4, 32.4-69.1 mm standard length, Gilan, Haviq River estuary (38°10'N, 48°54'E); CMNFI 1970-0519, 5, 29.6-39.2 mm standard length, Gilan, Chelvand River (ca. 38°18'N, ca. 48°52'E); CMNFI 1970-0525, 1, 20.4  mm standard length, Gilan, Safid River near Mohsenabad (ca. 37°22'N, ca. 49°57'E); CMNFI 1970-0526, 3, 32.9-39.1 mm standard length, Gilan, Safid River 5 km below Astaneh bridge (37°19'N, 49°57'30"E); CMNFI 1970-0527, 12, 27.4-69.7 mm standard length, Gilan, ditch near Kisom (37°12'N, 49°54'E); CMNFI 1970-0534, 16, 31.2-51.2 mm standard length, Gilan, Shafa River estuary (37°35'N, 49°09'E); CMNFI 1970-0535A, 3, 29.0-48.3 mm standard length, Gilan, Pir Bazar Roga (37°21'N, 49°33'E); CMNFI 1970-537, 16, 38.6-76.9 mm standard length, Markazi, Shah River near Manjil (36°44'N, 49°24'E); CMNFI 1970-0542, 37, 19.5-70.7 mm standard length, Gilan, Old Safid River estuary (37°23'N, 50°11'E); CMNFI 1970-0545, 14, 41.7-64.8 mm standard length, Gilan, Safid River (ca, 37°01'N, ca, 49°38'E); CMNFI 1970-0548, 33, 36.3-63.8 mm standard length, Mazandaran, Qareh Su (36°49'30"N, 54°03'30"E); CMNFI 1970-0549, 18, 14.5-22.8 mm standard length, Mazandaran, Qareh Su near Alm Emamzadeh (no other locality data); CMNFI 1970-0553, 13, 15.8-35.6 mm standard length, Gilan, Sowsar Roga (37°27'N, 49°30'E); CMNFI 1970-0554, 7. 17.5-33.8 mm standard length, Gilan, Pir Bazar and Sowsar Roga (37°21'N, 49°33'E and 37°27'N, 49°30'E); CMNFI 1970-0562, 1, 20.4 mm standard length, Gilan, Sowsar Roga (37°27'N, 49°30'E); CMNFI 1970-0566, 16, 19.5-39.9 mm standard length, Gilan, Old Safid River estuary (37°23'N, 50°11'E); CMNFI 1970-0567, 53, 15.9-38.3 mm standard length, Gilan, Pir Bazar Roga (37°21'N, 49°33'E); CMNFI 1970-0574, 5, 30.0-50.6 mm standard length, Gilan, Sowsar Roga (37°27'N, 49°30'E); CMNFI 1970-0575, 8, 24.8-32.9 mm standard length, Gilan, Pir Bazar Roga (37°21'N, 49°33'E); CMNFI 1970-0576, 3, 35.8-46.1 mm standard length, Gilan, Shafa River estuary (37°55'N, 49°09'E); CMNFI 1970-0579, 63, 26.2-71.9 mm standard length, Gilan, Old Safid River estuary (37°23'N, 50°11'E); CMNFI 1970-0580, 67, 31.1-48.5 mm standard length, Mazandaran, Iz Deh (36°36'N, 52°07'E); CMNFI 1970-0582, 1, 60.9 mm standard length, Mazandaran, Aliabad Reservoir (36°56'N, 54°50'E); CMNFI 1970-0587, 4, 46.4-53.1 mm standard length, Mazandaran, Babol Sar (36°43'N, 52°39'E); CMNFI 1970-590, 5, 24.5-70.0 mm standard length, Mazandaran, Shesh Deh River near Babol Sar (ca. 36°43'N, ca. 52°39'E); CMNFI 1979-0430, 2, 55.2-76.1 mm standard length, Mazandaran, river 1 km east of Now Shahr (36º39'N, 51º31'E); CMNFI 1979-0435, 1, 62.0 mm standard length, Gilan, stream 10 km west of Ramsar (36°57'N, 50°37'E); CMNFI 1979-0446, 4, 43.1-62.3 mm standard length, Gilan Astara River 2 km from Astara (38°26'30"N, 48°51'E); CMNFI 1979-0470, 8, 32.9-53.9 mm standard length, ; CMNFI 1979-0472, 9, 43.8-56.9 mm standard length, Mazandaran, stream 7 km west of Mahmudabad (36°37'N, 52°12'E); CMNFI 1979-0476, 1, 25.6 mm standard length, Mazandaran, Qareh Su 6 km from Kord Kuy (36°51'N, 54°05'E); CMNFI 1979-0479, 1, 47.6 mm standard length, Mazandaran, dam on Gorgan River 20 km north of Eimar (37°09'30"N, 54°41'30"E); CMNFI 1979-0626, 1, 49.5 mm standard length, Gilan, Safid River (no other locality data); CMNFI 1979-0685, 6, 26.8-44.2 mm standard length, Gilan, Safid River below Dehcha (ca. 37°22'N, ca. 49°57'E); CMNFI 1979-1215B, 5, 21.7-68.8 mm standard length, Gilan, Sowsar Roga (37°27'N, 49°30'E) ; CMNFI 1980-0117, 1, 47.4 mm standard length, Gilan, Golshan River (37°26'N, 49°40'E); CMNFI 1980-0122, 8, 34.4-47.9 mm standard length, Mazandaran, Nevissi River (36°38'N, 52°16'E); CMNFI 1980-0130, 28, 33.2-42.5 mm standard length, Mazandaran, Iz Deh (36°36'N, 52°07'E); CMNFI 1980-0132, 32, 25.8-52.8 mm standard length, Gilan, Safid River at Kisom (37°12'N, 49°54'E); CMNFI 1980-0136, 6, 31.0-65.9 mm standard length, Mazandaran, Fereydun Kenar River estuary (36°41'N, 52°29'E); CMNFI 1980-0138, 1, 31.8 mm standard length, Gilan, Safid River estuary (ca. 37°28'N, ca. 49°54'E); CMNFI 1980-0140, 1, 33.9 mm standard length, Gilan, Astara Mordab (ca. 38°26'N, ca. 48°53'E); CMNFI 1980-0916, 1, 53.3 mm standard length, Gilan, Nahang Roga (no other locality data); CMNFI 1991-0159, 1, 68.5 mm standard length, Mazandaran, Talar River (36°27'N, 52°48'E); CMNFI 1993-0140, 1, 44.3 mm standard length, Mazandaran, Shir River, Ramsar (36°51'48"N, 50°48'E); CMNFI 2008-0111, 1, 62.8 mm standard length, Gilan, Caspian Sea coast near Hendeh Khaleh (37°23'N, 49°28'E); CMNFI 2008-0112, 4, 26.6-51.4 mm standard length, Gilan, Caspian Sea coast near Hendeh Khaleh (37°23'N, 49°28'E); CMNFI 2008-0122, 2, 34.5-62.3 mm standard length, Gilan, Anzali Mordab (no other locality data).

Cobitis linea
(Heckel, 1847)

Common names

sagmahi-ye jonubi (= southern dogfish).

[southern spined loach]

Systematics

The type locality of Acanthopsis linea is "Bäche um Persepolis" according to Heckel (1847b). Persepolis is at 29°57'N, 52°52'E in Fars.

The holotype described by Bănărescu and Nalbant (1966) is one of 7 specimens assumed to be syntypes. The catalogue in Vienna lists 1 specimen in one column and 6 in the adjacent column. The type series is in poor condition being dried with scales lost and colour mostly faded except for one specimen. Bianco and Nalbant (1980) cite a specimen as the holotype, a female, 67.0 mm standard length, housed in the Naturhistorisches Museum Wien (NMW 48560 - this includes all 7 fish) and the type locality as Persepolis, presumably following Bănărescu and Nalbant (1966). The dried specimens measure 41.2-62.9 mm standard length and the undried one 61.3 mm standard length. My measurement is at variance with these authors.

Placed in the subgenus Bicanestrinia Băcescu, 1962 by Bianco and Nalbant (1980) and originally described in the genus Acanthopsis Agassiz, 1832 (see Eschmeyer (1990) for further details on this genus). Bohlen et al. (2006) place this species in their Bicanestrinia lineage I using the cytochrome b gene. Bicanestrinia has a derived character state in the duplication of the Canestrini scale or lamina circularis on the pectoral fin of males, the primitive condition being a single lamina circularis as in, e.g. Cobitis s.s. The separation between Bicanestrinia and Cobitis s.s. occurred 12-17 MYA when the land connection between Central Europe and Anatolia broke.

Material identified as Cobitis turcica (Hanko, 1924) from the "River Kor near Persepolis" in Nalbant and Bianco (1998) is C. linea based on my examination of IZA 7829-30. Fricke et al. (2007) have it in the Tor (sic, meaning Kor) River system of Iran, the same error. The species was described originally from Eregli in Anatolian Turkey and its presence in Iran needs confirmation. Erkakan et al. (1999) review the Turkish species of Cobitis but do not mention the occurrence of C. turcica in Iran.

Key characters

This species is distinguished by the dark brown lateral spots being reduced or absent, males have 2 Canestrini's scales at the upper bases of the unbranched and first branched pectoral fin rays, the laterocaudal branch of the suborbital spine is reduced or absent (although the spine itself is detectable by touch in even the smallest fish), and 14 branched caudal fin rays.

Morphology

Dorsal fin unbranched rays 2 and branched rays 6-7, anal fin unbranched rays 2-3 and branched rays 5, pectoral fin branched rays 7-9, and pelvic fin branched rays 5-6.

Meristic values for specimens examined by me are:- dorsal fin branched rays 6(16), anal fin branched rays 5(16), pectoral fin branched rays 8(6), pelvic fin branched rays 6(6), and total vertebrae 41(1).

Scales are embedded and have a reduced and eccentric focus. Johal et al. (2006) and Esmaeili and Niknejad (2006-2007) give scanning electron micrographs of the scales. The lateral line does not pass the end of the pectoral fin. The swimbladder capsule is globular but has ventral indentations anterolaterally on both sides. There are small keels on the upper and lower caudal peduncle. The anterior nostril is a short tube. The upper lip has fine furrows, the lower lip is thick and folded with a pair of medial lobes.

Sexual dimorphism

Males have Canestrini's scales as detailed above.

Colour

Overall colour is a yellowish-white with a golden iridescence especially on the operculum. Spots, blotches and dots are blue-grey or brown. The flanks are variably spotted and striped, the variability not related to sexual dimorphism. In some fish there is an upper flank, uniform stripe closely following the line of the back, below it a longitudinal series of irregular spots and blotches, a third stripe incompletely developed, and a fourth mid-lateral stripe of spots and blotches. In others the third and fourth stripes are an indistinguishable row of speckles. The mid-line of the back has a row of rounded spots numbering 7-11 (usually 8) predorsally, 0-3 (usually 2) at the dorsal fin base, and 5-11 (usually 8) postdorsally. The upper head is covered with many minute dots, extending onto the snout and upper head sides. The iris is golden-yellow. There is a blackish band from the eye to the mouth corner. There is a minute black spot on the caudal fin base. Fins are translucent. There are numerous dots scattered on the rays and membranes of the dorsal and caudal fins forming up to 9 irregular rows on the former and 8 more regular rows or bars on the latter, but still not very clearly defined. Spots on the dorsal edge of the caudal fin are clearly defined and regular; there are no spots on the ventral edge. The pelvic and anal fins are immaculate while the pectoral fin has some minor and irregular spots.

Size

Reaches 89.6 mm standard length.

Distribution

This species is found in the Kor River basin and the upper Kul River drainage of the Hormuz basin (Bănărescu and Nalbant, 1966; Bianco and Nalbant, 1980).

Zoogeography

This species may be related to Cobitis simplicispina Hankó, 1925 from Anatolia and other nominal species, all placed in the subgenus Bicanestrinia characterised by two Canestrini's scales. Members of this complex of species possibly reached western Asia from eastern Asia in the early Miocene. The disjunct distributions seen today were probably produced by Pleistocene climatic changes (Bianco and Nalbant, 1980).

Habitat

This species favours muddy bottoms. Specimens were caught in about 5-6 cm of mud, or at the foot of a muddy bank, with aquatic vegetation in the form of reeds.

Age and growth

Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 108 fish measuring 3.35-8.96 cm standard length. The a-value was 0.0089 and the b-value 3.060 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases).

Food

Unknown.

Reproduction

Adult females described by Bianco and Nalbant (1980) measuring 80.0-88.6 mm standard length have ripe eggs 0.8-1.1 mm in diameter. A specimen from the Lapui spring caught on 30 June carries eggs of this size.

Parasites and predators

Unknown.

Economic importance

This species is too small and rare to be of direct economic importance.

Conservation

Since this species is known only from a limited number of specimens, further studies should be undertaken to ascertain its abundance and distribution. It seems to favour muddy habitats and may have a restricted distribution in the rocky streams of Fars on this account.

Further work

See above.

Sources

Type material: See above (NMW 48560).

Iranian material:  CMNFI  1979-0292, 1, 69.8 mm standard length, Fars, Lapui Spring in the Kor River basin (29º48'N, 52º39'E); IZA 7829, 4, 41.9-85.5 mm standard length, Fars, Pulvar River 15 km north of Persepolis (29º59'N, 52º59'E); IZA 7830, 1, 27.1 mm standard length, Fars, springs of Kul River near Darab (no other locality data).

Cobitis sp.

Common names

None.

Systematics

 

Key characters

Morphology

Dorsal fin unbranched rays ? and branched rays ?, anal fin unbranched rays ? and branched rays ?, pectoral fin branched rays ?, and pelvic fin branched rays ?.

Sexual dimorphism

Males have Canestrini's scales as detailed above.

Colour

Size

Reaches ? standard length.

Distribution

This species is found in the Tigris River basin of Iran and Iraq. Iranian localities include ?

Zoogeography

Habitat

Unknown.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

This species is too small and rare to be of direct economic importance.

Conservation

Since this species is known only from a limited number of specimens, further studies should be undertaken to ascertain its abundance and distribution.

Further work

This taxon needs diagnosis and description, including DNA data, to distinguish it from other Cobitis in Southwest Asia.

Source

Iranian material: CMNFI 1979-0285, 1, 44.9 mm standard length, Kermanshahan, Qareh Su drainage (34º26'N, 46º37'E); CMNFI 1993-0127, 1, 50.5 mm standard length, Kermanshahn, Sarab-e Maran (34º44'N, 46º51'E); CMNFI 2008-0102, 1, 54.1 mm standard length, Kermanshahan, sarabs near Kermanshah (34º19'N, 47º04'E).

 

Misgurnus
Lacepède, 1803

Misgurnus anguillicaudatus
(Cantor, 1842)

Recorded from the Karakum Canal and Kopetdag Reservoir of Turkmenistan by Shakirova and Sukhanova (1994) and Sal'nikov (1995), this exotic species from China may eventually reach the Tedzhen River and Caspian Sea basins of Iran. No Iranian record.

Sabanejewia
Vladykov, 1929

Sabanejewia is distinguished from Cobitis Linnaeus, 1758 by having modally 12 rather than modally 14 branched caudal fin rays, a stronger suborbital spine, developed mental lobes which may be unfringed or well-fringed, large imbricated or unimbricated scales with a relatively large and central focus, and males have a protuberance on each side of the body in front of the dorsal and pelvic fins and lack the lamina circularis (a bony process at the base of the second pectoral fin ray)(see Vladykov (1929), Nalbant (1963, 1994), Sawada (1982), Vasil'yeva (1995b) and Perdices and Doadrio (1997, 2001) for further details). Krupp (1985c) does not consider Sabanejewia to be a distinct genus. Perdices et al. (2003) using mtDNA demonstrate that Caucasian-Caspian lineages are the sister group of a Danubian-Balkan lineage. Tang et al. (2008) using the cytochrome b gene found that ancestral Sabanejewia might have been the first cobitids to cross Siberia and invade the Euro-Mediterranean zoogeographic subregion.

Sabanejewia aurata
(De Filippi, 1863)

Qarasu, Aras River basin, August 2011, courtesy of K. Abbasi
Qarasu, Aras River basin, August 2011, courtesy of K. Abbasi

Common names

sagmahi-ye khardar (= spiny loach), sagmahi-e-jooibari.

[gizili iliskan in Azerbaijan; opi-dzug in Armenia; peredneaziatskaya shchipovka or Hither Asia spined loach, zolotistaya shchipovka or golden spined (spiny) loach, both in Russian].

Systematics

Kottelat (1997) tentatively considers that this species is restricted to Iran and possibly adjacent waters and does not occur in Europe.

The lectotype of Cobitis aurata (MZUT N.674), designated by Tortonese (1961), is stored in the Istituto e Museo di Zoologia della R. Università di Torino and 2 paralectotypes (MSNG N.365) from the Collezione di Universitá di Genova are in the Museo Civico di Storia Naturale di Genova (Tortonese, 1940; Tortonese, 1961). Eschmeyer et al. (1996) give the paralectotypes' catalogue number as ?MSNG 42727 (ex Univ. Genoa 365).

The type locality is "un fiumicello presso Sartschem" (De Filippi, 1863; not 1865 as in Berg (1948-1949), Banarescu and Nalbant (1964) and Reshetnikov et al. (1997) and not 1862 as in Tortonese (1940) and Eschmeyer et al. (1996)). The type locality is possibly Sarcham-e Sofla (37°07'N, 47°54'E) in the Qezel Owzan River drainage of the Caspian Sea basin in Iran.

Cobitis hohenackeri Brandt in Kessler, 1877 from the Kura River basin of Transcaucasia (Azerbaijan) is a synonym with 6 syntypes in the Zoological Institute, St. Petersburg (ZISP).

Key characters

Distinguished from other cobitids in northern Iran by having modally 12 branched caudal fin rays, large dark spots along the flank, and above them speckles which do not tend to form a stripe.

Morphology

Dorsal fin unbranched rays 2-3, branched rays 5-8, predominately 6 in the Caspian basin, anal fin unbranched rays 2-3, branched rays 4-8, usually 5, pectoral fin branched rays 5-9, predominately 7 in the Caspian basin (but see below), pelvic fin branched rays 4-8, usually 5 (but see below), and vertebrae 39-43.

Meristics for Iranian specimens:- ? check dorsal rays on x-rays, dorsal fin branched rays 6(11) or 7(4); anal fin branched rays 5(15); pectoral fin branched rays 7(1) or 8(14); pelvic fin branched rays 5(1) or 6(14); caudal fin branched rays 11(2) or 12(13); and total vertebrae ?.

Scales minute but visible to the naked eye, ca. 170-200. Dermal crest or adipose fins are variably developed behind the dorsal and anal fins. Barbels are longer than in C. keyvani, the mouth corner barbels reaching back to the posterior eye margin. Karyotype is 2n=50 (Klinkhardt et al., 1995).

Sexual dimorphism

The second pectoral fin ray in males is not enlarged as in Cobitis but there is a lateral distension of the body in front of the dorsal fin in mature males.

Colour

The back is a brownish olive-green with darker marbling. The flanks are golden-brown with a row of dark brown spots, less conspicuous than those in C. keyvani. These number 9-18, modally 11-13, in the Caspian basin. An upper row of spots is absent. The number of spots may be related to habitat, those fish from calm waters having few large spots. The back has several dark blotches along its mid-line. In the Caspian basin there are 7-16 back blotches, modally 9-10. There is a bar at the caudal base, sometimes with a small central gap. Fins are slightly pink.

Size

Attains 13.8 cm, but most fish are less than 10.0 cm.

Distribution

Found in the basins of the Baltic, Aegean, Black and Caspian seas and in the Tedzhen and Murgab rivers of Afghanistan and Turkmenistan according to most authors (Aliev et al., 1988; Nalbant and Bianco, 1998). A distinct subspecies is found in the Aral Sea basin (see below). Kottelat (1997) however considers that this species is restricted to Iran and possibly adjacent waters. It is found along the Caspian Sea coast of Iran including the Anzali Mordab and its tributaries, the Safid River at Kisom, lower Tonekabon, Chalus, Haraz and Babol rivers, Nakhurde (= ? Noqreh Deh), and in the Kashaf River of the Tedzhen River basin (Holčik and Oláh, 1992; Abbasi et al., 1999; Abdoli, 2000; Jolodar and Abdoli, 2004; Abdoli and Naderi, 2009).

The record of this species in the Tigris-Euphrates basin at Basrah, Iraq (BM(NH) 1920.3.5:9) in Nalbant (1963) and Bănărescu and Nalbant (1966), and repeated in Banister (1980), is probably an error of labeling (Bănărescu, 1973).

Zoogeography

The subspecies, Cobitis aurata aralensis Kessler, 1877, is reported from the Karakum Canal and the Uzboi lakes in Turkmenistan by Shakirova and Sukhanova (1994) and Sal'nikov (1995) and may well enter both the Tedzhen River and Caspian Sea basins of Iran eventually.

Habitat

This is a nocturnal species, hiding during the day under gravel and boulders of flowing rivers. If exposed, it will make jerking motions to the nearest cover. Some authors state that it also hides in sand. It prefers shallow and clear water. It occurs with C. keyvani but is commoner in faster water in the upper and middle reaches of rivers, from 5 to 150 cm water depth. However it may form populations in still water left behind after floods.

Age and growth

Males tend to be slightly smaller than females.

Food

Bottom-dwelling invertebrates are the main food items including larval insects such as mayflies, dragonflies and caddisflies as well as nematodes, copepods, chironomids, fish eggs, algae and detritus.

Reproduction

Fecundity reaches 14,700 eggs and egg diameter 0.85 mm. Eggs are shed over plants from April to August and this species may spawn in batches.

Parasites and predators

Unknown.

Economic importance

None.

Conservation

Lelek (1987) lists this species as rare to vulnerable in Europe. Kiabi et al. (1999) consider this species to be conservation dependent in the south Caspian Sea basin according to IUCN criteria. Criteria include few in numbers, habitat destruction, limited range (less than 25% of water bodies), absent in other water bodies in Iran, and present outside the Caspian Sea basin. The 2000 IUCN Red List lists this species as DD (Data Deficient).

Further work

The biology of this species in Iran has not been examined and its systematics needs clarification.

Sources

Vasil'eva and Vasil'ev (1988) give details on variation in meristics and colour of this species.

Iranian material: CMNFI 1970-0508, 4, 39.2-57.1 mm  standard length; Gilan, Safid River at Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1970-0545, 1, 33.9 mm standard length, Gilan, Safid River (ca. 37º01'N, ca. 49º38'E), CMNFI 1979-0448, 1, 70.1 mm standard length, Azarbayjan-e Khavari, Ahar Chay (38º18'30"N, 48º22'E); CMNFI 1980-0131, 1, 57.4 mm standard length, Iran, Caspian Sea basin (no other locality data); CMNFI 1980-0132, 1, 49.8 mm standard length, Gilan, Safid River at Kisom (37º12'N, 49º54'E); CMNFI 1980-0155, 7, 38.5-63.5 mm standard length, Azarbayjan-e Khavari, Qareh Su at Ardabil (ca. 38º15'N, ca. 48º18'E); CMNFI 2008-0221, 2, 56.0-61.9 mm standard length, Gilan, Safid River (no other locality data).

Sabanejewia caspia
(Eichwald, 1838)

Common names

mahi roshtegar talaee (= golden dustman fish), rofteghar mahi, sagmahi-ye Khazari (= Caspian loach), sag mahi khardar (= spiny loach), mar mahi (= snake fish), sagmahi-e-jooibari.

[xazar iliskani in Azerbaijan; Kaspiiskaya shchipovka or Caspian spined (spiny) loach in Russian].

Systematics

This species was described from "in sinu mardofiensi prope castellum Lencoranicum" (i.e. the Caspian Sea in the ? Mardofiensi Gulf near the Lenkoran fortress). The type specimen is apparently lost (Vasil'yeva, 1995b).

Key characters

Distinguished from other cobitids in northern Iran by having modally 12 branched caudal fin rays, no large dark spots along the flank but an irregular stripe, and above this stripe speckles which do not tend to form a stripe.

Morphology

Dorsal fin unbranched rays 2-3, branched rays 6-7, anal fin unbranched rays 1-2, branched rays 5-6, pectoral fin branched rays 6-7 (but see below) and pelvic fin branched rays 4-6. Vertebrae 41-42. A crest is well developed on the lower caudal peduncle but only posteriorly on the upper edge. Barbels are shorter than in C. keyvani, the posterior ones reaching beyond the posterior eye margin. Karyotype is 2n=50 (Klinkhardt et al., 1995).

Meristics for Iranian specimens:- dorsal fin branched rays 6(83) or 7(1); anal fin branched rays 5(80) or 6(4); pectoral fin branched rays 7(6), 8(72) or 9(6); pelvic fin branched rays 4(3), 5(67) or 6(14); caudal fin branched rays 7(1, but deformed), 9(1), 10(3), 11(4), 12(74) or 13(1); total vertebrae ?.

Sexual dimorphism

The second pectoral fin ray in males is not enlarged as in Cobitis but there is a lateral distension of the body in front of the dorsal fin an mature males.

Colour

Live specimens, especially young, are almost transparent. The adult has a pronounced dark line along mid-flank indicating the separation of the upper and lower muscle masses. Above this line the upper flank is yellowish with irregular dark grey pigment or brown speckles near the back. The lower flank has irregular grey pigment or brown speckles. The mid-line of the back has a more or less pronounced dark line. The belly and lower head surface are pale yellowish without grey pigment. The dorsal and caudal fins have yellow-orange rays bearing 3-4 series of dark grey spots. The base of the caudal fin has 2 dark dots although these are not as marked as in some Cobitis keyvani. The pectoral, pelvic and anal fins are transparent though larger fish may have elongate grey spots along the rays. The iris is golden.

Note that Vasil'yeva (1995b), based on original data and that of Derzhavin (1934), found that the dark longitudinal band and the dark caudal fin base may not be pronounced.

Size

Reaches 9.2 cm standard length.

Distribution

Principally found in the southern Caspian Sea basin. Records from the northern Caspian Sea are apparently in error and the range is from the Kura to the Babol rivers including the Anzali Mordab at "Khalkai" for example, the Safid, Chowbar, Tonekabon, Chalus, Shahzadeh, "Laidschana", Meshedessera (= ? Babol Sar), Haraz, (Holčik and Oláh, 1992; Vasil'yeva, 1995b; Abbasi et al., 1999; Abdoli, 2000; Jolodar and Abdoli, 2004; Abdoli and Naderi, 2009).

Zoogeography

This species is endemic to the Caspian Sea basin (Nalbant and Bianco, 1998).

Habitat

This species is found in both fresh and brackish waters, in slow or still water with aquatic vegetation, in the lower reaches of rivers and near river mouths, and in brackish bays.

Age and growth

Unnown.

Food

Unknown.

Reproduction

Spawning takes place in April in the brackish lagoons of the Lenkoran, Azerbaijan. Up to 955 eggs of up to 0.85 mm diameter are carried by each female.

Parasites and predators

None reported for Iran.

Economic importance

None.

Conservation

Holčík and Oláh (1992) report the loss of this species from the Anzali Mordab where it was once found. Reasons for this loss are unknown. Kiabi et al. (1999) consider this species to be conservation dependent in the south Caspian Sea basin according to IUCN criteria. Criteria include possibly few in numbers, habitat destruction, limited range (less than 25% of water bodies), absent in other water bodies in Iran, and absent outside the Caspian Sea basin. Vulnerable in Turkey (Fricke et al., 2007).

Further work

The biology of this species in Iran requires study.

Sources

Iranian material: CMNFI 1970-0517, 2, 24.4-31.7 mm standard length, Caspian Sea basin (no other locality data); CMNFI 1970-0537, 6, 25.4-53.6 mm standard length, Markazi, Shah River near Manjil (36°44'N, 49°24'E); CMNFI 1970-0542, 16, 25.0-59.1 mm standard length, Gilan, Old Safid River estuary (37°23'N, 50°11'E); CMNFI 1970-0550, 2, 18.7-27.2 mm standard length, Caspian Sea basin (no other locality data); CMNFI 1970-0555, 4, 20.9-38.0 mm standard length, Caspian Sea basin (no other locality data); CMNFI 1970-0566, 40, 26.6-58.7 mm standard length, Gilan, Old Safid River estuary (37°23'N, 50°11'E); CMNFI 1970-0567, 27, 26.3-62.0 mm standard length, Gilan, Pir Bazar Roga (37°21'N, 49°33'E); CMNFI 1970-0570, 2, 19.2-27.1 mm standard length, Gilan, Hasan Kiadeh (37°24'N, 49°58'E); CMNFI 1970-0579, 57, 24.7-60.4 mm standard length, Gilan, Old Safid River estuary (37°23'N, 50°11'E); CMNFI 1970-0580, 1, 63.4 mm standard length, Mazandaran, Iz Deh (36°36'N, 52°07'E); CMNFI 2008-0122, 49.7 mm standard length, Gilan, Anzali Mordab (no other locality data)..

Sabanejewia caucasica
(Berg, 1906) CMNFI

Reported from the Anzali Mordab and lower reaches of the Safid, Toenkabon, Chalus, Heraz and Babol rivers in Iran (Abdoli, 2000) and mapped from the Caspian coast of Iran at Babol by Kottelat and Freyhof (2007). Berg (1948-1949) notes that this species is found in the northern Transcaucasus and is not reported from the Kura River in the southwestern corner of the Caspian Sea. Its presence in Iran needs confirmation by specimens. Formerly in the genus Cobitis.

Nemacheilidae

CMNFI 1970-0558, 4, ?mm standard length, Ghasemlou Chay ()?brandtii;  CMNFI 1970-0560, 3, ?, mm standard length, Azarbaijan-e Bakhtari, Mamiyand Chay near Mamiyand (ca. 36º59'N, ca. 45º39'E)?brandtii

 

 

 

CMNFI 1979-0026, 8, 22.8-46.6 mm standard length, Fars, Shapur River at Shapur (29º47'N, 51º35'E);  CMNFI 1979-0155, 2, 30.7-30.9 mm standard length, Fars, spring at Gavanoo (28º47'N, 54º22'E); CMNFI 1979-0167, 41, 20.6-48.3 mm standard length, Kerman, qanat at Bam (29º06'N, 58º20'E); CMNFI 1979-0168, 1, 47.2 mm standard length, Kerman, qanat at Shahabad (29º07'N, 58º16'E); CMNFI 1979-0169, 11, ? mm standard length, Kerman, qanat 10 km from Mahan (30º08'30"N, 57º17'E); CMNFI 1979-0170, 1, 45.0 mm standard length, Kerman, qanat at Baghin (30º12'N, 56º48'E); CMNFI 1979-0172, 18, 33.8-45.8 mm standard length, Kerman, qanat on Kerman to Bandar Abbas road (29º51'N, 56º14'E); CMNFI 1979-0184, 2, 27.7-28.4 mm satndard length, ?; CMNFI 1979-0186, 2, ? mm standard length, Hormozgan, stream at Sar Khun (ca. 27º24'30"N, ca. 56º25'E); CMNFI 1979-0192, 7, ? mm standard length, Fars, qanat 2 km east of Rostaq (28º26'30"N, 55º04'E); CMNFI 1979-0193, 1, ? mm standard length, Fars, river 8 km from Darab (28º45'N, 54º27'30"E); CMNFI 1979-0194, 2, 37.6-45.6 mm standard length, Fars, jube 15 km from Darab (28º45'30"N, 54º24'E); ; CMNFI 1979-0208, 15, 31.1-58.8 mm standard length, Fars, qanat on road to Qatru (ca. 29º11'N, ca. 54º40'E); CMNFI 1979-0213, 7, ? mm standard length, Kerman, stream in Kharan River drainage (29º15'N, 56º25'E); CMNFI 1979-0219, 13, ? mm standard length, Kerman, jube 14 km west of Jiroft (28º37'N, 57º41'E); CMNFI 1979-0276, 14, 36.3-45.8 mm standard length, Lorestan, Chamesk River (ca. 33º19'N, ca. 47º53'30"E); CMNFI 1979-0284, 3, 38.7-41.9 mm standard length, Kermanshahan, Marek River at Mahidasht (34º16'N, 46º48'30"E); CMNFI 1979-0288, 1, 32.4 mm standard length, Ilam and Poshtkuh, Gangir River at Juy-e Zar Eivan (33º50'N, 46º18'E); CMNFI 1979-0306, 11, 20.0-49.3 mm standard length, Kerman, qanat 33 km from Sirjan (29º13'N, 54º33'E); CMNFI 1979-0307, 4, 31.0-42.3 mm standard length, Kerman, river at Sartal (ca. 29º17'N, ca. 56º38'E)? distinct species; CMNFI 1979-0316, 1, ? mm standard length, Baluchestan, stream on road to Chah Bahar (26º48'N, 61º02'E); CMNFI 1979-0341, 10, 25.5-45.6 mm standard length, Kerman, Tahrud west of Bam (29º23'N, 57º52'E); CMNFI 1979-0365, 7, 32.0-40.5 mm standard length, Khuzestan, stream in Doveyrich drainage (32º25"N, 47º36'30"E); CMNFI 1979-0366, 1, 36.0 mm standard length, Khuzestan, stream 17 km west of Dehloran (32º45'30"N, 47º05'30"E); CMNFI 1979-0367, 1, 41.3 mm standard length, Khuzestan, Meymeh River 11 km north of Dehloran (32º44'30"N, 47º09'30"E); CMNFI 1979-0371, 2, 44.5-55.6 mm standard length, Khuzestan, stream in Karkheh River drainage (32º05'N, 48º19'E); CMNFI 1979-0374, 2, 41.3-41.7 mm standard length, Khuzestan, stream tributary to Bala River (32º40'N, 48º15'E); CMNFI 1979-0389, 1, 32.4 mm standard length, Khuzestan, Zard River 1 km south of Bagh-e Malek (31º31'N, 49º53'30"E); CMNFI 1979-0390B, 3, 29.1-39.4 mm standard length, Khuzestan, stream 3km south of Bagh-e Malek (31º29'N, 49º54'30"E); CMNFI 1979-0395, 1, 30.7 mm standard length, Khuzestan, stream in Marun River drainage (ca. 30º57'N, ca. 49º51'E); CMNFI 1979-0411, 3, 21.9-25.9 mm standard length, Hormozgan, Minab River near Rudan (27º24'N, 57º12'E)?cf bampurensis; CMNFI 1979-0419, 19, 32.4-58.7 mm standard length, Fars, stream 7 km from Rostaq (28º29'N, 55º01'E); CMNFI 1979-0423, 4, 43.2-45.7 mm standard length, Boyer Ahmadi-ye Sardsir va Kohkiluyeh-Fars border, river in Beshar River drainage (30º31'N, 51º31'E); CMNFI 1979-0459, 4, ? mm standard length, Hamadan, stream 2 km south of Kazan (35º22'N, 49º02'E); CMNFI 1991-0156, 1, ? mm standard length, Hamadan, Gav Masiab River (34º16'N, 48º10'E); CMNFI 2007-0037, 5, 45.2-54.9 mm standard length, Kerman, Hosseinabad and Gamatabad qanats at Bam (29º06'N, 58º21'E); CMNFI 2007-0038, 1, 39.0 mm standard length, Kerman, Mehtiabad qanat at Bam (29º06'N, 58º21'E); CMNFI 2007-0039, 6, 32.4-41.7 mm standard length, Kerman, Tahrud River (ca. 29º23'N, ca. 57º63'E); CMNFI 2007-0043, 9, 33.0-56.7 mm standard length, Kerman, qanat at Emamzadeh Sultan (ca. 29º40'N, ca. 56º45'E); CMNFI 2007-0047, 2, 36.8-58.2 mm standard length, Kerman, qanat at Hoshum (29º14'N, 56º19'E); CMNFI 2007-0051, 5, 28.6-36.3 mm standard length, Hormozgan, upper Kol River basin (28º19'N, 55º55'E); CMNFI 2007-0055, 3, 36.1-40.3 mm standard length,-0103 Hormozgan, stream in Minab River basin (27º47'N, 57º12'E); CMNFI 2007-0075, 6, 41.3-47.9 mm standard length, Hamadan, Malayer River 5 km from Malayer (ca. 34º17'N, ca. 48º47'E); 

 

 CMNFI 2007-0089, 7, ? mm standard length, Azarbayjan-e Khavari, Ahar Chay at Ahar (38º28'N, 47º03'E); CMNFI 2007-0093, 8, ? mm standard length, Azarbayjan-e Bakhtari, Qotur River south of Khvoy (38º30'N, 44º58'E); CMNFI 2007-0100, 4, 44.2-58.4 mm standard length, Azarbayjan-e Bakhtari, Kalwi Chay near Piranshahr (ca. 36º44'N, ca. 45º10'E); CMNFI 2007-0105, 13, ? mm standard length, Kordestan, Zarineh River basin south of Saqqez (ca. 36º06'N, ca. 46º20'E); 

 

CMNFI 2007-0116, 2, 40.6-44.8 mm standard length, Kermanshahan, Gav Masiab River basin west of Sahneh (ca. 34º28;'N, ca. 47º36'E); CMNFI 2007-0118, 15, 24.7-58.8 mm standard length, Kermanshahan, Bid Sorkh River between Sahneh and Kangavar (ca. 34º23'N, ca. 47 52'E); CMNFI 2007-0119, 8, 26.9-37.9 mm standard length, Kermanshahan, Gav Masiab River basin near Kangavar (ca. 34º31'N, ca. 48º03'E); CMNFI 2008-0102, 1, 41.6 mm standard length, Kermanshahan, sarabs near Kermanshah (34º19'N, 47º04'E); CMNFI 2008-0117, 3, 48.4-51.2 mm standard length, Kermanshahan, Sarab-e Yavari (34º28'N, 46º56'E); CMNFI 2008-0120, 1, ? mm standard length, Khuzestan, Rud Zard at Rud Zard (31º22'N, 49º43'E); CMNFI 2008-0130,1, 22.8 mm standard length, Khuzestan, Kupal (31º15'N, 49º10'E);  BWC 1997-0005, BWC 2000-0001, CMNFI 2008-0161, 10, ? mm standard length, Khuzestan, A'la River at Pol-e Tighen (31º23'20"N, 49º52'44"E), CMNFI 2008-0163, 1, ? mm standard length, Khuzestan, Marun River at Chahar Asiab (30º40'28"N, 50º09'34"E); ?nielseni CMNFI 2008-0167, 54, ? mm standard length, Khuzestan, stream above Diuni Darreh (32º37'42"N, 48º41'40"E); CMNFI 2008-0170, 7, ? mm standard length, Khuzestan, Zard Rud (31º22'28"N, 49º43'15"E); CMNFI 2008-0175, 8, ? mm standard length, Lorestan, Kahman River at Dow Ab-e Aleshtar (33º47'N, 48º12'E);

New species: CMNFI 1979-0217, 6, 30.1-41.8 mm standard length, Kerman, Kharan River drainage (ca. 28º59'30"N, ca. 56º51'30"E);

CMNFI 2007-0045, 6, 28.8-39.2 mm standard length, Kerman, Kharan River drainage at Baft (29º14'N, 56º38'E)checkID

ID?persaCMNFI 1970-0540, 1, 49.5 mm standard length, ?; CMNFI 1979-0019, 2, 28.9-33.7 mm standard length, Fars, Barm-e Baba Hajji (29º23'N, 52º40'E); CMNFI 1979-0027, 9, 31.0-41.3 mm standard length, Fars, Chehel Chashmeh (ca. 29º43'N, ca. 52º04'E); CMNFI 1979-0073, 10, 31.1-56.1 mm standard length, Fars, Mand River (ca. 29º42'30"N, ca. 52º01'30"E); CMNFI 1979-0086, ?; CMNFI 1979-0111, 5, 43.2-52.7 mm standard length, Fars, stream on Shiraz to Bushehr road (29º37'30"N, 52º21'E); CMNFI 1979-0157, 3, 34.8-37.7 mm standard length, Fars, qanat at Hadiabad (28º52'N, 54º13'E); CMNFI 1979-0206, 3, 24.9-36.1 mm standard length, Fars, qanat 1 km from Runiz-e Pa'in (29º12'N, 53º40'E); CMNFI 1979-0399, 1, 26.2 mm standard length, Fars, stream near Basht (30º19'30"N, 51º15'E);

 

 

The hillstream, mountain or river loach family has had various family names - see Menon (1987), Mirza (1989b), Hieronimus (1990; 1991), Ng and Lim (1991), Kottelat (1988, 1991), Nelson (1991), Thorne (1992), Anonymous (1993a) and Bănărescu and Nalbant (1995) for discussions. The Iranian species were classified in Cobitidae in older works and more recently in Homalopteridae and Balitoridae. Mirza (1989b), Nalbant (1998), Tang et al. (2006) and Šlechtová et al. (2007) consider that the correct name for this family is Noemacheilidae (in Mirza) or Nemacheilidae. Nemacheilids are aligned with the main stem of cobitoid fishes rather than homalopterids as proposed by Sawada (1982) on osteological grounds. Nalbant (1998) considers the similarities observed by Sawada between nemacheilids and homalopterids to be homoplasies. Homalopterids are more closely related to cyprinids and psilorhynchids.

The family is found throughout Eurasia with a single species in northeast Africa. There are about 72 genera and about 793 species (Berra, 2001; Nelson, 2006; Eschmeyer and Fong, 2011), with more being described regularly. Iranian species belong to the subfamily Nemacheilinae.

Conway et al. (2010) point out that although there are no morphological synapomorphies for the family, molecular data does support monophyly. Prokofiev (2010) gives osteological characters for the subfamily Nemacheilinae.

Most species in Iran were placed in the genus Nemacheilus Bleeker, 1863 but this name contains only species from Southeast Asia (Kottelat, 1997). Noemacheilus Kuhl and van Hasselt in van Hasselt, 1823 is a nomen nudum since there are no taxonomic characters accompanying the original description. The next available name is Nemacheilus Bleeker, 1863 (see Kottelat, 1987). Nemachilus Günther, 1868 is an incorrect spelling. Much of the earlier literature on Iranian members can be found under the name Nemacheilus or its variant spellings.

These loaches have been placed in several genera or subgenera including the following relevant to Iran: Orthrias Jordan and Fowler, 1903 (see Banarescu, Nalbant and Balik (1978); Orthrias = Barbatula Linck, 1790 of authors, see Bănărescu and Nalbant (1995) for reasons advocating the later name over the earlier one; Kottelat (1997) gives Linck as 1789), Adiposia Annandale and Hora, 1920 (see Annandale and Hora (1920); see also Bănărescu and Nalbant (1995) where it is synonymised with Paracobitis), Triplophysa Rendahl, 1933 and Hedinichthys Rendahl, 1933 (see Bănărescu and Nalbant (1966); see also Bănărescu and Nalbant (1995) where the latter is synonymised with the former), Oxynoemacheilus Bănărescu and Nalbant, 1966 (see Bănărescu and Nalbant, 1966; synonymised with Orthrias in Bănărescu and Nalbant (1995) and with Barbatula in Bogutskaya and Naseka (2004)), Paracobitis Bleeker, 1863 (see Bănărescu and Nalbant, 1966), Schistura McClelland, 1839 (see Mirza, Nalbant and Banarescu, 1981; regarded as polyphyletic by Bănărescu and Nalbant (1995)), and Seminemacheilus Bănărescu and Nalbant, 1995 (q.v.). Views on the generic validity of these names conflict between authors and between the same author at different dates (see Krupp, 1985c; Eschmeyer, 1990; Eschmeyer et al., 1996; Eschmeyer's "Catalog of Fishes"). Kottelat (1984) retained Nemacheilus until a revision of all species was complete but there has been a tendency to use the above genera and this is followed here. Prokofiev (2004, 2007, 2009, 2010) has revised some of the Nemacheilidae, in particular the group of nemacheiline loaches that lack the preethmoid I in the skull, using osteology and morphology. This includes most of the species found in Iran and is now being followed by such authors as Esmaeili et al. (2010).

The table below summarises some of the various allocations and includes names used in Iran but not necessarily occurring there (see text below for further details:-

Species Original genus^ Other generic allocations^ Current genus Current species
angorae Nemacheilus Barbatula, Orthrias Oxynoemacheilus angorae
araxensis Orthrias Barbatula Oxynoemacheilus araxensis
bampurensis Nemacheilus Schistura Paraschistura bampurensis
bergianus Nemachilus Barbatula, Orthrias Oxynoemacheilus bergianus
brandtii Nemachilus Barbatula, Orthrias Oxynoemacheilus brandtii
cristatus* Nemacheilus Paracobitis, Schistura Metaschistura cristata
frenata* Cobitis Barbatula, Nemacheilus, Orthrias Oxynoemacheilus frenatus
iranica Paracobitis - Paracobitis iranica
kermanshahensis Noemacheilus Orthrias, Barbatula Oxynoemacheilus kermanshahensis
kessleri Nemachilus Schistura Paraschistura kessleri
kiabii Oxynoemacheilus - Oxynoemacheilus kiabii
kosswigi Turcinoemacheilus - Turcinoemacheilus kosswigi
longicauda Cobitis Adiposia, Nemacheilus Paracobitis longicauda
longipinnis Ilamnemacheilus - Ilamnemacheilus longipinnis
malapterura Cobitis Nemacheilus Paracobitis malapterura
nielseni Schistura - Paraschistura nielseni
persa Cobitis Barbatula, Nemacheilus, Orthrias Oxynoemacheilus persa
rhadinaeus* Nemachilus Adiposia Paracobitis rhadinaea
sargadensis Nemacheilus Schistura Paraschistura sargadensis
smithi Noemacheilus - Paracobitis smithi
stolickai Cobitis Nemacheilus Triplophysa stolickai
tongiorgii Seminemacheilus - Oxynoemacheilus tongiorgii
vignai Paracobitis - Paracobitis vignai

* original trivial name, the suffix of which may change if genus changes to masculine or to feminine; ^ Nemacheilus sometimes spelled Noemacheilus or Nemachilus by various authors

A number of nemacheilid species have been described from waters confluent with Iran, particularly from the Helmand River basin in Afghanistan. They have no Iranian records but are listed here as they may be relevant to revisionary studies.

The dating of the paper by Bănărescu and Nalbant as 1967 in various works may follow Nalbant and Bianco (1998). The Bănărescu and Nalbant paper states in Danish on page 186 "Reprints released the 31 December 1966" and this is presumably the correct date, as Art. 21.2 of the Code of Zoological Nomenclature states "Date specified. The date of publication specified in a work is to be adopted as correct in the absence of evidence to the contrary", and no evidence to the contrary has been presented (N. G. Bogutskaya, pers comm., 28 April 2011). This changing of dates here has no nomenclatural significance.

Members of this family in Iran are characterised by an elongate and weakly compressed and almost cylindrical body, head not compressed but rounded, scaleless or body covered in minute scales (too small for scale counts to be commonly or easily made), lateral line complete or incomplete, a small and inferior mouth, lips thick, fleshy and papillose, lower lip interrupted in the middle, 2 pairs of barbels on the snout and 1 pair at the mouth corners (8 pairs in some non-Iranian species), no collapsible spine under the eye (sometimes present in non-Iranian species but distinguishes members of the related Cobitidae in Iran), eyes small to minute, usually not visible from the underside of the head, reduced gill opening, short to moderate dorsal fin without spines, short anal fins, vent a short distance in front of anal fin origin, swimbladder enclosed partially or entirely in a bony capsule, certain osteological characters such as the shape of bones in the Weberian apparatus used in sound transmission from the swimbladder to the ear, gut short or long, a dermal crest or adipose fin may be present, caudal fin truncate, rounded or slightly forked, and often distinctive colour patterns of bars, stripes and blotches. Iranian species may lack scales, may have an adipose fin, and have a single unbranched ray leading the pectoral and pelvic fins.

Krupp (1985c) reviews prior works by P. M. Bănărescu and co-authors on Levantine nemacheilids and regards them as unsatisfactory. This calls into question works on Iranian species by this author. Krupp (1985c) lists characters important in studying Nemacheilus sensu lato and those which are individually variable or develop independently, much in contrast to characters favoured by Bănărescu. Morphometric characters can vary with nutritional status and ecological factors. Stable characters were head length, interorbital width, caudal peduncle length and depth, and predorsal length. Fin lengths are dependent on sex in some species, less so in others. Allometry is a problem in fin positions and measurements involving such characters can only be used when comparing fish of equal size. Mouth width and digestive tract shape are good characters but lip shape and development of the processus dentiformis are not. The swimbladder capsule form, including the presence or absence of a continuous collar between the two hemispheres and the shape of laminae, is an important character. Reduced laminae and wide recesses on the hemispheres are derived characters. The dorsal adipose fin development is stable in some species, variable in others, and is independently derived in different phyletic lines, thus being of limited value. The shortening and deepening of the caudal peduncle is derived in one Levantine species. Scale characters such as size and position of the focus and general scale structure are very variable and not characters easily quantified. Only specimens of the same size are comparable and numerous scales must be examined because aberrant ones are common. The lateral line length is a good character, although juvenile fish may have a shortened one. A reduced lateral line is a derived character. Colour patterns are subject to variation and both spotted and striped forms can be found within one species. Nevertheless, patterns can be important in distinguishing species. Thickening of pectoral fin rays is a derived feature but absence of this character is a symplesiomorph condition and cannot be used to relate species.

The species in this family are often difficult to identify and many literature reports are undoubtedly mis-identifications. While some of these may be corrected based on material deposited in museums, others have no voucher material and cannot be re-identified. Identification is problematical because scale counts are not available (too minute), fin ray counts are often very similar, and unique structures uncommon. Colour patterns can be used but are notoriously variable and many types in museums are decoloured making comparisons difficult. Morphometric characters require good series of adult fish of both sexes, from various localities in the species range, preferably even from the same locality taken over several years to allow for local variations in habitat which may conceivably affect shape. Menon (1987) considers that many species in this genus are from very similar habitats, the stressful one of running water, and have been constantly selected to fit this niche. Valid species resemble one another closely. Such characters as position of the anal opening, the dorsal fin origin and barbel length have been used in species definitions but Menon (1987) found these to vary with growth. Swimbladder structure depends on the habitat where the fish live and scale coverage on the physico-chemical nature of the water. Menon (1987) found lateral line character, number of branched dorsal fin rays, caudal fin shape, secondary sexual characters in males and, despite the above, body colour and anal opening position to be useful. Prokofiev (2010) reviewed the morphological classification of loaches and points out that changes in morphology occur with growth and species with wide ranges show large variations in morphology associated with the various biotopes. He gives extensive reviews of characters and their importance in defining genera and species, e.g. scale cover and scale characters are not considered important in phylogeny and are of only accessory import for genus and species diagnosis; colour, sexual dimorphism, adipose keel presence and extent, fin ray numbers, head and body shapes, abdominal axillary lobe presence and size, variations in the seismosensory system, nostril size and position, anal opening position, intestine shape, swimbladder and its capsule structure, and general osteology are all important characters.

Examination of Iranian species, where good series of fresh material was available, tend to confirm the observations of Krupp and Menon on characters. Position of the dorsal fin origin is variable within a species among morphometric characters used as distinctive, extent of the lateral line is also variable, there is marked sexual dimorphism, and colour patterns can be useful but also vary with the individual, the habitat and the temperature (fish kept in ice water have strong colour patterns while those preserved immediately from murky waters have faint patterns).

Two main problems exist in identifying certain Iranian loaches. These are determining appropriate characters which are not individually variable and which are apomorphic, and applying existing names to fresh material in comparison with poorly-preserved types.

A number of species remain to be described and are currently under study. Some named species are probably distinct taxa, e.g. "Nemacheilus" tigris (Heckel, 1843) (sagmahi-ye Dajleh) is recorded from the Karun River basin in Khuzestan (ZISP 24098) by Berg (1949) but specimens from this part of Iran differ from Heckel's types. The colour pattern on the fish figured by Berg (1949) from the Karun River in Iran is atypical according to Bănărescu and Nalbant (1966) - it has only 4-5 bars on the posterior part of the body. The type locality of Cobitis Tigris is "Flüsschen Kueik bei Aleppo" (Haleb, Syria) according to Heckel (1843b). Prokofiev 92009) places Cobitis tigris in the genus Paracobitis.

Sawada (1982) thought that this family dispersed by two routes from Southeast Asia, one through Siberia and one through South Asia to reach what is now Iran. Menon (1987) considers that the land mass between East Africa and the west coast of India has submerged only recently, probably simultaneous with the birth of the Ganges and Indus. Connections of the Pleistocene fore-deep of the Himalayas with the Tigris-Euphrates basin in what is now the Persian Gulf could have existed, allowing movement of "Nemacheilus" species along a continuous route from Yunnan to Anatolia. Menon (1987) further suggests a series of waves, spreading "Nemacheilus" westwards into Southwest Asia from a South China origin. The Triplophysa wave is the first wave of evolution, in which the earliest stock from Yunnan spread through Tibet in the late Miocene and early Pliocene before the major rise of the Himalayas. By the end of the Tertiary, particularly in the Pleistocene, the Tibetan Plateau had risen causing a dry and cold climate with increased solar radiation and torrential rivers. This change in the environment caused rapid evolution, leading to such taxa as Triplophysa and Hedinichthys. The rupecola wave took place in the late Pliocene along the southern face of the Himalayas through Iran to Anatolia and even northeast Africa. Some of the criticisms listed under the cyprinid genus Garra, whose distribution Menon (1964) also attributes to waves, may be apposite here too.

These are small fishes, up to about 200 mm in size although one species (not in Iran) reaches 482 mm (Prokofiev, 2010). They are quite secretive, hiding under stones or in mud. This common and stressful habitat may have led to a general similarity in body form among the various species. Some are known only from caves, including one Iranian species. Despite their small size, they are regarded as a delicacy in India (Hora, 1956). Barbatula angorae (and presumably other species) is a potential fishing bait for predatory fishes such as Sander lucioperca and has been examined experimentally for this purpose in Turkey (Kuşat et al., 1995). They are generally known as سگ ماهي (sag mahi meaning dog fish, but this is presumably the equivalent of loach in English), لوچ (= louch meaning loach) or mar mahi (= snake fish, presumably in reference to the elongate shape) in Farsi. These general names are not repeated below.

Genus Ilamnemacheilus
Coad and Nalbant, 2005

This genus is characterised by a high, laterally compressed body; large head with small eyes and mouth; anterior lip lacking an interruption in the middle; posterior lip with widened mental lobes, small round papillae covering only the mental lobes, the rest of the lips being unfurrowed; the processus dentiformes absent; lateral line complete and terminating slightly before the posterior margin of the caudal peduncle; scales small with a quite large and eccentric focus, sparsely present on the rear half of the body; stomach syphonal and intestine straight without loops; gas bladder with two encapsulated chambers united by a short encapsulated duct; paired fins very long; dorsal fin long; and caudal fin well forked.

The type species is Ilamnemacheilus longipinnis by original designation and monotypy.

Ilamnemacheilus longipinnis
Coad and Nalbant, 2005

Common names

None.

Systematics

The holotype and only known specimen is CMNFI 1979-0366 (79-966 is a lapsus), 36.0 mm standard length, Iran, Meymeh River, formerly a tributary of the Tigris River, 17 km west of Dehloran and about 21 km east of the Iraqi border, 32º45'30"N, 47º05'30"E, 28 January 1978, B. W. Coad and S. Coad. This species was described by the late T. T. Nalbant on material collected by me. I first saw the species description when I received reprints and had no input into this paper. The extensive vertebral fusions account in part for the unusual body form.

Key characters

Characters are those of the genus.

Morphology

Dorsal fin with 3 unbranched and 10 branched rays, anal fin with 2 unbranched and 5 branched rays, pectoral fin with 9 branched rays and pelvic fin with 5 branched rays. Total vertebrae 28 or 29 including the ural centrum (vertebral fusions present), and some centra have two neural and haemal arches. ?check vertebral counts against other loaches - very low ? fusions Other characters are listed above under the genus and Coad and Nalbant (2005) give some measurements.

Sexual dimorphism

Unknown.

Colour

The sole preserved specimen is a overall a pale brown with 3-4 indistinct greyish blotches in the middle of the second half of the body. All fins are pale but the caudal fin has faded greyish lines along the marginal rays. In life it was an olive-green overall with orange fins.

Size

Reaches 36.0 mm standard length.

Distribution

Endemic to Iran and found in Tigris River basin at a single locality (see above).

Zoogeography

An endemic genus in the Tigris-Euphrates basin (along with Turcinoemacheilus, not in Iran). This species may be related to an undescribed species from the Orontes River basin in Syria.

Habitat

The sample site was a small stream, 20 m wide with a maximum depth of 1 m. Altitude was 210 m. Capture depth was 30 cm in a medium current. The bottom was a mix of pebbles and mud with some encrusting algae. Water temperature was 14ºC, pH was 6.0 and conductivity was 1.65 mS. The cyprinids Cyprinion macrostomum and Garra rufa were caught with the loach.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

None.

Conservation

Known only from a single specimen, abundance and wider distribution unknown.

Further work

More collections are needed to record information on biology and distribution and provide a more detailed description.

Sources

The holotype and sole known specimen (see above).

Genus Metaschistura
Prokofiev 2009

This genus has a single, small species and the characters of the species are those of the genus. Prokofiev (2009) gives further osteological details.

Metaschistura cristata
(
Berg, 1898)

Common names

sagmahi-ye Torkomani or Turkmeny (= Turkmenian crested loach), sagmahi-ye kakoldar-e Torkomani.

[Turkmenskii grebenchatyi golets or Turkmenian crested loach in Russian].

Systematics

This species was described under Nemacheilus in Latin from "Habitat in flum. Tedschent, prope Aschabad, in provincia Transcaspica". The type locality is presumably the Tedzhen River in Turkmenistan although Ashkhabad is not on the Tedzhen River. Berg (1948-1949) notes "not in the Tedzhen!". Syntypes are reported to be in the Zoological Museum of Moscow State University (MMSU) by Eschmeyer et al. (1996).

Placed in the taxon Paracobitis by Bănărescu and Nalbant (1966) and later in Schistura by Bănărescu and Nalbant (1995) and by Nalbant and Bianco (1998).

Key characters

The dorsal fin branched ray count of 8 and unique colour pattern distinguishes this crested loach from other crested loaches in northeast Iran and adjacent regions. In addition the crest, or adipose fin, is shorter and thicker than in P. malapterura.

Morphology

Dorsal fin unbranched rays 2-6 (several minute, embedded in flesh and only visible on x-rays) and branched rays 7-9, modally 8, anal fin unbranched rays 2-4 (most anterior minute and embedded in flesh), and branched rays 5, pectoral fin branched rays 8-11, usually 9-10, and pelvic rays 6-8, usually 7. The dorsal fin origin lies midway between the snout tip and caudal base or nearer the caudal base. There is a fleshy pelvic axillary process but it does not extend beyond the end of the pelvic fin base. Caudal fin slightly emarginate in Turkmenistan but Hari Rud fish have a conspicuous fork. The two lobes of the caudal fin may be equal or either may be longer. The lobes are pointed or rounded. The dorsal adipose fin or crest begins shortly in front of the anal fin origin level and reaches the root of the caudal fin. There is a small gap between the dorsal fin insertion and the origin of the crest, and the depressed dorsal fin partially occupies this gap and the slight rise of the crest origin. It is strongly developed, high and thick. The crest is supported by 22-25 procurrent rays of the caudal fin (only visible on x-rays). Large fish have scales posteriorly on the body. Scales are small, oblong and widely separated. Bănărescu and Nalbant (1966) did not find scales on fish from the Harirud (confirmed for these fish (ZMUC P 2798, 2799, 27100, a fourth specimen P 2797 is lost; these catalogue numbers corrected from Bănărescu and Nalbant (1966) where incorrect; 63.1-68.3 mm standard length) from Obeh, Afghanistan; and not found on 3 fish, 38.1-60.9 mm standard length from Sarakhs on the Hari Rud) and this population may be distinct. Scales may be completely absent or only on the crest (Prokofiev, 2009). The lateral line extends nearly to the caudal base. Caudal peduncle short, 5-6 times in standard length (6.1-6.9 for 3 Iranian fish, 38.1-60.9 mm standard length). Lips are thick and furrowed, the upper lip with a slight median gap and the lower lip widely interrupted. The dentiform process on the upper jaw is weakly developed. The anterior nostril has a large flap developed posteriorly and postero-dorsally. Large fish have bulging cheeks. The posterior part of the intestine is straight (Bănărescu and Nalbant, 1966) or with a single loop (Bănărescu and Nalbant, 1995).

Meristics for Iranian specimens:- dorsal fin branched rays 8(4), anal fin branched rays 5(4), pectoral fin branched rays 9(2), 10(1) or 11(1); pelvic fin branched rays 7(4); and total vertebrae 36(1), 37(1) or 38(1).

Sexual dimorphism

There is no sexual dimnorphism.

Colour

Somewhat similar to P. malapterura but with a dark spot at the anterior base of the dorsal fin, on the unbranched and first branched rays. A similar spot may be present at the anterior base of the anal fin on the first two rays. The anterior part of the body is dark and the posterior half has 4-8 broad, brownish bands, separated by narrower light bands and reaching close to or to the ventral part of the body. Some smaller fish have anterior bands on the flank, the total number of bands reaching as high as 17. Bands vary in size from equal to twice the width of the pale interspaces. The last 3-4 bands extend onto the adipose fin. There is also a much darker and narrower band at the caudal base. Bănărescu and Nalbant (1995: Fig. 11A) illustrate a fish where this last band is only developed on the lower half of the caudal peduncle. The bar may also be composed of two blotches above and below the flank midline plus blotches on the bases of the outermost caudal rays both dorsally and ventrally. There 3-4 saddles anterior to the dorsal fin. All fins have thin bands comprised of spots irregularly lined up on the fin rays. The caudal fin may have three such bands while this fin and others may have no clear bands. Some fish have pigment on rays of the dorsal fin almost forming a single broad band extending from mid-fin half way towards the margin. There may be a second smaller spot on the last unbranched dorsal fin ray above the one on the anterior base.

Size

Attains 8 cm.

Distribution

Found in streams on the northern slope of the Kopetdag, Turkmenistan. Some of these streams presumably have headwaters in Iran and the species may be found there although this has yet to be confirmed by specimens. Also reported from the drainage of the Tedzhen River in Afghanistan which forms part of the Iran-Afghanistan border as the Hari Rud or Hari River (Bănărescu and Nalbant, 1966). This species is confirmed for Iran from Sarakhs on the Hari Rud, and probably the Kashaf River, a Hari River tributary (Abdoli, 2000).

A record from Lake Topiatan in the Uzboi River drainage in Turkmenistan north of the Iranian border mentioned in Nikol'skii (1947) is housed at the Zoological Institute, St. Petersburg (ZISP 25788). Berg (1948-1949) corrects the locality as the Sumbar River, an Atrak River tributary in the Caspian Sea basin. This specimen was examined by me and it is not this species but P. malapterura.

Zoogeography

Bănărescu and Nalbant (1966) consider that the closest relative of this species is Nemacheilus tigris (sic) from the Tigris-Euphrates basin rather than other crested loaches such as P. malapterura and P. rhadinaea found in neighbouring river systems. Note that Paracobitis tigris was described from Aleppo (= Haleb, Syria) and its occurrence in the Tigris-Euphrates basin is in question (see family account).

Habitat

A rheophilic species but further details of habitat preferences are unknown.

Age and growth

Patimar et al. (2011) studied fish from the Zanglanlou River in Khorasan Razavi Province. They found a maximum age of 4+ years, most fish (44%) were 1+, fish showed positive allometric growth, and male:female sex ratio was significantly different at 1:1.2, thought to be due to higher survival rate of adult females. Safai and Naserizadeh (2013) examined fish in the Radkan River of northeastern Iran. The sex ratio was 1:1, maximum age for both sexes was 6+ years, length-weight relationships were W=0.005166TL3.225 for males and W=0.006192TL3.125 for females, von Bertalanffy growth parameters were L=354.9 mm, k=0.0038, t0=-26.82 for males and L=339.0 mm, k=0.0043, t0=-24.88 for females, and growth performance indices were Φ’=6.17 for males and 6.20 for females.

Food

Gut contents of one fish, 60.9 mm standard length, from the Hari Rud were chironomid larvae.

Reproduction

Patimar et al. (2011) found the highest reproductive activity was in March to May, maximum number of eggs was 1246, mean absolute fecundity was 350.8 eggs, relative fecundity reached 1285.71 eggs and the mean was 287.78 eggs, and egg diameter reached 1.46 mm. Safai and Naserizadeh (2013) found the gonadosomatic index indicated peak reproduction in April-June with highest average values in May. Oocyte diameter was 0.09-1.58 mm, mean 0.54 mm.

Parasites and predators

Unknown.

Economic importance

None.

Conservation

Population numbers or trends are unknown.

Further work

More collections are needed to record information on biology and distribution.

Sources

Iranian material: CMNFI 2008-0138, 3, 38.1-60.9 mm standard length, Khorasan, Hari River at Sarakhs (36°32'N, 61°11'E); BM(NH) 1914.1.1:32, 1, 54.5 mm standard length, Khorasan, Kashaf River (no other locality data).

Comparative material: SMF 17133, 2, 34.4-98.3 mm standard length, Afghanistan, Hari River near Herat (34º21'N, 62º14'E); BM(NH) 1897.7.5:40, 51.4 mm standard length, Turkmenistan, Ashkhabad (no other locality data).

Genus Oxynoemacheilus
Bănăraescu and Nalbant, 1967

Members of this genus are found from Bulgaria east to Iran. There are 41 or more species with ?8 recorded from Iran. Characters below are based on Freyhof (2011).

These small loaches are scaled or scaleless, have a body with spots which tend to form bands or a stripe along the lateral line and there is no dark black spot on the base of the anterior dorsal fin rays. The dorsal fin origin is in front of or above the vertical from the pelvic fin origin and the anus is an eye diameter or less in front of the anal fin. Prominent black spot absent from the base of the unbranched, and sometimes first and second branched, dorsal fin rays (a saddle may extend onto the anterior base of the dorsal fin but there is no distinctive, very dark spot). There is usually no strong caudal peduncle crest. A crest, when present, is best developed at the caudal fin base and supported by procurrent rays. A small pelvic fin axillary lobe, variably developed, is present (but see Prokofiev (2009) where it is stated, erroneously, to be absent - Freyhof et al. (2011)). When a well-developed crest is present, it is usually highest at the caudal fin base and is supported by procurrent caudal fin rays. The caudal fin is very slightly emarginate to deeply emarginate or truncate. A forked caudal fin is found in a species with a cylindrical body and caudal peduncle. Males do not develop inflated or muscular cheeks in the spawning season but tubercles are found on the sides of the head and on the pectoral fins but not as fine brush-like aggregations. A suborbital flap or groove is present or absent in males, and may vary from present to absent within species. Nostrils are closely spaced or slightly separated, the mouth is semi-lunar with simple lips which are almost smooth to plicate and the dentiform process is weak to moderate but sometimes absent (O. angorae). The lateral line is usually complete but if incomplete as in O. kermanshahensis it reaches beyond the level of the dorsal fin origin or to the dorsal fin level (O. kiabii). The dorsal fin has 7-9 (11 in one species) branched rays and the anal fin 5 branched rays. Prokofiev (2009) lists various osteological characters such as 4 radial bones in the pectoral fin, usually 6 hypurals in the caudal fin support (O. brandtii has 5), the epural bone is present in the caudal fin support, the manubrium in the swimbladder is well-developed and the bony capsule is full divided by it, no preethmoid I, the supratemporal commissure is open from above, the fork of cst and cio is not fused to the skull, and others.

Oxynoemacheilus angorae
(Steindachner, 1897)

This taxon has been widely reported from northern Iran and several subspecies have been described under it. These taxa are now recognised as distinct species below. Freyhof et al. (2011) doubt that O. angorae s.s. occurs in the Caspian Sea basin and it may not be present in Iran at all. If O. angorae is restricted to Turkey, then the species for the western Caspian is lenkoranensis. Kura-Aras basin fish would then be araxensis but see below where this taxon could be in the Tigris-Euphrates basin (although adjacent headwaters could harbour the same taxon.

Nemacheilus angorae was originally described from Angora (= Ankara, Turkey). Sixteen syntypes of Nemacheilus angorae from Tabakane-Sir and Tschibuk-Tschai, Turkey are in the Naturhistorisches Museum Wieen according to Eschmeyer et al. (1996). Nemacheilus bergi Gratsianov, 1907 from the  Akstapha (= Akstafa) River, a right tributary of the Kura River, Azerbaijan is regarded as a synonym (types are lost) (Kottelat, 2012).

Some taxa or potential taxa peripheral to Iranian waters have been described namely:- Nemachilus angorae alasanicus Elanidze, 1983 was described from the upper reaches of the Alazani River at Alvani village, Georgia, a left bank Kura River tributary. It is here attributed to Elanidze (1983) since it is the only taxon listed in that book without an author; and the lower Kura River basin in Azerbaijan (of which the Aras is part) may have another subspecies in which about 40% of the fish have 7 branched dorsal fin rays, a shallower caudal peduncle than O. a. araxensis, and a colour pattern similar to O. angorae angorae (Banarescu, Nalbant and Balik (1978); see also description in Abdurakhmanov (1962) for measurements).

N. bergi and N. alasanicus may be synonyms of O. araxensis or O. lenkoranensis according to Freyhof et al. (2011) although N. bergi is an earlier name.

Oxynoemacheilus araxensis
(Banarescu and Nalbant, 1978)

Orthrias angorae araxensis Banarescu and Nalbant, 1978 in Banarescu, Nalbant and Balik (1978), is described from the Aras River basin of Turkey (type locality given below) (this subspecies was formerly referred to as Nemacheilus angorae bureschi (Drensky, 1928) by Banarescu and Nalbant (1964) and Banarescu (1968)). Nalbant and Bianco (1998), Fricke et al. (2007) and Freyhof et al. (2010) elevate this taxon to a species.

The holotype of Orthrias angorae araxensis, 62.0 mm standard length, from the "Kandili Karassu, oberes Araxes-Becken, Osttürkei" is in the Zoologischen Instituts und Zoologischen Museums der Universität Hamburg (ZMH 4827). Four paratypes, 45.0-65.2 mm standard length, from the same locality are under ZMH 5951 and 2 other paratypes, 51.2-59.3 mm standard length, also from the same locality are in the Institutul de Stiinte Biologice, Bucuresti, Romania (ISBB 2617). Five paratypes, 38.8-52.0 mm standard length, are from the "Oberlauf des Araxes Flusses bei Aras-Nehri, Hasankale" under ZMH 4826 and 3 paratypes, 41.0-45.0 mm standard length, from the same locality are under ISBB 2618 (Banarescu et al., 1978; Wilkens and Dohse, 1993). Note that Kottelat (2012) states that Kandili is a city on the upper Karasu in the Tigris-Euphrates basin

Oxynoemacheilus argyrogramma
(
Heckel, 1847)

Reported from the Tigris-Euphrates basin in Iraq but no Iranian record except by Saadati (1977).

Oxynoemacheilus bergianus
(Derzhavin, 1934)

Common names

agmahi-ye Safidruds, sehkhareh (?), sagmahi-e-jooibari.

[Sefidrudskii golets or Sefidrud loach in Russian; Safidrud stone loach].

Systematics

The type locality of Nemachilus bergianus in Latin from Derzhavin (1934) is "Systema fluminis Sefidrud" (= Safid River system). Berg (1948-1949) gives "Sefid-rud basin: Kisum village; Shah-rud R., falling into the Sefid-rud". The former is at Kisom at 37°14'N, 49°51'E or 37°12'N, 49°54'E in a gazetteer.

Bănărescu and Nalbant (1966) and Banarescu, Nalbant and Balik (1978) place this species as a subspecies of what is now called Oxynoemacheilus angorae (Steindachner, 1897). Nalbant and Bianco (1998) considered it to be a distinct species in Orthrias. Abdurakhmanov (1962) compares fish from Lenkoran (his O. angorae lenkoranensis) with O. bergianus. Head length and depth, predorsal distance, body depth, caudal peduncle depth, dorsal fin height, and pectoral and pelvic fin lengths are all greater in lenkoranensis while caudal peduncle length, pectoral-pelvic fin distance, interorbital width and eye diameter are all greater in bergianus.

The problems of the systematics of such a widespread and variable taxon as O. angorae are too complex to resolve here and I have retained O. bergianus as a distinct species until the problem has received a full study. Note also the confusion of some materials of this species with O. persa dealt with under that species description.

The holotype of Nemacheilus bergianus is 41.6 mm standard length and is in the Zoological Institute, St. Petersburg (ZISP 25433) and is from "Basin R. Sefid-Rud, c. Kissum, River Shahrud, tributary Sefid-Rud", collected by A. N. Derzhavin, 20.V.1922. The specimen is faded but there is a figure in Berg (1948-1949, Fig. 619).

Key characters

This species differs from related taxa by the following combination of characters (note that this is based mostly on very limited, often old material, and literature sources (often contradictory) - a thorough comparison of these fishes needs to be carried out):-

 
Character/Taxon araxensis bergianus brandtii lenkoranensis
Distribution Upper Aras River basin in Turkey Caspian, Namak and Orumiyeh basins in Iran Upper and middle Kura River and Aras River, Georgia and Azerbaijan (and Aras River in Iran?) Lenkoran, Azerbaijan (and western Caspian Sea basin in Iran?)
Caudal fin margin almost straight truncate, slightly emarginate deeply emarginate, pointed lobes, strongly forked deeply emarginate, forked
Caudal peduncle length/depth 2.3 in type examined by me (ZISP 36846) 3.3 (Derzhavin, 1934); 2.6 in type examined by me (ZISP 25433) 2.2-3.0 (Berg, 1948-49), 2.9-3.2 in Abdurakhmanov (1962); 1.7 in small type (19.3 mm SL) examined by me (BMNH 1897.7.5:39)) 2.0-2.6 (Abdurahkmanov, 1962)
Standard length/body depth 5.1-6.5 (Banarescu et al., 1978); 6.3 in type examined by me 6.5-8.5 (Derzhavin, 1934); 5.6 in type examined by me 6.2-6.6 (Berg, 1948-49); 4.9-7.6 (Banarescu et al., 1978); 5.7 in type examined by me -
Suborbital flap in males present present present absent

Morphology

Dorsal fin with 3-4 unbranched (the anterior 2-3 rays minute and not readily visible) and 8-9 branched rays, usually 8, anal fin with 3 unbranched and 5 branched rays, pectoral fin with 8-11 branched rays, and pelvic fin with 6-8 branched rays. Total vertebrae 35-37. The pelvic fins are separated by almost the width of a pelvic fin base. Caudal fin slightly emarginate. The upper lip is indented at the mid-point with a bony projection underneath. The lower lip is interrupted and also has a bony projection underneath. There are no adipose fins. The flank is minutely scaled and the lateral line is well-developed along the whole flank.

Meristics for Iranian fish: dorsal fin branched rays 8(57) or 9(1), anal fin branched rays 5(58), pectoral fin branched rays 8(1), 9(12), 10(41) or 11(4), pelvic fin branched rays 6(4), 7(48) or 8(6), and total vertebrae 35(10), 36(26) or 37(4).

Sexual dimorphism

Males have the anterior 2-3 rays of the pectoral fin expanded, with broad bands of fine tubercles overlapping on the rays and membranes, becoming less extensive medially. Less well-developed tubercles are present on the pelvic fin membranes. In "high" males fine tubercle are found all over the head and body, concentrated in some below the lateral line posterior to the first third of the pectoral fin level back to before the the pelvic fin level, generally lining scale margins (up to 6 per scale). the flap at the antero-ventral margin of the eye is well-developed in males, extending to below the nostril and sometimes as a thin band over the antero-dorsal eye margin. The edge of the flap may have fine tubercles.

Colour

The flank has several irregular dark grey to brown blotches. Some mid-flank blotches centred on the lateral line appear bar-like or form rough squares from the level of the dorsal fin origin posteriorly, but often they are less well-defined and much smaller. Posteriorly the flank blotches may give a barred appearance. The back has four transverse, dark, squarish spots or saddles behind the dorsal fin. The saddles may or may not extend to the mid-flank line but when they do the most posterior saddles are the most evident. The back anterior to the dorsal fin usually has no clear pattern although there is a saddle or arc of dark pigment over the back just in front of the fin. Some fish do bear saddles of pigment in front of the dorsal fin but these are much less obvious than the very dark ones behind the dorsal fin. The base of the caudal fin has a dark blotch, sometimes in the from of a vertical zig-zag pattern. The back and upper flank have a pinkish tinge. The lower flank is pale yellowish or whitish. The belly and lower head are white. The iris is silvery on the lower part and golden on the upper, strikingly so. The dorsal, pectoral and caudal fins are pinkish with 2-5 rows of elongate, dark grey to brown spots along the rays forming thin bars. Pigmentation on the pectoral fin is often along the rays and not in the form of rows. The leading edge of the dorsal fin may have elongated bars of dark pigment related to the thin bars on the rays, although often the most ventral, longest and darkest  may have no corresponding fin bar. The caudal fin spots are horizontal rather than forming bands as in Oxynoemacheilus brandtii. The upper base of the caudal fin is yellowish rather than dark as in O. brandtii and has 3-7 spots or elongate patches of pigment along its length, some of which line up with the thin bars. The pelvic and anal fins are colourless or slightly yellowish-pink without spots, although in some fish the pelvic fin has one row of spots. The barbels are typically without pigment except occasionally at the base of the second pair.

Size

Reaches 6.8 cm with Jolodar and Abdoli (2004) giving 7 cm total length.

Distribution

This species is found in the middle Aras River and its tributary the Qareh Chai (?or is araxensis), the Safid, Shafa, Siyah, Anzali Mordab and the Masouleh River in that basin, upper Safid River basin (Qezel Owzan and Shahrud), and the Karaj, Jaj, upper Shur or Abhar, middle and upper Qareh Chai and upper Qom rivers of the Namak Lake basin, southern and eastern tributaries of Lake Orumiyeh (Zarineh and Tata'u, and Talkheh rivers), and possibly in the eastern Dasht-e Kavir basin (Bănărescu and Nalbant, 1966; Saadati, 1977; Holčík and Oláh, 1992; Abbasi et al., 1999; Abdoli, 2000; Jolodar and Abdoli, 2004; Abdoli and Naderi, 2009)

Zoogeography

See family account.

Habitat

Unknown.

Age and growth

Females are mature at 5.0 cm. Heydarnejad (2009) gave the length-weight relationship for an Iranian sample ?where as W = 0.0078TL3.055. Golzarianpour et al. (2011) give a length-weight relationship for fish from the Jaj River identified as O. angorae ?check with a (intercept) being 0.008 and b (slope) 3.01.

Food

Unknown in detail but gut contents include aquatic insects. Abdoli (2000) lists Trichoptera, Ephemeroptera and Chironomidae for fish identified as ?.

Reproduction

Unknown, although fish with well-developed eggs have been captured as late as 15 July.

Parasites and predators

Unknown.

Economic importance

None.

Conservation

Kiabi et al. (1999) consider this species to be of least concern in the south Caspian Sea basin according to IUCN criteria. Criteria include few in numbers, habitat destruction, limited range (less than 25% of water bodies), present in other water bodies in Iran (sic), present outside the Caspian Sea basin (sic).

Further work

The biology and systematics of this species needs study. It is currently recognised as present in three basins although other nemacheilids (and cyprinids) are similarly distributed and are recognised as distinct taxa in these basins. A detailed study involving DNA might determine whether these basins harbour distinct taxa.

Sources

Type material: Holotype of Nemacheilus bergianus (ZISP 25433), see above.

Iranian material:  

Caspian:- CMNFI 1970-0521, 1, 33.4, mm standard length, Gilan, Safid River near Lulaman (no other locality data); CMNFI 1970-0527, 8, 33.1-42.7 mm standard length, Gilan, Safid River near Kisom (37º12'N, 49º54'E); CMNFI 1970-0537, 24, 33.3-50.4 mm standard length, Markazi, Shah River near Manjil (36º44'N, 49º24'E); CMNFI 1970-0545, 3, 34.7-44.4 mm standard length, Gilan, Safid River from Emamzadeh Hashem to Lulaman (ca. 37º01'N, ca. 49º38'E); CMNFI 1970-0556, 17, 22.6-58.6 mm standard length, Iran, Caspian Sea basin (no other locality data); CMNFI 1971-0341A, 13, 28.6-44.3 mm standard length, Gilan, Safid River (no other locality data); CMNFI 1979-0447, 2, 51.2-59.8 mm standard length, Gilan, stream 7 km east of Namin (38º23'N, 48º28'E); CMNFI 1979-0452, 2, 42.1-42.8 mm standard length, Azarbayjan-e Khavari, Qezel Owzan River 6 km from Mianeh (37º23'N, 47º45'E); CMNFI 1979-0453, 1, 34.2 mm standard length, Zanjan, Zanjan River (37º06'N, 47º56'E); CMNFI 1979-0692, 1, 27.7 mm standard length, Iran, Caspian Sea basin (no other locality data); CMNFI 1980-0132, 26, 18.5-44.1, Gilan, Safid River at Kisom (37º12'N, 49º54'E); CMNFI 1980-0160, 1, 46.4 mm standard length, Iran, Caspian Sea basin (no other locality data); CMNFI 2007-0081, 10, 42.2-58.6 mm standard length, Zanjan, Zanjan River basin near Soltaniyeh (ca. 36º27'N, ca. 48º45'E); CMNFI 2007-0082, 7, 31.2-53.8 mm standard length, Zanjan, Zanjan River basin near Zanjan (ca. 36º36'N, ca. 48º32'E); CMNFI 2007-0106, 17, 42.2-59.9 mm standard length, Kordestan, Qezel Owzan River basin near Divandarreh (ca. 35º52'N, ca. 47º05'E); CMNFI 2007-0107, 8, 34.7-57.0 mm standard length, Kordestan, Qezel Owzan River basin near Bijar (ca. 35º54'N, ca. 47º20'E).

Namak:- CMNFI 1979-0253, 7, 31.2-49.5 mm standard length, Markazi, Qareh Chay west of Baqerabad (34º52'N, 50º49'E); CMNFI 1979-0458, 1, 48.2 mm standard length, Markazi, Khar River 6 km north of Ab-Garm (35º47'N, 49º20'E); CMNFI 1979-0462, 4, 32.0-45.6 mm standard length, Markazi, Mazdaqan River (35º06'30"N, 49º40'30"E); CMNFI 1979-0463, 2, 41.0-51.6 mm standard length, Markazi, Qareh Chay (34º53'N, 50º26'E); CMNFI 1979-0465, 1, 37.1 mm standard length, Markazi, Qom River at Neizar (34º18'30"N, 50º32'E); CMNFI 1993-0152, 4, 34.3-48.4 mm standard length, Markazi, Sharra River near Khosbijan (34º07'N, 49º23'E); CMNFI 1993-0157, 2, 48.9-49.2 mm standard length, Markazi, Sharra River near Far (34º03'N, 49º19'E); CMNFI 1993-0159, 8, 36.8-48.8 mm standard length, Markazi, Sharra River near Pol-e Do Ab (34º03'N, 49º21'E); CMNFI 1993-0160, 3, 34.9-43.2 mm standard length, Markazi, Sharra River at Pol-e Do Ab (34º03'N, 49º22'E); CMNFI 1980-0154, 28, 10.9-26.8 mm standard length, Markazi, Karaj River below village (35º47'N, 50º58'E); CMNFI 1980-0156, 80, 29.9-50.8 mm standard length, Markazi, Karaj River near village (35º47'N, 50º58'E); CMNFI 2007-0074, 10, 25.6-50.8 mm standard length, Markazi, Qareh Su 32 km west of Arak (34º03'N, 49º21'E);CMNFI 2007-0121, 6, 32.0-54.1 mm standard length, Hamadan, Qareh Su basin north of Razan (ca. 35º25'N, ca. 49º02'E); CMNFI 2008-0152, 1, 43.5 mm standard length, Markazi, Namak Lake basin (no other locality data).

Orumiyeh:- CMNFI 1970-0559, 3, 14.2-37.3 mm standard length, Azarbaijan-e Bakhtari, Barunduz Chay (ca. 37º25'N, ca. 45º10'E); CMNFI 1970-0561, 29, 18.3-30.0 mm standard length, Azarbayjan-e Bakhtari, Cheagi Chai (no other locality data); CMNFI 2007-0084, 2, 28.5-43.4 mm standard length, Azarbayjan-e Khavari, Talkheh River basin west of Sarab (ca. 37º56'N, ca. 47º19'E); CMNFI 2007-0085, 12, 42.3-56.5 mm standard length, Azarbayjan-e Khavari, Talkheh River basin east of Sarab (ca. 37º56'N, ca. 47º41'E); CMNFI 2007-0091, 9, 36.2-49.9 mm standard length, Azarbayjan-e Khavari, Zilber Chay west of Marand (38º30'N, 45º23'E); CMNFI 2007-0095, 2, 47.5-53.3 mm standard length, Azarbayjan-e Bakhtari, Shahr Chay southwest of Reza'iyeh (ca. 37º27'N, ca. 44º56'E); CMNFI 2007-0096, 6, 36.5-51.3 mm standard length, Azarbayjan-e Bakhtari, Qasemul River south of Reza'iyeh (ca. 37º25'N, ca. 45º10'E); CMNFI 2007-0097, 4, 39.2-47.8 mm standard length, Azarbayjan-e Bakhtari, Baranduz Chay basin south of Reza'iyeh (ca. 37º16'N, ca. 45º08'E); CMNFI 2007-0099, 9, 24.6-43.8 mm standard length, Azarbayjan-e Bakhtari, Kalwi Chay west of Mahabad (ca. 36º35'N, ca. 45º25'E); CMNFI 2007-0103, 3, 36.4-55.8 mm standard length, Kordestan, Zarineh River basin north of Saqqez (36º18'N, ca. 46º16'E); CMNFI 2008-0279, 5, 52.3-57.6 mm standard length, Azarbayjan-e Khavari, "Kozem" or "Rozem" Chai (37º39'N, 45º36'E); BM(NH) 1899.9.30:131-137, 7, 33.3-35.8 mm standard length, Azarbayjan-e Khavari, Elinja Chai (= Alinja Chai, no other locality data).; BM(NH) 1899.9.30:138-140, 4, 41.8-53.4 mm standard length, Azarbayjan-e Bakhtari, Ula on the Zola Chai (= Zowla River, ca. 38º11N, ca. 44º51'E).

Oxynoemacheilus brandtii
(Kessler, 1877)

Qarasu, Aras River basin, August 2011, courtesy of K. Abbasi
Qarasu, Aras River basin, August 2011, courtesy of K. Abbasi

Common names

sagmahi-ye Kura.

[Kur cilpagcasi in Azerbaijan; Kurinskii golets or Kura loach in Russian].

Systematics

A syntype of Nemacheilus Brandtii from the upper Kura River at Tbilisi (= Tiflis), Georgia is in the Natural History Museum, London (BM(NH) 1897.7.5:39, 19.6 mm standard length, (small and decoloured), formerly in ZISP; other syntypes are in the Zoological Institute, St. Petersburg (ZISP) (Eschmeyer et al., 1996).

Nemachilus brandtii gibbusnazus is a subspecies with the author given as Elanidze in Elanidze (1983). The only apparently new taxon in Elanidze (1983) is Nemacheilus (= Oxynoemacheilus) angorae alasanicus (q.v.). This taxon is unique in this book as a species without an author name after it. On this basis it was attributed to Elanidze (1983). The date of N. b. gibbusnazus may also be 1983 but it may have been described in an earlier paper by Elanidze not yet located by me. Both these subspecies are not mentioned in Eschmeyer et al. (1996) nor in the online version (downloaded 26 August 2007). The distribution of N. b. gibbusnazus is given as "in fact found before in the R. Khrami (1947), then in the R. Kura (1962), in its lower course at Kukheti, in the upper reaches at Vardziya - Toloshi, in the R. Alazani - in the upper reaches at Alvani", all in Georgia and the drainage of the Kura River.

Placed in the genus Orthrias by Nalbant and Bianco (1998) and in Oxynoemacheilus by Freyhof et al. (2011).

Key characters

This species is distinguished by a more forked caudal fin with more pointed lobes. The caudal peduncle is said to be somewhat shorter and much lower than in O. araxensis (with which species it occurs in the Aras River basin): length of caudal peduncle 18.4-23.4% and depth 8.0-9.4% of standard length in brandtii, 18.1-21.8% and 10.2-12.7% in araxensis respectively. Other body proportions are given in Banarescu, Nalbant and Balik (1978). The colour pattern is much darker and the caudal fin margin is almost vertical in O. araxensis.

Morphology

Dorsal fin unbranched rays 3-4 and branched rays 7-9, usually 8, anal fin unbranched rays 2-3 and branched rays 5, pectoral fin branched rays 9-12, and pelvic fin branched rays 6-8. Scales minute. A weakly developed dentiform process on the upper jaw.

Sexual dimorphism

Unknown.

Colour

Typical fish from the Kura-Aras basin lack a stripe on the flank, having brown spots which are either large, more or less triangular and fairly well-defined or broken into many small speckles forming a reticulate pattern. Dorsal spots are better defined than in B. angorae especially behind the dorsal fin where they fuse completely or incompletely with the lateral flank spots to form bands. All spots are brown, never blackish as in B. angorae araxensis. There are several rows of speckles on the caudal fin and two rows on the dorsal fin.

Size

Reaches 8.5 cm.

Distribution

Found in the upper and middle Kura and Aras River basins, and presumably in the Iranian reaches of the latter. It is also reported from the Namak Lake basin by Saadati (1977) citing a manuscript report by V. D. Vladykov, ? confusion with bergianus and from the Arnar Chay, Azarbayjan.

Zoogeography

See family account.

Habitat

Unknown.

Age and growth

Age at maturity is about 2 years. Golzarianpour et al. (2011) give a length-weight relationship for fish from the Zarrineh River with a (intercept) being 0.009 and b (slope) 2.97.

Food

Eggs of other fishes may be a food item as well as aquatic insects.

Reproduction

Fecundity reaches 17,409 eggs and egg diameter 0.95 mm.

Parasites and predators

Unknown.

Economic importance

None.

Conservation

Endangered in Turkey (Fricke et al., 2007).

Further work

The biology and systematics of this species need study.

Sources

Type material: Syntype of Nemacheilus Brandtii ((BM(NH) 1897.7.5:39), see above.

IDs to check:- CMNFI 1979-0448, 9, 31.1-62.0 mm standard length, Azarbayjan-e Khavari, Ahar Chay 8 km from Ardebil (38º18'30N, 48º22'E); CMNFI 1979-0450, 3, 38.7-40.9 mm standard length, Azarbayjan-e Khavari, stream near Kivi (ca. 37º37'N, ca. 48º17'E); CMNFI 2007-0083, 2, 28.7-33.1 mm standard length, Azarbayjan-e Khavari, Qaranqu River basin west of Sar Eskand Khan (ca. 37º25'N, ca. 46º55'E); CMNFI 2007-0086, 3, 42.5-59.1 mm standard length, Azarbayjan-e Khavari, Qareh Su basin near Nir (ca. 38º02'N, ca. 48º00'E).

Oxynoemacheilus cyri
(Berg, 1910)

Nemacheilus tigris cyri Berg, 1910 is described from "Fontes fl. Cyrus, Caucasus". This species was placed in the subgenus or genus Paracobitis by Banarescu (1968) and Prokofiev (2009) while Fricke et al. (2007) recognise the subspecies as a distinct species in Orthrias. Freyhof et al. (2011) place it in Oxynoemacheilus. Six syntypes of N. tigris cyri (31.4-51.5 mm standard length) are in the Zoological Institute, St. Petersburg (ZISP 13291) from "Okam village Gel'skaya Plain, Karka Oblast", K. Satunin, 6.IX.1901. This is in the upper Kura River basin of Turkey. No Iranian record.

Oxynoemacheilus frenatus
(Heckel, 1843)

Common names

See genus account.

[lakh or telay (= bowed head according to Heckel (1843b) at Mosul), both in Arabic; banded Tigris loach].

Systematics

The type locality of Cobitis frenata is "Tigris", presumably at Mosul (Heckel, 1843b). Five syntypes are in the Naturhistorisches Museum Wien (NMW 48552) although the catalogue lists only 4 specimens. A lectotype designated by F. Krupp in 1984 is 70.0 mm standard length, the remaining specimens being small, 27.2-43.4 mm standard length.

Nemacheilus frenatus afrenatus Battalgil, 1942 described from "un petit ruisseau à Diyarbakir" is also from the Tigris River basin but in Turkey. This subspecies lacks the "frein à la bouche" (presumably the band across the snout) of B. f. frenata and its dorsal fin is higher than long (as measured at the base) while in B. f. frenata it is as high as long.

Bănărescu and Nalbant (1995) place this species in the genus Orthrias. Bănărescu and Nalbant (1966) consider this taxon to be a "doubtful species" but illustrate it in Bănărescu and Nalbant (1995: Fig. 20).

Key characters

The colour pattern is distinctive and there is no dermal crest or adipose fin behind the dorsal fin.

Morphology

Dorsal fin unbranched rays 2-3, branched rays 7-8, anal fin unbranched rays 2, branched 5, pectoral fin branched rays 10-13, and pelvic fin branched rays 6-7. Scales are present over the whole body but not readily visible without magnification. The anterior pectoral fin rays are thickened. Caudal peduncle thick (depth 80-90% of length) according to Saadati (1977). The bulb of the swimbladder capsule has large ovoid to circular perforations and the anterior and posterior lamina or wings are only moderately developed (Krupp, 1985c). Lips are not strongly plicate and the dentiform process is well-developed. The gut has a posterior loop.

Meristics for Iranian specimens: dorsal fin branched rays 8(1), anal fin branched rays 5(1), pectoral fin branched rays 10(1) and pelvic fin branched rays 6(1) based on lectotype.? add my material

Sexual dimorphism

Bands of tubercles are found on the pectoral fin rays of males, including the first, declining in breadth and extent on the smaller rays. Tubercles are also present on the pelvic and anal fin rays but are much less well developed. The head is covered in fine tubercles. Flank scales, particularly anterior ones, are lined anteriorly with tubercles. There is an elongate tuberculate swelling anterior to the lower eye margin on the snout.

Colour

Overall colour is yellowish, mottled with fine but irregular brown or black dots or blotches, some flank blotches being quite large. The rear of the body and the caudal fin in particular are mottled with brown, tending to form bars. A black band is continuous from the front of one eye, across the snout and round to the other eye. It may be diffuse on the snout or well-defined in fish from the same locality. The dorsal and caudal fins have thin but irregular bands made up of spots on the rays (up to 3 on the dorsal and 4 on the caudal fin), bands are faintly present on the anal and pelvic fins, and only a few are visible on the pectoral fins. The dorsal fin has an anterior basal spot at its origin, variably developed in individuals. There are distinct, dark spots at the base of the caudal fin above and below the body mid-line.

Size

Reaches about 9.2 cm (Heckel, 1843b).

Distribution

Found in the Quwayq and Tigris-Euphrates rivers. Abdoli (2000) records it from the upper Karun, middle and lower Dez, Kashkan and Simarreh rivers in the Tigris River basin of Iran.

Zoogeography

See family account.

Habitat

Known to inhabit both rivers and lakes, the environmental requirements of this species are unknown.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

None.

Conservation

The biology of this species, its distribution, numbers and habitat requirements are unknown so no comments on conservation can be made. Near threatened in Turkey (Fricke et al., 2007).

Further work

The biology and conservation status of this species need investigation.

Sources

Type material: Syntypes of Cobitis frenata (NMW 48552), see above.

Iranian material: BWC 95-30, mm standard length, .

Comparative material: BM(NH) 1974.2.22:1449-1477, 30, 37.7-55.5 mm standard length, Iraq, branch of Khalis River (); BM(NH) 1974.2.22:1478-1757, ca. 279, 19.7-53.3 mm standard length, Iraq, Khalis (); BM(NH) 1974.2.22:1758-1772, 15, 27.2-34.1 mm standard length, Iraq, Khalis (); BM(NH) 1972.2.22:1773-1776, 4, 28.3-34.0 mm standard length, Iraq, Khalis (); BM(NH) 1974.2.22:1777-1778, 2, 27.5-31.4 mm standard length, Iraq, Khalis ().

Oxynoemacheilus kermanshahensis
(Bănărescu and Nalbant, 1966)

Dehno Stream, Gamasiab River, Hamadan, courtesy of K. Golzarianpour
Dehno Stream, Gamasiab River, Hamadan, courtesy of K. Golzarianpour

Common names

sagmahi-ye Kermanshah.

[Kermanshah loach].

Systematics

This species was tentatively placed in the subgenus Orthrias Jordan and Fowler, 1903 but is regarded as "aberrant" by Bănărescu and Nalbant (1966). However, Nalbant and Bianco (1998) later place it in Orthrias.

The holotype of Noemacheilus kermanshahensis (ZMUC P 2787, 46.4 mm standard length) and the 7 paratypes (ZMUC P 2788-94, 25.5-61.8 mm standard length) are stored in the Zoological Museum of Copenhagen (Nielsen, 1974). The type locality is "Kermanshah in the drainage of the Karun River, a tributary of the lower Euphrates, Western Iran" (Bănărescu and Nalbant, 1966). The type series was collected on 5 February 1937 by E. Kaiser. The type locality is poorly defined - the city of Kermanshah is on the Qarasu River and lies in the drainage of the Karkheh River which drains towards the Tigris River. Nalbant and Bianco (1998) correct their original type locality description to the Karkheh River drainage and "Quareh Su, Kermanshah, River Shimarek", probably referring to the Qareh Su-Simareh drainage of the upper Karkheh River.

Key characters

Bănărescu and Nalbant (1966) consider that this species differs from most South and West-Asiatic loaches by the longitudinal pigment pattern on the flank but not all fish have this pattern. Most other species have bars, vertical patches of pigment. The longitudinal arrangement is found in O. angorae but kermanshahensis has smaller eyes, a shorter lateral line (almost complete in angorae), a more anteriorly placed vent, and scale shape comprising an almost central focus, vertical sub-oval outline and radii on all fields widely and evenly spaced. Characteristically the caudal peduncle is short and deep, depth being 92.9-114.3% of length (Saadati, 1977). The dorsal fin branched ray count of 7 is also a key character.

Morphology

Dorsal fin with 3 unbranched and 7 branched rays, anal fin with 2 unbranched and 5 branched rays, pectoral fin with 8-10 branched rays, and pelvic fin with 6-7 branched rays. Scales only on the posterior part of the body, well-developed on the caudal peduncle. The lateral line reaches the level of the middle or posterior half of the dorsal fin or over the front half of the anal fin. The origin of the dorsal fin is variable in relation to the snout tip and caudal fin base (Bănărescu and Nalbant, 1966). The caudal peduncle is short and deep and often lacks a crest although a weak crest may be developed just anterior to the caudal fin. The anus is somewhat anterior to the anal fin origin. Lips are thick and fringed, the lower lip being interrupted in the middle. The intestine is simple with the posterior part straight.

Meristics for Iranian material, including the type series: dorsal fin branched rays 7(10), anal fin branched rays 5(10), pectoral fin branched rays 8(3), 9(2), or 10(4), pelvic fin branched rays 6(3) or 7(7), and total vertebrae 38(7), 39(2) or 40(1).

Sexual dimorphism

No sexual dimorphism was noted by Bănărescu and Nalbant (1966) as their large specimen is female and others immature. Freyhof et al. (2011) note the presence of a groove at the antero-ventral margin.

Colour

The overall colour is yellowish with continuous, dark brown blotches on the flank my fish. A series of connected blotches line the upper flank, most obvious as blotches postdorsally, and separated from the mid-flank stripe by clear areas. A ventral stripe composed of blotches is also present and more clearly defined than indicated below. There is a bar at the caudal fin base. A broad bar is seen in some fish in the middle of the caudal fin with a less broad bar distally. A stripe may be present on the mid-line of the back and break into spots posteriorly, or the back may be uniformly pigmented.

The colour pattern in alcohol-preserved specimens dating from 1937 is yellowish with 3 wide, brownish stripes along the flank. The dorsal stripe is continuous anteriorly but breaks up into spots posteriorly in most specimens. The central, mid-flank stripe is the widest and is continuous although width is variable. The ventral stripe is absent from smaller specimens and consists of small and irregular spots. The upper parts of the head have small, irregular brownish spots. The caudal fin has 3 rows of spots and the dorsal fin 2 rows of spots, apparently concentrated on the fin rays (Bănărescu and Nalbant, 1966).

Size

Reaches 6.3 cm standard length. Golzarianpour et al. (2011) record a total length of 8.4 cm.

Distribution

Known only from Tigris River basin drainages of Iran (see above). Abdoli (2000) reports it from the Marun, upper Karun and lower Khersan, Dez, Simarreh, Qareh Su and Gav Masiab rivers.

Zoogeography

Relationships may lie with B. angorae according to Bănărescu and Nalbant (1966) based on colour pattern and general body shape, or with other species not found in Iran having stripes along the flank. This seems insufficient evidence on which to base relationships.

Habitat

Unknown.

Age and growth

Unknown in detail. Golzarianpour et al. (2011) give a length-weight relationship for fish from the Gamasiab River with a (intercept) being 0.015 and b (slope) 2.89.

Food

Unknown.

Reproduction

Unknown, although fish caught on 11 May had well-developed eggs..

Parasites and predators

Unknown.

Economic importance

None.

Conservation

No measures are being undertaken for this poorly known species.

Further work

Studies should be carried out to determine the numbers of this species, its distribution and its ecological requirements in Iran to ascertain if conservation measures should be taken.

Sources

Type material: The holotype (ZMUC P 2787) and paratypes (ZMUC P 2788-94) of Noemacheilus kermanshahensis, see above.

Iranian material: CMNFI 1993-0128, 1, 50.0 mm standard length, Kermanshahan, Sarab-e Sabz 'Ali Khan (34º25'N, 46º32'E); CMNFI 2008-0117, 510 mm standard length, Kermanshahan, Sarab-e Yavari (34º28'N, 46º56'E).

Oxynoemacheilus kiabii
Golzarianpour, Abdoli and Freyhof, 2011

Gamasiab River, Karkheh River basin, January 2010, courtesy of K. Abbasi
Gamasiab River, Karkheh River basin, January 2010, courtesy of K. Abbasi

Common names

None.

Systematics

The holotype ZFMK 41847, 54.5 mm standard length, is in the Zoologisches Forschungsmuseum Alexander Koenig, Bonn and is from Hamadan, Dehnoo stream, 3 km west of Nahavand, 34º10'N, 48º24'E. Paratypes are FCKG (Fish Collection, K. Golzarianpour) 8658, 8 specimens, 31.4-50.7 mm standard length, same locality as holotype, FCKG 8659, 7, 33.6-53.3 mm standard length, same locality as holotype, and FSJF (Fischsammlung J. Freyhof, Berlin) 3004, 2, 38.4-44.4 mm standard length, Babarostam, Gamasiab River tributary, 34º10'18"N, 48º21'23"E. The species is named after Bahram H. Kiabi for his conservation work, especially on fishes.

Key characters

This species is distinguished from other members of the genus by having two, rarely one or three, lateral pores (on each side) in the supratemporal head canal and no central pore; no suborbital flap and no groove in males; an incomplete lateral line reaching the level of the dorsal fin; a slightly emarginate caudal fin; a short pectoral fin not reaching the pelvic fin origin; a shallow adipose crest on the caudal peduncle dorsally from the level of the last dorsal fin ray base to anterior part of caudal fin (looks more extensive on photographs in type description); a long head (26-30% standard length); and colour pattern.

Morphology

A stout and deep-bodied species with a long and wide head. Many fish have a distinct hump at the nape. The compressed caudal peduncle is about as deep as long. Dorsal fin branched rays 8, anal fin branched rays 5, pectoral fin rays 10 and pelvic fin rays 8. The pelvic fin base has a short, triangular axillary lobe completely attached to the body. The incomplete lateral line has 18-29 pores. The body is scaled except on the back predorsally and the belly. On fish had three lateral pores in the supratemporal head canal and two fish had one left pore and two right pores. Lips are thick and well-furrowed, the lower lip interrupted medially. Processus dentiformis small and pointed. There is a shallow ventral crest on the caudal peduncle.

Sexual dimorphism

Males have slightly longer pectoral fins than females.

Colour

Overall body colour is brown in life and grey in preserved fish. The flank has 10-19, dark brown or black, large bars or blotches, most evident posterior to the dorsal fin level, and more irregular anterior to the dorsal fin level. The flank bars or blotches are not connected to predorsal back blotches. Blotches below the lateral line are smaller than those above it. A bar at the tail base may also take the form of two half-moon blotches. Head spotted and blotched, sometimes marbled. A tesselated pattern on the caudal fin rays forms 2-3 irregular bars and a similar pattern on the dorsal fin forms 3 bands. Other fins have a few dark brown or black spots on the rays.

Size

Reaches 56.5 mm standard length.

Distribution

Found in the Karkheh River drainage at the localities listed above for the type series.

Zoogeography

See family account.

Habitat

Found in clear waters of streams with a swift current.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

None.

Conservation

Known only from a restricted area with no data on numbers and threats.

Further work

The biology of this species and its conservation status need study.

Sources

The type description.

Oxynoemacheilus lenkoranensis
(Abdurakhamanov, 1962)

Lankaran cilpaxcasi (or cilpagcasi), both in Azerbaijan; Lenkoranskii golets or Lenkoran loach, both in Russian.

Nemacheilus angorae lenkoranensis Abdurakhamanov, 1962 (incorrectly lenkoranica in Bănărescu and Nalbant (1966)) is described from "rivers of the Lenkoran coast; Lenkoranchai, Vilyazhchai, Kumbashichai, Tangyaru, Astarinka" in the southern Caspian Sea basin.

A series of 18 fish in the Zoological Institute, St. Petersburg catalogued as ZISP 35701 from "Reka Lenkoran, Azerbaidzhan SSR" collected by Yu. Abdurakhmanov, 22.IX.1954 are probably the type series of Nemacheilus angorae lenkoranensis although they are not marked as such on the jar. One fish is a cobitid, 31.0 mm standard length (probably Cobitis keyvani), while the rest measure 24.9-33.3 mm standard length.

Oxynoemacheilus namiri
(
Krupp and Schneider, 1991)

Recorded from the Tigris River basin in Turkey by Nalbant (1998) but no Iranian record. Originally described in the genus Nemacheilus.

Oxynoemacheilus persa
(Heckel, 1847)

Kor River basin courtesy of H. R. Esmaeili, A. Parsi, Z. Gholami and M. Tahami
Kor River basin courtesy of H. R. Esmaeili, A. Parsi, Z. Gholami and M. Tahami

Common names

sagmahi-ye Fars (= Persian loach).

[Persian loach].

Systematics

The type locality for Cobitis Persa is "Quellen um Persepolis" according to Heckel (1847b). Kähsbauer (1964) reports a syntype of this species in the Naturhistorisches Museum Wien under NMW 48567. It measures 47.6 mm standard length and is probably the holotype as the Vienna catalogue lists only 1 specimen (and the Vienna card catalogue examined in 1997 concurs). This specimen is in poor condition and not readily comparable to fresh material. It is scaled although scales are not imbricate, the caudal fin is broken off, the body is collapsed so its shape cannot be determined, and it is decoloured.

Kessler (1877) refers to a Heckel species Nemachilus persicus, presumably this taxon judging from Kessler's page number reference. The trivial name is spelled persa as it is a noun in apposition (see Catalog of Fishes; Kottelat (2012)). Kotteleat (2012) gives the publication date as 1848.

Günther (1899) recorded Nemacheilus persa from the Lake Orumiyeh basin ("Zola Chai near Ula" - see materials under O. bergianus) and the upper Aras River basin ("Elinja Chai" - see materials under O. bergianus) but these fish were possibly O. angorae or O. bergianus. Berg (1948-1949), Bănărescu and Nalbant (1966) and Nalbant and Bianco (1998) also refer Günther's material to Nemacheilus persa but Abdurakhmanov (1962) suggests that these fish are O. angorae. Saadati (1977) places Lake Orumiyeh fish close to O. angorae which makes more sense geographically. The dorsal fin origin is closer to caudal base than the snout tip in the Lake Orumiyeh fish, interorbital width is greater, the head is longer, the caudal fin less emarginate, and there are more and darker spots on the body (Saadati, 1977). Evidently new material from the Kor River basin wherein lies Persepolis, the type locality for Cobitis persa, and the Lake Orumiyeh basin are required to resolve this problem - the Lake Orumiyeh sample examined by Bănărescu and Nalbant (1966) comprised only 3 males. The loach from the Lake Orumiyeh basin is now referred to O. bergianus, q.v.

Orthrias farsicus Nalbant and Bianco, 1998, named for Fars Province, collected from the "River Kor near Persepolis" in Fars is a synonym (Freyhof et al., 2011).

The holotype of Orthrias farsicus is in the Department of Zoology, Naples University under IZA 7823. It measures 45.8 mm standard length (51.8 mm standard length when measured by me) and was collected from the "River Kor near Persepolis" in Fars on 30 May 1976 by P. G. Bianco (note that IZA is the acronym for Dipartimento di Scienze Ambientali, Universita' Degli Studi Dell'Aquila L'Aquila, Italy, where the material was previously stored). Paratypes were all collected by P. G. Bianco and include 64 specimens, 39.0-57.0 mm standard length from the same locality as the holotype (IZA 7824), 61 specimens, 38.5-60.0 mm standard length from the same locality as the holotype but taken on 7 July 1975 (IZA 7825), 7 specimens, 42.0-51.3 mm standard length, same locality as the holotype but taken on 7 July 1975 stored in the Institutul Stiinte Biologice, Bucharest (ISSB 3451), 31 specimens, 43.2-56.3 mm standard length from the River Shur, tributary of the River Mand, near Dasht-e-Arzhan (Shiraz) 25 May 1976 (IZA 7826), 7 specimens, 47.0-57.0 mm standard length from the latter locality (ISBB 3446), 7 specimens, 47.0-58.0 mm standard length from River Qom, Qom Town 5 May 1975 (IZA 7845) and 7 specimens, 36.2-47.3 mm standard length from the latter locality (ISBB 3442). Three specimens from each of IZA 7824 (40.2-50.5 mm standard length) and 7826 (46.6-50.1 mm standard length) are in the Canadian Museum of Nature, Ottawa (CMNFI 2012-0005 and 2012-0006 respectively).

Bănărescu and Nalbant (1966) place this species in their subgenus Oxynoemacheilus but later (Bănărescu and Nalbant (1995) and also Nalbant and Bianco (1998)) place it in Orthrias. Freyhof et al. (2011) place the species in Oxynoemacheilus.

Key characters

This is the only known Oxynoemacheilus species from the Kor River basin and colour pattern and tuberculation are distinctive.

Morphology

This fish has long paired fins and a short, deep and compressed caudal peduncle. Lips and barbels have small tubercles and fine furrows. The upper lip has a median incision and the lower lip lacks mental lobes. The processus dentiformis in the upper jaw is well-developed but lacks a gap in the lower jaw. Dorsal fin with 2-4 unbranched (anteriormost very small and only visible under x-rays hence literature range) and 7-9, usually 8, branched rays, anal fin with 2 unbranched and 4-5, usually 5, branched rays, pectoral fin with 8-12 branched rays, pelvic fin with 6-8 branched rays, and total vertebrae ?. The type description of O. farsicus (a synonym) has dorsal fin branched rays 7-9 (90% with 8 rays in the type description, n=38), anal fin branched rays 4-5 (95% with 5 rays), pectoral fin branched rays 9-11 (69% with 10 rays) and pelvic fin branched rays 6-7 (65% with 7 rays). The lateral line is straight, on the mid-flank and ends just anterior to the caudal fin base. The cephalic canal system is well-developed laterally. Scales are present on the anterior and posterior flank according to Banarescu and Nalbant (1964) and Nalbant and Bianco (1998) for O. farsicus state that the rear half of the body is scaled completely and the anterior half has scalation well-developed but not as closely spaced as posterior scales. Scales are oval with a large focus. Esmaeili and Niknejad (2006-2007) give scanning electron micrographs of the scales along with a description. The caudal fin is slightly emarginate to forked. The barbels are hair-like thin (Heckel, 1847b).

Meristics for Iranian specimens including the holotype of O. persa and the holotype and 6 paratypes of O. farsicus: dorsal fin branched rays 8(47) or 9(2), anal fin branched rays 5(49), pectoral fin branched rays 9(1), 10(25), 11(21) or 12(1), pelvic fin branched rays 6(13), 7(35) or 8(1), and total vertebrae 35(2), 36(13), 37(23) or 38(2).

Sexual dimorphism

Banarescu and Nalbant (1964) report that pectoral fin rays 2-5 are widened and thickened in males. The appearance of a subocular pad in Bănărescu and Nalbant (1966:Fig. 3) is an error and was apparently meant to represent a small groove (Bănărescu and Nalbant, 1995). Nalbant and Biamco (1998) describe fish identified as O. farsicus (a synonym) as follows with additional comments by me in parentheses. Males have numerous fine tubercles on the dorsal surface of the pectoral fin rays in bands (following the rays but extending onto the membranes), and the anterior fn rays are expanded. There are fewer rays and less expansion when progressing proximally. There is a groove running anteriorly at a slight downward angle from the eye to the antero-ventral corner below the nostril, fading out under the anterior nostril. Near the eye and dorsal to this groove, fine tubercles form a narrow band. The sides and top of the head are finely tuberculate. The pelvic and dorsal fin rays have tubercles following the rays (not in bands as on the pectoral fin). Fine tubercles line the margin of the scales (and are especially evident on the anterior lower flank).

Colour

The overall colour is yellowish with brown to black blotching. There are 5-7 brown bars or blotches (shape varies) on the posterior body and irregular brown to black blotches on the anterior body, numbering up to about 12, being less well-defined anteriorly. The back has 7-11, modally 9, rounded dark-brown saddles, most apparent post-dorsally. The dorsal head is speckled brown and there is a dark-brown stripe from the eye to the snout tip, sometimes extending beyond the eye. The dorsal fin has 3-4 rows of elongate spots, the pectoral fin has up to 6 rows (not always in clearly defined rows but extending along the whole ray) and the caudal fin has 4-5 rows. The rows may be poorly defined with pigment spread along rays and not on the membranes. The anal and pelvic fins are immaculate although some fish may have pigment on the pelvic rays but not in rows. The description of O. farsicus is as follows. The flanks and back have roundish brown spots on a marbled background and the head and body are a pale yellow ventrally. Dorsal and caudal fins have rows of brown spots. The peritoneum is a dull brown.

Size

Reaches 67.7 mm standard length. Golzarianpour et al. (2011) give 7.1 cm total length.

Distribution

Found in the Kor River and Lake Maharlu basins (Esmaeili et al., 2011?). Abdoli (2000) maps the Kor and Pulvar rivers. Nalbant and Bianco (1998) report it (as O. farsicus) from the Namak Lake basin (Qom River at Qom - this is remote from the type locality of O. persa and needs further investigation), Kor River basin (Kor River near Persepolis) and Gulf basin (Shur River near Dasht-e Arzhan in the Mand River drainage). Records by Bănărescu and Nalbant (1966) in the Lake Orumiyeh basin are in error (see above).

Zoogeography

See family account.

Habitat

Unknown.

Age and growth

Unknown. Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 32 fish measuring 2.94-6.32 cm standard length. The a-value was 0.0211 and the b-value 2.784 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases).  Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 34 fish identified as O. farsicus (a synonym) measuring 3.15-6.77 cm standard length. The a-value was 0.0126 and the b-value 3.084 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases). Golzarianpour et al. (2011) give a length-weight relationship for fish from the Kor River with a (intercept) being 0.012 and b (slope) 2.88, the difference with the previous study attributed to small sample sizes.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

None.

Conservation

Conservation requirements are unknown but it may be relatively common as a secretive species.

Further work

The abundance and habitat requirements of this species need to be determined.

Sources

Type material: The holotype of Cobitis persa (NMW 48567), see above, and the holotype (IZA 7823) and paratypes (IZA 7824, 7825, 7845) of Orthrias farsicus, see above.

Iranian material: CMNFI 1979-0028, 5, 48.3-55.6 mm standard length, Fars, Kor River drainage (no other locality data); CMNFI 1979-0059, 28, 32.8-57.0 mm standard length, Fars, Pulvar River 8 km south of Sivand (30º01.5'N, 52º57'E); CMNFI 1979-0061, 3, 50.2-66.5 mm standard length, Fars, stream tributary to Pulvar River 14 km south of Sa'adatabad (30º04'N, 53º01'E); CMNFI 1979-0067, 1, 55.8 mm standard length, Fars, qanat at Zarqan (ca. 29º46'N, ca. 52º43'E);  CMNFI 1979-0070, 2, 38.8-39.9 mm standard length, Fars, Pulvar River near Naqsh-e Rostam (29º59'N, 52º54'E); CMNFI 1979-0117, 2, 42.1-50.6 mm standard length, Fars, Pulvar River at road bridge to Naqsh-e Rostam (29º59'N, 52º54'E); CMNFI 2008-0255, 47.9-51.3 mm standard length, Fars, Kor River (30º00'N, 52º45'E).

Oxynoemacheilus tongiorgii
(Nalbant and Bianco, 1998)

Common names

sagmahi-ye Hormuz.

[Hormuz loach].

Systematics

This species is named for Professor Paolo Tongiorgi, University of Modena. The holotype measures 23.7 mm standard length and is from " large water spring near Darab town, Kul river basin", collected on 2 June 1976 by P. G. Bianco and is stored in the Department of Zoology, University of Naples (IZA 801 but not located in a 2002 visit by me). One paratype, 20.0 mm standard length, is from the same locality (Institutul Stiinte Biologice, Bucharest, ISBB uncatalogued).

It differs from an Anatolian species, Seminemacheilus lendlii (Hankó 1925), mostly by colour pattern and possession of scales. It was originally described in the genus Seminemacheilus Banarescu and Nalbant, 1995 but Freyhof et al. (2011) transfer it to Oxynoemacheilus on unpublished DNA evidence. Kottelat (2012) gives a publication date of 1999 but the journal is dated November 1998.

Key characters

This species has a short and deep body, deeper than in related species in Iran, and a deep and short caudal peduncle. Scales are scattered and non-imbricate.

Morphology

The body is stout with a short and deep caudal peduncle. Caudal peduncle depth in length is 0.9-1.2, mean 1.0 for 23 fish), and larger fish tend to have a deeper caudal peduncle (this character may be difficult to measure as dorsal and ventral crests may vary in development and be collapsed). The short caudal peduncle means the anal fin reaches, or almost reaches, the caudal fin base. The crest on the caudal peduncle is low and originates at the caudal fin base, decreasing in height anteriorly and not reaching the dorsal fin insertion. A small ventral keel is also present. In the original description the body is said to be almost entirely scaleless except for minute scales on mid-flank but flanks and back are all scaled in samples seen here. Scales are scattered and non-imbricate. The belly anterior to the pelvic fins is scaleless except for some non-imbricate scales scattered immediately anterior to the fins. Radii are present on all fields, radiating from an almost central focus on anterior flank scales. The lateral line is short and may not reach beyond the end of the pectoral fin tip level but can extend back under the dorsal fin level. Esmaeili and Niknejad (2006-2007) give scanning electron micrographs of the scales along with a description. The processus dentiformis is given as absent in the original description but is weakly to well-developed in material seen here. The lips are furrowed, particularly the lower lip. The gas bladder capsules are strongly united. The dorsal fin has 3 unbranched and 7-9, usually 8, branched rays, the anal fin has 3 unbranched and 5-6, usually 5, branched rays, the pectoral fin has 2 unbranched and 9-11 branched rays and the pelvic fin has 1 unbranched and 5-7 branched rays. Total vertebrae 34-35. The pelvic fin insertion is opposite the dorsal fin origin. The caudal fin is very slightly emarginate. The head is wide with large eyes and nostrils just anterior to the eyes. The mouth is arched and the lower lip has large lobules but the mental lobes are reduced. The snout is blunt. Nostrils are widely spaced and equal in size. The gut has a short oesophagus and the stomach and intestine have one loop. Prokofiev (2009) lists various osteological features including the manubrium in the swimbladder being absent or weakly developed and the bony capsule swollen, 4 elongated-cylindrical radial bones in the  pectoral fin, no preethmoid I bone in the skull and 5 hypurals but no epural in the caudal fin support, among others.

Meristics for Iranian specimens: dorsal fin branched rays 7(1), 8(27) or 9(2), anal fin branched rays 5(28) or 6(2), pectoral fin branched rays 9(2), 10(22) or 11(6), pelvic fin branched rays 5(2), 6(18) or 7(10), and total vertebrae 34(11) or 35(10).

Sexual dimorphism

A small slit or groove is found at the antero-ventral eye margin. Sexual dimorphism consists of the anterior 2-3 rays in the pectoral fin being expanded, and pectoral fins having fine tubercles on rays and membranes, extending to all rays except the innermost three. Very fine tubercles are found on the dorsal and lateral head. Male pectoral fins are larger than in females.

Colour

The overall colour is yellowish-white to brown. The head is finely mottled dorsally and laterally. A short bar extends between the eye and snout tip. There is a row of grey to brown or black blotches along the mid-flank and these may line up vertically to form the appearance of thin to broad bar, especially on the rear half of the body. The bars may become weak saddles on the back and keel behind the dorsal fin. The lateral line anteriorly is white. The dorsal and caudal fins may have usually 3-4 rows of spots on the rays and membranes but these do not always line up in neat rows. A fish photographed in Freyhof et al (2011) has about 7 rows on the caudal fin. The dorsal fin may have pigment on the rays and patches of pigment proximally on the membranes. The caudal fin can have to 6 large and darker spots on its dorsal margin and up to 4 spots on the ventral margin, not necessarily lined up with the fin rows. The pectoral and pelvic fins bear extended patches of pigment on the rays and to some extent on the membranes, but not arranged in neat rows. There is an irregular bar at the caudal fin base. The peritoneum is silvery-white with scattered melanophores.

Size

Attains 59.3 mm standard length.

Distribution

Known from Iran in the upper reaches of the Hormuz basin near Darab, in the Kor River basin and in northwestern Iran (Nalbant et al., 2010; Esmaeili et al., 2011). Note that Freyhof et al. (2011), based on collecting expeditions by Hamid Reza Esmaeili, Shiraz University, consider that the Darab (Kol River) locality could be in error and Kol and Kor were confused.

Zoogeography

The limited distribution of this endemic and lack of information on relationships preclude zoogeographical analysis. See family account.

Habitat

Details are unknown.

Age and growth

Unknown. Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 45 fish measuring 2.95-5.93 cm standard length. The a-value was 0.0264 and the b-value 2.904 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases).

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

None.

Conservation

Population trends and numbers unknown.

Further work

More collections are needed to record information on biology and distribution.

Sources

Iranian material: CMNFI 1979-0285, 20, 29.6-49.8 mm standard length, Kermanshahan, Marek River basin (34º26'N, 46º37'E); CMNFI 1979-0286, 1, 51.8 mm standard length, Kermanshahan, Ravansar River at Ravansar (34º43'N, 46º40'E); CMNFI 1979-0292, 8, 29.0-45.6 mm standard length, Fars, Lapu'i spring (29º48'N, 52º39'E); CMNFI 1991-0155, 1, 41.7 mm standard length, Hamadan, Gav Masiab River (34º12'N, 48º20'E); CMNFI 1993-0128, 1, 58.3 mm standard length, Kermanshahan, Sarab-e Sabz 'Ali Khan (34º25'N, 46º32'E); CMNFI 2007-0118B, 17, 23.9-58.0 mm standard length, Kermanshahan, Bid Sorkh River between Sahneh and Kangavar (ca. 34º23'N, ca. 47º52'E).

ID?CMNFI 2007-0115, 1, 41.5 mm standard length, Kermanshahan, Qareh Su basin (ca. 34º34'N, ca. 46º47'E); CMNFI 2007-0115, 2, 49.7-54.7 mm standard length, Kermanshahan, Qareh Su basin (ca. 34º34'N, ca. 46º47'E); CMNFI 1993-0125, 1, 47.8 mm standard length, Kermanshahan, Sarab-e Nilufar (34º24'N, 46º52'E); CMNFI 2008-0236, ?

Genus Paracobitis
Bleeker, 1863

This genus comprises about 17, comparatively large species, some over 28 cm, in western Asia. Six species are reported from Iran but more may exist. One is mentioned here as specimens exist from the Kor River basin in Fars; a record of a Paracobitis from Iraq (see Coad, 2010) in the Tigris River basin is not matched with specimens from adjacent parts of Iran as yet.

The body is elongate, thick anteriorly and posteriorly compressed. The head is strongly depressed. Nostrils are closely spaced or weakly separated. The caudal peduncle is long, low and bears a distinctive, elongate and high crest dorsally from the dorsal fin to the caudal fin and, often, a more or less well-developed ventral adipose crest. However, the crest is not supported by the procurrent rays of the caudal fin unlike in Metaschistura and Oxynoemacheilus. The crest is not connected to the anterior part of the caudal fin (cf. Oxynoemacheilus species, when crest present). The caudal fin is slightly emarginate or truncate, rarely rounded. There is a well-developed pelvic axillary lobe reaching beyond the vertical of the posterior edge of the pelvic fin base. Scales are present or absent anteriorly and the lateral line is complete. The processus dentiformis is usually strongly developed (weak to absent in P. vignai) but its notch on the lower jaw is reduced. Lips are smooth or grooved. The gut is short with a single loop. The colour pattern is variable comprising bars or irregular spotting. There is no intense black spot at the base of the anterior dorsal fin rays. Males and females show no external differences except males have more muscular cheeks. Prokofiev (2009) gives further osteological details of this genus including cst having a bony roof, the fork of cst and cio is fused to the skull in adults (the cst is the supratemporal commissure running across the top of the head at the rear and cio is the temporal part of the infraorbital head canal), the free part of the swimbladder is reduced or absent, right and left lobes of the bony capsule are fully divided by the manubrium, and the fifth trunk vertebra does not take part in support of the capsule. There are 4 radials in the pectoral fin skeleton and the caudal fin support has 6 hypurals and an epural is present. There is no preethmoid I bone in the skull.

Paracobitis boutanensis
(McClelland, 1842)

Probably described from the neighbourhood of the Bolan Pass, Helmand River drainage of Afghanistan in the Sistan basin according to Hora (1929) and Bănărescu and Nalbant (1966). No Iranian record.

Paracobitis ghazniensis
(Bănărescu and Nalbant, 1966)

Described from "Ghazni, on the Ghazni River, tributary of the Ab-i-Istadah Lake, Helmand drainage; East Afghanistan" (Bănărescu and Nalbant, 1966). Ghazni is at 33°33'N, 68°26'E in the Sistan basin. No Iranian record.

Paracobitis iranica
Nalbant and Bianco, 1998

Common names

sagmahi-ye irani.

[Iranian loach].

Systematics

The species is named for Iran. The holotype is 79.8 mm standard length (66.0 mm standard length when measured by me) and is from "River Qom near the town of Qom" collected on 6 May 1976 by P. G. Bianco and stored in the Department of Zoology, University of Naples (IZA 7831). The locality in the jar is "Qom River near Qom (a little salt river 3.5 p.p.t.) near Qom at the bridge, 950 m, 5 June 1976", the date being at variance with the published date. Paratypes number 5 specimens, 59.0-92.0 mm standard length (IZA 7832)(only two present in 2002 visit by me), and 4 specimens, 47.0-72.0 mm standard length (Institutul Stiinte Biologice, Bucharest, ISBB uncatologued). Kottelat (2012) gives a publication date of 1999 but the journal is dated November 1998.

Key characters

This is the only known Paracobitis species in the Namak Lake basin. No single character is uniquely definitive (see below) and locality is key.

Morphology

The body is elongate with a compressed posterior region and a depressed head. The caudal peduncle is elongate and has a long dorsal adipose crest. The body is scaled. The dorsal fin has 7-8 branched rays (90% with 7 rays, n=9 in type description), the anal fin has 5 branched rays in all specimens examined in the type description, the pectoral fin has 9 branched rays in all specimens and the pelvic fin has 5-7 branched rays (78% with 7 rays, n=9). The unbranched dorsal and anal fin rays are often 4, the anterior two minute. The lateral line is straight and extends to the caudal fin base. The body is minutely scaled with a relatively eccentric and large focal zone. The head is conical with eyes in the anterior half. The nostrils are just in front of the eyes. The mouth has a strong arch with short barbels and well-furrowed lips interrupted in the middle. The processus dentiformis is well-developed. The stomach is syphonal and the intestine straight. The gas bladder capsule has a relatively short duct.

Multivariate analyses using 21 morphometric and 6 meristic characters to compare 47 fish from the Namak Lake basin with 19 fish from the Caspian Sea basin showed no unique character or small suite of unique characters that would separate these fishes. When all characters were used, a Discriminant Function Analysis separated 99% of individuals by basin. Nalbant and Bianco (1998) separate this taxon from P. malapterura by body colour pattern and a larger adipose crest. However, the colour pattern figured by them is seen in P. malapterura and Namak Lake basin fish may lack the posterior ventral row of spots or blotches. The adipose crest development is also mirrored in P. malapterura. The only field identification character is collection site.

Meristics for non-type material: branched dorsal fin rays 6(1) or 7(46), branched anal fin rays 4(1) or 5(46), branched pectoral fin rays 8(4), 9(35) or 10(8), branched pelvic fin rays 6(8) or 7(38), predorsal fin vertebrae 13(10) or 14 (37), abdominal vertebrae 25(12), 26(29), 27(5) or 28(1), caudal vertebrae 14(2), 15(25), 16(17) or 17(1), and total vertebrae 40(6), 41(28), 42(11) or 43(2).

Sexual dimorphism

Unknown.

Colour

The overall colour is yellowish-white with the flank and back marbled or with blotches of varying shape, often sub-oval and sometimes coalesced. The blotches extend onto the adipose crest. Blotches are irregularly arranged anteriorly but may line up in upper, mid and lower flank rows posteriorly The lower row may be reduced or absent. The head is covered by dark grey dots. The dorsal, caudal and pectoral fin are strongly pigmented on the rays, sometimes appearing as 3-4 elongate strips lining up as bands, or as almost continuous strips. The anal and pelvic fins lack pigment. There is a wide irregular bar at the caudal fin base.

Size

Attains 92.0 mm standard length.

Distribution

Known from the Namak Lake basin, an Iranian endemic.

Zoogeography

This species may have its origins from its Caspian congener as is seen with other taxa in this family and Cyprinidae.

Habitat

Details are unknown. The type series was collected in a small salt river (3.5‰).

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

None.

Conservation

Population trends and numbers unknown.

Further work

More collections are needed to record information on biology and distribution. Molecular studies would help define its distinction from P. malapterura.

Sources

Type material: The holotype (IZA 7831) and paratypes (IZA 7832) of Paracobitis iranica, see above.

Other material: CMNFI 1979-0252, 1, 53.7 mm standard length, Markazi, jube at Baqerabad (34º55'N, 50º50'E); CMNFI 1979-0253, 1, 58.9 mm standard length, Markazi, Qareh Chay west of Baqerabad (34º52'N, 50º49'E); CMNFI 1980-0154, 8, 45.8-80.8 mm standard length, Markazi, Karaj River below village (35º47'N, 50º58'E); CMNFI 1980-0156, 24, 28.4-63.6 mm standard length, Markazi, Karaj River near village (35º47'N, 50º58'E); CMNFI 1993-0155, 2, 53.4-58.7 mm standard length, Markazi, Sharra River near Khosbijan (34º07'N, 49º23'E); CMNFI 2007-0121, 5, 45.3-98.3 mm standard length, Hamadan, Qareh Su basin north of Razan (ca. 35º25'N, ca. 49º02'E); CMNFI 2008-0152, 1, 71.9 mm standard length, Namak Lake basin (no other locality data); USNM 205921-22, 6, 25.9-34.6 mm standard length, Markazi, Baragon River (ca. 36º00'N, ca. 50º50'E).

Paracobitis longicauda
(Kessler, 1872)

Common names

sagmahi-ye kakoldar-e sharqi (= eastern crested loach).

[vostochnyi grebenchatyi golets or eastern crested loach in Russian].

Systematics

May be a subspecies or synonym of P. malapterura. Has been placed in the genus Adiposia Annandale and Hora, 1920. Banarescu and Nalbant (1964) restrict P. longicauda to basins in Central Asia (e.g. the Amu Darya) and not Iran. Cobitis longicauda was originally described from the Ak-Darya in the Zeravshan River basin of Uzbekistan and the holotype is in the Zoological Institute, St. Petersburg under ZISP 2686 (Eschmeyer et al., 1996).

Bănărescu and Nalbant (1995) place this species in Paracobitis.

Key characters

This species is distinguished from P. malapterura by distinct scales and larger size (including a longer caudal peduncle).

Morphology

Dorsal fin unbranched rays 2-3, branched rays 7, anal fin unbranched rays 2-3, branched rays 5, pectoral fin branched rays 9-10,and pelvic fin branched rays 7. Total vertebrae 42-44. Caudal fin truncate. Large fish are scaled on the flanks, dorsal crest and belly, scales being visible to the naked eye in contrast to the few scales needing a microscope to be visible in P. malapterura and in P. rhadinaea. Caudal peduncle long, about 4.5-4.6 in standard length.

Sexual dimorphism

None reported.

Colour

Colour is very similar to P. malapterura with spots sometimes loop-shaped and sometimes rounded.

Size

Reaches 20.0 cm.

Distribution

Found in the Tedzhen and Murgab rivers of Afghanistan and Turkmenistan and in the Aral Sea basin, and in the Tedzhen (= Hari) River basin of Iran.

Zoogeography

See family account.

Habitat

Unknown.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

 

Unknown.

Economic importance

None.

Conservation

This species is too poorly known in terms of biology and numbers for an effective conservation assessment.

Further work

The biology of this species in Iran needs study.

Sources

Comparative material: ZISP 2686, 1, 140.8 mm standard length, Uzbekistan, Samarkand (); ZISP 4482, 1, 145.7 mm standard length, Uzbekistan, Zeravshan (?); ZISP 4483, 4, 107.2- 125.8 mm standard length, Uzbekistan, Zeravshan (); ZISP 4484, 8, 62.2-96.7 mm standard length, Uzbekistan, Zeravshan (); ZISP 10362, 4, 31.8-112.1 mm standard length, Turkmenistan, Tedzhen River (); ZISP 10963, 1, 93.7 mm standard length, Uzbekistan, Samarkand (); ZISP 13300-13301, 3, 93.7-157.6 mm standard length, Uzbekistan, Pul-e Khatun (?); ZISP 14510, 3, 119.4-173.4 mm standard length, Uzbekistan, Samarkand (); ZISP 33105, 9, 30.4-98.6 mm standard length, ?, Ak-Darya ().

Paracobitis malapterura
(Valenciennes, 1846)

Common names

sagmahi-ye kakoldar-e gharbi (= western crested loach), sagmahi-ye juibari or sagmahi-e-jooibari.

[lakh in Arabic; zapadnyi grebenchatyi golets or western crested loach in Russian].

Systematics

The type locality of Cobitis malapterura is given as "Syrie" in Cuvier and Valenciennes (1846) but has not been found there since (Berg, 1948-1949); it does occur in the Tigris-Euphrates basin however (Coad, 1991b). The specimen was collected by Aucher-Éloy who visited Iran and the specimen may in fact be from there. It is possible that the type locality is in the Caspian Sea basin of Iran although Banarescu and Nalbant (1964) give the Tigris-Euphrates basin which extends though Syria for the distribution of P. malapterura malapterura. Kottelat (2012) notes that the specimens were "sent from Syria" and not necessarily collected there. Bănărescu and Nalbant (1995) places this species Paracobitis.

Material from the Kor River basin may be this species but given that these are south of the Namak Lake basin where a distinct species occurs (P. iranica, q.v.), it seems likely to be distinct. Available material is limited to three specimens with no material for DNA (see below).

Cobitis longicauda Kessler, 1872 is possibly a synonym. Banarescu and Nalbant (1964) restrict P. longicauda (q.v.) to basins in Central Asia (e.g. the Amu Darya) and not Iran. Nemacheilus macmahoni Chaudhuri, 1909 has been advanced as a synonym or subspecies (Nikol'skii, 1947; Berg, 1948-1949; 1949; Banarescu and Nalbant, 1964) but see the review in Bănărescu and Nalbant (1966) and below under P. rhadinaea. Banarescu and Nalbant (1964) consider fish from Sistan and the Caspian Sea basin of Iran to be P. malapterura macmahoni.

Filippi's (1865) record of Cobitis merga (Krynicki, 1840) from "fiumicelli di Sartschem e di Sainkalé" was P. malapterura; these localities being in the Safid River basin near the falling of the Zanjan River into the Qezel Owzan of Safid Rud (Berg, 1948-1949) presumably at Sarcham-e Sofla (37°07'N, 47°54'E) and possibly Sa'in Qaleh (36°18'N, 49°04'E) in the Namak Lake basin.

Two syntypes are in the Muséum national d'Histoire naturelle, Paris under MNHN 3962 and B.3070 (formerly MNHN 3962) (Eschmeyer et al., 1996) and measure 125-145 mm total length (Bertin and Estève, 1948).

Key characters

This is the only Paracobitis in the Caspian Sea basin and locality is key (see above under P. iranica). The caudal peduncle is shorter than in P. longicauda, 5.6-6.3 times in standard length (versus about 4.5-4.6).

Morphology

Dorsal fin with 2-3 unbranched and 6-8 branched rays, anal fin with 2-3 unbranched and 5 branched rays, pectoral fin branched rays 10-12 (Nikol'skii (1947) gives 7-10) and pelvic fin branched rays 5-7. Caudal fin slightly emarginate. There is a well-developed dermal crest or adipose fin behind the dorsal fin to the caudal fin base. Scales are scattered on the posterior body in large fish but need magnification to be seen. The lateral line extends almost to the caudal fin. The dentiform process of the upper jaw is well-developed and fits in a lower jaw groove. The lips, especially the lower one, are strongly plicate. The eyes are small and widely spaced. The adipose crest is of variable height, lower than or similar to that figured in the description of P. iranica. Caudal peduncle short, 5.6-6.3 times in standard length. The gut is straight posteriorly.

Meristics for Iranian material: branched dorsal fin rays 7(19), branched anal fin rays 5(19), branched pectoral fin rays 8(4), 9(35) or 10(8), branched pelvic fin rays 6(6) or 7(13), predorsal fin vertebrae 13(3) or 14 (16), abdominal vertebrae 25(4), 26(14) or 27(1), caudal vertebrae 15(9), 16(9) or 17(1), and total vertebrae 40(1), 41(11), 42(5) or 43(2).

Sexual dimorphism

The cheeks are distended in some fish and this is believed to be a sexual character.

Colour

The top and sides of the body and head are mottled or reticulated with grey and some yellowish pigment. The reticulations may be very fine, giving a more mottled appearance. A reticulated pattern is found in larger fish; young fish have a less reticulated pattern with more blotches on the flank, including in some fish on the lower posterior flank (cf. P. iranica description). The belly and lower head surface are white. The flank reticulations or blotches extend onto the caudal peduncle crest. When touching the dorsal margin of the crest, the reticulations make the crest there dark, otherwise the crest is a light creamy colour along the margin and, in some fish without reticulations reaching the margin, the whole edge is light. The lateral line is white, sometimes in marked contrast to the rest of the flank. The dorsal fin has darkly pigmented rays, sometimes broken into series of spots, but in larger fish the whole ray is pigmented, except distally so the fin margin is white. The caudal fin has a series of 4-5 small spots elongated along the rays, the middle series being the blackest. Large fish have continuous pigment along the rays, except distally so the margin of the fin is white. The dorsal margin of the caudal fin may have 2-4 isolated spots. The pectoral fin has dark pigment along the rays or 2-3 series of small spots. The pelvic and anal fins have 1-2 series of grey spots and the pelvic fin may have only 1-2 spots. Pelvic and anal fins may be immaculate. At the front or along the base of the anal fin there is a broad pigmented band. There may be a dark, broad zig-zag bar at the caudal base, merging dorsally and/or ventrally with flank botches. The barbel bases are all darkly pigmented especially the second pair. The iris is silvery.

Size

Reaches 15.0 cm.

Distribution

This species is found in the Caspian Sea basin of Iran in rivers from the Safid to the Atrak including the Qezel Owzan, Sardab, Haraz, Ramian, Neka, Chalus, Tonekabon, Shahrud, Babol, Tajan, Karasu, Madar Su, Gorgan rivers, Sarchem near the Zanjan-Safid river junction, Bandar Gaz in Gorgan Bay (Laptev, 1934; Nikol'skii, 1947; Kiabi et al., 1994; 1999; Jolodar and Abdoli, 2004; Abdoli and Naderi, 2009; A. Abdoli, pers. comm.).

Note that limited material from the southern Kor River basin (three fish, see below) may represent a distinct species.

Zoogeography

See family account.

Habitat

This species appears to favour deeper water and stronger current than other sympatric loaches.

Age and growth

Tabiee and Abdoli (2005) found a sex ratio of 4:1 (male:female) in the Zarringol River of Golestan Province. Average total length was 59.07 mm for males and 82.42 mm for females. Condition factor was 2.6 for females and 1.97 for males. Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 48 Iranian fish measuring 2.67-7.43 cm standard length. The a-value was 0.0126 and the b-value 2.628 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases). Another study in the Zarringol River by Patimar et al. (2009) found maximum ages of 3+ years for males and 4+ years for females, length-weight relationship was W = 0.020L2.62 for males, 0.002L3.81 for females and 0 .008L3.08 for the population, growth was positively allometric, and sex ratio was 1:1.27 in favour of females (cf. Tabiee and Abdoli, 2005). The difference with Tabiee and Abdoli (2005) who found negative allometric growth was attributed to environmental conditions changing between years of sampling. Golzarianpour et al. (2011) give a length-weight relationship for fish identified as this species from the Jaj River with a (intercept) being 0.007 and b (slope) 3.01 and from the Gharechay River with a (intercept) being 0.001 and b (slope) 2.819.

Food

The Zarringol fish were carnivorous with chironomids making up 65.3% of the diet (Tabiee and Abdoli, 2005) or had a diet dominated by Plecoptera (72.4% and Trichoptera (20.8%) (Patimar et al., 2009).

Reproduction

The Zarringol River examined by Patimar et al. (2009) reproduced in April-June with highest GSI in May. Spawning was prolonged, over about 2 months. Absolute fecundity reached 1180 eggs with a maximum diameter of 2.8 mm.

Parasites and predators

A Diplostomum species is recorded from the muscles of this nemacheilid in the Shirud of Mazandaran (Alghmandi and Dalimi Asi, 2000).

Economic importance

Kiabi et al. (1999) consider this species to be of least concern in the south Caspian Sea basin according to IUCN criteria. Criteria include medium in numbers, habitat destruction, widespread range (75% of water bodies), present in other water bodies in Iran, and present outside the Caspian Sea basin.

Conservation

Little is known of the biology and population numbers so no conservation assessment can be made.

Further work

The systematics of this species and its relatives need further study.

Sources

Iranian material: CMNFI 1979-0481, 1, 77.9 mm standard length, Mazandaran, stream 3 km west of Ghalahleekesh (37º18'30"N, 55º31'E); CMNFI 1979-0486, 18, 12.7-63.2 mm standard length, Mazandaran, stream in Atrak River drainage (37º44'N, 56º18'E); CMNFI 1979-0489, 2, 27.4-45.3 mm standard length, Mazandaran, stream in Atrak River drainage (37º50'N, 55º53'E); CMNFI 1980-0160, 3, 35.4-40.0 m standard length, Caspian Sea basin (no other  locality data); CMNFI 1991-0157, 2, 70.4-73.0 mm standard length, Mazandaran, Ramian River (36º58'N, 55º07'E); CMNFI 1993-0145, 2, 53.4-57.0 mm standard length, Mazandaran, Qareh Su (no other locality data); CMNFI 2008-0109, 2, 83.4-83.9 mm standard length, Mazandaran, Gol-e Ramian 35 km east of Gorgan (37º01`N, 55º09È).

Comparative material: ZISP 25788, 1, 62.2 mm standard length, ?, Sumbar River at Aidere (?).

Paracobitis rhadinaea
(Regan, 1906)

Common names

See genus account. Fowler and Steinitz (1956) refer to a fish from Sistan known locally as mahrmahé (sic, presumably mar mahi, snake fish) and this may refer to this species which has an elongate body.

Systematics

Bănărescu and Nalbant (1995) and Nalbant and Bianco (1998) place this species in Paracobitis. The Catalog of Fishes spells the trivial name as rhadinaeus but Kottelat (2012) notes the name is an adjective.

Nemacheilus macmahoni Chaudhuri, 1909 described from the "affluents of the Helmand" is a synonym according to Bănărescu and Nalbant (1966) who refute the opinions of Nikol'skii (1947) and Berg (1948-1949; 1949) who consider macmahoni to be identical to P. malapterura. Earlier Banarescu and Nalbant (1964) placed fish from Sistan and the Caspian Sea basin of Iran as Nemacheilus malapterurus macmahoni. P. malapterura has both lips strongly furrowed, pelvic fin origin under the dorsal fin origin rather than behind, better developed scales which are also present on mid-flank, and a colour pattern of numerous oblique bands.

Affluents (=headwaters) is an error for effluents (= delta) and the species is not from the upper reaches of the Helmand River basin in Afghanistan as suggested by Kottelat (2012) but from the Helmand delta in Sistan.

P. rhadinaea is distinguished from macmahoni by Annandale and Hora (1920) in having an extremely short posterior diverticulum and minute vesicle in the swimbladder, by the absence of scales, a more elongate body, smaller, narrower and less flattened head, and by differences in the profile of the body.

A syntype of Nemacheilus rhadinæus (ZSI F1240/1) is in the Zoological Survey of India, Calcutta under the name Adiposia rhadinaea and the holotype of Nemacheilus macmahoni (ZSI F1222/1) is also there under the name Adiposia macmahoni (Menon and Yazdani, 1968). Two syntypes listed as Nemacheilus rhadinaeus from "Sistan" are in the Natural History Museum, London (BM(NH) 1905.11.29:28-29, 2, 137.8-209.1 mm standard length).

A specimen in the Zoological Institute, St. Petersburg (ZISP 24413, 76.5 mm standard length) is from "Helmand delta, northwest of Jalalabad, Seistan, XI 1918, Indian Mus. (Dr. Hora)" according to Berg (1949) and could be a syntype of Nemacheilus macmahoni (Eschmeyer's "Catalog of Fishes", downloaded 20 May 2008). However, this taxon was described from a single specimen ? . The jar bears a label on the outside reading "N. malapterurus XI 1918 Indian Museum S. L. Hora Delta of Helmand near Jalabad" and another label reads "Adiposia macmahoni Randa stream 4 mls N.W. of Jalabad Seistan. N. Annandale coll."

Adiposia Annandale and Hora, 1920 (type species Nemachilus macmahoni Chaudhuri, 1909, and including Nemacheilus longicaudus and N. rhadinaeus according to Annandale and Hora (1920)) is a synonym of Paracobitis.

Key characters

The body bears some scales and there are modally 10 branched pectoral fin rays in contrast to P. vignai.

Morphology

macmahoni:- ? check The head is broad and flattened and the small eyes are dorsal. The dorsal profile is slightly convex behind the head but the dorsal and ventral profiles soon become straight and nearly parallel, or thicker in front and tapering behind the dorsal fin. Body depth 5.8 in total length. Caudal fin rounded. Oval scales are found on the posterior part of the body in adults (not in rhadinaea in Regan (1906) and Annandale and Hora (1920)) or scales very small and deciduous all over the body. Nostrils are nearer to the eye than the snout tip. The two anterior pairs of barbels reach back to the nostril level and the posterior pair to the anterior or to middle or to the rear of the eye. The upper lip is minutely tubercular and the lower lip is widely interrupted in the middle. Scale radii are few and widely spaced, on all fields. Material identified originally as macmahoni has the pelvic origin behind that of the dorsal fin origin level, the caudal fin is slightly emarginate, the dorsal fin edge is straight, the anus is some distance in front of the anal fin, a well-developed adipose dorsal ridge runs from the dorsal fin to the caudal fin base, the lateral line is almost complete, scales are restricted to the last third of the body, being small, rounded and far apart, lips are almost smooth, and the intestine has a single loop posteriorly. The description of rhadinaea is short. The snout is longer than the postorbital distance, body depth is 7-10 times in body length, head length 5.0-5.5 times in body length, the mouth cleft extends to below the nostrils, lower lip interrupted medially, outer rostral barbel as long as maxillary barbel reaching back to or beyond nostrils, no scales, dorsal fin origin nearer tip of snout than caudal fin base, caudal fin slightly emarginate, caudal peduncle 2.0-2.75 as long as deep, 5.0-5.3 in length of fish.

Dorsal fin with 2-4 unbranched (first two minute and not readily visible) and 7, rarely 8, branched rays, anal fin with 2-4 unbranched (first two minute) and 5 branched rays, pectoral fin branched rays 9-10, usually 10, and pelvic fin branched rays 6-8, usually 7. Scales are highly deciduous and not always present on old preserved material. The dorsal fin rounded. There is a well-developed post-dorsal fin crest and a slight ventral crest on the caudal peduncle. The pelvic fin origin lies just in front of the mid-point of the dorsal fin base. There is an adipose tissue flap at the pelvic fin base. The anterior nostril is a tube followed immediately by a horizontal slit. The bony upper jaw has a slight protuberance and the lower jaw is curved and not indented.

Meristic values for Iranian specimens including types and macmahoni are:- dorsal fin branched rays 7(33) or 8(1), anal fin branched rays 5(34), pectoral fin branched rays 9(2) or 10(32), pelvic fin branched rays 6(2) or 7(32), and total vertebrae 41(1), 42(1), 43(4), 44(6), 45(6) or 46(2)..

Sexual dimorphism

None reported, the pectoral fin being identical in both sexes in material identified as macmahoni (Bănărescu and Nalbant, 1966).

Colour

Living fish identified as macmahoni are pale olivaceous fading to silvery-white on the belly. The head and upper part of the body are irregularly spotted and darker. Some fish are pale yellowish without markings or with faint marbling. All fins are tinged a dull red, most evidently on the caudal fin, and are obscurely marked with small dark spots. There is a dark band at the caudal base. Preserved material identified originally as macmahoni is whitish with 9-21 irregular, brownish spots along the flank, other spots are present dorsally and smaller ones between the dorsal and lateral rows. There are 3 rows of minute spots on the dorsal and caudal fins and 2 rows on the pectoral fin (Bănărescu and Nalbant, 1966). Types of macmahoni are brown all over, darker dorsally, barbels a lighter brown, dorsal and caudal fins with darker bands, pectoral also slightly banded but anal and caudal uniform light brown. P. rhadinaea has large oblong or rounded dark spots on the back and sides, dorsal and caudal fins with small spots and red tinged, lower fins pale and immaculate although pectoral, and to a lesser extent pelvic, fins may be red tinged.

Size

Attains 28.8 cm as macmahoni.

Distribution

This species is probably restricted to the Sistan basin of Iran and presumably Afghanistan. Bănărescu and Nalbant (1966) place this species in the Atrak and Safid rivers of the Caspian Sea basin, the Abkhar River of central Iran, probably most of Iran, the Helmand drainage and the Tedzhen River, evidently confusing it with P. malapterura and P. iranica. Abdoli (2000) lists as questionably from the Bejestan, Kerman-Na'in and Dasht-e Lut basins, from the middle and lower Halil and Bampur rivers of the Hamun-e Jaz Murian basin, and from the Simish and the river to its north in the Mashkid River basin.

Zoogeography

The closest relatives of this species are P. malapterura and P. longicauda (q.v.) in Iran (Bănărescu and Nalbant, 1966).

Habitat

Annandale and Hora (1920) note that Adiposia macmahoni was healthy buried some inches in mud when cyprinids died in foul water above.

Age and growth

Unknown.

Food

Stomach contents include cyprinid fish remains and mayfly larvae (Annandale, 1921).

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

Unknown.

Conservation

?

Further work

?

Sources

Type material: Syntypes of Nemacheilus rhadinæus (BM(NH) 1909.11.29:28-29), see above.

Iranian material: CMNFI 1979-0222, 11, 16.8-28.8 mm standard length, Sistan, jube 2 km south of Lutak (30º46'30"N, 61º24'E); CMNFI 1979-0223, 1, 23.9 mm standard length, Sistan, ditch 1 km south of Lutak (30º45'N, 61º24'E); CMNFI 1979-0228, 3, 55.1-119.8 mm standard length, Sistan, ditch 1 km from Zabol (31º02'30"N, 61º31'E); CMNFI 1979-0229, 3, 87.4-115.5 mm standard length, Sistan, ditch 5 km from Zabol (31º03'N, 61º33'E); CMNFI 1979-0231, 1, 22.6 mm standard length, Sistan, jube 3 km from Zabol (31º01'N, 61º32'E); CMNFI 1979-0232, 2, 82.5-95.6 mm standard length, Sistan, jube 11 km from Zabol (ca. 30º58'30"N, ca. 61º36'E); CMNFI 1979-0233, 1, 68.7 mm standard length, Sistan, ditch at Deh Vazi (ca. 30º57'N, ca. 61º38'E); CMNFI 1979-0234, 1, 85.5 mm standard length, Sistan, effluent of Hirmand River near Zahak (30º54'N, 61º40'E); CMNFI 1979-0237, 2, 17.5-25.2 mm standard length, Sistan, ditch 18 km south of Zabol (30º53'N, 61º27'30"E); CMNFI 1979-0238, 2, 23.6-27.7 mm standard length, Sistan, ditch 11 km south of Zabol (30º57'N, 61º27'30"E); BM(NH) 1920.1.20:31-34, 4, 79.6-109.5 mm standard length, Sistan, northwest of Jalalabad (no other locality data); ZISP 24413, 1, 76.5 mm standard length, Sistan, Randa stream 4 miles northwest of Jalalabad (?).

Paracobitis smithi
(
Greenwood, 1976)


Dorsal view of head

Ventral view of head

Photograph courtesy of Amir Hosin Zalaghi, 11-19 May 2010

Photograph courtesy of Amir Hosin Zalaghi, 11-19 May 2010

Photograph courtesy of Amir Hosin Zalaghi, 11-19 May 2010

Three photographs above courtesy of Amir Hosin Zalaghi, 11-19 May 2010


Courtesy of Iraj Hashemzade

Common names

sagmahi-ye gharzi (= cave loach), mahi kurghar (= cave fish).

[blind loach].

Systematics

This species is named for Anthony Smith who collected the holotype. Accounts of Smith's searches for cave fishes in Iran are given in his two books (Smith, 1953; 1979). The holotype is in the Natural History Museum, London under BM(NH) 1976.6.28:1, was collected in April 1976, is apparently an immature male and is 35.5 mm standard length. Locality data is given below.

Greenwood (1976) placed this species in the catchall genus Noemacheilus (correctly spelled Nemacheilus) pending an adequate revision of the subfamily Nemacheilinae and since little purpose would be served by erecting a new genus for a fish with such features as eyelessness and depigmentation found in common with other unrelated cave-dwelling fishes. Greenwood (1976) considers its relationship to lie with species in the subgenus or genus Paracobitis as does Nalbant and Bianco (1998). Kottelat (2012) tentatively follows the latter authors.

Key characters

The only eyeless, depigmented nemacheilid reported from Iran.

Morphology

Dorsal fin with 3 unbranched and 7 branched rays, anal fin with 3 unbranched and 5 branched rays, pectoral fin with 10 branched rays and pelvic fin with 5-6 branched rays. In specimens seen by me dorsal fin with 7(5) branched rays, anal fin with 5(5) branched rays, pectoral fin with 10(5) branched rays and pelvic fin with 5(5) branched rays. The caudal fin is unusual in having only 14(5) branched rays. Vertebrae 37-38. check on my fish? The barbels are short with the middle pair the longest. There are no scales. The lateral line is interrupted irregularly and there are more pores on the posterior half of the body. A long adipose fin is present dorsally, most obvious in young, and a weaker ventral ridge is present. The mouth is subterminal, lips are weakly to moderately papillose, and there is a horny ridge on the dentaries and on the upper jaw. The middle pair of barbels is the longest. The internarial flap is long. Gill membranes are broadly attached to the isthmus. The gut is short with a single transverse loop.

Sexual dimorphism

Unknown.

Colour

A dead white with the red of blood visible as a pale pink cast and a deeper red at the gills visible through the gill cover. Small, straw-coloured fat globules are visible under the skin of formalin preserved fish over the whole head and body with concentrations in the orbits and the fin bases, particularly the dorsal and anal bases. The viscera are visible through the body wall. There is no peritoneal pigment.

Size

Attains 64.5 mm total length.

Distribution

Found only at "Kaaje-ru" above the garden "Bagh-e Loveh", "Lowa" or "Levan" (probably Loven at 33°04'N, 48°37'E) which is about 4 km from kilometre 382 on the railway from Bandar Shapur and approximately 12 km north of the railway station Tang-e Haft. The stream below the cave locality is the "Ab-e Serum" which runs into the "Ab-e Zezar" which is a tributary of the Dez River. The locality is at 33°04'38.6"N, 48°35'33.1"E in Lorestan Province. Further locality details are given in Bruun and Kaiser (1948) and under the cyprinid Iranocypris typhlops.

Zoogeography

Endemic to Iran, its relationships to other species are unknown. It shares the cave habitat with another eyeless species, Iranocypris typhlops, a member of the family Cyprinidae. This blind cave species is placed in a world-wide context by Proudlove (1997a; 1997b).

Habitat

Known only from a well-like but natural outlet of a subterranean system. The outlet overflows to form a small stream from January to May (Smith, 1979) during the snow-melt period in the Zagros Mountains but in April to June this flow ceases (the precise timing of flow and its cessation is estimated from villager's comments and scientific visits and also varies with precipitation). The pictures above show flowing water in May 2010. The well area is about 5 by 3 m and gradually decreases as the year progresses. Divers descended to a depth of 60 feet (= 18.3 m) in 1977 in the "well" until the resurgence narrowed (Farr, 1977). A rope was let down by R. Mehrani (pers. comm., 2000) and reached 23 m before the rope ran out and yet it was not at the bottom. Smith (1979) reports divers descending to 60-70 feet (18.3- 21.3 m). The pool shelves deeply under the cliff rearwards but the whole pool surface is exposed to light. There is no vegetation in the pool except for some encrusting algae on the rocky sides. The shale fragments forming the outermost floor of the pool have a thin layer of mud on them which may contain algae.

It seems probable that a complex of flooded but narrow and inaccessible passages is the habitat of this species and the well is merely the surface manifestation of this complex (Bruun and Kaiser, 1948; Smith, 1978; Banister, 1992). There is a smaller pool (about 2 m across narrowing rapidly inside) and flowing exit stream lower down the gorge, about 50 m away from the main locality, where an Iranocypris typhlops was seen but not caught in December 2000 (Smith (1979) also tentatively reports sighting a fish here). This is assumed to be evidence of the interconnectivity of subterranean passages. The main pool was not flowing at this time. The stream from the smaller pool increases in flow downstream, possibly tapping more groundwater, and eventually has a moderate flow. No fish were seen in it. The stream falls over a high waterfall (estimated at 10-15 m high by Smith (1979) which seems about right) so the well localities are isolated from the local fishes in the main river. The main river houses Garra rufa and Nemacheilus species s.l.. The stream shows evidence of recent higher flow which tends to confirm overflow from the main well.

Sampling in December 2000 recorded a water temperature of 18.5°C, pH 7.5 and a conductivity of 334 µS. More photographs of the habitat can be seen under the account of Iranocypris typhlops (Cyprinidae).

A captive specimen attempted to climb out of a glass tank, almost its whole body being out of the water before it slid back. It did not hide from or react to light and spent most of the time resting on the tank bottom, moving along the bottom or more often swimming actively around the tank (Greenwood, 1976). Amir Hosin Zalaghi reports that this species swims faster than Iranocypis tyhplops (pers. comm., 10 August 2010).

 

Photograph courtesy of Amir Hosin Zalaghi, 11-19 May 2010 
 Photograph courtesy of Amir Hosin Zalaghi, 11-19 May 2010

Two photographs above courtesy of Amir Hosin Zalaghi, 11-19 May 2010

Age and growth

Unknown.

Food

Unknown but a captive specimen was fed on mosquito larvae.

Reproduction

Unknown.

Parasites and predators

Unknown but there are probably no predators in the cave environment.

Economic importance

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in textbooks.

Conservation

A fine of 10,000 rials is imposed specifically for illegal fishing of this species (Anonymous, 1977-1978), more recently 100,000 rials (N. Najfpour pers. comm., 2008). It is on the IUCN 1994 Red List of Threatened Animals as one of two rare fish species from Iran (see also Iranocypris typhlops) and is on the 2000 IUCN Red List as VU D2 (Vulnerable, acute restriction in its area of occupancy, also on subsequent Red Lists; see also Proudlove (2001)). Coad (2000a), using 18 criteria, found this species to be one of the top 4 threatened species of freshwater fishes in Iran.

B. Sandford (in litt., 1979) considered this fish to be endangered. The cave appeared to be a recently collapsed system and the network of fissures could be quite small. Coupled with recent collecting the number of extant specimens may be quite low.

This species is much rarer than the co-occurring Iranocypris typhlops, by at least an order of magnitude. Including the holotype, about 14 specimens are known to have been collected (see below; plus 2 in Muze-ye Melli-ye Tarikh-e Tabi'i, Tehran (MMTT 1227-1228) and 6 collected by students of A. Abdoli, Shahid Beheshti University, Tehran in 2001).

Further work

?

Sources

Type material: Holotype of Noemacheilus smithi (BM(NH) 1976.6.28:1), see above.

Iranian material: CMNFI 2007-0123, 5, 24.1-52.9 mm standard length, type locality, 28 January 1977.

Further information on the habitat is in the account of Iranocypris typhlops.

Paracobitis vignai
Nalbant and Bianco, 1998

Sistan River, Zahak, January 2012, courtesy of K. Abbasi
Sistan River, Zahak, January 2012, courtesy of K. Abbasi

Common names

sagmahi-ye Sistan.

[Sistan loach].

Systematics

The species is named for Professor Augusto Vigna Taglianti, La Sapienza University, Rome. The holotype is 89.0 mm standard length (86.5 mm standard length when measured by me) collected from "Nahr Taheri, Zabol, Seistan" on 9 October 1977 by A. Vigna and is deposited in the (Department of Zoology, University of Naples (IZA 7838). Paratypes are from the same locality and number 24 specimens, 35.0-78.0 mm standard length (IZA 7839) and 7 specimens, 44.0-49.0 mm standard length (Institutul Stiinte Biologice, Bucharest, ISBB uncatalogued). One specimen is in the American Museum of Natural History (AMNH 40946), presumably a paratype from one of the preceding series and 3 specimens are in the Canadian Museum of Nature, Ottawa under CMNFI 2012-0007 (paratypes from IZA 7839, 39.0-63.4 mm standard length). Kottelat (2012) gives a publication date of 1999 but the journal is dated November 1998.

Key characters

This species is scaleless and modally has 9 branched pectoral fin rays in contrast to P. rhadinaea with some scales and modally 10 pectoral rays.

Morphology

Dorsal fin branched rays 6-8 (80% with 7 rays in original description, n = 20), anal fin with 5-6 branched rays (95% with 5 rays), pectoral fin with 8-10 branched rays (85% with 9 rays) and pelvic fin with 6-7 branched rays (95% with 7 branched rays). Total vertebrae number 44-45.The lateral line extends to the base of the caudal fin. The body is slender and compressed, particularly posteriorly. The head is long and the eyes are small. The dorsal fin origin is at mid-body (snout tip to caudal fin base). The mouth is arched with strongly furrowed lips that have few papillae. Mental lobes are reduced. The caudal fin is deeply forked. The dorsal adipose crest or keel on the caudal peduncle is well-developed. The stomach is syphonal and the intestine is straight. The anus is placed well anterior to the anal fin origin. The gas bladder capsule has globular chambers and a short duct.

Sexual dimorphism

Unknown.

Colour

The head and body are greyish, lighter ventrally. Dark spots along the back and flank may be occasionally fused. Fine spots are scattered in a reticulated pattern on the body. The dorsal and caudal fins have dots in rows while other fins are colourless. The caudal fin base has a distinct blackish bar.

Size

Reaches 89.0 mm standard length.

Distribution

Endemic to Sistan.

Zoogeography

Nalbant and Bianco (1998) consider this species to be an epigean form of the blind cave fish P. smithi. The differences are in head shape and mouth and lip morphology, apart from the eyes and pigment loss typical of ipogean P. smithi.

Habitat

Details are unknown.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

None.

Conservation

Population trends and numbers unknown.

Further work

More collections are needed to record information on biology and distribution. The distinction of another Paracobitis species in Sistan, an area of interconnected waterways through periodic flooding, could be questioned. The chief distinguishing features from P. rhadinaea are the absence of scales (a somewhat variable character in respect of presence and the scales are often deciduous) and a modal count of 9 pectoral fin branched rays (some rhadinaea have 9 rays). Fresh material for molecular studies would help re-affirm this taxon. P. ghazniensis from higher in the Helmand River basin is distinguishable on colour pattern (flank bars), pelvic fin rays (6 as opposed to 7 in rhadinaea and vignai), caudal peduncle length (mostly 6.4-7.8 in standard length, compared to 5.4-6.2 in rhadinaea and vignai), and is in a water body isolated at times from the main Helmand River basin.

Sources

Type material: The holotype (IZA 7838) and paratypes (IZA 7839) of Paracobitis vignai, see above.

Paracobitis sp.

Common names

None.

Systematics

Paracobitis species are known and described from the Caspian Sea and Namak Lake basins, far to the north of specimens found in the Kor River basin of Fars. The latter material represents a new taxon, as yet undescribed. Multivariatr analyses separate the few Kor River specimens from Caspian and Namak fish but with no single character or small suite of characters diagnostic and useable in a key.

Key characters

Distribution is the key character.

Morphology

Dorsal fin branched rays 7(3), anal fin branched rays 5(3), pectoral fin branched rays 9(1), 10(1) or 11(1), pelvic fin branched rays 5(3), predorsal fin vertebrae 14 (2), abdominal vertebrae 26(1) or 27(1), caudal vertebrae 17(2), and total vertebrae 43(1) or 44(1). 

Sexual dimorphism

Unknown.

Colour

Two specimens are faded but the material has evidence of blotches typically found in Paracobitis species, extending onto the adipose crest, an irregular and broad caudal fin base bar, pigment on the rays of the dorsal and caudal fins, and blotches extending to the lower flank posteriorly.

Size

Reaches 81.5 mm standard length, possibly larger as only three specimens were caught.

Distribution

This species is found in the Kor River basin of Fars Province, southern Iran. Localities include the Kor River proper and a tributary stream.

Zoogeography

The relationships of this taxon could lie to the north in the Namak and Caspian basins or possibly to the west in the Tigris-Euphrates basin. Material from the latter was not available for study but other taxa from the Kor River basin show affinities with Mesopotamian ones.

Habitat

Unknown in detail. Two fish were collected from a qanat stream at 21°C where the bottom was gravel and mud with encrusting and submergent plants.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

This species is too small and rare to be of direct economic importance.

Conservation

Since this species is known only from a limited number of specimens, further studies are needed to ascertain its abundance and distribution.

Further work

This taxon needs diagnosis and description. Molecular studies would help define its distinction from P. malapterura.Another Paracobitis species may well exist in the Tigris-Euphrates basin (Coad, 2010) but no material is available - it would throw light on the distinction of the Kor River population. More collections are needed to record information on biology and distribution.

Source

Iranian material: CMNFI 1977-0510A, 2, 74.2-81.5 mm standard length, Fars, stream in front of Naqsh-e Rostam (29°59`30`N, 52°54È); CMNFI 2008-0255, 1, 51.0 mm standard length, Fars, Kor River (30°00`N, 52°44`58È).

Genus Paraschistura
Prokofiev, 2009

This genus is recently described and not all species have been examined and ascribed to it or related genera. Many were formerly included in the genus Schistura McClelland, 1838.

The species in this genus are small and are distributed from interior water bodies of Turkmenistan and from Iranian Baluchestan east to the upper reaches of the Indus River in Afghanistan and Pakistan. Four species are reported from Iran.

The genus is characterised in particular by a dark black spot or strip always present at the base of the anterior dorsal fin rays, found only in one other genus in Iran, Metaschistura (which in contrast has a well-developed caudal peduncle crest). Note that other genera may have a dark brown blotch or part of a saddle on the anterior dorsal fin base but this is never the deep black spot of this genus.

The genus is also characterised by an elongate body with dorsal and ventral profiles almost parallel. The belly is not rounded. The body is compressed anteriorly and the head is compressed or depressed. The caudal peduncle is short and moderately deep. The snout is usually blunt. The upper lip is furrowed and is continuous or has a slight median interruption. Lips are simple, being plicate to almost smooth. The processus dentiformis on the upper jaw is well-developed in many species and there is a corresponding notch on the lower jaw but the processus can be absent. Nostrils are closely spaced. The dorsal fin usually has 7 branched rays and the anal fin 5 branched rays. The caudal fin is slightly to deeply emarginated. There is no adipose crest or, if present, it is weakly-developed near the caudal fin and not supported by procurrent rays of the caudal fin. The lateral line is incomplete, may be very short and rarely reaches the end of the anal fin level. Scales are absent or rarely present but weakly developed on the caudal peduncle. A pelvic axillary lobe is present but is short and does not usually extend beyond the pelvic fin base.

Prokofiev (2009) lists various osteological characters that help diagnose the genus including the absence of a bony bridge between the parietal and pterotic bones which separates the species from Schistura sensu lato. There is no preethmoid I bone in the skull. Externally, there is no bony roof to cst, the fork of cst-cio is not ossified and it is not fused to the skull. The cst is the supratemporal commissure running across the top of the head at the rear and cio is the temporal part of the infraorbital head canal. The free part of the swimbladder is absent, the right and left lobes of its bony capsule are fully divided by the manubrium, and the fifth trunk vertebra does not take part in support of the capsule.

There is no sexual dimorphism. Colour pattern is distinctive with dark bars of varying number and form being present. The bar at the caudal fin base is usually much darker than anterior bars, and is never in the form of a spot. There is no stripe along the flank although a dark line is rarely present.

Paraschistura alta
(Nalbant and Bianco, 1998)

Described from "Afghanistan, Kajkai, Helmand river drainage, north east of Girisk". No Iranian record. Kottelat (2012) gives a publication date of 1999 but the journal is dated November 1998, and he retains this species in Schistura.

Paraschistura baluchiorum
(Zugmayer, 1912)

Possibly a synonym of P. bampurensis according to Nalbant and Bianco (1998). The type locality is Panjgur on the Rakhshan River of the Hamun-i Mashkel basin in Pakistani Baluchistan and it is also recorded from Afghanistan in the Helmand River drainage. A cotype is in the Naturhistorisches Museum Wien under NMW 19851 and is 46.0 mm standard length. Not recorded from Iran by specimens although Abdoli (2000) reports it from the Simish River and a river just north of it, both in the Mashkel basin. Prokofiev (2009) places this species in Schistura.

Paraschistura bampurensis
(Nikol'skii, 1899)

Common names

sagmahi-ye Bampur.

[Bampur loach]

Systematics

Publication date is given as 1900 in Berg (1949), Bănărescu and Nalbant (1966) and Kottelat (2012). This may be correct if the volume appeared late as the volume of the publication is for the year 1899; but note that a footnote and the plate both bear the year 1899.

Nemacheilus baluchiorum Zugmayer, 1912 is possibly a synonym. (Berg (1949) places this species in synonymy with Nemacheilus montanus - see below). Bănărescu and Nalbant (1966) and Nalbant and Bianco (1998) place S. bampurensis in the subgenus or genus Schistura but Kottelat (2012) has it in Paraschistura.

Berg (1949) placed P. bampurensis in Nemacheilus montanus (McClelland, 1839) but Bănărescu and Nalbant (1966) recognise bampurensis as distinct on its incomplete lateral line, focal zone of scales larger, dorsal fin positioned more anteriorly, deeper body, and colour pattern without regular bands (Bănărescu and Nalbant (1966: fig 12) for three pattern varieties). Observations were made by me on the faded syntypes of N. montanus from Simla, 62.8-68.7 mm standard length (BM(NH) 1860.3.19:118-119) as well as ZISP 8298, 9 specimens 25.3-51.1 mm standard length. Additional characters which distinguish bampurensis and montanus are as follows. The preorbital process is very strongly developed in N. montanus, being almost as deep as the eye and extending almost an eye length below the lower orbit margin, extending almost twice the distance of the process in bampurensis types and being less curved and close under the eye. N. montanus has a slight but obvious keel on the back before the caudal fin while bampurensis are humped there but usually not keeled. The caudal fin is dark at the base in montanus, not so in bampurensis or not as solid, wide and dark. The flank bars are oblique with the top forward in contrast to bampurensis, and are less numerous than in bampurensis (5-10 versus 11-22 total, from under dorsal fin insertion to caudal fin but excluding any caudal fin base bar). N. montanus types have faded bars but are clearly fewer.

The syntypes of Nemacheilus bampurensis are in the Zoological Institute, St. Petersburg under catalogue numbers ZISP 11698 and 11699 at the localities, dates and number of specimens in Latin as follows respectively:- "Kjagur prope urb. Bazman. 4. VII (6)" and "Kaskin prope urb. Bazman. 6.VII (4)" (Nikol'skii, 1899). Berg (1949) gives both these localities as between Bazman and Bampur. However, ZISP 11698 comprises 9 specimens, 35.1-44.6 mm standard length and ZISP 11699 comprises 4 specimens not listed as types on the jar, 36.8-44.5 mm standard length.

Key characters

The male has a characteristic, moveable protuberance directed downward on the preorbital bone at the antero-ventral corner of the eye. It lies close to the eye, extends slightly below the lower orbit, and curves partly around the orbit.

Morphology

Dorsal fin with 2-4 unbranched rays (anteriormost embedded in flesh and difficult to see) and 6-7 branched rays, anal fin with 2-3 unbranched and 4-5 branched rays, pelvic fin branched rays 6-7, and total vertebrae 34-36. Lateral line incomplete, ending in front of the dorsal fin level. Scales well-developed but the anterior third to a quarter of the body is scaleless. The posterior nostril is ovoid, slanting postero-dorsally. Barbels are large and the third pair is usually the largest although in some fish the second pair is the largest (Saadati, 1977). The lower lip is divided, lips are corrugated, and the upper jaw process is rounded, overlapping the lower one. There is a fleshy pelvic axillary process. Caudal fin slightly emarginate.

The type series (ZISP 11698) has dorsal fin branched rays 6(3) or 7(6); anal fin branched rays 4(1) or 5(8); pectoral fin branched rays 9(8) or 10(1); pelvic fin branched rays 6(2) or 7(7); and flank bars 14(2), 15(3), 17(1), 19(2) or 22(1).

Meristic counts for Iranian material:- Dorsal fin branched rays 6(3) or 7(73), anal fin branched rays 4(1) or usually 5, pectoral fin branched rays 8(2), 9(59) or 10 (10), pelvic fin branched rays 6(7), 7(51) or 8(2), total vertebrae 34(19), 35(17) or 36(4), and flank bars 10(1), 11(7), 12(6), 13(12), 14(6), 15(8), 16(3), 17(1), 18(2), 19(2) and 22(1).

Sexual dimorphism

See above under Key characters. Males also have tubercles on the pectoral fin rays and on the operculum (see figure in Berg (1949)). The first branched pectoral fin ray is greatly expanded, 2-3 times broader than the second ray, which itself may be expanded a little.

Colour

Body yellowish to a light olive-green with 9-18 dark, brownish or chestnut-brown bands. The caudal fin has 3-4 dark wavy bars and a bar at its base, the dorsal fin 2-3 dark bars and a black spot at the anterior fin base. Fins are a light orange or pinkish, particularly the caudal fin.

Size

Reaches 5.3 cm. Golzarianpour et al. (2011) gives a total length 6.7 cm (but see below under Age and growth).

Distribution

Berg (1949) and Abdoli (2000) record this species from the Dasht-e Lut basin without specific localities, questionably from the Tigris River basin (presumably based on Bănărescu and Nalbant (1966), see below), the middle and lower Bampur and Halil rivers of the Hamun-e Jaz Murian basin, and the Sarbaz and Nikshahr rivers of the eastern Makran. Specific localities include Kjagur and Kaskin near Bazman (Nikol'skii, 1899), both between Bazman and Bampur (Berg, 1949), Shur Ab near Kuh-i Birk (Berg (1949) and the Bampur River near Bazman.

Bănărescu and Nalbant (1966) report this species from Shapur, 12 km north of Kazerun and from Shah Bazan on a tributary of the Ab-i-Diz, and "probably most of Iran". This seemed inherently unlikely as the type locality is in Baluchestan and the material was later described as a new species, P. nielseni q.v.

Zoogeography

See family account.

Habitat

Unknown.

Age and growth

Unknown. Golzarianpour et al. (2011) give a length-weight relationship for fish from the Maroon River with a (intercept) being 0.008 and b (slope) 3.11. Note that if this is the Marun River in the northern Persian Gulf basin, then the species is probably not the P. bampurensis of Baluchestan.

Food

Unknown.

Reproduction

A female measuring 4.8 cm and caught in late February carried fairly well-developed eggs (Berg, 1949).

Parasites and predators

Unknown.

Econmic importance

None.

Conservation

This species is reported from several localities in Baluchestan, including those remote from human influence, and does not seem to be in any danger.

Further work

The biology of this species needs to be studied.

Sources

Type material: Syntypes of Nemacheilus bampurensis (ZISP 11698 and 11699), see above.

Iranian material: CMNFI 1979-0312, 14, 24.9-41.5 mm standard length, Baluchestan, Bampur River 8 km west of Iranshahr (27º11'N, 60º36'E); CMNFI 1979-0313, 5, 31.1-39.9 mm standard length, Baluchestan, Bampur River at Bangharabad (27º20'N, 60º46'E); CMNFI 1979-0315, 17, 20.0-44.8 mm standard length, Baluchestan, Bampur River 2 km north of Karevandar (27º51'N, 60º46'E); CMNFI 1979-0317, 28, 19.9-45.8 mm standard length, Baluchestan, Sarbaz River at Bondan (26º35'N, 61º13'E); CMNFI 1979-0318, 1, 23.8 mm standard length, Baluchestan, Sarbaz River at Huvar (26º09'N, 61º27'E); CMNFI 1979-0323, 5, 20.7-31.4 mm standard length, Baluchestan, Sarbaz River (ca. 26º26'N, ca. 61º16'E); CMNFI 1979-0326, 12, 19.3-35.0 mm standard length, Baluchestan, stream in Oghin River drainage (ca. 26º35'N, ca. 60º02'E); CMNFI 1979-0327, 10, 21.2-37.4 mm standard length, Baluchestan, stream in Nikshahr River drainage (26º32'N, 59º57'E); CMNFI 1979-0329, 14, 21.1-33.4 mm standard length, Baluchestan, stream at Zaminbandan (27º02'N, 61º20'E).

Paraschistura chrysicristinae
(Nalbant, 1998)

Described from the Tigris River basin in Turkey but no Iranian record.

Paraschistura kessleri
(
Günther, 1889)

Common names

sagmahi-ye Bejestan, sagmahi-ye Kessler.

[golets kesslera or Kessler's loach in Russian; sundali in Pakistan].

Systematics

Nemacheilus prashari Hora, 1933 is possibly a synonym, along with its subspecies (q.v.). Banarescu and Mirza (1965) have comparisons of Nemacheilus lindbergi (which is later regarded as a subspecies of Nemacheilus (= Parachistura) prashari, q.v.) with P. kessleri where most characters overlap in varying degrees. Bănărescu and Nalbant (1966) place P. kessleri in the subgenus Schistura and Nalbant and Bianco (1998) place it in the genus Schistura. Berg (1949) refers P. prashari to a subspecies of P. kessleri. Kottelat (2012) has this species and P. prashari as distinct taxa.

Nemacheilus kessleri turcomanus Nikol'skii, 1947 is reported from Turkmenistan east of the Iranian border in the Murgab River drainage (Kushk River near Kushk). Bănărescu and Nalbant (1966) consider that the differences are too slight to warrant a separate subspecies.

Berg (1948-1949) considered P. kessleri to be close to P. sargadensis, differing in having the snout slanting steeply in front of the eyes (rounded in sargadensis - this is generally true but exceptions occur with sharply downturned snouts in sargadensis examined by me), dorsal origin midway between the snout tip and the caudal fin base (further back in sargadensis - but see below, same in both species; and in a series of sargadensis of varying size small fish have a longer predorsal distance than dorsal origin to caudal base distance, in large fish a shorter predorsal distance, and in intermediate-sized fish an about equal predorsal distance, so this character appears to be highly variable and unreliable for separating these species), rudimentary caudal peduncle crest (weak to clearly pronounced crest - ?check on my sargadensis), fewer and less sharply pronounced flank bars (10-12 total, broad bars in kessleri seen by me while sargadensis has 13 or more thinner bars - ?check on my sargadensis), longer intestine (?), and pelvic fin origin just behind the level of the dorsal fin origin (under dorsal fin origin or slightly in advance - kessleri types seen by me had the pelvic fin origin almost exactly under the dorsal fin origin). Generally P. sargadensis agrees with the description of P. kessleri except for pigmentation (?I need to run type measurements I have against sargadensis noting that Ghazni fish may be a different species and comparing gut length/shape and bladder capsule (see Mirza et al. (1981)).

Nemacheilus kessleri was described from Nushki in the Pishin Lora River basin of Afghanistan (Menon, 1987). Four syntypes are in the Natural History Museum, London (BM(NH) 1886.9.21:177-180, 38.3-48.4 mm standard length) and 4 syntypes (now 3) are in the Zoological Survey of India, Calcutta (ZSI 11487-11490) (Eschmeyer et al., 1996). The holotype of Nemacheilus kessleri turcomanus is in the Zoological Museum of Moscow State University (MMSU P.5734) with apparently a paratype under MMSU P.5734 and another paratype under MMSU P.5735 (Eschmeyer et al., 1996). The Zoological Museum of Moscow University (ZMMU; their acronym) has 3 specimens under P-5734 and 1 specimen under P-5735, all syntypes (Pavlinov and Borissenko, 2001).

Key characters

Characterised by Menon (1987) as a member of the subgenus Schistura with 7 branched dorsal fin rays, no scales, forked caudal fin, short lateral line not reaching mid-body and no sexual dimorphism, and additionally by and Mirza et al. (1981) and Bănărescu and Nalbant (1995) as a member of the genus Schistura part of the kessleri-lindbergi group with the above characters and also small size, rather compressed head, emarginate or slightly forked caudal fin and a reduced dentiform process.

Morphology

Dorsal fin unbranched rays 2-4 (anteriormost rays embedded in flesh and hard to distinguish), branched rays 7, anal fin unbranched rays 2-3, branched rays 5, pectoral fin branched rays 8-11, pelvic fin branched rays 6-7, usually 7, and total vertebrae 37-39. Dorsal fin origin midway between the nostrils and the caudal fin base according to Berg (1948-1949) but the types examined by me clearly have the dorsal fin origin closer to the snout (predorsal distance when stepped back from dorsal fin origin overlaps the rays of the caudal fin). Scales are absent. The lateral line ends under the dorsal fin (above the tip of the pectoral fin in Menon (1987) ? check on fish, Menon quote is correct). No adipose fin but a rudimentary crest may be evident. Caudal fin slightly emarginate to quite deeply forked. Pelvic fin origin somewhat behind the level of the dorsal fin origin. The snout is blunt; in the types it falls gently at the nostrils in 3 fish, abruptly in 1 fish; and in 3 fish from Ghazni, Afghanistan abruptly in 2 fish and less so in the third, all apparently independent of size. Both lips are fringed, the upper with a slight interruption, the lower with a wide interruption. The dentiform process is reduced (Berg (1949), well-developed in Menon (1987) reduced in Ban and Nal 1995 ? check on fish). Barbels are variably developed or absent in some individuals, a condition illustrated by Hora (1933). The posterior part of the intestine has a single loop.

Sexual dimorphism

There is no sexual dimorphism.

Colour

The back and flank are crossed by 10-14 irregular, brownish bands, becoming less band-like and more irregular on the caudal peduncle. There is a dark spot at the base of the 3 anteriormost dorsal fin rays. There are 1-2 bands or rows composed of small spots on the distal part of the dorsal fin, or the rows may not be clearly defined with pigment in elongate patches along rays and the fin margin clear. The anal fin may have an anterior basal spot. There is a narrow blackish band at the caudal fin base and 2-3 narrower bands on the fin itself. The sides of the head are minutely spotted. Bănărescu and Nalbant (1966) consider that the fish reported by Berg (1949: fig. 54) from Kelat Margh in Neh-i-Bendan do not have the typical colour pattern of this species. Berg (1949) describes 6-8 chestnut-brown bars on the posterior part of the body, none anteriorly.

Size

Reaches 8.7 cm total length (Patimar et al., 2010).

Distribution

Found from the Lora River basin of Pakistan through Afghanistan to the Murgab River basin of Turkmenistan and in eastern Iran at such localities as Keljate-Marg in Zirckuch region and the Neh-i Bendan district (?Berg check localities for modern spelling). For Iran, Abdoli (2000) lists middle and upper Kashaf River of the Tedzhen River basin, questionably in the Bejestan basin, and in the Simish and Mashkid rivers, and river north of them in the Mashkid River basin. Nalbant and Bianco (1998) also cite the Mashkel Lake, northern Sistan.

Zoogeography

See family account.

Habitat

Unknown.

Age and growth

Patimar et al. (2010) examined fish from the Zanglanlou River in northeast Iran  and found maximum age for both sexes was 4+ years. Growth was positively allometric and sex ratio was 1:1.2 in favour of females.

Food

Unknown.

Reproduction

Patimar et al. (2010) found reproduction occurred in April-May with the highest GSI in April. Maximum egg diameter was 1.46 mm, aboslute fecundity reached 1246 eggs and up to 1285.71 eggs in relation to total weight.

Parasites and predators

Unknown.

Economic importance

None.

Conservation

?

Further work

?

Sources

Type material: Syntypes of Nemacheilus kessleri (BM(NH) 1886.9.21:177-180), see above.

Comparative material: BM(NH) 1944.4.1:7-9, 3, 42.0-57.6 mm standard length, Afghanistan, Ghazni (ca. 33°33’N, 68°26’E).

Paraschistura lindbergi
(Banarescu and Mirza, 1965)

Found in the Mashkel (= Mashkid) River basin of Pakistan on the southeastern border of Iran (see P. prashari for further discussion). This species was described from a rivulet connected to the drainage of the Farah River, Siaw, between Farah and Dilaram, Afghanistan. No Iranian record although Nalbant and Bianco (1998) record a specimen from near the Iranian border in the "Mashkel Lake system". Originally described in the genus Noemacheilus.

Paraschistura nielseni
(Nalbant and Bianco, 1998)

Common names

sagmahi-ye Nielsen.

[Nielsen's loach].

Systematics

Note that P 27110 (33.7 mm SL, my measurement here and below) "Shah Bazan, W. Iran, 16. IV. 1937. E. Kaiser" is listed as the holotype in the Zoological Museum, University of Copenhagen (ZMUC), not 27109 as in the original, published description (observed in 1999). The Shah Bazar locality in the description is presumably a misprint for Shah Bazan. Paratypes are 27109 (25.8 mm SL) from "Shah Bazan..." etc as above (probably reversed with the holotype), and 27105 "lok 44" (30.7 mm SL), 27106 "lok 54" (12.7 mm SL), 27107 "lok 54" (29.5 mm SL), 27117 "lok 54" (37.3 mm SL) from "Shapur, 15.03.1937. E. Kaiser. W. Iran". The published description gives the locality of Shapur as "12 km north-west of Kazerun".

The size of one paratype (12.7 mm) is smaller than any cited in the type description. The catalogue number of one fish in a drawing in the original description (i.e. 27108) is not in that description nor in jars in ZMUC. There has evidently been some confusion over the type series.

Named for Dr. Jorgen G. Nielsen, Department of Ichthyology, Zoological Museum, University of Copenhagen. Kottelat (2012) gives a publication date of 1999 but the journal is dated November 1998.

This species was previously reported as Nemacheilus bampurensis by Bănărescu and Nalbant (1966) based on this type series. The dorsal fin spot is reported as absent or present in this paper and the type description respectively (see below). Presence of this spot is a character of Paraschistura species and needs to be clarified.

Key characters

This species has a relatively short body with a short head and blunt snout. The lateral line is incomplete ending just anterior to the dorsal fin origin. Scales have a large, eccentric focus. The caudal fin is distinctly emarginate. The body is yellowish with 7-17 dusky brown bars. The dorsal fin has 7 branched rays.

Morphology

Meristics from type series:- dorsal fin branched rays 7(5), anal fin branched rays 5(5), pectoral fin branched rays 9(4), 10(1), pelvic fin branched rays 6(1), 7(4), and vertebrae 33(1), 34(4), 35(1). The body is compressed, especially posteriorly. The eyes are small and positioned in the middle or anterior portion of the head. The mouth is slightly arched and the upper jaw has a reduced processus dentiformis. Lips are slightly furrowed and the mental lobes are reduced. The maxillo-mandibular barbels are the longest. The nostrils are near the eyes with a relatively long tube on the anterior opening. The dorsal fin origin is at the middle of the body or slightly posterior. The pelvic fin insertion is just below the dorsal fin origin. The caudal fin is emarginate or slightly forked with rounded lobes. The anterior part of the body is scaleless. The stomach is syphonal and there is a single intestine loop. The gas bladder capsule has two globular chambers connected by a relatively long duct.

Sexual dimorphism

A 37.3 mm standard length specimen from the type series (ZMUC 27117) has the first pectoral fin branched ray broadened with wide band of tubercles and rays 2-4 decreasingly tuberculate. This male has a preorbital prolongation or flap. Paratype ZMUC 27105 has tubercles on up to 6 branched rays of the pectoral fin in broad bands, gradually decreasing in extent on more medial rays.

Colour

The head and body are yellowish, darker and more intense along the upper flank. Bars on the body are sometimes irregular or interrupted and number 7-17. The caudal fin has a bar at its base. The anterior dorsal fin base has a round dark spot (Nalbant and Bianco (1998), appears as an extension of saddle, a character of Oxynoemacheilus in illustrations); or no spot or a small and pale brown spot (Bănărescu and Nalbant (1966), appears as a distinctive spot of the Paraschistura type in illustrations - both based on the same material).

Size

Reaches 54.9 mm total length and 45.0 mm standard length.

Distribution

This species is endemic to Iran, known from the Tigris River and Gulf basins as in the type description cited above.

Zoogeography

Nalbant and Bianco (1998) regard this species as the westernmost member of the genus Paraschistura (but see dorsal fin spot problem).

Habitat

Unknown. Golzarianpour et al. (2011) give a length-weight relationship for fish from the Sabzab River with a (intercept) being 0.030 and b (slope) 2.94.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

None.

Conservation

Population trends and numbers unknown.

Further work

More collections are needed to record information on biology and distribution.

Sources

Type material: The holotype (ZMUC P 27110) and paratypes (ZMUC P 27105, 27106, 27107, 27109, 27117) of Schistura nielseni, see above and note possible confusion in numbers.

Paraschistura prashari
(Hora, 1933)

Reported from the Mashkel (= Mashkid) River basin of Pakistan on the southeastern border of Iran and from the Farah River basin in the Sistan drainage of Afghanistan as Nemacheilus prashari lindbergi Banarescu and Mirza, 1965 (or as a distinct species, lindbergi, in Schistura or Nemacheilus in Mirza, Nalbant and Banarescu (1981) and Menon (1987)) and from the Helmand River basin in the Sistan drainage of Afghanistan as Nemacheilus prashari haarlovi Bănărescu and Nalbant, 1966 (Banarescu and Mirza, 1965; Bănărescu and Nalbant, 1966; Mirza, Bănărescu and Nalbant, 1969; Coad, 1981c; Mirza, Nalbant and Banarescu, 1981). Possibly a synonym of P. kessleri as suggested by Berg (1949). No Iranian record.

Paraschistura sargadensis
(Nikol'skii, 1899)

Common names

sagmahi-ye Sarhad.

[Turkmenskii golets or Turkmen loach in Russian; perhaps Sarhadd loach should be used in English, see below].

Systematics

The syntypes of Nemacheilus sargadensis are in the Zoological Institute, St. Petersburg under catalogue number ZISP 11700 with locality data in Latin as "Sija-Rischan in sargado. 20.VIII (6)" (Nikol'skii, 1899) and in Russian in Berg (1949) "Zia-rishan, Province of Sarhad in Kerman, S.W. Iran, border with Beluchistan, not far from Kuh-e Taftan volcano, 1 IX 1898, N. Zarudnyi". The catalogue data in ZISP give the date as 20.VIII. There are 31 fish, 31.0-53.7 mm standard length, in rather poor shape, some stained dark brown, others uniformally decoloured or drab. Sarhad is the "frontier" or "land at the upper boundary", an area of plains or broad valleys and isolated mountains around Khash, for example, and a term used generally for the mountainous plateau of Baluchestan: Saadati's (1977) statement that the Sarhad is unlocatable is in error.

Nemachilus turcmenicus Berg, 1932 is described from "einem Bache unweit von der Eisenbahnstation Gjaurs in Turkmenistan (Transkaspien)" near the Iranian border and alternatively from the Kelte-chinar River (Cherokh River) near Gyaurs (37°47'N, 58°44'E), Turkmenistan (Catalog of Fishes). Three syntypes, 31.7-33.8 mm standard length, are in ZISP 11064. Berg (1948-1949) synonymises turcmenicus with sargadensis but Bănărescu and Nalbant (1966) consider it to be a valid subspecies, adding a third, paludani, from the Kabul River drainage of Afghanistan. Later Mirza, Nalbant and Banarescu (1981) separate paludani as a distinct species in a different species group from P. sargadensis, since P. sargadensis has scales, an almost forked caudal fin, a feeble dentiform process and irregular bars on the flank, all in contrast to P. paludani. Given the remote localities of turcmenicus and sargadensis they are unlikely to be synonyms (Kottelat, 2012).

Berg (1948-1949; 1949) and Saadati (1977) consider this species to be close to P. kessleri (q.v.), possibly a synonym. Nalbant and Bianco (1998) place this species in the genus Schistura.

Key characters

The caudal peduncle is deeper than in Schistura bampurensis and it lacks the protuberance under the eye in males.

Morphology

Dorsal fin with 2-5 unbranched and 6-8 branched rays, anal fin with 2-3 unbranched and 4-6, usually 5, branched rays, pectoral fin branched rays 7-10, and pelvic fin branched rays 6-7 for P. sargadensis turcmenica (Berg, 1932a; 1948-1949; Nikolaev, 1993). Body scaleless (scaled according to Mirza, Nalbant and Banarescu (1981) based on fish from the Kul River, Hormozgan ? check on my fish as this would distinguish from kessleri). Caudal fin moderately emarginate (deeply emarginate, almost forked in Mirza, Nalbant and Banarescu (1981)). Caudal peduncle depth 1.6-2.2 in caudal peduncle length. Dorsal fin origin nearer caudal base than snout tip and slightly behind the pelvic fin origin. A dorsal adipose fin may be present or absent. Dentiform process on upper jaw, if present, rudimentary.

Meristics for Iranian specimens:- branched dorsal fin rays 6(1) or 7(105), anal fin branched rays 5(106), pectoral fin branched rays 9(11), 10(84) or 11(11), pelvic fin branched rays 6(1), 7(94) and 8(11), and flank bars 11(12), 12(8), 13(13), 14(11), 15(8), 16(5), 17(5) or 18(5).

Sexual dimorphism

Unknown.

Colour

Body colour is brown backed with greenish tinges or grey, with 13-14 dark bands, in some fish hardly visible (and irregular in form according to Mirza, Nalbant and Banarescu (1981)). The bands are broad, slightly broader than the interspaces. A dark stripe runs along the midline of the back. The caudal fin has 2 dark bars and the dorsal fin an anterior dark spot at its base. A dark band is present at the caudal base. Young fish have a row of dark speckles along the lateral line.

Size

Reaches 6.5 cm.

Distribution

Found in the streams of the northern slope of the Kopetdag in Turkmenistan as Nemacheilus sargadensis turcmenicus, possibly extending into northern Iran. In southern Iran it is found in the Hormuz, Hamun-e Jaz Murian, Hamun-e Mashkid and Makran basins (Nalbant and Bianco, 1998) including at Tangeh Sarreh, Baluchestan, Shah Abbas qanat at Assadabad and the Kul River near Darab.

Zoogeography

See family account.

Habitat

Unknown. Golzarianpour et al. (2011) give a length-weight relationship for fish from the Halil River with a (intercept) being 0.007 and b (slope) 3.12.

Age and growth

Maximum age in the subspecies turcmenica does not exceed 1.5 years (Nikolaev, 1993).

Food

Important food items in the subspecies turcmenica are larval mayflies, caddis flies, chironomids, dragonflies and detritus (Nikolaev, 1993).

Reproduction

Eggs in the subspecies turcmenica reach 1.24 mm in diameter (Nikolaev, 1993).

Parasites and predators

Unknown.

Economic importance

None.

Conservation

?

Further work

?

Sources

Type material: Syntypes of Nemacheilus sargadensis (ZISP 11700), see above.

Iranian material: CMNFI 1979-0210, 37, 21.0-35.4 mm standard length, Kerman, river west of Istor (29º21'N, 56º08'E)?to locate; CMNFI 1979-0215, 14, 27.7-51.7 mm standard length, Kerman, Kharan River drainage at Baft (29º14'N, 56º37'E); CMNFI 1979-0218, 8, 32.8-47.8 mm standard length, Kerman, Kharan River drainage (28º53'N, 56º55'E); CMNFI 1979-0220, 1, 37.3 mm standard length, Kerman, jube 2 km south of Jiroft (28º39'N, 57º43'E); CMNFI 1979-0221, 19, 24.3-46.8 mm standard length, Kerman, river in Halil River drainage (28º51'N, 57º52'E); CMNFI 1979-0308, 11, 22.2-40.1 mm standard length, Kerman, river 44 km from Baft (29º02'N, 56º50'E); CMNFI 1979-0338, 68, 17.0-38.6 mm standard length, Baluchestan, Tahlab River drainage 8 km from Mirjaveh (28º58'N, 61º24'E); CMNFI 1979-0339, 5, 23.7-30.9 mm standard length, Baluchestan, Tahlab River drainage 16 km from Mirjaveh (28º56'30"N, 61º21'E); CMNFI 1979-0340, 25, 15.9-37.5 mm standard length, Baluchestan, jube at Ladiz (28º55'N, 61º18'E); CMNFI 1979-0419, ?; CMNFI 2008-0138, ?.

Genus Triplophysa
Rendahl, 1933

This genus contains loaches of the High Asian region numbering over 110 species. The region centres on the Tibetan plateau with some species found in north China to the east and in Central Asia to the west. The largest nemacheilines are found in this genus. One species is reported from Iran. The date of publication of the genus is given as 1874 in Kottelat (2012), a lapsus.

The body is elongate with a rounded belly and the caudal peduncle is slightly compressed or highly depressed, and is often very elongate or whip-like. The upper lip is deeply furrowed and there is no processus dentiformis on the upper jaw. The dorsal fin has 6-9 branched rays and is emarginate. There is no adipose crest on the caudal peduncle although a small ridge may be present in some species. The caudal fin is slightly to deeply forked, or is truncate. There is no pelvic axillary lobe. Scales are absent and the lateral line is usually complete. The swimbladder is reduced to two lateral parts and a posterior chamber which is rudimentary or with a secondarily developed bladder lying free in the abdominal cavity. A reduced swimbladder is an advantage in fast water, reducing buoyancy. The gut is short or long, a simple s-shape or a spiral. The colour pattern is usually mottled rather than banded. Males have a raised tuberculate area below the nostrils extending from the lip corner to the anterior eye margin, separated by a groove from a lower tuberculate area. Pectoral fin rays have thickened and tuberculate pads on their dorsal surface.

Prokofiev (2009) summarises osteological characters in his key and these include possession of 3 pectoral fin radials, if 4 are present then the 2 external ones are more or less dilated, flattened and at least partially overlie each other, and the presence of the preethmoid I bone in the skull. Prokofiev (2010) adds other osteological characters.

Triplohysa farwelli
(Hora, 1935)

Described from the "Helmand river, Afghanistan" in the Sistan basin but without any definite locality; it may be remote from the Sistan lowlands of Iran (Hora, 1935). Originally described under Nemacheilus. No Iranian record.

Triplophysa griffithii
(Günther, 1868)

Described erroneously as from Assam but the type locality is the Arghandab River near Kandahar in the Helmand River drainage of the Sistan basin (Hora, 1929; 1935; Bănărescu and Nalbant, 1966). Originally described in Nemachilus (sic). No Iranian record but see T. stolickai.

Triplophysa stolickai
(Steindachner, 1866)

Courtesy of Stephane Ostrowski
Courtesy of Stephane Ostrowski

Common names

sagmahi-ye Pamir.

 

[Tibetskii golets or Tibetan loach, Pamirskii golets or Pamirs loach, both in Russian; singhat in Pakistan; Stoliczka's loach].

Systematics

Often spelt stoliczkae, (e.g. in Annandale and Hora (1920)), the original spelling is Stoličkai, which becomes stolickai in Latin where accents on letters are not used (Kottelat, 2012). This species is placed in the genus Triplophysa by Menon (1987) and Bănărescu and Nalbant (1995).

Nemacheilus akhtari Vijayalakshmanan, 1950 described from "Farakhollum, about 10 miles South of Gardan Diwar, Helmund river" (Farakhulm at 34°31'N, 68°08'E), Afghanistan in the Sistan basin may be a synonym of Nemacheilus (= Triplophysa) griffithii (Günther, 1868) according to Bănărescu and Nalbant (1966) or T. stolickai according to Prokofiev (2007). There is no Iranian record.

This is a highly variable species originally described from streams around Lake Tsumureri in Rupshu Province of western Tibet and also found in the upper reaches of such rivers as the Hwang, Brahmaputra and Yangtse. It is reported further west from the Aral Sea basin, the upper Indus River basin, the upper Helmand River basin in Afghanistan and probably in Sistan in Iran. Prokofiev (2007) considers the systematics of the loaches in the stolickai group to be the most complicated in High Asian species. Berg (1948-1949) gives a series of figures which plainly show the variability in body form and pigmentation of this species. Such a wide distribution and variability may be indicative of several taxa being confused under this name.

Regan (1906) refers specimens from Sistan to Nemacheilus stenurus Herzenstein, 1888 (= Triplophysa stenura). Annandale and Hora (1920) consider stenurus not to be a distinct species but Hora (1922) compares stenurus with Nemacheilus tenuis (see below) and notes that stenurus has a continuous and entire lower lip (slightly indented in ZISP 7355 examined by me - wide interruption and greatly plicated in tenuis) and the dorsal fin origin is closer to the snout than the caudal fin base (equidistant or nearer the caudal base in tenuis). Specimens from Sistan labelled as Nemacheilus stenurus by Regan (1906) are then misidentified T. stolickai according to Annandale and Hora (1920). Since N. stenurus was described from the sources of the Yangtse Kiang in China and is a more recent name, it is probably not relevant to Iranian loaches.

Berg (1948-1949) places Nemacheilus tenuis Day, 1877 in stolickai as var. tenuis but notes that it occurs together with the type form in the Gunt River of the Pamirs in Tadjikistan as well as in Sistan, and thereby the implication is that this variety is not a subspecies (since subspecies do not occur together). Berg (1948-1949) also has the subspecies Nemacheilus stenurus uranoscopus Kessler, 1872 described from the Zeravshan River in Uzbekistan as possibly occurring in the Helmand River basin, presumably the upper reaches, but he includes Nemacheilus tenuis Day under this subspecies too. Note that the publication by Day is dated 1876, apparently published 1877 according to Eschmeyer et al. (1996). N. tenuis is the type species of the subgenus Indotriplophysa Prokofiev, 2010 elevated to a genus by Kottelat (2012). Kottelat (2012) notes the widely disjunct localities of the types of this species suggesting that two species may be involved.

Annandale and Hora (1920) initially place Sistan specimens in Nemacheilus stolickai but later Hora (1922) places them in Nemacheilus tenuis. Hora (1922) restricts the name stolickai to fish from the Indus River basin and considers that the various wide-ranging reports of stolickai refer to various other species. See also Vijayalakshmanan (1950) for comparisons with some nominal Afghan species in the Helmand basin.

The fish from Sistan examined by Annandale and Hora (1920) and assigned to T. stolickai are in two forms - one with a thin caudal peduncle, the stenurus type which they do not recognise as specifically distinct, and one with a thick caudal peduncle which they place in var. leptosoma Herzenstein, 1888 (note that varieties are not recognised by the International Code of Zoological Nomenclature). The former has a minimum caudal peduncle depth in caudal peduncle length of 5.3-7.8, mean 6.8 based on 3 fish from a table in Annandale and Hora (1920) and the one fish seen by me, and the latter has values of 2.9-3.6, mean 3.1 from the table in Annandale and Hora (1920). Annandale and Hora (1920) point out that this character is dimorphic but is neither related to sex nor "race" and that gut loop development also varies but independently of the caudal peduncle character. The specimen I examined from Sistan has a stenurus form of caudal peduncle (value 7.6), the dorsal fin origin is about equidistant between the snout tip and caudal base and so agrees with neither stenurus nor stolickai (see table in Vijayalakshmanan, 1950)), the head length in standard length (4.5) falls within the limits of stolickai (4.2-4.8) and not stenurus (4.9-5.5) (see table in Vijayalakshmanan, 1950)), and the lip grooves resemble another species, Nemacheilus akhtari Vijayalakshmanan, 1950, described from the Helmand River at Farakhollum about 10 miles south of Gardan Diwar in Afghanistan. The conclusion I reach here is that an individual fish in this group of species or forms can have a mix of characters used by authors to separate and define species.

The resolution of this problem, the correct name for Sistan loaches, other than the distinctive P. rhadinaea and P. vignai, requires extensive material from the whole range of the nominal species involved for dissection and comparison of gut shape, for analysis of sexual dimorphism and individual variability in such morphometric characters as caudal peduncle breadth and thickness and position of the dorsal fin, as well as in determining new and, hopefully, definitive characters. Conservatively, I refer Sistan fish to stolickai, the oldest available name, while recognising that tenuis may be a distinct species or subspecies and be the fish found in Sistan, or even that the fish in Sistan are an unnamed taxon or taxa. The Iranian and other material available to me does not permit a resolution of this wide ranging problem.

Syntypes of Cobitis Stoličkai are in the Naturhistorisches Museum Wien under NMW 48436 (5 fish), NMW 48439 (1) and NMW 50477 (4).

A syntype, or at least material examined by Day, of Nemacheilus tenuis is in the Naturhistorisches Museum Wien under NMW 48477 (Eschmeyer et al., 1996).

A syntype of Nemacheilus stenurus, 135.2 mm standard length, is in the Zoological Institute, St. Petersburg, Russia under ZISP 7355. Other types are ZISP 7256 (4), 7354 (3, now 2), 7355 (1), and BM(NH) 1891.10.7.33 (1, 64.9 mm standard length) (formerly in ZISP).

SEE Prokofiev (2007) for tenuis and stoliczkai and other data ?

Key characters

The elongate yet rounded caudal peduncle without an adipose fin distinguishes this species from P. rhadinaea and P. vignai the only other nemacheilids recognised in Sistan. The depth of the caudal peduncle just behind the end of the anal fin is the same as the width.

Morphology

Dorsal fin with 2-4 unbranched rays (anteriomost ones small and hidden in flesh) and 6-9, usually 7-8, branched rays, anal fin with 2-3 unbranched and 5 branched rays, pectoral fin with 8-12 branched rays, and pelvic fin with 6-8 branched rays. Total vertebrae 44.  Lateral line complete and distinctive. Scales are absent. Caudal peduncle long and slender or short and deep (see above). Head length in standard length 4.0-4.8. Eye diameter in head length 4.8-5.5, mean 5.2 in Annandale and Hora (1922) for Sistan fish (note that Day (1876; 1878) gives eye diameter as 8 in head length but Steindachner (1866) in his original description gives 5.5). Snout blunt. Barbels are reported as the maxillary ones reaching behind the eye, the rostral ones shorter; the specimen from Sistan had the maxillary and longest rostral barbel about equal in length, the maxillary reaching back to about mid-eye level. Lips are rugose, deeply incised and may be fimbriated. The lower lip is fimbriate and interrupted in the middle. There is a narrow, elongate post-labial groove with well-marked ridges on each side extending back on the mid-line of the lower head (in one specimen seen by me). The pectoral fin tip is formed by the third and fourth branched rays. Dorsal fin origin nearer the snout tip than the caudal base in Iranian specimens or equidistant while elsewhere in the range of this species it is nearer the caudal base (but dorsal origin is individually quite variable in Nemacheilidae). Caudal fin slightly emarginate, lower lobe longer than upper. A well-marked groove between the anus and the anal fin is absent. The gut length is slightly longer than the fish itself with two loops. Chromosome number is 2n=50 for fish from Kyrgyzstan (Klinkhardt et al., 1995).

Sexual dimorphism

Mature males have the first 3-6 pectoral fin rays thickened and tuberculate. A raised tuberculate area below the nostrils extends from the lips to the eye and is separated by a groove from a tuberculate area below (see Fig. 2B in Bănărescu and Nalbant (1995)).

Colour

The flank bears irregular dark spots which may coalesce into a stripe, or is marbled dark green or black. There are dark brown saddles over the back. Fins have rows of pigment, best developed on the dorsal and caudal.

Size

Reaches 16.5 cm.

Distribution

Found from Tibet to northern Kashmir, Turkestan and Sistan. In Iran, it is reported from the Hirmand River delta of Sistan (Annandale, 1921).

Zoogeography

See family account.

Habitat

Unknown.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

None.

Conservation

Biology and numbers are poorly known so an assessment of conservation status cannot be made.

Further work

The biology of this species needs study.

Sources

Type material: Syntypes of Nemacheilus stenurus (ZISP 7355 and BM(NH) 1891.10.7.33), see above.

Comparative material: BM(NH) 1905.4.7:3, 1, 87.7 mm standard length, Sistan, Helmand River, collected by H. McMahon and from the Indian Museum, Calcutta (the size of this fish is at variance with the 6 specimens measured by Annandale and Hora (1920:181) but appears to be from the collections made by the Seistan Arbitration Commission of 1902-1904 and is possibly the seventh specimen referred to in the text. This specimen has been identified as Triplophysa griffithii by S. O. Kullander and has old labels as Nemacheilus stenurus and Nemacheilus tenuis).

Genus Turcinoemacheilus
Banarescu and Nalbant, 1964

This genus contains a single species and is only recently recorded from Iran. The characters of the genus are the same as the species.

Turcinoemacheilus kosswigi
Banarescu and Nalbant, 1964

Gamasiab River, Karkheh River basin, January 2010, courtesy of K. Abbasi
Gamasiab River, Karkheh River basin, January 2010, courtesy of K. Abbasi

Turkey, courtesy of Jörg Freyhof
Turkey, courtesy of Jörg Freyhof

Common names

None.

Systematics

Originally described from Kapozik Kadun, Hakkari, Turkey. The holotype is ZMH H1884 and 6 paratypes are ZMH H1885.

Key characters

The pelvic fin origin lies in front of the dorsal fin origin and the anus lies closer to the pelvic fins base than the anal fin base, distinguishing this species from all other Iranian nemacheilids.

Morphology

The body is of uniform depth and almost cylindrical. Dorsal fin with 2-4 unbranched rays and 7 branched rays, anal fin with 1-4 unbranched rays and 5 branched rays, pectoral fin with 7-9 branched rays, and pelvic fin with 5-7 branched rays. The lateral line is short with 18-19 pores, ending before the dorsal fin origin level. The lateral line pores are larger than in other species in Iran. The body is scaleless. Eyes are small and far apart. There is no processus dentiformis. All fins are small and more or less rounded. The dorsal fin origin is behind the level of the posterior margin of the pelvic fins. The caudal fin is weakly emarginate. The lips are simple and smooth, the head short and flat and there is no keel on the caudal peduncle. The greater distance between dorsal and ventral fin, longer snout length, lower body depth and narrow body width are viewed as adaptations to fast flowing streamlets (Golzarianpour et al., 2009). The pectoral fins are large and horizontal and act as vacuum suckers (and possibly the pelvic fins too) (Breil and Bohlen, 2001). There is a fairly large fleshy pelvic axillary lobe.

Sexual dimorphism

None.

Colour

Overall colour is brown to yellowish. The back has a brown line. There are a series of irregular flank blotches (ca. 12) extending from the back to the mid- or lower flank but not the belly. These are weakly expressed anteriorly or merge into the general dark pigmentation there. There is a bar or arc of dark pigmentation at the caudal fin base. Fins are not spotted.

Size

Attains 5.9 cm total length (Golzarianpour et al., 2011).

Distribution

Reported from the Sezar River, a tributary of the Dez River at 33°28'N, 49°03'E by Golzarianpour et al. (2009) and in the northern Gulf basin (Nalbant et al., 2010). Also found in the Tigris River and its tributaries and the upper Euphrates River basin including the Göksu River in Turkey (Breil and Bohlen, 2001; Golzarianpour et al., 2009).

Zoogeography

An endemic species in the Tigris-Euphrates basin, its relationships with other nemacheilids is uncertain.

Habitat

Gravel beds with a strong current are favoured, the slender body allowing this fish to exploit spaces between the gravel. Capture sites in Turkey had clear to muddy water. Aquarium specimens showed thigmotaxis, fixing themselves in place by erecting all fins and pressing against opposing structures. In a strong current, they attached to the substrate with the pectoral fins using suction. Fish even left the aquarium water, a few centimetres above the waterline, attached to the glass with only the tail tip in the water. Splashing with water moved the head and body but the pectoral fins remained attached (Breil and Bohlen, 2001).The Iranian specimens were caught at 1456 m altitude.

Age and growth

Unknown. Golzarianpour et al. (2011) give a length-weight relationship for fish from the Sezar River with a (intercept) being 0.008 and b (slope) 2.89.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Tajbakhsh et al. (2012) record metacercaria of Posthodiplostomum cuticola and Allocreadium sp. (Digenea) and Procamallanus sp. (Nematoda) in fish from the Tigris River basin in Iran.

Economic importance

None.

Conservation

Biology and numbers are poorly known so an assessment of conservation status cannot be made. It is widely distributed and probably more common than current knowledge indicates. Golzarianpour et al. (2009) point out that dam construction and human activities  may threaten the species.

Further work

The biology of this species needs study.

Sources

Based on Golzarianpour et al. (2009).

Bagridae

The bagrid catfishes are found in fresh waters of Africa and Asia. Some species attain 2 m in length. There are about 20 genera and about 213 species (Nelson, 2006; Eschmeyer and Fong, 2011). Only one species is known from Iran.

This family is characterised by a scaleless body; a depressed head and rounded to compressed elongate body; short dorsal fin with a strong, often serrated spine; a strong, serrated spine in the pectoral fin; a short to long adipose fin; anal fin short to long; caudal fin forked or deeply emarginate; a free margin to the gill membranes over the isthmus; the anterior and posterior nostrils are well separated; anterior nostrils tubular; mouth ventral and transverse or arched; 4 pairs of barbels with the nostril barbel on the posterior nostril, maxillary barbels can be very long, other barbels are mandibular and mental (chin); teeth on the prevomer, premaxillaries and mandible; and eyes often covered by skin. Maximum size is about 2 m.

These catfishes are generally nocturnal. Certain species are important food fishes and others are kept as pets in aquaria.

Genus Mystus
Scopoli, 1777

This is a catchall genus comprising numerous species in Asia. Roberts (1994) restricts the genus to 8 closely related species (see also Grant (2004)). Only 1 species is known from Southwest Asia including Iran. A history and usage of the name Mystus is given by Jayaram and Anuradha (1984).

This genus is characterised by an elongate body, rounded anteriorly and compressed posteriorly, a short and moderately depressed head, head smooth or rugose, an elongate cranial fontanelle extending posteriorly to the base of the occipital process and divided into anterior and posterior portions of nearly equal length by an epiphyseal bar (Roberts, 1994), small to moderate eyes set high and not visible from the ventral surface of the head, a free circular eyelid, a wide transverse, usually subterminal mouth, maxillary barbels very long, jaw teeth are in villiform patches, on the lower jaw as a curved or angular band interrupted at the mid-point, continuous and curved slightly in the upper jaw, total gill rakers 11-30, gill openings very wide and free from the isthmus, adipose fin high and very long, caudal fin deeply forked, upper caudal lobe often much larger than lower, 37-46 vertebrae about equally divided between abdominal and caudal ones, and branchiostegal rays 6-12.

Mystus pelusius
(Solander, 1794)

Photograph from Khuzestan, courtesy of A. Afzali

Photograph from Khuzestan, courtesy of A. Afzali

Photograph from Khuzestan, courtesy of A. Afzali
Above three photographs from Khuzestan, courtesy of A. Afzali

Common names

ابوزمير (abu-zummair or abu zumir, an Arabic name used in Khuzestan), mahi nish dor (an Abadani name, from www.abadan.net/abadanidictionary.html, downloaded 4 December 2003), sag mahi (= dog fish), گربه ماهي (= gorbeh mahi, meaning catfish), chamu.

[abu-zummair, abouz-zoumeir, abu-al-zamir, abu'l-zoumeir, abu al-zummayr al-'amiq, zugzug in Aleppo, jahudi in Mosul, all in Arabic; Tigris mystus (Fricke et al., 2007)].

Systematics

Synonyms are Bagrus halepensis Valenciennes in Cuvier and Valenciennes, 1840 from the "Couiac, qui est la rivière d'Alep" (= Quwayq at Aleppo, Syria) (see Bailey (1951) on the publication date), Macrones aleppensis Günther, 1864, Macrones colvillii Günther, 1874 from "Bagdad", and Mystus misrai Anuradha, 1986 described from "Lake Antioche, Syria". Hypselobagrus Aleppensis is a new combination by Lortet (1883). Günther (1864) proposed Macrones aleppensis on page 75 but withdrew the name on page 431 when he realised the species was the Silurus pelusius of Solander.

No types of Silurus pelusius described from the "River Kowick" (= Quwayq) are known (Eschmeyer et al., 1996). A syntype of Macrones aleppensis, 127.8 mm standard length, from the "R. Coic, Aleppo, Syria" is in the Natural History Museum, London (BM(NH) 1955.6.25:1). Six syntypes of Macrones colvillii are also in London, ca. 175-234 mm standard length, from "R. Tigris nr. Baghdad" collected by Colville (BM(NH) 1874.4.28:6-8, 1875.1.14:19-21; Roberts (1994) agrees with my observations although Eschmeyer et al. (1996) give BM(NH) 1875.1.14:19-20 for the latter, i.e. 2 fish only).

The holotype of Mystus misrai is in the Muséum d'Histoire Naturelle, Geneva under MHNG 603.95, measuring 123.1 mm standard length, with 1 paratype under MHNG 2231.84 measuring 117.6 mm standard length, 1 paratype in the Zoological Survey of India, Calcutta under FF2315 and 1 specimen missing (Eschmeyer et al., 1996).

Roberts (1994) indicates that more material should be examined to compare fish with short barbels, weakly serrate dorsal spine, short adipose fin and a highland distribution (= pelusius) while the contrasting fish are colvillii.

Key characters

The 4 pairs of barbels, a strong spine in both the dorsal and pectoral fins, elongate and strong adipose fin are distinctive. The head tapers but is not as flattened as in Heteropneustes fossilis, and the mouth is subterminal.

Morphology

The dorsal fin spine is smooth on most of the outer edge and rough on the inner edge. Serrae can be weakly developed or absent, or may be well developed. The dorsal fin spine has 1-4 serrae or notches at the anterior tip and 5-9 along the rear margin, apparently not related to fish size. The pectoral spine is stronger than the dorsal spine and is serrated with 14-23 antrorse teeth on the inner margin, the number increasing with size. The maxillary barbel extends back to the pectoral fin origin or, rarely, as far as beyond the anal fin.

Dorsal fin with 1-2 spines and 7-8, usually 7, branched rays, anal fin with 6-10 branched rays, pectoral fin with 1 spine and 7-9 branched rays and the pelvic fin with 5-6, usually 5, branched rays. Al-Hassan and Hassan (1993) have shown asymmetry in pectoral ray and gill raker counts in samples of this species from the Shatt al Arab, Iraq, possibly due to environmental stress. Total gill rakers 10-18 (Roberts (1994) gives 12(1), 13(7), 14(11), 15(11), 16(8), 17(4) and 18(1), reaching the second raker or further when appressed. Total vertebrae 42-46 (Roberts (1994) gives 42(10), 43(16), 44(3), 45(2) and 46(5). The gut has a large stomach followed by an intestine with about 4 loops.

In specimens examined by me dorsal fin with 2(11) spines and 6(1) or 7(10) branched rays, anal fin with rays difficult to separate into branched and unbranched (perhaps 4-6 unbranched and 6-10 branched rays), pectoral fin with 1(11) spine and 7(3), 8(5), 9(2) branched rays and the pelvic fin with 5(11) branched rays. Total gill rakers 12(1), 13(2), 14(2), 16(2). One specimen with only 7 gill rakers, was possibly abnormal. Total vertebrae 42(1), 43(1), 44(2) and 45(1).

Sexual dimorphism

Unknown.

Colour

Pale brown to olivaceous overall with fins and belly lighter, on a predominant dark silver. Some fish may be silvery-grey overall when fresh. A dark shoulder spot is present. There may be a black spot at the base of the dorsal fin. The dorsal and anal fins have melanophores on the rays and membranes and so are darker than the other fins. The margin of the adipose fin is narrowly black. The caudal fin has a black margin. There may be 3 (sometimes 2), narrow, white stripes on the flank, one along and one each above and below the lateral line. The stripe below the dorsal and adipose fins is narrower than the others. Barbels are whitish, somewhat darker dorsally. Peritoneum silvery to light brown. Jayaram and Sanyal (2003) report an albino specimen from Baghdad.

Size

Reaches 171.2 mm standard length or 22.9 cm total length (Günther, 1874) but possibly to 30 cm total length (Firouz, 2005) or 33.0 cm (Al-Rudainy, 2008).

Distribution

Found in the Orontes, Quwayq, and Tigris-Euphrates basins. In Iran this fish is found in the lower reaches of rivers in the Tigris River basin of Iran such as the Karun, Karkheh and Jarrahi rivers, including too the Zohreh River of the northern Gulf basin (Abdoli, 2000). Also recorded from Fars near Darab in the Rudbal River drainage which flows to the Straits of Hormuz, possibly an accidental introduction, although Esmaeili and Coad (2005) point out that there is no evidence of fish introductions from Khuzestan to Fars. It may simply be very rare outside the Tigris-Euphrates basin in Iran.

Zoogeography

Jayaram and Sanyal (2003) consider that Mystus is derived from an African Bagrus-like ancestor and the genus spread from west to east.

Habitat

Niazi (1976) observed this species in rivers, marshes and brackish waters in Iraq although summer kill resulted from very low water levels and increased salinity.

Age and growth

Al-Hassan et al. (1991) aged this species using eye lenses and vertebrae for a population from the Qarmat Ali River north of Basrah, Iraq. Fish up to 20 cm total length were examined and three age groups were determined, with considerable overlap of lengths for each group. Al-Shami (1998) however found 7 age groups (0+-6+) for the same river using vertebrae to age fish 54-223 mm total length. The highest growth was found in the first year and no significant differences were found between males and females in growth rate. The L was 225.75 mm and the length-weight relationship log W = -4.7516 + 2.8173 log L. The relative condition ranged from 0.94 in December to 1.22 in May. Heydarnejad (2009) gave the length-weight relationship for an Iranian sample as W = 0.0277TL2.999.

Food

Roberts (1994) found eggs in the branchial chamber and stomach apparently identical with those from the ovary. Other stomach items were fish fin pieces and cyprinid fish scales. Aquatic insects, crustaceans, detritus and plant remains are also found in stomach contents of fish examined by me and Al-Rudainy (2008) also mentions fish. Al-Shami (1998) found mean feeding activity and intensity in the Qarmat Ali River, Iraq to be higher in spring and summer, declining in autumn and winter. This fish was carnivorous, taking mainly crustaceans but also insects, fishes, molluscs and aquatic plants. Hussein and Al-Shami (2001) also reported that fish in the Garma Canal, Iraq had a diet dominated by crustaceans (the isopod Sphaeroma annandalei, amphipods, the decapod Elamenopsis kempi, and the prawns Metapenaeus affinis and Atyaephyra desmaresti), followed by aquatic insects (chironomids, corixids and dytiscids), fish (Alburnus sp. and Aphanius dispar), molluscs (the gastropod Lymnaea tenera euphratica) and aquatic plants. Feeding occurred year round with a peak in May and a lowest value in November. Al-Shamma'a (2005) found shrimp and insects to form 47% by volume of the diet of this fish at Al-Fuhoud, Hawr al Hammar, Iraq.

Reproduction

The Qarmat Akli River fish attained maturity in the first year of life with the smallest mature male 92 mm long and the smallest female 72 mm. Eggs were laid in May and June with a fecundity range of 1156-25,833 eggs for fish 105-180 mm total length and 11.88-49.29 g in weight. Relative fecundity was 97.3-524.1 eggs/g (Al-Shami, 1998). Al-Rudainy (2008) gives a relative fecundity of up to 541 eggs/g in Iraq

Parasites and predators

None reported.

Economic importance

This species is of no economic importance. Anglers may catch it on hook and line in Khuzestan but, being scaleless, it is not eaten.

Conservation

This species appears to be relatively common, although not often caught in large numbers, and its conservation status has not been assessed.

Further work

See under Systematics.

Sources

Description also based on Anuradha and Jayaram (1985) and Anuradha (1986).

Type material: See above under Macrones aleppensis (BM(NH) 1955.6.25:1) and Macrones colvillii (BM(NH) 1874.4.28:6-8, 1875.1.14:19-21).

Iranian material:- CMNFI 1979-0087, 1, 162.2 mm standard length, Khuzestan, Karun River at Ahvaz (31º19'N, 48º42'E); CMNFI 1979-0368, 1, 75.6 mm standard length, Khuzestan, Karkheh River (32<º24'30"N, 48º09'E); CMNFI 1991-0153, 1, 123.3 mm standard length, Khuzestan, Zohreh River (no other locality data); CMNFI 1993-0133, 1, 152.6 mm standard length, Khuzestan (no other locality data); BM(NH) 1905.10.14:57, 1, 140.1 mm standard length, Bushehr, Jarrahi River 140 miles northwest of Bushehr (no other locality data); ZMH 2524, 1, 137.1 mm standard length, Kermanshah, Karasu-Gamasiab-Seymarreh (no other locality data); ZMH 4339, 2, 100.5-100.6 mm standard length, Khuzestan, Karun River (no other locality data); uncatalogued material, 1, 57.7 mm standard length, Fars, Cheshmeh Golabi, 15 km west of Darab (28º47'N, 54º22'E)(Esmaeili and Coad, 2005).

Comparative material:- CMNFI 1980-1036, 2, 161.0-167.8 mm standard length, Turkey, Elazig, Keban Dam near Elazig (38º41'N, 39º14'E); CMNFI 1987-0017, 1, 156.8 mm standard length, Iraq, Hawr al Hammar (no other locality data); BM(NH) 1912.5.2:7, 1, 172.5 m standard length, Iraq, Shatt al Arab (no other locality data); BM(NH) 1920.3.5:5-6, 2, 95.4-101.0 mm standard length, Iraq, Basra (30º30'N, 47º47'E); BM(NH) 1936.3.10:3, 1, 56.1 mm standard length, Iraq, Euphrates River at Nasiriyah (31º02'N, 46º16'E); BM(NH) 1974.2.22:1781-2, 1, 52.3 mm standard length, Iraq, Khalis (33º49'N, 44º32'E); BM(NH) 1974.2.22:1783-4, 1, 31.0 mm standard length, Iraq, Khalis (33º49'N, 44º32'E); BM(NH) 1975.5.16:6, 1, 155.2 mm standard length, Turkey, Elazig, Euphrates River, Keban Dam Lake (no other locality data).

Siluridae

The sheatfishes are found in Europe and Asia. There are about 12 genera and about 99 species (Nelson, 2006; Eschmeyer and Fong, 2011) with 2 reported from Iran. The phylogenetic relationships within the family are examined by Bornbusch (1995).

This family is characterised by a scaleless and elongate body; a moderately compressed head; a non-protractile mouth; teeth on the jaws and palate; 1-4 pairs of barbels (nasal barbels usually absent; maxillary barbels 1-2 pairs, sometimes vestigial or absent); nostrils separate, anterior ones tubular; 4-21 branchiostegal rays; gill openings very wide; dorsal fin short and spineless (usually fewer than 7 rays and sometimes absent); anal fin very long (41 or more rays) and may be confluent with the caudal fin; adipose fin absent; pectoral fin with a spine, often serrated; and pelvic fins small to absent. The largest species is found in Iran (Silurus glanis).

Genus Silurus
Linnaeus, 1758

These catfishes comprise about 5 species found from Europe to China and India.

This genus is characterised by an elongate body, rounded anteriorly but compressed posteriorly; a depressed head; 2-3 pairs of barbels, the maxillary barbels well-developed and often as long as, or longer than, the head; a large and terminal or superior mouth; teeth in bands on the jaws and roof of the mouth; nostrils well separated; eyes small and not visible from the underside of the head; a very short and spineless dorsal fin; no adipose fin; anal fin very long and united to the rounded caudal fin; pectoral fin with a strong serrated spine; and branchiostegal rays 12-15. The genus has been revised by Kobayakawa (1989).

Krieg et al. (1999) isolated microsatellite loci in both S. glanis and S. triostegus and found that the species diverged less than 20MYA and/or high levels of genomic conservation. Krieg et al. (2000) investigated mitochondrial DNA in S. glanis but found no consistent pattern of geographic structuring in European populations, evidence that gene flow and migration between populations were possible until quite recently. Their study also included S. triostegus and the data was diagnostic for the two species.

A general Farsi name for these fishes is گربه ماهي (= gorbeh mahi, meaning cat fish).

Silurus glanis
Linnaeus, 1758

Aras River Dam, courtesy of Asghar Abdoli (measuring fish)
Aras River Dam, courtesy of Asghar Abdoli (measuring fish)
2.25 m, ca. 90 kg, 5 October 1994

Safid River, 2 km west of Astaneh, 4 June 1978, caught in a large dip-net, photograph by Brian W. Coad
Safid River, 2 km west of Astaneh, 4 June 1978, caught in a large dip-net, Brian W. Coad

Aras Reservoir, March 2012, courtesy of K. Abbasi
Aras Reservoir, March 2012, courtesy of K. Abbasi

Aras Reservoir, March 2012, courtesy of K. Abbasi
Aras Reservoir, March 2012, courtesy of K. Abbasi

Anzali, courtesy of K. Abbasi
Anzali, courtesy of K. Abbasi

Common names

mahi-e sebili, esbele or esbeleh in Gilaki (probably derived from sibil meaning moustache in reference to the barbels), esbele-ye orupaiye, Urupai or Europaiye; ispek (Floor, 2003), گربه ماهي روگاهي or gorbeh mahi rogahee, nake or naque in the Lake Orumiyeh basin, where they are known as "whales".

[naxa, nagka or nakki in Azerbaijan; loko in Armenia; som in Russian; European catfish, sheatfish, wels, wels catfish, Danube catfish].

Systematics

Silurus Glanis was originally described from lakes of Europe, Sweden. A skin is a syntype in the Natural History Museum, London under BM(NH) 1853.11.12:168 (Eschmeyer et al., 1996). An image of this type is available at http://acsi.acnatsci.org/base/image_show_wrapper.html?target=230943. Silurus chantrei Sauvage, 1882 is possibly a synonym (see under Silurus triostegus).

Key characters

This species differs from S. triostegus by having weaker and shorter teeth, the upper and lower jaws meet at an antero-dorsal position (dorsal and superior position in S. triostegus), a less serrate pectoral fin spine posteriorly, and a darker colour. Maxillary barbel length is much longer on average, although there is some overlap. Anterior mandibular barbels are alwayslonger than posterior mandibular barbels while in S. triostegus the posterior mandibular barbels are always longer (Ünlü and Bozkurt, 1996).

Morphology

The lower jaw is longer than the upper. Adults have one pair of maxillary barbels and two pairs of mandibular barbels, for a total of 6 barbels. The maxillary barbel is much longer than the head (equal to head length in Silurus triostegus). The pectoral fin spine is finely serrated or smooth on its inner surface and smooth on its outer surface. Vomerine teeth form a single broad patch, not two as in S. triostegus (Kobayakawa, 1989).

Dorsal fin branched rays 3-5, usually 4, anal fin branched rays 70-108, pectoral rays 12-18 with 1 spine (generally higher on average than in S. triostegus but still overlapping), and pelvic rays 1 unbranched followed by 9-14 branched rays (Coad and Holčík, 2000; Reshetnikov, 2002). Vertebrae 70-76 and total gill rakers 9-17 (counts of 9 and 10 may be lower arch rakers only). Total vertebrae 67-74. In specimens examined by me dorsal fin branched rays 3-4, anal fin branched rays 83-87, pectoral rays 15-16 with 1 spine, and pelvic rays 1 unbranched followed by 11-12 branched rays. Vertebrae 18-19 + 54-56 = 72-74. Total gill rakers 12, reaching the raker below when appressed. The gut has a large stomach and an intestine with 3 about loops. Chromosome number is 2n=60 (Ráb et al., 1994; Klinkhardt et al., 1995).

Sexual dimorphism

Abdurakhmanov (1962) reports that females have longer maxillary barbels, a longer postorbital length and a greater caudal peduncle depth than males in Azerbaijan.

Colour

The body is mottled with brown, green or dark grey, even ventrally, over the base colour. The back is dark, from olive-brown to a blue-black, the sides lighter and the belly greyish-white with bluish speckles. Fins are a dark red-brown to brown-violet. Paired fins have a yellowish streak in the middle. Iris yellowish with black speckles. This species can blend its colour with any bottom on which it lies in wait for prey (Fortunatova, 1961).

Size

Reaches legendary sizes of 5.5 m and 367 kg ( and possibly over 500 kg) but most are much smaller (Machacek (1983-2012), downloaded 27 July 2012). In the Volga Delta females reach 1.75 m and 31 kg and males 1.95 m and 41 kg (Orlova, 1988); in Dagestan specimens up to 1.93 m and 41.3 kg are recorded although fish weighing 3.2-4.8 kg predominate (Shikhshabekov, 1978).

In the Caspian Sea commercial fishery of Iran, this species ranges in size from 41 to 186 cm and 0.6 to 42 kg (Farid-Pak, no date). Sohrabi (1996a) reports the larger fish in Iranian waters usually weigh 10-40 kg and depicts two fish from Gilan. One caught in 1995 weighed 27 kg, the other caught in 1994 weighed 62 kg. The record fish from Iran, caught by a Mr. Haratonian weighed 120 kg and was 2.2 m long. Eastwick (1864) bought a specimen 4.5 feet long (1.37 m) in the Safid River which had a 7 lb (3.2 kg) fish in its stomach. De Mecquenem (1908) reported that they reached 2 m in the Lake Orumiyeh basin and Anonymous (1977) reports fish from there at 400 lbs (= ca. 182 kg). Asghar Abdoli of the Agricultural and Natural Resources University, Gorgan kindly sent me a photograph of a specimen from the Aras Dam caught 5 October 1994 which was 2.25 m long and weighed about 90 kg.

Distribution

This species is found in Europe, Central Asia and Southwest Asia. In Iran it is found from the Aras River and Dam, along the whole Caspian coast from the Astara to the Atrak rivers including the Anzali Mordab and its Siah Keshim Protected Region, the Safid River, the Manjil Dam on the Safid River, the Amirkelayee Lagoon near Lahijan, Gorgan Bay, and from the southwest Caspian Sea and south-central Caspian Sea; and in the Lake Orumiyeh basin including the Nowruzlu Dam, Shapur-e Avval Reservoir, Mahabad, Gader Chai at Ocksa (Günther, 1899; Nedoshivin and Iljin, 1929; Berg, 1936; Nümann, 1966; Holčík and Oláh, 1992; Nejatsanatee, 1994; Riazi, 1996; Abbasi et al., 1999; Kiabi et al., 1999; Jolodar and Abdoli, 2004; Abdoli and Naderi, 2009). Also reported from the Karakum Canal and Kopetdag Reservoir in Turkmenistan (Aliev et al., 1988; Shakirova and Sukhanova, 1994; Sal'nikov, 1995) and may enter Iranian waters of the Tedzhen (= Hari) River basin.

Abdoli (2000) maps this species from lower Gorgan, Neka, Babol, Heraz, Chalus, Tonekabon, and Safid rivers, the Anzali Talab, along the Caspian coast, and in the middle Aras River, and in the lower Talkheh and Zarrineh rivers of the Lake Orumiyeh basin.

Also said to occur in the Tigris-Euphrates basin in Iraq but needs specimens for confirmation (see Freshwater Fishes of Iraq).

Zoogeography

The isolated populations of this wide-ranging species have not been studied. It may well have an origin in the Ponto-Caspian basin from which it has dispersed.

Habitat

This large species is found in the larger water bodies over soft bottoms and can tolerate brackish water (even for spawning) and moderately low oxygen levels. Warm, deep waters with slow current are preferred. It is active at night. Adults are solitary and found under overhanging banks or submerged trees. This catfish overwinters in aggregations on river beds. Resumed activity in spring depends on the local water temperature regime, probably as early as March in Iran. Riazi (1996) reports that this species is native (resident) to the Siah-Keshim Protected Region of the Anzali Mordab. Knipovich (1921) reports this species from depths of 23.8-25.6 m in the Iranian Caspian Sea. Khodabandeh and Shahriari Moghadam (2007) examined the ultrastructure of ionocyte cells which play an active role in osmotic regulation in this fish.

Movements, such as hunting for food, are stimulated by such environmental factors as temperature, solar radiation, air pressure and turbidity after rain. It is sensitive to extra-aquatic sounds. The head canal system is very sensitive and can track prey over distances up to 55 times the length of the prey and follow signals up to 10 seconds old (Kottelat and Freyhof, 2007).

Age and growth

Life span is 22 years for males and 16 years for females in the Volga Delta. Growth is most intensive in the first years of life. After maturity the annual increase in length is 5-7 cm. Relative weight increase is very high (30%) while length increase is 6-10%. June-July is the period of greatest length increase while weight increase takes place mainly in autumn. Maturity begins at 3-4 years, 57-66 cm and 1.3-2.3 kg but may be as early as 2 years, 51-52 cm and 1.2-2.2 kg or as late as the sixth year (Orlova, 1988). Maximum life span is 80 years.

Abbasi and Valipour (2005) found 9 age groups in the Anzali Lagoon with females making up 68.5% of fish caught. Total length of 95 fish was 19.8-186.5 cm and weight was 47.7-30,000 g. Nezami Balouchi et al. (2007) found age groups 1+ to 7+ in the Amirkelayeh Wetland with an average total length of 50.6 cm (range 32.5-73.0 cm) and an average weight of 944.2 g (range 200-3000 g).

Food 

Food in the Volga Delta includes such fishes as Cyprinus carpio, Abramis brama, Scardinius erythrophthalmus and, prior to the regulation of the Volga, Rutilus rutilus (andéor R. caspicus) and herrings (Clupeidae). Crustaceans now form part of the diet there (Orlova, 1988). At one time this catfish in the Volga delta ate 62-68% of its annual food in one month in spring when the Caspian roach (Rutilus rutilus - this may be R. caspicus) arrived on its spawning run. In the Kura region, commercial fishes such as Cyprinus carpio, Abramis brama, Rutilus rutilus (andéor R. caspicus), Aspius aspius, Sander lucioperca, Chalcalburnus (= Alburnus) chalcoides, Barbus (= Luciobarbus) brachycephalus or Barbus (= Luciobarbus) capito, and Silurus glanis make up 30.27% by frequency and 20.18% by weight, non-commercial species such as Scardinius erythrophthalmus, Tinca tinca, Cobitis spp., Rhodeus amarus, Pungitius platygaster, Atherina boyeri (= caspia) Alburnus alburnus (= hohenackeri), Blicca bjoerkna, and Caspiomyzon wagneri make up 50.68% and 33.79% respectively, marine fishes such as Clupeidae, Mugilidae, and Gobiidae make up 12.90% and 8.60% respectively, and crustaceans 25.64% and 17.09% respectively (Mamedov and Abbasov, 1990). In Azerbaijan, Abdurakhmanov (1962) reports Gobio gobio to comprise 22.4% of the diet, Caspiomyzon wagneri 15.7% and eggs 2.2%, Chalcalburnus (= Alburnus) chalcoides 10.2%, Alburnus alburnus (= hohenackeri) 10.2%, Cobitis taenia (cf. keyvani) 9%, Barbus lacerta 7.8%, Barbus (= Luciobarbus) capito 6.7%, Chondrostoma oxyrhynchum 5.6%, Capoeta capoeta 3.4%, Blicca bjoerkna 2.2%, and loaches 4.5%.

In the Anzali Lagoon of Iran according to Abbasi and Valipour (2005), this species ate 78.6% bony fishes, 15.8% crustaceans, 4.13% insects, 0.9% amphibians and 0.5% bivalves. Carassius auratus dominated at 33.9%, followed by the crustacean Macrobrachium spp., at 14.22%, Neogobius kessleri (= Ponticola gorlap) at 4.59% and Proterorhinus marmoratus (= nasalis) at 2.75%. C. auratus dominated in spring and autumn, Macrobrachium spp. in summer and N. kessleri in winter. Consumption of fish increased with size, being 44.2% at 20-55 cm and 94.5% at 91-125 cm, Cannibalism was not observed and commercial fish stocks were not consumed. Catfish in the Amirkkelayeh Wetland fed principally on Tinca tinca and amphipods each at a frequency of 36.3%, along with minor amounts of Blicca bjoerkna, Pungitius platygaster, Perca fluviatilis, Proterorhinus marmoratus (= nasalis), Carassius auratus, Cobitis keyvani (identified as C. taenia), frogs, rats, birds, water bugs and beetles, plecopterans and odonatans (Nezami Balouchi et al., 2007). They were also cannibals.

Generally a wide variety of fishes is taken along with crayfish, frogs and even birds and small aquatic mammals. It is a voracious predator but stories of attacks on dogs and small children are more legendary than factual although human remains may be scavenged (Gudger, 1945b). Active feeding occurs at water temperatures above 8°C so winter feeding is minimal or absent. While feeding often occurs at night, catfish can be heard feeding in the evenings by the snapping of the mouth and tail strikes on the water. In cloudy water conditions they come into shallow water to take earthworms, grasshoppers and frogs washed in from nearby fields (Mihálik, 1982).

Young catfish feed on plankton, particularly Cladocera such as Daphnia, Chydorus, Alona and Bosmina among others. Later the diet involves mosquito larvae, larger crustaceans, organisms associated with the river bank, worms, snails and young fishes. Cannibalism occurs if food is short (Mihálik, 1982).

Reproduction

Behmanesh et al. (2009) found the maximum mean gonadosomatic index for Anzali Lagoon fish was in May and June, decreasing to September. Maximum values for gonad maturity were in April-May at water temperatures of 19.3-22.7°C. Spawning was from April to mid-June.

Non-intermittent spawning in Dagestan takes place in late May and continues to the middle of July when the eggs of females mature at a water temperature of 20-22°C. Spawning rarely occurs below 20°C. Males may actually have running milt 30-40 days earlier than this and also later, a longer potential spawning season. Adhesive yellow eggs are laid in depressions in weed beds, formed by the male pressing on the plants. Fecundity is up to 285,000 eggs with diameters around 2-3 mm (to 467,000 elsewhere). Elsewhere spawning may be intermittent (Shikhshabekov, 1978).

The male guards the incubating eggs, even during the day, moving his tail fin every 3-5 minutes to ensure adequate oxygen supplies. Nests in Europe may be on the fine roots of plants which hang freely in the water. The nest is in shallow, 40-60 cm, water. Males pursue females just under the water surface, an indication spawning will occur the same evening or the next day. Spawning usually occurs in the evening, often before a thunderstorm on warm and stifling days. The male nudges the female in the anal region, swims under her and may lift her so that her back is above water, the male wraps himself around the female for 10-12 seconds, the male and female separate and the female sinks slowly to the bottom and discharges eggs, the male following to release milt. This process can be repeated several times over 1.5-2.0 hours and the water around the nest is milky from sexual products. Eggs hatch after 2.5-3.0 days at 23-25°C. Larvae are light sensitive and die in direct sunlight ands also if water temperature falls below 13-14°C (Mihálik, 1982).

Parasites and predators

Mokhayer (1976b) records the digenetic trematodes Aphanurus stossichi and Bunocotyle cingulata, the nematode larvae Anisakis sp. and the nematode adults Cucullanus sphaerocephala, and the acanthocephalan Corynosoma caspicum. Ataee and Eslami (1999, www.mondialvet99.com, downloaded 31 May 2000) report the helminth Mazocea alaosa from the gastro-intetsinal tract of fish from the Anzali wetland. Masoumian et al. (2005) recorded the protozoan parasite Trichodina perforata from this species in the Aras Dam in West Azarbayjan. Khara et al. (2006a) record the eye fluke Diplostomum spathaceum for this fish in the Amirkalayeh Wetland in Gilan. Sattari et al. (2004; 2005) surveyed this species in the Anzali and Amirkelayeh wetlands, recording Raphidascaris acus, Raphidascaroides sp. and Eustrongyloides excisus. Barzegar et al. (2008) record the digenean eye parasite Diplostomum spathaceum from this fish. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Cyprinacea sp. and Ergasilus sp. on this species.

The Caspian seal, Pusa caspica, is a predator on this species (Krylov, 1984).

Economic importance

There is some opportunity for sport fishing for this species in the Anzali Mordab and Lake Orumiyeh basin where it will take spinners and spoons as well as frog live bait. It reputedly puts up a tremendous fight (Anonymous, 1977).

Nevraev (1929) reports on catches in various regions of Iran in the early years of the twentieth century. There were no evident trends of increase or decrease. In the Astara region from 1901-1902 to 1913-1914 the catch varied irregularly from 699 to 4031 fish, in the Anzali region from 1901-1902 to 1918-1919 the catch varied from 18,177 to 206,485 fish, in the Safid River region from 1899-1900 to 1917-1918 the catch varied from 3290 to 43,835 fish, in the Mazandaran region from 1906-1907 to 1913-1914 the catch varied from 5282 to 11,283 fish, and in the Astrabad region from 1902-1903 to 1912-1913 the catch varied from 3500 to 26,200 fish. The commercial catch in Iran from 1956/1957 to 1961/1962 varied between 4,913 kg and 37,630 kg (Vladykov, 1964) and from 1965/66 to 1968/69 varied from 11 to 31 tonnes (Andersskog, 1970) but in the 6 years from 1980 to 1985 catches were recorded by the Food and Agriculture Organization, Rome as respectively 2, 2, 0, 2, 3, and 0 tonnes. The catch has been as high as 107,593 kg for the Anzali region alone in 1934/1935 (Vladykov, 1964). Holčík and Oláh (1992) report a catch of 2663 kg in the Anzali Mordab in 1990, at 3.6% of the catch the sixth most important fish there, and from 1932-1964 reported catches varied from none to 12.6 tonnes annually. The Iranian fishery peaks in October and in April (Farid-Pak, no date). This species is of great commercial importance in Dagestan (Shikhshabekov, 1978). The roe has been used as a form of caviar and glue has been made from the swimbladder and bones. In the Lake Orumiyeh basin this fish was used as fertiliser since, being scaleless, it could not be eaten for religious reasons (De Mecquenem, 1908). Its export value is recognised and studies have been carried out on this species such as the effect of ascorbic acid and citric acid on the lipid stability and rancidity inhibition of fillets in frozen storage (Pourashouri et al., 2008a. 2008b; Pourashouri et al., 2009).

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use as food, for industrial processes (such as fertilizers, fish meal, pet food, novelty products although not specified for this species), in sport, and in textbooks. The swimbladders were used for isinglass in the Caspian Sea basin (Lönnberg, 1900a).

This species has been implicated in ichthyootoxism, the symptoms of which are summarised under the genus Schizothorax. The presence of venom associated with the pectoral fins (Coad, 1979b) needs definitive examination.

Conservation

Lelek (1987) classifies this species as rare to vulnerable in Europe because of habitat changes and angling pressure. Kiabi et al. (1999) consider this species to be of least concern in the south Caspian Sea basin according to IUCN criteria. Criteria include commercial fishing, sport fishing, medium numbers, habitat destruction, widespread range (75% of water bodies), present in other water bodies in Iran, and present outside the Caspian Sea basin. Vulnerable in Turkey (Fricke et al., 2007).

Further work

Population numbers need to be monitored carefully as it is fished for although it lacks scales. Populations in isolated basins have not been examined in detail for their relationships to see if they are distinct.

Sources

Kobayakawa (1989) revised the genus Silurus and his data are incorporated here. Mihálik (1982) reviewed the biology of this catfish.

Iranian material: CMNFI 1970-0509, 1, 131.2 mm standard length, Gilan, Safid River at Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1979-0685, 2, 63.9-107.7 mm standard length, Gilan, Safid River below Dehcha (ca. 37º22'N, ca. 50º06'E); CMNFI 1979-0788, 2, 293.5-335.6 mm standard length, Mazandaran, Gorgan River at Khvajeh Nafas (37º00'N, 54º07'E); CMNFI 1979-1236, 1, 269.0 mm standard length, Mazandaran, Gorgan River at Khvajeh Nafas (37º00'N, 54º07'E); CMNFI 1980-0123, 1, 225.3 mm standard length, Gilan, Safid River around Dehcha (37º22'N, 50º06'E); CMNFI 1980-0905, 1, 208.0 mm standard length, Mazandaran, Gorgan River at Khvajeh Nafas (37º00'N, 54º07'E); OSU 4278, 2, 227.2-280.2 mm standard length, Azarbayjan-e Bakhtari, Zarineh River (no other locality data).

Silurus triostegus
Heckel, 1843

NMW 92345, syntype, Tigris River near Mosul, ca. 582 mm standard length, photograph courtesy of
NMW 92345, syntype, Tigris River near Mosul, ca. 582 mm standard length, photograph courtesy of
B. Herzig, Naturhistorisches Museum Wien

Common names

esbele, and jirri, yeri, yery, yari or iry (= eel), esbele-ye beinolnahrein (= Mesopotamian catfish); sag mahi (= dog fish) in Khuzestan; gorbeh mahi (= catfish).

[djirri, jirri, girri or yerri (= eel) in Arabic; Mesopotamian catfish, Tigris catfish, Asian djirri].

Systematics

Silurus chantrei Sauvage, 1882, a species with 4 barbels, was described from the "Fleuve Koura à Tiflis (= Kura River at Tbilisi, Georgia) but was possibly based on material from the Tigris-Euphrates basin (Berg, 1948-49; Haig, 1952) and may well be a synonym of this species. Two syntypes of Silurus chantrei, 160-170 mm total length, are in the Muséum national d'Histoire naturelle, Paris under MNHN A.3932 (Eschmeyer et al., 1996).

Günther (1899), Banister (1980) and Hora and Misra (1943) considered that S. triostegus may not be distinct from S. glanis but Coad and Holčík (2000) detail differences.

The type locality of Silurus triostegus is the "Tigris bei Mossul" according to Heckel (1843) and the description was based on 4 specimens although the catalogue in Vienna lists 1 specimen in spirits and 2 stuffed specimens. The card index in Vienna in 1997 lists only NMW 92345 as a dried syntype. Coad and Holčík (2000) found only a single stuffed type, ca. 582 mm standard length. Eschmeyer et al. (1996) list 1 dried syntype of triostegus in the Senckenberg Museum Frankfurt (SMF 2623, formerly NMW) and this has a standard length of ca. 710 mm. An image of S. triostegus (SMF 2623) is available at http://acsi.acnatsci.org/base/image_show_wrapper.html?target=221708.

Key characters

This species differs from S. glanis by having robust and longer teeth (snaggly, catching on flesh), the upper and lower jaws meet at a dorsal and superior position (antero-dorsal in S. glanis), a distinctly and coarsely serrate pectoral fin spine posteriorly, and a lighter colour. Maxillary barbel length is about equal to head length while in S. glanis it is much longer on average, although there is some overlap. Anterior mandibular barbels (when present) are always shorter than posterior mandibular barbels while in Silurus glanis the anterior mandibular barbels are always longer (Ünlü and Bozkurt, 1996).

Morphology

Bears 4 barbels in some specimens as opposed to the usual 6 in S. glanis but there is evidence that the four-barbelled Silurus (sometimes placed in a distinct genus Parasilurus Bleeker, 1862, e.g. in Berg (1949), now synonymised with Silurus - see Eschmeyer (1990)) have 6 barbels when young and one pair of mandibular barbels is reabsorbed (see Haig, 1952). Specimens with one pair and with two pairs of mandibular barbels have been reported (Kobayakawa, 1989). Ünlü and Bozkurt (1996) record 4 mandibular barbels (2 pairs) in 3 specimens, 3 barbels in 1 specimen and 1 pair in another specimen for Turkish Euphrates fish. In Silurus triostegus, the adults apparently lose one pair of barbels (F. Krupp, in litt., 1992). Coad and Holčík (2000) found all S. glanis in their study had 4 mandibular barbels (2 pairs) while in S. triostegus 11 fish had 4 mandibular barbels and 12 fish lacked the posterior mandibular pair. There was no apparent trend in barbel loss associated with increase in body size.

The pectoral fin spine is strongly serrated on its inner surface and smooth on its outer surface. Vomerine teeth are in two patches, not one as in Silurus glanis (Kobayakawa, 1989), although Ünlü and Bozkurt (1996) record some specimens of S. triostegus with only one patch. The maxillary barbel reaches only to the end of the head, not much longer as in S. glanis. However, Ünlü and Bozkurt (1996) report that maxillary barbels are longer than the head and later that they reach the end of the head; it is suggested here that barbel length varies individually. The lower jaw is longer than the upper jaw. Teeth in both jaws are recurved, the band of teeth is wider than in S. glanis, and the teeth are stronger and longer. The eye is larger than in S. glanis.

Dorsal fin branched rays 3-4 (counts of 3 rays are more common than in S. glanis; 3 in ten fish and 4 in thirteen fish examined by Coad and Holčík (2000)), pectoral fin branched rays 11-14 with 1 spine, pelvic fin branched rays 8-13 after 1 unbranched ray, anal fin rays 77-94, vertebrae 16-17 + 52-53 = 69-70, and total gill rakers 12-17, reaching the one below when appressed (Coad and Holčík, 2000). The stomach is large, and apparently more elonagte than in S. glanis although this may be distortion due to food content. The intestine has about 3 loops.

Sexual dimorphism

Unknown.

Colour

The upper body is mottled pale yellow-brown and black. Overall colour may appear dark or light and yellowish. Generally much lighter than Silurus glanis. The belly and lower head are white with the belly having black spots. Maxillary barbels and margin of the lower jaw very dark brown.

Size

Reaches 1.5 m in the Syrian Euphrates (Gruvel, 1931) and to more than 2.0 m (Krupp, 1992) and more than 50.0 kg (Machacek (1983-2012), downloaded 27 July 2012).

Distribution

This species is found in the Tigris-Euphrates basin including its Iranian portion in Khuzestan and such rivers as the Arvand, Bahmanshir, and Jarrahi, and the lower Karun, Karkheh and Dez (Marammazi, 1995; Abdoli, 2000).

Zoogeography

This species is presumably a relative of S. glanis but its closest affinities may lie with species to the east. Differentiation of the two species may have occurred around the Middle to late Miocene but this requires further study (Coad and Holčík, 2000).

Habitat

van den Eelaart (1954) reports this species from open and vegetated lakes and marshes and rivers in Iraq. The larger fish are mostly confined to rivers, entering marshes and lakes only on floods. The young have a greater tolerance of high temperatures and low oxygen.

Age and growth

Al-Abood (1989) found age groups 2 to 7 years for fish from a marsh area north of Basrah, Iraq in 6 weight groups from 300 to 3900 g. Al-Hassan and Al-Sayab (1994) examined 600 specimens from the Al-Hammar Marsh north of Basrah, Iraq for age using vertebrae and eye lens diameter and found 6 age groups. Oymak et al. (2001) describe 11 age groups in Atatürk Dam Lake, Turkey with females having higher L (202.85 cm versus 113.98) and lower K (0.046871 versus 0.101972) values than males. Males matured at age 3 and females at age 4.

Food

Fish are an important food including Liza abu and Acanthobrama marmid in the Iraqi marshes and presumably those across the border in Iran (Al-Shamma'a and Jasim, 1993; Ünlü and Bozkurt, 1996; Dawood, 1997; personal observations). Aquatic insects are also taken but fish predominate. The food in Hawr al Hammar, Iraq was predominately fish (Liza abu, Aphanius spp., Aspius vorax, Thryssa spp., Acanthobrama marmid, Silurus triostegus (young less than 16 cm were eaten by adults longer than 35 cm), Barbus (= Mesopotamichthys) sharpeyi, Heteropneustes fossilis and Cyprinus carpio) followed by shrimps (mainly Metapenaeus affinis), frogs (Rana esculenta) and crabs (mainly Sesarma boulengeri) with relative importance indices of 70.8, 16.3, 6.4 and 4.9 respectively (Al-Daham and Al-Seyab, 2000). Liza abu was the most important fish through most of the year (except July and August when absent)(relative importance index 42.0, followed by C. carpio at 11.5), in numerical abundance and total weight. During July the prey was B. sharpeyi and C. carpio and in August prey was restricted to C. carpio, presumably opportunistic feeding. Other fish species were mostly young of the year and of minor importance. The diet at Al-Fuhoud in the Hawr al Hammar was 70.7% fish by volume (Al-Shamma'a, 2005). Aquatic insects are also taken but fish predominated. Dawood (1997) also studied diet in the southern Hammar Marsh and found fish to be the most important prey year round while shrimps (Metapenaeus affinis) and molluscs were important  in certain months (mostly absent April-August). The disappearance of shrimps probably relates to their migratory pattern. Aquatic insects were found mostly in the spring. There is a reverse relationship between the presence of fish and shrimps. Fish and shrimps increased in the diet with increase in size while aquatic insects, molluscs and small crustaceans decreased with size. Fish species eaten were Liza abu, Barbus (= Carasobarbus) luteus, Alburnus sp., Cyprinus carpio, Thryssa hamiltoni, Heteropneustes fossilis, Aphanius sp., Gambusia holbrooki and Silurus triostegus. Frogs, detritus and aquatic plants were also found in the gut contents. Feeding occurred more at night and with another peak in late afternoon. The index of fullness values increased in April-September when water temperatures and metabolic rate rose.

Reproduction

Spawning takes place in March in Iraq (van den Eelaart, 1954; Al-Hassan et al., 1990) to May and June in Turkey (Oymak et al., 2001). The highest condition factors were found in April in Atatürk Dam Lake, Turkey, the mean egg diameter was greatest in May at 1.937 mm and fecundity attained 120,300 eggs (Oymak et al., 2001). Al-Rudainy (2008) gives sexual maturity in Iraq at 3 years, 50 cm total length and 1 kg weight, spawning in May and continuing for a few months, eggs being deposited on vegetation and egg diameters up to 4.2 mm.

Parasites and predators

Mortazaei et al. (2000) report an infection rate of 33.3% (22 of 6 fish) with the parasitic worms Proteocephalus sp. in this species from Khuzestan marshes.

Economic importance

This species forms 8.5% of the total catch in Iraq (Das et al., 1978), the total catch in 1976 being 691 t (Petr, 1987), but it is not a popular food. As a scaleless fish it is not eaten by Shi'a Muslims. It was found to have potential for use as a protein concentrate.

Conservation

This species is not commonly collected in Iran but this may be a consequence of habitats sampled and gear used. It may be under some threat as it is fished for, at least in neighbouring Iraqi waters.

Further work

Population numbers have not been examined for fish in Iran nor are its biology or relationships well known.

Sources

Kobayakawa (1989) revised the genus Silurus and his data are incorporated here as is the data of Ünlü and Bozkurt (1996) for Turkish Euphrates specimens.

Type material: See above, Silurus triostegus (NMW 92345).

Iranian material: CMNFI 1993-0133, 1, 192.9 mm standard length, Khuzestan, probably Karun River at Ahvaz (31º19'N, 48º42'E); SNM-YR 6421, 1, 325.6 mm standard length, Khuzestan, Karun River at Ahvaz (31º19'N, 48º42'E); ZSM 21832, 1, 406.0 mm standard length, Khuzestan, Dez River at Harmaleh (31º57'N, 48º34'E); ZSM 21833, 1, 425.4 mm standard length, Khuzestan, Dez River at Harmaleh (31º57'N, 48º34'E); uncatalogued, 1, 317.7 mm standard length, Khuzestan, probably Karun River at Ahvaz (31º19'N, 48º42'E).

Comparative material: BM(NH) 1874.4.28:3-5 and 1875.1.14:8 (same jar), 4, 236.7-511.5 mm standard length, Iraq, Tigris River near Baghdad (ca. 33º21'N, ca. 44º25'E); BM(NH) 1888.5.17:1, 1, 336.2 mm standard length, Iraq, Fao, presumably a fish market (no other locality data); BM(NH) 1893.6.23:26-28, 3, 254.3-425.7 mm standard length, Iraq, Fao (no other locality data); BM(NH) 1892.9.1:26, 1, 270.8 mm standard length, Iraq, Fao (no other locality data); BM(NH) 1920.3.3:167-176, 13, 123.1-490.9 mm standard length, Iraq, Basrah (30°30'N, 47°47'E).

Sisoridae

The sisorid or sucker catfishes are found in Asia as far east as Borneo. There are about 18 genera with about 196 species (Nelson, 2006; Eschmeyer and Fong, 2011). They are mostly small (as small as 2 cm) although some are very large (2 m). Five nominal species are reported from the Tigris-Euphrates basin in Southwest Asia but the diversity there is very limited, compared to India for example. A single specimen of a sisorid catfish has been caught in the Yeşil Irmak of Anatolian Turkey at Taşova (40°46'N, 36°20'E). This Black Sea drainage specimen calls into question the utility of characters used in identifying and defining species in Southwest Asia. Its characters are a mixture of features shared by G. armeniacus and G. silviae. A wide range of specimens of both sexes, various age groups and from various localities is not available to assess variation and resolve the species composition of the Southwest Asian fauna (Coad and Delmastro, 1985)).

The sisorid catfish family is characterised by a rounded to compressed body and a flattened head; short dorsal fin positioned anterior to the level of the pelvic fins; a spine in the dorsal fin (absent in some non-Iranian species) and in the pectoral fin; a well-developed adipose fin; a short anal fin; paired fins horizontal; gill membranes generally united to the isthmus; anterior and posterior nostrils close together; distinct nasal barbel present; 4-6 pairs of barbels; body with small tubercles (unculi); and a distinct thoracic adhesive apparatus in Iranian species but absent in some other species.

These catfishes are found in mountain streams where they use the adhesive apparatus to maintain position in the current. In Iran they are reputed to lie on their backs in the water to take a rest! They are also very resistant and can live for 6 hours wrapped in wet cloth, reviving when placed in an aquarium (R. Mehrani, pers. comm., 2000).

Genus Glyptothorax
Blyth, 1860

This genus is characterised by a flattened head, an adipose fin of moderate length, a short dorsal fin with a strong spine, the spine serrated anteriorly or posteriorly, or smooth, pectoral fin spine serrated posteriorly and in some with plicate skin ventrally, 4 pairs of barbels, maxillary barbels broadly based, an inferior and transverse mouth, villiform teeth on the roof of the mouth in two patches, eyes small and partly obscured by skin, gill openings wide, gill membranes joined to the isthmus, and an adhesive apparatus on the chest formed from plaits or folds of skin, often with a central depression.

The validity of the described Glyptothorax species in the Tigris-Euphrates has not been adequately resolved. Three other nominal species occur in addition to the two reported here from Iran, namely G. armeniacus (Berg, 1918), G. cous (Linnaeus, 1766), and G. steindachneri (Pietschmann, 1913) (see Coad and Delmastro (1985) for a partial discussion of this problem). Resolution of the taxa found in Iran awaits more extensive material for a better understanding of individual and species variation.

Aglyptosternon Bleeker, 1862 (and such mispellings as Aclyptosteron, Enclyptosternum and variants- see Eschmeyer (1990) for details) are synonyms of Glyptothorax (Li, 1986; Eschmeyer, 1990).

Confusion over the family placement of these fishes in various literature sources is reviewed in Banister (1980).

They are known generally as گربه ماهي (= gorbeh mahi, meaning cat fish) or arteshi in Farsi. Arteshi (meaning soldier-like) may be from their tank-like appearance or for their pigmentation which is said to resemble camouflage. Another general name is سگ ماهي (sag mahi, meaning dog fish). General names are not repeated below.

Menon (1954) considered that the members of this genus had spread westwards along the Himalayas as late as the early Pleistocene. A centre of origin in western Yunnan and the southern slopes of the Himalayas is advocated by Li (1986). This author suggests that a Pliocene movement occurred westwards and that, as well, the distribution of Glyptothorax was influenced by Pleistocene glaciations. Their entry route into the Tigris-Euphrates basin is given as along the Amu Darya system.

Glyptothorax armeniacus
(Berg, 1918)


Sucker, ventral view

Found in the Euphrates River basin southeast of Erzurum in Turkey but no Iranian record.

Glyptothorax cous
(Linnaeus, 1766)

Reported from the Tigris River basin in Iraq but no Iranian record. Species identity in the Tigris needs confirmation by specimens. See Banister (1980) for a brief history of the confusion surrounding the name of this species in the literature.

Glyptothorax kurdistanicus
(Berg, 1931)


Ventral head showing sucker

Common names

gorbehmahi-ye Kordestan.

[Kordestan sisorid; Iran cat (Fricke et al., 2007)].

Systematics

This species was originally described in the genus Glyptosternum, an unjustified emendation of Glyptosternon McClelland, 1842 by Berg (1931a). See Eschmeyer (1990) for further details on this genus name.

The holotype of this species is in the Zoological Institute, St. Petersburg (ZISP 20780) and is in poor condition, the pectoral spines being damaged for example. It is an adult male. The collection date in Berg (1931a) is 10 July 1914 (or 27 June old style). The type locality is "in Kurdistan, at the village Germau (or Germav), at the height of 1500 m, during the works of the Turko-Persian delimitation commission. Germau (or Germav, Germaw) is situated in latitude 36°N south-east of Serdesht, on the western slope of the Sur-kei Range, in the basin of the river Bané, tributary to the Little Zab, which is tributary to the Tigris R.". Berg (1949) gives the variants Germab and Sar Dasht for the localities and Bané is probably Baneh. The village Germab could not be located in gazetteers or on maps but Sar Dasht (36°09'N, 45°28'E) and Baneh (35°59'N, 45°53'E) are evident and the locality is between them.

Key characters

Berg (1931a) separates this species from G. armeniacus by the broader than long adhesive apparatus which does not have pinnate lateral branches and these characters also contrast with G. silviae. The caudal peduncle is short (5.9-6.0 in standard length compared to 4.7-5.2 in G. silviae).

Morphology

Dorsal fin with 1 spine followed by 5-7 branched rays, anal fin with 2 unbranched rays followed by 7-9 branched rays (note that these fin ray counts in Berg (1931a) do not agree with his figure). Pectoral fin with 1 spine and 7-9 branched rays. Total gill rakers 7-9, moderately long and reaching beyond the base of the second raker below when appressed. The adipose fin is short, much shorter than the distance between the dorsal and adipose fins. There are oblique osseus striae under the skin of the upper surface of the first pectoral ray. Head and body covered with minute, elongate tubercles oriented longitudinally but without striae. Tubercles are also present on the base of the caudal fin rays, adipose fin, base of the dorsal and pectoral fins, on the pectoral spine along its whole length both dorsally and ventrally, a few on the base of the pelvic fin rays and few to none on the belly particularly anterior to the pelvic fins. Tubercles on the side of the head are more rounded. Berg (1931a) states that the upper jaw tooth patch has well-developed lateral rami, but these do not extend markedly from the main patch. The nasal barbel is short and does not extend back to the eye. The maxillary barbel is shorter than head length and the mandibular and mental barbels are progressively shorter. The gut is an elongate s-shape after a muscular stomach.

Sexual dimorphism

Unknown.

Colour

Overall colour grey to brown with large, obvious, round, black spots and blotches on the sides or with small round black spots about eye size. All fins with broad black central band and variably developed basal bar. Basal bar most evident on the caudal fin. Adipose mostly covered with a large dark spot but dorsal and posterior edges hyaline.

Size

Attains 267.2 mm total length.

Distribution

Found in the Tigris-Euphrates basin including that part in Iran. Abdoli (2000) has mapped it in the upper Karun, middle and lower Dez, middle and upper Karkheh, Kashkan, Simarreh and lower Gav Masiab rivers. Jawad et al. (2009) record it from the Garaf River of the lower Tigris River basin in Iraq

Zoogeography

The relationships of this species, as with other members of the genus, is presumably with the more diverse fauna to the east.

Habitat

Unknown in detail but it is assumed to be in rocky and gravelly rivers which provide hiding places.

Age and growth

Unknown.

Food

Unknown for Iran but Turkish specimens contained fish remains in the stomach.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

This species is not of any direct economic importance.

Conservation

This species is poorly known in Iran and may be rare enough to warrant conservation efforts.

Further work

See under G. silviae.

Sources

Type material: See above, Glyptosternum armeniacum (ZISP 20780).

Iranian material: None.

Comparative material:- BM(NH) 1974.2.22:1789, 346.6 mm standard length, Iraq (no other locality data); BM(NH) 1968.12.13:465-470, 4, 53.7-76.8 mm standard length, Syria, Euphrates at Mayadine (35º01'N, 40º27'E); SMF 23676, 4, 229.0-297.7 mm standard length, Syria, bei al-Hasaka (36º30'N, 40º44'E); SMF 23677, 2, 65.7-122.0 mm standard length, Syria, Wadi Furati (36º26'N, 40º52'E); ZMH 4430, 2, 129.8-133.3 mm standard length, Turkey, Kemaliye Karasu (no other locality data).

Glyptothorax silviae
Coad, 1981

Ventral head showing sucker

Common names

gorbehmahi-ye jonubi or jonub (= southern catfish).

[Sylvie's sisorid, southern sisorid].

Systematics

The holotype is in the Canadian Museum of Nature, Ottawa under CMNFI 1979-0390A and measures 67.6 mm standard length. It is from "Khuzestan, stream 3 km south of Bagh-e Malek, tributary to Rud-e Zard or Ab-e Ala in the drainage of the Jarrahi River, 31°29'N, 49°54'30"E". Two paratypes from the same locality measure 44.0-51.5 mm standard length and are under CMNFI 1979-0390B, a third paratype measuring 42.3 mm standard length under CMNFI 1979-0389 is from a "stream tributary to Rud-e Zard or Ab-e Ala, 1 km south of Bagh-e Malek, in the drainage of the Jarrahi River, 31°31'N, 49°53'30"E", and a fourth paratype measuring 134.8 mm standard length under CMNFI 1979-0280 is from "Lorestan, river at "Pol-e Chubee" in Kashkan River drainage on Khorramabad to Kermanshah road via Nurabad (either Kaka Reza River at 33°43'N, 48°15'E or its tributary at 33°47'N, 48°12'E)".

Key characters

The head and body dorso-laterally lack striated or elongate tubercles (present in G. kurdistanicus). The thoracic adhesive apparatus is longer than wide (the reverse in G. kurdistanicus). The caudal peduncle is long (4.7-5.2 in standard length, 5.9-6.0 in G. kurdistanicus).

Morphology

Dorsal fin spines are smooth and number 2, branched rays 6; anal fin branched rays 8; pectoral fin branched rays 7-9; pelvic fin branched rays 5; total gill rakers 6-9; retrorse pectoral fin spine teeth 7-16, the number increasing with size of the fish; and total vertebrae 35-38. The adipose fin is long, its length being about equal to the distance between the dorsal fin insertion and the adipose origin (0.9-1.1). The pectoral fin is short and does not extend back to the pelvic fin origin. The caudal peduncle is deep (depth 47-62% of caudal peduncle length). The "sucker" or thoracic adhesive apparatus has pinnate lateral branches and is markedly longer than wide with a wide and long central depression. The head and body are finely papillose, particularly on the ventral surface. Anterior to the adhesive apparatus the ventral head surface is strongly papillose, becoming less developed laterally. The upper lip is much more strongly papillose than the weakly papillose lower lip. The diploid chromosome number is 2n=52 and the karyotype consists of 9 pairs of metacentric, 14 pairs of submetacentric and 3 pairs of subtelocentric chromosomes. The arm number is 98 (Esmaeili et al., 2009).

Sexual dimorphism

Unknown.

Colour

The body is nearly immaculate but in live fish is mottled light lime- green, grey-green, brown or grey. All fins have a central black bar on a salmon-pink, peach or yellow coloured background. The thoracic adhesive apparatus is pink due to an underlying vascular supply. The eye is red.

Preserved fish may have very few scattered brown or black spots dorsally and laterally on an otherwise immaculate body. The overall body colour is brown becoming pale brown or cream on the belly. The base of the caudal fin has a wide black bar separated from a second distal bar by an unpigmented section of the fin rays. The central-most 4 rays of the caudal fin are variably black in the otherwise unpigmented bar. The postero-dorsal and postero-ventral corners of the caudal lobes are not pigmented but the margins of the lobes are black. The central portion of the adipose fin is black with the margins unpigmented in smaller specimens. There is no black pigment on the basal part of this fin. The paired fins and the anal fin are unpigmented distally but become yellowish with fleshy tissue proximally and then brown at their bases. A central band is not well-defined in small specimens. A light patch is found on the back at the dorsal fin origin and at the dorsal fin insertion. The dorsal fin is darkly pigmented and a central black band is apparent although poorly defined. The adult female is generally darker than the smaller male fish in the type series such that the caudal fin bars are not as well defined, The adipose fin is dark brown and the light patches at the dorsal fin are not distinctive. However the bars on the dorsal, anal and paired fins are more obvious.

Size

Reaches 13.5 cm standard length.

Distribution

Known only from rivers draining to the Persian Gulf in southwestern Iran. Abdoli (2000) has also mapped it in the upper Karun and middle to lower Khersan, and middle to lower Dez rivers in the Tigris River basin and in the Mand and Shur rivers of the Gulf basin (the latter tributary to the Dasht-e Palang River). Raissy et al. (2010) record it from the Armand River in Chaharmahal va Bakhtiari Province.

Zoogeography

This species is known from waters in southern Khuzestan and from a single specimen from the Gulf basin. This latter is probably a relict of the late Pleistocene when the Tigris-Euphrates flowed down a drained Gulf receiving tributaries now isolated by the post-Pleistocene rise in sea level (Coad and Krupp, 1983).

Habitat

Their ability to use their sucker for clinging to objects can be seen in plastic jars where small specimens will adhere to the sides out of the water.

Age and growth

Unknown. Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 10 fish measuring 5.60-10.66 cm fork length. The a-value was 0.0164 and the b-value 2.975 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases).

Food

Unknown.

Reproduction

The largest specimen in the type series is a female bearing eggs and was caught on 6 July. The breeding season is probably the summer months.

Parasites and predators

Raissy et al. (2010) found ichthyophthiriasis (infection with Ichthyophthirius multifilis - ich or white spot disease), which cause epizootics in wild and cultured fishes, in fish identified as this species from the Armand River in Chaharmahal va Bakhtiari Province.

Economic importance

This species is not of any direct economic importance.

Conservation

Collections with an electroshocker in 1995 showed this species to be common in the Rud-e Zard.

Further work

The distribution of this and related species should be determined by further field work using electroshocking equipment to extract specimens from under rocks. The validity of the nominal Tigris-Euphrates basin species, including this one, needs examination using large series of adult and young which are not yet available in collections. Variation in critical characters is poorly known because of this shortage of specimens. Molecular and chromosomal techniques may provide additional characters.

Sources

Type material: See above, Glyptothorax silviae (CMNFI 1979-0390A, CMNFI 1979-0390B, CMNFI 1979-0280 and CMNFI 1979-0389).

Iranian material: Type material.

Glyptothorax steindachneri
(Pietschmann, 1913)

Reported from the Tigris River basin at Mosul in Iraq. The two syntypes in the Naturhistorisches Museum Wien have not been located (1997 visit) and the brief description is without figures or details of the thoracic adhesive apparatus. Its validity is in question. Not reported from Iran.

Heteropneustidae

The stinging or airsac catfishes comprise a single genus with four species found naturally from Pakistan through India to Thailand Eschmeyer and Fong, 2011).

The family is characterised by an elongate and compressed body with a flattened head; the mouth is small and transverse with fleshy, papillated lips; villiform teeth present on the jaws and vomer; 4 pairs of barbels present (nasal, maxillary and 2 mandibular); the anterior nostril is tubular and the posterior nostril a slit; gill openings wide and gill membranes free from the isthmus; air sacs are present (see below); swimbladder very small; the dorsal fin is short and spineless; no evident adipose fin; very long anal fin confluent with the caudal or separated from it by a notch; pectoral fin with a strong and venomous spine; skin scaleless; and branchiostegal rays 7.

These fishes can live in stagnant water by breathing air. They are dangerous to man since the pectoral spine harbours a strong venom. Stinging catfishes nonetheless are an important food in the native range.

Genus Heteropneustes
Müller, 1840

The only genus in the family, its characters are given above.

Heteropneustes fossilis
(Bloch, 1794)

Common names

eshlambo or abu shalambo (note variants on this word are used for catfishes and mudskippers); dudeh, doodeh or dood in Khuzestan (= smoke, perhaps because it is blackish); bu shalambo in Khuzestan; گربه ماهي نيش زن (= gorbeh mahi-e nishzan); گربه ماهي هندي (= gorbeh mahi hendi, meaning Indian catfish).

[samaka, abu-al-hukum, abu al-hakim, abu-alhaka, samma, samaka samma, djirri lasseye or jamhoori (latter at Baghdad in reference to the then new republic or jamhooria (F. Kedairy, in litt., 21 December 2005)) in Arabic; singhi in Pakistan; Indian stinging catfish].

Systematics

Silurus fossilis was originally described from India. A syntype of this species is in the Museum für Naturkunde, Universität Humboldt, Berlin under ZMB 3074 (Eschmeyer et al., 1996).

Key characters

The 4 pairs of barbels, short and spineless dorsal fin, absence of an adipose fin, and the long anal fin are distinctive. The head is small and very flattened and tapers both dorsally and ventrally to a terminal mouth.

Morphology

There are two, tubular air sacs extending from the gill cavity almost to the caudal peduncle, enabling this catfish to breathe air. On capture, air from these sacs may escape and cause a peculiar squawking sound. The anatomy and function of these organs was reviewed by Datta Munshi (1993).

Dorsal fin with 6-8 rays but no spine, anal fin rays 60-79, pectoral fin branched rays 7-8 after a strong spine serrated on its inner margin, and pelvic fin branched rays 5-6. Fin rays are difficult to count without dissection or x-rays because of the fleshy and heavily pigmented nature of the fins. Iranian specimens generally fall within the ranges cited above from literature sources as far as can be determined. Spine serrations are more notch-like than toothed. Al-Hassan et al. (1990) have demonstrated that the level of asymmetry in pectoral fin ray and total gill raker counts increases with fish length. Gill rakers are elongate, reaching adjacent raker 5-7 when appressed and number about 25. Barbels are elongate, the snout barbel being the shortest at about head length, the inner mandibular barbel being head length or longer, and the mouth corner and outer mandibular barbel being much longer than the head. The gut is elongate with several posterior coils.

Zakaria (1964) gives details of the pectoral fin spine anatomy. Singhkohli and Goswami (1987) and Kaul and Rishi (1987) describe abnormalities in this species including an upturned tail, a forked tail and forked barbels.

Sexual dimorphism

Unknown.

Colour

Yellow or leaden to dark green, grey-brown, rust-brown or even black, occasionally with two yellowish stripes. The flanks may also be spotted. The barbels are darker than the adjacent body. The eye is yellow. Young specimens are reddish and have a pale belly with numerous melanophores.

Size

Males reach 24.2 cm, females 34.4 cm in India (Datta Munshi and Choudhary, 1996).

Distribution

First recorded from Iraq for 1960 by Khalaf (1961) and Zakaria (1964) when a strange fish was reported to have inflicted a "painful bite" on several victims. The species spread northward and also eastward into Iran from southern Iraq. One collection from Dezful (see below) is dated February 1960 so the spread into Iran must have been very rapid or the original Iraqi introduction some years earlier than documented. It is now known from the Turkish Tigris River near Diyarbakır, presumably dispersing upstream from southern Iraq (Ünlü et al., 2010).

Found from Turkey and Iraq eastward but not continuously through South Asia to Vietnam. It is common in rivers and marshes of Khuzestan including the Dasht-e Azadegan and the Arvand Dez, Karkheh, Karun and Jarrahi rivers (Abdoli, 2000). Berra (2001) omits their Middle Eastern distribution as they are thought to be introduced.

Zoogeography

An Iraqi biologist told me that this species was introduced to Iraq for mosquito control (sic) by local authorities although no one would later admit to it. A more reasonable assumption is that it was introduced to eat the snail Bulinus truncatus, a vector for the human parasite causing schistosomiasis. It was ineffective in this regard (L. A. J. Al-Hassan, in litt., 1995; Jawad, 2003). There has probably been no natural, large scale migration from Pakistan as envisaged by Banister and Clarke (1977) and Banister (1980). Some Sumerian names may refer to this species but this is by no means certain (Sahrhage, 1999).

Habitat

It is common in rivers, marshes, ponds and canals and is found in both fresh and slightly brackish waters. Al-Daham and Bhatti (1977) found this species to suffer a 25% mortality over 72 hours at 10.25‰  sea water. It was most abundant in polluted and stagnant areas in the lower Diyala River where it dominated catches or was the only fish present (Khalaf et al., 1986; Biro et al., 1988). It is common in swamps and can survive temperatures up to 39.8°C (Pethiyagoda, 1991). Specimens survived 3-6 hours in air. It air breathes every 3-5 minutes but the frequency varies with time of day and weather conditions. On hot and calm days it visits the surface more frequently than during a heavy shower. S. Cowton (pers. comm., 23 August 2005) has observed schools of this species gaping at the surface in the artificial lake around Al Faw Palace in Baghdad, presumably in response to high temperatures and low oxygen. Individuals were also seen rapidly swimming straight up to the surface, gulping and diving straight back down again. On especially sultry days it may float or swim near the surface. In the dry season of India it can live in semi-liquid mud or at the bottom of fissures where the mud has cracked (Jayaram, 1980). It makes nest holes in the sides of ponds about 1 foot below the water surface in the form of anastomosing tubes with several exits. Up to 364 fish can be found in one complex of holes (Datta Munshi and Choudhary, 1996). Tekrival and Rao (1999) report its aquarium preferences and habits as 22-25°C, pH 7.5-8.5, alkaline water, predator, not too bright lighting, bottom dwelling with stones, roots and crevices preferred and cave brooding reproduction. Zakaria (1964) noted aquarium specimens producing audible squeaks when excited and preferred the darker side of an aquarium.

Age and growth

Khalaf et al. (1987) gave length-weight relationships for Diyala River, Iraq fish in autumn as W = 9.12 L2.95 (r = 0.98, n = 58) and in spring W = 0.11 L2.15 (r = 0.84, n = 66) and condition factor was 0.0012 in autumn and 0.08 in spring. Islam et al. (1982) give the following length-weight relationships for fish from the Ashar Canal, Basrah, Iraq:- log W = -6.35211 + 3.53226 log TL (r = 0.93543), log W = -5.96765 + 3.42353 log SL (r = 0.93687), and log W = -1.35223 + 2.04705 log GL (r = 0.87876) (TL = total length, SL = standard length, GL = girth length). Tabrez Nasar (1993) studied populations in India and gives length-weight relationships, log W = 1.7661 + 3.035 log L for one population and log W = 1.805 + 2.615log L for another. The coefficient of condition varied from 1.582 to 2.151, mean 1.89. A pond population did not grow as well as a natural one. Life span was up to 4 years in Iraq at Qarmat Ali using ocular lens diameter and vertebral rings (Al-Hassan et al., 1992) and may be 4+ years in India (Datta Munshi and Choudhary, 1996).

Food

Al-Daham et al. (1977) studied the diel feeding of this species in the Ashar Canal, Basrah, Iraq. Two feeding peaks were observed - at 0500 hours and 1700 hours, dawn and dusk, but stomachs examined around the clock had food in them. Aquatic plants and detritus are the bulk of the diet, followed by entomostracans and aquatic insect larvae. Also present are fish parts, molluscs and non-aquatic organisms. Cannibalism is reported in India for young fish (Jayaram, 1980). Khalaf et al. (1987) studied this species in the Diyala River, Iraq and found young fish to take chironomids and worms while larger ones ate fish. However all sizes take aquatic insects in spring. There is some competition with Barbus (= Mesopotamichthys) sharpeyi, a commercial species (Jawad, 2003). In a study of the recovering Hawr al Hammar, diet was 47.2% insects, 22.1% shrimps and 20.8% fish, in the Hawr al Hawizah 33.9% shrimps, 25.8% fish, 20.8% insects and 19.2% snails and in the Al Kaba'ish (= Chabaish) Marsh 51.2% shrimps, 26.4% fish, 12.0% insects and 10.4% snails (Hussain et al., 2006).

Chili macaroni, corned beef casserole, mixed vegetables and salad are dietary items at Camp Liberty, a former palace in Baghdad, where American soldiers feed leftovers from a distinguished visitors dining hall to catfishes, apparently this species (www.estripes.com, downloaded 7 September 2006).

Small Iranian specimens contained insects in their guts including Notonectidae and Diptera larvae. Abdoli (2000) lists diatoms, Chlorophyceaae, fish remains, Corixidae, Hemiptera, cyclopoid Copepoda, termites, Isopoda, Chironomidae, Oligochaeta and Rotifera.

Reproduction

Sexual maturity in India is reached when fish are about 1 year old, at 8.5 cm for males and 12.0 cm for females. Fecundity reaches 12,000 eggs (Haniffa et al., 2008). Al-Rudainy (2008) reported sexual maturity at 2-3 years, 12 cm length and 40 g weight in Iraq. Spawning there took place continuously from June to August with eggs deposited in batches on vegetation. Females take care of the fry.

Eggs are laid in a shallow depression excavated by both the male and female in mud or sand. Eggs are light green. They hatch in about 2 days in Sri Lanka. The parents guard the eggs and young until the young fish are about a month old and able to look after themselves. Singh Kohli and Goswami (1987) describe spawning behaviour in aquaria after hypophysation using pituitary glands of Indian major carps. A pair of males circled each other in a figure 8 pattern until one established dominance. The dominant male chased the female, swimming underneath her or obstructing her path, and touching barbels. The male tried to bite the female in the chases and shivered its whole body while making lateral passes. The male arcs its body into a u-shape, the female touches the male's genital papilla and the pair remain motionless for 2-5 seconds. The female jerks and separates from the male releasing the eggs which settle to the bottom. The pair rest before mating agian. About 40-200 eggs are released after each mating. Mating acts number 20-100 and always occur near the surface of the aquarium. Spawning is more complete and egg fertilisation is better when there is one female and two males, the other male acting as a stimulator, with the spawning male quarrelling with the non-spawning male between mating with the female. Datta Munshi and Choudhary (1996) report similar behaviour. The male nudges the genital region of the female with his head, occasionally shaking it from side to side. Eventually the female is aroused and nudges the male genital region. This female action was necessary for mating to occur. Mating did not happen when spawners were of different sizes. Once the female has her snout below the male genitalia, the male twists his body to place his snout below her genitalia. The fish remain motionless for 2 seconds, then the male vibrates his body and the female convulses and releases eggs. This can happen 30-50 times for each couple at 2-3 minute and then later 5-10 minutes or longer intervals. About 100-150 eggs are extruded, the number decreasing over time. If more than one male is present, mating only occurs after one establishes dominance. Males may eat eggs. Spawners mate in the water column or near the surface.

Parasites and predators

Jalali et al. (2005) summarise the occurrence of Gyrodactylus species in Iran and record G. fossilis in fish from the Karun River.

Economic importance

An important food fish in India and Sri Lanka, where its flesh is reported to have invigorating qualities. Some fish are exported from Sri Lanka for the aquarium trade. V. D. Vladykov reported (in litt., 22 July 1963) that he had seen this species in pet shops in Tehran, on sale at about $2.00 each.

The pectoral spine can cause a serious wound because of the toxin content of the epidermal cells covering the spine. The histology of the pectoral spines was described by Bhimachar (1944) based on Indian material and the toxin was found to have both neurotoxic and haemolytic effects. The toxin is fatal to frogs (within 15-20 minutes of subcutaneous injection of glycerinated venom) and to other fishes.

Zakaria (1964) reported severe swelling involving the whole arm from a hand sting in Iraq. The swelling and pain recede after about a day but the puncture wound can take about two weeks to heal and some pain can be felt when applying pressure to the wound site up to six weeks later. Caras (1964) (probably based on a report in Farsi in Game and Nature, Tehran, ca. 1961) recorded a diminutive black fish found in the Shatt al Arab which reputedly killed 28 people with a venomous bite (sic). Death was said to be swift. This was presumably a garbled report on this species. Verbal and newspaper reports from Tehran (V. D. Vladykov, in litt., 26 August 1961) maintain that this species could cause death to cattle and humans although Vladykov (in litt., 30 September 1963) considered fatal cases "not well proved". I was stung in the thumb by this fish in Iran with no effect (although I did devote considerable time and effort into squeezing and sucking blood from the puncture site!). Freshly caught or netted fish swing the head from side to side and thus are active envenomators (despite knowing this I was still stung). Treatment is symptomatic and some relief can be obtained by immersing the sting site in water as hot as can be withstood and applying a meat tenderiser. These treatments serve to coagulate the protein toxin. The wound should be cleaned to avoid secondary infections such as tetanus (Halstead, 1967-1970; Coad, 1979b).

R. Beck (pers. comm., 2000) reports that this fish is now present in the Syrian Euphrates, its tributaries, and in irrigation canals. Incredible numbers occur near town sewage outlets and in weed beds. It is known to consume eggs of Tor grypus, a preferred food species

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in aquaculture, as food, in textbooks and because of its venomous nature.

Conservation

This is an exotic species and requires no conservation although it is is listed as Vulnerable in its natural habitats (Haniffa et al., 2008a).

Further work

The distribution of this species should be mapped as it is potentially hazardous to humans.

Sources

Iranian material:- CMNFI 1979-0087, 1, 189.0 mm standard length, Khuzestan, Karun River at Ahvaz (31º19'N, 48º42'E), CMNFI 1979-0359, 5, 96.9-114.2 mm standard length, Khuzestan, Karkheh River at Hamidiyeh (31º29'N, 48º26'E); CMNFI 1980-0909, 4, 113.2-165.0 mm standard length, ? Khuzestan (no other locality data); BM(NH) 1980.8.28:4-5, 2, 91.7-98.0 mm standard length, Khuzestan, Dezful (32º23'N, 48º24'E); ZSM 27369, 3, 111.9-139.0 mm standard length, Khuzestan, Karun River near Ahvaz (ca. 31º19'N, ca. 48º42'E).

Comparative material: BM(NH) 1962.7.26:80-83, 4, 127.8-189.0, Iraq, Baghdad (33º21'N, 44º25'E); BM(NH) 1974.2.22:1785, 1, 89.8 mm standard length, Iraq (no other locality data); BM(NH) 1974.2.22:1786-1788, 3, 102.3-165.3 mm standard length, Iraq (no other locality data); ZSM 19455-56, 2, 123.2-127.7 mm standard length, Iraq, Tigris River near Amara (ca. 31º43'N, ca. 47º06'E).

Salmonidae

This family comprises the salmons, trouts, charrs and whitefishes and contains 11 genera and about 213 species found in cooler water in the Northern Hemisphere (Nelson, 2006; Eschmeyer and Fong, 2011). The distribution map in Berra (2001) is too extensive in the south of Iran. There are only two species native to Iran but several others have been introduced with varying degrees of success. Whitefishes may be placed in their own family, Coregonidae, but views differ (see Nelson, 1994; 2006).

Members of this family are characterised by numerous, deeply embedded scales on the body but not the head; an adipose fin; a lateral line with relatively numerous and often quite small scales; 7-20 branchiostegal rays; 3 upturned vertebrae at the tail fin; a large swimbladder; usually numerous pyloric caeca (11-210); gill membranes free from the isthmus; a small dorsal fin with few rays (less than 17); a pelvic axillary process; young usually with parr marks (bars along the flank); and a tetraploid karyotype.

Family members are important in fish culture in all the cooler waters world-wide.

Genus Coregonus
Linnaeus, 1758

This genus is characterised by small, almost toothless mouth, 115 or fewer scales in the lateral line, gill rakers often long and slender, there is a single flap between the nostrils, and no bright and complex colour patterns in adults and no parr marks in young, forked caudal fin. Numerous named species in North America and Eurasia (FishBase gives 217 in Coregonus alone, August 2007), variously recognised as full species or not.

Coregonus lavaretus
(Linnaeus, 1758)

Common names

safid mahi (= white fish at Karaj Dam), safid mahi juibarye (= brook or rivulet white fish), koregon or coregon.

[European whitefish, powan, houting, skelly, gwyniad, lavaret, pollan].

Systematics

Salmo lavaretus was originally described from Lake Bourget, France (Kottelat, 1997). There are numerous subspecies and infrasubspecific names for populations of this fish in northern Europe (Berg, 1948-1949). The origin of the Iranian specimens is unknown so it is not clear which subspecies or species they belong to and accordingly information on biology is of a general nature.

Key characters

This species is the only one in its genus reported from Iran and is recognisable by the generic characters above.

Morphology

The mouth is small, inferior and lacks evident teeth. Lateral line scales are 84-105. The dorsal fin has 3-5 unbranched and 8-13 branched rays, the anal fin has 3-5 unbranched and 11-15 branched rays and the caudal fin has a strong fork. Gill rakers are sparse and short or long and dense, denticulated or not, and number 21-56, with means and form varying between subspecies. Vertebrae number 56-64. The chromosome number is 2n=80 (Klinkhardt et al., 1995).

Sexual dimorphism

Males may bear rows of elongate tubercles on the scales.

Colour

The back and upper head is a dark bluish-grey, blue-green or dark green, flanks are greenish-grey to silvery and the belly is silvery to yellowish-white. Fine speckles may be apparent on the body. The snout is black. The iris is white. The dorsal, anal, adipose and caudal fins are dark while the pectoral and pelvic fins are only dark at the tips.

Size

Attains 97.0 cm and 10.0 kg.

Distribution

The native distribution of this species is in northern Europe and Siberia, in particular the drainages of the Baltic, North, Barents and White seas with isolated populations in central Europe in mountainous areas.

Fingerlings were released into the Karaj and Latian reservoirs near Tehran from 1965-1968 after hatching from eggs imported from Europe (Armantrout, 1980). Walczak (1972) reports some existing still in the Karaj Reservoir but none were found in the Latian. Ahmadi et al. (2011) record this species as still present in the Karaj (= Amir Kabir) Dam reservoir, the only locality for it in Iran. It was also introduced to the Manjil Dam on the Safid River (Griffiths et al., 1972) but this reservoir is drained to remove excess silt and no fishery exists (J. Holčík, pers. comm., 1992). There is no evidence of reproduction. Saadati (1977) states that they were established in both reservoirs. Also reported from the Farahnaz Reservoir, Markazi. Abdoli (2000) depicts the Karaj River and the Abhar River as harbouring this species.

Zoogeography

An exotic species not naturally occurring in Iran.

Habitat

This species is typically an inhabitant of large and deep lakes where its oxygen requirments are quite high. However it is tolerant of warm water and even a measure of pollution. Spawning may occur in rivers tributary to the lake. Anadromous forms occur but are rare in full seawater.

In lakes during the day, it is found at depths of 20-30 m or on the bottom if water is shallower. At night it may rise as far as the surface following the migrating plankton or to the waters edge in the shallows. Schools form in sub-littoral areas during the spawning season and strandings may occur.

Age and growth

Maximum age attained is 20+ years although in some populations most fish are 1+ to 3+ years old while in others most fish are 7+ to 10+ years old. Under good conditions young can attain 10-12 cm after one year and 15-20 cm after two years of life. Maturity is attained at 3-5 years and 25-35 cm.

Food

Plankton is the principal food item, with benthic crustaceans being taken in brackish water and crustaceans, molluscs and insect larvae in rivers. This species cannibalises its own eggs and eats the eggs of other fishes. Feeding may vary through the year, planktonic crustaceans being taken in summer and benthic invertebrates in winter.

Reproduction

There is a spawning migration which may occur in summer but more usually peaks in autumn. Spawning takes place in summer, autumn or winter, varying with the subspecies or form. Summer spawning takes place in deeper water than winter spawning. In lakes, the adult males enter the spawning grounds at dusk, these grounds being gravelly shoals off headlands or on offshore reefs. Some populations spawn over sand or even mud. Females move in each night as they ripen. The yellow eggs are 2-3 mm in diameter and slightly adhesive. They stick to gravel and are protected from predators by falling in crevices between the gravel. Up to 82,250 eggs may be laid. Eggs incubate for 90-100 days during winter at an optimum temperature of 6°C or less.

Parasites and predators

None reported from Iran.

Economic importance

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in aquaculture and as food in Europe and Russia. It has been widely introduced to reservoirs. Ahmadi et al. (2011) collected fry from the Karaj Dam and fed them in rearing cages with three diets, rotifers (Brachionus plicatilis), a commercial salmonid diet, and a an equal mixture of both. Total feeding rate had a profound positive impact on survival for all three treatments. The mixed feed had an easier prediction of survival rate perhaps because it better satisfied nutritional needs.

Conservation

No conservation of this exotic species is advisable.

Further work

The survival of this species in Iran should be verified.

Sources

Based on general European literature.

Iranian material: None.

Comparative material: BM(NH) 1986.11.14:1-3, 3, ca. 312-330 mm standard length, England, Cumbria, Lake Haweswater (no other locality data).

Genus Oncorhynchus
Suckley, 1861

Members of this genus are found naturally in the Northern Pacific Ocean and its drainages, migrating between fresh and salt water. Some live permanently in fresh water. There are about 12 species and two have been successfully introduced to Iran.

This genus is characterised by a large mouth, in adults extending back to the level of the posterior eye margin or further, scales are small, ca. 100 or more in the lateral line, lateral line scales are as large or larger than scales in adjacent rows and overlap with scales in front and behind, evident teeth in adults, teeth are present on the head and shaft of the vomer bone in the roof of the mouth, usually dark parr marks in young but adults without snow-white leading edge to lower fins, and usually black spots on the body or caudal fin or both.

These salmons spawn once and then die. They are important commercially as food and as sport fishes.

Oncorhynchus gorbuscha
(Walbaum, 1792)

Introduced to the former Soviet Caspian Sea basin in 1963 but not observed subsequently (Baltz, 1991). No Iranian record.

Oncorhynchus keta
(Walbaum, 1792)

Common names

azad mahi keta (= keta free fish or keta salmon, azad mahi being the Farsi for trout or salmon), mahi-ye azad-e chum.

[chum salmon, dog salmon, keta, summer chum].

Systematics

Salmo keta was originally described from rivers of Kamchatka, Russia.

Key characters

This species is distinguished by having 124-153 lateral line scales, 13-17 principal anal fin rays, flanks and caudal fin without distinct black spots, short, stout gill rakers 18-26 and pyloric caeca 135-249.

Morphology

Dorsal fin with 10-14 principal rays, pectoral rays 14-16 and pelvic rays 10-11. Scales are horizontal ovals with a central focus and no radii. Circuli are fine but are lost or only partly developed on the flesh-covered exposed part of the scale. The gill rakers reach just beyond the first to just beyond the second raker when appressed. The pelvic axillary scale is very elongate. The gut is s-shaped. The chromosome number is 2n=74 (Klinkhardt et al., 1995).

Sexual dimorphism

Breeding males develop hooked jaws and large teeth and a slight hump in front of the dorsal fin.

Colour

Marine fish are steel-blue on the back and upper flank, silvery on the flank and silvery-white on the belly. The upper flank and back may have fine black speckles but no spots. The pectoral, pelvic, anal and caudal fins have dark edges. Spawning males in fresh water are dark olive to black above, greyish-red to brick-red on the flank with greenish to purplish bars or blotches and a dark grey belly. Calico is the characteristic colour of dominant males. The anal and pelvic fins are often tipped with white. Females are less strongly marked. Young chum are iridescent, mottled green on the back and silvery iridescent green on the flanks and belly. There are 6-14 parr marks which do not descend much below the lateral line and are narrower than the space between them. Fins are clear to white.

Size

Reaches 120.0 cm and 20.8 kg.

Distribution

Found from Alaska to California on the Pacific coast of North America and also in eastern Arctic Siberia, the Beaufort Sea and south to the Sea of Japan, the widest distribution of any Pacific salmon.

This species was introduced to the former Soviet Caspian Sea in 1962-1970 in an attempt to offset losses of Salmo caspius which was cut off from its spawning grounds by dams. It was considered as ideal because it spawns in the lower reaches of rivers and dams would not affect its spawning migration, it returns to spawn after 2 or 3 years and so some year-classes could escape detrimental conditions in any one year, and because it produces downstream migrants at a smaller size and age than the native salmonid (Magomedov, 1970; 1978; Baltz, 1991). 7.5 million fertilised eggs were transported from the Amur River and Sakhalin Island in the former Soviet Far East and incubated at the Samur Fish Farm, Dagestan. 5,850,000 fry were released.

Walczak (1972) reported capture of a specimen in Iranian waters in 1971, presumably a stray from the former Soviet stocking programmes. Holčík and Razavi (1992) record only two specimens from Iran, one in 1964 and one in 1972, taken between Bandar Anzali and the Astara River on the Iranian coast of the Caspian Sea. They also seem to indicate another specimen taken after 1972. Jolodar and Abdoli (2004) report it from Astara and the Anzali regions, indicating no large scale spread.

Zoogeography

A species introduced to Iranian waters by man.

Habitat

In North America, chum salmon enter streams on the spawning migration and often travel less than 150 km, stopping at the first barrier as they are not strong jumpers. Some fish even spawn in tidal areas. However some rivers, like the Yukon, have a run which travels about 3200 km and takes from early June to the end of September. A migratory speed of up to 115 km/day has been recorded with bursts of speed to 4.6 m/s. Most runs are in the fall but some are in the summer. Very rarely a chum will become trapped in a lake if an outlet stream dries up but normally adult chum are only found in fresh waters on the spawning run.

Age and growth

Reproduction occurs at ages 1+ to 2+, with spawning fish at 2+ predominating in Caspian populations from former Soviet waters. In their native streams in the Far East fish are in their fourth and fifth years of life. 1500 adults were caught returning to one river to spawn in the Caspian in 1966, considered impressive by former Soviet biologists. Accelerated growth and maturity in transplanted fish is well documented (Magomedov, 1970). There is a 15C° difference between the Caspian and native water temperatures for spawning and egg development. Life span in Canada is about 7 years, perhaps 9 years, with fish in British Columbia maturing at 1-6 years with age 3 fish dominant. They spend 2-7 years at sea before returning to spawn. Males grow faster and larger than females.

Food

Food in the sea includes crustaceans, worms, molluscs, squids, jellyfish and fishes. Young fish in the sea take zooplankton. Adults on the spawning run do not feed. Young fish in fresh water eat aquatic insects, such as chironomids, mayflies and caddisflies, as well as crustaceans, worms and terrestrial insects.

Reproduction

Transplantations into the former Soviet Caspian Sea basin were not very successful because of a lack of suitable spawning streams (McNeil, 1979). Spawning runs into some former Soviet streams were recorded from the first half of September to the end of October, the same periods as in the native habitat (Magomedov, 1970). Males arrived first, the mass run was composed more of females and there were more males at the end of the run. Since a single male will spawn with several females, early male arrival on the spawning grounds may promote successful fertilisation. The absolute fecundity of a 3+ salmon was 2739 eggs in the Caspian.

Females excavate a redd by lying on their sides and lashing the tail. The redd is a trough up to about half a metre deep and up to 3.2 m long and 2.1 m wide bordered by a ridge of gravel at the downstream end. In some cases no redd is excavated and eggs are shed over and between boulders. Females may excavate more than one redd and males may spawn with more than one female. A female and one dominant male lie in the redd, gape their mouths, vibrate and release eggs and perm. The dominant male may have several accessory males accompanying him. The female dislodges gravel at the upstream end of the redd to cover the eggs. The orange eggs are up to 7.8 mm in diameter (perhaps 9.5 mm when fertilised) and each female may shed up to 7779 in the Pacific basin. The adult fish die after spawning and may live only a week after first entering fresh water. The fry emerge from the gravel in March-May in North America, some remaining for several weeks in fresh water or immediately migrating to the sea.

Parasites and predators

Various insects, fish, birds and mammals prey on both young and adult chum.

Economic importance

It is too rare to have any economic importance in Iran. Total catches in the North Pacific Ocean have been as high as 69.2 million fish annually. In Japan it is very important in an ocean ranch industry.

Conservation

The presence of this species in Iranian waters is the result of former Soviet attempts to acclimatise it to the Caspian Sea as a potential replacement for declining stocks of Salmo caspius. The stocking programme lasted from 1962 to 1979 and rivers along the Dagestan coast showed mass spawning but this became rare. Pollution by wood from forestry operations in Dagestan, absence of suitable gravel beds, low salinity in the Caspian Sea and heavy surf on the Dagestan coast may be responsible for the heavy mortality (Magomedov, 1978; Holčík and Razavi, 1992). As an exotic species, there is no need for conservation of chum salmon.

Further work

Reports of this species in Iranian waters should be documented.

Sources

Bakkala (1970) summarised the biology of this species in North American waters and Salo in Groot and Margolis (1991) over its whole range.

Iranian material: None.

Oncorhynchus kisutch
(Walbaum, 1792)

Introduced to the Caspian Sea basin but not subsequently observed and no Iranian records (Holčík and Razavi, 1992).

Oncorhynchus mykiss
(Walbaum, 1792)

Lorestan fish farm specimen, north of Aleshtar (golden form), 3 December 2000,
Lorestan fish farm specimen, north of Aleshtar (golden form), 3 December 2000,
photograph Brian W. Coad

Duane Raver, U.S. Fish and Wildlife Service
Duane Raver, U.S. Fish and Wildlife Service

Common names

قزل آل (= gazalala, ghezel ala, qezel ala or kizil ala; probably in confusion with native Salmo caspius, these words meaning literally red spots or red spot fish but used for trout and salmon species in Iran),
قزل آلاي رنگين كمان (qezel ala-ye ranginkhaman, meaning rainbow trout).

[mykiss in Russian; rainbow trout, steelhead trout].

Systematicss

Salmo mykiss was originally described from Kamchatka, Russia.

Formerly known as Salmo gairdneri Richardson, 1836 (see Bailey and Robins, 1988; Smith and Stearley, 1989). Placed in the genus Parasalmo Vladykov, 1972 by Reshetnikov et al. (1997) and Mednikov et al. (1999) but Osinov (1999) has reservations since allozyme data show specific level differences with Oncorhynchus although further studies were deemed necessary.

Key characters

This species is distinguished by having 100-161 lateral line scales, 8-12 principal anal fin rays, the vomer bone in the roof of the mouth has teeth on its head and shaft, no red spots on the body but only small dark ones and radiating rows of black spots on the dorsal and caudal fins, and no teeth at the tongue base.

Morphology

Dorsal fin principal rays 10-12, pectoral rays 11-17 and pelvic rays 9-10. Gill rakers 16-22. Pyloric caeca 27-80. The chromosome number is 2n=58-62 (Klinkhardt et al., 1995).

Sexual dimorphism

Breeding males have an elongated snout, the lower jaw is hooked and the roof of the mouth is white.

Colour

Overall colour is very variable. Stream fish are darker and more colourful (rainbows) than lighter, silvery lake or sea fish (steelhead). Some sea-run and lake fish have small orange to red marks below the lower jaw. The back and upper flank are steel-blue, greenish, silvery-olive or even brown, flanks and belly are silvery, grey, white or yellow-green. The side of the head and the flank are characteristically pink. The flank has a broad pink to red or lilac stripe with small black spots. The adipose fin has a black margin and a few spots. Pectoral, pelvic and anal fins may have a few spots and are dusky without any strong markings. Pectoral and pelvic fins are often orange-red. Spawning fish are very dark and the flank stripe is dark red or purple. The young have 5-13 dark, oval parr marks centred on the lateral line with the spaces between the marks wider than the marks. There are 5-10 parr marks on the back in front of the dorsal fin. The upper flank has some dark spots. The dorsal fin is tipped white or orange and has a dark leading edge, sometimes broken up into spots. The adipose fin has a black margin. The anal fin has a white or orange tip. Some adults in streams do not lose their parr marks. A golden form occurs rarely on fish farms in Lorestan and elsewhere is farmed specifically.

Sizee

Reaches 122.0 cm, possibly 150 cm, and 35.0 kg as sea-run or lake fish but smaller in streams.

Distribution

Found in western Canada and from Japan and Alaska to Mexico. Widely introduced outside this natural range world-wide in suitable waters. They have been introduced to the former Soviet part of the Caspian Sea in 1973 and several hundred returning adults were reported in 1975 (McNeil, 1979; Baltz, 1991).

Rainbow trout were introduced to Iran about 1966 for hatchery production as a commercial product (MacCrimmon, 1971; 1972). They are now widespread in Iran, stocks being maintained by hatchery introductions and sometimes natural reproduction, wherever temperature regimes are suitable in the higher reaches of the Alborz and Zagros mountains (Walczak, 1972; Anonymous, 1977; B. Sandford, in litt., 1979; Y. Keivany, in litt., 1992; Ghorbani Chafi, 2000; personal visits to fish farms).

B. Sandford considered few populations became established because of competition with large populations of omnivorous cyprinids which also take fish eggs, presumably of trout too. Sandford cited viable populations in the Madar Su, a small stream in the former Mohammad Reza Shah Park (see below), Tar Lake in the Alborz (also recorded from Tar Lake by Riahi (1996)), Gahar Lake in the Zagros and the Qara Chai west of Hamadan. Introductions to Neuer or Neur Lake near Ardabil, Ghorighol Lake, Rebeshahr and the Ab-e Bazuft were all failures (although Walczak (1972), Saadati (1977) and R. Mehrani (pers. comm., 2000) report several successful stockings including the Ab-e Bazuft, Gahar Lake, Namrud, Dez River and Jajehrud).

Also reported from the Gorgan and Haraz rivers and Gorgan Bay (the latter escapees from cages)(Kiabi et al., 1999), Madar Su, Golestan National Park (Kiabi et al., 1994), from the Farahnaz Reservoir, Markazi, Namrud in Semnan, Golestan River, Karaj River, Amir Kabir Dam (www.iran-doe.org/Special/Alborz.htm), Shah Abbas Dam west of Esfahan, Darius-e Kabir Dam north of Shiraz, Karaj River, Zayandeh near Esfahan, Rebeshahr southeast of Yasuj (Anonymous, 1977; Y. Keivany, in litt., 1992). An attempt to culture this species in cages in the Avan River, 7 km from Alamut, in 1994 failed and fish escaped (Nialmir, 2001). Also in the Sardab Rud (Jalali et al., 2005).

An attempt to establish a run of steelhead, the migratory form, in the Chalus River from the Caspian Sea was apparently unsuccessful (Armantrout, 1980).

This species was stocked in the 1970s by the Iranian Department of the Environment in the Doogh or Madar Su, head stream of the Gorgan River in Golestan National Park and "Neur" Lake in East Azarbayjan near Ardabil in the Caspian Sea basin; in Ghorigol Lake (east of Tabriz) and the Liqvan Chay in the Lake Orumiyeh basin; the Karaj River, the Jajrud, the Namrud, Tar Lake northeast of Tehran, Lake Lasem near Tehran, the Lar River, probably in the Chashmeh Do Barare, Ab Kharsang, Varangarud and Baragon near Tehran, all in the Namak Lake basin; the Karun River, Mohammad Reza Shah Dam, Rebeshahr River southeast of Yasuj, the Ab-e Bazuft in 1975 (established), Gahar Lake in the headwaters of the Dez River and its outlet stream (established, caught by A. Abdoli in 1995), at farms in the Rostamabad area discharging into the Kuhrang stream of the Karun River basin, and an isolated section of the Dez River formerly fishless (established), all in the Tigris River basin; the Shah Abbas Dam west of Esfahan, in the Zianrud (presumably the Zayandeh River of the Esfahan basin) where a reproducing population was established; in the Dorudzan (Dariush-e Kabir) Dam near Shiraz (104,000 in Esfand 1350)(Surber, 1969; Anonymous, 1977; Armantrout, 1980; Petr, 1987; Abdoli, 1993b; Fadaeifard et al., 2012).

Qanats in Markazi Province were stocked with 930,000 trout (presumably this species) in 2006 (www.iranfisheries.net, downloaded 28 July 2006).The qanats were in Shazand (2 qanats), Arak (4), Delijan (1) and Khomein (1).

Abdoli (2000) lists the Dasht-e Kavir and Kerman-Na'in basins generally, the upper Kashaf River in the Tedzhen River basin, the Gorgan, lower Neka, middle and lower Babol, and Heraz rivers in the Caspian Sea basin; the upper Talkheh, Tatavi and Zarrineh rivers in the Lake Orumiyeh basin, the Abhar stretch of the Shur and the upper and middle Karaj rivers of the Namak Lake basin; and the Khersan, upper Dez and Kashkan rivers of the Tigris River basin, and the upper and middle Zayandeh River of the Esfahan basin.

Zoogeography

An exotic species introduced as a food fish.

Habitat

Rainbows are found in rivers or streams where there are pools and riffles. Some live in lakes and are called Kamloops trout in Canada while others run to sea and are called steelheads. They can tolerate temperatures up to 24°C, warm for a trout, but prefer temperatures below 20°C. Sea-run fish spend about 1-4 years usually in inshore waters at middle to surface depths after 1-4 years in fresh water. Some fish (half-pounders) return to streams after a few months at sea. Summer steelhead have spent one winter at sea and return in summer (April-October) to spawn next spring while winter steelhead are larger and return from December to April peaking in January to spawn in March and April. Aquaculture of trout in Iran is affected by drought conditions reducing water flow to farms. Production was higher in 1999 compared to the previous year but this was because 82 new farms were opened; average production per farm fell (Foghi, 2004).

Age and growth

Life span varies with habitat, up to 11 years in some lake fish but only 3-4 years in many streams and small lakes. Growth varies with habitat including such factors as length of sea, stream or lake life, years before spawning, available food supplies, latitude, altitude, temperature regime, competition with other salmonids, and so on. Ageing these fish may be difficult because of the complicated life history pattern of stream and lake residency. Maturity is also variable with habitat. Some males mature at 9 months in fish introduced to warm southern waters and some females only at 8 years, but generally maturity is reached at 3-5 years in Canada, for example, with males maturing a year earlier than females. Neur Lake fish had a growth rate of 17 lbs (7.72 kg) in 3 years, more than doubling in size, because of the excellent food supply of shrimp. In Tar Lake, where food resources were poor, mostly surface insects, fish grew to only 3-5 lbs (1.4-2.3 kg) in 2 years (Anonymous, 1977). In the Karaj Reservoir, one-year-old fish with a length of 8-10 cm grew to 25 cm in one year although the available food was only plankton.

Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 111 Iranian fish measuring 27.5-57.0 cm standard length. The a-value was 0.0161 and the b-value 3.044 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases).

Food

Food includes plankton, crustaceans, aquatic and terrestrial insects, snails, leeches, salmon eggs, and other fishes. The fish eaten enhance growth and the species taken depends on what is available. In Lorestan, this species thrives in rivers where Gammarus is numerous, e.g. near Dow Ab-e Aleshtar. Fish and squid are taken at sea along with some crustaceans. Abdoli (2000) lists Trichoptera, Plecoptera, Chironomidae, Ephemeroptera, Ceratopogonidae, Odonata, Simuliidae, Coleoptera, Decapoda and Amphipoda as food in Iran.

Reproduction

Spawning takes place from March to August but is usually in spring. North American Great Lakes fish may spawn from late December to late April. Water temperature for spawning usually exceeds 10°C but may be 5-13°C. A female excavates a redd by lying on her side and thrashing her tail. Redd excavation occurs during the day and night and dimensions are usually longer and deeper than the female's body. A male courts a female by rubbing his snout and body against her, by vibrating, by swimming over her in the redd and by pressing against her. Several males are found around each female but one male is dominant. The spawning act lasts 5-8 seconds with the pair parallel in the redd pressed together, both fish gape, arch and vibrate. Other males may shed sperm. The female covers the eggs with gravel by dislodging it from the upstream end of the redd. Most spawning takes place in the morning and evening and nests may be abandoned during the day. Females construct several redds and may spawn with several males. Eggs are orange or pink, 5.0 mm in diameter and up to 12,749 in number. However, egg numbers are usually a few hundreds to thousands per female. The eggs hatch in about 8 weeks and fry generally emerge in June to August from spring spawnings. Repeat spawning can occur for up to 5 years.

Nematollahi and Azari Takami (2002) studied the quality and quantity of seminal fluid in cultured male broodstock specimens of this species at the Karaj Fish Farm. Differences were found between stocks in sperm motility and concentration.

Parasites and predators

A wide variety of other fishes and birds feed on this species and there have been extensive studies on parasites and diseases of this commercially important species. Akhlagi (1999, www.mondialvet99.com, downloaded 31 May 2000) reports infectious pancreatic and haemotopoitic necrosis in fish from Bovir Ahmadi va Kohkiluyeh Province but not in Fars Province. Liver and intestinal submucosa degeneration and kidney necrosis were observed in diseased fish. Prearo and Ghittino (1993) record a case of lipid liver degeneration in cultured trout. Asadzadeh Mangili (2000) records ichthyobodosis in 3 cm fry from a farm in Bovir Ahmadi va Kohkiluyeh. The mortality rate was 40% and infection rate with Ichthyobodo was 100%. Rainbow trout fed on raw marine fishmeal powder in Fars Province were found to be more exposed to vibiosis (pathogenic Vibrio anguillarum)(Ghazi and Akhlaghi, 1998). Diplostomiasis (infection with Diplostomum spathaceum) is reported from cultured trout in West Azarbayjan (Asadzadeh Manjili and Ghorbanzad, 1999). Zorriehzahra (2002) reports on diseases in farmed fry and Zorriehzahra et al. (2002) record enteric redmouth disease in farmed fish around Tehran. Akhondzadeh et al. (2002) and Akhondzadeh Basti and Zahrae Salehi (2003) show that the psychotropic pathogen Listeria monocytogenes is found in market and fish farm samples of this species. Soltani and Rostami (1997) record a mortality level of about 16% for farmed trout in northern Iran infected with a Cytophaga/Flexibacter-like bacterium. Soltani et al. 1999) report a yersinosis-like infection in farmed Iranian rainbow trout. Jalali et al. (2005) summarise the occurrence of Gyrodactylus species in Iran and record G. derjavini from fish in the Sardab-rud. Asadzadeh Mangili (2001) describes gas bubble disease from fish farms in Kermanshah Province. Mortality occurred mostly in fish at 100-200g and symptoms included inappetition, skin darkening, aggregation of fish at inlets and outlets in the ponds, and swimming near the surface. In 20% of the samples, unilateral or bilateral exophthalmia was observed. High day-night fluctuations in the oxygen and carbon dioxide content of the pond water was observed through excessive plant growth. Abdi et al. (2006) recorded the first mortality from acanthocephalosis with Pomphorhynchus laevis in fish farms in Maku, West Azarbayjan. Ebrahimzadeh Mousavi et al. (2007) isolated 12 species of fungi from eggs in Mazandaran hatcheries. Barzegar et al. (2008) record eye parasites from this fish including the protozoan Ichthyophthirius multifilis and the digenean Diplostomum spathaceum. Miar et al. (2008) examined fish in Valasht Lake and the Chalus River, Mazandaran and found the protozoans Trichodina trutta, Ichthyophthirius multifilis and Chilodonella hexastica as well as the metazoan Gyrodactylus derjavini. Khosravi et al. (2010) found 5 species of Saprolegniaceae fungi on eggs in two hatcheries in Kermanshah Province, three species for the first time in Iran. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Lernaea sp. on this species. Mood et al. (2011) record Ichthyophthirius multifilis, Trichodina sp., Chilodonella sp., Gyrodactylus sp. and Dactylogyrus sp. in fish from coldwater fish farms of Mazandaran.

Economic importance

This species is commercially farmed in Iran for local sale. Experimental studies in the 220 ha "Neur", "Neuer" or "Noor" Lake near Ardabil have shown very good production of this species. A growth rate of 17 lbs (7.72 kg) in 3 years was reported (Anonymous, 1977). A yield of 160 kg/ha was obtained and a catch of 35,000 kg reported (Saadati, 1977) but this was unusual in that there were no other fish and food supplies were extensive. The lake is subject to winter-kill and requires stocking and/or helixers to maintain open water in winter (Bullock, 1971; Nehring, 1973b; 1973c; 1973d; 1974b; 1974c; 1974d; 1974e; 1974f; 1975b; 1975c; 1975d; 1975e; 1975f; 1975g; Nehring et al., 1974; Abbasi, 1974; 1975; Boettcher, 1974a; 1974b; Sanford, 1975; Harrington, 1976). Kohnehchahry and Heydarpur (1973) outline methods of raising trout using submerged cages which they believe would be suitable for Iranian waters. Cage culture has been tried in Gorgan Bay, the fish escaping during several storms and numbering in the millions (B. Kiabi, pers. comm., 1994; www.ramsar.org/ram_rpt_37e.htm, downloaded 28 July 2000). They have also been cage cultured in Valasht Lake, Mazandaran where 12 cages were expected to produce 22 tonnes of fish (Abzeeyan, Tehran, 5(5):IV, 1994). Nezami et al. (2000) give a total production of fish farms in Gilan, Mazandaran and Golestan provinces (along the Caspian Sea) as 851 t.

The "Kelardasht" or Kalerdasht" Fish Farm (= Shaheed Bahonar Trout Farm of Shilat, the Iranian Fisheries Company, near Kelardasht) has an annual production capacity of 2,500,000 fingerlings of 2 g. Various other fish farms in Iran produce rainbow trout for stocking (Surber, 1969; Krasznai, 1987; Petr, 1987). More recently, Bartley and Rana (1998b) give a figure for the Kelardasht Farm of 3 million fingerlings from a pool of 2000 brood fish at a cost of 600 million rials. White (1988) reported that 60 ha of trout ponds and raceways produced 1000 t a year (see also under Cyprinus carpio). The largest fish farm in Lorestan produced 500 t a year in 2000. Edwards (1989) reported production of 1750 t from 20 licensed farms with a further 1000 t from unlicensed farms. Fingerling "trout" production (probably rainbow trout) in government hatcheries was 0.57 million in 1984, 1.81 million in 1985, 1.57 million in 1986, 3.02 million in 1987, 0.50 million in 1988, 4.23 million in 1989, 4.34 million in 1990, 1.90 million in 1991, and 2.00 million in 1992 (Emadi, 1993a). Fingerling production in 1996 was 6 million (Bartley and Rana, 1998a). Production of this trout increased 27% per year betwen 1991 and 1996, the raceway area for trout increased from 80,000 to 166,000 sq m between 1992 and 1996 and production increased from 775 to 1900 t (Rana and Bartley, 1998a). For comparison, the figures for carps (probably Chinese carps) exceeded 100 million and for mahi safid (Rutilus frisii kutum) it was 145 million. However there are also private commercial hatcheries producing about 750 t of trout each year. The Jajrud Hatchery near Tehran produced 1 million fingerlings as early as 1968 as well as several thousand marketable sized fish (Surber, 1969); and there has been a considerable expansion of hatchery capacity in Iran since that time. Rainbow trout production is only about 5% of total aquaculture production although value per metric tonne is U.S.$14,000 while for carp it is U.S.$10,000. Trout production in Ilam Province was estimated at 3.3 million fish based on 21 farms (Tehran Times, 8 August 2005, downloaded 20 January 2006).

Rana and Bartley (1998a; 1998b) outline trout aquaculture methods and problems in Iran. Most culture of this trout takes place in the Alborz and Zagros mountains which are cooler because of altitude. Farms are small, producing less than 50 tonnes per year. Raceways are used for breeding, rearing of larvae and growing out. Eyed eggs may be imported from Scotland and Norway but this may be banned in future because of disease risks. Increased Iranian expertise and costs may also favour production of eyed eggs in Iran. Survival to the eyed stage attains 80% and to the alevin stage 70%. The private sector produced 10 million fingerlings in 1996. The Manzanarbad Trout Farm near Kelardasht sold 2 million fingerlings in 1997 and produced 25 t for local consumption from 2000 sq m of concrete raceways (Bartley and Rana, 1998b). The Mahi Sera Fish Farm at Karaj produces market sized fish from 20,000 brood stock (Bartley and Rana, 1998b). These are replaced every 2-3 years to maintain growth rates. About 5000 fish are saved each year as next year's brood stock based on size, form and colour. Milt from 4 males is used to fertilise eggs from one female. In 1997, the farm produced 185 t of fish each weighing about 250 g (= 800,000 fish). Sale price is 10,000-12,000 rials/kg. The production cycle is 9-10 months at 13°C.

Trout are fed commercial pellets with a conversion efficiency of 1:1.1-1.4 (wet:dry weight). Growth to marketable size (about 30 cm and at least 225 g) takes 9-14 months depending on temperature (2-13°C) at different localities. Feed deficiencies, poor genetic stocks and low temperatures contribute to slow growth in some farms. Production in raceways is nevertheless about 15kg/cu m, a good efficiency. Cost of fingerlings to grow out farms is 120 rials for a 1 g fry, 250 rials for a 5 g fry and 15 rials/g for heavier fish (Bartley and Rana, 1998b).

Effluents from rainbow trout farms have a significant adverse impact downstream on nitrate, nitrite and phosphate and on conductivity as shown by a study of the Zaringol Stream, a branch of the Gorgan River (Mirrasooli et al., 2012). Fadaeifard et al. (2012) also found impacts but observed that it was possible to construct fish farms at about 1.5 km intervals discharging into the Kuhrang stream in the upper Karun River basin, this distance being dependent on stream discharge and self-purification of the stream.

There is no consistent monitoring of released fish so the success of stocking is unknown. The Dorudzan Reservoir about 100 km from Shiraz has received 400,000 fry in 1970, 2 million in 1972, 300,000 from Australia in 1974, 100,000 in 1984 and 100,000 in 1985 (Petr, 1987). Experimental fishing in 1976 gave a poor return, perhaps through competition for space with native e Capoeta species, heavy poaching during migration into the inflowing river, and lowering of the reservoir level. Eggs are no longer imported and the supplies are spawned in Iran (Emadi, 1993b).

Artificial breeding of this species at the Shahid Motahary Fish Hatchery in Yasuj has been carried out 4-5 months earlier than normal using changes in light regime and hormones. The object of this work was to produce fingerlings throughout the year, increase production and save on production costs (Abzeeyan, Tehran, 7(5):IV, 1996). Dehghani and Akbari (1997) have successfully carried out gonadectomies on this species under field conditions.

A production of 5.5-8.0 tonnes of rainbow trout in rice paddies during winter has been recorded (Iranian Fisheries Research Organization Newsletter, 22:2, 2000). Production in Ilam Province was estimated at to be 3.3 million fish by the end of Iranian year 2005 (http://agriculturenews.faorne.net, downloaded 11 April 2006). However, some farms are not economical in Ilam (Rezaei and Darvishi, 2007).

Forghani et al. (2010) studied culture of rainbow trout fingerlings in shrimp ponds at Abadan and found a survival rate of 86.7% with a food conversion rate of 1.59.

Culture of this species in earthern ponds at Bafgh in Yazd Province used running, saline ground water unsuitable for drinking or agriculture. Production was 2-4 tonnes/ha in 0.40-0.75 ha ponds with salinity at 10-15‰, well water temperature 24ºC reduced to usually less than 20ºC, with a water replacement of 5-10% per day for a period of 5-6 months in autumn and winter. The highest growth performance was from fish fed 35% protein, 430 Kcal/100g energy diet and 20.6% lipid with P/E ratio of 81.4 mg protein/Kcal energy. Aeration had no significant effects on growth (Iranian Fisheries Research Oraganization Newsletter, 25:4, 2000; 54 & 55:4, 2009; Ahmadi and Alizadeh, 2004; Bahabadi, 2006). A total of 500,000 fingerlings in fish ponds were released in a two month period in 2005, expected to result in 220 t of product (www.growfish.org/Iranianreport.html). Gonad development was found to be accelerated in brackish water, males maturing two months earlier than in fresh water and ovarian development also being accelerated in Yazd fish (Fahlati Marvast et al., 2003; Fahlati Marvast et al., 2006).

Edwards (1989) gives details on culturing this species and the problems to overcome. Abtahi et al. (2002) found clove oil had no significant difference with MS222, another anaesthetic used in fish farms. Sajedi et al. (2003) have examined mtDNA in Iranian hatchery stocks to measure inter- and intra-populational diversity as an aid in fishery management by selection of the best broodstocks. Studies on the best foods and conditions for this trout in Iran have been undertaken by various authors. Kenari and Mirzakhani (2005) examined the effects of using Artemia urmiana from Lake Urmia (= Orumiyeh) enriched with n-3 HUFA, a fatty acid not present in the brine shrimp, as a larval food. Growth and survival was better than with non-enriched shrimp or with artificial feed. All-female diploid and triploid rainbow trout have been produced in Iran, such trout being unable to reproduce if escaping from fish farms (Johari, 2005). Ahmadi et al. (2005) used the Caspian Sea amphipod, Pontogammarus maeoticus, in rainbow trout food as a means of improving muscle pigmentation, the reddish colour from carotenoids in this dietary item enhancing market value. Carotenoid deposition was higher in males (R. Khodarahami in 5th International Symposium on Sturgeon, Iranian Fisheries Research Organization, 9-13 May 2005, Ramsar). Alavi Yeganeh et al. (2004) studied the use of gammarid powder as a dietary item and its effects in reducing stress in trout larvae, finding a 10% supplement to be effective. Supplementary levels of 2 and 4% of the amphipod were effective in improving flesh colour. Fani (2006a) found fishmeal to have higher digestible and metabolizable energy than soybean meal as feed. Fani (2006b) examined the use of oil by-product from soybeans on weight gain, feed conversion ratio and flesh chemical composition, recommending 10% oil by-product be added to the diet. Shadnoush (2006) showed that corn meal could be used as a nutrient in trout diet, improving weight and length. Kalbassi et al. (2004) induced tetraploidy using heat shock. Johari et al. (2006) investigated production of all-female trout using sex-reversed males and found this to be one of the best methods. Zamani et al. (2006) showed that chromosome manipulation (triploidy) had no effects on digestive enzyme activity.

The decline in caviar production has led to a project in Iran to use trout eggs, test being made on egg tenderness, colour, smell taste and shelf life ( (Iranian Fisheries Research Organization Newsletter, 58 & 59:1, 2009).

In addition to these reports, there have been a wide variety of studies in Iran on methods and treatments relevant to culturing rainbow trout, briefly summarised here. Others may be searched for in the Bibliography by common or scientific name of the fish. Sharifian (1998) studied increases in cortisol hormone levels in trout as a measure of stress caused by handling and induced spawning; Akhlaghi (2000c) on immunology of suspected viral diseases in cultured trout, namely infectious haematopoitic and pancreatic necrosis; Akhlagi et al. (2000) on using powdered clove tree as an anaesthetic; Mirvaghefi et al. (2000) on hydrogen peroxide for fungal control on eggs; Alizadeh et al. (2001) on the effects of dietary energy levels on growth and carcass composition in trout maintained in brackish water; Nafisi Behabadi et al. (2001) on using poultry by-product meal as a dietary replacement for fish meal fed to trout maintained in brackish water; Soltani et al. (2001) on the anaesthetic effect of clove flower extract under various temperatures and pH; Vahdati et al. (2001) on optimal conditions for phagocytic activity of granulocytes; Vahdati et al. (2001) on crowding stress and haematological parameters; Vahdati et al. (2001) on the effect of cortisol on phagocytic activity of macrophages; Akhlagi (2002) on vaccination against streptococcosis; Azari Takami et al. (2002) and Dorafshan et al. (2006) examined induction of gynogenesis by UV radiation; Dorafshan et al. (2002) induced spawning by use of a GnRH analogue; Soltani et al. (2002) on fry mortality syndrome in farmed fish in Tehran and Lorestan provinces; Yakhchali (2002) on the abundance of the parasite Ichthyophthirius multifilis on coldwater fish farms (which produce trout mostly); Fallahi et al. (2003) on isolating and identifying infectious haematopoietic necrosis virus from farmed fish; Farhangi et al. (2003) on the use of natural zeolites for reducing ammonia toxicity; Johari et al. (2003) on sperm traits and fertilisation performance of normal males and neomales; Ebrahimi (2004; 2005) on the deleterious effects of copper, cadmium and zinc, pollutants, on sperm anatomy and motility; Lorestany et al. (2004) on activating solutions and their effects on sperm motility and fertilisation rate; Sharifrohani (2004) on the use of plant essential oils as an antifungal agent for fish eggs in aquaculture; Sheikhi Moghaddam et al. (2004) on the use of alvita (sodium di-acetate) as a fungicide and bactericide; Bazyar Lakeh et al. (2005) on the effect of dietary astaxanthin on fertilisation rate; Azari Takami et al. (2005) on the nutritional effects on trout larvae of vitamin C-enriched Artemia urmiana nauplii in relation to growth, survival and environmental stress; Esmaeili et al. (2006) induced meiotic gynogenesisi by thermal shocks, the yield being best using shock 35 and 50 minutes after fertilisation; Fallahi et al. (2006) on the serological diagnosis of infectious haematopoietic necrosis disease in fry; Gholipour et al. (2006) found a density of 62 fish per square metre was optimal in concrete ponds for weight gain and feed conversion; Hosseini Najd Geramei and Irani (2006) studied the effects of photoperiod on growth, survival and and feeding of larvae; Johari and Kalbassi (2006) on alterations in the red blood cells of triploid trout; Mirzakhani et al. (2006) studied the feeding of n-3HUFA enriched Artemia nauplii as food for larvae used to increase resistance to environmental stress from pH and temperature; Mirzakhani and Mahmood Abad (2006) found feeding n-3HUFA enriched Artemia nauplii to larvae increased resistance to environmental stress (pH and temperature); Niksirat et al. (2006) found that unfertilised eggs can be held in coelomic fluid outside the body for at least 48 hours without any other treatment;  Rohani et al. (2006) on using Geranium herbarum essence to control fungi on eggs, Safari and Boldaji (2006) examined use of canola meal as a partial replacement for fishmeal to reduce cost of diets; Sear et al. (2006) evaluated sperm in relation to weight, length and condition factor of the fish; Tala et al. (2006) on gonad morphology after administration of 17α-methyl testosterone, female trout becoming functional males; Tala et al. (2006) on histological changes to the gonads after application of 17α-methyl testosterone, ranging from functional gonads to intersexes and sterility; Yarahmadi and Moghadasi (2006) on use of decapsulated cysts of Artemia urmiana in the larval diet; Zorriehzahra (2006) on fry mortality syndrome from 52 fish farms and hatcheries across Iran; Emtiazjou and Sheykhi Moghadam (2007) on the use of Alvita (sodium di-acetate) for removing fungi from eggs as an alternative to the carcinogen malachite green; Geramy et al. (2007) on photoperiod on growth, survival and feeding parameters in larvae; Hosseinie Najd et al. (2007) on initial different foods on subsequent growth of larvae; Kalbasi and Lorestani (2007) investigated the effect of various solutions on the duration of sperm motility, coelomic fluid without blood being the best; Khajeh and Peyghan (2007) evaluated blood serum parameters in fish cultured in earthen ponds and showed age-related changes; Latifi et al. (2007) found that citric acid retarded deterioration of trout during refrigerated storage; Lorestani et al. (2007) correlated various sperm quality measures of broodstocks with fish of different ages; Meshkat Rouhani et al. (2007) found that a lower protein content (40%) when the carbohydrate to lipid ratio in diet is 0.5 gave the most favourable conditions for growth; Mohagheghi Samarin et al. (2007) on the effect of temperature on oocyte quality and therefore stripping time; Naem et al. (2007) on treatment of diplostomiasis using praziquantel; Nafisi Bahabadi (2007) found that high energy feeds for trout reared in brackish water were more effective and commercial; Rezaei et al. (2007) on bacterial changes and the biogenic amines found in farmed trout stored on ice, Soltani et al. (2007, 2007) on vaccination against, and immunological responses to, Streptococcus iniae infection; Naji et al. (2007) on the effects of 17ß-estradiol valerat on sex reversal on sac fry larvae; Rahmati Leeshei et al. (2007) on the effect of phytase enzyme on  digestibility of  four vegetative foods; Safari and Boldaji (2007) on dietary lipid level effects on growth, feed utilisation and composition; Tabandeh and Akhlaghi (2007) on efficacy of conventional disinfectants on isolated Streptococcus iniae from diseased fish, sodium iodide and sodium hypochlorite being most efficient although biofilms lowered  efficiency; Varirizadeh et al. (2007)  on using the gonadotropin releasing hormone agonist (GnRHa) in emulsified form to stimulate onset of ovulation; Yousefian and Rezvani (2007) found that abnormalities were not significantly higher in brother-sister matings than controls; Zamani et al. (2007) on the activity of various digestive enzymes at initiation of exogenous feeding in comparison to Salmo caspius, activity being higher in rainbow trout; Zargarian et al. (2007) on using 2000 units/kg diets of phytase with at least 35% replacement of fish meal by soybean meal in farming trout; Akhlaghi and Sharifi Yazdi (2008) identified Yersinia ruckeri, the causative agent of enteric redmouth disease, in cultured fish from Fars Province; Alevi Yaganeh et al. (2008) on use of Gammarus powder as a dietary supplement for larvae; Alishahi (2008) on vaccinating trout against the ciliate Ichthyophthirius multifilis; Faghani et al. (2008) on the immunostimulatory effects of alginic acid and anti-streptococcus vaccine  on growth rate, feed conversion ratio, condition factor and survival rate; Johari et al. (2008) on red blood cell alterations in triploids; Karbassi et al. (2008) on the environmental management of aquaculture in raceways at Sarab Gerdu; Manouchehri and Akrami (2008) on a new automatic sorter for large amounts of fish; Mohagheghi Samarin et al. (2008) on eyeing and hatching rates under cold temperatures; Motalebi et al. (2008) on production of aflatoxins by molds in improperly stored fish meal used as food for trout; Nafisi Bahabadi and Soltani (2008) on dietary energy levels and feeding rates in fingerlings; Safari and Boldaji (2008) on replacing soybean meal with canola meal in diets; Shadnoush et al. (2008) on using acorn meal as a pellet binder and stabiliser; Shafaeipour et al. (2008) on the effects of dietary canola meal on physiology and biochemistry as it can be used to replace substantial levels of fish meal in food; Yousefian (2008) on genetic parameters of growth; Yousefian and Nejati (2008) on inbreeding depression shown as variations in percentage egg hatchability, survival of fry and weights; Zargar et al. (2008) on the distribution of infectious haematopoietic necrosis in five provinces; Iranian Fisheries Research Organization Newsletter (56:42, 2008) on use of the growth hormone somatotropin to increase body weight, daily weight gain and length; Akbari et al. (2009) on feeding larvae with HUFA and vitamin C enriched Artemia urmiana nauplii; Fazaeili et al. (2009) on deposition of pigments from plants in various tissues (feed conversion ratio and specific growth rate were affected by the experimental diets); Gashti (2009) determined that machine trimming of fillets produced less waste than hand trimming, although hand trimming was faster; Heydarnezhad and Purser (2009) assessed food anticipatory behaviour in fish hand-fed in small raceways; Imani et al. (2009) on feeding deprivation and re-feeding, showing that fish could tolerate and compensate for starvation; Jafarian et al. (2009) on the increase in growth performance and feeding efficiency in larvae fed probiotic larvae with Daphnia magna meal; Lorestani et al. (2009) on the interaction of male broodstock age and different dilutions on the production rate of eyed eggs; Mahmoudzadeh et al. (2009) on the effect of synthetic and alfalfa pigments on tissue pigmentation (feed conversion ratio and specific growth rate were affected by the experimental diets); Mousavi et al. (2009) on the antifungal activity on trout eggs of a combination of essential oils from various herbs; Mousavi et al. (2009) on identification of 7 species of Saprolegniaceae fungi from infected trout eggs; Omran and Hedayatifard (2009) on fatty acid composition; Pooramini et al. (2009) on use of yeast as a probiotic and its affect on growth, survival and carcass quality; Rafaei et al. (2009) on the bioaccumulation of copper, zinc iron and manganese in eggs during incubation, found to be lower than EPA standards; Shafaeipour et al. (2009) on effects of canola meal on growth, body composition and biochemical parameters and its use to replace fish meal in diets; Soltani et al. (2009) on the use of Zataria multiflora (Persian thyme) essential oil and its effects on egg hatchability and survival of larvae compared with hydrogen peroxide and malachite green used as anti-fungal agents; Soltani et al. (2009) on the experimental pathology of Streptococcus sp.; Iranian Fisheries Research Organization Newsletter (56:2, 2008 and 58 & 59:2, 2009) on culturing trout using Silo tanks which are easier to maintain and have higher production than cement tanks; Sourinezhad et al. (2009) on suppression of ovary development in triploids and increase in body growth; Tavakoli and Akhlaghi (2009) on changes in serum and blood factors after infection with the pathogen Aeromonas hydrophila; Akbary et al. (2010) comparing live food and an artificial diet on survival, growth and body composition in larvae, the former two factors being better with live food; Akhlaghi and Nouroziasi (2010) on identifying probiotic vibrios to combat pathogenic ones by stimulating immunity; Chegeni et al. (2010)  on formulated diet with Artemia nauplii in the diet to improve growth and survival of larvae; Dadgar et al. (2010) on cottonseed meal as a total replacement for soybean meal; Dorofshan et al. (2010) on using microsatellite makers to detect and asses hybridization with Salmo caspius; Fadaeifard et al. (2010) on iron and lead amounts in farmed fish, their food and influent water in Chahar Mahall va Bakhtiari being no threat to human health; Ghovati et al. (2010) on on screening broodstock using microsatellite markers associated with body weight; Haghparast et al. (2010) on the antioxidant properties of various sodium salts on refrigerated sticks; Kalbassi et al. (2010) on reduced sperm quality when broodstock are stripped weekly rather than biweekly; Mahboobi Soofiani et al. (2010) on starvation effects on morphology and haematology; Meshkini et al. (2010) on various combinations of Artemia decapsualted cysts, nauplii and concentrated feed on growth and survival of larvae, nauplii and nauplii plus feed being the best; Mohammadtaheri et al. (2010) used microsatellite markers to screen broodstock for resistance to the IHN virus. Moogouei et al. (2010) on the effects of physico-chemical parameters on growth in a raceway culture system; Zorriehzahra et al. (2010) on hematological and biochemical parameters of fry in western Mazandaran, helpful in diagnosing some diseases; Akbary et al. (2011) on fatty acid and vitamin C enriched Artemia nauplii fed to trout larvae and the effects on growth, survival and resistance to high temperature stress; Alinezhad et al. (2011) on contamination of pellet feed with fungi and aflatxoxin B1; Ansari et al. (2011) on the improving effects of natural (algal) over synthetic astaxanthin in food on egg quality; Badzohreh et al. (2011) on the supplement β-glucan (Macrogard) having a positive effect on growth and enhancing efficacy of anti-Streptococcus iniae vaccine; Banaee et al. (2011) on oral administration of the herbal medicine silymarin (an extract from milk thistle seeds) affecting blood parameters at higher doses (800 mg/kg); Faramarzi et al. (2011) on longer photoperiods increasing growth and feed utilization; Gholipour Kanani et al. (2011) on anaesthetic effects of methanesulfonate, clove oil and electroanaesthesia on lysozyme activity, with the first having the least effect; Golchinfar et al. (2011) on activity of digestive enzymes during fry development in relation to optimal aquaculture feeding; Haghighi et al. (2011) on the improving effects of oral recombinant bovine somatotropin on growth performance; Jafarian et al. (2011) on comparison of commercial microbial products and those isolated from sturgeon guts on growth and survival of larvae; Jalali et al. (2011) on gene expression; Kiaalvandi et al. (2011) on the use of different fish meals as the main protein sources for growth; Liaghat et al. (2011) on how the acquired immune status of survivors of natural Streptococcus iniae infection was not altered by immunization; Mavadati and Habibian (2011) on the comparative effects of clove oil and 2-phenoxyethanol as safe anaesthetics; Mohammadzadeh and Rezaei (2011) on dipping whole fish in green tea extract as an anti-oxidant to increase storage on ice; Noori et al. (2011) on long photoperiods being effective in growth and retardation of gonadal development in females; Oraei et al. (2011) on gamma irradiation and its lack of effect on fatty acid composition in fillets; Rahimi et al. (2011) on triploid hybrids with Salmo caspius not having salinity tolerance; Rami et al. (2011) on hormonal sex reversal using synthetic estrogen as all-female stocks are economically advantageous, growing larger and maturing later than males; Salamatdoustnobar and Ghorbani (2011) and Salamatdoustnobar et al. (2011) on using prebiotics to improve meat quality and growth; Sheikholeslami et al. (2011) on use of inulin as a prebiotic and the resulting increased resistance to Streptococcus bacteria; Tahmasebi et al. (2011) on dietary nucleotides havig positive effects on serum complements and resistance to Streptococcus iniae; Vahidi and Ghodratizadeh (2011) on the enhancement of immune response with a chitin-supplemented diet; Zolfaghari et al. (2011) on on determining shelf-life of fresh fillets (5 days at 4°C); Bakhtiyari et al. (2012) on varying dietary protein levels at different feeding times and the effect on nitrogenous waste production in a semi-recirculating rearing system; Moahmmadi Azarm et al. (2012) on soybean and egg lecithin effects on growth, survival and resistance to anoxia stress of alevins, with the latter being best; Moradyan et al. (2012) on low stocking density having better effects on alevin survival and growth; Saeedi Far et al. (2012) on the effects of diazinon, an organophosphate pesticide, on haematological indices of fry; Zarghami et al. (2012) evaluated oxytetracycline resistance of Aeromonas hydrophila isolated from reared trout, which resistant bacterial infection or resistant genes could be transferred to humans; etc. Numerous further studies on this economically important species in Iran are not all documented here.

The Department of the Environment has arranged fishing matches for this trout in the Lar Dam since 1994 (www.iran-doe.org/Special/LAR.htm, downloaded 29 December 2000).

Robins s et al. (1991) list this species as important to North Americans. Importance is based on its use in aquaculture and aquaria, as food, in sport and in textbooks. Rainbows have been used extensively as research animals as their requirements are well known and they are readily available from hatcheries. Hatchery fish often have reduced or absent fins and deformed mouths. Farmed fish are sold frozen world-wide, the most important trout in this regard. Rainbow trout are one of the top few sport fishes in North America and of great commercial importance because of the money spent on gear, accommodation, transport, etc. by anglers in pursuit of this fish. Many books and articles have been written on the methods and joy of catching this trout. The flesh is excellent eating fresh or smoked and may be red if food is mostly invertebrates or white if food is fishes.

Conservation

Alborz trout have escaped and made their way to the Caspian Sea but the numbers and impact are unknown (Bartley and Rana, 1998b). This species is an exotic and does not need conservation but escapees may affect survival of native species.

Further work

Bartley and Rana (1998b) make various recommendations for the aquaculture of this species in Iran. The establishment of reproducing populations in the wild should be carefully monitored because of the dangers for native species posed by this exotic. Triploid all-female trout should be considered.

Sources

Sources are summaries of biology in North America such as Scott and Crossman (1973). Observed on trout farms in Iran but there is little data on escaped Iranian populations.

Iranian material: no preserved material.

Genus Salmo
Linnaeus, 1758

Members of this genus comprise about 25 nominal species found in North America and Eurasia including the Black, Caspian and Aral seas basins and the upper, and cooler, reaches of neighbouring basins in Iran and Turkey including the Euphrates River, and the Orumiyeh and Namak lakes (Berg, 1948-1949). A single, variable species was recognised throughout Europe and east to Afghanistan, including Iran (Salmo trutta Linnaeus, 1758) but recent studies indicate a greater diversity (Turan et al., 2011).

These fishes have teeth on the shaft and head of the vomer bone in the roof of the mouth (may be lost with age), the jaw is long, reaching to the posterior eye margin or beyond, scales are small usually more than 100 in the lateral line, colour is pale silvery with dark markings or spottings, anal fin rays (all counted) number 15 or less (16 or more in Oncorhynchus), and there are various unique osteological characters.

These are famous sport fishes and were once caught in commercial quantities. Their biology has been studied extensively and numerous books and papers have been written about them.

Salmo salar
Linnaeus, 1758

Introduced to the Caspian Sea (Mamaev, 2002). No Iranian record.

Salmo caspius
Kessler, 1877

  Stuffed specimen, Bandar-e Anzali,Brian W. Coad
Stuffed specimen, Bandar-e Anzali, Brian W. Coad

Courtesy of H. Niksirat
Courtesy of H. Niksirat

Hevigh River, Caspian Sea basin, July 2010, courtesy of K. Abbasi
Hevigh River, Caspian Sea basin, July 2010, courtesy of K. Abbasi


Lar Dam, September 2008, courtesy of K. Abbasi


Chesli River, Gilan, June 2010, courtesy of K. Abbasi

Liqvan Chay trout, ca, 27.0 cm total length, 4 October 1994, courtesy of Asghar Abdoli
Liqvan Chay trout, ca, 27.0 cm total length, 4 October 1994, courtesy of Asghar Abdoli

Liqvan Chay trout, ca, 27.0 cm total length, 4 October 1994, courtesy of Asghar Abdoli
Liqvan Chay trout, ca, 27.0 cm total length, 4 October 1994, courtesy of Asghar Abdoli

Common names

آزاد ماهي (azad mahi or mahi-ye azad = free fish, used in Farsi for trout and salmon), ماهي آزاد (= mahi azad), azad mahi qezelala-ye daryacheh khazar (= Caspian Sea trout), mahi azad Darya-ye Khazar, mahi azad setareh-i Darya-ye Khazar, all for the subspecies caspius; قزل آلا (= gazalala, ghezel ala or kizil ala, meaning red spots), قزل آلاي خال قرمز (= qezelala-ye khalqermez; the Turkic qezel ala is used in Farsi for trout and hence this last means "trout with red spots" although literally it is a tautonym, "red spots with red spots"), mahi azad-ye kezel ala-ye ilbumi (= native salmon trout), qezel ala khalqermez yabumi, qezel ala-ye juibary (= brook trout), all apparently for freshwater residents; mahi azade khal sorkh; qezel ala-ye Liqvanchai (for trout in the Lake Urmia basin).

[gizilxalli or ala-balyk for freshwater residents and gizil balig or kizil-balyk for sea-run fish in Azerbaijanian; Kaspi azatmahysy or kumja in Turkmenian; ala-balukh Armenia; Kaspiiskii losos' or Caspian salmon for sea-run fish, and ruch'evaya forel or brook trout and pestrushka for freshwater residents in Russian; alabalik in Turkish; massialé in Kurdish; brown trout, brook trout, river trout, Caspian trout].

Systematics

Salmo Trutta was originally described from European rivers. Salmo caspius Kessler, 1877 was described from the Bozh'em-Promysl (or Bozhii Promysel) fishing grounds on the Kura River, Azerbaijan with 3 syntypes in the Zoological Institute, St. Petersburg (ZISP). Salmo spurius Pallas, 1814 from the Terek River in the Caspian Sea basin, Russia and Salmo lacustris var. Romanovi Kavraiskii, 1896 from Lake Tabiszchuri in Transcaucasia are usually regarded as synonyms of Salmo trutta. Salmo trutta ezenami Berg, 1948 is a subspecies from Lake Ezenam in Dagestan (ZISP 28356)(Dorofeyeva and Salmanov (2001)) though it may be deserving of species rank (Reshetnikov et al., 1997). Salmo trutta ciscaucasicus Dorofeyeva, 1967 (see also Dorofeyeva (1997) and Dorofeyeva and Salmanov (2001)) is described from the Keyranchay (ZISP 26244) and is found in drainages of the western shore of the Caspian Sea, except for the Kura River and, presumably, excepting Iranian rivers.

The native brown trout (freshwater populations) and Caspian salmon (sea-run populations) of Iran were both usually referred to Salmo trutta caspius Kessler, 1877 in the Caspian Sea and Namak Lake basins (Saadati, 1977). Derzhavin (1929b) examined two small fish from the Karaj River of the Namak Lake basin and recognised them as typical "brook trout" (Salmo trutta). Nümann (1969) considers that southern Alborz trout originate from Caspian stocks. Some authors call the species Salmo fario caspius Kessler, 1877, e.g. Dadikyan (1986). Fricke et al. (2007) and Naseka and Bogutskaya (2009) regard caspius as a full species.

Shirangi et al. (2011) used microsatellite DNA to compare spring and fall migratory forms in Iran. Spring fish are more fusiform and silvery than fall fish and this was thought to indicate a genetic difference. However, they found low genetic differentiation and conclude that only one population exists in the southern Caspian Sea.

Tortonese (1954), Kuderskii (1974) and Kottelat (1997) discuss the confused nomenclature of this species. Boulenger (1896), Berg (1948-1949) and Derzhavin (1929b) consider Namak Lake basin trout to be e Salmo trutta macrostigma (Dumeril, 1858), originally described from Algeria as a distinct species. Turan et al. (20120 note that Salmo macrostigma (Duméril, 1858) is restricted to Algeria. The Namak trout were supposedly derived from Mediterranean populations via Persian Gulf drainages but Saadati (1977) dismisses this on the grounds of a close similarity between Namak and Caspian trout and the apparent absence of trout from the Zagros Mountains and Persian Gulf drainages of Iran. Osinov (1988) could not determine a Mediterranean route based on electrophoretic evidence.

In the Lake Orumiyeh basin, apparently restricted to the "Lighvanchai" (probably the Liqvan Chay, the town of Liqvan being at 37°50'N, 46°26'E), is a population of brown trout which has been referred to as a distinct subspecies (Anonymous, 1977). It has not been formally described. Saadati (1977) considers all Iranian trout to be S. t. caspius, with those in the Namak Lake basin being a distinct "race", with some slight differences to typical S. t. caspius (see below), and those trout in the Lake Orumiyeh basin could be a new subspecies.

Earlier authors consider the Caspian basin species to be the same as the Atlantic salmon, Salmo salar Linnaeus, 1758, but blood serum electrophoresis (Ostroumova, 1970) support its position as Salmo trutta (and presumably caspius) as does karyology and osteology (Dorofeyeva, 1965; 1967) and enzyme electrophoresis (Osinov, 1984; 1988). Osinov and Bernatchez (1996) consider trout from the Caspian Sea basin to be part of their "Danubian" grouping, brown trout from the Black, Caspian and Aral sea basins as opposed to an "Atlantic" grouping from the Baltic, Barents and White sea basins (and also a tributary of the upper Volga River in the Caspian Sea basin), based on allozyme and mtDNA studies. Vera et al. (2011) concur. Novikov et al. (2008) examined allozyme variability in trout populations from Iran and found them to be similar but diverging significantly from other populations in the Caspian Sea basin. Vatandoust et al. (2008) demonstrated population differences between rivers in Mazandaran using meristic and morphometric characters. Turan et al. (2009) described two new species of trouts from northern Anatolian streams, one resident and the other migratory. A similar situation may occur in Caspian streams where there are the two life history types, although the authors caution that each situation should be investigated individually. They advocate a more cautious treatment of trout taxonomy as such results have important implications for management and conservation. Vera et al. (2011) found two main clusters, using microsatellite data, connected by gene flow among river basins in Iran, presumably by anadromous fish. Jamshidi and Kalbassi (2011) examined random amplified polymorphic DNA markers for spring- and fall-run trout from rivers along the Caspian shore of Iran, and recognised them as conspecific. Hashemzadeh Segherloo et al. (2012) used DNA and morphology to compare south Caspian, Orumiyeh (including Liqvan Chai fish) and Namak populations. They concluded that the Caspian and Orumiyeh populations did not differ significantly but the Namak population represented a unique haplotype. All haplotypes fell within the Danubian phylogenetic grouping as distinct members. The Namak haplotype may have a centre of origin in the Caspian basin.

A artificial hybrid of Salmo trutta caspius with Oncorhynchus mykiss is reported on by Pourgholam and Noruzy Moghadam (1996).

Key characters

The dark-spotted back, light halos around some of the dark-coloured flank spots, caudal fin not or only weakly spotted, teeth on the vomer shaft, and only 9-15 total anal fin rays are distinctive. ? from S. trutta?

Morphology

Salmanov (1990) has demonstrated that brook, lake and sea forms of trout from the Caspian Sea basin differ in head and body proportions and in fin shapes although no populations in Iran were considered or given taxonomic status. Osinov (1988) maintains that migratory and non-migratory populations are the same species.

In the Caspian Sea basin dorsal fin with 3-5 unbranched and 7-12 branched rays, anal fin with 2-5 unbranched and 6-10 branched rays, pectoral fin with 10-14 branched rays, and pelvic fin with 6-9 branched rays. Lateral line scales 108-134 (Salavatian et al., 2011; Salavatian et al., 2011). Scale rows between end of the base of the adipose fin and the lateral line 15-17, mode 16 (Turan et al., 2012). Gill rakers 15-23. Pyloric caeca 26-61. Vertebrae 55-61. Scales in lateral series for S. caspius 140-160, vertebrae 59-61 (Derzhavin, 1929b; Behnke, 1965). See also below under Zoogeography. The chromosome number is 2n=78-84 (Klinkhardt et al., 1995), 2n=80 (Annual Report, 1995-1996, Iranian Fisheries Research and Training Organization, Tehran, p. 61, 1997; Kalbassi et al., 2006) or 2n=80-82 (Dorofeeva, 1998). The chromosome formula is 14M + 10SM + 56T with NF=104 (Kalbassi et al., 2006).

Meristic values for Iranian specimens are:- dorsal fin branched rays 9(1), 10(5) or 11(1), anal fin branched rays 8(7), pectoral fin branched rays 12(7), pelvic fin branched rays 7(1) or 8(6), and total gill rakers 17(3), 18(3) or 19(1).

The Liqvan Chay fish are similar to S. caspius in all except colour, gill rakers being somewhat higher (18-22, mean 20.2) and lateral series scales being higher (132-154, mean 142) but sample sizes were small. Salavatian et al. (2011, 2011) found the Lar Reservoir population to be distinct morphometrically and meristically from fish from the Republic of Azerbaijan.

Sexual dimorphism

Abdurakhmanov (1962) reports on two populations form Azerbaijan where he found head length, predorsal distance, snout length and lower jaw length to be greater in males and postorbital length greater in females for both populations, while body depth, dorsal and anal fin heights, and pectoral and pelvic fin lengths are greater in males and eye diameter, pelvic-anal fin distance, head depth and interorbital width are greater in females for one population.

Colour

A specimen from the Anzali Mordab had a dark brown-grey back and top of the head with some round black spots, silvery-yellowish sides with 3-4 longitudinal rows of small red spots with whitish halos, a large black spot on the preoperculum and several smaller spots on the operculum. The belly and lower head surface were white with minute dark melanophores. The dorsal fin was a pale grey with 3-4 horizontal rows of black spots and 1-2 series of 4-8 ellipsoid red spots vertically. The adipose fin was overall yellowish with some grey pigment while the upper half of the fin was more orange in colour. The pectoral, pelvic and anal fins were a pale yellowish or nearly colourless with grey pigment. The anterior 2-3 rays were coloured orange. The caudal fin was greyish with the upper and lower marginal rays dark orange. All fins lacked spots except one red spot observed on the dorsal fin of a larger fish (19.8 cm total length). Smaller fish (13.7-16.2 cm fork length) had 12-14 parr marks. The red flank spots are recognised in the name "kizil ala". Generally in the sea, this species has numerous, dark, x-shaped spots on silvery flanks although some individuals have almost none. The fish caught by Holmes (1845) mentioned below had a dirty olive-green back, dark brown and red spots on the flanks, a roseate tint to the flanks, and a golden belly. The migratory form was dark blue on the back, silvery on the flanks and belly, and had dark spots on the flanks. Turan et al. (2012) state that this species has few, black spots on the body, ocellated with a narrow light ring, restricted to the back and upper part of the flank (about upper third in fish larger than 200 mm SL), and absent from the mid-dorsal area in front of thr dorsal fin.

Prosek (2003) illustrates in colour a trout from the Tigris River in Turkey, from the Caspian Sea basin in Turkey and Armenia, and from the Liqvan Chay.

The Liqvan Chay population has 54-400 red to orange spots along its flank (compared to only 30-50 generally in European brown trout) giving it a ruby-red sheen (Anonymous, 1977). Only one Liqvan Chay fish had less than 100 red spots based on 13 fish examined by Saadati (1977) compared to a range of 27-134 for 13 S. t. caspius from the Shah Neshin River near Ardabil, where only 2 fish had more than 100 spots. The spotting pattern consists of profuse black and red spots and is distinct from all other Salmo trutta sensu lato which have fewer and larger spots (Saadati, 1977). Colour photographs, courtesy of Asghar Abdoli of the Agricultural and Natural Resources University, Gorgan, of 3 fish from the Liqvan Chay dated 4 October 1994 with a total length of 27.0 cm showed two fish with about 87 red spots while the other had about 200 (accurate counts not possible because of body curvature and silvery reflections). There was considerable variation in spot size among the three fish, the one with most spots having generally smaller spots which give the sheen referred to above, while the other two fish have more discrete and larger spots. It is unknown whether the Liqvan Chay population has been contaminated with other stocks of Salmo trutta.

Size

The sea-run Caspian trout attains a larger size than freshwater populations. Attains 51.0 kg and 1.5 m but most seen in Iran were 10-15 kg (Walczak, 1972). Length reaches 118 cm and weight 21 kg in Iranian samples but such large fish are rare in Iran (Farid Pak, 1968b). Most Iranian catches are 55-105 cm and 1.8-12.7 kg (Farid-Pak, no date). Holmes (1845) reported one fish caught at the mouth of the "Mazzur" River weighing 16 lbs (7.3 kg) and measuring 38.5 inches total length (0.94 m). Nümann (1969) reports fish almost 1 m long in the Karaj Reservoir in 1967. A fish 78 cm long and weighing 8 kg was worthy of note in 1996 (Anonymous, 1996c).

Distribution

The Caspian salmon is found in the Caspian Sea and enters Iranian rivers to spawn as well as being resident in Gilan, Mazandaran and Golestan provinces (Sheil, 1856; Nedoshivin and Iljin, 1929; Kozhin, 1957; Nümann, 1966; R. J. Behnke files; Armantrout, 1980; Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 28, 1996; Abbasi et al., 1999; Abdoli, 2000). Formerly known from the Anzali Mordab and its tributaries (Holčík and Oláh, 1992). Reported prolifically from the Seh Hazar, Chalus and Babul rivers (Fortescue, 1920). Found in the Safid River, Kargan River near Hashtpar, Chalus and Babol rivers. Rare in the Aras (Berg, 1948-1949). Abundant in the upper reaches of the Gorgan (G. S. Karelin cited in Berg (1948-1949) although Karelin only visited the lower Gorgan himself). Reported from Gorgan Bay, the southeast Caspian Sea, southwest Caspian Sea and south-central Caspian Sea and from the Tajan, Babol, Haraz, Sardab, Shirin, Ashek, Kelyareh, Nav, Tonekabon, Pol-e Rud, Lar, Elarm, Ab-Safid, Kamardasht, Delichai, and Safid rivers (Kiabi et al., 1999; Vatandoust et al., 2008; Abdoli and Naderi, 2009; Salavatian et al., 2011 Shirangi et al., 2011). Jolodar and Abdoli (2004) give its distribution in the Caspian basin as in upstream waters from the Aras to the Tajan, absent in the Atrak and Gorgan rivers.

Also found in the Jajrud (Ouseley, 1819-1823), in the Karaj River (Fortescue, 1924; Derzhavin, 1929b; Berg, 1949), and from the vicinity of Damavand (? Damavand River) (Fraser, 1825), and numerous other localities in the Namak Lake basin (see Armantrout (1980) for localities from the 1970s and Abdoli (2000) for more recent general distributions). However Nümann (1969) seems to indicate that fish in the Karaj River were stocked about 100 years previously from Caspian Sea rivers.

Reports from the Zagros Mountains in Iran are uncertain and those from mountains near Kerman even more so! (Walczak, 1972). These last two records have not been confirmed by specimens and with introduction of exotic material may now be difficult to verify. Fraser (1834) reports that "Trouts are found in several of the streams of Azerbijan and Kurdistan". It may be worth noting that Heckel (1843b:995) reports that the collector Theodor Kotschy "did not find trout....in the mountains of Kurdistan. Since we know our collector's diligence, we doubt that trout occurs there" but in (1846-1849a:254) Heckel states "we must add one trout (Salmo), which tastes excellently according to the report presented by our traveller; that trout occurs relatively frequently in the mountains of Kurdistan, but has not been seen by us". This is merely confusing: did Kotschy later find trout or change his account, or did Heckel recall Kotschy's accounts incorrectly? Turan et al. (2011) describe a new species of trout, Salmo tigridis, from the Tigris River basin in Turkey and it is possible that this species occurs in Iranian tributaries of that basin. It has 19-20, mode 19, scale rows between the end of the adipose fin base and the lateral line (27-32, mode 28 in S. caspius), a deeper and stouter caudal peduncle with depth 11.5-12.6% SL, mean 11.8% (10.0-11.5, mean 10.4), fewer gill rakers 17-19 (19-21), a shorter maxillary in males at 8.7-9.2% SL, mean 8.9% (9.4-11.2, mean 10.4), and in general body colouration being grey in males and greenish to yellowish in female (brownish)

The Liqvan Chai in the Lake Orumiyeh basin contains a distinctive trout and Liqvan Chay trout were stocked in the Ab-e Bazuft of the upper Karun River (Tigris River) basin in the autumn of 1975 but this failed (B. Sandford, in litt., 1979; Prosek, 2003).

Zoogeography

The origin of Iranian trout is probably from the north via glacial lakes during the last or earlier ice ages and via the Volga River system or, possibly, from the west via the Black and Mediterranean sea basins (Tortonese, 1954; Nümann, 1969; Kuderskii, 1974; Osinov, 1984; Farid-Pak, 1991). The initial colonisation of the Caspian Sea by this trout apparently occurred much earlier than the late glacial period on electrophoretic evidence (Osinov, 1988). The question of a northern origin remains open however on present biochemical evidence (Osinov and Bernatchez, 1996). The division between their Atlantic and Danubian groupings occurred about 0.5-0.9 million years ago based on their data, perhaps 0.7-2.0 million years ago when the division of European and North American groups of salmonids are taken into account. These authors note that populations within the Caspian Sea basin have unique gene pools, suggestive of reproductive isolation, and these fish are not simply the subspecies S. trutta caspius. Bernatchez (2001) considers all Caspian trout to belong to the Danubian or Ponto-Caspian lineage, one of five lineages, and that the centre of origin was probably from drainages associated with the Cauacasus region of the Black Sea. The major demographic expansion of this lineage probably occurred about 270,000-290,000 years ago. Bernatchez (2001) considers that populations of each sea basin occupied by the Danubian lineage should be recognised as distinct evolutionary lineages but he does not give them names.

Trout in the Zagros Mountains may be e Salmo trutta macrostigma (Dumeril, 1858), if this is a valid subspecies, while some authors consider Zagros trout to be S. t. caspius on zoogeographical grounds but pure samples of this population have never been examined systematically (Behnke, 1965). Boulenger (1896) and Berg (1948-1949; 1949) consider trout in the upper Euphrates and in the Namak Lake basin to be S. t. macrostigma. Namak Lake trout are then Pliocene relicts of this Mediterranean subspecies from an invasion via the Persian Gulf region. However, the Namak Lake fishes generally show evident affinities with the Caspian Sea basin ichthyofauna and the trout themselves are very similar in appearance.

The Namak Lake basin trout are generally held in Iran to have been planted by the royal family in the nineteenth century (Saadati, 1977). The royal family had sporting camps on the Lar River and the "Varang-e Rud", a tributary of the Karaj River. Saadati (1977) compares 30 specimens from each of these two localities and found characters to be similar except for pyloric caeca (45-58, mean 51 in the Karaj sample, 35-46, mean 39 in the Lar sample) and in scales in lateral series (131-146, mean 138) and 121-134, mean 127 respectively). He concluded that Karaj River fish are native, of relatively recent origin from the Caspian Sea basin, and only slightly divergent.

Saadati (1977) also reports on a population of trout in the Mordugh Chai (probably Mordaq Chay) in the Lake Orumiyeh basin. This population was typical S. caspius with only 17-38 red spots on the flank. It may have been of recent origin via a headwater transfer as the tributaries to the Caspian Sea basin are found nearby. A human agency may have been involved, always a complicating factor in the distribution of fish which are of sporting or commercial interest.

Habitat

Mahi azad live in the Caspian Sea proper and migrate up rivers to spawn. Moosari (1996) describes migration into the Tonekabon River of Iran. Young mahi azad stay in rivers for 2 years. There are also land-locked populations such as that of the Lar River near Damavand Mountain and this species may also be found in cool to cold lakes. Areas with clean gravel are required for reproduction. Temperatures above 15°C cause egg mortality. Adults can survive up to 29°C, 18-24°C is the optimum range and 12.4-17.6°C the preferred range. Atai Mehr et al. (2006) found juveniles to survive salinities of 0 to 12.5 g/l over 120 hours, indicative of their adaptability to changing environments. The best weight and water salinity for release of cultured juveniles was 10 g and 8.0-12.5 g/l. Bourani et al. (2010) found that weight groups of 10, 15 and 20 g could carry out hypo-osmoregulation of water the salinity of the Caspian Sea but that this ability is not evident in 5 g fish.

In the sea, they inhabit coastal areas at 40-50 m with distinct stocks centred on the basins of the major rivers. Migrations occur from Iranian shores to Dagestan (Caspian Sea Biodiversity Database, www.caspianenvironment.org).

Age and growth

The Liqvan Chay population females only begin to mature in their third year and only 50% of males are mature in their second year. This population appears to be in good condition compared to other populations, e.g. in the Lar River, and these figures probably represent more natural values. A population estimate was 1246 fish per km compared to 11,380 per km in part of the Lar River where fish had concentrated to avoid severe drought. The life span in the Lar is 6 years (Nehring, 1975a).

There are stocks of dwarf males in Caspian rivers which do not descend to the sea.

Farid Pak (1968b) found the smallest sexually mature fish in the Caspian Sea was 53 cm long and the largest 105 cm. Most fish in the commercial catch of the Caspian Sea were 5-9 years old in the 1950s (Farid-Pak, no date). Caspian Sea fish may mature as early as 1 year for males under riverine conditions but most fish mature at 3-9 years, depending on the river (Caspian Sea Biodiversity Database, www.caspianenvironment.org). Afraei et al. (2000) examined 190 fish from the Tonekabon River and found age groups 0+ to 4+. Most fish were in the 2+ age group. Maximum size was 175 mm and 84.5 g. Condition factors were 1.268 for males and 1.257 for females. See also below under Reproduction.

Niksirat and Abdoli (2009) and Abdoli et al. (2011) found a decline in length from 77.6 cm in 1947 to 59.8 cm in 2007 (22.4%), in weight from 4880.2 g to 2486.8 g (49.5%), in absolute fecundity from 7041.8 in 1947 to 4526.1 in 1973 and 2941.2 in 1986, and in relative fecundity from 1451.4 per kg body weight in 1947 to 1372.6 in 1973 and 1199.8 in 1986 (only latter value significantly different in relative fecundity). Size classes over 75 cm have disappeared from the population over the past 60 years and mature samples were less than 50 cm. Various factors cause such declines, including fishing mortality and overfishing of prey such as Clupeonella spp. (and their loss from the exotic comb jelly), an important food which helps produce more eggs. Kheyrandish et al. (2010) examined fish from six rivers in Mazandaran and found five age classes (0+ to 4+) with the most frequent being 1+ and 2+. Condition factor ranged from 0.58 to 1.47. Females dominated over males in 5 of 6 rivers, the highest sex ratio being 6.7 females:1 male in the Pajimiane River and 0.8:1 in the Khojirood. There were significant correlations between fish length and depth and fish length and width of rivers surveyed.

Fazli et al. (2011) studied trout from the Lar Dam and found a significantly different male:female sex ratio at 1:1.84, a life span up to 6 years with age 4 the largest age group, a length-weight regression of W = 0.028L2.706 for females, W = 0.034L2.624 for males showing negative allometric growth for both sexes and W = 0.029L2.710 for the population, growth parameters were L = 45.0 cm, K = 0.27 yr-1, t0 = -0.23 yr, instantaneous coefficients for natural, total and fishing mortality were estimated to be 0.40 yr-1, 0.75 yr-1, and 0.35 yr-1and the exploitation rate was 0.47 These results indicate a fast growth and a population in good condition in the study year of 2004.

Nümann (1969) found Alborz trout to reach 26 cm at the end of 3 years, corresponding to the growth rate for central European fish.

Food

Freshwater populations are known to eat mayfly larvae in Iran as well as other aquatic insects and crustaceans and terrestrial insects taken at the surface. Larger specimens elsewhere are known to take fish and crayfish, and more rarely frogs, salamanders and rodents and this probably occurs in Iran too. Tonekabon River fish (Afraei et al., 2000) fed most intensively in spring and least intensively in autumn. The main prey was Simulium, Plecoptera and Ephemeroptera. Salavatian et al. (2011) examined fish from the the Elarm, Ab-Sefid, Kamardasht and Delichayi rivers in Lar National Park where food was Chironomidae pupae and larvae (91.5%), Coleoptera (6.4%) and others 2.1%. Cannibalism was rare

In the sea, young feed on amphipods, mysids and shrimps while adults take common and anchovy kilkas, silversides and shads.

Reproduction

This fish runs up rivers in October-November and March-May in Iran (Roux, 1961b). Holčík and Oláh (1992) record migrations into the Anzali Mordab and a run up the Pasikhan and Siahdarvishan rivers from late September to the end of December. A second run occurred in April-May but was much smaller. Holmes (1845) reports "salmon" in rivers of Gilan from June to the end of October. Bartley and Rana (1998b) report the spawning run on the Tonekabon River to be from September to November and spawning to be from November to December. Spawning occurs on riffles of sand and gravel beds. Spawners are 8-12 years old and are mature at 700 g. About 1500 eggs are produced for each kilogramme of female. The fish examined from the Tonekabon River (Afraei et al., 2000) had an absolute fecundity of 168-379 eggs, average 268 eggs. Maximum egg diameter is 6.1 mm, number of eggs per gram reaches an average of 14.7 and absolute fertility reaches 13,468 eggs in Iranian fish which are considerably smaller than Kura River fish in Azerbaijan (Farid Pak, 1968b).

Alborz trout are ripe at 2-3 years while Caspian Sea fish spend 2-3 years in rivers, then 2 years in the sea before re-entering rivers to spawn at 60-70 cm. Spawning can occur more than once in a life span (Nümann, 1969).

In the Kura River of Azerbaijan there are two races. One enters the river in October and travels no higher than the middle reaches 600-700 km from the mouth, is sexually mature and does not weigh more than 12 kg, and spawns in the same year. The second begins to enter the Kura in October but most fish run in November and December, this race spawns in the upper reaches about one year later (8-11 months) after a long migration of over 1000 km, and attains 51 kg. There are two races also in Iranian rivers (Berg, 1959). The Kura salmon may have up to 45,000 eggs of up to 6.5 mm diameter. Kura salmon spawn only once between ages 5 and 9 years. Some Kura salmon become smolts and migrate to sea in the first year of life but most descend in their second year.

Parasites and predators

Mokhayer (1976b) records the protozoan Trichodina from the gills of trout (? this species) in the Karaj River, the cestodes Eubothrium crassum and E. rugosum, the acanthocephalan Corynosoma caspicum, and the annelid Piscicola geometra. Jalali et al. (2005) and Malmberg et al. (2007) summarise the occurrence of Gyrodactylus species in Iran and record G. derjavini from fish in the Sardab-rud. Sattari et al. (2005) surveyed this species in the Chesli and Khorma rivers, recording Cystidicoloides ephemeridarum. Sattari et al. (2004) record the nematode Cystidicoloides ephemeridarum sp. from this species in Gilan.

Economic importance

In a survey of the Lar River in June, RaLonde and Walczak (1970b) found 22 fishermen had caught 222 trout with an average length of 19.85 cm, comparable with an earlier survey by Nümann (1964). The largest fish was 27 cm long. The catch was 2.56 fish per fisherman per hour, a good rate of success (but see older records below). This rate and the production of trout in this river was expected to decline drastically with construction of a dam on the Lar. Sport fishermen took 50,000 trout from the Lar River in 1967 (Surber, 1969). Malek-Eizadi (1993) gives a recent account, in Farsi, of the Lar trout and apparently confirms its decline. Floor (2003) cites older records of fishing in the Lar River. Salavatian et al. (2011) indicate that 17,862 licences were issued for summer 2006 fishing in Lar Reservoir.

Azad mahi were caught in weirs, nets and with long, 3-pronged forks year round but the principal seasons were spring and autumn (Holmes, 1845; Floor, 2003). The catch in the Safid River region was largest from February to April, with a maximum in the middle of March. In 1912-1913, a total of 1180 fish were caught in the Safid River, Anzali and Astara regions. Weights reached 10-12 kg, average 7.5 kg (Nedoshivin and Iljin (1929) cited in Berg (1948-1949)). Fortescue (1920) reported 1500-2500 "salmon" from the Seh Hazar at Shahsevar, averaging 8-10 lbs (3.63-4.54 kg) each and seldom exceeding 18 lbs (8.17 kg). Nevraev (1929) reports on catches in various regions of Iran in the early years of the twentieth century. In the Astara region from 1901-1902 to 1913-1914, the catch ranged in numbers from 236 to 1563 and in the Safid River region from 1916-1917 to 1917-1918 the range was 185-663 fish. Yearly catches of mahi azad in the Anzali region alone have reached as high as 3,037 kg in 1939/1940 and as low as 90 kg in 1946/1947 before commercial fishing was prohibited (Vladykov, 1964; Walczak, 1972). The Food and Agriculture Organization, Rome reported only 1 tonne of "salmonoids" caught in Iran for each of the years 1983 to 1985, presumably the Caspian salmon. Occasional catches are taken in sturgeon nets of fish 1-4 kg in the Anzali region and elsewhere along the Iranian coast (Holčík and Oláh, 1992). Many fish are taken by poachers in traps and nets used to block spawning streams. Salehi (2008a) notes a decline in catches of this species from 13 t in 1995 to less than 3 t in 2005 and most of the catch is based on stocking rather than natural reproduction.

There is a state supported stocking programme but growth rate is slow. High summer temperatures (28°C) in the Caspian Sea would affect cage culture and survival (Rana and Bartley, 1998a).

Salmo trutta is reported to be ichthyootoxic although there are no reports for S. caspius (Coad, 1979b).

Conservation

Pietro delle Valle, who traveled in Iran from 1616 for 7 years, reported trout from rivulets in Ardabil in the Caspian Sea basin (see Pinkerton, 1758-1826, volume IX:84), a city not now noted for so salubrious an environment for salmonids.

Ouseley (1819-1823) dined while at Tehran on dried and salted azad mahi two feet long (0.61 m) from the Caspian Sea for breakfast one day in November 1811 and on fresh trout (later referred to as "kizl-áleh") from the Jajrud for dinner.

Vigne (1842) recorded catching six or seven dozen a day in the Lar River near Tehran, Charles Murray reported in 1858 that the stream "abounds so much in trout that I frequently kill 50 in an hour with a fly" (recorded in Wright (1977)), Valentine Baker (cited in Prosek (2003)) caught 50-60 trout a day up to 4 lb in weight, and Mounsey (1872) caught 450 fish up to 1.5 lbs (0.68 kg) in 3 days using flies. Anderson (1880) found that the Lar River "abounded in trout". They were still numerous there in the 1970s (Nehring, 1975a). However this population was unusual in that growth rates and population densities fluctuated drastically from year to year, probably because spawning habitat and nursery areas were the only adequate features of this river. Features lacking were feeding areas, stable banks and stream bed, good plant cover along the banks to supplement productivity, good physical and chemical water quality and water levels, and presence of predators as controls on excess reproduction. As a result, recruitment of young-of-the-year was high and led to interspecific competition, earlier sexual maturity, and stunting. Overgrazing in the Lar Protected Area (formerly a National Park) has affected the survival of trout in this region since this reduces the availability of insect food, and in 2000 this situation was exacerbated by the prolonged drought (Imamai, 2000).

Numbers of this species have declined drastically as evidenced by catch records. The catch decreased 100 times since 1959 to the early 1970s and smaller fish taken in beach seines were used by fishermen to eat as their market value was low. In the 1970s, the Shahsavar River was assessed as the best in Iran for this species as the river was in good condition. However, even this river was blocked in at least two areas for capturing fish and almost all adults were taken for "culture" by Shilat. A take of 300 fish provide as many eggs as 20 should because of poor treatment (15°C, , Saprolegnia infections). The Anzali Mordab was surveyed half a mile from a guard station and 11 gill nets were found, all illegal, in a half mile stretch in half an hour (Carl Bond Archives, Oregon State University, Corvallis). Niksirat and Abdoli (2009) report gill nets used in estuaries catch migrating fish illegally and they are also caught as bycatch in the Rutilus frisii kutum fishery, but not released. About 2-3 times the the legal fishery for artificial reproduction is taken by poaching.

Attempts to increase natural reproduction by raising young to a size more likely to survive have been attempted in Iran (Andersskog, 1970). The "Kelardasht" (or Shaeed Bahonar) Fish Farm (part of Shilat, the Iranian Fisheries Company) on the "Sandar" River has a production capacity of 100,000 fingerlings of 15-20 g but this has never been achieved (Woynarovich, 1985; Krasznai, 1987). It takes about 2 years before fingerlings reach 15-20 g and 10-15 cm, large enough to release. Brood stock are collected from the Tonekabon River during the September to November spawning migration. Spawners are 8-12 years old and mature at 700 g. New brood stock are collected each year although stock from previous years are kept as backup. In 1997, 500 fish were caught in a 3:1 ratio of males to females. Eggs from two females are mixed with milt from two males. Egg yield is about 1500 eggs/kg. Growth rate is slow in this species, compounded by low rearing temperatures of 2-17°C. Fingerlings take 10 months to reach 10 g (Bartley and Rana, 1998b). In a study of the Tonekabon River migration pattern, the following conditions are found to be favourable for release of cultured fish: release size should be at least 15 g, release point 500-700 m away from the estuary, release time March and October, and for one-year-old fish smolt release is favoured over parr as the former would migrate to the sea (Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 28, 1996). Fingerling production by government hatcheries was 0.01 million in 1986, 0.05 million in 1988, 0.10 million in 1989, 0.16 million in 1990, and 0.20 million in 1991 and in 1992 (Emadi, 1993a). Fingerling production in 1995 was 0.8 million and in 1996 was 0.42 million (Bartley and Rana, 1998a; 1998b). In 2002 and 2003, 344,000 and 325,000 fingerlings were released at a cost of 2125 and 2288 million rials respectively (Salehi, 2008a). The "Gharasoo" Research Station in Sari is using cage culture of this species (Madbaygi, 1993b). Smolts weighing 25 g on average were released into cages and fed on locally prepared food (Iranian Fisheries Research and Training Organization Newsletter, 4:5, 1994). Culture of a rapid-growing triploid sea trout has been studied in Iran (Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 32, 1996; Annual Report, 1995-1996, Iranian Fisheries Research and Training Organization, Tehran, p. 67, 1997). Sayyad Borani et al. (2006) studied the effects of weight on the osmoregulatory ability of juveniles in an attempt to determine the ideal weight for release (see also above under Habitat). Weight classes were 5, 10, 15 and 20 g and the three larger classes were capable of osmoregulation in Caspian Sea water. Mojazi Amiri et al. (2005) detailed the histology of the digestive tract from hatching to parr stage in order to better understand the digestive system capabilities and enhance rearing. Asaeian et al. (2006) studied brood fish taken from the early mid and late migrations, finding those from the early period were more suitable for offspring production, passing through smoltification more rapidly. However, they noted that fish from all periods should be used to maintain genetic diversity. Bahrkazemi et al. (2006) traced the histochemical changes in the digestive tract from hatching to parr stage. Javaheri Baboli et al. (2006) examined the effects of n-3 HUFA enriched Artemia nauplii as a starter food and showed it was no better than newly hatched Artemia. Sarvi et al. (2006), Hatef et al. (2007), Niksirat et al. (2007) and Sarvi Moghanloo et al. (2007), have studied cryopreservation of sperm, sperm and seminal plasma composition, and in vitro storage of unfertilized ova in Iran as means to help conserve wild populations. Research into osmotic regulation determined that fingerlings 10 g or above have a higher chance of survival (Iranian Fisheries Research Organization Newsletter, 53:4, 2008). Ataei Mehr et al. (2007) studied the effects of weight and salinity on the number and size of Malphigian bodies in juvenile kidneys which has relevance for release of cultured fish in fresh and salt water. Saber et al. (2007) found that for larvae a diet with 50% protein and 4600cal/g energy was appropriate. Jamalzadeh et al. (2008) established haematological and serum biochemical indices for smolts, juveniles and breeders. Zamani et al. (2007) found differences in activity of some digestive enzymes in the stomach, pyloric caeca and intestine of parr and of smolt. Bahrekazemi et al. (2009) studied the relation between egg viability, egg and ovarian fluid composition and time of stripping in fish at the Kelardasht Hatchery. Eyeing and hatching rate declined with over-ripening time  although larval abnormalities remained constant. The best time to take eggs was up to 10 days post-ovulation. Mahmoudi et al. (2009) fed different levels of dietary nucleotides showed positive effects on growth performance and a decrease in liver demolition. Ramezani (2009) fed fish experimental diets and found a better feed conversion ratio and specific growth rate at lower protein levels, 50% protein supporting maximum growth. Bahre Kazemi et al. (2010) investigated the use of biochemical and histological parameters as egg quality biomarkers. Hajirezaee et al. (2010a; 2010b, 2010c) examined the chemical and physical properties of seminal fluid and its effect on sperm motility, and effects on semen characteristics of various stripping frequencies. Oulad et al. (2010) looked at the effects of different levels of dietary nucleotides on osmoregulation of pyloric caeca in in juveniles. Rahbar et al. (2010, 2011) examined three female broods (4, 5 and 6 years old) and found that the 6 year olds showed maximum average fertilisation (97.5%), survival rate until eyed stage (92%), hatching (93%) and survival rate until absorption of the yolk sac (94.5%). Ahmadi et al. (2011) experimentally infected trout with the bacterial pathogen Aeromonas hydrophila and studied blood and tissue changes. Maniei et al. (2011) investigated the selective use of body fatty acids by parr under feeding, starvation and re-feeding regimes. Sotoudeh et al. (2011) found that 4% dietary phosphatidylcholine had positive effects on growth, nutrition factors and liver fatty acid composition of alevins. Amiri Moghaddam et al. (2012) found starvation had negative effects on ion regulatory capacity of parr and re-feeding did not fully compensate for this. Sourinejad et al. (2012) examined the effects of milt volume equalization, which did not result in a balanced contribution of breeders and also produced large variations in contribution to progeny. Toorchi et al. (2012) found that larger juveniles performed better in response to salinity challenges, concluding that releasing large juveniles in stocking programmes in estuaries was safe but that smaller juveniles should be released into rivers or be acclimatised to estuarine conditions before release there.

Salehi (2008a) carried out a cost factor analysis for fingerling production at the Kelardasht Hatchery in Iran and found labour costs to be 52% and feed 16%. The average cost of production of a fingerling was U.S.$0.84 (6269-7123  rials), more than that for sturgeon (1753-2028) and kutum (54-121). Higher costs are probably associated with lack of broodstock, lower production quantities and the scale of the hatchery. The rate of return at age 3-4 years was estimated at 0.5% of those released annually.

RaLonde and Walczak (1970b) and Walczak (1972) lists the following reasons for decline of this species: lowering of the Caspian Sea level making spawning migrations difficult, habitat changes including siltation, pollution, irrigation dams, and unscreened irrigation intake canals, and poaching. Both adults and freshly released fingerlings are heavily poached (Petr, 1987). In the Lar and Karaj rivers about 1960 professional fishermen took trout using nets, chemicals and explosives until laws regulating seasons and equipment were passed and game wardens hired to enforce them (Surber, 1969). Streams running into the Caspian have their lower 1-3 km dried up by May through irrigation demands and trout are unable to reach the sea. Stronger enforcement of laws by game guards, education of the public, particularly local people, habitat protection and improvement, and a rational exploitation system are required to protect this species. Firouz (1974; 1976) reported that the sea-run form was a major commercial species 25 years prior to his account but the deteriorating environment coupled with dynamiting, netting and trapping severely reduced populations. This fish is now completely absent from the Anzali Mordab (Holčík and Oláh, 1992). Waters where populations survived were designated "Protected Rivers" in an effort to manage this species effectively. There are special licence requirements, bag limits and seasons which anglers must observe. A fine of 10,000 rials is imposed specifically for illegal fishing of this species (Anonymous, 1977-1978) and each fisherman is now limited to 3 fish per day with the season lasting from 11 July to 10 September for two days a week (Salavatian et al., 2011).

The Liqvan Chay population has been confined to this single river by, it is believed, destruction of habitat through agriculture and domestication of sheep and goats (Anonymous, 1977). De Mecquenem (1908) said trout were abundant in the upper reaches of rivers in the Lake Orumiyeh basin. In 1975, 500 specimens of this subspecies were transplanted into the Ab-e Bazoft near Shiraz in an effort to conserve it (Anonymous, 1977).

Caspian salmon used to enter the Kura River basin, with 20% running up the Aras River on the Iranian border. Most spawning beds on the Aras became inaccessible in the 1950s with the construction of the Bagramtapinskaya Dam. The Mingechaurskaya Dam on the Kura cut off 80% of that rivers spawning beds in 1951. The former Soviets built salmon hatcheries to offset the losses but these proved to have a low efficiency. The young fish had a high mortality because of release at unsuitable high temperatures, release in poor feeding areas, and exposure to water intakes. In addition, the released fish were too small or too few to undergo significant smoltification and migration to ensure subsequent reproduction. Under natural conditions only a small proportion of juveniles undergo smoltification (Bakshtanskii et al., 1973).

Kiabi et al. (1999) and Jalali et al. (2009) consider S. caspius to be critically endangered in the south Caspian Sea basin according to IUCN criteria. Criteria include sport fishing, few in numbers, habitat destruction, medium range (25-75% of water bodies), absent in other water bodies in Iran, and absent outside the Caspian Sea basin. Kiabi et al. (1999) also consider S. t. fario to be vulnerable in the south Caspian Sea basin according to IUCN criteria. Criteria include sport fishing, medium numbers, habitat destruction, medium range (25-75% of water bodies), present in other water bodies in Iran, and present outside the Caspian Sea basin. Coad (2000a), using 18 criteria, found this species to be one of the top 4 threatened species of freshwater fishes in Iran. Nezami et al. (2000) consider this species to be endangered because of overfishing, habitat destruction and spawning ground degradation. Mostafavi (2007) lists it as vulnerable in the Talar River, Mazandaran.  Critically endangered in Turkey (Fricke et al., 2007). Jalali et al. (2009) recommend reduction in pollution, waters where fish are released should be designated Protected Rivers, cage culture should be enhanced, spawners from different populations should be used to enhance genetic diversity in restocked populations, and gamete storage protocols could be applied to commercial aquaculture or to a biological conservation programme. Shirangi et al. (2011) note that successful propagation is of fall run fish and the numbers of spring run fish have declined.

Osinov and Bernatchez (1996) note that populations within the Caspian Sea basin have unique gene pools, suggestive of reproductive isolation, and these fish cannot be managed as a single subspecies, S. trutta caspius. Re-introductions and artificial maintenance of native populations could lead to loss of diversity. Yousefian et al. (2012) found a high heritability of body weight and length using broodstock of different age and weight.

Further work

Togan et al. (1995) show that two Turkish populations of brown trout are genetically distinct, at a level often found between species. Genetical analysis of suspected surviving pure populations of Iranian Salmo caspius should be carried out to determine which stocks should be receive special attention for conservation.

Sources

Iranian material:- CMNFI 1970-0551, 5, 109.9-166.3 mm standard length, Gilan, Ghaleh River near Fowman (37º13'N, 49º19'E); CMNFI 1979-0086, 2, 76.5-145.5 mm standard length, Mazandaran, Hasanabad River tributary to Chalus River (no other locality data); CMNFI 1980-0133, 3, 116.8-139.2 mm standard length, Markazi, Karaj fish hatchery (no other locality data); CMNFI 1980-0158, 1, 93.3 mm standard length, Markazi, Polurd River at Damavand Mountain after Ab-e Ali (35º51'N, 52º04'E); CMNFI 2007-0125, 3, 91.2-169.3 mm standard length, Markazi, Luniz River in Karaj River basin (no other locality data); CMNFI 2007-0126, 7, 95.2-158.3 mm standard length, Azarbayjan-e Khavari, Aldarvish River, Sabalan Mountain (no other locality data); BM(NH) 1877.7.5:1, 7, 168.1-194.7 mm standard length, Tehran (no other locality data); BM(NH) 1908.8.7:27-28, 2, 129.0-145.7 mm standard length, north slope of Elburz Mountains near Tehran (no other locality data).

Salmo tigridis
Turan, Kottelat and Bektaş, 2011

This species is recently described from the Tigris River basin of Turkey (Çatak Stream - and see below under Salmo trutta for some biology). It probably exists in Iranian waters but requires confirmation by specimens. The photograph below is a fish from the Tigris River basin.

Bazoft River, upper Karun River basin, August 2009, courtesy of K. Abbasi
Bazoft River, upper Karun River basin, August 2009, courtesy of K. Abbasi


Bazoft River, upper Karun River basin, August 2009, courtesy of K. Abbasi

Salmo trutta
Linnaeus, 1758

European Salmo trutta, Duane Raver, U.S. Fish and Wildlife Service
European Salmo trutta, Duane Raver, U.S. Fish and Wildlife Service

Common names

See above under Salmo caspius where general names for trout will encompass this species.

[Brown trout, German brown trout, European brown trout, sea trout, brownie, Loch Leven trout, Von Behr trout, spotted trout and liberty trout].

Systematics

See above under Salmo caspius.

Key Characters

?This species is a relative of the Atlantic Salmon and is distinguished from it by the upper jaw extending beyond the eye in adults (and below rear half of the eye in young rather than the centre in young), the gill cover has many spots, dorsal fin principal rays usually 10-14, and orange to rusty-red spots are often present on adult flanks.

Morphology

Principal anal fin rays 9-12, pectoral rays 13-14 and pelvic rays 9-10. Lateral line scales 110-136 and pyloric caeca 30-60. Males develop a hooked lower jaw when spawning and the brown colour becomes more intense and golden.

Sexual dimorphism

Males develop a hooked lower jaw when spawning and the brown colour becomes more intense and golden.

Colour

Overall colour is a light to golden brown with silvery flanks and a white to yellowish belly. The back, flanks, side of the head and dorsal and adipose fins bear black or dark brown spots, often with a lighter halo of orange, pink or red, and the flank has pink or red spots. Only brown trout have both light and black spots on the flanks. The caudal fin may have spots restricted to its upper lobe but lacks the overall radiating spots of rainbow trout. The adipose fin is orange to orange-red, the only family member with this colouration. Sea run or lake fish are more silvery, obscuring some of the spots. Spots may be x- or y-shaped. The red flank spots usually have blue halos. Young have 7-14, narrow parr marks and a few red spots along the lateral line. The adipose fin is orange with a light margin.

Size

Reaches 1.4 m and reputedly 50 kg for sea-run fish. Note that as the taxonomy of trouts changes this maximum may not apply.

Distribution 

Brown trout were artificially planted in Gahar Lake of the upper Dez River of the Tigris River basin where viable populations existed in the 1970s in both the upper and lower lake, and more recently (B. Sandford, in litt., 1979; R. Mehrani, pers. comm., 2000). European brown trout were planted in the Caspian Sea and Namak Lake basin and established south of Dorud in the Zagros, and in the Zayandeh River dam but their origin is unknown (Coad and Abdoli, 1993). Trout were also introduced to the Karun River basin and the Zayandeh River Dam (Y. Keivany, in litt., 1992). Trout are also recorded from the the Lake Orumiyeh basin in the upper Talkheh, Zarreineh and Tatavi rivers (Abdoli, 2000) but whether these are introduced is not certain.

Zoogeography

This species is introduced in Iran. 

Habitat

Brown trout are mostly stream and river dwellers although some are in lakes and ponds. They can tolerate warmer and more turbid waters than brook trout and only the tainbow trout is more tolerant among salmonids. Brown trout may survive in areas no longer suitable for Brook Trout because deforestation has increased stream temperatures and agriculture and industry have increased pollution and turbidity. Ideally there should be overhanging and submerged vegetation, coarse gravel and cover such as logs, boulders and undercut banks, growth is better in alkaline waters (20-200 mg L-1, total dissolved solids 800 mg L-1 for culturing this species), a depth of 7-58 cm is needed for spawning, a winter flow of less than 15 cm sec-1 is required, preferred temperatures are 10.0-17.6°C (other reports give 21.1°C and temperatures up to 24°C are tolerated) and for spawning 6.7-8.9°C, dissolved oxygen is optimal at 7-9 mg L-1 and pH range tolerated is 5.0-9.5 although the optimal range is 6.8-7.8.

Age and Growth

Life span is over 18 years with maturity attained at 2-4 years. Çetinkaya (1999) found the fish in the Çatak stream, of the upper Tigris River, Turkey to have an age range of 1-8 years, a fork length of 8.4-39.0 cm, a weight of 6.7-756 g, and a sex ratio 2.45:1 male:female. This population was described as S. tigridis (see below) and is not the same taxon as introduced populations in Iran.

Food

Food is aquatic and terrestrial insects, crustaceans, molluscs, various fishes, frogs, salamanders and even small mammals. Most food is taken as drift, the trout positioned in slower water behind a rock and darting out into faster current to seize prey. Predators include other fishes, birds, water-snakes and otters. Çetinkaya (1999) found the diet of fish in the Çatak stream, of the upper Tigris River, Turkey to be Trichoptera, Ephemeroptera, Gammarus, chironomid pupae and larvae, and Plecoptera.

Reproduction

Spawning occurs from October to January depending on locality, usually at 7-9°C but as low as 2°C or as high as 14°C. It usually takes place in gravel stream riffles but may occur on rocky shores of lakes. The female excavates a redd into which the spawning pair deposit eggs and sperm while gaping and quivering over a 4 second period. The female covers the redd with gravel after spawning to protect the eggs. Subsequent spawning occurs, usually after a 10 hour interval. Occasionally a community redd is excavated by a number of fish spawning close together. Eggs are up to 5.0 mm in diameter and each female can contain 20,865. Young may spend about 2 years in their natal stream before going to a lake but some fish remain permanently in streams.

Çetinkaya (1999) found the fish in the Çatak stream, of the upper Tigris River, Turkey, to be sexually mature at age 4 and 16-17 cm fork length with an individual fecundity of 2349 eggs.

Parasites and predators

None known in Iran.

Economic importance

This trout is a valuable and popular sport fish in Europe, The benefits of stocking brown trout include its difficulty of capture (and thus appeal to anglers), its longer contribution to the fishery than brook trout and rainbow trout, better survival and growth than other trouts, and its tolerance of warmer waters. There is an extensive European literature on angling methods and it is reputed to be a wilier fish than brook trout, for example. They tend to bite best in the late evening or at dawn, especially when large. These trout are caught on worms, crayfish, lures and flies. Brown trout have white to pink flaky flesh and are excellent eating. Flesh colour changes with age, to pink, and is related to diet.

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in aquaria and aquaculture, as food, in sport, in textbooks and because it has been widely introduced outside its natural range. Salmo trutta is reported to be ichthyootoxic although there are no reports for Iran or for S. caspius (Coad, 1979b; see also under the genus Schizothorax for symptoms of this egg poisoning).

Conservation

None required fro an exotic species.

Further work

The distribution and biology of this trout in Iran needs detailed investigation to separate it from native taxa.

Sources

Based on general accounts of the species.

Genus Salvelinus
Richardson, 1836

The charr genus comprises many species in Europe, northern Asia and North America whose systematics have not been fully worked out. FishBase lists 83 nominal species (August 2007). The charrs have numerous small scales 105 or more in lateral series), teeth on the jaws, palatines and tongue, teeth are present on the head but not the shaft of the vomer bone in the roof of the mouth, scales in the lateral line are smaller than surrounding scales and have little or no overlap with scales before and behind, the body has light pink, red or cream spots, and the lower fins have snow-white leading edges.

Salvelinus fontinalis
(Mitchill, 1814)

Duane Raver, U.S. Fish and Wildlife Service
Duane Raver, U.S. Fish and Wildlife Service

Common names

ماهي آزاد چشمه اي (= mahi azad cheshmehi or azad mahi cheshmehi, meaning spring free fish, i.e. spring salmon, free fish being used for salmon and trout species in Farsi), qezel ala-ye juibary (= brook trout).

[brook trout, brook charr, speckled trout].

Systematics

Salmo fontinalis was originally described from the vicinity of New York city.

Key characters

This species is characterised by 109-132 lateral line scales, 8-13 anal fin principal rays, light-coloured spots on the body, teeth on the head of the vomer bone in the roof of the mouth, pectoral, pelvic and anal fins with a white leading edge followed by contrasting black, truncate caudal fin, dorsal and caudal fins have wavy, dark lines and blotches and the back has dark or light green or cream, worm-track markings (vermiculations).

Morphology

Dorsal fin principal rays 9-14, pectoral rays 10-15 and pelvic rays 7-10. Scales are minute, horizontal and irregular ovals with a central focus, few circuli and no radii. The pelvic axillary scale is well-developed. Gill rakers 13-22, reaching the second adjacent raker whem appressed. Pyloric caeca 20-55. The gut is s-shaped. The chromosome number is 2n=84 (Klinkhardt et al., 1995).

Sexual dimorphism

Spawning males develop a hooked lower jaw or kype.

Colour

The back is olive-green to dark brown or blackish fading to a silvery-white belly. Flanks have a red to yellow tint. Flank spots are pale but there are also small, red spots with blue halos. The pectoral, pelvic and anal fins are yellow, orange, or reddish behind the white and black leading edges. Sea-run fish have a blue-green back and silvery flanks with a purplish tinge. Spots are obscured except for the red ones. Brook trout in large lakes are also more silvery than stream resident fish. The jaw tips and the roof of the mouth are blackish. Spawning males are much brighter in overall colour and have an orange-red lower flank and upper belly, bordered below by black on each side which delimits the white belly. Young have 6-12 brown parr marks, the widest equal to eye diameter, and small red, yellow or blue flank spots. The white leading edge to the lower fins is apparent.

Size

Attains a reputed 86.0 cm and 8.0 kg, possibly 9.39 kg, but most are smaller.

Distribution

In North America this species is found from the shores of Hudson Bay and Labrador south in marine waters to Cape Cod in the east and Georgia in the Appalachian Mountains, west through all the Maritime provinces, Québec and Ontario (except the extreme west) to northeast Manitoba. Widely introduced to western Canadian provinces, the U.S.A., South America, Europe, Asia and Australasia.

A private hatchery on the Jajrud imported over 1 million brook trout eggs which were raised to fingerling size only for most to be lost in a flood in 1968. Some were planted in the Jajrud and in the Latian Reservoir in the Namak Lake basin. Survival remains unknown. Also recorded from the Sardab and Chalus rivers of the Caspian Sea basin (Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 26, 1996) but this is possibly a misidentification for Oncorhynchus mykiss.

Zoogeography

An exotic species from North America, not closely related to native Iranian salmonids.

Habitat

Brook trout are found in cool waters of streams and lakes, usually less than 20°C, adults preferring 14-16°C. Spawning requires temperatures below 15°C, and below 9°C for optimal hatching. The upper lethal limit is 25°C for adults and 20°C for newly hatched fish. pH range is 4.1-9.5. Pools, underneath banks, under overhanging bushes or behind rocks are favoured spots. During summer months they retreat to deeper water, to about 8 m, in lakes. Some populations run to sea in Hudson Bay and Atlantic Canada. They stay in coastal waters, not moving more than a few kilometres from their natal stream. Populations in the North American Great Lakes live and feed mostly in the lake and run up natal streams to spawn.

Age and growth

Maximum life span is over 20 years but most reach only 5 years in North America. Maturity is attained at 2-3 years, some males at 1 year. Stunting is common in small streams while sea run fish grow faster than freshwater ones. Optimum growth temperatures are 10-19°C.

Food

Food in North America includes aquatic and terrestrial insects, molluscs, various fishes, frogs, and even snakes, mice, voles and shrews. Stream-dwelling fish feed heavily on drifting aquatic organisms during spring run-off but in summer as drift decreases surface insects become important. Most feeding occurs in the early morning and late evening although some food is taken throughout the day. Diet shifts in response to competition with other species. Sea run fish take various marine invertebrates and fishes. Sea run adults in spring and summer eat crustaceans and fish in lower estuarine areas while young are in the upper estuary eating crustaceans and insects. During the fall in the river adults eat little and during winter back in the estuary consumed mostly crustaceans. There is thus a division of food resources between young and adults.

Reproduction

Spawning in North America takes place from August to December, earlier in the north and later in the south. Sea-run charr enter their natal stream in spring and summer even though spawning occurs in fall. Each year they spend 1.5-3 months feeding in the sea. The spawning ground is usually gravelly streams but may be lake shoals if there is some current or spring outflow to keep eggs oxygenated. Spring flows are preferred even in streams. Males arrive on the spawning ground first and defend a territory. Both sexes will rush at other fish entering the redd area. The female cleans a redd of debris by turning on her side and lashing her tail. Redd depth between stones is tested by inserting the anal fin. Redd construction may take up to 2 days with work being carried out both by day and night. Courtship involves gentle pushes, touches and strokes of the female by the male. The female is ready to spawn when she crouches in the redd with her genital area between the stones. The male arches his body and may press the female against the redd bottom, both fish vibrate and eggs and sperm are shed. Accessory or sneaky males may rush in to shed sperm. The female lashes her tail to push eggs into the gravel and then dislodges gravel with her anal fin to cover the eggs to depths as great as 20 cm. Yellow-orange eggs are up to 5.0 mm in diameter and number up to perhaps 17,000 per female although averages range from the low hundreds to a few thousand. Both sexes may spawn again with other fish. The eggs develop over winter, taking 165 days at 2.8°C but only 47 days at 10°C. Temperatures above 11°C will kill the eggs.

Parasites and predators

Brook trout are cannibals on their eggs and young and are eaten themselves by other fishes, water snakes, turtles, various birds and otters.

Economic importance

Brook trout are very popular sport fish in North America caught on lures, live baits and flies. These trout are easier to catch than Salmo trutta sensu lato and take a wider range of lures. They fight well but are often quite small and do not leap spectacularly like some other members of the salmon family.

The ready availability of this salmonid has made it a useful experimental fish for various physiological, biochemical, toxicological and other studies. Some reared stocks however show deformed or lost fins and distorted mouths.

Conservation

This introduced species is not in need of conservation but in fresh water their preference for cool and clear water makes them susceptible to loss if waters are dammed, channelised and polluted or if banks are eroded deforested and overgrazed, conditions not uncommon in Iran.

Further work

The survival of stocks of this species in Iran should be verified and the effects of this exotic on native species researched.

Sources

Based on general summaries on North American populations such as those in Scott and Crossman (1973) and Becker (1983) as no Iranian material to hand. Iranian populations have not been studied.

Genus Stenodus
Richardson, 1836

The inconnu genus contains a single species found in waters draining to the Arctic Ocean in Eurasia and North America and in the Caspian Sea.

Its characters are essentially those in the species description; a small bone lacking a head canal on the outer side of the lower jaw at the joint of the articular and dentary is distinctive. Branchiostegals number 8-11.

Stenodus leucichthys
(Güldenstaedt, 1772)

Common names

safid mahi (= white fish), آزاد ماهي (azad mahi or free fish), azad mahi Volga (= Volga free fish, or Volga salmon, free fish being used in Farsi for salmon), زيبا (= ziba), mahi-ye ziba or ziba mahi (= beautiful fish, in northwest Gilan, e.g. at Astara), mahi-e-azad-e-ziba, inkonu.

[belorybitsa in Russian; nelma or azatmahy in Turkmenian; inconnu, conny, white salmon, Caspian inconnu].

Systematics

Salmo leucichthys was originally described from the Volga and Ural rivers, Caspian Sea and Kamchatcka, Russia.

The type subspecies in the Caspian Sea basin is the one found in Iran. The subspecies Stenodus leucichthys nelma (Pallas, 1773) of White Sea, Siberian and northwestern North American drainages is more common.

Key characters

The characters of the species are those of the genus.

Morphology

Dorsal fin branched rays 8-13 after 2-6 unbranched rays and anal fin branched rays 9-16 after 2-5 unbranched rays. Lateral line scales 88-121, scales above the lateral line 8-13, and scales below the lateral line 10-12. There is a pelvic axillary scale. Scales bear numerous fine circuli, the focus is slightly subcentral posterior, the anterior scale margin is wavy but irregular in outline and the scale is generally rounded. There are no radii. There is a single flap between the nostrils.Gill rakers number 17-27 and reach the fourth adjacent raker when appressed. Pyloric caeca number 191-193 and the gut is s-shaped. Vertebrae number 64-70.

Sexual dimorphism

Males develop tubercles on the head and sides of the abdomen during spawning.

Colour

Overall colour is silvery without spots.

Size

Reaches 1.5 m and 28 kg, rarely 40 kg, possibly to 48 kg. On the coast of Dagestan the average size was 7.6-10.8 kg in the 1920s (Berg, 1948-1949).

Distribution

The subspecies is restricted to the Caspian Sea and its drainage. Rare in Iranian waters with 5 specimens caught and preserved from 5-7 km east of Bandar-e Anzali in the Caspian Sea from 1984-1990. Several dozen fish have been caught each year in Iran, starting in the late 1960s, all from between the mouths of the Safid and Astara rivers (Holčík and Razavi, 1992). Reported from the southeast Caspian Sea, southwest Caspian Sea and south-central Caspian Sea (Kiabi et al., 1999; Abdoli and Naderi, 2009).

Zoogeography

This species is found naturally in the Caspian Sea but was generally believed to inhabit only the northern, and rarely the western, parts to the north of 40°N. Rare specimens were reported by Kazancheev (1981) from the coasts of Dagestan and Azerbaijan in early summer. It probably penetrated into the Caspian Sea from the Arctic Ocean basin during the last glacial period via ponded lakes.

Habitat

Inconnu live in the Caspian Sea proper at depths less than 60-65 m (optimally 25-45 m) and migrate up rivers to spawn, in the Volga for 3000 km. Fat content of fish at Astrakhan at the Volga mouth was 26% of total weight but by the time fish reached the spawning grounds in the Ufa River this had fallen to 14%, and after spawning to 1.5% for females and 1.6% for males. Off Iran they are caught mostly above depths of 25 m from October until the end of December. Some are caught in beach seines in shallow water (Holčík and Razavi, 1992) but elsewhere are known from depths down to 46 m. It is said to move into the central and southern Caspian in summer and returns north when water temperatures fall below 8-10°C in the first half of September (Berg, 1948-1949). This contrasts with the catch records for Iran. Fingerlings tolerate temperatures up to 25ºC in culture ponds but in the sea it is found at temperatures below 20ºC (Caspian Sea Biodiversity Database, www.caspianenvironment.org).

Age and growth

Life span is at least 12 years in the Volga population for females and 8 years for males. Iranian fish were up to 6 years old (Holčík and Razavi, 1992). Spawning females are predominately 6-8 years and males 5-6 years old (Letichevskiy, 1981). Females on the Volga spawning grounds are 85-105 cm, 5.5-9.5 kg while males are 75-100 cm, 3.5-7.5 kg.

Food

There is no feeding on the spawning migration, the fish relying solely on their stored reserves of fat. Fat declined in one study from 21% before, to 2% after, spawning. In the Caspian Sea it feeds on fishes such as gobies (Gobiidae), Rutilus rutilus (presumably R. caspicus), Atherina boyeri (= caspia), young Sander, common and anchovy kilkas, and shads. In the sea, adults feed on kilka and silversides (97-99%)(Caspian Sea Biodiversity Database, www.caspianenvironment.org). Young feed on zooplankton 7-10 days after hatching but are predatory in their second month. Fry descend to the Caspian Sea to feed until they are sexually mature.

Reproduction

Spawning takes place in October and November at 0.2-6.0°C after an ascent up north Caspian rivers. The Volga River run peaks in December and March. Fecundity reaches 390,000 eggs of up to 2.4 mm diameter. Eggs average 26% of the female's weight. The eggs are slightly adhesive and shed on the river bottom. Spawning occurs only once or twice in the life cycle of most fish although exceptional females may spawn 3 times. Spawning occurs at intervals of 2-3 years. Incubation takes 180-200 days and hatching takes place in spring. The young immediately descend to the sea.

Parasites and predators

Eggs are eaten by Gobio species and by Lota lota.

Economic importance

This species has no economic importance in Iran but was a significant member of former Soviet fisheries, taken both in the sea and on its spawning migration. Up to 110,000 fish were taken annually in the Volga-Caspian region. It has a rich, delicate flesh with a fat content of 18-26% and is a favoured smoked product (Caspian Sea Biodiversity Database, www.caspianenvironment.org).

Conservation

This species was listed as threatened in Southwest Asia by Holloway (1976) and endangered in the Caspian Sea by Fricke et al. (2007). Kottelat and Freyhof (2007) state that it is maintained only by stocking and is on the brink of extinction. The population of this species, spawning principally in the Volga, declined drastically after regulation of this river. In the 1960s, the population was estimated to be only 2000 fish but it has recovered to 17,000 fish through artificial rearing (Mina, 1992). In the two decades 1961-1980, fish hatcheries in the Volga Delta raised 72.4 million inconnu. Gravel spawning grounds were also constructed (Letichevskiy, 1981). It has been proposed for inclusion in the "Red Book of the U.S.S.R." which forms the basis for measures to protect species (Pavlov et al., 1985). Lelek (1987) classified it as endangered. Kiabi et al. (1999) consider this species to be data deficient in the south Caspian Sea basin according to IUCN criteria. Criteria include commercial fishing, sport fishing, few in numbers, bodies), limited range (less than 25% of water bodies), absent in other water bodies in Iran, and present outside the Caspian Sea basin.

Further work

Further records of this species should be recorded to document the presence and spread of this fish in Iranian waters.

Sources

Morphology based on Holčík and Razavi (1992) for Iranian specimens.

Iranian material: None.

Comparative material: BM(NH) 1878.12.26:23, 1, 199.5 mm standard length, Bremen (no other locality data); BM(NH) 1899.7.25:22, 1, 215.5 mm standard length, Russia, River Obi (no other locality data).

Esocidae

The pikes, pickerels and muskellunge are found in fresh waters of the Northern Hemisphere. They are moderate to large-sized fishes, up to 1.4 m. There are only 5 species in a single genus (Eschmeyer and Fong, 2011) with 1 species reported in Iran.

The family is characterised by a flattened, elongate, duck-billed snout; dorsal and anal fins far back on the body near the tail; no adipose fin; teeth on the tongue and on the basibranchial bones behind the tongue are small; jaws have large teeth; branchiostegal rays 10-20; nasal bones are present; the swimbladder is connected to the gut by a duct; intermuscular bones are forked or y-shaped; no fin spines; pelvic fins are abdominal; cycloid scales; the infraorbital sensory canal on the head has 8 or more pores; gill rakers are present as sharp denticles in patches; no pyloric caeca; the lateral line is complete; and the forked caudal fin has mostly 17 branched rays.

Pikes are predators on other fishes aided by the posterior dorsal and anal fins which facilitate rapid darts forward. They are important sport fishes, much sought after by anglers for their fighting ability, but are not very good eating because of the intermuscular bones.

Genus Esox
Linnaeus, 1758

The characters of this genus have been outlined above under the family.

Esox lucius
Linnaeus, 1758

Female, 64 cm and 3.5 kg, from drainage canal near Abkenar, Anzali Mordab, courtesy of S. Nouripanah
Female, 64 cm and 3.5 kg, from drainage canal near Abkenar, Anzali Mordab,
courtesy of S. Nouripanah


Anzali Wetland, June 2012, courtesy of K. Abbasi
Anzali Wetland, June 2012, courtesy of K. Abbasi

Anzali Wetland, June 2012, courtesy of K. Abbasi
Anzali Wetland, June 2012, courtesy of K. Abbasi

Anzali Wetland, June 2012, courtesy of K. Abbasi
Anzali Wetland, June 2012, courtesy of K. Abbasi

Courtesy of K. Abbasi
Courtesy of K. Abbasi

 

Common names

ordak mahi (= duck fish from the snout shape), shok, shuk or shook (in Gilaki), shook chehkhab, chekab.

[durnabaligi in Azerbaijan; shchuka in Russian; pike; northern pike].

Systematics

Esox Lucius was originally described from Europe.

Key characters

The broad and flat snout and the dorsal and anal fins set far back on the body are distinctive.

Morphology

Dorsal fin principal rays 15-19, about 6-10 unbranched and 13-18 branched, principal anal rays 12-16, about 4-8 unbranched and 10-15 branched, pectoral rays 11-17 and pelvic rays 7-13. The number of branched rays may be size-related as in smaller fish more anterior rays in the dorsal and anal fins are not branched. Lateral line scales 105-148, pored scales 42-56, but difficult to count accurately. Each scale on mid-flank is a rounded rectangle. The anterior margin is indented where 1-2 radii terminate. The radii split the scale so that the segments overlap. Circuli are very fine and the focus is posterior. Gill rakers are broad and spinulose, embedded in the arch skin with the tips of the spinules protruding. There are 9-11 pores on the lower jaws (usually 5 on each jaw). Vertebrae 56-65. The chromosome number is 2n=50 (Klinkhardt et al., 1995). The gut is an elongate s-shape.

Meristic values for Iranian specimens are:- dorsal fin branched rays 14(7) or 15(9); anal fin branched rays 11(1), 12(8) or 13(7); pectoral fin branched rays 14(9), 15(5) or 16(1); pelvic fin branched rays 8(1), 9(7), 10(6), 12(1) or 13(1); pores on each lower jaw 5(22).

Sexual dimorphism

There is no obvious sexual dimorphism. Attempts have been made to sex pike by characters of the urogenital region, but these are hampered by seasonal variations. Abdurakhmanov (1962) reports on fish from Azerbaijan where head length is greater in males while predorsal distance and interorbital width are greater in females.

Colour

The overall colour is dark with light spots, although there is variation over the vast range of this species in the details. The back and upper flank are dark green, olive-green or brownish, fading to a whitish belly. The flank has 7-9 rows of greenish, yellow to whitish blotches along it. Scales have a golden tip. The head sides have wavy, golden or yellow blotches and lines and the eyes are bright yellow to golden. The dorsal, anal and caudal fins are green, yellow, orange or pale red, blotched and barred irregularly with black. The pectoral and pelvic fins are dusky to orange. Young have 8-12, wavy, white or yellow bars which become the bean-shaped blotches in adults as they gradually break up. There is a gold to green stripe along the middle of the back in some fish but others are completely dark green. There is a stripe below the eye. The peritoneum is silvery.

Size

Attains 1.75 m and about 48.0 kg, possibly to 2.13 m and 65.78 kg, despite legends of pike up to 5.0 m (Tsepkin, 1986; Machacek (1983-2012), downloaded 27 July 2012).

Distribution

Across northern Eurasia and northern North America. Iranian populations are at the southern edge of the range for this species. Found in the Caspian Sea basin, from the Anzali Mordab and its drainage canals, tributaries and the Sia Keshim Protected Region to Gorgan Bay and its tributaries such as the Karasu, with other rivers including the Safid, Chalus, Tajan, Tonekabon, Babol and Haraz, and the Atrak River basin, Boojagh Wetland and the Amirkelaye Lagoon near Lahijan (Derzhavin, 1934; Berg, 1936; Armantrout, 1980; Holčík and Oláh, 1992; Nejatsanatee, 1994; Riazi, 1996; Abbasi et al., 1999; Kiabi et al., 1999; Abdoli, 2000; Jolodar and Abdoli, 2004; Abdoli and Naderi, 2009; Jamalzadfalah et al., 2011; Khara et al., 2011). However, Abdoli and Naderi (2009) do not report it from the Atrak River.

This species has been introduced to the the Avan River, 7 km from Alamut as early as 1956 (Nialmir, 2001), and to Valasht Lake near Marzanabad, Avan River and Evan Lake northeast of Qazvin, the upper Karaj and middle Shur (Abhar stretch) rivers of the Namak lake basin, the Avan River near Alamut (in 1956), Ghorigol Lake near Tabriz, Marivan Lake in Kordestan, and the Haft Barm Lakes west of Shiraz (Anonymous, 1977; Petr, 1987; Niamir, 2001).

Zoogeography

This widely-distributed species reaches its southern range limit in Iran.

Habitat

Pike are solitary and are found in lakes and rivers where the water is still or flowing slowly as well as marshes and ponds. They are found only in the lower reaches of rivers along the Iranian shore and do not penetrate upstream (Berg, 1948-1949). O'Donovan (1882) reported that a small stream in the Atrak River drainage had many large pike lurking under bushes, stupefied by foul water, and that the Cossacks in his escort caught many of them by striking with the point of their sabres or simply whisked them out of the water by the tail. Large numbers of Rutilus frisii were seen here too and presumably the pike thrived on this food source. Riazi (1996) reports that this species is native (resident) to the Siah-Keshim Protected Region of the Anzali Mordab. They are found in the more brackish areas of the Caspian Sea, at least in Kizlyar Bay of the north Caspian, at salinities up to 4‰ under the influence of fresh water from the Volga River. Here they are found in thickets of soft and rigid vegetation, as solitary predators but also feeding beyond the vegetation limit in the sea. They aggregate only in spring for spawning and in autumn prior to wintering on the bottom (Stolyarov and Abusheva, 1997).

In fresh water, vegetation is heavy and the water warm but they usually retire to deeper, cooler water at the height of summer. Temperatures above about 30°C are usually fatal to pike. However pike are active in winter, as anglers can testify, and at this time can tolerate dissolved oxygen concentrations lower than 0.1 mg/l. pH range is 5.0-9.5 although they have been recorded as spawning at 4.2-4.4 but embryos are then malformed (Mann, 1996). Summer distribution is usually within 300 m of shore and less than 4 m deep. On windy days, pike retreat offshore in surface waters. Pike tolerate brackish water, up to about 7‰ for reproduction and 10‰ for feeding and growth. Reproduction requires living or dead vegetation (either aquatic or flooded terrestrial vegetation) in shallow, still waters protected from strong winds. Vegetation is also important for recruitment of young pike. Pike deposit their faeces at specific locations far from their usual feeding area as the faeces contain alarm pheromones recognised, and avoided by, prey species.

Drainage canal near Abkenar, Anzali Mordab, courtesy of S. Nouripanah
Drainage canal near Abkenar, Anzali Mordab, courtesy of S. Nouripanah

Age and growth

Life span is up to about 26 years but is less than half this in fast-growing southern populations. Some aquarium fish have lived 75 years. Maturity, like growth, varies with latitude and habitat, and also with quality of food. Higher temperatures may inhibit growth. Males mature at 1-6 years (30-46 cm) and females usually at 2-6 years, rarely at 1 year (31-63 cm). Females grow larger, faster and live longer than males. Growth is best at 19-21°C and is very efficient.

Nezami Balochy et al. (2005) examined this species in the Zibakenar-Kiashar Bojagh lagoon on the Caspian coast of Iran. The 122 fish were 17.7-74.0 cm long, average total length being 33 cm, and weight was 38-1100 g (average 307.3 g). The age groups were 0-9 years. Nezami et al. (2004; 2006) examined  this species in the Amirkelayeh Lagoon and found ages 1+ to 6+ years, average total length 44.8 cm (range 15.6-63.0 cm) and average weight 717.9 g (24-1700 g). Valipour (1998) investigated the pike in four areas of the Anzali Lagoon and found relatively fast growth with fish above 2 years of age mature with an average length of 32 cm.

In the Kizlyar Bay of the north Caspian Sea, males mostly mature at age 2+ with a body length of 36-40 cm while females are 3+ and 45-50 cm. Some males are mature in the first year of life at 26-30 cm and some females in the second year of life at 33-36 cm. Maximum age is 11 years in this population (Stolyarov and Abusheva, 1997). In Lake Aksehir, Turkey most males and females are mature at 2 years of age (Karabatak, 1988). Altindağ et al. (1999) give growth features for a population in a Kesikköprü Dam lake, Turkey where females reached age 5 and males age 4.

Food

Food is initially zooplankton and aquatic insects but fish begin to predominate at 5.0 cm after about 1 month's growth. Over 90% of the diet of adults is fish, but frogs, crayfish, mice, muskrats and ducklings are taken. Both sexes fast during spawning but females have rations 1.5-2.3 times as much as males in summer and winter. The daily ration was high from May to August with a June peak and very low in winter in North America.

Pike have a highly mobile eye which enables them to spot prey in almost any direction and have sighting grooves along the snout to facilitate their judgement of depth and distance. Food is seized after a rapid dart from concealment. Cylindrical fish like perch are preferred over more deep-bodied species as being easier to swallow. The prey capture process can be summarised as follows: eye movements towards prey, turning of body towards prey, stalking, darting, capture, rotating prey head first in mouth, and swallowing. Prey is sucked into the mouth which is opened just as the pike reaches its prey. Prey size is usually about one-third to one-half the length of the pike.

Valipour (1998) found feeding intensity and growth coefficient of this species in the Anzali Lagoon decreased with age. Fish in the 0+ age group fed mainly on zooplanktonic mysids while older fish took Carassius auratus, Hemiculter leucisculus, Rhodeus amarus and larvae of Alburnus chalcoides. Apparently, the pike is not restricting populations of commercial species in the lagoon but has an important role in controlling non-commercial exotics such as Carassius auratus and Hemiculter leucisculus. Abdoli (2000) lists Gambusia holbrooki, Carassius auratus, Hemiculter leucisculus, Liza saliens, Atherina boyeri (= caspia) and Alburnus charusini (= Alburnus hohenackeri) as food items for Iranian pike. The Zibakenar-Kiashar Bojagh lagoon fish had Odonata (14%) as the most frequent food, followed closely by Syngnathus caspius (13.8%) and Ponticola gorlap (13.4%). Other fish eaten included Gambusia holbrooki and pike. Nezami et al. (2004) and Nezami Balouchi

. (2006) examined  this species in the Amirkelayeh Lagoon and found the diet to be 24% Tinca tinca, 16% Proterorhinus marmoratus (= nasalis), with Esox lucius, plecopterans, frogs and water beetles at 8%, and Syngnathus caspius, Carassius auratus and Gammarus at 4%. Diet varied with season, and age and sex of the pike.

In the Kyzylagach or Imeni Kirova Bay of Azerbaijan Abramis brama, Cyprinus carpio, Rutilus rutilus (presumably R. caspicus), Rutilus frisii, Vimba vimba (= V. persa), Mugilidae, and Atherina boyeri (= caspia) are taken (Kuliev, 1989). Pike are cannibals when food is short. They compete with other piscivorous fishes, such as Sander lucioperca and Perca fluviatilis, for food. In the Kizlyar Bay of the north Caspian, the food of 2-month-old pike 3-4 cm long is mainly Rutilus rutilus  (presumably R. caspicus) fry, of pike 9-15 cm long this species plus fry of Cyprinus carpio and Scardinius erythrophthalmus, and of adults mainly Cyprinus carpio, Rutilus rutilus  (presumably R. caspicus) and Clupeonella cultriventris (= caspia) (during its spawning migration in April and May), Scardinius erythrophthalmus, Tinca tinca, Blicca bjoerkna, Alburnus alburnus (= hohenackeri), Cobitis sp., some gobies, frogs and crayfish. Sturgeon fry were once an important diet item but this declined with the decline in sturgeon numbers (Stolyarov and Abusheva, 1997).

Reproduction

The spring spawning migration in Dagestan begins at ice melt and spawning takes place from mid-March to the beginning of April when water temperatures reach 5-7°C. One batch of eggs is laid over a period of 10-15 days (Shikhshabekov, 1978). Spawning runs occur in the late evening and early night, several days before spawning occurs. Spawning takes place during the day, often in mid-afternoon, in shallow bays or flooded fields just after ice melt, generally in late March to May. Water temperatures on the spawning run are as low as 1.1°C. Spawning itself takes place at temperatures several degrees warmer than this, up to 17.2°C. Each female is accompanied by 1-2 males as she swims over vegetation in the shallow water. Both sexes roll to bring their genital regions close together, vibrate and release 5-60 eggs and the sperm. Tail sweeps scatter the eggs. This 3-10 second process is repeated many times each day. Eggs are amber, up to 3.4 mm in diameter after fertilisation, adhesive and each female can produce up to 595,000, although usually much less. In the Kizlyar Bay of the north Caspian Sea, the oldest females produce 365,000 eggs (Stolyarov and Abusheva, 1997). Eggs hatch 12-14 days later and the fry attach to vegetation by an adhesive head gland until the yolk sac is absorbed 6-10 days later.

The Kizlyar Bay population begins to spawn in late February or early March, sometimes under ice or immediately after ice melt at 4-6°C. Eggs are laid at depths of about 0.5 m and may dry out during water surges caused by northwestern winds although the increase in sea level has lessened this. Eggs adhere to vegetation but fall off after 2-3 days but do not die because of the low water temperatures and favourable oxygen conditions. Eggs hatch in 7-18 days depending on water temperature (Stolyarov and Abusheva, 1997).

Parasites and predators

Eslami et al. (1972) found helminths in 78.9% of 109 pike examined from Iran, a very high rate of infestation. The species encountered were Triaenophorus crassus, Raphidascaris acus and Contracaecum osculatum baicalensis. This latter parasite can infest man if fish is eaten smoked, salted or fried at temperatures below 50°C. Mokhayer (1976b) records the digenetic trematode Rhipidocotyle illense, the nematode larva Eustrongylides excisus, and the acanthocephalan Acanthocephalus lucii. Molnár and Jalali (1992) report the monogenean Tetraonchus monenteron from pike in Lake "Sama" in the Alborz Mountains. Ataee and Eslami (1999, www.mondialvet99.com, downloaded 31 May 2000) report Asymphylodora tinca from the gastro-intestinal tract of fish from the Anzali wetland. Naem et al. (2002) found the monogenean trematode Tetraonchus monenteron on the gills of this species from the western branch of the Safid River. Khara et al. (2004) looked at pike from the Amir Kelaieh Lagoon and recorded Raphidascaris acus, Camallanus lacustris, Eustrongylides excisus, Triaenophorus crassus, Trichodina sp., Tetraonchus monenteron, Diplostomum spatheceum, Lernaea sp., Argulus sp. and Piscicola sp. and atrributed the diversity to its piscivorous diet. Khara et al. (2006a) record the eye fluke Diplostomum spathaceum for this fish in the Amirkalayeh Wetland in Gilan. Sattari et al. (2002) and Sattari (2004) records the presence of the nematode, Eustrongylides excisus. This parasite can damage muscles in commercial species and render them unsuitable for sale. Sattari et al. (2005) surveyed this species in the Anzali, Amirkelayeh and Boojagh wetlands, recording Raphidascaris acus, Eustrongyloides excisus and Camallanus lacustris. Khara et al. (2006b) record the nematode Raphidascaris acus from this species in the Boojagh Wetland of the Caspian coast. Khara et al. (2007) examined fish from the Chamkhaleh River and recorded Raphidoscaris acus, Camallanus lacustris, Diplostomum spathaceum, Tetraonchus monenteron, Triaenophorus crassus, Corynosoma strumosum and Lernaea sp. Sattari et al. (2004; 2007) record the nematodes Camallanus lacustrisRaphidascaris acus and Eustrongylides excisus, the digeneans Rhipidocotyle illense and Diplostomum spathaceum and the monogenean Tetraonchus monenteron in this species in the Anzali wetland of the Caspian shore. Khara et al. (2004) recorded parasites from fish in the Amir Kelayeh Lagoon including Raphidascaris acus, Camallanus lacustris, Eustrongylides excisus, Triaenophorus crassus, Trichodina sp., Tetraonchus monenteron, Diplostomum spathaceum, Argulus sp., Piscicola sp., and Lernaea sp., a diversity governed by its piscivorous diet.  Barzegar et al. (2008) record the digenean eye parasite Diplostomum spathaceum from this fish. Khara et al. (2008) found the eye parasite Diplostomum spathaceum in this fish from Boojagh Kiashar Wetland in Gilan. Miar et al. (2008) examined fish in Valasht Lake and the Chalus River, Mazandaran and found the metazoans Rhaphidoscaris acus and Tetraonchus menonteron. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Lernaea sp. on this species. Jamalzadfalah et al. (2011) found 10 parasite species in fish from the Anzali Wetland namely Argulus foliaceus and Lernaea cyprinacea (Crustacea), Eustrongylides excisus and Raphidascaris acus (Nematoda), Tetraonchus monenteron and Dactylogyrus sp. (Monogenea), Diplostomum spathaceum (Digenea), Trichodina sp. (Ciliata), Corynosoma strumosum (Acanthocephala), and Rhipdocotyle illense (Platyhelminthes). Khara et al. (2011) list the mongenean Tetraonchus moneteron, the digenean Diplostomum spathaceum and the nematode Raphidascaris acus from this fish in the Boojagh Wetland of the Caspian Sea.

Young pike are eaten by various other fishes including adult pike, birds and even large aquatic insects.

Economic importance

Nevraev (1929) reports a catch for the 1901-1902 to 1913-1914 period in the Anzali region was 1150 to 20,529 fish. Holčík and Oláh (1992) report a catch of 5836 kg in the Anzali Mordab in 1990, at 7.8% of the catch the fourth most important fish there, while annual reported catches from 1932-1964 varied from none to 98 tonnes. However it is not a favoured food fish in Iran (Vladykov, 1964).

There is some opportunity for sport fishing for this species in the Anzali Mordab and potentially in various lakes around the country where it has been introduced (Anonymous, 1977). Anglers catch this species in rivers along the Caspian shore such as in the Shazdeh River at Babol Sar (Noorbakhsh, 1993b). It is an important sport and commercial fish in other parts of its range. The catch in Turkey in 1981 was 796 tonnes and for the inland waters of the former U.S.S.R. in 1975 it was 16,101 tonnes. Stolyarov and Abusheva (1997) report a commercial stock of 2500 t with a recommended catch of 800-850 t in Kizlyar Bay in the north Caspian in the early 1990s.

The eggs or roe of this species are very poisonous as fresh extracts injected intravenously into rabbits have caused respiratory distress, convulsion and death within one hour (Halstead, 1967-1970). Curiously, Anzali lagoon "perk" eggs have been studied as a replacement for caviar (Iranian Fisheries Research Organization Newsletter, 58 & 59:1, 2009).

Conservation

Raat (1988) gives details of conditions which should exist to facilitate pike reproduction and growth, including such factors as vegetation, water levels, eutrophication, pollution, prey availability, intra- and inter-specific interactions, fishery, stocking, concentration of dissolved solids, pH levels, and temperature regimes. Vladykov (1964) noted a fish kill in the Anzali Mordab on 11 June 1962 where water chestnut (Trappa natans) had caused an oxygen deficiency in the shallow water.

Gilan Fisheries Research Station has cultured pike in earthern ponds. Pike caught in autumn and winter were injected with gonadotropic hormone from carp (4-7 mg/kg). Eggs were stripped after 48-72 hours and incubated for 10 days at 8-11°C and 7-8 days at 10-15°C. Absorption of the larval yolk sac took approximately twice as long as the incubation period.

Ramin (1999) reports on a project involving artificial spawning and raising of fry in earthen ponds. Spawning temperatures were 8-15°C from 4 February to 20 March. Males and females were a minimum of 3 years old and a maximum of 5 and 6 years respectively and weighed 0.75-4.0 kg. Eggs composed 10-20% of body weight, incubation lasted 120 degree days and yolk sac absorption 160-180 degree days. The rate of fertilization was 45-85% and swelled eggs were 2.5-3.5 mm. Absolute fecundity was 22,400-112,000. Survival from larvae to fingerlings in chicken manure enriched ponds was 20-22% over 50 days with growth to 7.4 g and 8.5 cm on average.

Pike from the Anzali Lagoon have been cultured with Chinese carps and were found to be mature in less than a year (Iranian Fisheries Research Organization Newsletter, 56:2, 2008).

Kiabi et al. (1999) consider this species to be conservation dependent in the south Caspian Sea basin according to IUCN criteria. Criteria include commercial fishing, sport fishing, medium numbers, habitat destruction, medium range (25-75% of water bodies), present in other water bodies in Iran, and present outside the Caspian Sea basin. Mostafavi (2007) lists it as conservation dependent in the Talar River, Mazandaran. Endangered in Turkey (Fricke et al., 2007).

Further work

The distribution and population numbers of of this species needs documentation in Iran.

Sources

Crossman and Casselman (1987) give a bibliography and Raat (1988) and Craig (1996) give synopses of biological data on this extensively studied species. General biology and characters are based on world-wide data. Sohrabi (1996b) gives an account of this species in Farsi.

Iranian material: CMNFI 1970-0510, 1, 250.3 mm standard length, Gilan, Golshan River (37°26'N, 49°40'E); CMNFI 1970-0535A, 1, 329.6 mm standard length, Gilan, Pir Bazar Roga (37°21'N, 49°33'E); CMNFI 1970-0542, 1, 39.0, mm standard length, Gilan, Old Safid River estuary (37°23'N, 50°11'E); CMNFI 1970-0543A, 1. 136.2 mm standard length, Gilan, Caspian Sea at Hasan Kiadeh (37°24'N, 49°58'E); CMNFI 1970-0553, 1, 136.7 mm standard length, Gilan, Sosar Roga (37°27'N, 49°30'E); CMNFI 1970-0579, 9, 71.9-111.2 mm standard length, Gilan, Old Safid River estuary (37°23'N, 50°11'E); CMNFI 1971-0343, 3, 63.8-74.7 mm standard length, Gilan, Langarud at Chamkhaleh (37°13'N, 50°16'E); CMNFI 1979-0685, 1, 149.2 mm standard length, Gilan, Safid River around Mohsenabad below Dehcha (no other locality data); CMNFI 1980-0123, 2, 126.3-209.3 mm standard length, Gilan, Safid River around Dehcha above Mohsenabad (no other locality data); CMNFI 1980-0138, 2, 148.2-164.3 mm standard length, Gilan, Safid River estuary (ca. 37°28'N, ca. 49°54'E); CMNFI 1993-0147, 1, 106.0 mm standard length, Iran (no other locality data).

Lotidae

The Lotidae or cuskfishes may be included within the cod family Gadidae. The family is found in temperate to cold marine waters in the Northern Hemisphere, with only one species always resident in fresh water. There are about 6 genera and 21 species in the family (Eschmeyer and Fong, 2011). Maximum size exceeds 1.5 m.

Cuskfishes ods are recognised by the single barbel usually present under the chin; 1-2 dorsal fins and 1 anal fin; the caudal fin usually extends around the dorsal and ventral tip of the caudal peduncle and is rounded; no fin spines; wide gill openings with the branchiostegal membranes free or narrowly attached to the isthmus; 6-8 branchiostegal rays; vomer bone in the roof of the mouth toothed; swimbladder not connected to the auditory capsules and with 2 slender, anterior processes; scales small and cycloid; an obvious lateral line; and the egg with an oil globule. Eggs and larvae are usually pelagic.

Genus Lota
Oken, 1817

This genus has a single species found in North America and Eurasia, rarely in Iran. The characters of the species are therefore the same as for the genus.

Lota lota
(Linnaeus, 1758)

Common names

mahi charb (= fat or greasy fish), lot.

[nalim in Russian; burbot, eelpout].

Systematics

Gadus Lota was originally described from European lakes. Iranian specimens are presumed to be the type subspecies with a long and low caudal peduncle and high meristic counts (Pivnička, 1970). Subspecies defined on numbers of pyloric caeca are not accepted since there is clinal and size variation (Kottelat, 1997).

Key characters

The chin barbel and 2 dorsal fins, both lacking spines, are distinctive.

Morphology

First dorsal fin rays 7-16, second dorsal fin rays 60-94, anal fin rays 58-86, pectoral fin rays 15-24, pelvic fin rays 5-10, gill rakers 4-12, short, rounded and spinulose, discrete or almost reaching the base of the adjacent raker when appressed, pyloric caeca 21-67, and vertebrae 50-72. The head is flattened, the anterior body rounded and the posterior body compressed. Scales on the body are minute, cycloid, embedded, with few circuli and no radii, and extend onto fin bases and gill covers, and to the level of the nostrils on the head. The second pelvic fin ray is elongated. The anterior nostril is a barbel-like tube. The gut has numerous pyloric caeca, and anterior and posterior loops. The chromosome number is 2n=48 (Klinkhardt et al., 1995).

Sexual dimorphism

Unknown.

Colour

Overall colour is a yellowish to olive-green to dark brown and may be black, with large light blotches on the head, body and vertical fins. The belly is yellow to white and may be finely spotted. All fins are strongly mottled. Mottling is most evident in younger fish; adults tend to become uniformly dark.

Size

Reaches 1.52 m and 32.0 kg, possibly 2.1 m and 36.0 kg (Machacek (1983-2012), downloaded 27 July 2012).

Distribution

Found across northern Eurasia and northern North America. Recorded from the lower Safid River in 1921 (Derzhavin, 1934; Berg, 1948-1949) and possibly waters in Gilan but apparently rare in Iran. Berra (1981) omits the distribution in Iran and the southern Caspian Sea basin of Azerbaijan.

Zoogeography

Populations in the Kura River of Azerbaijan may be disjunct from northern populations but their characters have not been thoroughly examined. Abbasov (1980) did not report this species from the Aras River on the northern border of Iran so the Iranian fish, if not strays, may be disjunct even from the southern populations in the Kura River.

Habitat

This species favours cool, clear rivers and lakes where it is most active at night. It hides under rocks and plant roots or in holes in river banks during the day. Young fish remain in shallow weeded areas or rocky streams. It is found only in the lower reaches of rivers along the Iranian shore and does not penetrate upstream (Berg, 1948-1949). This species can tolerate 6‰ and so may enter the Caspian Sea near river mouths. Embryos of this species suffer a 50% mortality if pH rises to 8.0 (Mann, 1996). Fat reserves in the liver are used up in summer when this species is less active.

Age and growth

Maturity is attained at age 2-7 years, varying with habitat, rarely at 1 year for males. Growth is faster in southern populations and maturity is earlier than for northern populations. Maximum life span is 22 years.

Food

Food when young is mostly aquatic insects, crustaceans and molluscs while older burbot feed voraciously on fishes and their eggs and on frogs. Adults do not feed during the spawning period. Most feeding occurs at night. It is reported to be cannibalistic. Glycogene and fat are stored in the liver during spring and autumn feeding periods, allowing the fish to cope with high water temperatures in summer when the fish is less active and feeding is low, even enabling some growth and gonad development (Kottelat and Freyhof, 2007).

Reproduction

Spawning takes place from November to March, even under ice, at temperatures of 0.5-4.0°C on gravel, sand or hard bottoms. Successive matings occur with the male and female swimming head down along the bottom until the male rotates and pushes his belly against hers, releasing eggs and sperm in a cloud. The female beats her tail, mixing the gametes and dispersing the fertilised eggs. One or two females may be surrounded by many males in a spawning ball. The eggs are easily moved by water currents because of a large oil globule in the yolk but gradually sink and lodge in interstices in gravel and sand. Up to 5 million eggs are produced with a diameter about 1.9 mm. About 80 "day degrees" are needed for the eggs to hatch and since they are laid in winter this takes some weeks, e.g. about 6 weeks at 2°C.

Parasites and predators

None reported from Iran.

Economic importance

This species is of economic importance in the former U.S.S.R. and the flesh is said to be excellent as is the liver. The eggs have been used as caviar but are also reported as toxic (Halstead, 1967-1970; Coad, 1979b; see under the genus Schizothorax for symptoms of ichthyootoxism). However it is too rare in Iran to be a food fish and a potential health hazard.

Conservation

Classified as rare to intermediate by Lelek (1987) for Europe. Abdurakhmanov (1962) gives data on only one specimen from the southern Caspian Sea basin, indicative of its rarity. It may be extirpated from Iranian waters.

Further work

Anglers and commercial fishermen should report any captures of this rare species in Iranian waters. Specimens should be preserved for comparison with northern populations.

Sources

Counts are taken from Pivnička (1970).

Mugilidae

The mullets or grey mullets are found world-wide in temperate to tropical coastal waters readily entering estuaries and even resident in freshwaters. There are about 20 genera and 72 species (Eschmeyer and Fong, 2011) but only 3 species are native to Iran in fresh and brackish waters and a further 2 species have been successfully introduced (the latter not mapped in Berra (2001) because they are exotics). Other species are recorded as entering the rivers of southern Iran from the Persian Gulf and Sea of Oman although identification is not always certain (see Marine List in Checklists in the Introduction). Maximum size is about 0.9 m.

This family is characterised by a compressed to subcylindrical body with a somewhat flattened head; moderate sized scales which may be cycloid but are ctenoid in most adults and extend onto the top and sides of the head; faint or no lateral line along the flank but pits or grooves on scales contain the sense organs; the eye may have an adipose or fatty eyelid forming a vertical, slit-like opening; vertebrae usually 24, rarely as high as 26; wide gill openings; gill rakers long and slender and increasing in number with growth; upper elements of the gill arch are specialised as a pharyngobranchial organ,;5-6 branchiostegal rays; a spiny and short first dorsal fin (4 spines), the second dorsal with 1 unbranched ray or spine and 6-10, usually 8, branched rays; anal fin with 2-3 spines and 8-12 branched rays; an abdominal pelvic fin with 1 spine and 5 soft rays; pelvic bones connected to the postcleithrum by a ligament; mouth transverse and small; teeth on jaws short, weak and flexible, and the lower jaw may be toothless; the stomach wall is strongly muscled (gizzard-like); and the gut is very long and coiled.

Mullets are schooling fish which feed on microscopic algae and the minute animals associated with the algae. They grub, gulp or suck (hence "mugil") bottom deposits, spitting out some of the debris and extracting nutrient from the remainder. The long gill rakers filter the food, the strong gizzard-like stomach crushes it and the long intestine (about 7 times body length) aids in digestion. Their bottom feeding leaves long patches of disturbed sediment readily visible from a distance. Eggs, larvae and young mullet are pelagic. Adults are found in large schools in coastal waters or on tidal flats. These fishes are very important economically as food eaten fresh, smoked or canned, as bait, and as cultured fish in ponds. The flesh is oily and rich but has few bones. Mugilid species in Khuzestan are thought to be the intermediate hosts of Heterophyidae flukes found in humans and carnivores (Massoud et al., 1981).

Recent studies have been carried out on culturing Mugil cephalus and Liza aurata in brackish water, in inland pools and salt and freshwater culture ponds in the northern and central parts of Iran (Emadi, 1993a; Iranian Fisheries Research and Training Organization Newsletter, 11:6, 1996; Azari Takami et al., 1997a; Yelghi et al., 2012). In April 1994, 20,000 Mugil cephalus fry weighing 0.5 g were imported from Hong Kong as part of this programme (Iranian Fisheries Research and Training Organization Newsletter, 4:8, 1994). Azari Takami et al. (1997a) successfully raised fry of Liza aurata and L. saliens at Bafgh-Yazd in central Iran in earthen ponds supplied with brackish well water. Fry in the 3-10 g range were cultivated for 100 days at 2200 fry/ha with pond fertilisation and with fertilisation and complementary feeding, giving 100 g and 120 g fish respectively. Experiments with different stocking densities and mixes of the two species were also carried out. Second year L. saliens reached 165 g and third year L. aurata reached 630 g and sexual maturity.

Experiments have been carried out on fishing methods for these fishes in Iran, comparing encircling gill nets with fixed gill nets and with purse seines (Annual Report 1992-93, Iranian Fisheries Research and Training Organization, Tehran; Annual Report 1993-94, Iranian Fisheries Research and Training Organization, Tehran). In 1994-1995, the total mullet catch was 4145 t, 85% of which was L. aurata. Catch per unit effort decreased by 50% over the preceding three years (Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 37, 1996).

Soltani and Rahanandeh (2001) describe a case of gastric bloat in Mugil capito in the Caspian Sea. Mugil capito Cuvier, 1829 is a synonym of Liza ramada (Risso, 1827), a species not known from the Caspian Sea.

The general name for mullets in Farsi is kefal or kafal.

Pillay (1972) gave a bibliography on mullets and Thomson (1997) reviewed the family, giving further details of anatomy than summarised here.

Genus Liza
Jordan and Swain, 1884

This genus is characterised by thin to moderately thick, terminal upper lip without papillae, the lower lip is directed forwards and is thin-edged, teeth are setiform, ciliiform or absent in the upper lip, ciliiform or absent in the lower lip, there is a symphysial knob to the lower jaw and the lower jaws meet at a 90º angle or more, the maxilla is bent down over the premaxilla and is either uniformly curved or is s-shaped, the maxilla end is visible when the mouth is closed, the anteroventral edge of the preorbital bone is serrate, weakly concave or kinked and ventrally it is broad and squarish, an adipose eyelid is present sometimes but is not well-developed being a narrow rim around the eye at all ages, the pharyngobranchial organ has two valves, pyloric caeca number 2-14, predorsal scales are unicaniculate, and the pectoral axillary scale is weak or absent. Thomson (1997) lists further characters but the genus is distinguished easily from Mugil (q.v.), the only other genus from Iran.

Liza abu
(Heckel, 1843)

 

Further illustrations

Liza abu, lateral view of head of above; Liza abu, dorsal view of head of above; both after Berg (1949)

Common names

بياح (biah), biah zury, zuri, kafal, shochy, do'kelki in Khuzestan (= with two sails, presumably referring to the two dorsal fins), ? derbak.

[maid, khishni, hishni, abu-khraiza, hosoon, hashsoun, abu-khraiza, or abu sukkanejn (= father of two anchors in allusion to the toothed suborbital bone according to Heckel (1843b)), all in Arabic; minghaj in Pakistan; abu mullet, freshwater mullet].

Systematics

Mugil pseudotelestes Pietschmann, 1912, described from the "Schatt el Arab bei Basra (Aschar)" (Ashar is at 30°31'N, 47°50'E), is probably a synonym (Coad, 1991b) based on the original description. Mugil hishni Misra in Hora and Misra, 1943 described from "Rivers and Hors, Iraq" (hors or hawrs are marshes) is also a synonym based on the description, an opinion concurred in with Ingham (no date) and Özdilek (2003). Thomson (1997) considers hishni to be a synonym and possibly pseudotelestes although he considers the description of this species insufficiently detailed. Mugil abu zarudnyi Berg, 1949 was described from "Ser-i-pul, 30 km from Malamir, between Deh-i-dez and Malamir, region of the Bakhtiars and Lurs, upper course of the Karun R., south-western Ian, 17 III 1904, N. Zarudnyi". Thomson (1997) considers this subspecies to be of doubtful validity given the variability of this species. Different stocks exist in the Tigris, Euphrates and Orontes rivers as evidenced by morphology (Turan et al., 2004).

Randall (1995b) places this species in the genus Chelon Rose, 1793; Thomson (1997) does not.

Material of Mugil abu, housed in the Naturhistorisches Museum Wien, is under NMW 9224-9230 (7 fish) and 67868 (2) and are syntypes. The type locality is the "Tigris bei Mossul" according to Heckel (1843b) and the catalogue lists 4 specimens.

Mugil (Liza) abu zarudnyi Berg, 1949 was described from Iranian tributaries of the Tigris River basin. The 5 syntypes of zarudnyi are in the Zoological Institute, St. Petersburg (ZISP 24336), measure 69.1-80.7 mm standard length. The date in the catalogue is 4.II.1904 (old style).

The holotype of Mugil hishni is in the Zoological Survey of India, Calcutta under ZSI F13626/1 with 1 paratype under ZSI F13627/1 (Menon and Yazdani, 1963; Eschmeyer et al., 1996).

Key characters

The high lateral scale count, long pectoral fins reaching almost level with the first dorsal fin origin when folded back (note fin tips often frayed, especially in preserved material, so not as apparent), short pectoral axillary scale, thin lips, 3 anal fin spines and 8 branched rays, relatively strong spines in the first dorsal and anal fins, and peg-like or setiform teeth (not tricuspid) in the upper jaw only, distinguish this species from other species in the genus Liza and from other mullets.

Morphology

The eye has fatty tissue covering as far as part of the iris. Lips are thin and the lower lip has a pronounced knob. Peg-like teeth are obviously present only on the upper lip, the lower lip having scattered ciliate teeth. The end of the maxilla has an s-shaped bend and its tip is exposed. The anterior edge of the denticulate preorbital is angular, bending down at the corner of the mouth. The pectoral axillary scale is weakly developed. Occasional antero-dorsal scales have double canals. Pectoral fin length is 75-78% of head length and this fin extends back to a level almost at the first dorsal fin origin. First dorsal fin spines 3-4, usually 4, second dorsal fin soft rays 5-10, usually 8 after 1-2 spines, anal fin spines 3 followed by 5-10, usually 8 soft rays, pectoral fin branched rays 13-17 and pelvic fin branched rays 5-6, usually 5. Vertebrae 21-25. Fusions and minor vertebral anomalies were described by Al-Hassan (1987) and Al-Hassan and Naama (1986b) from Iraqi fish. Lateral scales 39-53, usually 44-50. Scales are strongly ctenoid on the exposed part and the fish feels rough to touch when rubbed from tail to head. The embedded part of the scale has fine circuli. The focus is very posterior. Scale shape is rectangular with the anterior margin vertical or somewhat wavy. The anterior upper and lower corners are square or even pointed. The dorsal and ventral scale margins are straight and the posterior margin is rounded. There are a few radii (as few as 4) from the focus to the anterior margin and, where these intersect the margin, it may be slightly indented to form a wavy edge. There are 4 pyloric caeca and the gut is elongate and coiled. Gill rakers are short than the gill filaments and number 44-55. They bear teeth on the internal edge. Islam and Al-Nasiri (1978) give morphometric characters. Hoda (1978) describes the larva (as Mugil hishni) in Iraq. 2n = 50 (Balasem et al. (2000). Thomson (1997) notes that this is the most variable species in the family.

Al-Hassan (1984b) showed differences in counts of vertebrae and dorsal fin rays between fish from Basrah and the Karkheh River in Iran. Turan et al. (2004) also noted differences in morphometric data for stocks from the Orontes (= Asi), Tigris and Euphrates rivers of Turkey, attributing this to phenotypic adaptation as the stocks were genetically homogenous. Thomson (1997) notes that this is the most variable species in the family. Jawad (2004b) studied asymmetry in this species from the Shatt al Arab at Basrah and suggested pollution could be a cause. Jawad and Öktener (2007) record lordosis (axial spinal curvature) in this species in the Atatürk Dam Lake, Turkey. Jawad et al. (2010) describe a specimen from Karkheh River branch near Susangerd with a malformed caudal fin.

The nominal subspecies zarudnyi has 3-4 first dorsal fin spines, 1-2 spines and 5-7 soft rays in the second dorsal fin, the anal fin has 3 spines and 5-9 soft rays, usually 9, and scales in lateral series 41-46. The larger scales distinguish it from the typical form at Mosul in Iraq (Berg, 1949). My observations on the types show 4(4) first dorsal fin spines, 1(2) or 2(2) second dorsal fin spines and 7(3) or 8(1) soft rays, 7(1), 8(1) or 9(2) anal fin soft rays, 5(4) pelvic fin rays, 13(1) or 14(3) branched pectoral fin rays, and 39(1), 43(1), 44(1) or 45(1) scales in lateral series. One specimen is unusual in having a mix of abnormally low counts and normal counts: 3 first dorsal fin spines, 5 second dorsal fin soft rays, 5 anal fin soft rays, 5 pelvic fin rays, 15 pectoral fin rays, and 43 lateral series scales.

Meristic values for Iranian specimens, excluding the above: first dorsal fin spines 4(36); second dorsal fin soft rays 7(4), 8(31) or 9(1); anal fin soft rays 8(28) or 9(8); pectoral fin branched rays 13(1), 14(17), 15(13) or 16(4); pelvic fin branched rays 5(35) or 6(1); lateral series scales 44(5), 45(5), 46(5), 47(8), 48(2), 49(4), 50(1) or 51(1); and total vertebrae ?.

Sexual dimorphism

Unknown.

Colour

The back is a light brown or greyish or olive-green with the flanks and belly silvery or yellowish-silver. There may be an indistinct silvery stripe or 2 grey stripes along the flank. Generally the upper flank is dark, the lower flank pale, and the two clearly demarcated. The upper part of the lower flank may bear alternating light and dark stripes but these are sometimes poorly expressed. The second dorsal and caudal fins are dusky to brownish and lower fins are yellowish-white to hyaline. Scale margins are covered in small black spots. The top of the eye is lime-green and reddish-brown just below but still silvery around the pupil generally. Young fish have a reddish colour at the base of the pelvic, anal and caudal fins and have an evident mid-lateral stripe against a pale body.

Size

Attains 26 cm total length and 0.15 kg.

Distribution

Found in the Tigris-Euphrates and Orontes (= Asi) (Özdilek, 2003) river basins including the Tigris River basin in Iran in rivers and marshes such as Dez, Karkeh and Karun rivers and Shadegan Marsh, in lower reaches of rivers draining to the Persian Gulf, and in Pakistan. In Iran it is also recorded from Parishan (= Famur) Lake but it was not captured in a 1995 survey. A record from the Barm-e Shur near Shiraz is probably an introduction as this species was not caught there in the 1970s by me (Gh. Izadi, pers. comm., 2001).

Abdoli (2000) maps this species in the lower Hasan Langi, Kul, Gowdar and Mehran rivers at the Straits of Hormuz in the Hormuz basin; the lower Mand River and the lower Zohreh (= Hendijan) rivers in the Gulf basin, the Arvand, lower Karun, lower Karkeheh and the Simarreh rivers in the Tigris River basin. It is also reported from the upper Karun River as M. a. zarudnyi.

Zoogeography

As a freshwater member of a genus with many species in the sea, its origins lie with these marine relatives.

Habitat

This mullet is a freshwater species found in rivers, streams, channels, canals and drains, lakes, reservoirs and ponds, on fish farms of Iraq and neighbouring countries, occasionally entering estuaries. It usually occurs in schools. Epler et al. (2001) found it to be the dominant species of fish in lakes Habbaniyah, Tharthar and Razzazah, Iraq, comprising 72% of all fish collected. This was one of the most abundant species in the recovering marshes of southern Iraq in 2005-2006 (at almost 36% of 16,199 fishes collected)(Hussain et al., 2006) and in the marshes in the 1980's (Hussain and Ali, 2006). Nasir and Na'ama (1988) and Hussain and Naama (1989) report it from the Khawr az Zubayr in a marine environment, probably a consequence of human-induced environmental changes. It is the most abundant species in autumn in the Karun River, Iran, comprising 44.2% of the catch (Mokhayer, 1981c).

van den Eelaart (1954) reporting on Iraqi populations found this mullet in the surface waters and submerged vegetation of lakes and marshes, preferably where there is a gentle water flow. In December-January it enters rivers and deeper waters, especially in very cold winters. Ahmad et al. (1983; 1985) have shown this species can withstand abrupt increases of temperature up to 30°C and salinity up to 10‰ for 24 hours under experimental conditions. Mortality in water at 15‰ is low. Salinities up to 30‰ and temperatures up to 35°C are tolerated, presumably if increases are gradual. Ahmed et al. (2002) stated that it prefers salinities not exceeding 2‰ (later 5.6‰) but can survive abrupt transfer to 15‰. S. Cowton (pers. comm., 23 August 2005) has observed schools of this species gaping at the surface in the artificial lake around Al Faw Palace in Baghdad, presumably in response to high temperatures and low oxygen. It is often the dominant species left in small pools when marshes dry up, as in the Shadegan Marshes of Iran observed by B. W. Coad.

Age and growth

Hashemi et al. (2010a, 2010b) give a length at maturity (LM) of 6.7 cm, production per biomass (P/B) of 0.72, L of 289 mm, K of 0.36 yr-1 and t0 of -0.35 yr-1 for fish from the Shadegan Wetland in Khuzestan.

Al-Nasiri and Islam (1978) studied the age and growth of this species from commercial catches in the Shatt al Arab, Iraq. Specimens were aged by examining the otoliths and three age groups determined with 0+ fish being 8.9-10.9 cm, 1+ fish being 9.9-14.8 cm and 2+ fish being 13.6-18.2 cm total length. Length-otolith formulae were given. Al-Yamour et al. (1988) examined a population of this species in the Al-Daoodi Drain, Baghdad and found an age span up to 7 years using scales to age fish. Older age groups (5 and 6) grow more slowly than Diyala River fish, perhaps because of higher salinity in the Drain and the enriched conditions in the Diyala River due to sewage. The length-weight relationship was W = 0.034 L2.6, for males W = 0.062 L2.396,and for females W = 0.031 L2.642. The condition factor was 6.308 for males and 3.155 for females. Khalaf et al. (1986) studied a population in the Diyala River, Iraq which was polluted with sewage from the Rustamiyah treatment plant. Khishni is the second dominant species from December to March after Chondrostoma regium and in August the dominant species in catches. Fish were heavier and in better condition in the more polluted areas, but probably not as good for human consumption! Growth declines after age group 5 with few fish in age group 6. Length-weight relationships were W = -3.39 L2.64 for males and W = -2.96 L2.50 for females. Mhaisen and Yousif (1989) examined a population of khishni in Mehaijeran Creek, a side branch of the Shatt al Arab south of Basrah, Iraq. Only 3 age groups (0+ to 2+) are reported since the creek water level is affected by the tide and older fish tend to move into the deeper waters of the adjacent Shatt al Arab. Growth is faster in the creek than in other Iraqi locations. The length-weight relationship was log W = 2.794 log TL - 1.64840 for males and log W = 3.337 log TL - 2.25728 for females. Mhaisen and Al-Jaffery (1989) report 6 age groups for fish from Babylon Fish Farm west of Hilla, Iraq with a total length-weight relationship log W = 2.96057 log L - 11.19599 (and for standard length log W = 2.87574 log SL -10.24769). Fish were in good condition as they were taking food meant for cultured carp. The relative condition factor was 0.98-1.04, similar to values obtained by other authors. The von Bertalanffy growth equation was TLt = 409.6 {1 - exp [-0.06314 (t - 3.13868)]}. Al-Shamma'a et al. (1995) examined fish from the Tigris River at Al-Za'faraniyah, Iraq and found log total weight = -2.033 + 3.119 log total length (r= 0.955). Epler et al. (2001) found the oldest age groups to be 4+ in Iraqi lakes Habbaniyah and Razzazah and 5+ in Lake Tharthar. The mean condition factor was 0.97, 1.05 and 1.09 in lakes Habbaniyah, Tharthar and Razzazah respectively. Syzpuła et al. (2001), studying age and growth in the lakes Habbaniyah, Tharthar and Razzazah in 1981 and 1982, found this species grew fastest in Lake Tharthar. A decrease in length growth was noted in the second year of life and a decrease in body mass in the last years of life.

The effect of starvation over 35 days on the proximate chemical composition of this species was studied by Yesser et al. (1999). They found a sharp decline in lipid content (4.38 to 0.98%), a slight drop in protein and an increase in both moisture and ash. Both condition factor and viscera somatic index declined gradually, 1.2 to 0.83 and 10.18 to 3.63% respectively. Body weight fell by 30.59%. Lipid reserves in muscle were therefore an important form of energy storage.

Ünlü et al. (2000) report age groups of 1+ to 4+ in the Turkish Tigris River. Hussain et al. (1987) found condition factor to range from 1.12 to 1.64 for different length groups for Karkheh River fish from Iran near the Iraq border and Hawr al Hawizah.

Food

Al-Nasiri et al. (1977) studied the feeding ecology of this species in the Shatt al Arab, Iraq. Aquatic plant parts and phytoplankton are food items but the vast bulk of material is sand grains and organic detritus, the former presumably ingested while searching for detrital food. Phytoplankton is made up of diatoms (50%), green algae (36%) and blue-green algae (14%), with diatoms the most abundant numerically. Plant parts are fragments of Vallisneria leaves and rarely leaves and stems of Potamogeton and Polygonum. Islam et al. (1981) reported on the seasonal patterns of feeding in Rashdiyah Reservoir, Baghdad. However it should be noted that only 10 fish were examined for each month. Aquatic plant parts, organic debris and phytoplankton are dominant foods followed by zooplankton and aquatic insects. The diet is diverse, 48 species of Chlorophyceae, 60 species of Bacillariophyceae, 9 species of Myxophyceae and 3 of Euglenophyceae. Heavy feeding occurs in late winter, spring and fall with peaks in April and November. Empty stomachs are more common in summer. Islam and Khalaf (1983) studied diel feeding patterns in the same reservoir and found peaks at 0600 and 2100 hours. Naama and Muhsen (1986) studied the feeding periodicity of this species in the Hawr al Hammar, Iraq and found the main food to be algae, mixed with incidental sand grains. There is a single feeding peak each day at 1730 hours and feeding stops at 0200 hours. This species is a day feeder. In another study of the recovering Hawr al Hammar, diet was 76.49% algae and 20.3% diatoms with amounts of crustaceans and plants being less than 10% each, in the Hawr al Hawizah 74.3% algae and 22.5% diatoms, with amounts of plants and crustaceans being less than 10% each, and in the Al Kaba'ish (= Chabaish) Marsh 75.8% algae and 20.0% diatoms with plants and crustaceans at less than 10% each (Hussain et al., 2006). Ahmed and Hussain (1982) examined the food of young fish near Basrah, Iraq and found organic detritus to be an important food followed by phytoplankton. The smallest fish are consuming eggs in large quantities. Al-Shamma'a and Jasim (1993) studied feeding in the Hawr al Hammar in Iraq during the flood period and reports small quantities of adult insects, chironomid larvae, trichopteran larvae, other insect larvae, molluscs and worms, while copepods, cladocerans, and rotifers are important foods. This variation from the diet studies reported above may be due to the flood not allowing diatoms and algae to settle out and become available as food. Typical foods and gut contents such as organic detritus, algae, diatoms, plant tissues and sand grains are also present. Sand grains may help in trituration of food and therefore are not an accidental inclusion. Al-Shamma'a et al. (1995) examined fish from the Tigris River at Za'faraniyah and found phytoplankton ranked first in the diet followed by detritus. Debris comprised more than 37% of stomach contents on the points method. Epler et al. (1996) found detritus to be the main food in three Iraqi lakes. Epler et al. (2001) found detritus to be 83.7% of the diet in Lake Tharthar, mineral parts (presumably sand) 12.7% and plants 3.6%. In Lake Habbaniyah detritus was 73.0%, mineral parts 16.5% and plants 8.4%. In Lake Razzazah detritus was 84.4%, mineral parts 14.7% and plants 0.9%. Detritus was the principal food year round with no seasonal variations. Hussain and Ali (2006) examined feeding relationships among fishes in the Hawr al Hammar and found this species to be a herbivore, 39.6% of the diet being plants and algae, with 18% detritus.

Hussain et al. (1987) examined the food of khishni in the Karkheh River, Khuzestan, where it is the dominant species in terms of numbers caught, by two orders of magnitude. However capture techniques would affect this assessment. Food was similar to that reported from Iraqi fish. Phytoplankton is the most important food for fish 3.0-3.9 cm long, followed by organic detritus, while in larger fish this order is reversed. Silt is an important content of the gut in all fishes studied. Eighteen species of Bacillariophyceae, 6 Chlorophyceae and 4 Cyanophyceae species were recorded from gut analyses. Abdoli (2000) reports Daphnia, Hemiptera, Navicula, Nitzschia, Amphora and Cymbella species for Iranian specimens.

Reproduction

Marammazi (1994) considered it likely that this species spawns twice each year in the Zohreh River, Iran draining to the northern Persian Gulf. Chelemal et al. (2009) studied reproduction in fish from Dasht-e Azadegan west of Ahvaz. Females predominated over males (2.7:1) and maximum gonadosomatic indices were found in March for both sexes with a minimum in August, suggesting spawning in April with a spawning season extending from February to June. Details were given on histological development of the ovaries with spawning females having eggs 470.3μ in diameter.

van den Eelaart (1954) gave the spawning season in Iraq as end of February to mid-March. In the Al-Daoodi Drain at Baghdad and the Hawr al Hammar of southern Iraq spawning starts in March and continued through May (Al-Yamour et al., 1988). Mean number of eggs reaches 59,793, increasing with size and weight. In the study by Khalaf et al. (1986), all age group 1 fish are mature and females outnumber males in age groups 2 to 6 although this may be due to sampling bias. Ünlü et al. (2000) report maturity at age 1 in the Turkish Tigris River based on gonad development, with no spawning in the period August to February. Mhaisen and Yousif (1988) examined a population of khishni in Mehaijeran Creek, a side branch of the Shatt al Arab south of Basrah. Fish are ripe from January to March and females partly spent in April and spent in May. The fish spawn once. All males and females were mature at 16 cm in the creek while females mature at 13 cm in the Hawr al Hammar. Some males matured at 10.5 cm and some females at 10.6 cm in the creek. Naama et al. (1986) studied reproduction in the Hawr al Hammar and found a prolonged breeding period from November to April during which two batches of eggs could be shed. Maturity is attained at the end of the first year of life for both sexes when they are about 10 cm total length, with all males mature at 16 cm and all females at 13 cm. Egg diameter is up to 0.75 mm. Epler et al. (1996) record a relative fecundity of 359,873 to 756,118 eggs for fish of age groups 1+ to 3+ from three lakes in Iraq. Mature eggs were found from January to May in one lake and from November to March in another, maturity being two months earlier in saltier water. Daoud et al. (1998) examined reproductive biology in fish from the Tigris River at Al-Khadhmia north of Baghdad. They found a sex ratio of 1:1.45 (male:female), significantly different from 1:1, a breeding season from December to March, sexual maturity in all fish longer than 10 cm total length, smallest mature male 10.4 cm, smallest mature female 10.6 cm, absolute fecundity 12,033-63,836 eggs and relative fecundity 1237 eggs/g. Epler et al. (2001) studied reproduction in lakes Tharthar and Habbaniyah and found both sexes to achieve maturity in the first year of life at 14.2 cm. Spawning occurred in May and fecundity was 652-791 thousand eggs/kg body mass. Eggs are shed on, and adhere to, vegetation.

Rashid (19940 gives details of the histological changes in the ovaries for fish from the Qarmat Ali canal, along with several indices. The gonosomatic index showed no significant change from October to December but increased from 3.03 in December to 9.43 in January and 16.4 in February, decreasing subsequently suggesting a gradual spawning took place in March and into April. The hepatosomatic index showed a significant increase in February to 3.34 and a significant decrease in March to 1.32, suggesting the liver was involved in vitellogenin synthesis necessary to oocyte development. The somatic index increased from 0.65 in October to 1.15 in December as the fish lay down reserves, decreasing to 0.53 in February and rising throughout March and April.

Parasites and predators

Bykhovski (1949) reports a new species of monogenetic trematode, Ancyrocephalus fluviatilis, from this species in the Karkheh River, Iran. Mokhayer (1981c) reports a high infestation rate with larvae of the nematode Contracaecum in Karun River fish. This has health consequences for humans if fish are eaten raw, or are inadequately salted, smoked or cooked. There has been no report yet of this parasite from humans in Iran.

Moghainemi and Abbasi (1992) record a wide range of parasites from this species in the Hawr al-Azim in Khuzestan. Mortazaei et al. (2000) report the worm Neoechinorhynchus tylosuri in 54.8% of fish examined from Khuzestan marshes. Farahnak (2000) and Farahnak et al. (2002) record Anisakidae from this species in Khuzestan. Barzegar et al. (2008) record the digenean eye parasites Diplostomum spathaceum and Tylodelphys clavata from this fish. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Caligus lacustris, Lamproglena sp. and Ergasilus sp. on this species.

Economic importance

It appears in stores as a regular food fish in Ahvaz, Khuzestan, and is an important food fish in southern Iraq (Al-Nasiri and Islam, 1978) with 212,850 kg marketed at Basrah from October 1975 to June 1977 (Sharma, 1980). It forms about 29% of the total annual catches of freshwater fishes in southern Iraq and about 7.4% of the total annual catch for the whole country despite its small individual size (Mhaisen and Yousif, 1989; Mhaisen and Al-Jaffery, 1989). Its caloric value is 162.42 Kcal/100g (Mahdi et al., 2005). Salim (1962) records this species jumping into boats in Iraq when attracted by lights. It is important on fish farms in Iraq where it competes for food with cultured carp (Al-Shamma'a and Jasim, 1993) and Mhaisen and Al-Jaffery (1989) believed it could be farmed successfully because of its high growth rate. Taken also in the subsistence fisheries of the Indus River in Pakistan.

van den Eelaart (1954) gave the fishing season for this species as December-January (peaking in January) in rivers, and December-March (January) for lakes and marshes in Iraq.

Balasem et al. (2000) showed this species to be more sensitive to pollution than other Iraqi species tested, namely Barbus (= Carasobarbus) luteus, Cyprinus carpio, Carassius carassius (sic, probably C. auratus) and Gambusia affinis (sic, probably G. holbrooki). The LC50 of L. abu to copper after 72 hours exposure was 0.4 p.p.m. while the other species showed no mortality at this level. L. abu was also more sensitive to arsenic than these species and also Barbus (= Mesopotamichthys) sharpeyi, Barbus (= Luciobarbus) xanthopterus, Barbus (= Tor) grypus and Ctenopharyngodon idella (Ali et al., 1999). Wahab (1999) also tested the toxicity of four organophosphate insecticides to young fish, namely Dursban, Superacid, Malathion and Nogos with LC50 at 0.102. 0.125, 0.875 and 1.2 5 mg/L respectively. Salinity has a significant effect on the toxicity. This mullet has also been used to assess the impact of gas oil from oil spills in the Shatt al Arab on ionic regulation, on the chloride cells and on the gills (Faddagh et al., 2004). It was found to be sensitive to this form of pollution and suitable as an indicator species. Velayatzadeh et al. (2010) found the heavy metals copper and zinc in muscle tissues of fish from the Karun, Bahmanshir and Dez rivers. Beheshti et al. (2011) examined fish from the Dez River for copper, iron, zinc and manganese and found iron and manganese to be present at contaminating levels according to international standards. Maghtouie et al. (2011) examined fish for mercury levels from the Arvand River and found no health threat for consumption by humans.

Conservation

Mhaisen and Yousif (1988) recommend banning fishing from mid-January to mid-May during the spawning season and instituting an increase in fishing net mesh size to 23.2 mm.

Further work

Variation in this species should be re-examined to determine if nominal species and subspecies are valid. Biology and stock assessments for Iranian waters need to be carried out.

Sources

Type material: Liza abu zarudnyi (ZISP 24336).

Iranian material: CMNFI 1979-0240, 3, 57.1-70.7 mm standard length, Fars, Lake Famur (ca. 29º31'N, ca. 51º50'E); CMNFI 1979-0304, 12, 21.5-50.0 mm standard length, Fars, Lake Famur (ca. 29º31'N, ca. 51º50'E); CMNFI 1979-0352, 5, 11.4-22.6 mm standard length, Khuzestan, marsh 57 km east of Abadan (30º33'30"N, 48º48'E); CMNFI 1979-0353, 3, 15.7-18.3 mm standard length, Khuzestan, Karun River (30º22'30"N, 48º17'E); CMNFI 1979-0354, 6, 18.4-25.2 mm standard length, Khuzestan, Karun River tributary (30º31'N, 48º19'E); CMNFI 1979-0355, 3, 16.7-29.4 mm standard length, Khuzestan, Karun River tributary (30º35'N, 48º22'E); CMNFI 1979-0360, 12, 17.7-35.8 mm standard length, Khuzestan, canal branch of Karkheh River (31º40'N, 48º35'E); CMNFI 1982-0369, 11, 39.3-57.9 mm standard length, Khuzestan, Karkheh River (no other locality data); CMNFI 1993-0149, 2, 106.0-142.2 mm standard length, Khuzestan, Karun River (no other locality data); ZSM 25387, 4, 56.6-90.9 mm standard length, Khuzestan, Karun River at Ahvaz (31º19'N, 48º42'E); ZSM 25700, 6, 41.0-57.4 mm standard length, Khuzestan, fish pond near Ahvaz (no other locality data); uncatalogued 1, 169.8 mm standard length, Khuzestan (no other locality data).

Comparative material: BM(NH) 1971.4.2:1, 1, 130.2 mm standard length, Iraq, pond 25 km from Mosul (no other locality data); BM(NH) 1973.5.21:179-184, 6, 41.4-51.4 mm standard length, Iraq, Tigris River at Jadriyah, Baghdad (no other locality data); BM(NH) 1974.2.22:10-13, 4, 107.3-119.5 mm standard length, Iraq, Najab Bazar (no other locality data).

Liza aurata
(Risso, 1810)

Anzali Shore, January 2012, courtesy of K. Abbasi
Anzali Shore, January 2012, courtesy of K. Abbasi

Common names

kefal-e auratus, mahi kefal-e talaee (= golden mullet fish), kefal-e zarin, kafal.

[gizili kefal in Azerbaijanian; gatykelle, singil, orsbalyk in Turkmenian; singil' in Russian; golden mullet, golden grey mullet, long-finned mullet].

Systematics

Mugil Auratus was originally described from Nice, France. No major relevant synonyms.

Ghodsi et al. (2011) examined fish from Gomishan and Miankale regions in Golestan using microsatellite markers but found conspicuous genetic variation. There was evidence of a genetic bottleneck in the populations.

Key characters

This species is distinguished from its relative in the Caspian Sea by each head and antero-dorsal flank scale having only one pit or groove (occasionally double), 6-11 pyloric caeca of about equal length or gradually becoming longer from ventral to dorsal, scales on the snout ending anteriorly as a single row of small scales, and the oral edge of the preorbital bone is moderately concave. The upper jaw reaches posteriorly to a level with the posterior nostril, the only species in the genus with this character. Young have two vertical dark lines at the origin of the caudal fin rays and a herring bone pattern on the flanks (Harrison in Miller, 2003).

Morphology

The first dorsal fin has 4 strong spines, the last weakly developed compared to the first three. The second dorsal fin has 1-2 weak spines (more resembling unbranched rays than spines) and 6-9, usually 7 or 8 soft rays (the second unbranched ray or spine may be branched near the tip - the mode of 8 may include this ray in literature sources). The anal fin has 3 spines and 7-10, usually 9 branched soft rays. The pectoral fin has 2 unbranched rays and 13-17 branched rays and the pelvic fin has 1 spine and 5 branched rays. Scales number 40-49 in lateral series. There is no pectoral axillary scale. There is an elongate first dorsal fin axillary scale and a pelvic axillary scale. Scales have an almost vertical anterior margin with the anterior dorsal and ventral corners square to pointed. This margin is slightly indented where radii intersect it. The dorsal and ventral scale margins are parallel and the posterior margin is rounded. The focus is posterior and the posterior field has weak ctenii. Circuli are fine and numerous and radii are limited to the the anterior field, numbering up to 13 in fish examined here but probably size dependent. The adipose membrane of the eye is rudimentary. The gill rakers are fine and numerous, apparently finer and more numerous than in Liza saliens, exceeding 100 in the larger fish examined here. Raker number is probably size dependent. Rakers are slender, compressed and serrated medially. When appressed, a raker reaches the thirteenth raker below. The gut is elongate with several anterior and posterior loops after a muscular stomach. The chromosome number is 2n=48 (Klinkhardt et al., 1995).

Meristic values for Iranian specimens are:- first dorsal fin spines 4(30); second dorsal fin with 1(3) or 2(27) "spines" or unbranched rays and 6(1), 7(27), 8(1) or 9(1) branched rays; pectoral fin unbranched rays 2(30) and branched rays 15(7), 16(20) or 17(3); pelvic fin branched rays 5(30); lateral line scales 41(2), 42(4), 43(8), 44(3), 45(8), 46(4) or 47(1); and total vertebrae ?.

Sexual dimorphism

Unknown.

Colour

The body is a dusky grey to blue-grey with a silvery flank and belly. The back and upper flank have a series of dark to golden stripes. There is a golden blotch on the operculum. The peritoneum is dark brown to black. See also Key characters.

Size

Commercial catches in Iran during the 1950s weighed 0.3-0.8 kg, sometimes as much as 1.5-1.8 kg (Farid-Pak, no date). Reaches 59 cm and 2.5 kg on the coast of Turkmenistan. The Fisheries Research Centre in Mazandaran caught an exceptional specimen of this species weighing 1.5 kg and measuring 54 cm long (Iranian Fisheries and Research Training Organization Newsletter, 3, 1994).

Distribution

Found in the Mediterranean and Black seas and from the British Isles south to South Africa.

First reported from Iranian waters in 1933 by Shukolyukov (1937a) and by Dmitriev (1946), this species and Liza saliens became acclimated to the Caspian Sea over a period of 30-35 years and are regarded as naturalized from the latter half of the 1960s onward in Soviet waters (Marti, 1940; 1941; Khoroshko, 1982). It was first introduced to the Soviet Caspian Sea in 1930-1931 (Baltz, 1991). Kaplin in Mandych (1995) indicates that these grey mullets entered from the Black Sea during a Caspian transgression but this is incorrect.

In Iran, this species is found in the lower reaches of rivers along the Caspian coast, the Anzali Mordab and its outlets where numbers will probably increase with increasing salinity (Holčík and Oláh, 1992), the Safid River, Gorgan Bay, the southeast Caspian Sea, southwest Caspian Sea and south-central Caspian Sea (Abbasi et al., 1999; Kiabi et al., 1999; Abdoli and Naderi, 2009).

Zoogeography

This species is an exotic in Iran.

Habitat

This mullet inhabits the sea and enters the lower reaches of rivers, and is occasionally found in nearby lakes. It is recorded from depths of 5-700 m (Caspian Sea Biodiversity Database, www.caspianenvironment.org). Eggs develop in the open sea, larvae migrate to the coast and young feed along the shore and in bays. It winters along the Iranian shore in coastal waters, has a feeding migration in surface waters beginning in March to shallow areas of the middle Caspian and, by September-October, mature fish migrate over deeper water (300-700 m) of the middle and south Caspian to spawn. Feeding ceases at 6-8ºC and death occurs below 1.5ºC. The optimum temperature is 23-25ºC but young can be found in shallow water at 37.5ºC. It occurs from fresh water to salinities of 57‰ with mass mortalities recorded at 65‰ (Caspian Sea Biodiversity Database, www.caspianenvironment.org). Amini Ranjbar and Sotoudebnia (2005) assessed heavy metal (Zn, Cu, Pb and Cd) concentrations in fish from the Caspian Sea in Mazandaran and found higher than normal levels of Pb and Cd in muscle tissues.

Age and growth

Life span is up to 12 years in the Caspian Sea (Khoroshko, 1982). Maturity is attained at 38-45 cm and an age of 3-4 years for males and 5-6 years for females (RaLonde and Walczak, 1972). Fish taken from commercial catches in Iran are mostly 3-7 years old, 30.2-49.1 cm long and weigh 354-1266 g (Razivi et al., 1972). Fazli (1998) studied this species in the southeastern Caspian Sea of Iran and found that both scales and opercula could be used in ageing, although the  latter were better. Maximum growth occurred at 3 years of age. The b value in the relation between length and weight was 3.019 and sex ratio was 1:1.3 with females dominating. Ghaninejad et al. (2010) sampled fish from along the Iranian coast captured in 2007-2008. The overall male/female sex ratio at 1:1.22 differed significantly from 1:1 although only fish from Golestan were significantly different, size range was 19.0-50.2 cm, mean 32.7 cm, weight was 67 g to 1475 g, mean 411 g, age range was 2 to 10 years with mean age 4.42 years, age groups 3,4 and 5 years constituted 62% of the age composition, peak spawning was in October off Gilan but in November off Mazandaran and Golestan, average absolute fecundity was 700,881 eggs with a range of 200,112 to 2,282,862 eggs, and length of females at 50% maturity was estimated at 28.4 cm.

Food

This species is reported to enter the Anzali Mordab to feed from January to March (Holčík and Oláh, 1992), a result of the increasing salinity of this lagoon. Food items are small benthic invertebrates and detritus with some insects and plankton. Most stomach contents contain numerous sand grains. Adults scrape periphyton from rocks, silt and artificial structures. The predominant food items found by Ghadirnejad and Ryland (1996) from fish taken in the southern Caspian Sea were bivalves, foraminiferans and calanoid copoepods. Other food items were ostracods, eggs, nematodes, Nereis and cyclopoid copepods. Juveniles fed on ostracods and copepods while adults preferred bivalves, foraminferans and Nereis. This fish consumed more foraminferans than its relative and less copepods.

Reproduction

This mullet spawns in more northerly waters of the Caspian than its relative Liza saliens. Spawning begins in the middle of July in the middle Caspian and ends in the south Caspian from the middle to the end of October. Intensive spawning in the middle Caspian takes place in August-September at 20-22°C and a transparency of 7-11 m. Spawning depth is in the upper 0.5 m. The main spawning areas are at least 50-60 miles (80-96 km) from the coast (Avanesov, 1972) and this prevents the spawning area being easily exploited. Fecundity exceeds 4.4 million eggs in the Caspian Sea (Khoroshko, 1982); in Mazandaran average absolute fecundity reaches 772,350 eggs (Abdoli et al., 1996). The eggs are pelagic and have a diameter of 0.9 mm. In Iran, Gorgan Bay is believed to be a very important nursery and this is the major species caught by fishermen in winter (www.ramsar.org/ram_rpt_37e.htm, downloaded 28 July 2000). Shabanipour (1995) and Shabanipour and Heidari (2004) carried out histomorphological studies on the ovary of this species in Iran. Fazli (1998) found populations in the southeastern Caspian Sea of Iran spawned in September-November based on the gonadsomatic index, with most spawning in October. Males are ready to reproduce earlier than females. Absolute fecundity was 586,165 eggs. Fazli et al. (2008), again examining populations in the southeastern Caspian Sea, report a reproductive season of October to December, a male:female ratio of 1:1.42, sexual maturity at 26.0 cm fork length, mature gonads in 20% of fish at age 3, 63% at age 4, 88% at age 5 and 97% at age 6, individual absolute fecundity from 113,386 to 1.47 million eggs with a mean of 451,963. They concluded that the overlap between the fishing and spawning seasons required a delay of one month in the former to protect stocks. Heidari et al. (2009) studied late oogenesis in fish from Anzali. Recently spawned oocytes were lemon in colour and up to 0.8 mm in diameter.

Parasites and predators

Mokhayer (1976b) records the trematode Saccocoelium obesum from this species on the Iranian coast. This is a parasite of the intestine, with contamination rates of 90.4%, 88.6%, 81% and 34% for autumn, winter, spring and summer respectively. Gills carry Microcotyle mugilis at a rate of 2.0-2.3% and fish are also infected with Ancylodiscoides (sic) at a rate of 19.6%, 43%, 72% and 43.3% for autumn, winter, spring and summer respectively (Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 27-28, 1996; Annual Report 1993-94, Iranian Fisheries Research and Training Organization, Tehran, p. 49-50, 1995). Naem et al. (2002) found species in the monogenean treamtode genus Ligophorus on the gills of this species from the western branch of the Safid River.

This species is eaten by Sander spp., Silurus glanis and Huso huso.

At Ziba-kenar beach, Gilan in February 2004, this species showed erratic swimming and belly-up posture. A post-mortem showed gas accumulation and distension in the swimbladder, a yellowish liver, liquefaction of the gall bladder, excess micro-sand in the the caecum and hyperaemia of the intestine. The cause is thought to be an infectious agent closely related to a betanovirus or nodavirus (Iranian Fisheries Research Organization Newsletter, 38:3, 2004; Zorriehzahra et al., 2005). Soltani et al. (2010) identified a betanodavirus from mass mortalities in 2006-2008.

Economic importance

The catch in Iran for this species and Liza saliens from 1956/1957 to 1961/1962 varied from 166,197 kg to 960,282 kg (Vladykov, 1964) and from 1965/66 to 1968/69 varied from 490 to 1916 tonnes (Andersskog, 1970). Vladykov (1964) and RaLonde and Walczak (1972) cite figures for the Bandar-e Anzali region for Liza aurata from 1937/38 to 1961/62 of 14 kg to about 113 t and for 1969/70 a catch of 1,085 t was recorded, while for the whole Northern Shilot (Fisheries Company) in the period 1965/66 to 1968/69 catches varied between 548 t and 1916 t. Holčík and Oláh (1992) report a catch of only 3 kg in the Anzali Mordab in 1990. The Food and Agriculture Organization, Rome records the following catches for the 6 years from 1980 to 1985 for mullets in Iran, presumably from the Caspian Sea:- 150, 400, 1500, 2733, 2135, and 2200 t respectively. Moghim et al. (1994) give a total biomass of 2400 tonnes for this species in the southern coastal area of the Caspian Sea with a maximum sustainable yield of 960 tonnes (figures for Liza saliens are 7000 tonnes and 2900 tonnes respectively). Caspian Sea catches of this species in Iran for the year 1993-1994 was 22% of the bony fish catch with Liza saliens comprising only 4% (the major part of the catch was safid mahi, Rutilus frisii, at 62%)(Annual Report 1993-94, Iranian Fisheries Research and Training Organization, Tehran, p. 83, 1995). Catches in former Soviet waters of the Caspian Sea are about three-quarters or more this species and one-quarter Liza saliens (Khoroshko, 1982). The main season for fishing in the Caspian Sea off Iran is from December to February, peaking in January (Farid-Pak, no date). Abdoli et al. (1996) note that beach seiner cooperatives started fishing a month earlier than usual in 1993, coinciding with the spawning season in October. Annual catch variations occur in Iranian waters, this species having an abundance of 86.5% in 1994 decreasing to 38% in 1995 (Iranian Fisheries Research and Training Organization Newsletter, 9:5, 1995). The catch of this species declined from 76.9% to 41.2% of the total mullet catch from 1993-1994 to 1994-1995 through overfishing of juveniles (Iranian Fisheries Research and Training Organization Newsletter, 11:7, 1996). Abdolmalaki (2001) records a catch per unit effort for beach seining as 93 kg in 1991, falling to 33 kg in 1997 through overfishing. The mean length of the catch in Guilan decreased from 1991 to 1997 while in Mazandaran it increased. Fazli and Ghaninejad (2004) give the year 2001 as that of the maximum mullet catch in their survey of the years 1993-2001, the catch increasing through this period. Catch per unit effort in each seine declined from 114 kg in 1993 to 43 kg in 1996, increasing to 66.4 kg and 78.4 kg in 2000 and 2001. For the years 2000 and 2001, L. aurata had a mean fork length of 32.7 and 32.3 cm, a weight of 418.3 and 419.8 g, and a K value of 0.61 and 0.93. They concluded that the harvest of this species is at a sustainable level. Heydatifard et al. (2002) state that its catch is greater than that of other bony fishes (presumably excluding kilka) in the southern Caspian Sea. S. M. E. J. Zorriehzahra (pers comm., 19 January 2011) noted a decline in catches from 6442 mt in 2002 to 2400 mt in 2010.

Hosseini et al. (2004) determined the shelf life of this species on ice in Iran was 10 days. Mazandaran fish have an average fat composition of 9.25%, 72% being unsaturated. Omega-3 fatty acids constitute 18.7% of the total fatty acids with a significant ratio to omega-6 fatty acids of 1.97 (Bedayatifard et al., 2002). Akhondzadeh Basteh et al. (2006) found the bacterial pathogens Listeria monocytogenes and Staphylococcus aureus in salted Liza aurata and Vibrio haemolyticus in fresh and salted L. auratus. Hedayatifard and Yousefian (2009; 2010) determined the comparative fatty acid composition in filleted fish that were fresh or dry-salted and found this fish to be one of the best sources for omega-3 essential fatty acids compared to other fishes in the Caspian Sea basin. Razavilar and Tavakoli (2006) studied the the prevalence of human toxigenic Clostridium botulinum and the need for food safety control measures.

Conservation

The successful introduction of this species and Liza saliens in the Caspian Sea has apparently led to a decline in various clupeid species (Alosa) and of Sander marinum (Baltz, 1991). Beach seines with mesh sizes of 28 mm, 33 mm and 36 mm caught non-standard fish as 43.3%, 35% and 2.2% of the total catch respectively. Seines with a cod end of at least 36 mm are recommended for stock protection (Iranian Fisheries Research and Training Organization Newsletter, 6:8, 1994) (note the report names the fish as Mugil cephalus but the catch was probably a mix of Liza aurata and L. saliens). Contamination of this species in Iranian waters with Pb, Ni and Zn was examined by Fazeli et al. (2005). The highest contamination was in the southwest part of the Caspian followed by the south centre and the southeast. The highest concentrations observed were 17.51 mg/kg for lead, 6.23 mg/kg for nickel and 647.28 mg/kg for zinc. Pazooki et al. (2009) found the heavy metals cadmium and chromium in fish from Bandar Anzali, more in the skin than in muscle and more in females than males, but not enough to constitute a health hazard.

Further work

As an important fisheries species, population numbers and biology should be carefully monitored on an ongoing basis.

Sources

Distinguishing characters based on Trewavas and Ingham (1972); Sakri (1993) gives an account in Farsi on the spawning and catch of mullets in the Iranian Caspian.

Type material: None.

Iranian material: CMNFI 1970-0509, 1, 124.2 mm standard length, Gilan, Safid River (37º24'N, 49º58'E); CMNFI 1970-0535, 1, 137.2 mm standard length, Gilan, Shafa River (37º35'N, 49º09'E); CMNFI 1970-0543, 3, 33.5-38.5 mm standard length, Gilan, Caspian Sea at Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1970-0543A, 4, 33.1-60.5 mm standard length, Gilan, Caspian Sea at Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1970-0563, 19, 52.8-87.0 mm standard length, Gilan, Caspian Sea, Kazian Beach (ca. 37º29'N, ca. 49º29'E); CMNFI 1970-0565, 4, 52.2-93.0 mm standard length, Gilan, Safid River estuary (ca. 37º28'N, ca. 49º54'E); CMNFI 1970-0581, 2, 147.7-151.5 mm standard length, Gilan, Caspian Sea near Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1970-0587, 1, 78.9 mm standard length, Mazandaran, Babol River (36º43'N, 53º39'E); CMNFI 1971-0326A, 20, 30.0-46.0 mm standard length, Gilan, Caspian Sea near Bandar-e Anzali (37º28'N, 49º27'E); CMNFI 1979-0430, 1, 42.0 mm standard length, Mazandaran, river 1 km east of Now Shahr (36º39'N, 51º31'E); CMNFI 1979-0470, 19, 29.4-57.4 mm standard length, Mazandaran, stream 21 km west of Alamdeh (36º35'N, 51º43'E); CMNFI 1979-0471, 3, 56.7-65.8 mm standard length, Mazandaran, Caspian Sea 14 km west of Alamdeh (36º35'N, 51º48'E); CMNFI 1979-0686, 2, 59.1-87.3 mm standard length, Gilan, Safid River near Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1980-0117, 2, 99.7-114.5 mm standard length, Gilan, Golshan River (37º26'N, 49º40'E); CMNFI 1980-0127, 6, 29.5-36.9 mm standard length, Gilan, Caspian Sea near Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1980-0146, 1, 135.7 mm standard length, Mazandaran, Gorgan Mordab at Ashuradeh-ye Kuchak(36º50'N, 53º56'E); CMNFI 1980-0149, 1, 38.4 mm standard length, Gilan, Chabak River (37º21'N, 49º50'E).

Liza saliens
(Risso, 1810)

Anzali Wetland, October 2010, courtesy of K. Abbasi
Anzali Wetland, October 2010, courtesy of K. Abbasi

Common names

kefal-e saliens, kafal-e poozehbarik, kafal.

[sivriburun kefal in Azerbaijanian; vitibutum or gatykelle in Turkmenian; ostronosik in Russian; sharpnose mullet, small mullet, grey mullet, leaping mullet].

Systematics

Mugil Saliens was originally described from Nice, France. No major relevant synonyms.

Key characters

This species is distinguished from its relative in the Caspian Sea by each head and antero-dorsal flank scale having 2-7 or more grooves (mostly 2 grooves), pyloric caeca in two groups, 3-5 short and 3-4 long (total 6-9), scales on the snout ending anteriorly as numerous rows of small scales, and the oral edge of the preorbital bone is deeply notched. Young lack the vertical dark lines at the origin of the caudal fin rays and the herring bone flank pattern seen in L. aurata (Harrison in Miller, 2003).

Morphology

The first dorsal fin has 4 spines, the last one weakly developed compared to the first three. The second dorsal fin has 1-3, usually 1or 2, spines (more resembling unbranched rays than true spines) and 6-9, usually 8 or 9 soft rays (the second unbranched ray or spine may be branched near the tip - modes of 8 or 9 in literature may include this ray). The anal fin has 3-4, usually 3, spines and 7-10, usually 9, soft rays. The pectoral fin has 2 unbranched rays and 14-17 branched rays and the pelvic fin has 1 spine and 5 branched rays. Scales number 42-50 in lateral series. There is no pectoral axillary scale. There is an elongate first dorsal fin axillary scale and a pelvic axillary scale. Scales have an almost vertical anterior margin with the anterior dorsal and ventral corners square to pointed. This margin is slightly indented where radii intersect it. The dorsal and ventral scale margins are parallel and the posterior margin is rounded. The focus is posterior and the posterior field has weak ctenii. Circuli are fine and numerous and radii are limited to the the anterior field, numbering up to 10 in fish examined here but probably size dependent. The gill rakers are fine and numerous, but not as fine and numerous as in Liza aurata, exceeding 70 in the larger fish examined here, and probably size dependent. When appressed, a raker reaches raker 6 or 7 below. Rakers have a serrated medial edge. Vertebrae 23-25. The gut is elongate with 2 posterior loops after a muscular stomach. The chromosome number is 2n=48 (Klinkhardt et al., 1995).

Meristic values for Iranian specimens are:- first dorsal fin spines 4(30); second dorsal fin with 1(6), 2(22) or 3(2) "spines" or unbranched rays and 6(1), 7(20) or 8(9) branched rays; pectoral fin unbranched rays 2(30) and branched rays 14(2), 15(27) or 16(1); pelvic fin branched rays 5(30); lateral line scales 42(2), 44(6), 46(6), 47(8), 48(4) or 49(4); and total vertebrae ?.

Sexual dimorphism

Males are smaller than females.

Colour

The back is dark grey to blue-grey or grey-brown, the belly pale to silvery or yellowish, and the silvery-grey flanks have about 7 bluish stripes. There is a golden blotch on the operculum. The peritoneum is brown to black.

Size

Commercial catches in Iran during the 1950s weighed 0.3-0.8 kg, sometimes as much as 1.5-1.8 kg (Farid-Pak, no date). Reaches 40.0 cm.

Distribution

Found in the Mediterranean and Black seas and along the Atlantic coast to South Africa.

First reported from Iranian waters by Dmitriev (1946), this species was introduced to the Soviet Caspian Sea in 1930-1931 (Baltz, 1991). In Iran it is reported from the lower reaches of such rivers as the Gorgan, Gharasu, Tajan, Talar, Babol, Haraz, Sardab, Pol-e Rud, and Safid, the Anzali Mordab and Gorgan Bay, the southeast Caspian Sea, southwest Caspian Sea and south-central Caspian Sea (Roshan tabari, 1987; Abbasi et al., 1999; Kiabi et al., 1999; Abdoli and Naderi, 2009).

Zoogeography

This species is an exotic in Iran.

Habitat

This is a pelagic species found mostly in inshore areas and entering lagoons and rivers. It is found in surface waters over depths of 5-700 m at salinities of 4-13‰. It migrates to the southern Caspian in autumn when water temperatures fall (Caspian Sea Biodiversity Database, www.caspianenvironment.org). Generally it tolerates fluctuations of 5-27Cº and 11-28‰ although fry need time to adjust to lower salinities (Harrison in Miller, 2003).

Age and growth

Life span is up to 10 years in the Caspian Sea (Khoroshko, 1982). Males mature at 3 years and females at 4 years (Caspian Sea Biodiversity Database, www.caspianenvironment.org). The maximum growth rate in fish sampled between Babolsar and Bandar-e Torkeman in the Iranian Caspian was observed in fish age 3 years. The operculum gave better ages than scales (Fazli, 2000). Patimar (2008a) examined fish in Gorgan Bay and found maximum ages of 6+ for males and 7+ for females. Both sexes grew allometrically (b = 2.478 for males, 2.545 for females) and rapidly during their first year of life, reaching more than 40% of their growth. Age and growth were in contrast to other populations in the southeast Caspian Sea. The von Bertalanffy growth functions were Lt = 30.415(1-e-0.275(t+0.645)) for males and Lt = 34.832(1-e-0.211(t+1.009)) for females. Males predominated at smaller sizes and females at larger sizes but see below.

Food

Diet is very similar to that of Liza aurata, comprising detritus, periphyton and small benthic organisms Stomach contents contain sand grains in Iranian fish. The predominant food items found by Ghadirnejad and Ryland (1996) from fish taken in the southern Caspian Sea were bivalves, foraminiferans and calanoid copoepods. Other food items were ostracods, eggs, nematodes, Nereis and cyclopoid copepods. Juveniles fed on ostracods and copepods while adults preferred bivalves, foraminferans and Nereis.

Reproduction

Spawning starts at the end of May and the beginning of June in the southern Caspian Sea and continues until the end of September and the beginning of October, most intensively between June and August off the Turkmenistan coast according to Avanesov (1972). Iranian fish spawned in July-August and males ripen earlier than females (Fazli, 2000; Yousefian et al., 2003). Yousefian et al. (2003) give details of oogenesis, detailing the stages in gonad development. Spawning occurs over depth ranges of 5-700 m but most spawning in the southeastern Caspian is at 5-7 m. Temperatures are between 17 and 29°C with the most intensive spawning when sea surface temperature is 25-29°C. Water transparency is 1-9 m. The greatest aggregations of eggs are found 5-7 miles (8-11 km) from shore in the eastern Caspian. Fecundity exceeds 2.1 million eggs in the Caspian Sea (Khoroshko, 1982). Egg diameter is up to 0.8 mm. Females predominate in Iranian samples examined by Fazli (2000) at a ratio of 1:3.14. Patimar's (2008a) Gorgan Bay fish reproduced in May-July with an average fecundity reaching 389,790 eggs in 7+ fish and relative fecundity 4281 eggs/g. The low absolute fecundity contrasted with other populations in the southeast Caspian Sea. Ghaninejad (2011) found fish in markets illegally (May to September) and in legal commercial fisheries (October to April) off Gilan to have a significant 1:2.23 sex ratio (male:female), a peak spawning in July, an average absolute fecundity of 207,050 eggs (range 29,700-512,900 eggs), and length of females at 50% maturity was estimated at 23 cm.

Parasites and predators

This species has a worm parasite in the intestine, Saccocoelium obesum, with contamination rates of 62%, 10.7%, 12% and 22.2% for autumn, winter, spring and summer respectively, lower than in Liza aurata. Gills carry Microcotyle mugilis at a rate of 2.0-2.3% and fish are also infected with Ancylodiscoides (sic) at a rate of none, none, 1.6% and 4% for autumn, winter, spring and summer respectively, much lower than in Liza aurata (Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 27-28, 1996; Annual Report 1993-94, Iranian Fisheries Research and Training Organization, Tehran, p. 49-50, 1995).

Economic importance

The main season for fishing in the Caspian Sea off Iran is from December to February, peaking in January (Farid-Pak, no date). Some catch information is summarised above under Liza aurata which is 5-6 times more abundant in catches. However a report on the 1994-1995 catch showed a severe reduction of 2500 tonnes, as Liza saliens replaced L. aurata as the main part of the catch. L. aurata declined from 76.9% of the mullet catch in 1993-1994 to 41.2% in 1994-1995, the cause being overfishing of juveniles (Iranian Fisheries Research and Training Organization Newsletter, 11:7, 1996). Moghim et al. (1994) give a total biomass of 7000 t for this species in the southern coastal area of the Caspian Sea with a maximum sustainable yield of 2900 t (figures for Liza aurata are 2400 tonnes and 960 tonnes respectively). Annual catch variations occur in Iranian waters, this species having an abundance of 13.5% in 1994 increasing to 62% in 1995 (Iranian Fisheries Research and Training Organization Newsletter, 9:5, 1995). Abdolmalaki (2001) records a catch per unit effort for beach seining as 78 kg in 1991, falling to 15 kg in 1997 through overfishing. The mean length of the catch in Gilan and Mazandaran did not decrease from 1991 to 1997, the former in contrast to Liza aurata. Fazli and Ghaninejad (2004) give the year 2001 as that of the maximum mullet catch in their survey of the years 1993-2001, the catch increasing through this period. Catch per unit effort in each seine declined from 114 kg in 1993 to 43 kg in 1996, increasing to 66.4 kg and 78.4 kg in 2000 and 2001. For the years 2000 and 2001, L. saliens had a mean fork length of 27.5 and 25.1 cm, a weight of 224.7 and 179.1 g, and a K value of 0.91 and 0.71.

Conservation

See above under Liza aurata. Mercury levels at 0.0102-0.108 mg kg -1 w.w. in this species from the Iranian shore were lower than the WHO guideline of 0.5 mg kg-1 w.w. (Agah et al., 2007).

Further work

As an important fisheries species, population numbers and biology should be carefully monitored on an ongoing basis.

Sources

Distinguishing characters based on Trewavas and Ingham (1972).

Type material: None.

Iranian material: CMNFI 1970-0510, 34, 38.1-72.8 mm standard length, Gilan, Golshan River (37º26'N, 49º40'E); CMNFI 1970-0507, 9, 23.6-46.8 mm standard length, Gilan, Caspian Sea at Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1970-0508, 1, 50.6 mm standard length, Gilan, Safid River at Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1970-0509, 4, 31.9-66.8 mm standard length, Gilan, Safid River at Hasan Kiadeh(37º24'N, 49º58'E); CMNFI 1970-0528, 16, 43.6-64.4 mm standard length, Mazandaran, Tajan River estuary (36º49'N, 53º06'30"E); CMNFI 1970-0535, 1, 153.3 mm standard length, Gilan, Shafa River (37º35'N, 49º09'E); CMNFI 1970-0539, 43, 25.1-102.0 mm standard length, Gilan, Caspian Sea off Bandar-e Anzali (37º28'N, 49º27'E); CMNFI 1970-0543A, 2, 76.7-79.9 mm standard length, Gilan, Caspian Sea at Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1970-0549, 1, 48.1 mm standard length, Gilan, Qareh Su near Alb Imamzadeh (no other locality data); CMNFI 1970-0565, 3, 91.8-108.8 mm standard length, Gilan, Safid River estuary (ca. 37º28'N, ca. 49º54'E); CMNFI 1970-0577, 13, 22.1-38.8 mm standard length, Gilan, Caspian Sea near Astara (ca. 38º26'N, ca. 48º53'E); CMNFI 1970-0581, 4, 41.0-71.0 mm standard length, Gilan, Caspian Sea near Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1970-0586, 8, 27.3-50.4 mm standard length, Mazandaran, Gorgan Mordab at Ashuradeh-ye Kuchak(36º50'N, 53º56'E); CMNFI 1970-0587, 165, 15.7-47.9 mm standard length, Mazandaran, Babol River at Babol Sar (36º43'N, 52º39'E); CMNFI 1970-0590, 11, 28.6-54.0 mm standard length, Mazandaran, Shesh Deh River near Babol Sar (ca. 36º43'N, ca.52º39'E; CMNFI 1979-0470, 17, 16.6-43.4 mm standard length, Mazandaran, stream 21 km west of Alamdeh (36º35'N, 51º43'E); CMNFI 1979-0471, 1, 59.1 mm standard length, Mazandaran, Caspian Sea 14 km west of Alamdeh (36º35'N, 51º48'E); CMNFI 1979-0477, 1, 16.7 mm standard length, Mazandaran, Gorgan Mordab at Bandar-e Shah (36º54'N, 54º02'30"E); CMNFI 1979-0788, 1, 22.7 mm standard length, Mazandaran, Gorgan River at Khadje Nafas (37º00'N, 54º07'E); CMNFI 1980-0120, 9, 48.8-73.2 mm standard length, Mazandaran, Babol River at Babol Sar (36º43'N, 52º39'E); CMNFI 1980-0126, 2, 217.5-244.2 mm standard length, Gilan, Caspian Sea near Bandar-e Anzali (37º28'N, 49º27'E); CMNFI 1980-0127, 5, 22.8-52.8 mm standard length, Gilan, Caspian Sea near Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1980-0129, 18, 28.3-64.2 mm standard length, Mazandaran, Tajan River estuary (36º49'N, 53º06'30"E); CMNFI 1980-0136, 13, 35.9-76.8  mm standard length, Mazandaran, Fereydun Kenar River estuary (36º41'N, 52º29'E); CMNFI 1980-0146, 2, 88.8-96.6 mm standard length, Mazandaran, Gorgan Mordab at Ashuradeh-ye Kuchak(36º50'N, 53º56'E); CMNFI 1980-0149, 10, 36.0-83.2 mm standard length, Gilan, Chabak River (37º21'N, 49º50'E); CMNFI 1980-0157, 77, 25.7-50.1 mm standard length, Mazandaran, Gorgan River estuary (36º59'N, 53º59'30"E); CMNFI 1993-0144, 2, 78.4-92.2 mm standard length, Mazandaran, Neka Power Plant (36º51'48'N, 53º23'24"E).

Liza subviridis
(Valenciennes, 1836)

Courtesy of J. E. Randall
Courtesy of J. E. Randall

Common names

biah or biah arabi at Bushehr, garriz at Bandar Abbas; meid or maid (for small specimens); kefal posht sabz.

[biag, meid or maid, biah akhtar, beyah akhtar in Arabic; greenback mullet, greenback grey mullet, silver mullet].

Systematics

Originally described from Malabar, India. Synonyms are Mugil dussumieri Valenciennes in Cuvier and Valenciennes, 1836 and Mugil jerdoni Day, 1876. Placed in the genus Chelon Rose, 1793 in Randall (1995b).

Key characters

The preorbital bone does not occupy the whole space between the eye and upper lip as in all other mullets.

Morphology

The head is wide and flattened with head length 23-27% of standard length. There is a well-developed adipose eyelid, covering about half of the iris, but this is absent in young fish. The lower lip has a weak to marked symphysial knob (authors differ) and both lips are thin. The upper jaw extends back on a level with the anterior nostril to the anterior eye rim. Upper jaw teeth are in several rows and ciliiform lower jaw teeth in one row. The preorbital bone is strongly notched anteriorly and only occupies three-quarters of the space between the eye and upper lip. Scales are regularly arranged and are dense on the second dorsal and anal fins. The second dorsal fin origin lies over the anterior half of the anal fin base. The pectoral axillary scale is rudimentary or absent. The pectoral fin does not reach back on a level with the first dorsal fin origin and is 74-76% of head length.

The first dorsal fin has 4 spines, the second dorsal fin has 1 spine and 8-9, usually 8, soft rays and the anal fin has 3 spines and 8-9, usually 9, soft rays. Scales in lateral series 27-32. Transverse scales 10-11, usually 11. Pyloric caeca 4-5.

Ahmed and Hussain (2003) describe the larvae of this species from Khawr az Zubayr and the Shatt al Arab estuary. Al-Hassan and Madhi (1989) studied enzyme polymorphisms in populations from the Shatt al Arab, Khawr az Zubayr and Kuwait but found no evidence of sub-population differentiation. A single stock is found in the northern Persian Gulf.

Sexual Dimorphism

Unknown.

Colour

The back is dark to light green or greyish-green, the flanks silvery to white and the belly white. There may be 3-7 blackish stripes along the flank but these may not always be obvious. The caudal fin is bluish and has a black margin, other fins are hyaline.

Size

Attains 39.5 cm total length and 270 g (Al-Hassan and Al-Seyab, 1992).

Distribution

Found in the Persian Gulf and eastwards to India, China, northern Australia and Polynesia. Reported from the Shatt al Arab and Hawr al Hammar, Iraq near Iranian Khuzestan by Al-Hassan et al. (1989), Al-Hassan and Al-Seyab (1992) and Hussain et al. (1997; 2001). Records of Liza dussumieri from the Tigris River at Amara and Baghdad, the Euphrates River and the Shatt al Arab by Kennedy (1937), Khalaf (1961), Mahdi (1962), Al-Nasiri and Hoda (1976) and Hussain et al. (1989), if correctly identified, are referred to this species (Thomson and Luther in Fischer and Bianchi, 1984).

Recorded from the lower Mand River in Bushehr Province at 28°14'N, 51°51'E, 28°11'N, 51°32'E and 28°07'N, 51°24'E by M. Rabbaniha (pers. comm., 1995) and from Khuzestan in the Zohreh and Bahmanshir rivers and the Hawr al Azim by Najafpour (1997). Records from Iran could be L. vaigiensis (q.v.). No specimens have been examined by me to confirm these anecdotal and literature records.

Zoogeography

A widely distributed marine species, this mullet also enters and is resident in fresh waters but is dependent on reproduction of marine populations to maintain the freshwater ones.

Habitat

This species lives in the sea, particularly in coastal waters, lagoons and estuaries, but regularly enters fresh water. At a freshwater station on the Shatt al Basrah Canal, Iraq with salinities up to 3.5‰, Al-Daham and Yousif (1990) found this species to be dominant, comprising 59.6% by number and 40.0% by weight. Eggs of this species and L. klunzingeri were most abundant in the inner part of the Shatt al Arab estuary in February with larvae most abundant in February and April. The temperature range for several sites was 12.5-22ºC and salinity 32-38‰. Temperature is the most important factor influencing spawning (Ahmed and Hussain, 2000). This was one of the more abundant species in the recovering Hawr al Hammar of southern Iraq in 2005-2006 and in the marshes in the 1980's (Hussain and Ali, 2006).

Age and Growth

Shadi et al. (2011) give a length-weight relationship for fish from the northern Persian Gulf off Khuzestan with a (intercept) = 0.017 and b (slope) = 2.819 for fish 7.2-16.5 cm total length, mean 11.5 cm.

In the Shatt al Basrah Canal of southern Iraq, populations of this species are comprised of age groups 1 to 6 but most fish are in age groups 1 and 2 (Al-Daham and Wahab, 1991). Age 2 dominates from February to April and age 1 from June to January. Females are slightly longer in each age group and have a longer life span than males. The oldest females are age 6 and 303 mm and the oldest males age 4 and 251 mm. Males are less numerous than females with a sex ratio 1.0:1.4. Maturity is attained at age 1 and the smallest mature male is 137 mm and the smallest mature female is 142 mm. Ali et al. (1999) examined the stock of this species in Khawr Abdullah and found the infinity length (L) to be 34.3 cm total length (TL), the growth coefficient (K) 0.31, annual total mortality (Z) 1.06, natural mortality (M) 0.70 and fishing mortality (F) 0.366. The exploitation rate (E) was 0.34 indicating that the stock was slightly exploited during September 1989-August 1990. The length-weight relationship was W = 0.0000208 L2.985. Mohamed et al. (1998) found, for fish from the northwest Arabian (= Persian) Gulf, a length-weight relationship of W = 1.34 x 10-5 L2.961, condition factor ranged from 0.98 to 1.21 and the growth model was Lt = 308 (1-e-0.225 (t + 0.618)).

Ali et al. (1999) summarise annual growth of this species from their own and other studies as follows, attributing differences to differing methodologies:-

Author

sex

L

K

1

2

3

4

5

6

Wahab (1986) (Shatt al Basrah)


29.4
40.2

0.39
0.151

14.6
15.5

19.3
20.5

22.1
23.0

24.0
25.2

-
27.3

-
29.0

Al-Hisnawi (1990) (Khawr az Zubayr)


25.2
48.9

0.474
0.141

11.4
11.35

16.55
16.78

19.84
20.92

21.85
24.31

-
27.82

-
30.5

Mohammed et al. (1998) (Northwest Arabian Gulf)

♂ + ♀

30.8

0.225

9.4

13.6

17.0

19.7

21.9

23.7

Ali et al. (1999) (Khawr Abdullah)

♂ + ♀

34.3

0.31

8.9

15.5

20.37

23.97

26.65

28.64

Muhsin (1988) determined protein, lipid and ash content of fish taken monthly from the marine Khawr az Zubayr and found fluctuations to follow the reproductive cycle. Lipid content of the liver increased in winter and of the carcass in winter and spring, the pre-spawning period, along with increases in the gonadosomatic and hepatosomatic indices. The latter indices peaked in March and decreased sharply until June. Total protein decreased during winter and spring from the  carcass and in spring from liver and ovaries as they were used for energy and to support maturation of the ovaries.

In Malaysia, maturity is attained at 95-115 mm for males and 105-115 mm for females (Chan and Chua, 1980).

Food

Food items are algae, diatoms and detritus extracted from ingested mud and sand. Al-Hassan and Madhi (1987) include higher plants as part of the diet and Al-Hassan and Al-Seyab (1992) copepods for Iraqi waters. Mohamed et al. (1998) found this species to be a detritivore in the northwest Persian Gulf. Hussain and Ali (2006) examined feeding relationships among fishes in the Hawr al Hammar, Iraq and found this species to be a herbivore, 75.2% of the diet being plants and algae. In another study of the recovering Hawr al  Hammar, diet was 80.84% algae and 15.66% diatoms with amounts of crustaceans and plants being less than 10% each (Hussain et al., 2006).

In Malaysian estuaries and coastal areas, Chan and Chua (1979) working in Malaysia found this species to feed only on zooplankton when less than 12 mm standard length, becoming bottom feeding at 16-20 mm on zooplankton, diatoms and detritus and by 24 mm zooplankton was absent but filamentous algae was added where available. The proportion of the various food items varied with the locality.

Reproduction

Spawning occurs in the sea, with all eggs released at once in a Malaysian population (Chan and Chua, 1980). Al-Daham and Wahab (1991) were unable to locate eggs and larvae in fresh waters of Iraq. They believe that adults leave the Shatt al Basrah Canal to spawn in the sea. However, Al-Hassan and Madhi (1987) give a reproductive season of February through April in the Shatt al Arab, Iraq and Ahmed and Hussain (2003) give March to June in the offshore area near the mouth of the Khawr az Zubayr and Shatt al Arab. Muhsin (1988) determined April-June to be the spawning season, with mature ova released in May, in the Khawr az Zubayr (see above). Male and female gonads are best developed in February to March and spawning is deduced to have occurred from March to May, possibly offshore in the Persian Gulf. In Malaysia, spawning can occur from June to November and in the off-season (Chan and Chua, 1980). In Iraq, fry 27-40 mm in length are captured from April to June in fresh water. The adult mullet return to fresh water after spawning. Fecundity was 133,224-295,065 eggs for fish 182-243 mm total length, this being higher than in Malaysia at 40,000-145,000. Fecundity is somewhat lower than in other mullets but may be attributed to year round spawning at some localities (Chen and Chua, 1980). Al-Hassan and Madhi (1987) give a fecundity of 549,278 eggs with egg diameter up to 0.65 mm and Al-Hassan and Al-Seyab (1992) give 580-590,000 eggs for their large specimen from Hawr al Hammar.

Parasites and predators

Unknown.

Economic importance

This species forms part of coastal fisheries in shallow waters and is caught with a variety of nets, particularly seines, stationary traps, gill nets and cast nets. It forms a substantial part of brackish water fisheries in southern Iraq, being the most abundant species in estuaries, and economically the most important mullet (Al-Daham and Wahab, 1991). Ali et al. (1999) gave a catch of 35 t in Khawr Abdullah, Iraq, almost 6% of the marine catch. Mohamed et al. (1998) gave a value of 3.4% of the commercial catch for this species with catch rates in the northwest Arabian (= Persian) Gulf of 0.15 kg/h in June to 0.75 kg/h in October. This species is potentially important in fish culture as it is herbivorous. Consequently ponds can be fertilised inexpensively and plant food supplements such as rice bran and peanut meal can be given. Chan and Chua (1979) recommend that fish for stocking should be at least 30 mm standard length as their bottom feeding habit is then most suitable for pond culture.

The proximate chemical composition measured in fish from the Basrah fish market was moisture 76.16%, protein 16.82%, fat 5.1% and ash 1.82% and the general condition was a maximum of 1.78 (Ali et al., 2004). It is characterised as a fatty fish according to a lipid content 9-14% by wet weight of muscle in autumn (Hantoush et al., 1999).

Conservation

Population numbers and trends are unknown so conservation requirements cannot be ascertained.

Further work

Materials of freshwater mullets should be collected in Iranian waters and studied for their identity and biology.

Sources

Description based on literature sources, particularly Blegvad and Løppenthin (1944), Fischer and Bianchi (1984), Talwar and Kacker (1984) and Thomson (1997).

Liza vaigiensis
(Quoy and Gaimard, 1824)

Common names

[massiyat or biah in Iraq; square-tail mullet, diamond-scaled mullet, diamond-scale grey mullet, brown-banded mullet].

Systematics

Originally described from Waigeo, Indonesia. Placed in the genus Ellochelon Whitley, 1930 in Randall (1995b).

Key characters

This species is uniquely characterised in this area of the northen Indian Ocean by the caudal fin being only slightly emarginate. All other local mullets have an obviously forked tail fin.

Morphology

The preorbital bone is notched anteriorly and is wide, filling the space between the eye and the upper lip. The head is broad and flattened on top, with head length 24-30% of standard length. The eye has adipose tissue around its margin. Lips are thin but the lower lip has a high symphysial knob. There are ciliate teeth on the lips, in 1-2 rows on the upper lip and 1 row on the lower lip. Larger fish lose the upper lip teeth at >250 mm standard length and the lower lip teeth at ca. 100 mm standard length. About half of the anal fin base lies anterior to the level of the second dorsal fin origin. The pectoral axillary scale is rudimentary or absent. The pectoral fin does not reach back on a level with the first dorsal fin origin. The second dorsal and anal fins are densely scaled. Hoda (1978) describes the larva in Iraq.

First dorsal fin spines 4, second dorsal fin with 1 spine and 7-9 soft rays, anal fin with 3 spines and 8 soft rays, pectoral fin with 15-18 branched rays, lateral series scales 24-29, transverse scales (from first dorsal fin origin to pelvic fin origin) 9-11, and predorsal scales 15-16. Gill rakers 40-61.

Sexual Dimorphism

Unknown.

Colour

The back is olive brown to pale olive, the flanks silvery and the belly white with yellowish tinges. Upper flank scales have characteristic brown blotches which appear to run together to form about 5 stripes. The iris has yellow patches. Fin margins are dusky to black but the body of each fin is yellowish. Young fish have black pectoral fins but in adults the lower half is yellow. The caudal, anal and pelvic fins are a bright yellow in young fish while the second dorsal fin is dark on its distal half.

Size

Reaches 63 cm.

Distribution

This species is widespread in the tropical Indo-Pacific Ocean from Africa to Japan and Tahiti including the Persian Gulf. It also enters fresh waters. Reported from the Shatt al Arab, Iraq near Iranian Khuzestan by Al-Nasiri and Hoda (1976). A single specimen from Khuzestan, Iran confirms its presence in fresh water. Akbari (1998) records it from coastal and estuarine waters in Hormozgan. Other records from Iranian rivers draining to the Persian Gulf (e.g. Dez, Karun, Dalaki and Mand rivers) may be this species but could also be L. subviridis.

Zoogeography

A widely distributed marine species, this mullet also enters and is resident in fresh waters but is dependent on reproduction of marine populations to maintain the freshwater ones.

Habitat

This species is found in coastal waters over sand, in lagoons and over reef flats and commonly enters fresh waters. It apparently enters fresh water during the rainy season elsewhere in its range, ascending only to tidal limits.

Age and Growth

Unknown.

Food

Food items are algae, diatoms and detritus whih are ingested along with associated sand and mud.

Reproduction

Fry are known to occur along the sea coast.

Parasites and predators

None reported from Iran.

Economic importance

This mullet is common in commercial catches along the west coast of India. It is caught in castnets, stakenets and beach seines.

Conservation

Population numbers and trends are unknown so conservation requirements cannot be ascertained.

Further work

Materials of freshwater mullets should be collected in Iranian waters and studied for their identity and biology.

Sources

Description based on literature sources, particularly Fischer and Bianchi (1984), Talwar and Kacker (1984) and Thomson (1997).

A single specimen from Khuzestan, Iran was examined measuring ?.

Genus Mugil
Linnaeus, 1758

This genus is characterised by a strongly flattened head, a transverse mouth with the lateral cleft short, thin lips, lower jaw with a symphysial knob and lower jaws meeting at symphysis with an acute angle, the maxilla end is not visible when the mouth is closed, the anteroventral edge of the preorbital is serrate and straight and the ventral edge is slender and pointed, no true teeth in the jaws, a very well-developed adipose eyelid reaching to, or nearly to, the pupil, two pyloric caeca, pharyngobranchial organ with a single valve, no vomerine or palatine teeth, scales cycloid or feebly ctenoid, and pectoral axillary scale long and pointed. Thomson (1997) and Harrison in Miller (2003) give a wide range of generic characters but since only one species is found in Iranian waters, and that rarely, the species description suffices here.

Mugil cephalus
Linnaeus, 1758

Common names

kefal or kafal, biah-e sarpahn (= flathead mullet), كفال خاكستري (= kafal-e khakestari).

[biah, biyah, biaha Ramadiyah, anubah or asfatiya in Arabic; striped mullet, loban, common, grey, jumping, flathead grey, sea, or river mullet; mighach, minghaj or karul in Pakistan].

Systematics

Mugil Cephalus was originally described from the Atlantic Ocean at the shores of Europe, entering rivers. No major relevant synonyms. Durand et al. (2012) indicate, based on DNA data, that this wide-ranging species may include up to 14 species so the Iranian taxon may eventually bear a distinct name.

Key characters

This species is distinguished from Liza species by having only 2 pyloric caeca, the posterior end of the maxilla is straight rather than curved down behind the premaxilla and visible behind the corner of the closed mouth, adipose tissue on the eye reaches the pupil in adults, and the tip of the jaw end is on the line of the gape.

Morphology

There is a prominent adipose eyelid which leaves only a narrow slit over the pupil and extends twice as far behind the eye as in front of it.

First dorsal fin with 4 spines, second dorsal fin with 1-2 spines and 6-9, usually 8or 9, soft rays, anal fin with 3 spines and 7-9, usually 8, soft rays (below 5 cm there are 2 spines and 9 soft rays), pectoral fin branched rays 14-17, and pelvic fin branched rays 5.

Lateral series scales 36-46 with 3-5 on the caudal fin, transverse scale rows 14-15, predorsal scales 24-26 and cheek scales in 3-4 rows. Scales on the head extend forward to the tip of the snout. Scales are found on the caudal fin rays and at the bases of the anterior dorsal and anal fin rays. The pectoral axillary scale is very long, about one-third the length of the fin, and there is a pelvic axillary scale. Scales bear an anterior mid-margin notch, the anterior corners are very square and sharp, upper and lower margins are parallel and the posterior margin is rounded. There are few radii (as few as 5) running from a subcentral posterior focus to the notch. Circuli are numerous and fine and the exposed scale surface is coarse. Gill rakers 24-80, long, fine and crowded, count increasing with growth of fish. Pyloric caeca number 2 and the gut is long with numerous loops, complexly coiled. The chromosome number is 2n=48 (Klinkhardt et al., 1995).

There is a distinct axillary scale at the pectoral fin, 33-36% of pectoral fin length. The pectoral fin reaches the tenth postopercular scale but not the level of the first dorsal fin origin; folded forward it reaches the eye but not the pupil.

A single Iranian specimen has 4 first dorsal fin spines, 8 soft second dorsal fin rays, 8 anal fin soft rays, 15 pectoral fin branched rays, 5 branched pelvic fin rays, 43 scales in lateral series, and 24 total vertebrae.

Sexual dimorphism

Females grow larger and weigh more than males.

Colour

Back bluish-grey to olive-green or greyish-bown, flanks silvery and belly silvery to white. Each scale has a dark spot which line up to give the impression of 6-12 indistinct brownish or grey stripes. The upper rear corner of the operculum has a golden spot. There is dark purple blotch at the pectoral fin base. The dorsal and pectoral fins are a dark blue-grey while the anal and caudal fins are a yellowish-green. The dorsal and caudal fins have dusky margins. Young are a bright, iridescent silver. Estuarine and freshwater fish may be duller overall than marine fish.

Size

Attains 1.2 m and 7 kg.

Distribution

Found world-wide between about 51°N and 42°S. Introduced into the Caspian Sea in 1902, it has not been observed there since and there appear to be no valid Caspian records for Iran (Dmitriev, 1946; Baltz, 1991). The introduction probably failed because the pelagic eggs were not buoyant at the low Caspian salinities (Baltz, 1991) although Harrison in Miller (2003) notes spawning has been induced in fresh water. However Kiabi et al. (1999) report it from the southeast Caspian Sea and the species is being cultured at Sari on the Iranian coast of the Caspian Sea (Iranian Fisheries Research Organization Newsletter, 40 & 41:3, 2004; and see below). Kolodar and Abdoli (2004) and Yelghi et al. (2012)  report it from the Gomishan Lagoon but only in farms there.

Reported from the Shatt al Arab, Iraq near Iranian Khuzestan by Al-Nasiri and Hoda (1975b). Akbari (2002) records this species from creeks and coastal waters of Hormozgan. It enters rivers of southern Iran from the Persian Gulf and Sea of Oman, notably the Sarbaz River.

Zoogeography

A widely distributed marine species, this mullet also enters and is resident in fresh waters but is dependent on reproduction of marine populations to maintain the freshwater ones.

Habitat

This mullet is found in surface waters of the sea and frequently ascends rivers. It is common in coastal and estuarine waters. It favours marine waters where the average monthly temperature is above 16°C and where summer temperatures rise above 18°C. Temperature ranges of 5-37°C are tolerated. It can live in salinities up to 126‰. It is a fast swimmer and may leap out of the water.

Age and growth

Age at maturity for both sexes varies with water temperatures, at 1 year in the warmest waters, such as Florida, and as late as 6-8 years in the Black Sea for example. Black Sea fish were 31-37 cm long at maturity while in the Gulf of Mexico females were 25.8 cm and males 24.0 cm and larger. Life span lies between 11 and 21 years. Growth rate varies with locality and ceases in warm temperate waters during mid-winter.

Food

Diet is comprised of microscopic organic matter taken in with mouthfuls of sand. The sand probably helps grind up food in the gizzard-like stomach. The food items are diatoms, blue-green and green algae, foraminifers, small crustaceans and detritus, and sometimes plankton. Rarely, swarming marine worms are eaten rather than microscopic items, perhaps as an additional energy source prior to spawning. Juveniles eat invertebrates.

Reproduction

In the sea, spawning occurs in surface waters near the edge of the continental shelf over deep water after a migration from rivers. In the Sea of Marmara, Turkey, this takes place in October and in southern India in October-May. Eggs and larvae drift with ocean currents until they are 20-30 mm about 2-3 months after hatching. Fry make their way into estuaries and juveniles live in estuaries and lower reaches of rivers. Each female is attended by several males during spawning. The males nudge and press against the female's abdomen. The fish may quiver and cease swimming momentarily, sometimes rising to the surface. Fertilisation is external. Spawning occurs once yearly and some females only spawn in alternate years after first maturity. Fecundity is up to 4,800,000 non-adhesive, straw-coloured eggs and egg diameter reaches 1.08 mm (some reports cite 7 million eggs). Hatching occurs in about 2 days.

Parasites and predators

Eslami and Anwar (1971) record the cestode Caryophyllaeus fimbriceps from this species on the Caspian coast of Iran. In the absence of valid records for this fish in Iranian Caspian waters, the host may have been another mullet species.

Economic importance

Fisheries take place around coasts world-wide as the mullet merge into migratory schools. In Hormozgan, is is caught in coastal waters by cast net and stake-net but only at 1.33% by weight and 0.19% by frequency compared to other edible fishes and with a frequency of 5% compared to other mugilids. World-wide, it is sold fresh, frozen or salted and the roe is made into a type of caviar. This species is also widely cultured, for example in Israel and India, with yields up to 2434 fish per hectare. It has been reported as being ciguatoxic (intermittently poisonous through feeding on toxic food) (Bagnis et al., 1970) and as being ichthyoallyeinotoxic (hallucinogenic fish poisoning) (Halstead, 1967-1970; 1978). The latter is a sporadic and mild form of poisoning. This species is rare in Iranian freshwaters and unlikely to be health hazard there (Coad, 1979b).

Fingerlings from Taiwan, Egypt and Hong Kong have been cultured in enclosed ponds in Mazandaran and the Gomishan Wetland (A. Matinfar, pers. comm., 1995; Annual Report 1993-94, Iranian Fisheries Research and Training Organization, Tehran, p. 43-44, 1995; Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 31, 1996; Annual Report, 1995-1996, Iranian Fisheries Research and Training Organization, Tehran, p. 39, 42, 1997; Iranian Fisheries Research and Training Organization Newsletter, Tehran, 18:5, 1997; Iranian Fisheries Research Organization Newsletter, 40 & 41:3, 2004; Iranian Fisheries Research Organization Newsletter, 53:3, 2008). 20,000 fingerlings, weighing about 0.5 g, were imported to the Caspian Sea Ecology Research Center in Sari in 1993. Fry were cultured for 21 days on plankton grown by manuring ponds with wheat and rice bran and soya. Growth in salt water averaged 235 g and 28.4 cm in the first year and 544 g and 40 cm in the second year. Growth in fresh water averaged 216 g and 28.5 cm in the first year and 668 g and 38 cm in the second year. Experiments on the mass production of Nannochloropsis oculata algae used in the culture of rotifers and Mugil cephalus larvae have been carried out in Golestan Province (Iranian Fisheries Research Organization Newsletter, 37:3, 2003). Mirhashemi Rostami et al. (2008) has examined mortality factors in fry from 9 year old broodstock in Iran.

Sharifpour et al. (2003), Ghelichi et al. (2004), Yeganeh et al. (2005), Mirhashem Rostamy et al. (2006) and Tehrani (2006) give further details on culturing this mullet in Iran. Peak spawning was mid to late January with the best time for artificial breeding using hormones being December. Up to 2.6 million eggs could be produced from a female with fertilisation varying from 10 to 95% and hatching rate between 0.008 and 88.9%. Ghelichi and Jorjani (2004) state that this species is expected to play an important future role in fish culture in Iran. They studied induced spawning in this species and found 17-α hydroxyprogesterone to be effective. Gelichi et al. (2007) studied oocyte maturation after stimulation with hormones and also found 17-α hydroxyprogesterone to be the most effective steroids in final maturation.

Conservation

Population numbers and trends are unknown so conservation requirements cannot be ascertained.

Further work

The biology and frequency of occurrence of this species in fresh waters of Iran is unknown.

Sources

Thomson (1963) gives a synopsis of the biology of this species and the above account is based mostly on his work. Distinguishing characters are based on Trewavas and Ingham (1972).

Iranian material: OSU 4281, 1, 234.3 mm standard length, Baluchestan, Kalani, Sarbaz River (25º17'N, 61º24'E).

Atherinidae

The silversides or sand smelts are found in coastal areas of temperate to tropical seas and in fresh water. There are about 14 genera and 68 species (Nelson, 2006; Eschmeyer and Fong, 2011) with only one reported from the Caspian Sea and Iran. Most are small fishes with a maximum length of 60 cm.

These small, silvery fishes have a moderately elongate body; usually cycloid and moderately large scales; no lateral line but sometimes a pit on each scale; small teeth in the jaws and sometimes on the roof of the terminal, upwardly-directed mouth; wide gill openings with branchiostegal membranes free of the isthmus; 5-6 branchiostegal rays; gill rakers usually long and slender; two well-separated dorsal fins, the first with 3-10 unbranched but flexible rays, and the second with 1-2 unbranched and the rest branched rays; anal fin long; pectoral fins high on the flank; no pyloric caeca; egg membranes with filamentous outgrowths; back bluish to greenish with small melanophores but translucent; and a silvery stripe along the flank, often distinctively outlined with black.

Silversides can occur in vast schools in inshore waters and are an important item in the diet of other fishes. They have been used as bait but are too bony to be much used as food. Their food is plankton. Eggs are large and greenish. The sticky egg filaments entangle with plants, rocks or sand as anchors until hatching.

Genus Atherina
Linnaeus, 1758

Members of this speciose genus are found in fresh and brackish waters with a single representative in Iran. The record of Atherina hepsetus Linnaeus, 1758 in the Caspian Sea by Quignard and Pras in Whitehead et al. (1984-1986) is an error (Vasil'eva, 1994).

The body is compressed with a rounded belly, the mouth large and terminal, jaws large, reaching back to the anterior eye level or beyond, sides of the upper jaw are straight and premaxillaries protractile, the dentary bone has a high central portion, a preopercular notch is absent, rows of setiform teeth on the jaws, vomer and palatines, cycloid scales extending onto the head, short pectoral fins, vent nearer the pelvic fin origin than the anal fin, numerous gill rakers, 5-10 flexible rays in the first dorsal fin, and the second dorsal fin is similar in length to the anal fin with its origin above the anal fin.

Atherina caspia
Eichwald, 1831

Anzali Shore, December 2009, courtesy of K. Abbasi
Anzali Shore, December 2009, courtesy of K. Abbasi

Common names

shisheh mahi (= glass fish), گل آذين ماهي (gol azin mahi, meaning flower decoration fish), atrinka (from the Russian).

[aterinka or xazar aterini in Azerbaijan; kaspi aterinasy in Turkmenian; Kaspiiskaya aterinka or Caspian silverside in Russian; Caspian sand smelt, Boyer's sand smelt, big-scale sand smelt].

Systematics

Formerly called Atherina mochon pontica natio caspia Eichwald, 1831 or Atherina mochon caspia in Caspian Sea literature. Atherina mochon Cuvier, 1829 was described originally from Ivasa and Atherina presbyter pontica Eichwald, 1831 from near Odessa, Ukraine. The Caspian Sea taxon was also known as Atherina boyeri Risso, 1810. Atherina Boyeri was originally described from the sea shore and lower course of rivers in the Département du Var, France. Reshetnikov et al. (1997) give the date for this species as 1826. Atherina presbyter var. caspia Eichwald, 1831 described from "in Caspii maris littore australiore, sinu balchanensi" (i.e. southern shore of the Caspian Sea, Balkhan Bay) was considered a synonym. No types are known. Naseka and Bogutskaya (2009) consider caspia to be a full species.

Syntypes of A. boyeri are in the Muséum national d'Histoire naturelle, Paris under MNHN A-4342 (2) and MNHN B-860 (1) (Eschmeyer's "Catalog of Fishes", downloaded 29 August 2007).

Bamber and Henderson (1985) demonstrated how the morphology of this species around Britain varies according to temperature and salinity during embryonic development. Isolation of populations results in local selection and random genetic drift and thus recognisable morphologies but these were not recognised as distinct. Kottelat (1997) reviews literature reports of variation in this species in the Mediterranean and Black seas and neighbouring fresh waters and concludes that lacustrine populations should be called A. boyeri while marine populations are A. mochon. The status of Caspian populations is not commented on.

The silverside in the Caspian Sea was referred to as Atherina boyeri caspia by Savenkova and Asanov (1991) and by Vasil'eva (1994; 1996b) and this seemed a reasonable step as the population is isolated from other populations in the Atlantic Ocean and Mediterranean Sea. The subspecies was characterised by a reduced number of infraorbital bones (3 bones in Caspian Sea fish as opposed to 4 in Black Sea fish; apparently infraorbital bones 2 and 3 fuse), a reduced number of gill rakers (generally 19-27 as opposed to 27-37), and the form of the maxilla (the inferior margin in Caspian Sea fish is always smooth while in the Black Sea and Sea of Azov a "wing" is usually developed). However Kiener and Spillman (1972) allocated Caspian populations to this subspecies on the basis of larger size, hardly an adequate criterion. Vasil'eva (1994) makes the allocation based on a reduced number of gill rakers, reduction of the number of bones around the eye to three (preorbital, infraorbital and postorbital) and a smooth lower margin to the maxilla without a wing protuberance.

Dobrovolov and Ivanova (1999) studied two non-enzymatic proteins and 11 enzymes for putative Atherina boyeri and A. mochon pontica. They concluded that these are distinct species and indeed the Black Sea fish are a distinct species, A. pontica. They diverged 2.316MYA. These authors did not examine Caspian Sea material. Miller in Miller (2003) followed a conservative approach, regarding the various named and wide-ranging populations as representing a single polymorphic species.

Key characters

The two dorsal fins, cycloid scales, pectoral fin high on the flank and the vent remote from the anal fin are characteristic.

Morphology

First dorsal fin spines 5-10; second dorsal fin with 1-2 spines and 8-15 soft rays; anal fin with 1-2 spines and 9-18 soft rays; pectoral branched rays 10-17, usually 12-15; pelvic fin with 1 spine 5-7, usually 5, branched rays; lateral series scales 37-53, possibly to 61, see Vasil'eva (1994); total gill rakers 19-27 in the Caspian Sea, to 39 elsewhere, spinulose on the interior surface, and long and reaching about 7 rakers along the arch when appressed; and vertebrae 39-52. The anus is 4-5 scales in advance of the anal fin. Scales are higher than broad, with slight indentations on the otherwise straight dorsal and ventral margins, a rounded posterior margin, and a wavy to rounded anterior margin with a protuberant central point. There are no radii. Circuli are restricted to the anterior third of the scale with a central and vertical roughened area posterior to the circuli, presumably made up of fragmented circuli. The focus is central. The lower jaw symphysis fits into a notch in the upper jaws. The haemal arches of the anterior 4-7 caudal vertebrae are expanded around the swimbladder. Gut s-shaped. The chromosome number is 2n=48 (Klinkhardt et al., 1995).

Meristic values for Iranian specimens are: first dorsal fin spines 8(28), 9(5) or 10(1); second dorsal fin soft rays 11(10), 12(21), 13(2) or 14(1); anal fin soft rays 13(7), 14(15), 15(9) or 16(3); pectoral fin branched rays 12(1), 13(12), 14(14) or 15(6); pelvic fin branched rays 5 (34); caudal fin branched rays 13(1), 15(30), 16(2) or 17(1); lateral series scales 50(1), 51(10), 52(5), 53(3), 54(8), 55(3), 56(3) or 57(1); predorsal scales 17(1), 18(2), 19(12), 20(8), 21(3), 22(3), 23(1), 24(3) or 26(1); caudal peduncle scales 12(31), 13(2) or 14(1); transverse scales from anal fin antero-dorsally 10(2), 11(6), 12(23) or 13(3); total gill rakers 24(3), 25(4), 26(14), 27(8), 28(4) or 29(1); total vertebrae 45(1), 46(19), 47(7) or 48(7).

Sexual dimorphism

Females tend to be larger than males and there are some differences in morphometrics and meristics (Ghoorbanalidoost et al., 2003).

Colour

General colour is given in the family account. The lateral band is strongly developed and bright in Caspian Sea specimens. The belly is white, fins are pale to translucent grey. Lagoon specimens are brownish or grey-brown on the back. The peritoneum is brown to black, eggs being encased in a black peritoneum while the abdomen wall is a light brown.

Size

Reaches 11.8 cm and 17.6 g in the Atrak River (Savenkova and Asanaov, 1991), to 14.5 cm standard length in the Caspian Sea basin generally, and elsewhere to 19.0 cm standard length (Henderson and Bamber, 1987). Kiener and Spillman (1972) found a maximum of 15.4 cm in their Caspian Sea sample.

Distribution

Found from the shores of Europe to the Black and Caspian seas, and in the Uzboi Valley of Turkmenistan. In Iran, it is reported along the whole Caspian Sea coast, and from the the Safid River, Talar River, Anzali Mordab, and its outlet the Sowsar Roga seasonally, the Bandar Anzali breakwater and beach, Farahbad, Mazandaran, and Gorgan Bay (Abbasi et al., 1999; Holčík and Oláh, 1992; Roshan Tabari, 1997; Kiabi et al., 1999; Abdoli and Naderi, 2009).

 

Zoogeography

This species probably entered the Caspian Sea from the Black Sea during Khvalyn transgression (10-70,000 years B.P.) via the Kumo-Manych Depression (Kosarev and Yablonskaya, 1994). Berg (1948-1949) contends that it entered the Caspian from the Black Sea in post-glacial times while most other Caspian fishes are relicts of earlier transgressions or migrants from northern waters.

Habitat

The Caspian silverside is a schooling fish found at depths exceeding 100 m but is concentrated at 10-20 m. Salinity tolerance is high, up to 77‰ (Miller in Miller, 2003), up to 60‰ in the Atrak River (Savenkova and Asanov, 1991). Reproduction can occur up to 42‰> while preferred levels are 3-12‰. A wide temperature range is tolerated, 0-31ºC (Miller in Miller, 2003). It is also found in lagoons and river mouths, and enters rivers to spawn, against currents up to 1.2 m/sec. It is the dominant species in the Gomishan Wetland in spring, summer and autumn (Patimar et al., 2009). Holčík and Oláh (1992) report its apparent recent occurrence in the Anzali Mordab in response to increased salinity there. It is also known from fresh water in Lenkoran.

Age and growth

Maturity is attained in the first year and life span varies from 1 to 5 years, 4-5 years usually in the Caspian Sea basin (Henderson and Bamber, 1987). Miller in Miller (2003) summarises growth and population dynamics from populations outside the Caspian Sea. Ghoorbanalidoost et al. (2003) found populations in the south Caspian Sea to be 73.82 mm long on average and 3.15 g in weight, with age  groups 1-3 years, average 1.7 years, total length and length-weight relationship W = 0.00000615L3.02 and a sex ratio of male:female = 0.47:0.53. Heydarnejad (2009) gave the length-weight relationship for an Iranian sample as W = 0.0326TL3.033.

Food

Savenkova and Asanov (1991) report plankton, eggs and juvenile fishes to be food items in the Atrak River of Turkmenistan. Some populations also eat benthic organisms such as amphipods, worms and molluscs. Iranian specimens contained encysted cladocerans and beetles in their guts. It has a trophic plasticity, adapting to whatever conditions obtain (Miller in Miller, 2003).

Reproduction

Spawning in this species is intermittent and occurs along the coast from May to July, peaking in mid-May to mid-June (Savenkova and Asanov, 1991) or from April to August (Henderson and Bamber, 1987). Preserved Iranian fish samples have relatively large eggs from at April to September, e.g. 1.4 mm on 27 April, 1.6 mm on 14 May. Savenkova and Asanov (1991) also studied the annual spawning migration into the Atrak River (lower reaches in Turkmenistan, upper reaches shared with Iran). Fish are caught in a fish ladder 18-19 km from the sea and in the river. The first schools appear in the Atrek mouth as early as mid-February at water temperatures of 8-10°C but the mass migration takes place in mid to late March and the first half of April at 14-22°C. Spawning occurs in March, April and May but most intensively in April at flood water temperatures of 13.7-23.0°C. There is a larger migration in high-water years. Sex ratio is about 1:1. A female may be able to spawn 5-6 times in one season so that egg numbers and diameters vary within each individual for different generations of eggs. Egg deposition in the Caspian Sea is associated with the alga Cladophora to which the eggs are attached or entangled by long filaments (up to 15) and this plant is present in the Atrak. Egg diameters are up to 2.0 mm (3.5 mm in Europe, but possibly these are water hardened) and the light yellow eggs number up to 568 (Gon and Ben-Tuvia, 1983) although the Caspian Sea Biodiversity Database (www.caspianenvironment.org) gives 5500 eggs per female (presumably total seasonal fecundity). Only one gonad develops. Larvae are pelagic but may school close to shore.

Henderson and Bamber (1987) and Gon and Ben-Tuvia (1983) review the reproductive biology of this sand smelt. It can rapidly adjust its life history to a range of environments, from fresh water to coastal water.

Parasites and predators

The Caspian seal, Pusa caspica, is a predator on this species (Krylov, 1984) as are the larger fishes such as Stenodus leucichthys and Alosa saposchnikowii (Lönnberg, 1900a).

Economic importance

This species is food for a number of other fishes including such economic species as sturgeons, Sander and the predacious shads. It has been caught as a bycatch in kilka seine nets and used in fishmeal production. In Europe it has been sold fresh or canned.

Conservation

Kiabi et al. (1999) consider this species to be of least concern in the south Caspian Sea basin according to IUCN criteria. Criteria include abundance in numbers, widespread range (75% of water bodies), absent in other water bodies in Iran, and present outside the Caspian Sea basin.

Further work

The biology of this species in Iranian waters has been investigated by Patimar (1995) (not seen).

Sources

Some meristics are taken from Altun (1999) and Trabelsi et al. (2002).

Iranian material: CMNFI 1970-0507, 11, 22.2-56.3 mm standard length, Gilan, Caspian Sea at Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1970-0509, 5, 49.5-101.7 mm standard length, Gilan, pond at Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1970-0543A, 7, 52.9-83.1 mm standard length, Gilan, Caspian Sea at Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1970-0563, 18, 39.8-112.5 mm standard length, Gilan, Kazian Beach (ca. 37º29'N, ca. 49º29'E); CMNFI 1970-0581, 5, 47.6-54.8 mm standard length, Caspian Sea near Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1971-0343, 2, 65.9-72.5 mm standard length, Gilan, Langarud at Chamkhaleh (37º13'N, 50º16'E); CMNFI 1979-0081, 3, 74.1-77.3 mm standard length, Mazandaran, Caspian Sea, 3 km west of Chalus (36º41'N, 51º24'E); CMNFI 1980-0146, 4, 92.1-103.6 mm standard length, Mazandaran, Gorgan Mordab at Ashuradeh-ye Kuchak (36º50'N, 53º56'E); and CMNFI 1980-0160, 2, 49.2-73.8 mm standard length, Iran, Caspian Sea basin (no other locality data).

Adrianichthyidae

Genus Oryzias
Jordan and Snyder, 1906

Oryzias latipes
(Temminck and Schlegel, 1846)

Introduced to the Tedzhen River basin and Karakum Canal in Turkmenistan (Aliev et al., 1988; Sal'nikov, 1995) but no confirmed Iranian record. It is reported as reproducing in Iran by the Food and Agriculture Organization, Rome (www.fao.org/scripts/acqintro/query/retrive.ide, downloaded 28 July 1999) but this record is presumably in error and needs confirmation by specimens.

Cyprinodontidae

The tooth-carps, killifishes or pupfishes are small fishes found in fresh, brackish and sea water. There are 9 genera and about 121 species found in tropical to warm temperate climates almost world-wide (Eschmeyer and Fong, 2011). Iran has 9 described species, with more to be discovered.

This family is characterised by a moderately elongate and compressed body, rather cyprinid-like but with jaw teeth (hence the common and scientific names), body and head covered with scales, no spines in the fins, no barbels, the mouth is very protractile and armed with comb-like teeth, the lower jaw is strong and robust as the dentary is expanded medially, 4-6 branchiostegal rays, gill membranes free from the isthmus, lateral line absent or reduced to points, caudal fin rounded or truncate, pectoral fins set low on the body (in contrast to Poeciliidae), dorsal and anal fins short, no adipose fin, no pyloric caeca, and no gonopodium (egg layers, in contrast to the livebearer subfamily in the Poeciliidae).

Wildekamp (1993) gives a general account of cyprinodontid species. These fish can be maintained in aquaria, and are very popular with aquarists. Specific requirements where known are given under each species. Moderate to hard fresh water is used and for water with a total hardness less than 10dGH, sea salt can be added (about one teaspoon per 10 litres). Summer temperatures of 20-25°C are good and although higher temperatures make the fish more active, they also age the fish more rapidly. Winter temperatures below 20°C imitate nature and are recommended. Weekly partial water changes are suggested and live and frozen fish foods and even flake foods are eaten. Breeding occurs throughout the year in aquaria kept above 15°C, eggs are deposited on fine-leaved plants (thread algae, Java moss or yarn mops) or even on aquarium filter sponges and gravel. Egg development takes 8-20 days, fry take about 2 days to absorb their yolk sac and will then consume baby brine shrimp. These fish can be kept in outdoor ponds in milder climates (http://ark.aka.org/Eurasian.htm, downloaded 1 December 2003).

Genus Aphanius
Nardo, 1827

These tooth-carps are found around the shores of the Mediterranean, in Southwest Asia and as far as northeastern India and Somalia. There are about 25 known species.

Sethi (1960) advocated placing Aphanius in a separate family, Aphaniidae, but this did not find general acceptance.

The genus Aphanius Nardo, 1827 has been used for these tooth-carps for many years. However, Lazara (1995) designated Lebias fasciata Valenciennes, 1821 as the type species for Lebias Goldfuss, 1820, making Lebias a subjective senior synonym of Aphanius. On this basis Lebias must be used rather than Aphanius but Lazara's type species designation is invalid (Kottelat and Wheeler, 2001). Since Lazara (1995) could have chosen another species as type species, the change involves a large number of species and these species are threatened and listed in various legislation, a petition is before the International Commission on Zoological Nomenclature to suppress Lazara's designation (Kottelat, 1997; Kottelat and Wheeler, 2001; Wildekamp, 2001; Villwock et al., 2002). Wildekamp et al. (1999) present evidence that Lebias is a synonym of Cyprinodon.

Literature on these Mediterranean and Southwest Asian fishes may appear under either of these names or under Cyprinodon Lacepède, 1803, the latter now restricted to American species.

Bănărescu (1995) disagrees with Parenti's (1981) relationship of Aphanius to South American Orestias (see also Parker and Kornfield (1995), Stevenson (1997) and Costa (1997) for conflicting views).

This genus is characterised by a thick oval body, large to moderate cycloid scales, the head flattened on top, a small, superior mouth with tricuspid teeth, the upper jaw bordered by the premaxillaries only, lateral line system present only on the head, dorsal fin positioned somewhat posteriorly with1-2 unbranched rays and 7-13 branched rays, anal fin rays 1-2 unbranched and 7-14 branched, dorsal and anal fins larger in males than in females, dorsal fin inserted opposite the anal fin origin (in contrast to Gambusia), and colouration of males and females distinct. Seyfali et al. (2002) has 2n=48 for fish from the Damghan basin which Coad and Abdoli (2000b) suggest may be a distinct taxon. Kamal et al. (2009, 2009) studied some biological characteristics of the Damghan population.

Aphanius is the only genus in the family currently recognised in Iran. However Parenti (1981) distinguishes derived members of the genus Aphanius as `Aphanius' without formally describing a new genus. One of the distinguishing features of `Aphanius' is the reduction of cephalic sensory pores to neuromasts, a character found in A. mento, A. sophiae and A. vladykovi in Iran.

Wildekamp et al. (1999) review the Turkish species which show scale reductions not seen in Iranian populations. Hrbek et al. (2002) outline the historical biogeography of the species complex in central Anatolia, Turkey using mtDNA, testing the hypothesis of geographic speciation driven by early Pliocene orogenic events that may well be paralleled in Iran. Hrbek and Meyer (2003) studied the phylogeny of Eurasian tooth-carps using mtDNA and observed a western Tethys Sea clade (all species except those listed below) with a middle Oligocene divergence into Iberian Peninsula and Atlas Mountains, and Turkey and Iran sections. Late Miocene orogenic events were correlated with a large amount of genetic differentiation in Turkey (and presumably Iran). An eastern Tethys Sea clade (dispar, ginaonis, mento, sirhani) had an Oligocene divergence into a freshwater clade inhabiting the Arabian Peninsula and neighbouring areas and a euhaline clade inhabiting coastal areas from Pakistan to Somalia. Speciation is predominately vicariant-based but ecological factors played a significant role.

These fishes are known generally as gour-e khar in Farsi (= literally "striped donkey" which means zebra here although usually in Iran this term refers to the wild ass or onager), kapurdandandar or kopurdandandar (= tooth-carp), or even آفانيوس (= Aphanius). Saefali (1999) is a recent study of these fishes in Farsi.

Iranian Aphanius away from coastal drainages are thought to be relicts of the Tethys Sea (Kosswig, 1955a; 1967; Bianco, 1995), having been trapped by the rising Iranian Plateau, rather than invaders from the coast (Krinsley, 1970). Fossil tooth-carps, Aphanius kirgisicus and A. longipinnis have been described from Kyrgyzstan in Miocene deposits, far inland from the distribution of living species (Yakovlev, 1959). Priem (1908) describes a fossil cyprinodont from the Miocene of Iran.

The systematics of the Iranian populations would repay careful study using biochemical and genetic techniques coupled with aquarium studies on behaviour and cross breeding. Certain populations, isolated from others in discrete drainage basins, appear to be different on the basis of colour patterns yet show little morphological variation, e.g. the Damghan population, see Coad and Abdoli (2000b), Saefali (1999) and Seyfali et al. (2002), the most northerly and easterly population in Iran characterised by large spots in females; populations in Fars outside the Lake Maharlu and Kor River basins; and populations in the Namak Lake basin - all yet to be investigated in detail (Coad, 2000b). Since these patterns are used in mate recognition, the populations could be distinct taxa. Huber (1996) has noted in cyprinodonts from Africa that many taxa can only be separated on the basis of colour in life, other characters overlapping. He suggests a cyprinodont species definition which includes possession of at least one stable phenotypic characteristic, and this could be colour of the male fish. He does recommend that genetic isolation be demonstrated where possible by karylogy, biochemical techniques or breeding experiments.

Seifali et al. (2003) compare A. sophiae and A. vladykovi meristically and morphometrically. Saifali et al. (2004) found a suite of morphometric characters to be different between samples of A. vladykovi and fish from Qomy-Abad in Tehran Province, the Namak Lake basin.

The mosquitofish (Gambusia) is a niche competitor and may well eliminate these tooth-carps. Generally, Aphanius prefers springs, lakes, marshes, sea shores and hot springs. Flowing water is usually avoided but they can be found near the source of springs flowing into salt lakes, constrained by hypersaline waters downstream. However they are tolerant of high salinities. They occur in schools but males are aggressive to other males. Males are brightly coloured, often striped, while females are subdued with spots or faint bars.

A related Mediterranean species is reputed to be ichthyootoxic but this has not been demonstrated for Iranian species (Coad, 1979b). Symptoms of this egg poisoning are summarised under the genus Schizothorax.

These small fishes are used in the aquarium trade as they tolerate a wide range of temperatures and salinities and are particularly colourful. As early as 1912, J. P. Arnold in Germany noted the survival of imported Iranian Aphanius (possibly A. farsicus) in aquaria at temperatures of 14-27°C, with a variety of plants such as Myriophyllum, Nitella, Riccia, Cabomba, Salvinia and filamentous algae, with daphnia, cyclops, enchytraeids and red mosquito larvae as food, and egg laying apparently preferred on the Salvinia with 8-12 days for hatching at 27°C.

Aphanius sp. from the Namak Lake is recorded as having the parasite Clinsotomum complanatum (Hosseinie, 1987). Aphanius from Damghan were infected with the trematode Tetracotyle sp. (Gholami et al., 2011). 

Kamal et al. (2007; 2009) and Bakhtiyari et al. (2011) give some life history data on the Damghan and Shour River of Eshtehard (west of Tehran) populations. Food in the Shour River was mostly chironomids while crustaceans including Daphnia were eaten at Damghan. Stable environmental conditions at Damghan allowed more prey selection and a higher condition factor. The Damghan fish lived longer and had a higher egg diameter while Shour River fish had a higher L and a shorter reproductive period, again probably reflecting environmental constraints.

Aphanius arakensis
Teimori, Esmaeili, Gholami, Zarei and Reichenbacher, 2012 

 

Common nameses

Kapour-e-dandandar-e-Arak.

[Arak tooth-carp].

Systematics

The holotype is a male, 31.5 mm SL from Arak, Namak Lake basin, 34º00’N, 49º50’E, in the Zoological Museum of Shiraz University, Collection of Biology Department (ZM-CBSU 10999), and paratypes are 35 males, 22.6-32.7 mm SL, and 35 females, 22.5-34.1 mm SL, same locality as holotype (ZM-CBSU  11000). The species is named for the city of Arak, the capital of Markazi Province..

Key characters

This species is distinguished by distribution and by pigmentation, males having 12-19, commonly 15-16, flank bars and females with prominent pigmentation along the mid-flank.

Morphology

Total dorsal fin rays 11-14, total anal fin rays 10-12, branched pectoral fin rays 14-18, branched pelvic fin rays 6-8, lateral series scales 27-32 (also given as 28-32), caudal peduncle scales 10-13, and total gill rakers 8-10. Lateral series scales and flank bar numbers are statistically different from other species. The otolith has a short but high antirostrum, a wide excisura and a ventral rim with some small, drop-like processes. Molecular evidence also defines the species with 19 apomorphies.

Sexual dimorphism

See colour description.

Colour

Lighter flank bars in males are narrow with the interspaces broader than the bars and becoming wider posteriorly. The head is grey on top and the body overall is dark from numerous melanophores although the belly is pale. The dorsal, anal and caudal fins have a white margin with some darker pigment proximally making a strong contrast. The first dorsal fin rays are dark and membranes may all be dark too in some fish. The anal fin also can have darkened membranes but this not as pronounced as in the dorsal fin. The bulk of the caudal fin is pale or whitish but membranes may be lightly pigmented. The pectoral and pelvic fins are yellowish. Most fish have dark blotches at the dorsal and anal fin bases. Females are greyish on the back with the belly and lower head surface light. The chin and sides of the head are speckled with melanophores. There is a line of relatively dark melanophores below the eye. All fins are white. The flanks have dark, irregular blotches or spots from the body middle to the caudal peduncle. Small falnk spots may line up along the mid-flank as a row from behind the head to the caudal fin base. A lozenge-shaped spot is found at the central caudal peduncle base.

Size

Attains 34.1 mm standard length.

Distribution

Zoogeography

This taxon is the sister-species of A. sophiae from the Kor River basin to the south.

Habitat

The type series was collected from a small, shallow polluted pond, 6 x 4 m, fed from a nearby spring. The shores had Juncus and Typha but there was no vegetation in the pond. The pond bottom was muddy with some gravel.  The water was almost stagnant, with a temperature of 23°C. This was the only fish species in the pond and the tooth-carp was also present in neighbouring springs.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

None at present in Iran but a possibly a species of interest to aquarists.

Conservation

This species is known from a restricted area and the type locality has anthropogenic pollution. It may well be threatened.

Further work

The distribution of this taxon in the Namak Lake basin should be explored further and threats to its existence assessed.

Sources

Based on Teimori et al. (2012).

Type material: See above.

Aphanius dispar
(Rüppell, 1829)


Male

Female, courtesy of Z. Gholami, H. R. Esmaeili and M. Ebrahimi
Female, courtesy of Z. Gholami, H. R. Esmaeili and M. Ebrahimi

Male, courtesy of J. E. Randall
Male, courtesy of J. E. Randall

Female, courtesy of J. E. Randall
Female, courtesy of J. E. Randall


Mehran River, courtesy of H. R. Esmaeili

Common names

gour-e khari, kopurdandandar-e balehbolband (= tooth-carp with striped fin), kapurdandan-e balmband, mahi dom parchami.

[harsun. batrikh or batrikh motakayer in Arabic; nambal in Pakistan; high-finned pupfish, Arabian killifish, mother of pearl fish].

Systematics

A detailed synonymy is given by Villwock et al., (1983), Wildekamp et al. (1986) and Wildekamp (1993). Hoedeman (1951) proposed a new genus for this species, Aphaniops, based on the absence of a dermal sheath or genital pouch around the anterior anal fin rays, only 8-9 dorsal fin rays in contrast to 10-14 rays in Aphanius and 7-8 pelvic fin rays in contrast 5-7 rays. It has not found general acceptance.

The type subspecies is found in all Iranian drainages (Krupp, 1983) although Aphanius dispar stoliczkanus (Day, 1872) has been recorded in earlier literature as the subspecies of these drainages except the Tigris River basin for which Aphanius dispar richardsoni (Boulenger, 1907) is reported (Berg, 1949). Aphanius dispar richardsoni is limited to the Dead Sea valley of Israel and western Jordan by Villwock et al. (1983) and Wildekamp (1993). Cyprinodon stoliczkanus Day, 1872 is regarded as a synonym of Aphanius dispar dispar by Krupp (1983) while Berg (1949) places Bampur River, Baluchestan fish in this subspecies as a southern representative of Aphanius dispar with 6-9, rarely 10 dorsal fin rays as opposed to 9-10 rays in Aphanius d. dispar.

Hrbek and Meyer (2003) note that, based on their mtDNA study, the monophyly of A. dispar is strongly rejected and it does not constitute a species in terms of the phylogenetic species concept. Reichenbacher et al. (2009) found that isolated populations in Saudi Arabia may be capable of evolving into new species in a short period of time, based on otolith morphology - evidently there is still much to be learned about this nominal species.

Hybrids with Aphanius dispar richardsoni are reported from Israel (Goren and Rychwalski, 1978).

The lectotype of Lebias dispar as designated by Villwock et al. (1983) is in the Senckenberg Museum Frankfurt under SMF 821 with paralectotypes SMF 1988 (10). Further paralectotypes in the Natural History Museum, London under BM(NH) 1860.11.9:152 listed as from Abyssinia (3 fish, 44.1-49.6 mm standard length) (Eschmeyer et al., 1996) but a label in the jar states that "status as types doubtful, types in SMF range from 26.1-35.8 mm SL, locality in description and types given as 'Red Sea'".

Types of Cyprinodon stoliczkanus Day, 1872 are in the Zoological Survey of India, Calcutta (ZSI 1477, ZSI 1478), the Natural History Museum, London (BM(NH) 1889.2.1.2065-74, originally 21 fish, now 14 fish, 14.4-29.4 mm standard length when examined in September 2007) and the Australian Museum, Sydney (AMS B.7730-7731) (Whitehead and Talwar, 1976; Eschmeyer et al., 1996; Ferraris et al., 2000). This species was originally described from a "stream at the village of Joorun, and also at Lodai, along the edge of the Rann" (= the salt water Rann of Cutch, Sind, India).

The lectotype of Cyprinodon richardsoni as designated by Krupp and Schneider (1989) is in the Natural History Museum, London under BM(NH) 1856.5.2:4 with paralectotypes under BM(NH) 1856.5.2:5 (8 fish) (12.9-25.0 mm standard length, the largest being the lectotype) (Eschmeyer et al., 1996; personal observations).

Krupp (1983) did not find the variation in teeth numbers between different populations and subspecies observed by Berg (1949). Keivany et al. (2011) examined 16 populations from southern Iran meristically and found significant differences in 10 of 11 characters but their wide overlaps prevented population differentiation. Teimori et al. (2012) used otolith morphology to show vicariance events in the Persian Gulf area during the last glacial maximum (21,000-18,000 B.P.), dispersal during the Early Holocene rise in sea level (11,000-8000 B.P.), and Holocene to Present interruption of gene flow at the Strait of Hormuz have shaped intraspecific differentiation of this species.

Key characters

The colour pattern is distinctive.

Morphology

Scales along the flank 24-35. Scales are squarish with an almost vertical anterior margin and protruding anterior corners, parallel dorsal and ventral margins and a rounded posterior margin with tiny teeth. The anterior margin has a small central protuberance with shallow indentations above and below or almost straight margins to each anterior corner. Radii are present on the anterior field, numbering 10-20 but mostly 14-16, and are almost horizontal and parallel. The exposed part of the scale has dimples rather than circuli. Circuli are few. The focus is subcentral posterior. There is no pelvic axillary scale. Total dorsal fin rays 7-11, usually 9-10, total anal fin rays 8-12, usually 10-11 (the number of unbranched rays in the dorsal and anal fins varies from 1-3), total pectoral rays 12-18 and total pelvic rays 6-7, usually 7. Vertebrae 24-29. Al-Hassan (1982b) reports on abnormalities in the vertebrae including loss of the posterior part of the centrum and fusions. Teeth are tricuspid with the central cusp concave at its tip and only slightly longer than the lateral cusps (Goren and Rychwalski, 1978), although in some fish seen by me the tip is rounded. There are 12-20 teeth per jaw (Krupp, 1983). Gill rakers 11-22, modally 13-16, reaching the second adjacent raker when appressed. Some gill raker counts are difficult to make accurately as those at the anterior arch end are minute and those at the dorsal end are partially concealed in flesh. The gut is coiled ?see my ginaonis paper. Reichenbacher et al. (2007, 2009) give descriptions of otolith morphology. Chromosomes number 2n=48 (Klinkhardt et al., 1995; Esmaeili et al., 2008). Karyotype 16Sm + 32St and arm number 32 (Esmaeili et al., 2008).).

Meristic values for Iranian specimens:- total dorsal fin rays 8(1), 9(39) or 10(10); total anal fin rays 9(2), 10(42) or 11(6); total pectoral fin rays 14(2), 15(16), 16(28), 17(3) or 18(1); total pelvic fin rays 6(6) or 7(44); lateral series scales 24(2), 25(5), 26(11), 27(17), 28(7), 29(2), 30(1), 31(2), 33(2) or 35(1); total gill rakers 15(13), 16(16), 17(16) or 18(3); and total vertebrae ?.

Sexual dimorphism

Males have longer fins than females and are more brightly coloured. The dorsal fin is twice as long in the male and reaches the caudal fin when appressed. When expanded it is widely flared and distinctive as is the enlarged anal fin.

Colour

Breeding males are brown-grey, grey or black-brown with iridescent blue-white flank spots and white and brown to light orange or light blue, irregular, narrow bars. The head has blue and orange tinges and in particular there is an orange spot on the operculum postero-dorsally. Lips are blue-white. The flank over the pectoral fin has electric blue spots. The anterior belly becomes blue with pearl spots. Scales have a dark margin. The dorsal fin is spotted light blue on a bright orange background and is barred. Barring may be irregular and in overall view in breeding males appears as speckling. Pectoral, anal and pelvic fins are lemon-yellow. The anal fin has barring on the posterior 4 rays in breeding males. The caudal fin has 2-3 dark and light blue alternating bars, the last bar being yellow. The bars are crescent-shaped, with concave side posterior. Males outside the breeding season are less brightly coloured with silvery on the flanks with a grey or black-brown back and irregular flank bars. Male specimens from the Bampur River basin in Iran have a body speckled silvery-white to light blue, weak tail bars, blue and orange tinges to the head, an orange spot on the operculum and an orange-yellow edge to the anal fin.

Young have pale brown flank bars on the caudal peduncle. Some young may have a few spots on the flank rather than bars. Females are brown-grey to silvery with 8-20 narrow flank bars and a dark-brown back. The flank bars are dark brown and may have a tinge of orange, the interspaces silvery. Flank bars may number as few as 6 or 7 (Talwar and Jhingran, 1991; Keivany and Ghorbani, 2012). Females also have the orange operculum spot and the blue spots over the pectoral fin. Female fins are hyaline. The peritoneum is brown to black.

Ross (1980) notes that Arabian specimens are brighter in waters with algae and plants; those found over sand and mud are duller, an effective camouflage. He also records variant colour patterns with blue tails, or an overall gold colour, or with green or turquoise flank spots.

Size

Reaches 8.0 cm, but this may be an error as Krupp (1983) points out. He found fish only to 5.2 cm standard length although in Carpenter et al. (1997) 8 cm is given as a maximum. Wildekamp (1993) records fish to 7.0 cm for males and 6.5 cm for females in total length. Fish from Ain Al Adhari, Bahrein reach 7.3 cm total length. Generally fish in fresh water are smaller than marine specimens.

Distribution

Found from Egypt, eastern Sudan and Ethiopia, and in the Red Sea through southern Southwest Asia and east to Rajasthan. Also in the eastern Mediterranean Sea (Kornfield and Nevo, 1976). The subspecies Aphanius dispar richardsoni is found in the Dead Sea rift valley.

A specimen in the Naturhistorisches Museum Wien from the hot spring at Ginao (NMW 13799) may be mislabelled as several other collections from this hot spring are all clearly A. ginaonis.

This species is found in Iranian fresh waters from the Tigris basin to the Makran basin in water bodies draining to the Persian Gulf and Sea of Oman, islands in the Gulf such as Qeshm (OSU 8120), and in the endorheic Hamun-e Jaz Murian basin. It is reported from the Shapur and Dalaki rivers (Gh. Izadpanahi, pers. comm., 1995;Bibak et al., 2012), Gudar River east of Anveh, Mehran River below Bastak, Sarageh River north of Bandar Abbas, Galashkerd Lake near Minab, and the Kol. Shur, Hasan Langi and Ghale-gazii rivers in Hormozdgan. Abdoli (2000) illustrates distributions in the Arvand, lower Zohreh (= Hendijan), lower Hilleh, Mand, Kul, Hasan Langi and Minab rivers, lower reaches of rivers in the Makran basin from the Jagin to Bahu Kalat, the lower Halil and Bampur Rivers in the Hamun-e Jaz Murian basin, and the endorheic Mashkid River. Keivany and Ghorbani (2012) map distribution from Khuzestan to Baluchestan in waters draining to the Persian Gulf, as well as the Hamun-e Jaz Murian, Mashkid and southern Lut internal basins. They also document a record more remote from these distributions, 50 km northwest of Ilam, in the Sartang-e Bijar River of the Tigris River basin (33°41'24"N, 45°51'48"E).

Zoogeography

Villwock et al. (1983) contend that an ancestor of A. dispar was present in the middle Miocene Transgression connecting the Mediterranean Sea to the Indian Ocean. The mesohaline conditions, which developed in the Mesopotamian basin, were a good prerequisite for the development of A. dispar since, apart from some euryhaline species, there would be no competitors. A. dispar spread from Mesopotamia through the Persian Gulf to India and the Red Sea. During the Pleistocene, it colonised inland waters.

Habitat

Highly tolerant of salinity, this species is found from the sea to fresh water habitats, including landlocked basins. Slightly saline waters in aquaria seem to discourage parasites (Andrews, 1983). Salinities as high as 145‰ are tolerated (Lotan, 1971) and this fish can be moved abruptly from fresh to hypersaline water. Adaptation to high salinity waters involves a reduced osmotic permeability of the gills and an increase in transportation of sodium chloride by the intestine (Skadhauge and Lotan, (1974). Lotan (1969, 1971, 1973) and Skadhauge and Lotan (1974) describe sodium, chloride and water balance, and osmoregulation in this species. Plaut (1999) found that salinities above ~250% seawater seem to decrease swimming capabilities and routine activity rate and a decrease in resting metabolic rate indicated stress. This species has been reported to survive in waters 38.4°C in Oman and only began to die at 45.1-46.0°C (Haas, 1982). Fish survived up to 40°C in aquaria without acclimation (Lewis, 1970). Populations survive in isolated sections of streams with little food because of histological changes in the stomach (Ba-Omar et al. (1998). Aphanius dispar occurs in shallow water and among vegetation over sand, rock or soft detritus bottoms.

Frenkel (1995) describes the effects of environmental factors on growth and reproduction in this species in Israel. Frenkel and Goren (1997) showed fish to be tolerant of a wide range of environmental variables as measured against the gonado-somatic index (GSI) and ovary maturation stages. The GSI was not affected by temperature over a range of 18 to 37°C while maturation increased from 18 to 27°C but remained the same between 27 and 37°C. For salinities between 0‰ and 56‰ differences were only found in mean GSIs at the two extremes and at 0‰ ovaries contained only primordial germ cells. GSIs and maturation stages only differed in feeding experiments when fish were deprived of food or fed at equal or greater than 1% of body weight per day. A decrease in photoperiod from 14L:10D (14 hours light, 10 hours dark) to 10L:14D caused a decrease in oocyte maturation stages and ovaries were no longer suitable for spawning. In aquaria, a pH of 7.5-8.5, temperatures above 27°C and water hardness greater than 200 p.p.m. and the addition 4 heaped tablespoons of kosher salt per ten gallons of water (38 litres) favoured reproduction (Mackowiak, 1988). Alkahem (1989) found 50% mortality in 96 hours when fish were exposed to pH levels of 3.5-3.9.

Feulner (1998) reports that United Arab Emirate fish "hover" in the water column with the tail slightly curved to one side, presumably a feeding behaviour. Males and females in aquaria swam in separate schools (Al-Daham et al., 1977).

Age and growth

Sexual maturity in ideal conditions may be attained as early as 7 months but full size takes a year. The length-weight relationship in southern Iraq was W = 1.2301 x 10-4 L2.4544 for males and W = 2.8357 x 10-6 L3.5112 for females. The average condition factor for males was 1.8900 and for females 1.6952. For a given length group males are heavier than females and condition factors are higher for males (Huq et al. 1977). Muhsin (1987) studied the effect of differing food supply on the weight and chemical composition in females of this species taken from the Al-Asafiah River, Basrah, Iraq but examined in tanks. Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 41 Iranian fish measuring 2.37-3.56 cm total length. The a-value was 0.0012 and the b-value 3.214 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases). Alavi-Yeganeh et al. (2011?check published paper date) give length-weight data on this species from two localities in Iran with a (intercept) being 0.0108 and b (slope) being 3.332 for all localities and sexes combined.

Food

Iranian specimens have guts packed with filamentous algae and detrital remains. Al-Daham et al. (1977) found food in southern Iraq to be predominately filamentous algae and diatoms, with some copepods, rotifers and, in one instance, a small winged insect. In contrast, Shafi and Shalli (1986) report a diet of beetles, ephemeropteran nymphs, rotifers, filamentous algae and plants in southern Iraq. Younis et al. (2001b) found Shatt al Arab, Iraq fish fed on amphipods and copepods (29.9%), organic detritus (11.6%), and algae and diatoms (7.1%). In aquaria, Al-Daham et al. (1977) found this species to prefer animal foods in contrast to Aphanius mento which preferred plant foods. Al-Akel et al. (1987) have observed the selective feeding behaviour of this species at Al-Khurj, Saudi Arabia. Certain species of filamentous algae are selected over others, e.g. the blue-green alga Oscillatoria, the green algae Ulothrix, Spirogyra and Crucigenia, the desmid Cosmarium, and the diatoms Cocconeis, Diatoma, Navicula and Rhoicosphania, and algal spores and zygotes. Haas (1982) observed this species in Oman to pick at rocks and other substrates, take items at the surface, chase small fish and eat insect larvae. Gut contents are mainly desmids and diatoms, some filamentous algae and rarely insect larvae and one snail. This species is an omnivorous surface feeder (Carpenter et al., 1997).

Reproduction

A female in spawning condition approached a male and appeared to feed on a plant or plant-encrusted rock in fish studied by Haas (1982) in Oman. Feulner (1998) reports fluttering of the tail by male fish. The male places his chin on her nape and 2-3 eggs are spawned rapidly. The female presses her vent against the plant or rock vertical surface, the male arches his back into an s-shape and presses his body and anal fin against hers and clasps her with his dorsal fin. Eggs are released singly with a quiver. The time elapsed from chin on nape to egg release is as short as 1-2 seconds. Reproduction occurs throughout the day and throughout the year in Oman with 69% of females having ripe eggs (up to 41) in April-June. Peak spawning in southern Iraq is April to June when only a small proportion of eggs in the single ovary are fully developed eggs (Al-Daham et al., 1977). Shafi and Shalli (1986) record up to 73 mature eggs in southern Iraq specimens with a diameter of 220 microns. Breeding season was May-July. In Saudi Arabia mature and developing oocytes are observed in fish during the whole year. The egg cycle may be more than one year with spawning in March and April (El-Hawawi and Al-Imam, 1984). On the Mediterranean coast of Egypt, spawning occurs from March to September with a peak in July and August. Maximum egg diameter is 2.2 mm and size at maturity for females is 30 mm total length (Lotan and Ben-Tuvia, 1996). Iranian specimens have large eggs (2.0 mm) on 16 March but young fish (8.9 mm standard length) were caught on 26 November suggesting reproduction is almost year round.

Bibak et al. (2012) examined reproduction in the Dalaki River, Bushehr and found females to predominate (390 fish versus 140), males had a maximum length of 49 mm and females 39 mm, and gonadosomatic indices indicated a spawning period of April to June, with peaks in each of these months of 6.449 and 6.632.

Spawning areas have some water flow and vertical surfaces of plants or algae-encrusted rocks. Males defend a territory about 30 cm wide against other males and will display with erect fins if approached. Parallel swimming, head to head confrontations, rapid circling, and flank bites were observed by Haas (1982). Two males will swim and chase each other in circles until one swims away with fins depressed. Males will change their territory after one day defending it. In Israel, males of Aphanius dispar richardsoni do not show any territorial behaviour (Goren and Rychwalski, 1978). This fish will spawn in a few months of birth in aquaria (Ross, 1987). Three pairs of fish yielded about 40 eggs per day in aquaria and these eggs hatched in two weeks at 27°C (Mackowiak, 1988).

Parasites and predators

This species is reported to be heavily infested with metacercariae of the trematode Clinostomum complanatum in eastern Saudi Arabia, up to 86.6% incidence with up to 41 metacercariae per fish, the infestation being higher in females than males (Kalantan et al., 1987). Gholami et al. (2011) record the trematode Clinostomum complanatum from fish in the Mehran River near Rudan and Gholami et al. (2011) report the cestode Ligula intestinalis in fish from the Mehran River near Bastak.

Economic importance

Al-Akel et al. (1987) suggest the use of this species as a control agent for filamentous algae.

Steven (1913) reported that this fish was responsible for the absence of mosquito larvae from streams in Karachi which otherwise appeared to be good breeding places. Christophers and Shortt (1920-1921) thought that tooth-carps in southern Mesopotamia were a significant factor in lowering the malaria rate, presumably mostly this species. Sharma and Al-Daham (1979) compared the efficiency of this tooth-carp with the mosquitofish for mosquito control under experimental conditions. The tooth-carp is more efficient when males and females were together than mosquitofish since males of the latter expend time and energy defending territory. Mosquitofish are more active in pursuit of mosquito larvae than the tooth-carp when filamentous algae was present. Cloudy conditions and lower temperatures slow consumption and both species prefer pupae to larvae of mosquitos, with females taking more pupae than males. The tooth-carp could consume nearly twice the amount of mosquitos than the mosquitofish over a given period of time. Overall the tooth-carp compares favourably with the mosquitofish as a destroyer of the malaria-carrying mosquito. Homski et al. (1994) under laboratory conditions found that Aphanius dispar is more successful than the mosquitofish in preying on the third and fourth instars and on pupal stages of mosquitos while the mosquitofish, with a smaller relative mouth size, is more successful on the first two instars. Small fish of both species eat first instars exclusively. However, Aphanius dispar eats more second instar larvae under a vegetation cover since the larger Aphanius are more capable of penetrating shallow water than mosquitofish. Aphanius dispar occurs in shallower water and among more vegetation than the open water mosquitofish. The two species can be used to complement each other in control of mosquitos.

Ataur-Rahim (1981) found that this species will eat mosquito larvae in water storage tanks in Riyadh, Saudi Arabia. Fifteen fish consumed all larvae at a mean density of 5/100 sq cm water surface area in a tank 3 by 2 m in a day. One fish, 5.2 cm long, ate 61 mosquito larvae. Haas (1982) fed this species mosquito larvae in the presence of abundant algae as an alternative food. An average of 96 larvae are eaten each day. Etemadfar et al. (1983) studied the potential use of this species for malaria control in southern Fars, Bushehr and Hormozgan provinces. Louis et al. (1988) detail a control strategy against malaria using this species in Djibouti as do Fletcher et al. (1992) in Ethiopia and Fareed and Baban (1995) in Yemen. Fletcher et al. (1992) showed a decrease in sites harbouring mosquito larvae from 34% without the fish to 1.6% with the fish stocked. Chandra et al. (2008) briefly reviews the use of this species in biocontrol.

The studies of Frenkel and Goren (1997) in Israel show that the reproduction of this species can be controlled such that it has potential for mass production and could be used in mosquito control. Carpenter et al. (1997) state that this species is used in mosquito control. Ross (1987) gives details of aquarium care and breeding of this species.

Conservation

Al-Johany and Yousuf (1993) show that Gambusia affinis has a wider range of temperature tolerance than this species and is better suited for a desert habitat in Saudi Arabia. The exotic Gambusia is therefore a threat to A. dispar. Al-Kaheh-Al-Balawi et al. (2008) found that Saudi Arabian populations are declining from reduced availability of food, habitat degradation, chemical contamination, introduction of exotics and exploitation and some of these factors doubtless obtain in Iran.

Further work

This widespread species has been reported from a number of springs in southern Iran and, like A. ginaonis, may be distinct at some level. Molecular work would be useful in determining the taxa involved and evolution in hot springs. The utility of this species in mosquito control in smaller water bodies of Iran where it also occurs naturally could be explored. Al-Daham et al. (1977) found that this species in aquaria ate mosquito larvae "eagerly" compared to "very eagerly" for Gambusia holbrooki and "not eaten" for Aphanius mento.

Sources

Type material:

Iranian material: NMW 13799, 1, ?; CMNFI 1979-0129, 1, 31.3 mm standard length, Fars, spring about 2 km north of Farrashband (28º54'N, 52º04'E); CMNFI 1979-0137, 106, 8.9-37.7 mm standard length, Fars, stream outside Lar (27º40'N, 54º32'30"E); CMNFI 1979-0138, 1, ? mm standard length, Fars-Hormozgan border, Rasul River drainage (ca. 27º32'N, ca. 54º58'30"E); CMNFI 1979-0139, 2, ? mm standard length, Fars-Hormozgan border, Rasul River drainage (ca. 27º25'30"N, ca. 54º59'E); CMNFI 1979-0140, 3, ? mm standard length, Hormozgan, Kul River drainage (27º14'N, 55º46'30"E); CMNFI 1979-0141, 6, ? mm standard length, Hormozgan, Kul River at road bridge (27º17'30"N, 56º03'30"E); CMNFI 1979-0142, 10, ? mm standard length, Hormozgan, Baghu River at Baghu (27º17'N, 56º28'E); CMNFI 1979-0143, 118, ? mm standard length, Hormozgan, Hasan Langi River drainage (27º21'N, 56º50'30"E); CMNFI 1979-0145, 1, ? mm standard length, Hormozgan, Geru River (26º55'N, 57º01'30"E); CMNFI 1979-0148, 21, ? mm standard length, Hormozgan, Sarzeh River (27º30'30"N, 56º15'30"E); CMNFI 1979-0151, 45, ? mm standard length, Hormozgan, Shur River drainage near Kahkom (28º02'N, 55º53'E); CMNFI 1979-0177, 37, ? mm standard length, Hormozgan, Sarzeh River drainage (27º33'N, 56º14'E); CMNFI 1979-0179, 13, ? mm standard length, Hormozgan, Sarzeh River drainage (27º36'N, 56º15'E); CMNFI 1979-0181, 9, ? mm standard length, Hormozgan, Kul River drainage (27º17'30"N, 56º03'30"E); CMNFI 1979-0182, 31, ? mm standard length, Hormozgan, stream on road to Bandar-e Lengeh (27º14'N, 55º46'30"E); CMNFI 1979-0185, 24, ? mm standard length, Hormizgan, stream in Rasul Rievr drainage (27º06'N, 55º45'E); CMNFI 1979-0187, 2, 26.2-33.2 mm standard length, Hormozgan, Sar Khun oasis (27º23'30"N, 56º26'E); CMNFI 1979-0312, 7, ? mm standard length, Baluchestan, dam on Bampur River (27º11'N, 60º36'E); CMNFI 1979-0313, 1, ? mm standard length, Baluchestan, Bampur River at Bangharabad (27º20'N, 60º46'E); CMNFI 1979-0328, 18, ? mm standard length, Baluchestan, jube 2 km south of Bampur River near Bampur (27º10'30"N, 60º21'E); CMNFI 1979-0352, 2, ? mm standard length, Khuzestan, marsh in Jarrahi River drainage (30º33'30"N, 48º48'E); CMNFI 1979-0354, 8, ? mm standard length, Khuzestan, Karun River tributary (30º31'N, 48º19'E); CMNFI 1979-0355, 1, ? mm standard length, Khuzestan, Karun River tributary (30º35'N, 48º22'E); CMNFI 1979-0362, 2, ? mm standard length, Khuzestan, jube in Karkheh River drainage (31º42'N, 48º33'E); CMNFI 1979-0379, 4, ? mm standard length, Khuzestan, Dez River (32º12'N, 48º27'E); CMNFI 1979-0381, 1, ? mm standard length, Khuzestan, stream west of Shushtar (ca. 32º10'N, ca. 48º35'E); CMNFI 1979-0384, 7, ? mm standard length, Khuzestan, Ab-e Shur drainage (32º00'N, 49º07'E); CMNFI 1979-0403, 25, check size range ?19.7-37.0 mm standard length, Bushehr, stream 23 km south of Kaki (ca. 28º11'N, ca. 51º43'E); CMNFI 1979-0406, 48, 15.6-35.8 mm standard length, Hormozgan, stream near Bandar-e Charak (26º48'N, 54º18'E); CMNFI 1979-0407, 3, ? mm standard length, Hormozgan, stream 30 km from Barvedun (26º51'N, 54º37'E); CMNFI 1979-0408, 106, 14.6-48.4 mm standard length, Hormozgan, Mehran River (27º04'N, 54º35'E); CMNFI 1979-0410, 10, ? mm standard length, Hormozgan, Mehran River (26º53'N, 55º17'E); CMNFI 1979-0414, 12, ? mm standard length, Hormozgan, pool on road to Tiab (27º05'N, 57º02'E); CMNFI 1979-0415, 1, ? mm standard length, Hormozgan, stream 18 km south of Genu (27º17'30"N, 56º20'E); CMNFI 2007-0053, 4, ? mm standard length, Hormozgan, Sarzeh River (ca. 27º36'N, ca. 56º15'E); CMNFI 2007-0057, 16, ? mm standard length, Hormozgan, Mehran River 4 km below Bastak (ca. 27º05'N, ca. 54º05'E); CMNFI 2007-0058, 3, ? mm standard length, Fars, headwaters of Gowdar River (ca. 27º24'N, ca. 54º16'E); OSU 8129, 1, 54.5 mm standard length, Hormozgan, Qeshm Island salt spring (no other locality data); BWC95-30, uncatalogued, 3, ?, Khuzestan, Kupal (31º15'N, 49º10'E); BWC98-6, uncatalogued, 24, ?, Hormozgan, Ab-e Garm-e Khamir (ca. 26º59'N, ca. 55º35'E); BWC2000-6, uncatalogued, ?, ?, Khuzestan, Dez River (32º14.663'N, 48º20.112'E); BWC2000-15, uncatalogued, ?, ?, Khuzestan, stream south of Shushtar (31º41.867'N, 48º59.581'E).

Comparative material:- BM(NH) 1920.3.3:193-202, 12, 21.0-30.5 mm standard length, Iraq, Basra (30º30'N, 47º47'E); BM(NH) 1920.3.6:9-10, 3, 17.1-26.8 mm standard length, Iraq, Basra (30º30'N, 47º47'E); BM(NH) 1949.7.21:1-11, 12, 18.4-42.4 mm standard length, Iraq, Bahr-el-Milk (32º35'N, 43º50'E); BM(NH) 1974.5.22:30, 1, 35.2 mm standard length, Iraq, Basra (30º30'N, 47º47'E); BM(NH) 1974.5.22:31-32, 2, 37.4-44.4 mm standard length, Iraq, Al Faw ().

Aphanius farsicus
Teimori, Esmaeili and Reichenbacher, 2011


Male


Female


Cyprinodon persicus male after Jenkins (1910)
Cyprinodon persicus male after Jenkins (1910)

Male, courtesy of Z. Gholami, A. Gholamifard and R. Zarei
Male, courtesy of Z. Gholami, A. Gholamifard and R. Zarei

Female, courtesy of Z. Gholami, A. Gholamifard and R. Zarei
Female, courtesy of Z. Gholami, A. Gholamifard and R. Zarei

Common nameses

Kapour-e dandandar-e Farsi (Farsi tooth-carp (sic), correctly Fars tooth-carp as it was named for the province not the language), gour-e khar, kopurdandan-e Irani (= Iranian tooth-carp as persicus).

[Persian pupfish or toothcarp, but see also A. sophiae].

Systematics

Cyprinodon blanfordii Jenkins, 1910 is potentially a synonym by authors (see Coad, 1996a). Hanko (1924) placed Cyprinodon blanfordi and C. persicus in Cyprinodon chantrei Gaillard, 1895 but this is an error.

Jenkins (1910) described Cyprinodon blanfordii ("East of Shiraz, South Persia") and C. persicus ("Spring on the edge of Shiraz Lake, Southern Persia"). The first nominal species has so generalised a locality as to be of very uncertain provenance. It could conceivably be in the Shiraz valley, the Kor River valley or outside either of these (see below under A. sophiae where it is argued that is is a synonym of this species). The second nominal species is clearly from the valley of Shiraz and its salt lake Maharlu, an endorheic basin.

The valid name for Lake Maharlu basin Aphanius was given previously as persicus in this website. Gaudant (2011) showed that Brachylebias Priem, 1908 is a junior synonym of Aphanius Nardo, 1827. Teimori et al. (2011) show that Brachylebias persicus Priem, 1908, a Late Miocene species from northwest Iran, is distinct from the modern species and introduce A. farsicus as a replacement name for A. persicus (Jenkins, 1910).

The Jenkins types in Calcutta have not been examined by me. Cyprinodon blanfordii has page priority but the locality is vague and the fish have a pigment pattern description and illustration which is spotted on the flank with a lozenge-shaped spot at the caudal fin base, thus appearing to be a female A. sophiae.

Material described and illustrated by Villwock (1960) as A. sophiae are this species.

Three syntypes of Cyprinodon blanfordi (ZSI F9416 to ZSI F9418), two syntypes of Cyprinodon persicus (ZSI F9403 and ZSI F9404) are in the Zoological Survey of India, Calcutta (Menon and Yazdani, 1968).

Key characters

Females of this species are barred where all other Aphanius species in this area of southern Iran have spotted females. The distribution is apparently limited to the Lake Maharlu basin near Shiraz.

Morphology

Although populations are isolated in springs and streams around Lake Maharlu, Coad (1996a) found them to be relatively homogenous.

Lateral line scales 24(3), 25(34), 26(69), 27(176), 28(100) or 29(22). Scales above the lateral line 4-6, scales between lateral line and the anal fin 4-8, scales between the lateral line and the pelvic fin 5-9, and scales around the caudal peduncle 12-17. Esmaeili et al. (2007) determined a chromosome number of 2n=48 with 11 pairs of submetacentric and 13 pairs of subtelocentric chromosomes and an arm number of NF=70. Karyotype 16Sm + 32St and arm number 32 (Esmaeili et al., 2008). Total dorsal fin rays 8(1), 10(8), 11(97), 12(204), 13(89), or 14(7); total anal fin rays 9(3), 10(111), 11(225), 12(64) or 13(3); total pectoral fin rays 13(9), 14(82), 15(211), 16(94), 17(9) or 18(1); total pelvic fin rays 4(11), 5(181) or 6(213); total gill rakers 9(5), 10(51), 11(220), 12(108), 13(19) or 14(3); abdominal vertebrae 9(1), 10(1), 11(27), 12(355), or 13(22); and caudal vertebrae 12(1), 14(16), 15(285) or 16(104).

Sexual dimorphism

Most apparent in colour and pigmentation detailed below. Females are longer and heavier than males (Esmaeili and Shiva, 2006).

Colour

"The basic colouration is bright shining silvery white. The entire body, with the exception of the dorsal crest and the belly, is tigered with small brown spots; in some specimens these spots seem to be arranged in three irregular longitudinal rows of which the uppermost extends as a band across the operculum and to the eyes; all fins are uniformly yellow with a reddish border. In males the spots are more dense, in females they are paler and scarcer, only the 2-3 at the base of the caudal fin are dark brown" (Heckel (1847b) on A. sophiae). "The colour of this small fish with its enviable crystal teeth is uniformly brown, at least it is now in ethyl alcohol, with a silvery-white throat and belly; laterally on the tail there are a few scattered, darker spots, and a deeper black spot is located on the last scales anteriorly of the caudal fin. All fins are blackish without any markings" (Heckel (1847b) on A. crystallodon). "Silvery, brownish on back, brown on top of the head, with numerous small spots of brownish, arranged in three or more, more or less irregular, longitudinal series, the median of which, on the lateral line, commonly ends in a black spot on the middle base of the caudal. On some this median series is entirely composed of blackish spots. Rarely the spots are somewhat confluent into longitudinal bands. Brownish specimens show a longitudinal band of silvery along the middle of the flank. The form described as A. crystallodon has spots only towards the base of the caudal, around the black spot usually ending the median series" (Garman, 1895).

The flank in females bears numerous alternating light and dark bars, the light bars varying in width from about one half to twice that of dark bars. The bars gradually merge with background pigmentation anteriorly on the flank and while clearly defined on the rear flank are difficult to distinguish anteriorly.

The caudal fin spot in females can be oval, tear-drop shaped or elongate but is usually in the form of a lozenge. Occasionally, single, smaller, dark, subsidiary spots may be found antero-dorsally and antero-ventrally to the basal spot or scattered spots may be found irregularly before and behind the basal spot.

Males have a pigmentation very similar to that of A. sophiae and the description here is identical. Some minor observed variation is attributed to variation in size and maturity of the fish compared. Males have light flank bars half the width or much narrower than alternating dark bars. The margins of the dorsal, anal and caudal fins are clear while the rest of these fins is dark. Some fish have up to 3, but usually 2, thin, light bars on the basal half of the caudal fin; these are generally larger fish. The margin of the lower half of the pectoral fin has concentrations of pigment on the membranes such that this area is darker than the rest of the fin. The anal fin is darkest posteriorly where pigment is concentrated on membranes. The distal third of the fin is pigmented to form a dark band, becoming lighter proximally. The dorsal fin is the darkest fin (except for the clear margin) and the anterior base is the darkest part of the fin. Bands are not always evident but pigment spots are large proximally. Some fish have 2, sometimes 3, thin light bars at the base separated by thin dark bars and paralleling the body profile while others have none. The dorsal fin base may have instead a series of lighter spots, sometimes irregular and not paralleling the body profile.

Size

Reaches 59.5 mm standard length (Esmaeili and Shiva, 2006).

Distribution

This species is restricted to the Lake Maharlu basin near Shiraz.

Zoogeography

This species is found only within the endorheic Lake Maharlu basin, Coad (1996a) suggests that tooth-carps in inland waters may have risen with the post-Pliocene uplift of the Zagros Mountains rather than being the result of relatively recent inter-basin dispersal.

Habitat

This species is found in fresh streams and springs and in springs of varying saline content or saline influence from hypersaline chloride Lake Maharlu. The lake at 124‰ is an impassable barrier to dispersal but is very shallow and dries out periodically, e.g. in 1967 (Cornwallis, 1968a). The larger springs and their streams can then meet and transfer fishes on the dried-out lake bed. Some springs are 27 km apart along the lake margin and have probably had no contact or fish exchange over many generations. This is especially the case for the smaller springs which emerge from the ground with a diameter of only 1 m and restricted flow. When lake water rises, lower springs are inundated with hypersaline water and the fishes are killed. The spring is presumably recolonised from a higher spring when lake level falls.

Fish placed in pure lake water die within minutes. Springs discharging directly into the lake contain tooth-carps but the fish do not venture into the lake. Some fish, if disturbed or deliberately harassed, will swim out for a short distance into the hypersaline lake water but they blanch and rapidly retreat to fresher water (Coad, 1996a).

Esmaeili and Shiva (2006) found the bottom of these water bodies to be muddy but the water was clear and slow-running. Conditions in October at three different sites were 16.9-19.0ºC, pH 6.70-6.74, dissolved oxygen 3.96-6.11 mg/l, nitrate 0.9-1.6 mg/l, nitrite 0.029-0.062 mg/l, phosphate 0.35-0.65 mg/l and ammonium 1.55-2.60 mg/l.

Age and growth

In the studies of Esmaeili and Shiva (2006), the sex ratio was 1.67:1 for females:males, highly significant, the age groups were 0+ to 3+ years, and positive allometric growth was determined (b value significantly greater than 3).The condition factor for females was highest in February and March and lowest in September while in males it was lowest in December and increased until February. Fishes smaller than 25 mm had an equal sex ratio, suggesting selective predation on males, or better survival or longevity of females. Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 62 fish measuring 1.86-4.27 cm standard length. The a-value was 0.0222 and the b-value 3.395 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases). Alavi-Yeganeh et al. (2011?check published paper date) give length-weight data on this species from Kaftarak spring with a (intercept) being 0.0097and b (slope) being 3.303 for sexes combined.

Food

Unknown but presumably similar to other tooth-carps.

Reproduction

Esmaeili and Shiva (2006) found the gonadosomatic index in females to increase from November to June, decreasing slowly from late June to November. The reproductive period lasted six months. Males had two peaks, in April and August. Egg diameters reached 1.71 mm, absolute fecundity ranged from 45 to 250 eggs (average 115.7 eggs) and relative fecundity was 21.6 to 244.1 with a mean of 90.01 per gram body weight. Even small fish had hydrated eggs and, with the extended reproductive period, shows adaptation to an unstable habitat. Monsefi et al. (2009) found that this species is a batch spawner in the Barm-e Shur spring with a spawning period from April to November. This species lives in unstable environments such as temporary lagoons or very small pools and batch spawning of relatively large eggs gives a greater chance of survival.

Parasites and predators

Mokhayer (1989) reported metacercariae of the eye fluke, Diplostomum spathaceum from Aphanius sophiae, probably this species, in Iran. The fluke can cause complete blindness and death in commercially important species of fish. This fish was also infested with yellow grub, Clinostomum complanatum. González-Solís et al. (1997) report Contracaecum sp. larvae from this species in the Lake Maharlu drainage, Fars.

Economic importance

This species has no current economic importance although it would make an excellent aquarium fish. Villwock (1959) gives details of maintaining what seems to be this species under aquarium conditions, gradually transferring them from a mixture of salty and magnesium sulphate water to stagnant tap water. They prefer some admixture of sodium chloride and magnesium sulphate which matches their natural habitat (2% synthetic sea water plus a tablespoon of epsom salts per 25 litres water). Aquatic plants such as Myriophyllum and Cabomba are also necessary for breeding as the eggs are attached to them but artificial substitutes can be used. Yellowish transparent eggs are 1.5 mm and are deposited usually individually, rarely in small groups. Larvae feed on Artemia salina nauplii and adults on daphnia, red mosquito larvae and tubifex. Male-male interactions include mock fights with spread fins and close swimming. The loser hides in the algae and is not strongly pursued by the victor. These interactions, and those between sexes, are not strongly developed and aquaria can be stocked at 15-20 fish per 25 litres. Y. Keivany notes (pers. comm., 2004) that this species is difficult to maintain in aquaria.

Conservation

The populations are found in a variety of springs and streams around Lake Maharlu. Some of these localities are threatened by water abstraction, road construction and pollution but a number of localities are small and/or saline and not under threat.

Further work

Studies on aquaria maintenance would enable it to become established in the aquarium trade and help its conservation.

Sources

Type material: None seen.

Iranian material: CMNFI 1979-0017, ?47, ?17.9-35.5 mm standard length, Fars, stream at Pol-e Fasa (29º29'N, 52º38'30"E); CMNFI 1979-0018, 75, ? mm standard length, Fars, Barm-e Shur (29º28'N, 52º41'30"E); CMNFI 1979-0033, 1, ? mm standard length, Fars, Barm-e Shur (29º28'N, 52º41'30"E); CMNFI 1979-0034, ?124, 14.8-38.7 mm standard length, Fars, spring on shore of Lake Maharlu (29º27'N, 52º44'E); CMNFI 1979-0035, 17, ? mm standard length, Fars, spring south of Barm-e Shur (29º28'N, 52º41'30"E); CMNFI 1979-0038, 16, ? mm standard length, Fars, spring near Barm-e Shur (29º28'N, 52º41'30"E); CMNFI 1979-0039, 91, 15.9-37.9 mm standard length, Fars, spring on shore of Lake Maharlu (29º27'N, 52º44'E); CMNFI 1979-0041, 61, 20.8-41.6 mm standard length, Fars, spring on shore of Lake Maharlu (29º23'N, 52º48'E); CMNFI 1979-0042, 28, ? mm standard length, Fars, spring on shore of Lake Maharlu (29º23'N, 52º48'E); CMNFI 1979-0046, 5, 18.1-32.2 mm standard length, Fars, qanat at Barm-e Dalak (ca. 29º35'N, ca. 52º38'E); CMNFI 1979-0047, 41, 11.3-32.1 mm standard length, Fars, spring source of Ab-e Paravan marshes (ca. 29º34'N, ca. 52º42'E); CMNFI 1979-0048, ?45, ?15.3-39.9 mm standard length, Fars, spring on shore of Lake Maharlu (ca. 29º32'N, ca. 52º48'E); CMNFI 1979-0049, 12, 17.2-38.5 mm standard length, Fars, spring on shore of Lake Maharlu (ca. 29º32'N, ca. 52º48'E); CMNFI 1979-0050, 27, 18.3-28.9 mm standard length, Fars, spring on shore of Lake Maharlu (ca. 29º31'30"N, ca. 52º49'30"E); CMNFI 1979-0051, 3, ? mm standard length, Fars, spring on shore of Lake Maharlu (ca. 29º30'N, ca. 52º52'E); CMNFI 1979-0052, 16, ? mm standard length, Fars, spring on shore of Lake Maharlu (ca. 29º30'N, ca. 52º52'E); CMNFI 1979-0064, 10, ? mm standard length, Fars, spring on shore of Lake Maharlu (ca. 29º31'30'N, ca. 52º49'30"E); CMNFI 1979-0065, 8, ? mm standard length, Fars, spring on shore of Lake Maharlu (ca. 29º31'N, ca. 52º50'E); CMNFI 1979-0066, ?31, 15.6-42.9 mm standard length, Fars, spring on shore of Lake Maharlu (ca. 29º32'N, ca. 52º48'E); CMNFI 1979-0110, 2, ? mm standard length, Fars, lake in Park-e Shahr, Shiraz (29º38'N, 52º32'E); CMNFI 1979-0112, 24, 11.9-45.9 mm standard length, Fars, stream at Pol-e Fasa (29º29'N, 52º38'30"E); CMNFI 1979-0118, 12, ? mm standard length, Fars, Barm-e Shur (29º28'N, 52º41'30"E); CMNFI 1979-0119, 75, ? mm standard length, Fars, steram at Pol-e Fasa (29º29'N, 52º38'30"E); CMNFI 2004-0006 (GenBank AY593484, AY593493), 15, ?22.2-41.8 mm standard length, Fars, Nasrabad Spring near Imamzadeh Ibrahim (29º35'09"N, 52º39'08"E);

Aphanius ginaonis
(Holly, 1929)


Male

Common names

gour-e khar, kopurdandan-e Genu or kapurdandan-e Geno (= Genow tooth-carp).

[Holly's pupfish, Genow pupfish, Gheno pupfish].

Systematicscs

Cyprinodon ginaonis was originally described from "Hei$e Quelle vom Djebel Ginao nordlich von Bender Abbas, südöstliches Persien". This is the Ab Garm-e Ganow at 27°26-28'N, 56°18-20'E, north of the Iranian port of Bandar Abbas at the Straits of Hormuz.

Berg (1949) places this species in the synonymy of Aphanius dispar stoliczkanus (see A. dispar above) and Villwock et al. (1983) regard it tentatively as a synonym of Aphanius dispar. Wildekamp (1993) is of the opinion that it may be a subspecies of Aphanius dispar. Hrbek and Meyer (2003) using mtDNA found this species to be deeply nested within the A. dispar clade. Villwock (2004) has used cross-breeding experiments that demonstrate this taxon and A. dispar are comparable to intraspecific crosses in other taxa. However, fin ray counts are non-overlapping and Reichenbacher et al. (2007; 2009) demonstrate differences in otolith morphology and affirm its validity as a species. Holly (1929b) reports a specimen of Cyprinodon dispar from the same spring. I was unable to verify the co-occurrence of two species in the hot spring through my own samples. Holly's specimen of A. dispar is in the Naturhistorisches Museum Wien (NMW 13799) but there are no field notes to confirm its locality with accuracy. It may have been collected in an adjacent stream. A. ginaonis is regarded here as a good species on morphological grounds (see below).

Four syntypes are in the Naturhistorisches Museum Wien (NMW 13800-13803). Holly (1929a) in his original description refers to 3 syntypes. One of these 4 fish was selected as a lectotype by F. Krupp (NMW 13800) and measures 21.8 mm standard length while the remainder measure 17.8-23.2 mm standard length.

Key characters

This species is distinguished from Aphanius dispar, the only other tooth-carp in this part of southern Iran by its locality and by having fewer total dorsal fin rays (5-7 as opposed to 8-11, mostly 9-10 in A. dispar from neighbouring drainages).

Morphologygy

Total dorsal fin rays 5(16), 6(38) or 7(6). Gut coiling is highly variable between individuals and is complex (illustrated in Coad (1980b)). Reichenbacher et al. (2007, 2009) give descriptions of otolith morphology. Chromosome number 2n=48, karyotype 14Sm + 34St and arm number 31 (Esmaeili et al., 2008). Esmaeili and Gholami (2007) give details of scale ultrastructure.

Saefali (1999) has D4-7, Pelvic5-7, Pect 13-17, A6-10, ll25-32,vert 26-30, bars10-22, gr13-21 SEE DATA SHEETS

Sexual dimorphism

Females have a higher mean numbers of anal fin rays, total vertebrae and flank bars, and longer pelvic fins (Coad, 1980b). Seifali and Sheidai (2001) found rostrum to pelvic fin distance, rostrum to anal fin distance, weight, total distance (? total length), head depth, head distance (? length) and body depth to differ between the sexes. Esmaeili and Gholami (2007) found that female scales were larger than those in males.

Colour

Males have a brighter colouration than females although colouration is individually variable within sexes. The back is mottled black. The dorsal, anal, pectoral and pelvic fins are a light orange. Flank bars are alternately black tinged with orange and grey-white, light blue or light orange. The side of the head anteriorly is iridescent light blue to green and the opercle is dorsally orange, postero-ventrally blue. The iris is iridescent green and the anterior ventral head surface is light blue. The peritoneum is black.

Preserved specimens have two bars on the male caudal fin with the fin margin hyaline. These bars are usually straight. Some fish have 1-3 bars or a forked bar. Females lack these bars and their caudal fin rays are lightly speckled. The male dorsal fin has 4-5 horizontal rows of spots, fading dorsally. These are not always well-developed and then consist of irregular, black pigmentation on the rays and membranes. The dorsal fin pigmentation in females is weak. The anal and pelvic fins have little or no pigment. The pectoral fins of males carry pigment concentrated on the rays, weakly speckled in females. Flank bars in females vary from weak to obvious, the light intermediate bars not as highly coloured as in males nor as well defined.

Size

Reaches 4.5 cm total length.

Distribution

This species was described by Holly (1929a) from the Ab Garm-e Ganow at 27°26-28'N, 56°18-20'E (various reports give differing latitude-longitudes for this well-known locality, hence the range). Ab Garm literally means "hot water" in Farsi and is used to denote a hot spring. Ganow means "foul water". The spring lies at an altitude of about 135 m (Moghadam, 1974) on the slopes of the Ganow Mountain (= transliterated as Ginao in German), hence its scientific name.

Zoogeography

The occurrence of sympatric A. dispar would confirm the species distinction of this species.

Habitat

The species is common within the bounds of the hot spring stream, numbering in the thousands in the 1970s, but no population trends have been recorded (but see below). Fish are found particularly along the stream margin and in many minor subsidiary springs which emerge a few metres from the main spring. Some fish are found in water as shallow as 1 cm but temperatures are 37-40°C (when shade air temperature was 20°C at 1400 hours on 27 January). The main spring itself issues from the ground at 30 litres second-1 and 41°C and drains as a stream about 10-15 m wide. A fault, over which the spring stream pours, isolates the fish from those downstream. Coad (1980b) summarises water chemistry and Reichenbcher et al. (2009) and Golmoradizadeh et al. (2012) give recent descriptions of the habitat. The water is clear and colourless but there is a strong sulphur odour. The stream bed is composed of stones and pebbles covered by lime-green to dark blue-green algae. Moghadam (1974) gives details of the algae species.

Age and growthth

Essentially unknown although fish are larger later in the year (11.8-31.4 mm standard length in January and 15.6-34.7 mm in March). Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 33 fish measuring 1.70-3.17 cm standard length. The a-value was 0.0210 and the b-value 3.375 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases).  Alavi-Yeganeh et al. (2011?check published paper date) give length-weight data on this species with a (intercept) being 0.0076 and b (slope) being 3.384 for sexes combined. Golmoradizadeh et al. (2012) gave growth parameters of W(t) = 0.012TL3.42 for females and W(t) = 0.0101TL3.38 for males. Females showed an asymptotic total length (TL) of 53.03 mm, a growth coefficient (K) of 0.15 year-1, t0 1.01 year, and natural mortality coefficient (M) 0.49 year-1. Males showed an asymptotic total length (TL) of 48.83 mm, a growth coefficient (K) of 0.21 year-1, t0 0.44 year, and natural mortality coefficient (M) 0.62 year-1 (note that the values for M are reversed in the abstract compared to the text by sex - the text value is given here). Females attained an age of 5+ years and males 4+ years. Maturity was attained at age 2+ years or older. Sex ratio was 1:1.33 (male: female) and significantly different.

Food

Gut contents are the lime green algae typical of the hot spring and chironomids.

Reproduction

Temperature is constant and daylength may then be important in triggering reproduction, although fishes caught on 27 January and 21 March have mature eggs of similar size (ca. 1.6-1.7 mm) suggesting an extended reproductive period. There was a strong correlation between absolute fecundity and weight (Golmoradizadeh et al., 2012).

Parasites and predators

Unknown.

Economic importance

None except related species are important in the aquarium trade and this is a potential economic use.

Conservation

Coad (1980b) listed this species as rare although it should be classed as vulnerable under the revised IUCN Red List Categories (IUCN, 1996) according to Coad (1998a).

The site is easily accessible by an asphalt road and is close to the large city of Bandar Abbas. Two hammams (bath houses) are in operation, one for men and one for women, the spring water is drained off to irrigate date palms, and local people also use the site for bathing, washing kitchen utensils and food containers, and washing clothes so there is input of soap and food debris (personal observations; A. R. Zinaei and H. R. Esmaeili, Payam-e Noor University, Bandar Abbas, 6 August 1997). The area is not under direct management by environment officials although in 1999 press reports referred to planned research studies aimed at preventing extinction of this taxon (IRNA, 29 September 1999) and the spring does lie in the Genu Protected Area (Biosphere Reserve) described by Zehzad et al. (1997).

The habitat could deteriorate if too much soap and food debris enter the stream. A single catastrophic event, such as a chemical spill, water diversion or construction activity, could eliminate the species. Recent construction has severely affected the population with only 1-2 specimens being caught and the population being replaced by A. dispar (H. R. Esmaeili, pers. comm., September 2007).

Reichenbcher r et al. (2009) suggest that A. dispar specimens have been added to the spring by local people to increase the population size. Otolith differences between samples taken in 201 and 2009 suggest hybridisation.

Local people using the spring should be educated on its ecological significance and alternative facilities for bathing and washing established. Roadside access to the spring should be prevented. The population structure and reproductive regime of the fishes could be analyzed to determine if they are susceptible to abrupt changes.

Further work

This species differs from A. dispar in a number of characters but this may merely be a consequence of the unusual environment of the hot spring. Cross-breeding experiments would be very revealing and would demonstrate or disprove reproductive isolation and hybrid fertility. The population is very important as the only species in a unique habitat, lending itself to studies of speciation, adaptation and variation in response to high temperatures.

Sources

Type material: see above, NMW 13800-13803.

Topotypes: CMNFI 1979-0175, 77, 11.8-31.4 mm standard length. check size range; CMNFI 1979-0416, 7, ? mm standard length, Hormozgan, Ab Garm-e Ganow below falls (ca. 27º26'N, ca. 56º20'E); CMNFI 1979-0417, 44, ? mm standard length, Hormozgan, Ab Garm-e Ganow, side springs and edge of mainstream (ca. 27º26'N, ca. 56º20'E); CMNFI 2007-0054, 13, ? mm standard length, Hormozgan, Ab Garm-e Ganow (ca. 27º26'N, ca. 56º20'E);

Aphanius isfahanensis
Hrbek, Keivany and Coad, 2007

Courtesy of M. Ebrahimi, A Teimori and Z. Gholami
Courtesy of M. Ebrahimi, A Teimori and Z. Gholami

Courtesy of M. Ebrahimi, A Teimori and Z. Gholami
Courtesy of M. Ebrahimi, A Teimori and Z. Gholamimi

Common names

كپوردندان اصفهان (= kapurdandan-e Esfahan or Esfahan tooth-carp).

Systematics

The holotype is in the Canadian Museum of Nature, Ottawa under CMNFI 2004-0001, male, 25.0 mm SL, Esfahan Province, Zayandeh Rud (Zayandeh River) at Varzaneh Bridge, 32º25'32"N, 52º39'14"E, 1 July 2002, Y. Keivany and S. Asadollah and paratpes are under CMNFI 2004-0002, 18 males, 20.8–30.9 mm SL, 18 females, 22.0–38.4 mm SL, of 49 total (13 not used in meristic and morphometric analyses, total range in size 12.1-38.4 mm standard length), same locality as holotype; AMNH 233639, 1 male, 25.2 mm SL, 1 female, 21.6 mm SL, same locality as holotype; MRAC 2004-05-P-01-02, 1 male, 25.2 mm SL, 1 female, 21.6 mm SL, same locality as holotype; GenBank accession numbers AY593488, AY593489, AY593497, and AY593498.

Key characters

Distribution and colour pattern identify this species. Males can be distinguished from those of all other Iranian species by having distinct black edge on the dorsal, anal, and pelvic fins. The dorsal fin is covered with a high density of black blotches. Females can be distinguished from females of A. sophiae and A. vladykovi by having flank-bars rather than spots. It can also be distinguished from A. farsicus by less well-defined bars terminating at a mid-flank stripe and a relatively light gray stripe at the caudal-fin base rather than a black spot or blotch. Flank-bars are also characteristic of females of Aphanius dispar, however.

Morphology

Morphometric data are given by Hrbek et al. (2007) but these characters cannot be used for a simple distinction from other Aphanius in Iran but only in multivariate space. This species is clearly distinguished at the genetic level from all other species of Aphanius. It has 82 molecular apomorphies – 19 transversions, two transversions/transitions (depending on comparison), and 61 transitions – that show fixed character state differences to homologous characters analyzed in A. sophiae, A. farsicus (formerly persicus), and A. vladykovi from Iran. Thirty-seven of these character states are also apomorphies when compared to A. anatoliae, A. danfordii, A. villwocki, A. asquamatus, and A. fasciatus from Turkey.

Dorsal fin rays 11-14, anal fin rays 10-13, pectoral fin rays 13-16, pelvic fin rays 4-6, lateral line scales 25-29, total gill rakers 10-13, precaudal vertebrae 10-13 and caudal vertebrae 15-18. Chromosome number 2n=48, karyotype 12Sm + 36St and arm number 30 (Esmaeili i et al., 2008).

Sexual dimorphism

See colour below.

Colour

Male flank bars number 8-12, mean 10.2, significantly less than in A. sophiae at 10-21, mean 14.4. The bars are broad with interspaces about equal or slightly narrower. The bars extend from behind the head to the tail. Anterior bars fade on the belly, whereas, posteriorly on the caudal peduncle, they encircle the body. Dorsally, the head is dark gray and the body is lighter but still heavily pigmented with melanophores; the belly lacks pigmentation. The sides of the head are densely speckled with melanophores, more thinly on the ventral side; in most specimens the chin is darker than the rest of the ventral head surface. The eye is bounded ventrally and postero-ventrally by a thin line of black pigment. The defining male coloration is the black margins of the pelvic, anal, and dorsal fins. The dorsal and anal fins may present a halo effect, the margins being so dark in relation to the rest of these fins. The tips and outer margin of the pelvic fin are blackish. The anal fin has a broad, blackish margin with the rest of the fin light cream-colored. The dorsal fin has the blackest margin. The rest of the dorsal fin is variably blotched, the blotches being much lighter than the fin margin. Most specimens have a contrasting pigmentless area just below the fin margin. The pectoral fin has sparse pigmentation along rays and ventrally on the interradial membranes but lacks the concentrated black pigmentation seen on the pelvic fins. The caudal fin rays and membranes are sparsely pigmented, and the whole margin may be blackish but in most fish pigment is restricted to the upper and lower margins, the lower margin only, or is absent. Large females have a grayish dorsal surface to the head, a lighter back and upper flank covered with scattered melanophores, and a mid-flank stripe terminating on the caudal peduncle in a blackish, short stretch covering up to three scales (Fig. 2b). This short stretch of pigment is present in all females, faint in some, rarely forming a blotch and in some small fish tapering anteriorly. The flank stripe may be broken into a series of blotches in some smaller females, or it may be continuous as in large fish. Starting anterior to the belly there is a ventrolateral series of thin bars (up to ten) separated by cream colored interspaces 1–3 bars wide. These are absent in some smaller females, which may only have blotches at various levels present in this region. Even some large females have faint flank pigmentation so that bars and the stripe are weakly expressed. At about the origin of the anal fin, the flank bars may continue onto the caudal peduncle in regular form or become irregular, breaking up into blotches. Anteriorly the bars terminate ventrally at about the level of the lower edge of the pectoral fin. The belly and lower head have sparse pigment although the chin and sides of head are speckled with melanophores. The eye is bounded ventrally and posteroventrally by a thin line of black pigment. Fins lack any distinctive pigment pattern. Fin rays are outlined with melanophores, and interradial membranes of the caudal and anal fins have melanophores at varying degrees of density. The dorsal fin has the most interradial pigmentation, particularly near, but clear of the fin base.

Size

Attains 38.4 mm standard length, 42.8 mm total length (Alavi-Yeganeh et al., 2011?check published paper date).

Distribution

The type locality is near the town of Varzaneh on the lower reaches of the Zayanadeh River, about 30 km upriver from the terminal sump, the Gav Khuni marsh.

Zoogeography

An estimate of a 4.8 MYA divergence of A. isfahanensis and A. sophiae + A. persicus (now farsicus) is given by Hrbek et al. (2007). This divergence time is in accord with the hypothesis of a near-simultaneous diversification of ~5 MYA of organisms occupying different geological units of central Iran.

Habitat

The type locality had a water temperature of 27ºC, pH was 6.7, the water was brackish, conductivity was 10.9 mS, dissolved solids were 5450 ppm, dissolved oxygen was 12.3 mg/L, river width was 50 m, and capture depth was 0.5 m. Current was slow, and there was no cover. Gambusia holbrooki was captured at the same locality. Bleher (20110 notes  that the type locality was dry at Varzaneh Bridge and only found specimens in an artificial circulating spring channel 50 m long in southern Esfahan.

Age and growth

Unknown. Alavi-Yeganeh et al. (2011?check published paper date) give length-weight data on this species from two localities with a (intercept) being 0.0120 and b (slope) being 3.035 for all localities and sexes combined.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Gholami et al. (2009) record the trematode Diplostomum spathaceum from the eyes of this fish.

Economic importance

None at present but a potential aquarium species. Breeding and aquarium conditions are similar to A. mento although it is reputedly difficult to breed.

Conservation

Known only from the type locality, which is not protected, this species needs a conservation assessment.

Further work

The biology of this species requires study.

Sources

Type material: See above, CMNFI 2004-0001, CMNFI 2004-0002, AMNH 233639, MRAC 2004-05-P-01-02.

Aphanius mento
(Heckel, 1843)

Male

Common names

gour-e khar, kopurdandandar-e Arvand (= Arvand River tooth-carp), kopurdand-e Irani.

[batrikh qabras in Arabic; Persian pupfish, Persian minnow, Black Persian minnow, Persian killie, Arvand pupfish, chin killifish, pearl-spotted killifish].

Systematics

Lebias Cypris Heckel, 1843 is a synonym. Krupp (1984a) gives a detailed synonymy of this species but considered cypris to have priority. Klee (1967) and Lazara (1989) point out that Garman (1895), in a work dated in July of that year, as first revisor placed cypris in mento. This decision has priority over Gaillard (1895), a work with no exact date of publication, in which mento was placed in cypris, according to the Zoological Code of Nomenclature (Ride et al., 1985). The type locality for both Lebias mento and Lebias cypris is "Mossul" according to Heckel (1843b).

Hybrids with h Aphanius dispar richardsoni are reported from Israel (Goren and Rychwalski, 1978).

Parenti (1981; 1984) places this species in the genus "Aphanius", i.e. distinct from true Aphanius without defining and naming a new genus. This genus is more closely related to the Anatolian Kosswigichthys Sözer, 1942 and the South American Orestias Valenciennes in Cuvier and Valenciennes, 1846 than true Aphanius. Hrbek and Meyer (2003) note that, based on their mtDNA study, this species diverged from the ancestor of the A. dispar clade at an early date, 26.87±1.31 MYA. They speculate that the ancestor of mento invaded the northern perimeter of the Arabian plate as it was abutting Laurasia some time after the separation of their eastern and western clades of Aphanius (see above) before the closing of the Tethys Sea. It later spread through the present-day Tigris-Euphrates and into the Levant and southern Turkey.

The 5 syntypes of Lebias cypris are in the Naturhistorisches Museum Wien (NMW 59598) with 1 specimen selected as a lectotype by Krupp and Schneider (1989), the other 4 being paralectotypes. The syntypes of Lebias mento are (NMW 62105 (4 fish), 59042 (5) and 22624-22632 (9). Eschmeyer et al. (1996) list NMW 21699-704 (6) and NMW 59832 (21) as possible syntypes. The Vienna catalogue lists 5 specimens of Lebias cypris and 8 of Lebias mento.

Key characters

The adult colour pattern is distinctive and dorsal fin ray counts are usually higher than in A. dispar.

Morphology

Scales along side of body 23-28, total dorsal fin rays 9-14, usually 12-14, total anal fin rays 9-13, pectoral rays 12-16, and pelvic rays 4-6, usually 5. Frontal scalation is E-type (Hoedemann, 1958). Flank scales are squarish to a vertical oval. The former shape has a vertical anterior margin which may be wavy or slightly convex, upper and lower anterior corners rounded but square-cut, dorsal and ventral margins parallel, and the posterior margin rounded. The oval scales have all margins rounded. The focus is central to subcentral posterior, circuli are fine although coarser on the posterior field, radii are restricted to the anterior field and are almost horizontal and parallel. There is no pelvic axillary scale. Total gill rakers 11-15, rakers being spinulose and reaching beyond the adjacent raker when appressed. Vertebrae 25. Teeth are tricuspid with a long and pointed central cusp (Goren and Rychwalski, 1978). There are 14-16 teeth on the lower and 10-12 on the upper jaw (Berg, 1949). The gut is s-shaped. Reichenbacher et al. (2007) give a description of otolith morphology. Chromosome number 2n=44 or 48 (Wildekamp, 1993; Klinkhardt et al., 1995).

Sexual dimorphism

See below under Colour.

Colour

Adult, breeding males are a dark blue-black to dark brown or almost black with iridescent blue-white to silvery spots regularly-arranged on the fins as curved lines, and irregularly on the body (sometimes as irregular vertical bars and sometimes the spots are vertically elongate) (Richter, 1989; preserved material from Iraq; Goren and Rychwalski, 1978). The edges of the gill cover are orange-red (Wildekamp et al. (1999). Male colour fades in low light conditions and in winter males have silvery flanks with a dark brown back. Spots are silvery-blue on the upper flank and are not as numerous as in the spawning male. Females are grey-brown or grey-white to silvery with large golden blotches or silvery to blue spots and dark dots. Scales along the mid-flank usually have a dark border (Wildekamp et al. (1999). Fins in females are hyaline. Body colour is reportedly heightened in brackish water (Grimes, 1974). The peritoneum is silvery with dense but fine melanophores.

Sizeze

Reaches 8.4 cm total length or more (Güçlü and Küçük, 2008).

Distribution

Found in the Orontes (= Asi) and Tigris-Euphrates basins, the Levant in coastal and Dead Sea basins, western Jordan, and in southern Turkey in Mediterranean basins as well as in central Turkey. Tigris-Euphrates records are relatively rare (see map in Wildekamp (1993)). The species was described from Mosul in northern Iraq and there are records from the region of Basrah in southern Iraq near the border with Khuzestan. Wildekamp (1993) has one record apparently mapped in southern Iran near the border with Iraq. This is the only record from Iran although Abdoli (2000) indicates the Arvand River on a map.

Zoogeography

See Systematics above and under the genus.

Habitat

This species has been reported to survive in waters up to 38°C and as low as 0°C. pH at 7.0-8.5 has been reported as ideal for aquaria maintenance. Aquarists report it as a robust species favouring hard and alkaline water with some salt added (Thiermann, 1978; Kostich, 1979; Phillips, 1985). Grimes (1974) considers it to be quarrelsome in aquaria like other Aphanius and Allen (1988) also noted that males are very aggressive. It inhabits fresh or slightly brackish water of springs, streams and lakes, usually near shore where males establish territories in vegetation. It generally prefers habitats dense in vegetation.

Age and growth

Life span is about 3 years with maturity in aquaria at 4-6 months. The length-weight relationship in southern Iraq was W = 6.0001 x 10-5 L2.6403 for males and W = 2.3028 x 10-4 L2.235 for females. The average condition factor for males was 2.0499 and for females 2.3121. The condition factor increases with increase in length in females and decreases in males (Huq et al. 1977). Güçlü and Küçük (2008) examined a population of this species in a spring in the Mediterranean Sea basin of Turkey and found fish in age groups 0 to 7. They also give length-weight relationships and a von Bertalanffy growth equation.

Food

Al-Daham et al. (1977) found filamentous algae and diatoms were the most important foods in southern Iraq. Shafi and Shalli (1986) report a diet of beetles, ephemeropteran nymphs and algae in southern Iraq. Aquarium specimens prefer meaty foods such as brine shrimps over flake foods (Grimes, 1974) but flake food is taken (Allen, 1988). Curiously, Al-Daham et al. (1977) for their southern Iraqi fish in aquaria found this species to prefer plant food. Males approached food items individually and prevented other fish from taking the food. Females were also solitary though less aggressive than the males. However Gambusia holbrooki out-competed male A. mento for food, on one occasion even snatching a Gambusia embryo from the mouth of an A. mento. Güçlü and Küçük (2008) found their Turkish fish to feed principally on Gammarus sp and Palaemon sp.

Reproduction

Anderson (1966), Lutman (1982) and Richter (1989) observed spawning in aquaria. There is a dominant male which is more deeply and richly coloured than other males. The dominant male defends a spawning territory, fighting other males. Males will defend separate spawning mops if these are made available in aquaria. The dominant male will pursue a female and places his larger dorsal fin over the her back. Several spawnings will follow and females will spawn with more than one male. Spawning can occur near the surface. These fish can spawn at six months of age. Peak spawning in southern Iraq is April to June when only a small proportion of eggs in the single ovary contained fully developed eggs (Al-Daham et al., 1977). However, Shafi and Shalli (1986) state that it breeds in southern Iraq in May-July. There are up to 71 highly-adhesive, eggs of 170 microns diameter. Aquarium spawned eggs hatch in 7-14 days, depending on temperature, 10 days at 25°, 3 days in the 80s°F, or 6-7 days in the low 70s°F (accounts vary). Allen (1988) found egg production to be higher in aquaria when the equivalent of seven tablespoons of salt was added per gallon at a water temperature of 25°C. A minimum of 20 eggs per day were produced.

Byniak (1979) describes the effects of day length and temperature on the reproductive cycle of females in this species in Israel.

Parasites and predators

Unknown.

Economic importance

An aquarium fish with breeding details given by Zipay (1961), Anderson (1966), Lutman (1982), Richter (1989), Semeit (1999) and www.killi-data.org/ (downloaded 21 September 2007). The species is hardy and fry are easily raised, given large enough tanks and an easily defensible area for the male to guard eggs. However, populations stop breeding in aquaria after 6-8 generations (www.killi-data.org/, downloaded 21 September 2007). Blaustein and Byard (1993) show that this species will prey on mosquito larvae in a laboratory situation and has potential in control of this vector of malaria, at least in vegetated habitats.

Conservationon

The presence of this species in Iran needs confirmation by field work and until this is done and its distribution, population numbers and biology worked out, its conservation status cannot be assessed. Endangered in Turkey (Fricke et al., 2007).

Further work

See above.

Sources

Iranian material: None.

Comparative material: BM(NH) 1920.3.3:203-222, 22, 24.9-36.0 mm standard length, Iraq, Basra (30º30'N, 47º47'E); BM(NH) 1949.7.21:12, 24.4 mm standard length, Iraq, Bahr-el-Milh Lake east of Karbala (32º35'N, 43º50'E); BM(NH) 1981.10.6:19-23, 5, 26.6-31.8 mm standard length, Iraq, Qarmat Ali, Basra (30º30'N, 47º47'E); BM(NH) no catalogue number, 4, 19.7-31.7 mm standard length, Iraq, Basra Liwa (no other locality data); uncatalogued, 1, 31.2 mm standard length, Iraq, Khawr az Zubayr (no other locality data). ?Iraq fish from Hussain

Aphanius mesopotamicus
Coad, 2009

 

Male paratype 21.7 mm SL

 

Female holotype
Male paratype 21.7 mm SL above, female holotype below

 

 From Basrah, Iraq (ZISP 25393) after Berg (1949), female 30 mm

 

From Basrah, Iraq (ZISP 25393) after Berg (1949), male, 29 mm

From Basrah, Iraq (ZISP 25393) after Berg (1949). Female (above), 30 mm, and male, 29 mm

 

Common nameses

None.

Mesopotamian tooth-carp.

Systematics

The holotype is a female, 29.3 mm SL from Khuzestan, canal branch of Karkheh River, 31º40’N, 48º35’E (CMNFI 1979-0360A) and paratypes are 37 specimens, 14.6-29.1 mm SL, same locality as above (CMNFI 1979-0360B) and 6 specimens, 15.1-20.5 mm SL, from Khuzestan, Karkheh River branch at Abdolkhan, 31º52’30”N, 48º20’30”E (CMNFI 1979-0364). The species is named for Mesopotamia (the land between the rivers) referring to the Tigris-Euphrates basin where the species is found. .

Key characters

This species is distinguished by distribution and by pigmentation, males having clear margins to the unpaired fins, no bars on the caudal fin and 10-15 clearly defined flank bars, females bear irregular blotches or spots on the flank.

Morphology

Total dorsal fin rays 11-13, total anal fin rays 10-12, branched pectoral fin rays 13-15, branched pelvic fin rays 5-6, lateral series scales 25-29, total gill rakers 10-14, precaudal vertebrae 11-13 and caudal vertebrae 14-17.

Sexual dimorphism

See colour description.

Colour

Males have clear margins to the unpaired fins, no bars on the caudal fin and have 10-15 clearly defined flank bars. The dorsal surface of the head and the upper flank are more heavily pigmented with melanophores than more ventral areas. The belly and lower head are unpigmented. The chin and snout have dense melanophores and a rim of melanophores underscores the eye in both sexes. The dorsal, anal and caudal fins in males have wide clear margins. This is also seen in the material from Basrah, Iraq (BM(NH) 1982.9.2:326-328). The material from Basrah figured by Berg (1949) has distinct light margins in some printed versions of the figures. The caudal fin is darker just proximal to the clear margin, lighter in mid-fin and dark again at the base. The dorsal fin has irregular pigmentation on the membranes and, to a lesser extent, on the rays. The pigmentation may involve an overall darker colour in contrast to the light margin or may have some pattern to it. The pattern is often elongate and short blotches with no regular arrangement and sometimes may appear as up to 5 wavy and oblique bands. Dark pigmentation is found just behind the first ray on the fin membrane. The anal fin is darkest just proximal to the clear margin. Up to the last 6 membranes of the anal fin are dark and this pigment may be broken up in as many as 4 elongate bars along each membrane. A similar pattern is found in some dorsal fins and the general effect on both fins is that the postero-dorsal (anal fin) and postero-ventral (dorsal fin) parts of these fins are the darkest. The dorsal, anal and caudal fins generally have more pigment on the membranes than the rays and in some this is quite distinctive, making the rays stand out.

The pectoral and pelvic fins in males are generally clear or somewhat milky and opaque and lack melanophores. The distal parts of the membranes between the last 5 rays of the pectoral fin and the small membrane area of the pelvic fins can be pigmented. Males have flank bars circling the caudal peduncle and reaching the anal fin base but fading ventrally on the lower part of the anterior flank, not reaching the ventral margin of the belly and becoming progressively shorter and less distinct the more anterior they are. Bars are 2-5 times broader than the pale interspaces.

Females have a similar head and dorsal and ventral body pigmentation but it is much lighter than in males. Fins have little or no pigmentation. The proximal third to half of the dorsal fin rays and membranes, particularly the anterior ones, have pigment in some fish but this is weakly expressed compared to the condition in males. Some fish have a few faint melanophores lining the anal fin rays.s.

The most distinctive feature in females is a spot, oval to lozenge-shaped, at the central base of the caudal fin. Its concentration of melanophores is much higher than any other feature. The flank in females can have up to 14 thin, dark, wavy and irregular vertical patches of pigment. These patches may be interrupted in their vertical extent and are weakly expressed anteriorly. They fade ventrally, ending above the lower edge of the caudal peduncle and above the anal fin base, and are absent on the lower anterior flank. The patches are light and not as contrasting with the lighter interspaces as the bars found in males. Patches are thin, half to one third of the width of the interspaces. Very often the patches are broken up into spots and elongate blotches of various sizes, and a regular barred appearance is not usual. The spots and blotches are all smaller than the eye size by at least half. Material from Basrah, Iraq figured by Berg (1949) has the spots emphasised but material from Basrah (BM(NH) 1982.9.2:326-328) examined for this study has a more blotchy appearance and spots are not well-defined.

Size

Attains 29.3 mm standard length.

Distribution

Found in the southern Karkheh River basin of Iran and at Qarmt 'Ali at the northern part of Basrah on the Shatt al Arab, southern Iraq. Possibly also at Shalili-ye Bala or Shalili-ye Pa'in (ca. 31º58'N, 48º53'E), near Shushtar, Karun River basin of Iran from a record in Berg (1949) from Shellali (ZISP 25446). Keivany et al. (2012) captured fresh material from a Karkheh River branch 10 km west of Hoor-Al-Azim Wetland (31º32'59.4"N, 48º09'48.3'E, not shown on map below).

Distribution map of Aphanius in Mesopotamia and adjacent areas. Solid squares type series of A. mesopotamicus (lower one CMNFI 1979-0360A and B, upper 1979-0364), open squares additional, non-type material of A. mesopotamicus (BM(NH) 1982.9.2:326-328 and ZISP 25393 (lower left) and ZISP 25446 (adjacent to type series)), open circles A. vladykovi, stars A. isfahanensis, solid circles A. sophiae, and diamond A. farsicus (map modified after Hrbek et alal. (2006)).

Zoogeography

This species is found in the Tigris River basin and its morphology shows evidence of relationship with species of internal basins in Iran.

Habitat

The habitat of the new species is known only from field notes made at the time of capture in Iran on the Khuzestan plains. The two localities are a river and a canal branching from that river. The 25 m wide river had a water temperature of 22°C at 1205 hours, pH 6.0, conductivity 1.0 milliSiemens, a mud bottom and the principal plant materials were rushes and reeds. The 30 m wide canal had a water temperature of 15°C at 0930 hours, pH 6.0, conductivity 1.8 milliSiemens, a mud bottom and the principal plant material was filamentous green algae. Keivany et al. (2012) captured material in a river branch with calm water flow on a sandy bed with Ceratophyllum, Typha and Phragmites, water temperature 24.4°C, dissolved oxygen 8.8 mg/l at 106% saturation, pH 8.23, conductivity 2367 µS and total dissolved solids 1181 mg/l.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

None recorded in Iran.

Economic importance

None at present in Iran but a possibly a species of interest to aquarists.

Conservation

This species is known only from museum collections and recent efforts to capture fresh material were unsuccessful. These attempts should be repeated and the numbers and biology of the species assessed.

Further work

The biology and continued presence of this species in Iranian waters requires study.

Sources

Based on Coad (2009).

Type material: See above.

Comparative material: BM(NH) 1982.9.2:326-328, 4, 20.2-25.3 mm SL, Iraq, Qarmat `Ali, Basrah (30°34’N, 47°46’E).

Aphanius pluristriatus
(
Jenkins, 1910)

Male, courtesy of H. R. Esmaeili
Male, courtesy of H. R. Esmaeili

Female, courtesy of H. R. Esmaeili
Female, courtesy of H. R. Esmaeili


Female and male, Mand River basin courtesy of  H. R. Esmaeili

Cyprinodon pluristriatus male after Jenkins (1910)
Cyprinodon pluristriatus male after Jenkins (1910)

Common names

Kapour-e-dandandar-e-Mond.

[Multi-striped (Mond) tooth-carp]

Systematicscs

Cyprinodon pluristriatus Jenkins, 1910 was often referred to the synonymy of A. sophiae but is found "East of Shiraz, stream running to Fussa, Southern Persia, 5,000 feet". Assuming this to be Fasa and using topographic maps, the locality is in the Mand River basin draining to the Persian Gulf. Recently, Esmaeili et al. (2012) have re-discovered and re-described this species from the Zarjan Spring system in the Mand (or Mond) River basin of Fars near Fasa, presumed to be the type locality.

Four syntypes of Cyprinodon pluristriatus (ZSI F9408 to ZSI F9411) are in the Zoological Survey of India, Calcutta (Menon and Yazdani, 1968), although Jenkins (1910) has ZSI F9408-9412.

Key characters

Distribution, a smaller caudal peduncle index, higher number of flank bars on average, lower number of gill rakers and higher J-scale index separate this species from other southern Iranian Aphanius species (Esmaeili et al., 2012).

Morphology

Morphological and meristic characters overlap with other Aphanius species examined by Esmaeili et al. (2012), q.v. for details, although the both sexes of this species are separated by discriminant function analyses. For example, gill rakers number 8-11 with mean values of 9.84 for males and 9.58 for females (ranges for other species are 8-13 for A. farsicus, means 10.47 for males and 10.43 for females; 10-13 for A. isfahanensis, 10.78 and 11.16; and 9-12 for A. sophiae, 10.66 and 10.71).

Sexual dimorphism

See colour description.

Colour

The dorsal, caudal and anal fins in males have a pale margin, proximal to which the fin is dark with the bulk of the fin lighter in colour. The main part of these fins has numerous rows of spots. The lower edge of the pectoral fin is black. The dark fin margins and spots are absent in females. Males have alternating thin white and brown flank bars, the white bars numbering 11-17 with a significantly higher mean (13.84) than A. farsicus (9-16 bars, mean11.56), A. isfahanensis (9-13, 10.67) and A. sophiae (8-15, 11.89) (Esmaeili et al., 2012). Females have thin or thick, dark, wavy and irregular short bars or blotches of pigments on the flank, some approaching a spot shape. An oval to lozenge-shaped dark spot is at the central base of the caudal fin and this may be broken up into several spots. Both males and females have small and irregularly arranged spots on the operculum.

Size

Not given

Distribution

Specimens were known only from the Zarjan Spring system in the Mand River basin of Fars near Fasa, but possibly exist in a qanat near Jahrom (Esmaeili et al., 2012).

Zoogeography

Esmaeili et al. (2012) suggest that this species diverged from an Aphanius population in the ancient Kor River system. In the Quaternary, a Palaeo-Kor River crossed the type locality of A. pluristriatus and drained to the Persian Gulf via the Mand River. In the late Quaternary (20,000-10,000 years ago) or Holocene (6000-2000 years ago), this exorheic drainage was closed due to tectonic uplift of the Zagros Mountains. The Kor River basin became endorheic and isolated from the exorheic Mand River basin. The Kor and Mand Aphanius populations diverged to become sophiae and pluristriatus respectively.

Habitatat

Unknown in detail.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

None recorded.

Economic importance

None at present in Iran but a possibly a species of interest to aquarists.

Conservation

This species is known only from the Zarjan Spring system but has not been collected post-2003 because of drought and competition with such exotics as Xiphophorus hellerii (Esmaeili et al., 2012). It could be extinct and a thorough survey of all possible habitats should be made.

Further work

See above under Conservation.

Sources

Esmaeili et al. (2012), Zarjan being at 29°03'39.44N, 53°36'59.23E (H. R. Esmaeili, pers. comm., 5 March 2012).

Aphanius sophiae
(Heckel, 1847)


Male


Female

Cyprinodon blanfordii female after Jenkins (1910)
Cyprinodon blanfordii female after Jenkins (1910)0)

Male courtesy of H. R. Esmaeili, A. Parsi, Z. Gholami and M. Tahami
Male courtesy of H. R. Esmaeili, A. Parsi, Z. Gholami and M. Tahami

ale courtesy of H. R. Esmaeili, A. Parsi, Z. Gholami and M. Tahami
Male courtesy of H. R. Esmaeili, A. Parsi, Z. Gholami and M. Tahami

Female courtesy of H. R. Esmaeili, A. Parsi, Z. Gholami and M. Tahami
Female courtesy of H. R. Esmaeili, A. Parsi, Z. Gholami and M. Tahami

Common names

gour-e khar, kopurdandan-e safiyeh or kopurdandan-e sufieh (= sufi or sophy tooth-carp).

[batrikh sophiae in Arabic; Persian minnow, Cypris pupfish, but this seems inapt].

Systematics

The type locality of Lebias Sophiae is in "lauen Salzquellen bei Persepolis" according to Heckel (1847b).

Lebias punctatus Heckel, 1847 is listed from "Nemek-Deria oder Salzsee, in welchen sich unter Schiraz die Quellen des Saadi ergiessen" (Nemek-Deria or salt lake below Shiraz into which the Saadi springs flow), i.e. Lake Maharlu in the Shiraz Valley, the spring at Sa'adi's Tomb being a tributary, if somewhat distant (Heckel, 1847b). Lebias crystallodon is from "grossen Salzsee Nemek-Deria, unter Schiraz" (Heckel, 1847b). Both punctatus and crystallodon are synonyms (Günther, 1859-70; Garman, 1895; Gaillard, 1895; Hanko, 1924; Berg, 1949; Coad, 1996a).

The original catalogue, handwritten by Heckel in Vienna, gives the type locality of sophiae as "Salzquellen bei Persepolis" which is in agreement with Heckel's publication. The catalogue is more equivocal for the two nominal species, punctatus and crystallodon, listing only "Nemek Deria". The labels in jars containing L. crystallodon and L. punctatus were written under Franz Steindachner's curatorship and are not original (H. Ahnelt, in litt., 1987). Nemek Deria (= salt lake in Farsi) is a general term employed for the type of terminal water body seen in both the Maharlu and Kor River basins. The situation was further confused by Berg (1949) who listed the locality of punctatus as "Lake Niriz in Shiraz". Lake Niriz (= Neyriz or Bakhtegan) is the terminal sump for the Kor River.

All material of Aphanius from the Kor River basin examined fresh by me comprises fish with bars along the flank and fish with spots. The barred fish are males and the spotted fish females as confirmed by dissection. Males and females do not segregate in the field and it seems very unlikely that the material examined by Heckel from several collections would be comprised solely of males (I found sample sizes as low as 5 to contain both males and females). The males and females may have been separated and allocated different collection localities. The material described as L. punctatus is then female L. sophiae and I strongly suspect that it was misunderstood as coming from the Shiraz valley and the locality in Heckel (1847b) is incorrect. All female Aphanius from the Shiraz valley are barred and the name punctatus is singularly inappropriate and would not be applied to them (see illustrations in Villwock (1960; 1977)). L. crystallodon could be a female L. sophiae with weak spotting on the flanks, again mislabelled as from the Shiraz valley. Günther (1866) considers crystallodon to be a post-spawning female of L. sophiae and Gaillard (1895) also considers it to be a female L. sophiae. Berg (1949) places L. sophiae as the male and L. punctatus and L. crystallodon the females of L. sophiae.

Cyprinodon blanfordiiii Jenkins, 1910 may well be a synonym of A. sophiae (see above under A. farsicus for other views). The locality is vague, "East of Shiraz, South Persia" and could lie in the Lake Maharlu basin (occupied only by A. farsicus, the females of which are barred on the flank), the Kor River basin (A. sophiae only), or outside either of these, in rivers draining to the Persian Gulf. The illustration shows a finely-spotted fish with a lozenge-shaped larger spot at the tail base, pigmentation found in A. sophiae.

Parenti (1981; 1984) places Aphanius sophiae in the genus "Aphanius", i.e. distinct from true Aphanius without defining and naming a new genus. This genus is more closely related to the Anatolian Kosswigichthys and the South American Orestias than true Aphanius.

The syntypes of Lebias sophiae are in the Naturhistorisches Museum Wien and include a wide range of material of which the following is part:-

Lectotype and paralectotypes (as labelled in 1997): NMW 14496 (6 fish measuring 25.8-30.4 mm standard length, one of which measuring 29.3 mm standard length is the lectotype as designated by F. Krupp, 24 October 1984 according to a jar label), NMW 22616-22623 (8, 26.1-29.0 mm standard length), NMW 60327 (8, 25.2-31.1 mm standard length), NMW 68283 (8, 18.9-27.6 mm standard length, partially dried and distorted), NMW 75067 (7, 24.7-27.4 mm standard length), and probable syntypes NMW 14760 (67) and NMW 15056 (7). The catalogue in Vienna lists 40 fish in one column and 12 fish in the adjacent column, the latter possibly meant to be the type series as it is written more boldly but which fish these were cannot now be ascertained. All these fish are males.

Eschmeyer et al., (1996) list syntypes under NMW 14496 (6), NMW 33616-23 (8) (not noted above), NMW 60327 (8), NMW 68283 (8), NMW 75067 (7) and also 9 fish in the Museum für Naturkunde, Universität Humboldt, Berlin under ZMB 31377 (not noted above). The Berlin material is listed as ex 16140, is all male and 20.3-26.7 mm standard length.

Syntypes of Lebias punctatus comprise the following material:-

Lectotype and paralectotypes (as labelled in 1997): NMW 76509 (7 fish, measuring 29.9-36.0 mm standard length, one of which measuring 36.0 mm standard length is the lectotype as designated by F. Krupp, 24 October 1984 according to a jar label), NMW 15070 (5, ca. 29.9-35.6 mm standard length, dried and distorted), NMW 15156 (14, 19.2-32.1 mm standard length), NMW 59837 (5, 26.6-35.7 mm standard length), and probable syntypes NMW 59609 (6, ca. 21.4-34.7 mm standard length, smallest dried). The Vienna catalogue lists only 12 fish opposite this name. These fish are all females.

The holotype of Lebias crystallodon measuring 40.1 mm standard length is NMW 15175 as the description and catalogue agree there is only one fish. The specimen is decoloured and in poor condition.

Key characters

The large, dark, usually lozenge-shaped spot at the caudal fin base in females with fine spotting on the flanks is distinctive for tooth-carps in this area of southern Iran.

Morphology

Lateral line scales 25(1), 27(7), 28(14), 29(17), 30(12) or 31(2). Scales above the lateral line 4-7, scales between lateral line and the anal fin 5-8, scales between the lateral line and the pelvic fin 6-9, and scales around the caudal peduncle 15-20.

Total dorsal fin rays 11(1), 12(11), 13(27), 14(12) or 15(2); total anal fin rays 10(1), 11(18), 12(27) or 13(7); total pectoral fin rays 14(1), 15(7), 16(31), 17(10), 18(3) or 19(1); total pelvic fin rays 5(9) or 6(44); total gill rakers 10(2), 11(28), 12(21) or 13(2); abdominal vertebrae 11(11), 12(37), or 13(5); caudal vertebrae 15(3), 16(33) or 17(17); and total vertebrae 27(8), 28(29) or 29(16).

Cephalic sensory pores are reduced to a series of neuromasts. Older literature on chromosome number was 2n=48 but is based on fish from outside the Kor River basin to which the species is restricted here (Klinkhardt t et al., 1995). Esmaeili et al. (2007) determined a chromosome number of 2n=48 with 14 pairs of submetacentric and 10 pairs of subtelocentric chromosomes and an arm number of NF=76. Karyotype 8Sm + 40St and arm number 280 (Esmaeili et al., 2008).

Sexual dimorphism

Males have more bands on the flank and are darker and more intensely coloured than females. Male dorsal, anal and caudal fins are a deep black with narrow, silvery-white borders and the anal fin has a few silvery spots at its base. Females have brown dorsal, anal and caudal fins with 3-4 rows of black spots and the anal fin also has 2 cross rows of white silvery spots near the base. The caudal fin spot in females can be oval, tear-drop shaped or elongate but is usually in the form of a lozenge. Occasionally, single, smaller, dark, subsidiary spots may be found antero-dorsally and antero-ventrally to the basal spot or scattered spots may be found irregularly before and behind the basal spot. Male flank bars 10(4), 11(1), 12(2), 13(3), 14(5), 15(4), 16(6), 17(3), 18(1) or 21(1).

Eleven of 21 morphometric characters are significantly different between the sexes including head length, head and body depths, fin lengths, and caudal peduncle shape (Coad, 1998i).

Colour

"The body colouration is basically dark brown; the body, with the exception of the forward dorsal portion and the belly, is marked by 12-17 white, silvery shining vertical lines or narrow bands. Those individuals which have the most bands and are also darker and more intensively coloured seem to be males; their vertical fins are deep black with narrow silvery-white borders, and only the anal fin has a few silvery spots at the base. The lighter individuals which are also slightly higher and wider, we believe to be the females; they have brown vertical fins with 3-4 cross rows of black spots; their anal fins are marked with 2 cross rows of white silvery spots at the base" (Heckel, 1847b).

The caudal fin spot in females can be oval, tear-drop shaped or elongate but is usually in the form of a lozenge. Occasionally single, smaller, dark, subsidiary spots may be found antero-dorsally and antero-ventrally to the basal spot or scattered spots may be found irregularly before and behind the basal spot.

The flank spots in females are much lighter than the caudal base spot and are from a half to much less in size, usually much smaller than half size. Spots are usually rounded but can be oval and vertically elongate. Spots are independent of scale arrangement on the lower half of the flank, particularly posteriorly and may form 2-3 poorly defined to distinct longitudinal rows. Upper flank scales have a crescent of pigment in mid-scale, leaving the margin and anterior base mostly free of pigment, or pigment may fill each exposed scale surface almost entriely except for the posterior margin, and thus appear as spots. Females have purplish tints on mid-flank and yellowish tints on the lower flank, upper flank, head top and chin. Recent collections and photographs (see above) show females from the Kor River basin without the characteristic small spots of the type series and collections made in the 1970s. It is not known whether the is natural variation or the result of translocation of specimens.

Males have a pigmentation very similar to that of A. farsicus and the description here is identical. Some minor observed variation is attributed to variation in size and maturity of the fish compared. Males have light flank bars half the width or much narrower than alternating dark bars. The margins of the dorsal, anal and caudal fins are clear while the rest of these fins is dark. Some fish have up to 3, but usually 2, thin, light bars on the basal half of the caudal fin; these are generally larger fish. The margin of the lower half of the pectoral fin has concentrations of pigment on the membranes such that this area is darker than the rest of the fin. The anal fin is darkest posteriorly where pigment is concentrated on membranes. The distal third of the fin is pigmented to form a dark band, becoming lighter proximally. The dorsal fin is the darkest fin (except for the clear margin) and the anterior base is the darkest part of the fin. Bands are not always evident but pigment spots are large proximally. Some fish have 2, sometimes 3, thin light bars at the base separated by thin dark bars and paralleling the body profile while others have none. The dorsal fin base may have instead a series of lighter spots, sometimes irregular and not paralleling the body profile.

Sizeze

Reaches 5.8 cm total length (Alavi-Yeganeh et al., 2011?check published paper date).

Distribution

This species is restricted to the Kor River basin in Fars.

Zoogeography

As noted in the genus account, Aphanius species in Southwest Asia are regarded as relicts of the Tethys Sea. The mountain populations of A. sophiae (and A. vladykovi) may well have risen with the post-Pliocene uplift of the Zagros Mountains. Since Aphanius species have a low dispersal ability (Kosswig, 1967) it is considered unlikely that these small fishes dispersed from lowland populations into an existing, high mountain range represented today by the Zagros.

Bobek (1963) suggests that there may have been an outflow from the Kor River basin to the Persian Gulf at the south-east corner which was cut off at the end of the Pleistocene by alluvial fans. Krinsley (1970) maintains that any outlet was closed by the late Pliocene. Evidence for the isolation time of Kor River basin tooth-carps is equivocal.

Habitat

Tooth-carps identified as this species has been reported to survive in waters up to 39°C and salinities as high as 130‰. In aquaria lives at 5-37°C. It is found in springs, streams and small pools and lakes in fresh and saline waters.

Age and growth

Unknown. Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 64 fish measuring 2.18-4.64 cm standard length. The a-value was 0.0343 and the b-value 2.787 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases). Alavi-Yeganeh et al. (2011?check published paper date) give length-weight data on this species from six localities with a (intercept) being 0.0087and b (slope) being 3.333 for all localities and sexes combined.

Food

Unknown in detail.

Reproduction

Unknown. Incubation is said to be 10 days at 25°C but this is a general citation for Aphanius spp. in aquaria.

Parasites and predators

Unknown.

Economic importance

This species has importance as an aquarium species, being colourful and hardy.

Conservation

Construction of irrigation canals and drying of some springs because of man-induced changes in the water table may threaten populations of this species.

Further work

The distribution of this species and its population numbers should be documented by field work. Molecular and chromosomal techniques would prove useful information for relating this species with others from neighbouring basins and throughout central Iran. Its biology is poorly known under natural conditions.

Sources

Type material: See above, Lebias sophiae (NMW 14496, 22616-22623, 60327, 68283, 75067, 14760 and 15056, Lebias crystallodon (NMW 15175), and Lebias punctatus (NMW 76509, 15070, 15156, 59837 and 59609).

Iranian material:- CMNFI 1979-0025, 1, 31.6 mm standard length, Fars, Kor River at Marv Dasht (29º51'N, 52º46'30"E); CMNFI 1979-0059, 30, 18.1-39.3 mm standard length, Fars, Pulvar River 8 km south of Sivand (30º01'30"N, 52º57'E); CMNFI 1979-0061, 3, 23.2-32.1 mm standard length, Fars, stream tributary to Pulvar River (30º04'N, 53º01'E); CMNFI 1979-0062, 6, 24.0-42.0 mm standard length, Fars, spring 17 km south of Sa'adatabad (30º05'N, 53º00'E); CMNFI 1979-0067, 1, 25.7 mm standard length, Fars, qanat at Zarqan (ca. 29º46'N, ca. 52º43'E); CMNFI 1979-0071, 6, ? mm standard length, Fars, qanat stream on road to Ramjerdi (ca. 30º00'N, ca. 52º38'E); CMNFI 1979-0091, 3, 23.2-32.1 mm standard length, Fars, stream tributary to Pulvar River (30º04'N, 53º01'E); CMNFI 1979-0117, , mm standard length, ; fish? CMNFI 1979-0155, 2, ? mm standard length, Fars, spring at Gavonoo (28º47'N, 54º22'E);ID? CMNFI 1979-0292, 6, 26.5-34.4 mm standard length, Fars, Lapu'i spring near Zarqan (29º48'N, 52º39'E); CMNFI 1979-0342, 1, 39.4 mm standard length, Fars, Kor River at Band-e Amir (29º49'N, 52º51'E); CMNFI 1979-0498, 6, 17.4-25.2 mm standard length, Fars, spring in Kor River basin (30º05'N, 52º27'E); CMNFI 2004-0003 (GenBank AY593483, AY593492), 47, ?20.7-28.8 mm standard length, Fars, Abdolmahdi Spring (30º06'12"N, 52º58'38"E); CMNFI 2004-0004 (GenBank AY593482, AY593492), 38, ?19.5-34.0 mm standard length, Fars, Malasskuh Spring (29º52'04"N, 52º29'20"E); uncatalogued (GenBank AY593481, AY593490), 6, 25.7-37.0 mm standard length, Fars, Dolatabad Spring (29º43'05"N, 52º50'11"E);

Aphanius vladykovi
Coad, 1988

Male

Female

Common names

gour-e khar, mahi-e gour-e khari (= "striped donkey" or zebra fish), kopurdandan-e Zagros (= Zagros tooth-carp), mahi parchami (= flag fish).

[Zagros pupfish].

Systematics

The type locality is in the Shahrestan-e Bakhtiari va Chahar Mahall in a large pool, 3 km west of Boldaji at 31°57'N, 51°01'E. The male holotype, 36.6 mm standard length, is in the Canadian Museum of Nature, Ottawa under CMNFI 1979-0247 with 35 male and 16 female paratypes from the same locality under CMNFI 1979-0247A and 1 male paratype from a stream 3 km east of Boldaji at 31°55'N, 51°05'E under CMNFI 1979-0248. Villwock (2004) has used cross-breeding experiments that demonstrate this species, A. sophiae and A. dispar are genetically isolated and distinct species.

Key characters

This species is distinguished from all other Aphanius in Iran and Southwest Asia by the higher number of normal-sized lateral line scales (36-47). Other members of Aphanius, and the related Kosswigichthys, have 35, usually 30, or less in the lateral series. The high number of scales is matched only by the genus Anatolichthys Kosswig and Sozer, 1945 of southwest Anatolia. Anatolichthys is polymorphic for scale counts which reach as high as 55 and shows scale loss, perhaps associated with the bitter, salty lakes in which it lives (Coad, 1988i). Male dorsal fin colour is much darker and the anal fin is light compared to A. sophiae, which most closely resembles A. vladykovi. Females lack the typical large spot at the base of the caudal fin found in A. sophiae.

Morphology

Scales are relatively small and numerous over the whole body as exemplified by scale counts. Scales are regularly arranged, embedded and imbricate. Anterior flank scales are a vertical oval with circuli and radii restricted to the posterior field. Numbers of circuli and radii are body and scale dependent and so are fewer in this species than in A. sophiae. There are up to 21 teeth in the upper jaw and teeth are in a single row in each jaw. Gill rakers are short, just reaching the adjacent raker when appressed. All fins are rounded distally and the anal fin is enclosed by a fleshy sheath, most apparent in large females but is also present in males. The gut has a single large loop. Cephalic sensory pores are reduced to a series of neuromasts. Keivany (2003) gives details of cephalic osteology. Pazooki et al. (2008) and Mardani Karani et al. (2008)  examined interlocality morphological variation, finding one of four populations to be more distinctive and 55 of 72 characters different respectively. Esmaeili et al. (2009) give a chromosome number of 2n = 48 and a karyotype of 8 submetacentric and 40 subtelocentric chromosomes with an arm number of 28.

Lateral line scales 36(2), 37(8), 38(15), 39(6), 40(10), 41(6), 42(3), 43(1), 46(1) or 47 (1). Scales above the lateral line 6-9, scales between lateral line and the anal fin 8-12, scales between the lateral line and the pelvic fin 11-15, and scales around the caudal peduncle 20-25.

Total dorsal fin rays 11(4), 12(21), 13(27) or 14(1), total anal fin rays 11(13), 12(25) or 13(15), total pectoral fin rays 14(4), 15(18), 16(25) or 17(6), total pelvic fin rays 5(9), 6(42), 7(1) or 8(1), total gill rakers 10(8), 11(41), or 12(4), and total vertebrae 27(2), 28(24), 29(26) or 30(1). Saefali (1999) records 8-11 gill rakers, 9-14 anal rays, and 9-15 dorsal rays.

Sexual dimorphism

Colour is sexually dimorphic and is outlined below. Ten of 21 morphometric characters are significantly different between the sexes including head length, head and body depths, fin lengths, and caudal peduncle shape (Coad, 1998i). Females attain larger sizes than males (76 mm versus 58 mm total length (Keivany and Soofiani, 2004a; b)).

Colour

Live adult males bear creamy bars on yellowish flanks. The pelvic fins and the edge of the caudal fin are yellow, the anal fin has an orange-yellow edge and the pectoral fins are orange-yellow. The dorsal fin is white with a wide blue-black band in the centre and a narrow blue-black band at the base. In some males the basal band is absent. Live adult females have a slight bluish tinge to their flanks and flank spots are brown.

In preserved specimens, the dorsal fin bands are dark and remain darker in preservative than the pigmentation seen in Aphanius sophiae which is more diffuse, tending to several horizontal rows of speckles, and which is not formed into a strongly distinct band or bands. Other fins are lightly pigmented, being finely speckled with melanophores with the dorsal and pectoral fins the darkest. The anal fin is fleshy and not hyaline but pigmentation is light and not dark as in A. sophiae. There is some tendency to formation of one or two narrow bars on the anal fin, paralleling the base, but these are weakly developed or absent in most fish. A mid-dorsal stripe is not apparent, perhaps because the dorsal flanks and back are darker than in females. Flank bars are light alternating with darker broad bars. The lighter bars become broader posteriorly. There is a light, wide bar at the base of the caudal fin which is not included in the count of male flank bars. Male flank bars 9(3), 10(8), 11(8), 12(10), 13(3), or 14(5). The sides of the head and upper and lower jaws are pigmented with scattered melanophores while the underside of the head is not. The belly is also free of extensive melanophores.

The most characteristic feature of females is the large number of scattered spots on the flank, extending from behind the head to the base of the tail. These spots do not extend to the dorsal flank or the back. The distinctive, large and often lozenge-shaped spot on the central caudal fin base of A. sophiae is absent. The upper back and flank are more heavily pigmented than the flanks (except for the flank spots). The lower surface of the head and the belly are melanophore free as in males and the sides of the head and the upper and lower jaws are heavily pigmented. The dorsal fin is finely pigmented on the rays and membranes and is generally darker than the caudal fin which is also finely pigmented. The other fins have very little pigment but all are fleshy and not hyaline. There is some suggestion of a thin horizontal stripe near the base of the dorsal fin in the largest specimens but this is never well developed.

Teeth are hyaline or discoloured with brown in patches. The peritoneum is dark brown.

Size

Reaches 7.6 cm (Keivany and Soofiani, 2004b).

Distribution

This species is found near Boldaji and in the Chaghakor Wetland in the upper reaches of the Karun River basin. Abdoli (2000) indicates its presence on a map in the upper Marun River and the upper Khersan River (the latter in the Karun basin).

Zoogeography

The origins of this mountain species have been discussed above under the account for A. sophiae.

It is found only in the uppermost reaches of the Karun River basin which drains to the head of the Persian Gulf. In this locality it is isolated from populations of Aphanius in the lowlands by a series of tangs which block gene flow. Oberlander (1965) in studying the origin of Zagros drainage patterns found that streams did not follow the line of least resistance but often incise the higher parts of mountain ridges which are anticlines composed of Miocene Asmari limestone. This situation arises because the Zagros ridges were eroded by a superposed drainage pattern already antecedent to the exhumation of overlying sediments of later Miocene marls and evaporites. The trend of the Zagros ridges is in a NW-SE direction and streams often cross this direction at right angles. This has resulted in the development of the tangs, narrow clefts in the Zagros ridges with very swift waters, which may be up to 2400 m deep with vertical sides rising from the water surface for 300 m, effective barriers to small fishes.

Habitat

This species is found high in the Zagros Mountains (the type locality lies at an altitude of about 2380 m). It is found in small pools, streams and marshes, in fresh water. The type locality is an artificially dammed pool about 300 m wide with cloudy water and a mud and pebble bottom. Capture depth was 40 cm. The shore was grassy and the pool contained large amounts of Myriophyllum and marginal rushes. At 1515 hours on a warm, windy and sunny day, water temperature was 29°C, pH was 6.5 and conductivity was 0.2 mS. A second locality in Coad (1988i) was a mud-bottomed stream 3-4 m wide with pools up to 1 m deep. At 1545 hours water temperature was 22°C, pH was 6.2 and conductivity was 0.45 mS. Current was slow and the stream had moderate amounts of Myriophyllum. Keivany and Soofiani (2004b) reporting on the habitat at the Madar-Dokhtar spring near Gandoman, found water conductivity of 240-280 μS, pH 6.9-8.5 and total dissolved solids125-138 mg/l. The bottom was muddy, interspersed with pebbles, and water is clear (but cloudy in the river). Myriophyllum and Potamogeton were used for spawning. Fish survive 30‰ and 30°C in aquaria.

Pazooki et al. (2008) found females to be more numerous than males, and the sexes lived in mixed schools. Occasionally larger males would chase smaller males but then both would rejoin the school. Males are territorial and defended against fish and beetles up to twice their size. The species preferred areas free from vegetation. These authors found the species easily tolerated to 33°C in aquaria for long periods, and the optimum temperature for the species is probably 21-24°C.

A mass mortality of this species was observed by A. Abdoli in late March or early April 1994 at Chagh-khor wetland near Boldaji.

Bagheri (1999) describes the macrofauna and environment of the Chaghakhour Lagoon where this species occurs.

Age and growth

Keivany and Soofiani (2004b) show maximum age is 2+ years. Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 319 fish measuring 1.46-4.64 cm standard length. The a-value was 0.0309 and the b-value 3.062 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases). Alavi-Yeganeh et al. (2011?check published paper date) give length-weight data on this species from four localities with a (intercept) being 0.0087and b (slope) being 3.414 for all localities and sexes combined.

Food

Abdoli (2000) reports Daphnia and Gammaridae as food. The main food in the Madar-Dokhtar spring is Gammarus, but a specimen from a river also had some algae, Cyclops sp., snail eggs and fish larvae. Mouth structure would lead to the conclusion that this species is a surface feeder but it concentrated on the dominant crustacean in the Madar-Dokhtar spring (Keivany and Soofiani, 2004b).  Pazooki et al. (2008) found chironomids, dipterans, Diaptomus, Daphnia, diatoms and filamentous algae in guts and considered this species to prefer an animal diet.

Reproduction

Mature males chase the female in aquaria, pushing her into a corner, and showing pectoral fin flipping and shivering. Eggs were laid in a batch of 3-10. Absolute fecundity in the Madar-Dokhtar spring examined by Keivany and Soofiani (2004b) was 220-650 eggs, relative fecundity was 45-155 per gram body mass and working fecundity (free and ripe eggs) was 110 eggs absolute and 73 relative. Size of ripe eggs was 0.8-1.2 mm. Eggs are adhesive and attch to plants in small patches, from 3 to 30. Early April was the spawning peak. Eggs hatched at 9-13 days at 21-22ºC. Young reached 11.0-27.2 mm after three months in an aquarium.

Parasites and predators

Barzegar et al. (2004) examined this species for parasites in fish from the Beheshtabad river in Chahar Mahall va Bakhtiari Province and found Lernaea cyprinacea. Keivany and Soofiani (2004b) found it to be susceptible to trichodiniasis and ichthyophthiriasis in aquaria. Barzegar et al. (2008) record eye parasites from this fish including the digeneans Diplostomum spathaceum, Tylodelphys clavata and Ornithodiplostomum sp. Raissy (2008) also investigated parasites in this species from the Gandoman Lagoon. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Lernaea cyprinacea on this species.

Economic importance

This pretty little fish has great potential as an aquarium species (Coad and Keivany, 1998; Keivany and Soofiani, 2004b). It has been maintained in an aquarium for several months, tolerating temperatures of 30°C but probably favouring 20-22°C. pH values were 8-9. Water changes with their associated variations in conditions had no affect on the fish. The pupfish survived 3-4 hours longer than a Capoeta species when the latter was treated with formalin in a shared tank for a Saprolegnia infection and the tank was inadvertently left for 6-8 hours without a water change. Male pupfish are very aggressive and frequently attack other fishes of the same size or even larger. They ate all the coexisting guppy larvae in the aquarium, even those aged 2-3 weeks. The pupfish was fed crushed, dried freshwater shrimp and sometimes live shrimp although they had difficulty handling live shrimp. They were also observed biting and destroying vegetation in the aquarium, although it is unclear whether they actually ate any. It is now available in aquarium stores in Germany (Oliver Lucanus, pers. comm., 23 January 2004) and is kept during summer months in garden ponds, reproducing there successfully (Thomas Schulz, pers. comm., 14 January 2005).

Conservation

The distribution of the species is apparently quite limited, in waters neighbouring the towns of Boldaji and Shahr-e Kord. This should be confirmed by extensive field work in this poorly explored area of the Zagros Mountains. A limited distribution renders this species liable to significant loss or even extinction if habitats are disturbed or destroyed. The type locality, it should be noted, is an artificially dammed pond.

Significantly, a mass mortality of this species was observed by A. Abdoli in late March or early April 1994 at Chagha Khur wetland although the cause is unknown. M. Raissy (pers. comm., 1 July 2009) reports parasitic infections (7 species, including the digenean Ornithodiplostomum in most organs) as a cause for depopulation in Choghakhor (Chagha Kur) lagoon.

An additional threat is escapes of cultured rainbow trout (Oncorhynchus mykiss). Trout culture is being rapidly developed in this region and the trout are known predators on native fishes.

Further work

The distribution of this species in the Zagros needs to be clarified.

Sources

Type material: See above, CMNFI 1979-0247, CMNFI 1979-0247A and CMNFI 1979-0248.

Iranian material: CMNFI 2004-0005, ?, ?, mm standard length, ; uncatalogued (GenBank AY593486, AY593495), 4, 25.6-37.0 mm standard length, Chahar Mahall va Bakhtiari, Ebrahimabad (31º52'30"N, 51º10'10"E); uncatalogued (GenBank AY593487, AY593496), 2, 21.5-28.7 mm standard length, Chahar Mahall va Bakhtiari, Taqanak Bridge (32º12'35"N, 50º49'29"E); uncatalogued (GenBank AY593485, AY593494), 5, 31.0-49.8 mm standard length, Chahar Mahall va Bakhtiari, Madar-Dokhtar spring (31º52'12"N, 51º08'29"E).

Poeciliidae

Poeciliids, including the livebearers, are found in fresh and brackish waters from the eastern United States south through the Caribbean to northeastern Argentina and in Africa. There are about 41 genera with about 343 species (Nelson, 2006; Eschmeyer and Fong, 2011) and one or two species of livebearer has been widely introduced around the world as a control agent for malaria and is found in Iran, along with a released aquarium species.

This family is characterised by an anteriorly rounded and posteriorly compressed body and a depressed head; large cycloid scales on the head and body; supraorbital pores with neuromasts in fleshy grooves; a small, terminal mouth with the lower jaw projecting; teeth incisor-like or villiform on the jaws; gill membranes free from the isthmus; 4-6 branchiostegals; no spiny fin rays; a small dorsal fin; pectoral fins set high on the side (in contrast to Cyprinodontidae anterior pelvic fins; no adipose fin; and the anal fin in males is developed into a copulatory organ with rays 3-5 modified while females have a rounded anal fin (in livebearers only, the species found in Iran, in contrast to Cyprinodontidae).

The gonopodium is used for transferring sperm packets into the female. It is moveable to the side or forward to allow copulation to occur. The sperm packets release sperm when placed in the female and some can be stored for future use. Details of gonopodium anatomy are important in identifying and classifying species. The gonopodium is composed primarily of the third, fourth and fifth anal rays with various hooks, serrae and spines. Young are born alive, a condition known as ovoviviparity where the eggs develop and hatch in the mother.

Livebearers are found in habitats from mountain streams to brackish coastal marshes and river mouths. Food is mostly encrusting algae and the associated, small invertebrates. Males tend to be smaller than females and more brightly coloured. These colours are best seen during the courtship display.

Some species are important in the aquarium business, such as the guppy and swordtail, while the mosquitofish has been used world-wide as a predator on aquatic mosquito larvae, the adult fly being a carrier of malaria. They have also been used extensively in genetics research, research on tumours and in immunology. Some livebearers are all-female species and egg development is stimulated by spermatozoa from another species, without any genetic contribution. Young are identical to the mother. This unusual form of reproduction is called gynogenesis. Others have superfetation where eggs are at different developmental stages within the mother and are born over a period of several days rather than all at once.

Genus Gambusia
Poey, 1854

This genus is characterised by large scales, short dorsal and anal fins, the anal fin in males anteriorly placed and modified into a long intromittent organ (or gonopodium), dorsal fin inserted behind the anal fin origin (in contrast to Aphanius, Cyprinodontidae), and dorsal and caudal fins are spotted.

Gambusia affinis
(Baird and Girard, 1853)

Gambusia holbrooki was considered to be a subspecies of this species. Iranian mosquitofish were G. holbrooki but it is possible that some populations consist of G. affinis although none were seen by me or by Holčík and Oláh (1992). Shakirova and Sukhanova (1994) report this species from the Atrek River in Turkmenistan, which flows into Iran, as well as Kopetdag streams on the northern border of Iran. However identification of these taxa may be confused. The source of most (? all) Middle East mosquitofish is Europe and Kottelat and Freyhof (2007) record that there is no confirmed presence of G. affinis in Europe.

Duane Raver, U.S. Fish and Wildlife Service
Duane Raver, U.S. Fish and Wildlife Service

Gambusia holbrooki
Girard, 1859

Common names

گامبوزيا (= gambusia).

[gambuzi, zoory, or zurri in Arabic; gambuziya in Azerbaijan; gambuziya in Russian; mosquitofish; eastern mosquitofish (for holbrooki, western mosquitofish for affinis); plague minnow, in reference to its ecological impact].

Systematics

Gambusia holbrooki was originally described from eastern Florida and South Carolina, U.S.A. Wooten et al. (1988) distinguish the eastern and western mosquitofish in the U.S.A. based on morphology, biochemistry and distribution. However, extensive hybridization occurs in the native habitat and the composition of all the introduced Iranian populations has not been studied. Introduced populations world-wide are generally referred to the taxon holbrooki.

Key characters

Males are easily recognised by the anal fin rays 3-5 being specially modified into an elongate gonopodium for intromittent fertilisation. G. holbrooki has 8 dorsal and 11 anal total fin rays while G. affinis has 7 dorsal and 10 anal total fin rays (Walters and Freeman, 2000).

Morphology

Dorsal fin with 5-9 rays, usually 7 (with the last two counted as one), anal fin with 7-11 rays, usually 10, and pectoral fin with 11-14 rays. Lateral scale rows number 26-33. In males, the posterior edge of the joints of the first elongate ray is serrated (smooth in affinis), the cusps of the posterior branch of the second elongate ray are short and almost straight (long and curved in affinis), and the apical hook of the posterior branch of this ray is short with 2-3 joints (very long with 4-6 joints). Vertebrae 28-34. The karyotype is 2n=48. Detailed counts on Iranian specimens were not made; examination of cleared and stained material indicates that the species in Iran and Iraq is G. holbrooki.

Sexual dimorphism

Males reach a smaller adult size than females (see below). Males have the end of the anal fin base well ahead of the beginning of the dorsal fin. Females lack a gonopodium and the end of the anal fin is under the beginning of the dorsal fin. In addition various morphometric characters differ widely (see Abdurakhmanov (1962) for details).

Colour

Males are translucent grey to light olive with a blue, green or purplish sheen on the sides and opercle. The back is olive-brown to yellowish-brown and the belly silvery or yellowish. A dark bar passes through the eye. The iris has a purple sheen. The flanks may appear spotted as pigment margins the scales to form a diamond pattern on the body. Dorsal and caudal fins are spotted and a dusky, light tan but other fins are clear. Adult females have a large, triangular, bluish-purple blotch on the lower flank behind the pectoral fin (called the gravid spot). The black peritoneum can be seen through the body wall.

Size

Reaches 63.0 mm in females and 45.4 mm in males (Tabibzadeh et al., 1970a; Vargas and de Sostoa, 1996), perhaps to 8 cm in the largest females (Reshetnikov, 2002).

Distribution

The natural distribution is from New Jersey southward to northern Mexico but it has been introduced to all continents except Antarctica. This species was first introduced to the Ghazian marshes of the Caspian littoral of Iran in 1922-1930. Some mosquitofish may have spread from the Lenkoran District of Azerbaijan, reaching the Safid Rud in 1937 (Shukolyukov, 1949). From the Ghazian stock, additional samples were collected in 1966 for introduction around the country via raising ponds to over 3000 permanent water bodies (Tabibzadeh et al., 1970a, 1970b; Spillman, 1972; Emadi, 1996c). Many of these contained fishes but some were previously fishless or at least now appear to contain only mosquitofish, e.g. springs at Kahurak and Hormak in Sistan and Baluchestan Province. Over 1.5 million fish were distributed in 1969 alone (Tabibzadeh et al., 1970b). This species is now the most widely distributed fish in Iran, found in all basins (Abbasi et al., 1999; Kiabi et al., 1999; Abdoli, 2000).

Zoogeography

This species is an exotic, originating in eastern and southern North America and introduced to Iran from stocks introduced to Italy and from Baku in Azerbaijan.

Habitat

Mosquitofish are normally inhabitants of clear and weedy streams and ditches, weedy margins of large rivers and lakes, marshes and brackish coastal lagoons, usually over mud or sand bottoms. They prefer more open waters with less vegetation than Aphanius dispar, for example. Males and females in aquaria swam together in a mixed school in contrast to the sexually segregated schools of Aphanius dispar (Al-Daham et al., 1977). In the Anzali Mordab, dense schools are found in surface waters of areas covered by submerged and floating vegetation while deeper water is fishless (Holčík and Oláh, 1992). In Iran, it is one of two most abundant species in Caspian wetland areas along with Carassius auratus (Iranian Fisheries Research and Training Organization Newsletter, 19:4, 1998).

Al-Daham and Bhatti (1977) found G. affinis, probably this species, to have a complete tolerance of 10.25‰ sea water and 90% of fish survived 20.5‰ for 24 hours and even as high as 58‰. This species has been reported to survive in waters up to 42°C and as low as 0.5°C. Summer air temperatures of 45°C and winter temperatures several degrees below zero are survived as long as the water is deep enough (Tabibzadeh et al., 1970a). Upper and lower thermal tolerance increases with body size (Al-Habbib and Yacoob, 1993). At temperatures below 15-18°C, mosquitofish grow but do not mature or breed. They prefer a temperature of 31°C.

Muddy and polluted conditions, acid to alkaline water, and dissolved oxygen less than 1 mg/l are survived by this species. pH range survived is 4.46-10.2 and dissolved oxygen levels as low as 0.2-0.4 mg/l as long as water surface access is available. They can take advantage of the oxygen rich surface layer as they position their bodies with flattened head and back immediately adjacent to t\he air-water interface (Lewis, 1970).

Their success in Iran and other waters world-wide is attributed to the following factors according to Meffe and Snelson (1989):- 1) abundant in original range, 2) polyphagous, 3) short generation time, 4) a single female can colonise since livebearer, 5) broad physiological tolerances, 6) closely associated with man, 7) high genetic variability, 8) specialized reproduction with moderate numbers of advanced young several times a year, and 9) high aggression levels.

Age and growth

Numbers of each sex in a population vary between localities, mostly females predominate over males. Males are more sensitive to temperature extremes, starving and overcrowding and this will affect such things as size and age at maturity. Males do not grow much after sexual maturity is attained but females have indeterminate growth, although growth slows as energy is put into egg production. The average condition factor for males was 1.2369 and for females 1.0906 in southern Iraq. The condition factor slowly decreases as length increases in the male and vice versa for females (Huq et al. 1977). Sexual maturity can be reached in only 2-3 months but such fish die before winter. Minimal size for maturity is 16-28 mm total length. In the laboratory under ideal conditions sexual maturity can be attained in 3 weeks. Life span seldom exceeds 15 months, although the range is 6-24+ months depending on when in the spawning season the fish was born. In Spain, maximum ages are 2+ for females and 0+ for males (Vargas and de Sostoa, 1996). Fish born late in the year may not mature until the following year.

Growth to maturity can be so rapid that a 50-fold increase in numbers can occur over a 10-week period. Densities may reach over half a million fish per hectare. Under ideal conditions it is theoretically possible for 10 pregnant females to produce 5 million fish in 6 months.

Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 35 Iranian fish measuring 1.98-3.27 cm standard length. The a-value was 0.0190 and the b-value 3.214 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases).

Patimar et al. (2011) evaluated fish from the Tajan River in the Caspian Sea basin of Iran and found maximum observed ages of 0+ for males and 1+ for females. Negative allometric growth occurred in males (b = 2.442) and positive allometric growth in females (b = 3.232), males having a lower fitness therefore. Sedaghat and Hoseini (2012a) examined fish from the Dinor River, Kermanshah and found positive allometric growth for females where W = 0.06L3.49 and isometry for males here  W = 0.05L2.87.

Food

Generally this species is regarded as an opportunistic omnivore. Swanson et al. (1996) refer to this fish as an aggressive predator and Al-Daham et al. (1970) for southern Iraqi fish considered it to be a carnivore, feeding on aquatic and terrestrial insects including spiders, ants, beetles and mites, and also filamentous algae. Females are cannibals. Food includes diatoms, algae, worms, crustaceans including zooplankton, insects, snails, other fishes and amphibian larvae. The mosquitofish is dependent on sight to detect and attack prey. Selection of zooplankton can lead to an increase in phytoplankton numbers (and thus affect water quality) and to changes in size frequencies of zooplankton populations.

Food is taken at or near the surface and by grazing on plants and rocks. Tabibzadeh et al. (1970a, 1970b) record diet as molluscs, aquatic insects and various forms of mosquito larvae. They report up to 94 pupae or 104 mosquito fourth-stage larvae eaten per day by this fish and effective clearance of springs and marshes. Large-sized mosquitofish took more than 500 second instar mosquito larvae per day and medium- and large-sized mosquitofish consumed 22-64 fourth instar larvae per day in an Iraqi study under laboratory conditions (Mohsen et al., 1989). Abdoli (2000) lists Diptera and Chironomidae as food items. Meffe and Snelson (1989) report predatory behaviour on the young of other species of fish and females attack other fishes, shredding fins and sometimes causing mortality.

Reproduction

Patimar et al. (2011) found Iranian Caspian fish matured between February and July with the highest gonadosomatic index in May. Egg diameters reached 3.0 mm and absolute fecundity varied from 7 to 57 eggs. Females dominated in the sex ratio (1:2.15). An increase in female size was correlated with a decrease in relative fecundity.

Na'ama and Al-Hassan (1989) report G. affinis (presumably G. holbrooki) to have a brood size up to 48 from Baghdad specimens and 51 from the Basrah area, Iraq. The smallest reproductive female was 1.7 cm total length. Weight is the most accurate predictor of brood size. In Iran, Tabibzadeh et al. (1970a) found that females mature and reproduce within 2 months at temperatures above 15°C. In less favourable conditions this may take 8-10 months. A minimum photoperiod of 12.5-13 hours is necessary to stimulate reproduction although year-round temperatures above 20°C can offset this light requirement. About 17 days after fertilisation, the female gives birth to as many as 428 live fish over a period of about 1 month after 3-8 weeks gestation. Each fertilisation can give 2-3 broods and each female can produce up to 9 broods during her life, although 2-5 is more usual.

The breeding season extends from April to November in favourable conditions, May to September in more temperate conditions. Several months pass between successive spawnings. Eggs are up to 1.8 mm in diameter when mature and embryos are about 6-8 mm at birth. The young are protected within the female and are independent when born, with resulting low mortality. Environmental breeding requirements are simple and with low mortality results in the successful spread of this exotic.

The small male approaches the female from behind and with a rapid motion inserts his gonopodium tip into the female. The gravid spot is a releasing stimulus. It indicates a receptive female and also is a target for the male gonopodium. Sperm are transferred in a spermatophore. During mating the gonopodium is angled forward at 140-150°.

Most males copulate without courtship. When courtship occurs it involves the male assuming an s-shaped position with its body and vibrating in front of, or at the side of, the female. Males are aggressive and dominate smaller males to restrict access to females. Females may be inseminated by several males and sperm can be stored to be used up to 10 months later. A single transplanted female can populate a new habitat.

Parasites and predators

Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Lernaea sp. on this species (as G. affinis).

Economic importance

Introduced widely to combat malaria by consuming the aquatic larvae and pupae of the carrier mosquito. However to be effective, this fish must have aquatic plants cleared from its habitat. The mosquitofish is also a predator on the eggs and young of other, native fishes and a competitor for food, and it alters ecosystems by greatly reducing rotifer, crustacean and insect populations allowing phytoplankton to increase dramatically (Myers, 1965; Hurlbert et al., 1972; Tabibzadeh et al., 1970a; Edrissian, 2006; Chandra et al., 2008). Native species might well be more effective predators on mosquito larvae (see account of Aphanius dispar and Ahmed et al. (1988) and Beidas and White (1982)). Muhaisen et al. (1985) studied control of this species in Iraq as larvae on fish farms and found potassium permanganate, copper sulphate, acriflavine and malachite green to be effective.

Haas and Pal (1984) point out that Gambusia may favour schistosomiasis, a human parasite, by altering the ecology of freshwaters, perhaps by reducing the numbers of natural predators on vectors of this parasite. They also note that it is considered a pest by fish farmers.

Tabibzadeh et al. (1970a, 1970b), Motabar (1978) and Swanson et al. (1996) give methods for raising and distributing mosquitofish. Also used as a research species and sometimes seen in aquaria. Al-Nasiri and Sharma (1978) used this species in Iraq to study the toxicity of the agricultural insecticide Ekatin 25 to fish and Mohsen et al. (1989) studied the predatory efficiency and tolerance to mosquito larvicides.

Conservation

The most widespread fish in Iran and, as an exotic, not in need of any protection. However this distribution undoubtedly affects the conservation status of native species since mosquitofish have a strong impact in small and non-diverse habitats such as the small springs found throughout Iran. Muus and Dahlstrøm (1999) mention that it tends to oust native toothcarps.

Further work

The effects of this species in competition with native fishes should be investigated although it is so widespread that it could never be eradicated.

Sources

Meffe and Snelson (1989) give the ecology and evolution of the family and Rauchenberger (1989) the systematics of the genus. Swanson et al. (1996) and Pyke (2005, 2008) review biology, culture and mosquito control and are a source for general information above.

Iranian material: The following is a sub-sample of Iranian material as this species is very common and material extensive (see map for distributional information): CMNFI 1970-0515, 2, 20.6-30.2 mm standard length, Gilan, Shafa River near estuary (37º35'N, 49º09'E); CMNFI 1970-0519, 5, 14.1-19.5 mm standard length, Gilan, Chelvand River (ca. 38º18'N, ca. 48º52'E); CMNFI 1970-0590, 3, 25.1-29.2 mm standard length, Mazandaran, Shesh Deh River near Babol Sar (ca. 36º43'N, ca. 52º39'E); CMNFI 1971-0343, 1, 26.0 mm standard length, Gilan, Langarud at Chemkhaleh (37º13'N, 50º16'E); CMNFI 1979-0019, 26, 15.4-29.0 mm standard length, Fars, Pol-e Fasa (29º29'N, 52º38'30"E); CMNFI 1979-0025, 2, 28.7-33.8 mm standard length, Fars, Kor River at Marv Dasht (29º51'N, 52º46'30"E); CMNFI 1979-0026, 1, 21.6 mm standard length, Fars, Shapur River at Shapur (29º47'N, 51º35'E); CMNFI 1979-0126, 53, 14.9-33.1 mm standard length, Fars, Dasht-e Arjan springs (29º37'N, 51º58'E); CMNFI 1979-0173, 13, 24.7-39.6 mm standard length, Hormozgan, qanat at Hajjiabad (28º19'N, 55º54'E); CMNFI 1979-0192, 2, 24.9-26.4 mm standard length, Fars, qanat 2 km east of Rostaq (28º26'30"N, 55º04'E); CMNFI 1979-0226, 65, 23.1-39.2 mm standard length, Sistan, 3 km east of Kuh-e Khajeh (30º57'N, 61º17'E); CMNFI 1979-0230, 36, 7.2-44.4 mm standard length, Sistan, Jehil-e Puzak (ca. 31º15'N, ca. 61º42'E); CMNFI 1979-0283, 1, 35.9 mm standard length, Kermanshahan, Qareh Su basin (34º21'N, 47º07'E); CMNFI 1979-0288, 7, 26.8-29.4 mm standard length, Ilam and Poshtkuh, Gangir River (33º50'N, 46º18'E); CMNFI 1979-0309, 5, 15.4-32.8 mm standard length, Kerman, Fahraj River (28º57'N, 58º42'E); CMNFI 1979-0328, 2, 24.3-24.8 mm standard length, Baluchestan, jube near Bampur (27º10'30"N, 60º21'E); CMNFI 1979-0360, 13, 17.8-25.9 mm standard length, Khuzestan, Karkheh River canal (31º40'N, 48º35'E); CMNFI 1979-0478, 2, 17.8-28.2 mm standard length, Mazandaran, ditch 29 km from Eimar (37º09'N, 54º36'E); CMNFI 1979-0498, 2, 22.1-35.8 mm standard length, Fars, spring on road to Dariush Dam (30º05'N, 52º27'E); CMNFI 1980-0122, 25, 14.3-26.8 mm standard length, Mazandaran, Nerissi River (36º38'N, 52º16'E); CMNFI 1980-0128, 7, 23.6-35.6 mm standard length, Mazandaran, Qareh Su (36º49'30"N, 54º03'30"E). CMNFI 1980-0132, 5, 18.4-30.2 mm standard length, Gilan, Safid River at Kisom (37º12'N, 49º54'E); CMNFI 1980-0145, 75, 32.6-40.4 mm standard length, Gilan, Bandar-e Anzali (37º28'N, 49º27'E).

Comparative material: BM(NH) 1974.5.2:38-40, 3, 17.3-25.1 mm standard length, Iraq, Baghdad (33º21'N, 44º25'E); BM(NH) no catalogue number, 6, 10.7-37.5 mm standard length, Iraq, Baghdad Liwa (no other locality data); BM(NH) no catalogue number, 4, 28.3-38.8 mm standard length, Iraq, Baghdad Liwa (no other locality data); BM(NH) no catalogue number, 8, 16.8-32.5 mm standard length, Iraq, Kut Liwa (no other locality data); BM(NH) no catalogue number, 2, 20.3-21.9 mm standard length, Iraq, Hilla Liwa (no other locality data).

Genus Xiphophorus
Heckel, 1848

This genus comprises the platyfishes and swordtails and is found in Mexico and central America with 25 or more species. these are small fishes, 14 cm or less, with several members and hybrids important in the aquarium trade. They are also used in genetic studies. They have a gonopodium in males as in other poeciliids with a unique structure in detail (Rosen, 1960). Male swordtails may have a characteristic elongate lower caudal fin lobe, the "sword" but this is not always developed in all species and all individuals.

Xiphophorus hellerii
Heckel, 1848

Aquarium swordtails from Wikimedia Commons
Aquarium swordtails from Wikimedia Commons

Male and female specimens from Jahrom
Male and female specimens from Jahrom

Common names

dom shamshiri.

[swordtail, green swordtail]

Systematics

The trivial name is often spelled d helleri.

Key characters

The swordtail is characteristic along with details of the gonopodium.

Morphology

Dorsal fin branched rays 11-15, anal fin branched rays 8-10, pectoral fin branched rays 12-13 and pelvic fin branched rays 6. Lateral line scales 26-30. Vertebrae usually 30-31.

The swordtail is medium to large with a long straight caudal appendage. The terminal segment of gonopodial ray 3 is produced into a crescent-shaped hook and blade, pointed distally  Ray 4a curves strongly backward over the blade at an angle greater than 90°. Distal serrae of ray 4p are reduced in size and number, and proximal serrae are rather slender; The terminal segment of ray 5a is produced into a claw, several times larger than the distal serrae of ray 4p.

scale rows along the middle of the body are green to blue; belly is white and there is a s in all populations (regional forms). In a few variants, there can be one or two additional lines above and one or two below this line. In almost all populations, the dorsal fin has spots or flecks of red. This coloration can also appear in the caudal fin. The sword of males is bright yellow and edged more broadly in black below than above. Old fish (3 to 4 years old) can have a totally black sword from the caudal fin all the way to the tip of the sword.

Sexual dimorphism

The male sword may not always develop in aquarium specimens but usually appears at 3-4 weeks of age. Mature females have a gravid spot in front of the anal fin.

Colour

The overall colour is olive-grey to olive-green. There is a light red, dark red dark, violet or dusky brown longitudinal line or rarely a zig-zag stripe on the greenish-yellow flank, and there may be 1-2 reddish stripes above and 1-2 reddish stripes below. Scale rows along the mid-flank are green to blue, edged with brown and net-like in appearance. The dorsal fin is spotted red or brownish in median and below the margin rows. Fins are generally a yellow-green. The belly is white. The sword is a bright yellow or orange-yellow edged more broadly in black below than above. The black lower edge is carried forward onto the caudal peduncle. Fish 3 to 4 years old can have a totally black sword. Aquarium specimens may be red, green, black and albino, and various combinations of these.

Size

Attains 16.0 cm total length, females larger than males.

Distribution

The native distribution is from southern Mexico to Guatemala, Honduras and Belize. Introduced specimens were caught in Iran at Jahrom (28º29'N, 53º31'E) and Kashan 34º04'N, 51º31'E) (Esmaeili et al., 2010).

Zoogeography

An exotic species presumably released from aquaria.

Habitat

In its native habitat this species is found in streams and rivers but can also occur in canals, ponds, lagoons, ditches and in warm springs. The Iranian specimens were caught in a small pool, part of a qanat system, and in a spring. Vegetation can be none to common. The water may be clear to polluted and  current none to moderate. Adults prefer current in streams while the young remain in quieter shore areas. It occurs at both low and relatively high elevations, up to ca. 1500 m. In aquaria this species prefers slightly alkaline, medium hard water and temperatures in the 22-28ºC range.

Age and growth

Sexual maturity is reached after eight to twelve months in some reports, three months in others. Life span exceeds 4 years.

Food

Food is crustaceans, insects, worms and plant matter.

Reproduction

This species is a livebearer and is easy to breed in aquaria. Females can produce more than 180 young at one time, at intervals of about a month. Adult males can be aggressive to other fishes. A well-developed male sword stimulates females to mature and inhibits maturity in younger males. The female swordtail can undergo sex reversal, becoming  male under certain environmental conditions.

Parasites and predators

None recorded in Iran.

Economic importance

None at present in Iran but a possible source of aquarium specimens. It is a popular aquarium species, easy to maintain and is sold world-wide and it has been used in genetic studies. There are various cultivated forms of this species, hybrids with related species. It does adversely affect native species when introduced.

Conservation

None required as an exotic. 

Further work

The biology of this species in Iranian waters, and its effects on native species, requires study. It should be extirpated, if possible.

Sources

General aquarium literature and Esmaeili et alal. (2010).

© Brian W. Coad (www.briancoad.com)