Freshwater Fishes of Iran, Species Accounts - Cyprinidae

Freshwater Fishes of Iran

Species Accounts

Cyprinidae
Garra to Vimba

Revised: 09 November 2010

Genus Garra
Hamilton, 1822

The genus Garra is found throughout Southwest Asia and from Africa to southeast Asia. There are about 73 species and 4 are recognised from Iran. The genera Discognathus Heckel, 1843 and Discognathichthys Bleeker, 1859 are synonyms. Hora (1921) presents anatomical arguments for including Discognathus in Garra.

This genus is characterised by a small to moderate-sized body, elongate and almost cylindrical, a rounded snout with the mouth inferior and crescent-shaped, the lower jaw has a horny edge, the upper lip is usually fringed and continuous with the snout, the lower lip and chin area modified into a suctorial disc with free posterior margin (in Iran, elsewhere a some have a callous pad (Stiassny and Getahun, 2007) and the smallest specimens lack full disc development), the anterior disc margin free or adherent (species with the latter condition were placed in a separate genus, Discognathus or Discognathichthys), 1 or 2 pairs of short barbels (species with the former condition were placed in a separate genus, Discognathus or Discognathichthys), vomero-palatine organ vestigial or regressed, eyes small, usually large scales, lateral line complete, small dorsal and anal fins without thickened rays, pectoral and pelvic fins placed horizontally on the body, first two or more pectoral fin rays prominent and often unbranched, pharyngeal teeth in 3 rows (typically 2,4,5-5,4,2) with hook-shaped tips and spoon-shaped crowns, vent may be midway between pelvic and anal fin bases or nearer the latter, elongate and coiled gut, a black peritoneum, and 2n=50.

The Farsi common name used generally for these fishes is gel cheragh (= mud-eater, mud-grazer), سنگ ليس (sang lis) and ماهي سنگي (mahi sangi), not repeated in each Species Account.

These fishes are found in mountains streams and other flowing waters, maintaining position with their suctorial disc, reduced swimbladder, flattened belly and large, splayed and horizontal paired fins. Also known to occur in slow-moving or stagnant waters. They scrape algae from rocks. These are oily fishes which are eaten in India (Hora, 1956).

Menon (1954) considers that the members of this genus spread westwards along the Himalayas as late as the early Pleistocene. Kosswig (1952) indicates their presence in the Araxes (= Aras) of Turkey but this seems to an error.

Garra persica
Berg, 1913

Common names

Persian stone lapper.

Systematics

This species is recognised only as a subspecies of Garra rufa by Bianco and Banarescu (1982) while Menon (1964) and Karaman (1971) synonymise it with Garra rufa. Karyotype analysis separates this species from Garra rufa (Esmaeili et al., 2009).

The syntype specimens are in the Zoological Institute, St. Petersburg under catalogue numbers ZISP 11707 (6 specimens from the "River Bampur in Eastern Persia. N. Zarudnyi 1898, 15-27.VII") and 11706 (1 specimen from "Kiabad in Zirkuh (Eastern Khorassan). N. Zarudnyi 1898, 3.V") according to Berg (1913) where the original description is founded on these fish, implying all are types. The latter is also given as "settlement Kiabad between Zirkuh province and Sistan" in the catalogue (this locality may be at or near Kuh-e Ziri at 32°48'N, 59°50'E according to Coad (1981d)). These dates are old style and corrected to new in Berg (1949) (27.VII-8.VIII and 15.V respectively). In St. Petersburg under ZISP 11707 there are 10 fish 24.0-46.5 mm standard length and ZISP 11706 is not listed as a type in the catalogue nor in Berg (1949). Berg (1949) lists 10 fish in 11707 too. These specimens were formerly identified as Discognathus lamta by Nikol'skii (1899) who lists 1 fish in 11706 and 6 in 11707. Three syntypes are in the Zoological Survey of India, Calcutta (ZSI F11101/1) listed under Garra rufa obtusa and received from the Zoological Institute, St. Petersburg, Russia on exchange (Menon and Yazdani, 1968). There are more apparent types available than those listed by Berg (1913).

Key characters

Two pairs of barbels are present, the adhesive disc is well developed with a free posterior margin, the dorsal fin has 7 branched rays, and the caudal fin modally 16 branched rays. The caudal fin ray count is unique in cyprinids from Iran and very rare elsewhere. Almost all cyprinids show a strong mode of 17 branched caudal fin rays.

Morphology

Dorsal fin with 4 unbranched and 6-8 branched rays, anal fin with 2-3 unbranched and 4-6 branched rays, branched pectoral fin rays 12-16, pelvic fin branched rays 6-8, caudal fin branched rays number 15-17 with a strong mode at 16, lateral line scales 28-38, total gill rakers 15-22 with lower counts in smaller fish, pharyngeal teeth usually 2,4,5-5,4,2, and total vertebrae 34-35.

Scales are regularly arranged and only lacking on the anterior isthmus in some fish. A pelvic axillary scale is present but is not always well-developed. Scales have numerous radii on all fields with the focus broken up into a network of lines. There are 4 short barbels. Upper lip well-developed, the rostral fold weakly fringed. The adhesive disc and rostral fold are greatly papillose although the disc centre is not as papillose as the margins. The gut is very elongate and greatly coiled. Chromosome number 2n=48 with 15m, 8Sm and 1St (Esmaeili et al., 2009).

Counts for 12 Iranian topotypic specimens from the Bampur River are as follows:- dorsal fin with 4(12) unbranched and 7(12) branched rays; anal fin with 3(12) unbranched and 4(3) or 5(9) branched rays; branched pectoral fin rays 12(1), 13(6) or 14(4); caudal fin branched rays number 15(1), 16(8) or 17(1) in the type series and 15(1), 16(10) or 17(1) in topotypes; lateral line scales 29(2), 30(1), 31(1), 32(2), 33(2), 34(2) or 35(2); total gill rakers 15(6); pharyngeal teeth 2,4,5-5,4,2(5); and total vertebrae 34(9) or 35(3).

Meristic values for topotypes and other material: dorsal fin branched rays 6(3), 7(115) or 8(4); anal fin branched rays 4(5) or 5(118); branched pectoral fin rays 12(8), 13(40), 14(49), 15(20) or 16(5); branched pelvic fin rays 6(9), 7(100) or 8(13); caudal fin branched rays number 15(10), 16(113) or 17(9); lateral line scales 28(1), 29(5), 30(5), 31(77), 32(18), 33(28), 34(20), 35(22), 36(11), 37(4) or 38(2); total gill rakers 15(16), 16(13), 17(18), 18(17), 19(11), 20(4) or 21(2); pharyngeal teeth 2,4,5-5,4,2(5); and total vertebrae 32(4), 33(44), 34(34) or 35(4).

Sexual dimorphism

Males develop numerous breeding tubercles around the snout, between the nostril and the eye and between the nostrils. There is a transverse depression anterior to the nostrils on the snout. A postspawning individual from the Hormuz basin measuring 148.7 mm SL has small tubercles under the eye running forward onto the snout as a band, the most evident tuberculation. The swollen snout tip bears no tubercles. The top of the head has tubercles but these are smaller and sparser than the band under the eye. Scattered large tubercles are present on the gill cover. Evident tubercles line the dorsal, anal, caudal and pectoral fin rays (pelvic fins not present on specimen), the largest being those on the anal fin.

Colour

The back and flanks are an orange-brown to golden-brown. There is a blue spot on the flank near the postero-dorsal edge of the operculum (dark black in preserved fish). The dorsal fin bears elongate blotches on the posterior half of each fin membrane, usually fading distally, but in some fish occupying the whole membrane. Proximally there is a gap between these blotches and 3-5 bars which originate at the posterior edge of the base of branched ray three and succeeding rays, and extend distally across the ray and then along the ray and the membrane to the gap. These bars are much more heavily pigmented than the dorsal blotches. There is a bluish tinge or spot around the pectoral fin base, sometimes developed as a bar along the edge of the gill cleft, becoming dark blue dorsally. There is a dark bar or a roundish, poorly-delimited spot on the caudal peduncle at the base of the caudal fin. Fins are generally pink to light orange. The caudal fin pigmentation is individually variable. Some are blotched irregularly on both rays and membranes, in others there is a trace of a band in mid-fin extending from the dorsal to the ventral margin following the posterior outline of the fin, while others have pigment heavily concentrated only in the mid-fin clear of the margins. The pelvic fin has little or no pigment and the anal fin has a very few irregular light blotches on both rays and membranes. The pectoral fin is pigmented near the dorsal base with some pigment on anterior rays and membranes. In live fish the paired fins are a light orange and other fins show less marked orange tinges. The peritoneum is black.

Size

Attains 7.5 cm total length (Berg, 1913).

Distribution

This species is found in the Hormuz, Makran and Hamun-e Jaz Murian basins and possibly the Sistan and Kerman-Na'in basins (Bianco and Banarescu, 1982; Abdoli, 2000).

Zoogeography

The species of the genus Garra are thought by Menon (1964) to have colonised Iran from a centre of origin in southern China by a series of "waves". The earliest wave arrived in the Miocene and is represented in Iran today by the species rossica and variabilis, characterised by the primitive condition of a weakly-developed adhesive disc without a free posterior border, the posterior chamber of the swimbladder is cylindrical and well-developed, there is no proboscis on the snout and the vent is close to the anal fin. A second wave is represented in Iran by rufa (and by implication persica) characterised by a well-developed disc and a tuberculated snout marked off by a transverse groove. There were 6 "waves" all told but only the first two are relevant to Iran. Karaman (1971) criticises this complex interpretation on two grounds. He maintains that it is unlikely that fishes would immigrate from southern China to Iran but leave no extant forms between these two remote places and that the species assigned to the various waves show no characteristics which would make them more adaptive and capable of replacing earlier wave members. The characters of rossica and variabilis (one pair of barbels, weak disc, reduced squamation) could equally be loss characters and a more recent specialisation rather than the primitive condition.

Habitat

Kiabi and Abdoli (2000) found this species to have the widest altitudinal range in Hormozgan Province.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

None reported from Iran.

Economic importance

None.

Conservation

This species is widely distributed in various smaller water bodies in eastern Iran and does not appear to be under threat.

Further work

The biology of this species remains unknown and would repay study. It is a significant component of a variety of water bodies.

Sources

Type material: See above, ZISP 11707 and 11706.

Iranian material: CMNFI 1979-0138, 1, 25.7 mm standard length, Fars-Hormozgan, Rasul River drainage (ca. 27º32'N, ca. 54º58'30"E); CMNFI 1979-0139, 1, 30.6 mm standard length, Fars-Hormozgan, Rasul River drainage (ca. 27º25'30"N, ca. 54º59'E); CMNFI 1979-0144, 1, 27.3 mm standard length, Hormozgan, Minab River at Minab (27º09'30"N, 57º04'E); CMNFI 1979-0145, 4, 14.8-25.4 mm standard length, Hormozgan, Geru River drainage (26º55'N, 57º01'30"E); CMNFI 1979-0149, 7, 29.0-49.4 mm standard length, Hormozgan, stream north of Bandar Abbas (27º36'N, 56º14'E); CMNFI 1979-0152, 1, 62.2 mm standard length, Hormozgan, Shur River drainage (28º09'N, 55º43'E); CMNFI 1979-0178, 23, 25.1-66.9 mm standard length, Hormozgan, Sarzeh River draiange (27º36'N, 56º15'E); CMNFI 1979-0180, 1, 42.7 mm standard length, Hormozgan, stream 3 km east of Essin (27º19'N, 56º17'30"E); CMNFI 1979-0181, 1, 44.0 mm standard length, Hormozgan, Kul River drainage (27º17'30"N, 56º03'30"E); CMNFI 1979-0186, 8, 30.2-64.6 mm standard length, Hormozgan, stream and pools at Sar Khun (ca. 27º24'30"N, ca. 56º25'E); CMNFI 1979-0187, 9, 32.1-57.9 mm standard length, Hormozgan, stream and pools at Sar Khun (ca. 27º23'30"N, ca. 56º26'E); CMNFI 1979-0312, 10, 26.6-35.6 mm standard length, Baluchestan, dam on Bampur River (27º11'N, 60º46'E); CMNFI 1979-0315, 1, 23.8 mm standard length, Baluchestan, Bampur River 2 km north of Karevandar (27º51'N, 60º46'E); CMNFI 1979-0324, 1, 29.6 mm standard length, Baluchestan, Bampur River at Sa'idabad (27º11'N, 60º22'E); CMNFI 1979-0329, 2, 25.4-30.8 mm standard length, Baluchestan, stream at Zaminbandan (27º02'N, 61º20'E); CMNFI 1979-0411, 1, 60.4 mm standard length, Hormozgan, Minab River (27º24'N, 57º12'E); CMNFI 1979-0412, 9, 22.9-39.3 mm standard length, Hormozgan, spring at Saras (27º30'N, 57º34'E); CMNFI 1979-0416, 39, 15.1-46.8 mm standard length, Hormozgan, Ab Garm-e Ganow (ca. 27º26'N, ca. 56º20'E); CMNFI 2007-0051, 10, 29.5-43.7 mm standard length, Hormozgan, upper Kol River basin (28º19'N, 55º55'E); CMNFI 2007-0055, 5, 30.9-44.6 mm standard length, Hormozgan, Minab River basin (27º47'N, 57º12'E); CMNFI 2007-0056, 2, 32.1-54.2 mm standard length, Kerman, qanat at Kahnuj (27º58'N, 57º45'E); CMNFI 2007-0058, 7, 36.7-51.7 mm standard length, Fars, headwaters of Gowdar River (ca. 27º24'N, ca. 54º16'E).

Garra rossica
(Nikol'skii, 1900)

Common names

mahi-e sangi, mahi-e sang lis, anjarak.

[diskognat in Russian; pathar chat or patherchatta in Pakistan].

Systematics

Discognathus phryne Annandale, 1919 described "from Seistan" is a synonym. Menon and Yazdani (1968) give Nasratabad, Seistan as the type locality. G. rossica has been placed in the genera Discognathus Heckel, 1843 and Discognathichthys Bleeker, 1859, here considered to be synonyms.

Discognathus rossicus var. nudiventris Berg, 1905 was described from "Schiwar" in Iran for specimens with a naked abdomen, thoracic region and groove on the back anteriorly. The distribution of these specimens overlaps with that of the type form and they are not given independent taxonomic recognition here. Berg (1949) later placed them as an infraspecies.

The syntypes of Discognathus rossicus are in the Zoological Institute, St. Petersburg (ZISP 10365), the type locality in Latin on page 239 being "Flum. Tedschent in prov. Transcasp. Zarudnyi. 1892 (4)" while on p. 240 are the localities "Habitat in flumine Tedshent in provincia Transcaspiensi, nec noc in Persia orientale ad Kirmanum orientale" (Nikol'skii, 1900), and confirmed by me. However there were 3 fish in the jar (45.0-54.5 mm standard length) although 4 are listed in the catalogue and in the type description. Berg (1905) lists 3 fish but in Berg (1949) lists only 2. Other materials listed by Nikol'skii (1900) from eastern Iran and Kerman (ZISP 11113, 11703, 11704, 11705, 11708) are apparently not types of rossicus although Berg (1949) indicates that 11704 ("Neizar in Seistano") and 11705 ("Ljabeab in Seistano") are part of the type series from Iran. Eschmeyer et al. (1996) also list ZIL (= ZISP) 10665 (4) as part of the type series, perhaps a misprint for ZISP 10365.

The types for the var. nudiventris are in ZISP 11113, listed by Berg (1905) as 2 fish from Schivar, by Berg (1949) as being 4 fish, not numbered in the ZISP catalogue and with 5 fish in the jar (45.6-66.2 mm standard length). ZISP 11113 appears to have been renumbered in part as 11703 and 11708 according to Berg (1949). ZISP 11708 is listed in Berg (1949) as 13 specimens and 11703 seems to be also 13 specimens. The type localities for var. nudiventris are for ZISP 11113 "Shivar, north of Nikh (Nekh), north-east Kerman, basin of L. Hamun, 23 VI 1896, N. Zarudnyi", for ZISP 11708 "Podaghi, 28°08'N, north-north-east of Bazman, eastern Kerman, 6 VII 1898, N. Zarudnyi" and 11703 is probably "Neizar in Seistan, N. Zarudnyi" according to Berg (1949) (the catalogue number 11703 does not appear under the description of materials in Berg (1949), possibly omitted in error, and it is deduced here that it should have preceded the locality cited). This is not critical as the variety or infraspecies has no taxonomic status.

The holotype of Discognathus phryne is in the Zoological Survey of India, Calcutta (ZSI F9787/1) (Annandale, 1919b; Menon and Yazdani, 1968), a syntype (listed as a cotype) measuring 42.2 mm standard length from "Baluchistan" with the annotation "Ind. Mus. Ex. F 9789/1" is in the Natural History Museum, London (BM(NH) 1919.8.16:1) with another syntype (cotype) from "Quetta" measuring 32.2 mm standard length with the annotation "Ind. Mus. Ex. F 9790/1" (BM(NH) 1919.8.16:2).

Key characters

The single pair of small maxillary barbels (sometimes an anterior pair), absence of a free anterior margin to a weakly developed adhesive disc on the lower head surface, gill raker count, and distribution distinguishes this species. It is separated from the closely related, but geographically separated G. variabilis by smaller size, head length longer than caudal peduncle length, head length equal to or longer than pectoral fin length, distinctly emarginate caudal fin, and dorsal fin origin mid-way between snout tip and caudal fin base or closer to caudal fin base (Berg, 1949).

Morphology

Dorsal fin with 2-3 unbranched and 6-7, usually 7, branched rays, anal fin with 2-3 unbranched and 5 branched rays, pectoral fin with 11-16 branched rays and pelvic fin with 7-8 branched rays. Lateral line scales 33-46. The mid-line of the back, and the chest and belly are naked in some populations. Some fish have only 3-4 rows of scales below the lateral line and this character is extremely variable (Berg, 1949). Scales are a vertical ovoid with an anterior or subcentral anterior focus. The posterior scale margin is rounded and elongate, the dorsal and ventral margins are rounded and merge into the posterior margin and the anterior margin has slight indentations above and below a shallow, rounded central protuberance. However some scales may be squarish with rounded corners and scale shape can be very variable. Circuli are fine. Radii are found on all fields, moderate to very numerous although this is individually variable. A pelvic axillary scale may be present or absent. Gill rakers on the lower arm 9-11, 10-12 total (but see below), almost reaching the adjacent raker when appressed. Pharyngeal teeth 2,4,5-5,4,2, 3,4,5-5,4,3, 2,4,5-5,4,3, 3,4,5-5,4,2, 2,4,5-5,3,2, 2,4,5-5,4,1, 2,4,4-4,4,2, 2,3,5-5,3,2 or 1,3,5-5,3,1. Teeth are conical to flattened with oblique but flattened crown which is slightly concave. Rarely crowns are blade-like and lack the flattened crown. Specimens from the Hari River basin have scales on the thorax and ridge of the back in front of the dorsal fin while those from Sistan lack scales in these areas (Annandale and Hora, 1920; Saadati, 1977). The anterior pair of barbels are usually absent but may be present and minute or even moderately well-developed. Some fish may lack barbels entirely. The gut is very elongate and coiled.

Meristics for Iranian specimens:- dorsal fin branched rays 6(5) or 7(54); anal fin branched rays 5(59); pectoral fin branched rays 11(2), 12(14), 13(14), 14(15), 15(8) or 16(4); pelvic fin branched rays 7(39) or 8(14); lateral line scales 34(3), 35(6), 36(17), 37(10), 38(4), 39(7), 40(1), 43(1), 44(1), 45(1) or 46(1); total gill rakers 11(7), 12(18), 13(18), 14(9), 15(3) or 16(2); pharyngeal teeth 2,4,5-5,4,2 (15), 2,4,5-5,3,2(2), 2,4,5-5,4,1(1), 3,4,5-5,4,2(1) or 2,4,5-5,4,3(1); and total vertebrae 34(11), 35(37) or 36(11).

Sexual dimorphism

Males in spawning condition bear small but evident tubercles on the operculum and the head above the operculum, fine tubercles on top of and anteriorly on the head, small tubercle son the pectoral fin rays following the branching of the rays and on the first unbranched ray, and there are a few minute tubercles on anterior flank scales.

Colour

The upper flanks and back are dark to greyish-brown, greenish-brown or golden-brown and there may be large spots on the upper flanks. The body is silvery overall. There is often a dark bar at the base of the tail. Fins are colourless and the belly and lower head surface are white to yellowish-white. The belly and lower head may be bright yellow as are the neighbouring fin bases. The bases of dorsal fin branched rays 3-5, and sometimes 6, have small dark spots. Young have a bluish mid-lateral stripe along the flank. The peritoneum is black. Colour is darker in clear than in muddy water.

Size

Reaches 9.5 cm.

Distribution

This species is found in the Mashkel (= Mashkid) River basin of Pakistan (Mirza, 1992), the Tedzhen and Murgab River drainages of the former U.S.S.R., and the Tedzhen River, Bejestan, Sistan, Dasht-e Lut, Hamun-e Jaz Murian and Makran basins of Iran (Nikol'skii, 1900; Berg, 1905; Berg, 1949; Menon, 1964; Spillman, 1972; Abdoli, 2000).

Records of Discognathus variabilis Heckel, 1843 from Sistan by Nikolskii (1899) and Regan (1906) are G. rossica (Menon, 1964).

Zoogeography

See under Garra persica. G. rossica is related to Garra variabilis of the Tigris-Euphrates basin.

Habitat

Found in pools and slow-flowing ditches and channels and in reed beds in Sistan.

Age and growth

Sexual maturity is attained at 2-3 years. Females grow more rapidly than males.

Food

Gut contents include green filamentous algae, higher plant fragments and sand grains.

Reproduction

Spawning occurs in the summer and up to 984 eggs are produced. Some fish still contain undeposited eggs in late July. Egg diameter is up to 1.06 mm (Nikol'skii, 1945). In southern Iran, fish with 1.1 mm eggs have been collected on 14 November and on 2 December eggs were 1.3 mm. On 8 May eggs were 1.2 mm suggesting either early spawning or prolonged retention of eggs. The most tuberculate males were found in both November and May.

Parasites and predators

None reported from Iran.

Economic importance

None.

Conservation

This species is widely distributed in eastern Iran and does not appear to be under significant threat.

Further work

Further studies on its biology would clarify details of life history. Its wide distribution in isolated basins may have led to some divergence and speciation that is not readily detected with morphological studies and molecular work on this species might be rewarding.

Sources

Type material: See above, Discognathus rossicus (ZISP 10365), var. nudiventris (ZISP 11113), and Discognathus phryne (BM(NH) 1919.8.16:1 and BM(NH) 1919.8.16:2).

Iranian material: CMNFI 1979-0091, 5, 24.6-58.3 mm standard length, Khorasan, qanat at Nehbandan (31º32'N, 60º02'E); CMNFI 1979-0226, 5, 37.2-49.0 mm standard length, Sistan, pool near Kuh-e Khajeh (30º57'N, 61º17'E); CMNFI 1979-0227, 6, 48.1-61.1 mm standard length, Sistan, naizar at Kuh-E Khajeh (30º57'N, 61º16'E); CMNFI 1979-0230, 2, 29.2-43.4 mm standard length, Sistan, Jehil-e Puzak at Gohoor-ghoori (ca. 31º15'N, ca. 61º42'E); CMNFI 1979-0236, 7, 18.3-47.3 mm standard length, Sistan, irrigation ditch 27 km from Zabol (ca. 30º52'N, ca. 61º22'E); CMNFI 1979-0238, 11, 15.3-29.2 mm standard length, Sistan, irrigation ditch 11 km south of Zabol (30º57'N, 61º27'30"E); CMNFI 1979-0315, 2, 24.9-50.3 mm standard length, Baluchestan, Bampur River 2 km north of Karevandar (27º51'N, 60º46'E); CMNFI 1979-0316, 9, 35.7-53.4 mm standard length, Baluchestan, stream 68 km south of Iranshahr (26º48'N, 61º02'E); CMNFI 1979-0326, 1, 39.0 mm standard length, Baluchestan, Ughin River south of Pip (ca. 26º35'N, ca. 60º02'E); CMNFI 1979-0327, 5, 37.3-46.7 mm standard length, Baluchestan, stream 26 km south of Pip (26º32'N, 59º57'E); CMNFI 1979-0330, 68, 14.7-65.1 mm standard length, Baluchestan, stream 22 km west of Qaleh-ye Zaboli (27º02'30"N, 61º26'E); CMNFI 1979-0336, 30, 22.4-31.6 mm standard length, Baluchestan, qanat 7 km from Khash (28º10'N, 61º15'E); CMNFI 1979-0339, 3, 40.6-51.0 mm standard length, Baluchestan, Tahlab River drainage 16 km from Mirjaveh (28º56'30"N, 61º21'E); CMNFI 2007-0025, 8, 36.6-47.1 mm standard length, Khorasan, qanat south of Birjand (ca. 32º24'N, ca. 59º49'E); CMNFI 2007-0026, 19, 36.3-62.9 mm standard length, Khorasan, qanat at Shusf (31º48'N, 60º01'E); CMNFI 2007-0027, 13, 31.4-60.7 mm standard length, Khorasan, qanat at Khvansharaf (31º34'N, 60º06'E); CMNFI 2007-0028, 13, 36.3-58.9 mm standard length, Khorasan, qanat at Khunik-e Pa'in (31º28'N, 60º06'E); CMNFI 2007-0029, 7, 35.2-60.7 mm standard length, Baluchestan, qanat at Hormak (29º58'N, 60º51'E); CMNFI 2007-0031, 2, 36.0-45.5 mm standard length, Baluchestan, headwater of Bampur River (27º51'N, 60º46'E); CMNFI 2007-0035, 9, 28.9-50.1 mm standard length, Baluchestan, stream west of Zaboli (ca. 26º58'N, ca. 61º27'E); USNM 205905, 6, 30.2-36.9 mm standard length, Baluchestan, small springs in upper Sarbaz River basin (no other locality data).

Garra rufa
(Heckel, 1843)

Common names

mahi-e sangi, mahi-e sang lis, shirbot, gel khorok, gel ra (and even gararufa or gara in Farsi).

[djulake; kokur ahmar or karkoor ahmar; garagoor; gassur achmar (= red gassur) or gassur hadjari (gassur of the pilgrims) at Aleppo (= Haleb, Syria) according to Heckel (1843b), all in Arabic; red garra].

Systematics

Discognathus obtusus Heckel, 1843 described from "Aleppo" and "Mossul", Discognathus crenulatus Heckel, 1849 described from the "Confluenten des Araxes, als aus den Quellen des Saadi und dem Kara-Agatsch" (= probably includes the Pulvar (= Sivan) River near Persepolis; Sa`di at 29°37'N, 52°35'E, now within the city of Shiraz; and the Qarah Aqaj River; all in Fars), Garra rufa gymnothorax Berg, 1949 from "Kulihan, Karun R. basin, 6 IV 1904, N. Zarudnyi" and Garra rufa turcica Karaman (1971) from the Ceyhan River basin in Turkey are synonyms (Krupp, 1985c). Records of Garra lamta (Hamilton, 1822) from Iran are in error (Menon, 1964).

The types of Discognathus rufus are from "Aleppo" according to Heckel (1843b). The syntypes of Discognathus rufus according to Krupp (1985c) are in the Naturhistorisches Museum Wien under NMW 53240, 8 specimens, 59-108 mm standard length from Aleppo and 1 syntype is in the Senckenberg Museum Frankfurt under SMF 553 (formerly NMW), 103 mm standard length and also from Aleppo. The catalogue in Vienna lists 6 specimens. One specimen from NMW 53240, 112.3 mm standard length (as measured by me), was designated as the lectotype and 7 fish, 60.2-97.5 mm standard length as paralectotypes by F. Krupp, 29 October 1984, and published in Krupp and Schneider (1989).

Four syntypes of Discognathus obtusus, 46-134 mm standard length, are under NMW 53238 and 2 syntypes, 65-92 mm standard length, are under SMF 5408 (formerly NMW and also numbered SMF 447)(65.1-93.1 mm standard length). A further 10 fish under NMW 53257 and measuring 31.5-106.2 mm standard length are also indicated on the jar in Vienna as syntypes but this is probably in error as the catalogue lists 6 fish. Eschmeyer et al. (1996) give only 1 syntype under NMW 53257, 4 syntypes under NMW 53238, 1 dried syntype under NMW 79372, 1 syntype in the Senckenberg Museum Frankfurt under SMF 447 and 2 syntypes under SMF 5408 (formerly NMW).

The syntypes of Discognathus crenulatus are in the Naturhistorisches Museum Wien under NMW 53236 (14 specimens) from the Qarah Aqaj River and 53237 (6) from Sa`di measuring 33-79 mm standard length (Kähsbauer, 1964). The 14 specimens under NMW 53236 measure 24.0-75.9 mm standard length and 7 (not 6) specimens under NMW 53237 measure 35.4-56.6 mm standard length according to my observations. Neither the record of Kähsbauer (1964) nor my own data from jars on the shelves accord with the catalogue in Vienna which gives 10 or 8 and 6 or 5 specimens respectively for these two syntype localities.

The syntypes of Garra rufa gymnothorax are in the Zoological Institute, St. Petersburg (ZISP 13214), there being 6 fish in the catalogue and 6 in the jar although Berg (1949) lists 7 in his description. They measure 30.5-44.9 mm standard length. The date in Berg (1949) is 6.VI.1904 while in the catalogue it is 4.III.1904 and in the jar 24.III.1904, variations not accountable by old and new styles of dating (13 days apart). A further collection listed by Berg (1949), (ZISP 24435), is not listed as type material in the text nor in the jar but the catalogue suggest that they are (Eschmeyer et al. (1996) list these 10 fish as syntypes). ZISP 24354 is from "Ziaret-Seid-Hasan, Mesopotamiya". The type series may be only ZISP 13214 as in the text. Eschmeyer et al. (1996) list ZIL (= ZISP) 13215, 17 fish, and ZIL 24436, 3 fish, as syntypes also.

Bianco and Banarescu (1982) referred their material from the Hablehrud and Mand River to Garra rufa crenulata as these fish had fewer scales (29-32 for Mand and 33-34 for Hablehrud versus 35-38 from the Tigris-Euphrates) and fewer gill rakers (15-21 versus 25-27 in Tigris-Euphrates specimens). Possible syntypes of crenulata had intermediate scale counts (31-34) between Mand and Hablehrud fish. These authors suggest that there may be distinct subspecies in these two rivers. Their sample sizes are too small in my opinion to warrant subspecies recognition. Berg (1949) was uncertain of the status of this taxon.

A Principal Components Analysis on 448 fish from the Hormuz, Lake Maharlu, Gulf and Tigris River basins and Sa`di's Tomb using 20 morphometric and 5 meristic characters did not separate any of these groups. Note that fish form the Hormuz basin, rivers draining to the Straits of Hormuz, modally had 7 branched dorsal fin rays (as in G. persica, see table below) but branched caudal fin rays were modally 17 (not 16 as in G. persica). There may be some introgression in this region of Iran.

Menon (1964) and Karaman (1971) consider Garra persica to be a synonym but this species is regarded here as distinct. Both these authors refer specimens from the Tigris River basin of Iran (and therefore all Iranian specimens) to a subspecies, Garra rufa obtusa, distinguished from the type subspecies in Syria, Lebanon, Israel and Jordan by having a variable number of dorsal fin branched rays (7-8), the anal aperture further forward, and the anal fin origin half way between the pelvic fin base and the caudal fin base as opposed to closer to the pelvic base. Krupp (1985c) synonymises Garra rufa obtusa with the type subspecies.

Fish lacking scales on the breast were named by Berg (1949) as Garra rufa gymnothorax but this is a variable character widespread among cyprinid fishes in Iran and is unlikely to be of systematic significance.

Key characters

Two pairs of barbels are present, the adhesive disc is well developed with a free anterior margin, the dorsal fin has 8 branched rays modally, and the caudal fin 17 branched rays modally.

Morphology

Dorsal fin with 2-3, usually 3, unbranched and 7-9 branched rays with a very strong mode at 8, anal fin with 2-3 unbranched and 4-6 with a very strong mode at 5 branched rays (99.6% of 534 specimens from Iran). Pectoral fin branched rays 12-14 (in literature, but see table), pelvic fin branched rays 7-8 (but see table). Lateral line scales 31-38 (but see table), scales from the dorsal fin origin to the lateral line 3-6, from the lateral line to the pelvic fin origin 2-5, predorsal mid-line scales 9-13, and scales around the caudal peduncle 12-17 with a strong mode at 16. Pharyngeal tooth formula 2,4,5-5,4,2 or 2,4,4-4,4,2 (3,3,5-5,3,3 in Heckel (1843b)). Teeth are hooked at the tip. The short gill rakers number 16-21, 12-17 on the lower arm. In Iranian specimens the range is 14-26 (a range only is given since rakers are difficult to count on the arch ends with accuracy and may be related to age). The upper lip is delicately fimbriated. Pharyngeal teeth in Iranian specimens? Total vertebrae in Iranian specimens 32-37 (see table). The chromosome number is probably 2n=52 (Klinkhardt et al., 1995) although Ünlü et al. (1997) give 2n=38 for Turkish specimens with 26 meta- to submetacentric chromosomes and 12 telo- to subtelocentric chromsomes (NF=64) and Gözükara and Çavaş (2004) gave 2n = 44 for Turkish specimens with 22 metacentric and 20 submetacentric chromosomes and 2 acrocentric ones (NF=85).

Some fish are very rounded in cross section while others are more terete.

Meristics for Iranian specimens:-

Basin/Dorsal fin branched rays

7

8

9

x

S.D.

Hormozgan

27

6

7.2

0.39

Lake Maharlu

90

2

8.0

0.15

Sa`di's Tomb

5

20

7.8

0.41

Gulf

16

173

1

7.9

0.29

Tigris River

15

176

3

7.9

0.30

Basin/Pectoral fin branched rays

10

11

12

13

14

15

16

18

x

S.D.

Hormozgan

1

3

18

10

1

14.2

0.78

Lake Maharlu

1

2

15

50

23

1

14.1

0.78

Sa`di's Tomb

5

16

2

2

1

13.2

1.24

Gulf

1

4

57

84

42

1

13.9

0.81

Tigris River

1

1

10

54

93

33

2

13.8

0.88

Basin/Pelvic fin branched rays

6

7

8

9

x

S.D.

Hormozgan

20

12
7.4 0.49

Lake Maharlu

8

84
7.9 0.28

Sa`di's Tomb

9

16
7.9 0.27

Gulf

1

110

79
7.4 0.50

Tigris River

2

51

138

3
7.7 0.50

Basin/Lateral line scales

25

26

27

28

29

30

31

32

33

34

35

36

37

38

40

x

SD

Hormuz

2

7

19

10

1

33.0

0.87

Lake Maharlu

1

4

13

21

32

30

9

31.2

1.54

Sa`di's Tomb

2

6

6

7

4

33.1

1.39

Gulf

1

1

5

16

25

24

25

42

36

19

5

31.2

2.06

Tigris River

2

2

24

44

59

45

9

2

1

1

33.9

1.34

Basin/Total vertebrae

32

33

34

35

36

37

x

S.D.

Hormozgan
7 24 7 3 34.2 0.78

Lake Maharlu
6 38 21 3 34.3 0.70

Sa`di's Tomb
14 9 34.4 0.50

Gulf
2 15 36 34 9 34.3 0.93

Tigris River
23 52 76 17 2 34.5 0.92

Sexual dimorphism

Large males become heavily tuberculate on the front and sides of the snout and in a band from the eye to the nostril and across to the other nostril and eye as illustrated in Fowler and Steinitz (1956) for a specimen from `Ain Umm Keishik in Iraq. A deep, tubercle-free groove is apparent between the upper band of tubercles through the nostrils and the tubercles on the snout above the mouth.

Colour

Overall colour is brownish-olive to dark green with darkly mottled flanks and a yellowish to whitish belly. The head and flanks may be a rusty-red, bronze or golden. A dark or bluish-green band runs along the whole flank ending in a spot on the caudal fin base. Much of the body may be blackish with only the belly creamy. Others are a light olive-green with lime-green highlights giving an iridescent effect especially on upper anterior flank scales. There is a black, greenish-blue, lime-green or dusky-blue spot behind the upper corner of the gill opening, sometimes extending as a bar to the pectoral fin base where the skin is also blue. Fins can be yellowish with darker margins. The pectoral fins can be orange-pink dorsally, grey-white or slightly orange-pink ventrally. The pelvic and anal fins may be orange with the fin rays yellow posteriorly in the anal fin but yellow mesially in the pelvic fin. The bases of the pectoral, pelvic and anal fins are orange-red in breeding males and the caudal fin is orange. The caudal fin can be orange to red ventrally and yellow dorsally. There is a black spot at the caudal fin base and the upper caudal lobe may have a few dark grey spots. The dorsal fin is dark green with reddish pigment at its middle. There is usually a dark spot at the bases of each of the middle 4-5 dorsal fin rays. In some specimens the dorsal fin is orange with yellow posterior rays. The pectoral, dorsal and caudal fin rays may be olive to black rather than yellow or orange. The iris is bright yellow, orange or red.

There is variation in colouration. Some fish are pale while others are very dark; the spots on the dorsal fin may extend two-thirds of the way up the fin rather than being restricted to the base; and the flanks may not be mottled. Fish from muddy water are a sickly grey with the body mottled and the lower caudal lobe dark. Their colour darkens and becomes brighter after immersion in ice water. Fish from deep in qanats are very pale.

Size

Attains 24 cm total length in the Tigris River in Iraq (Rahemo, 1995). Reaches 15.9 cm (Krupp, 1985c), over 17 cm according to Heckel (1843b). Fish up to 18.5 cm total length are known from Khuzestan.

Distribution

Found in the Tigris-Euphrates basin and the Ceyhan, Orontes (= Asi), Quwayq and Jordan river basins and coastal drainages of the eastern Mediterranean as well as much of southern Iran.

In Iran it is found in the Tigris River, Gulf, Lake Maharlu, and Kor River basins and the Hormuz basin (Berg, 1949; Menon, 1964; Bianco and Banarescu, 1982; Gh. Izadpanahi, pers. comm., 1995; M. Rabbaniha, pers. comm., 1995; Abdoli, 2000).

Zoogeography

The wide distribution in Southwest Asia and inadequate examination of variation may mask distinct taxa, although this is not apparent on morphometric and meristic grounds. If such variation is valid, then taxa may reflect vicariant events.

Habitat

This is the commonest species in catches in southwestern Iran, followed by Cyprinion macrostomum. In areas under human influence in Lorestan, such as the lower reaches of rivers and near cities, it is more common than in higher, pristine waters. As well as grazing on exposed rock surfaces in streams, it can be found under pebbles on the stream bed.

Age and growth

Rahemo (1995) reported fish up to age 7 from the Tigris River, Iraq in a parasitological study. Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 291 Iranian fish measuring 2.28-11.82 cm standard length. The a-value was 0.0265 and the b-value 2.919 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases).

Food

Gut contents include diatoms, algae and large quantities of sand in fish examined from Iran. Younis et al. (2001b) found Shatt al Arab fish feeding mainly on organic detritus, followed by diatoms and algae, with arthropods ranking third. . A study by Yalçin-Özdilek and Ekmekçi (2006) in the Asi (= Orontes) River in Turkey demonstrated that this species is a grazer on aquatic plants, mostly consisting of benthic cyanobacteria, chrysophytes and phytoplankton with included rotifers and protozoans. Both season and location in a stream affected the composition of the diet with season the most important factor.

Reproduction

Bardakci, Ozansoy and Koptagel (at www.epress.com/w3jbio/vol5/bardakci/paper.htm, downloaded 29 March 2001) note depression of vitellogenesis in a hot spring population in Turkey, perhaps due to temperature and starvation. A nearby stream population has a higher gonadosomatic index. Ovaries increased in size and weight from May to July in both localities although the hot spring had fewer mature oocytes and more atretic oocytes at various development stages. High temperatures and poor food conditions in some Iranian habitats may be limiting factors in reproduction for this species.

Parasites and predators

Jalali and Molnár (1990a) record two monogenean species, Dactylogyrus spp., from this species in the Dez River. Gussev et al. (1993b) describe two new species, Dactylogyrus rectotrabus and D. acinacus, from this species in the Dez River, Khuzestan. Jalali et al. (2005) summarise the occurrence of Gyrodactylus species in Iran and record G. sp. from fish in the Helleh River.

Eaten by Silurus triostegus at Baghdad (notes on a specimen in the Field Museum of Natural History, Chicago (FMNH 51251)).

Economic importance

Ündar et al. (1990) identify this species and Cyprinion macrostomum as the "doctor fish" of the Kangal hot spring in Turkey (Warwick and Warwick, 1989; Kürkçüoğlu and Öz, 1989; Bardakci, Ozansoy and Koptagel at www.epress.com/w3jbio/vol5/bardakci/paper.htm, downloaded 29 March 2001, Bilke, 2004; Anonymous, 2007; and various newspaper and television reports). High water temperatures around 35°C reduce the amount of plankton available as fish food and the fish nibble away infected skin of humans who bathe in these waters. This fish is known as "licker" (and Cyprinion macostomum as "striker") from its behaviour in the spa pools. The healing properties are linked to the high level of selenium (1.3 p.p.m.) in the water, selenium being beneficial in some skin diseases and possibly UV light. The fish facilitate the action of the selenium and UV light by softening and clearing away psoriatic plaque and scale, exposing the lesions to the water and sunlight. However some lesions are made worse and the fish can cause some new ones.

Conservation

A common species with a wide distribution and not under any specific threat. Vulnerable in Turkey (Fricke et al., 2007).

Further work

Meristic counts summarised above show overlapping but significant variation between drainage basins and may reflect recognisable taxa. Molecular studies would help clarify this situation but is should be noting that variation of this nature is to be expected in a wide ranging species and does not always warrant taxonomic distinction.

Sources

Type material: See above, Discognathus rufus (NMW 53240 and SMF 553), Discognathus obtusus (NMW 53238 and SMF 5408), Discognathus crenulatus (NMW 53236 and 53237), and Garra rufa gymnothorax (ZISP 13214), and see comments on other possible types.

Iranian material: CMNFI 1970-0540, 7, 22.2-41.4 mm standard length, Fars, qanat south of Kazerun (no other locality data); CMNFI 1979-0018, 48, 21.5-64.9 mm standard length, Fars, Barm-e Shur (29º28'N, 52º41'30"E); CMNFI 1979-0019, 4, 28.9-35.4 mm standard length, Fars, Barm-e Baba Haji (29º23'N, 52º40'E); CMNFI 1979-0026, 2, 21.5-22.3 mm standard length, Fars, Shapur River at Shapur (29º47'N, 51º35'E); CMNFI 1979-0027, 1, 50.5 mm standard length, Fars, Chehel Cheshmeh (ca. 29º43'N, ca. 52º04'E); CMNFI 1979-0033, 34, 23.7-72.0 mm standard length, Fars, Barm-e Shur (29º28'N, 52º41'30"E); CMNFI 1979-0036, 1, 36.4 mm standard length, Fars, Shapur River at Shapur (29º47'N, 51º35'E); CMNFI 1979-0045, 16, 20.0-59.2 mm standard length, Fars, spring at Sa'adi's Tomb, Shiraz (29º37'N, 52º35'E); CMNFI 1979-0046, 1, 43.7 mm standard length, Fars, qanat at Barm-e Dalak (ca. 29º35'N, ca. 52º38'E); CMNFI 1979-0047, 2, 36.2-67.6 mm standard length, Fars, spring source of Ab-e Paravan marshes (ca. 29º34'N, ca. 52º42'E); CMNFI 1979-0048, 1, 40.1 mm standard length, Fars, spring and marsh northeast side of Lake Maharlu (ca. 29º32'N, ca. 52º48'E); CMNFI 1979-0075, 31, 12.5-57.1 mm standard length, Fars, Mand River at Pol-e Kavar (29º11'N, 52º41'E); CMNFI 1979-0109, 1, 74.3 mm standard length, Fars, Mand River at Shahr-e Khafr (28º56'N, 53º14'E); CMNFI 1979-0111, 8, 30.7-62.0 mm standard length, Fars, stream on Shiraz-Bushehr road (29º37'30"N, 52º21'E); CMNFI 1979-0112, 5, 55.0-77.1 mm standard length, Fars, stream draining Soltanabad Marshes (29º29'N, 52º38'30"E); CMNFI 1979-0113, 4, 40.8-63.7 mm standard length, Fars, spring at Sa'adi's Tomb, Shiraz (29º37'N, 52º35'E); CMNFI 1979-0115, 5, 57.7-66.5 mm standard length, Fars, spring at Sa'adi's Tomb, Shiraz (29º37'N, 52º35'E); CMNFI 1979-0120, 5, 27.6-66.3 mm standard length, Bushehr, Dalaki River near Konar Takhteh (29º28'N, 51º21'E); CMNFI 1979-0125, 2, 99.0-121.4 mm standard length, Bushehr, Dalaki River near Dalaki (ca. 29º28'N, ca. 51º21'E); CMNFI 1979-0128, 6, 26.6-32.1 mm standard length, Shur River between Atashkadeh and Firuzabad (28º51'N, 52º31'E); CMNFI 1979-0129, 43, 24.8-46.9 mm standard length, Fars, spring 2 km north of Farrashband (28º54'N, 52º04'E); CMNFI 1979-0131, 10, 16.9-46.6 mm standard length, Fars, Ab-Arak River 24 km from Qir (28º38'N, 52º49'E); CMNFI 1979-0132, 11, 23.1-49.5 mm standard length, Fars, Shur River 54 km from Firuzabad (28º35'N, 52º58'E); CMNFI 1979-0155, 11, 27.1-42.9 mm standard length, Fars, spring at Gavanoo village (28º47'N, 54º22'E); CMNFI 1979-0156, 20, 33.8-56.7 mm standard length, Fars, qanat at Rashidabad (28º47'N, 54º18'E); CMNFI 1979-0157, 10, 40.6-88.6 mm standard length, Fars, qanat at Hadiabad (28º52'N, 54º13'E); CMNFI 1979-0158, 13, 35.3-54.2 mm standard length, qanat over Qasook River (28º54'N, 53º53'30"E); CMNFI 1979-0161, 11, 43.1-92.2 mm standard length, Fars, qanat on Shiraz-Neyriz road (29º10'30"N, 53º41'E); CMNFI 1979-0193, 3, 22.8-27.1 mm standard length, Fars, river 8 km from Darab (28º45'N, 54º27'30"E); CMNFI 1979-0195, 3, 44.3-59.4 mm standard length, Fars, jube stream of road to Fasa (ca. 28º54'N, ca. 53º53'30"E); CMNFI 1979-0199, 4, 37.3-45.0 mm standard length, Fars, qanat 18 km from Jahrom (ca. 28º23-25'N, ca. 53º31-40'E); CMNFI 1979-0200, 5, 27.6-43.6 mm standard length, Fars, Mand River tributary 13 km from Jahrom (28º36'N, 53º36'30"E); CMNFI 1979-0202, 9, 19.1-25.1 mm standard length, Fars, Mand River (29º01'N, 53º00'E); CMNFI 1979-0206, 2, 50.9-51.9 mm standard length, Fars, qanat near Runiz-e Pa'in (29º12'N, 53º40'E); CMNFI 1979-0241, 6, 35.7-49.2 mm standard length, Fars, Shapur River at Shapur (29º47'N, 51º35'E); CMNFI 1979-0271, 8, 34.7-56.7 mm standard length, Lorestan, Kashkan River drainage (33º39'N, 48º32'30"E); CMNFI 1979-0273, 15, 40.9-58.2 mm standard length, Lorestan, Kashkan River drainage 5 km from Khorramabad (33º26'N, 48º19'E); CMNFI 1979-0274, 3, 24.4-32.8 mm standard length, Lorestan, Kashkan River drainage 20 km from Khorramabad (33º27'N, 48º11'E); CMNFI 1979-0275, 7, 38.6-60.5 mm standard length, Lorestan, Kashkan River near Ma'mulan (33º25'N, 47º58'E); CMNFI 1979-0276, 10, 42.1-70.0 mm standard length, Lorestan, Chamesk River (ca. 33º19'N, ca. 47º53'30"E); CMNFI 1979-0277, 1, 128.4 mm standard length, Lorestan, Kashkan River drainage (33º30'N, 47º59'30"E); CMNFI 1979-0278, 12, 53.9-89.3 mm standard length, Lorestan, Kashkan River draiange, Sarab Dowrah (33º34'N, 48º01'E); CMNFI 1979-0279, 9, 33.5-117.3 mm standard length, Lorestan, Khorramabad River (33º37'N, 48º18'E); CMNFI 1979-0288, 43, 22.9-102.3 mm standard length, Ilam and Poshtkuh, Gangir River at Sarab Ewan (33º50'N, 46º18'E); CMNFI 1979-0289, 5, 57.2-101.9 mm standard length, Kermanshahan, Diyala River drainage (34º28'N, 45º52'E); CMNFI 1979-0290, 6, 24.1-54.9 mm standard length, Kermanshahan, Diyala River drainage in Qasr-e Shirin (34º31'N, 45º35'E); CMNFI 1979-0291, 7, 24.9-66.8 mm standard length, Kermanshahan, Diyala River draiange 15 km from Qasr-e Shirin (34º24'N, 45º37'E); CMNFI 1979-0293, 8, 92.1-101.3 mm standard length, Fars, Mand River at Kavar (29º11'N, 52º41'E); CMNFI 1979-0304, 2, 41.6-43.0 mm standard length, Fars, Lake Famur (ca. 29º31'N, ca. 51º50'E); CMNFI 1979-0350, 4, 28.1-33.5 mm standard length, Khuzestan, Marun River near Marun (30º39'30"N, 50º02'E); CMNFI 1979-0364, 4, 36.0-50.6 mm standard length, Khuzestan, Karkheh River branch at Abdolkhan (31º52'30"N, 48º20'30"E); CMNFI 1979-0365, 2, 32.1-32.8 mm standard length, Khuzestan, stream in Doveyrich River drainage (32º25'N, 47º36'30"E); CMNFI 1979-0366, 2, 22.7-29.2 mm standard length, Khuzestan, stream 17 km west of Dehloran (32º45'30"N, 47º05'30"E); CMNFI 1979-0367, 1, 38.0 mm standard length, Khuzestan, Meymeh River 11 km north of Dehloran (32º44'30"N, 47º09'30"E); CMNFI 1979-0368, 4, 25.3-47.7 mm standard length, Khuzestan, Karkheh River (32º24'30"N, 48º09'E); CMNFI 1979-0369, 4, 24.9-37.7 mm standard length, Khuzestan, Shush River at Shush (32º12'N, 48º14'30"E); CMNFI 1979-0371, 1, 37.3 mm standard length, Khuzestan, stream in Karkheh River drainage (32º05'N, 48º19'E); CMNFI 1979-0374, 1, 62.2 mm standard length, Khuzestan, stream tributary to Bala River (32º40'N, 48º15'E); CMNFI 1979-0375, 10, 37.9-71.1 mm standard length, Khuzestan, stream tributary to Bala River (ca. 32º45'N, ca. 48º14'30"E); CMNFI 1979-0378, 1, 49.3 mm standard length, Khuzestan, stream tributary to Karkheh River (ca. 32º48'N, ca. 48º04'E); CMNFI 1979-0379, 1, 63.1 mm standard length, Khuzestan, Dez River (32º12'N, 48º27'E); CMNFI 1979-0382, 3, 34.7-38.0 mm standard length, Khuzestan, Karun River at Shushtar (32º03'N, 48º51'E); CMNFI 1979-0383, 5, 31.9-53.0 mm standard length, Khuzestan, stream in Ab-e Shur drainage (31º59'30"N, 49º06'E); CMNFI 1979-0384, 1, 63.3 mm standard length, Khuzestan, river in Ab-e Shur drainage (32º00'N, 49º07'E); CMNFI 1979-0385, 2, 44.9-46.8 mm standard length, Khuzestan, stream in Ab-e Shur drainage (ca. 32º01'N, ca. 49º07'30"E); CMNFI 1979-0387, 2, 48.2-64.5 mm standard length, Khuzestan, stream in Jarrahi River drainage (31º25'N, 49º38'E); CMNFI 1979-0388, 1, 55.2 mm standard length, Khuzestan, Zard River 21 km north of Ramhormoz (31º19'N, 49º44'E); CMNFI 1979-0389, 2, 43.8-57.7 mm standard length, Khuzestan, Zard River at Bagh-e Malek (31º31'N, 49º53'30"E); CMNFI 1979-0390B, 10, 33.8-69.7 mm standard length, Khuzestan, stream tributary to Zard River (31º29'N, 49º54'30"E); CMNFI 1979-0391, 1, 39.7 mm standard length, Khuzestan, stream in Marun River drainage (31º28'N, 49º51'E); CMNFI 1979-0392, 1, 42.6 mm standard length, Khuzestan, Zard River 25 km north of Ramhormoz (ca. 31º32'N, ca. 49º48'E); CMNFI 1979-0396, 3, 22.7-35.1 mm standard length, Kheyrabad River 20 km from Behbehan (30º32'N, 50º23'30"E); CMNFI 1979-0397, 2, 52.4-59.5 mm standard length, Khuzestan, stream tributary to Kheyrabad River (30º30'N, 50º28'E); CMNFI 1979-0398, 1, 37.6 mm standard length, Boyer Ahmadi-ye Sardsir va Kohkiluyeh, stream in Zohreh River drainage (30º24'30"N, 50º37'30"E); CMNFI 1979-0399, 1, 41.0 mm standard length, Fars, stream near Basht in Zohreh River drainage (30º19'30"N, 51º15'E); CMNFI 1979-0497, 2, 52.4-60.3 mm standard length, Fars, Mand River at Band-e Bahman (29º11'N, 52º40'E); CMNFI 1979-0501, 8, 24.7-48.9 mm standard length, Fars, Mand River at Kavar (29º11'N, 52º41'E); CMNFI 1987-0217, 5, 35.8-55.6 mm standard length, Khuzestan, Karun River at Kut Abdollah (31º13'N, 48º39'E); CMNFI 2007-0063, 5, 40.1-65.9 mm standard length, Fars, Mand River tributary near Jahrom (28º36'N, 53º37'E); CMNFI 2007-0065, 1, 81.2 mm standard length, Fars, Barm-e Dalak (ca. 29º35'N, ca. 52º38'E); CMNFI 2007-0066, 2, 47.1-55.3 mm standard length, Fars, qanat under Sa'di's Tomb, Shiraz (29º37'N, 52º35'E); CMNFI 2007-0100, 2, 48.4-54.8 mm standard length, Azarbaijan-e Gharbi, Kalwi Chay near Piranshahr (ca. 36º44'N, ca. 45º10'E); CMNFI 2007-0109, 10, 54.7-78.3 mm standard length, Kordestan, Qeshlaq River basin (ca. 35º16'N, ca. 47º01'E); CMNFI 2007-0110, 1, 84.9 mm standard length, Kordestan, Yuzidar River basin (ca. 35º05'N, ca. 46º56'E); CMNFI 2007-0111, 11, 26.5-64.5 mm standard length, Kermanshahan, Alvand River near Sar-e Pol-e Zahab (ca. 34º36'N, ca. 45º56'E); CMNFI 2007-0112, 19, 43.1-54.3 mm standard length, Kermanshahan, Kerend River basin near Shahabad-e Gharb (ca. 34º06'N, ca. 46º30'E); CMNFI 2007-0116, 4, 25.8-31.7 mm standard length, Kermanshahan, Gav Masiab River basin west of Sahneh (ca. 34º28'N, ca. 47º36'E).

Comparative material: BM(NH) 1931.12.21:9-10, 2, 94.9-95.5 mm standard length, Iraq, Mosul (36º20'N, 43º08'E); BM(NH) 1973.5.21:187-188, 2, 67.6-73.1 mm standard length, Iraq, Tigris River at Jadriyah (no other locality data); BM(NH) 1974.2.22:1418, 1, 67.4 mm standard length, Iraq, Khalis (33º49'N, 44º32'E); BM(NH) 1974.2.22:1441-1444, 4, 48.9-86.3 mm standard length, Iraq, Baghdad (33º21'N, 44º25'E); BM(NH) 1986.2.14:2-3, 2, 64.8-91.8 mm standard length, Iraq, Baghdad (33º21'N, 44º25'E); NMW 91123, 5, ?, Turkey, Cheilani bei Cizre, Tigris basin (ca. 37º20'N, ca. 42º10'E).

Garra variabilis
(Heckel, 1843)

Common names

See under genus account.

[karkoor mit-la'oon, gassur diseileki or gassur isivid (gassur = colour of strawberries, isivid referring to the spotted, almost black fish according to Heckel (1843b), all in Arabic; variable garra].

Systematics

Menon (1964) considers this species to be the most primitive in the genus. It has been placed in the genus Discognathichthys Bleeker, 1860, e.g. by Berg (1949).

Heckel (1843b) gave the type localities as "Mossul" and "Aleppo". The syntypes of Discognathus variabilis are in the Naturhistorisches Museum Wien under NMW 53239, 8 specimens, 38-112 mm standard length and in the Senckenberg Museum Frankfurt under SMF 403 (formerly NMW), 4, 72-86 mm standard length, all from Aleppo (Krupp, 1985c). In Vienna I made counts on types as listed below under Sources. Material under NMW 53238 (3) from Aleppo may also be types. Eschmeyer et al. (1996) list NMW 532339 (= an error for 53239) (8), NMW 53260-69 (1, 2, 2, 2, 3, 2, 2, 2, 2, 2), NMW 53272 (4), SMF 403 (4) (formerly NMW) and in the Museum für Naturkunde, Universität Humboldt, Berlin, ZMB 3301 (3) (formerly NMW; 82.6-99.2 mm standard length measured in February 2006).

Key characters

The single pair of maxillary barbels, absence of a free anterior margin to a weakly developed adhesive disc on the lower head surface without papillae on the rear part, scaled back, chest and belly, gill raker count and distribution distinguishes this species. It is separated from the closely related but geographically separated G. rossica by larger size, head length shorter than caudal peduncle length and pectoral fin length, slightly emarginate caudal fin, and dorsal fin origin closer to snout tip than the caudal fin base (Berg, 1949).

Morphology

Dorsal fin with 2-3, usually 3, unbranched and 6-8, modally 7, branched rays, anal fin with 2-3 unbranched and 5 branched rays. Pectoral fin branched rays 11-14, pelvic fin branched rays 7-8. Lateral line scales 32-40, scale from the dorsal fin origin to the lateral line 4-6, scales below the lateral line to the pelvic fin origin 3-4, scales around the caudal peduncle usually 16, and predorsal scales in mid-line 12-14. The chest and belly are scaled. The upper lip is not fimbriate. There may be 2 pairs of barbels in some larger fish. Pharyngeal tooth formula 2,4,5-5,4,2 (2,3,5-5,3,2 in Berg (1949), 3,3,5-5,3,3 in Heckel (1843b)) and the short gill rakers number 13-20, on the lower arm of the gill arch only. Gut very elongate and coiled.

Sexual dimorphism

Specimens from NMW 91121 had the top and sides of the head finely tuberculate and scales on the back before the dorsal fin with fine tubercles lining the scale margins. The upper lip, lip sides and sucker (except for a naked central area) have keratinised tubercles. Tubercles line the dorsal surface of pectoral fin rays, fading medially and following the ray branching in single rows.

Colour

Overall colour is olivaceous brown or greyish with darker mottlings. The flanks may have large, irregularly-arranged dark spots. The upper corner of the operculum may have a black spot. The belly is reddish-yellow. The middle 3-4 rays of the dorsal fin each have a small, black spot at their bases. There is a black spot at the caudal fin base. The lateral line may occasionally have a double row of black spots as in certain Alburnoides spp. Young fish may have a dark lateral stripe. Peritoneum black.

Size

Reaches 15.5 cm or according to Heckel (1843b) 5 Zoll (= about 21 cm).

Distribution

Found in the Quwayq (= Kueik), Orontes (= Asi) and Nahr al-Kabir rivers of the Levant and the Tigris-Euphrates basin (Menon, 1964; Krupp, 1985c). In Iran, this species is found in the Tigris River basin. Abdoli (2000) maps the Jarrahi, middle Karun, lower Dez, Karkheh, lower Simarreh, and lower Kashkan rivers.

Keyserling (1861) recorded this species from Sistan but this is a misidentification. Records of this species (as Discognathus variabilis) from Sistan by Nikolskii (1899) and Regan (1906) are G. rossica (Menon, 1964).

Zoogeography

This species is related to Garra rossica of eastern Iran. See also under Garra persica. Krupp (1985c) considers this species to belong to the indo-asiatic line of Garra.

Habitat

Garra rufa and this species seem to exclude each other, variabilis being more common in faster water (F. Krupp).

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

None reported from Iran.

Economic importance

None.

Conservation

This species appears to be very rare in Iran and if specific localities are found they should be protected.

Further work

The presence of this species in Iranian waters and its biology there need study.

Sources

Type material: See above, Discognathus variabilis (NMW 53239, SMF 403 and ZMB 3301) and see comments also on other possible types.

Iranian material:- None.

Type material used for counts: NMW 53260, 1, 40.6 mm standard length, NMW 53261, 2, 87.6-97.8 mm standard length, NMW 53262, 2, 51.4-52.1 mm standard length, NMW 53263, 2, 101.8-103.1 mm standard length, NMW 53264, 3, 52.3-54.2 mm standard length, NMW 53266, 2, 87.6-92.7 mm standard length, NMW 53267, 2, 61.5-68.3 mm standard length, NMW 53268, 7, 43.9- 71.4 mm standard length (dried at some point), NMW 53269, 2, 100.3-109.1 mm standard length, NMW 53272, 4, 84.9-92.1 mm standard length, all previous Tigris at Mosul; NMW 53239, 8 36.9-111.0 mm standard length, Aleppo.

Comparative material: BM(NH) 1935.9.12:27-40, 5, 60.8-69.1 mm standard length, Iraq, Karasu (no other locality data); BM(NH) 1935.9.12:53, 1, 76.0 mm standard length, Iraq, Tchaiy Su (no other locality data); BM(NH) 1986.2.14:2-5, 2, Iraq, Baghdad (33º21'N, 44º25'E); BM(NH) 1968.12.13:290-297, 8, 30.7-71.6 mm standard length, Syria, Tigris River at `Ayn Diwar (37º17'N, 42º11'E); BM(NH) 1968.12.13:298-304, 7, 44.4-66.7 mm standard length, Syria, Quweiq River at Masslemiyeh (no other locality data); NMW 91121, 10, 71.9-115.4 mm standard length, Turkey, Wadi Mahmedian Tschai (ca. 38°20'N, ca. 40°45'E).

Genus Gobio
Cuvier, 1816

The gudgeon genus includes about 24 species found from the British Isles through Europe and northern Asia to Korea. There is one species in Iran and two other species are now in the genus Romanogobio.

The body shape is distinctive, being elongate and fusiform with moderately large scales (36-51), the throat is naked or scaled, the mouth is inferior or terminal, horseshoe-shaped and has a barbel at each corner, the lower lip is thin like the upper lip but is interrupted medially, gill rakers are short and widely spaced, pharyngeal teeth are in 2 rows (usually 5 in the main row and 2-3 in the second) and are obviously hooked at the tips, both dorsal and anal fins are short and spineless, the gut is short and the peritoneum silvery, and the vent is remote from the anal and pelvic fin origins.

Gobio gobio
(Linnaeus, 1758)

Common names

mahi kopur kafzi (= bottom-dwelling carp fish), گاو ماهي (= gav mahi, probably in error for Neogobius and related gobies).

[Turkestanskii peskar' or Turkestan gudgeon in Russian].

Systematics

Cyprinus Gobio was originally described from England.

Bungia nigrescens Keyserling, 1861 described from "Fluss Heri-Rud bei Herat" (the Harirud at Herat in Afghanistan, formerly part of Persia) is a synonym. Bungia Keyserling, 1861 is a synonym of Gobio Cuvier, 1816 (Eschmeyer, 1990).

This species is represented in Iran by the subspecies Gobio gobio lepidolaemus Kessler, 1872, originally described as Gobio fluviatilis var. lepidolaemus from Ak-darja and Chodshaduk in the Zeravshan River basin, Uzbekistan and the Syr Darya at Khodzhent, Tajikistan. This subspecies is distinguished from the typical gudgeon by having a scale-covered throat, deep body, deep and short caudal peduncle, slightly notched or emarginate caudal fin and small size (Amanov, 1972). Berg (1948-1949) considers that these characters would be sufficient to distinguish this taxon as a full species but there are intermediate forms. Interestingly, Reshetnikov and Shakirova (1993) list Gobio lepidolaemus as a full species.

Also in the western Caspian Sea basin as Gobio gobio lepidolaemus natio holurus Berg, 1914 (the Terek gudgeon or Terskii peskar') but not recorded from Iranian waters and considered to represent intergrades between G. gobio gobio and G. gobio lepidolaemus by Bănărescu in Bănărescu (1999).

A syntype from Khodzhent of Gobio gobio lepidolaemus measuring 49.7 mm standard length is in the Natural History Museum, London under BM(NH) 1897.7.5:26, formerly in St. Petersburg University, a syntype is in the Zoological Institute, St. Petersburg under ZISP 2078 or 2076, and a possible syntype is in the Zoological Museum of Moscow State University under MMSU P.1052. Svetovidova (1978) refers to ZISP 2078 as the holotype on page 257 and ZISP 2076 as the holotype on page 262 (Eschmeyer et al., 1996).

Key characters

See above. This is the only gudgeon in eastern Iran and is separated from other cyprinids by the meristic characters, presence of barbels, absence of fin spines, mouth not transverse or crescentic but horseshoe-shaped, and colour pattern. It is separated from the other related species in Iran (Romanogobio macropterus and R. persus) by having the body and caudal peduncle compressed (caudal peduncle depth at anal fin insertion greater than caudal peduncle width) and by well-defined spots on the dorsal and caudal fins. This subspecies is also characterised by all members of the population having scales on the breast, absent or variably developed in other subspecies and populations to the west.

Morphology

Usmanova (1975) found differences between populations of this species in Uzbekistan associated with habitat. Fish from more stable habitats have deeper and wider bodies and reduced fin sizes.

Dorsal fin branched rays 6-8, usually 7, after 2-4 unbranched rays, anal fin branched rays 5-8, usually 5 (Amanov, 1972 - an error, see below) or 6 (Usmanova, 1975; Banarescu and Nalbant, 1973) after 2-3 unbranched rays, pectoral fin branched rays 13-17 and pelvic fin branched rays 7-8. Lateral line scales 33-46, usually 37-42 and averaging less than 40 (Bănărescu in Bănărescu (1999)). Scales have posterior radii only (or, if present, very few anterior radii) and the scale focus is subcentral anterior but not very eccentric in small fish, very eccentric in large fish. The anterior scale margin is rounded to wavy. There is a pelvic axillary scale. The anus is separated from the anal fin origin by 5-6 closely overlapping scales and is near the end of the pelvic fins but underneath them. Pharyngeal teeth 3,5-5,3 usually but Pipoyan (1998) found 21 variant counts for 141 Gobio gobio in Armenia with 3,5-5,3 (34.0%), 2,5-5,2(22.7%), 2,5-5,3(10.0%) and other combinations at about 2% or less. The anterior teeth are blunt with small hooks, conical and short and are followed by long, thin, strongly hooked teeth (the description of Bungia nigrescens may be in error in stating that there is only one row of teeth but Pipoyan (1998) notes that Gobio gobio in Armenia are exceptionally uniserial). Total gill rakers 1-7, only developed rakers being counted and anterior rakers reduced to bumps not included. Developed rakers are stubby and may or may not touch the adjacent raker when appressed. Total vertebrae 33-42 (this wide literature range may reflect the wide species range but could also include specimens counted in varying ways). The gut is an elongate s-shape with a slight anterior loop. The chromosome number is 2n=50 (Klinkhardt et al., 1995; Bănărescu in Bănărescu, 1999).

Meristics in Iranian fish are as follows: branched dorsal fin rays 7(4), branched anal fin rays 6(4), branched pectoral fin rays 15(2), 16(1) or 17(1), branched pelvic fin rays 7(4); lateral line scales 37(1), 38(2) or 39(1), scales around caudal peduncle 14(2) or 16(2); total gill rakers 5(1) or 6(2); pharyngeal teeth 3,5-5,3(1), 3,5-4,3(1) or 3,4-5,2 or 3(1); and total vertebrae 38(2) or 39(1).

Sexual dimorphism

Snout length, depth of the head at the occiput, eye diameter, greatest body depth and thickness, predorsal distance, pectoral-pelvic fin distance, and barbel length are greater in females than males while caudal, pectoral and pelvic fin lengths and height of dorsal fin are greater in males. Females reach larger sizes than males (Usmanova, 1975). Males darken in the spawning season. Fine tubercles develop on the side and upper surface of male heads, on the upper flank and back anteriorly, and the 8 outer rays of the pectoral fin.

Colour

The top of the head, the back and the flanks above the lateral line are dark brown and may have a greenish tinge. The lower flanks are paler and may have a silvery tinge or be a light yellow. The mid-flank bears a row of 6-13 dark spots which may merge into a line, merge in pairs or form a lattice. The back may have 4-5 longitudinal dark bands with a variegated pattern. The dorsal and caudal fins have 3-5 rows of spots, the pectoral fins have several rows of small spots and the pelvic and anal fins may also have 2-6 rows of spots but are often colourless. The peritoneum is silvery.

Colour may vary with habitat, being more uniform and darker on a monotonous background and more spotted and lighter on a gravelly background.

Size

Attains 20.0 cm total length for the species, 11.1 cm for lepidolaemus.

Distribution

Found from the British Isles and north of the Pyrenees through much of Europe north of the Alps to Siberia. The subspecies considered here is found only in the Tedzhen or Hari River basin in Iran (Abdoli, 2000). It is also found in north flowing rivers of the Kopetdag in Turkmenistan and eastwards in Central Asia to the Chu River in Kyrgyzstan. The natio holurus is found in the Kuma, Terek and Sulak rivers of the western Caspian shore but not as far south as waters neighbouring Iran.

Zoogeography

This is a widely distributed and very variable species with many taxa listed as subspecies, potentially species if further investigated. As part of a gobionine fauna found across Eurasia, this wide distribution may be suggestive of further work that could be done to clarify relationships of these fishes.

Habitat

This gudgeon is found in rivers near the bank where there is a sand or fine gravel bottom and in inlets, backwaters and weed beds of irrigation canals in the Kashkadar'ya of Uzbekistan (Usmanova, 1975). It prefers stagnant water in the Surkhandar'ya of Uzbekistan (Amanov, 1972). Gudgeons are generally resistant to pollution and varying environmental conditions although there is little information on the eastern Iranian subspecies.

Age and growth

Sexual maturity is reached at the age of 1 year and a length of 3.9-4.3 cm in females from the Kashkadar'ya in Uzbekistan, at 4.6-5.3 cm in the Surkhandar'ya, or at 2-3 years and 3-7 cm in reservoirs in Uzbekistan; also reported as 2-3 years in the Issyk Kul' and the Chu River (Usmanova, 1975). Life span is over 4 years (Amanov, 1972).

Food

Food is mainly benthic invertebrates, chiefly insect larvae such as chironomids in Iranian fish, but varied items may be taken depending on the water body and food availability. Abdoli (2000) lists Chironomidae, Ephemeroptera, Plecoptera and Trichoptera. Some insects falling on the water surface are taken. Remains of terrestrial plants, green algae and detritus have also been recorded (Usmanova, 1975). Detritus and vegetation (mostly diatoms) dominate in some gut samples, followed by insect larvae and benthopelagic crustaceans, and occasionally fish eggs. Other samples show chironomid larvae to be the main diet item or cladocerans. Vegetation fragments are apparently seized accidentally with such foods as chironomids (Amanov, 1972).

Reproduction

Spawning is intermittent and takes place from May to August in Uzbekistan, May-July in the Issyk Kul' and May-June in the Chu River (Usmanova, 1975). Up to 12,900 eggs are laid on a clay-sand bottom at 18-20°C (Amanov, 1972). Well-developed eggs are present in Iranian fish caught on 10 November. Elsewhere gudgeons slap the water surface with the rear part of the body, and males and females rub their bodies together while releasing eggs and sperm (Bănărescu in Bănărescu, 1999).

Parasites and predators

None reported from Iran.

Economic importance

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in aquaria and textbooks. Gudgeons in Europe have been used as bait fish and as food.

Conservation

Lelek (1987) classifies populations of this species in Europe as intermediate to rare.

Further work

The biology of this species in Iran requires study.

Sources

Type material: See above, a syntype of Gobio gobio lepidolaemus (BM(NH) 1897.7.5:26).

Iranian material: CMNFI 2007-0014, 3, 81.5-85.0 mm standard length, Khorasan, Kuh-e Sang Park, Mashhad (ca. 36º18'N, ca. 59º36'E).

Comparative material: SMF 17137, 5, 61.3-84.1 mm standard length, Afghanistan, tributary of the Harirud near Herat (34°21'N, 62°14'E).

Genus Hemiculter
Bleeker, 1859

This genus contains several species with a native distribution in the Amur River basin of the Russian Far East and in China, Taiwan and Viet Nam. A single species has been accidentally introduced to Iran.

The sawbellies are characterised by an elongate body; moderate-sized scales with a deeply-decurved lateral line only 1-3 scales above the mid-ventral line; a scaleless keel from the pectoral fins to the vent; a short dorsal fin with a spine and an elongate anal fin; pharyngeal teeth in 3 rows; gill rakers short and numerous; and pelagic eggs.

Hemiculter leucisculus
(Basilewsky, 1855)

Common names

tizeh kuli or teez-e-kooli (= sharp or spiny fish, kuli being any small fish), shakam tiz (= ?sharp belly), kuli-e mordab (= lagoon fish).

[vostrobryushka or sharpbelly in Russian; common sawbelly, knifefish].

Systematics

Culter leucisculus was originally described from Peking, China. Hemiculter eigenmanni (Jordan and Metz, 1913) is a synonym.

Key characters

The sharp keel, lacking scales, extending from the vent or anus to the throat below the pectoral fin on the mid-ventral surface is distinctive, especially when combined with the dorsal fin spine and the three rows of pharyngeal teeth.

Morphology

The last dorsal fin unbranched ray is a sharp spine with a flexible tip. The lateral line curves downward from its origin on the head to the level of the end of the pectoral fin and then parallels the lower body margin eventually to curve upward (sometimes sharply) at the end of the anal fin level and run along the middle part of the caudal peduncle. There is a tubercle on the lower jaw which fits into a notch on the upper jaw. Dorsal fin branched rays 6-8, usually 7, after 2-3 unbranched rays and anal fin branched rays 10-18, mostly 13-14 (but see below), after 3 unbranched rays. Lateral line scales 43-55, scales above the lateral line 8-11, and scales below the lateral line to the pelvic fin origin 1-3. Scales bear numerous fine circuli and a few posterior radii. Total gill rakers 17-29, reaching the second raker below when appressed. Pharyngeal teeth, 2,4,5-5,4,2, 2,4,5-4,4,2, 2,4,4-5,4,2, 2,4,4-4,3,2, 3,4,5-4,3,2 or 2,3,5-4,3,2. Teeth are hooked at the tip with an elongate and narrow grinding surface. Total vertebrae 42. The gut is an elongate s-shape. The chromosome number is 2n=48.

Meristic values for Iranian specimens are:- dorsal fin branched rays 6(1) or 7(6), anal fin branched rays 11(1), 12(5) or 13(1), lateral line scales 47(1), 48(1), 50(1), 52(1), 53(1) or 54(2), total gill rakers 17(1), 18(4) or 19(2), and pharyngeal teeth 2,4,5-5,4,2(2), 2,4,4-5,4,2(2), 2,4,5-4,4,2(2) or 2,4,4-4,3,2(1).

Sexual dimorphism

Unknown.

Colour

Overall colour dark above, silvery on the flanks and whitish on the belly. There is a dark stripe along the upper flank. The lips are dark. The dorsal, caudal and anterior pectoral and anal fin ray edges and their fin membranes are lightly pigmented with melanophores. The peritoneum is silvery with some melanophores giving it a brownish pigmentation in preserved fish.

Size

Reaches 25 cm.

Distribution

The native range of this species is from Maritime Russia south through China to Korea and Viet Nam. First reported from the Anzali Mordab by Holčík and Razavi (1992) and apparently not uncommon there. Abbasi et al., (1999), Kiabi et al. (1999), Abdoli (2000) and Gasmi and Mirzaei (2004) record this species from the lower Safid River and the Anzali Talab, and in the middle Aras River. Patimar et al. (2002a; 2002b; 2008) and Patimar (2008) report it from the International Wetlands of Alma-Gol, Adji-Gol and Ala-Gol. Found in ab-bandans along the Caspian shore of Iran (Jolodar and Abdoli, 2004). Also reported in the Tedzhen River, Karakum Canal and Kopetdag Reservoir of Turkmenistan (as H. eigenmanni)(Aliev et al., 1988; Shakirova and Sukhanova, 1994; Sal'nikov, 1995) and so may eventually be found in the Tedzhen (= Hari) River basin of Iran. Now recorded from the Hawizah Marsh in southern Iraq (Coad and Hussain, 2007).

Zoogeography

This species is introduced to Iran, probably by accident along with commercial shipments of Chinese major carps from Central Asia in the former U.S.S.R. and/or Rumania in 1967. The Chinese major carps in Central Asia came from the Amur River basin in the Far East and sawbellies were accidentally transferred with them in the 1950s-1960s (Holčík and Razavi, 1992).

Habitat

Found in rivers, lakes, small ponds and swamps but little appears to be known about its habitat requirements.

Age and growth

Patimar et al. (2002a; 2002b; 2008) report six age groups in the International Wetlands of Alma-Gol, Adji-Gol and Ala-Gol. The smallest mature specimens there were 2 years old. It is an 'r' strategist, forming dense stunted populations in any new environment.

Food

Macrophytes, fish, crustaceans and insects are eaten by this species and young fish feed on zooplankton. Iranian fish contain large plant fragments and filamentous algae.

Reproduction

Up to 1,180 eggs are produced. Fish from a swamp near Hendeh Khaleh in Gilan taken on 9 August contained well-developed eggs but Patimar et al. (2002a; 2002b; 2008) report a peak spawning in March in the International Wetlands of Alma-Gol, Adji-Gol and Ala-Gol.

Parasites and predators

Sattari et al. (2007) record the digenean Diplostomum spathaceum and the monogenean Diplozoon sp. in this species in the Anzali wetland of the Caspian shore. It is food for Sander lucioperca, Silurus glanis and Aspius aspius in Turkmenistan (Aliev et al., 1988).

Economic importance

This species has potential as a food fish and is canned in China but this is probably outweighed by its competition with native species for food and the possibility of predation on fish eggs and young. The sawbelly is easily able to switch from one food to another as conditions warrant (Holčík and Razavi, 1992) and is known to show more rapid growth and higher fecundity than under native conditions. Welcomme in Courtenay and Stauffer (1984) regards this species as a pest when introduced.

It is found on the fish market at Bandar Anzali and a catch of 41 kg is reported from the Anzali Mordab in 1990 (Holčík and Oláh, 1992).

Conservation

None required for an introduced species.

Further work

The spread of this species and its effects on native species and habitats should be monitored.

Sources

Based on data in Holčík and Razavi (1992) for 5 Iranian specimens from the Anzali Mordab (134.5-143.4 mm standard length).

Iranian material: Uncatalogued material, 2, 99.6-113.1 mm standard length, Gilan, swamp near Hendeh Khaleh (37°23'N, 49°28'E).

Comparative material: CMNFI 2006-0028, 2, 112.5-123.6 mm standard length, Iraq, Hawizah Marsh (31º38'30"N, 47º35'21"E and 31º36'02"N, 47º33'09"E).

Genus Hemigrammocapoeta
Pellegrin, 1927

This genus comprises 5 species of which 1 is found in Iran. The genera Tylognathoides Tortonese, 1938, Neotylognathus Kosswig, 1950 and Hemigarra Karaman, 1971 are synonyms (Krupp, 1985c; Krupp and Schneider, 1989).

The genus is characterised by having a distinct rostral flap without separate lateral lobes, underslung mouth, horny covering to the upper and lower jaws, no disc on the chin behind the lower lip but this area papillose, no or up to 2 pairs of barbels (mouth and lip structures can be quite variable), short dorsal and anal fins, complete or incomplete lateral line, and pharyngeal teeth in 3 rows with somewhat hooked tips and spoon-shaped crowns, count uniquely 2,4,5-5,4,2 (but see below). Most species are found in the Levant. The genus is closely related to Garra according to Krupp (1985c).

Hemigrammocapoeta elegans
(Günther, 1868)

Common names

None.

Systematics

Originally described in the genus Tylognathus Heckel, 1843 and also placed in the genus Hemigarra Karaman, 1971. Tylognathus elegans was described from "Mesopotamia?" (sic). Six syntypes are in the Natural History Museum, London (BM(NH) 1850.10.21:31-36), one specimen (number 36) being an alizarin preparation. One of the specimens had been designated as a lectotype but was loose in the jar, and the jar contains a total of 17 fish measuring 36.0-44.5 mm standard length.

Key characters

Characters of the genus serve to identify this species in Iran, particularly around the mouth region.

Morphology

Dorsal fin with 3 unbranched and 6-8, usually 8 branched rays, anal fin with 3 unbranched and 5 branched rays, pectoral fin with 11-17 branched rays and pelvic fin with 8 branched rays. Last unbranched dorsal fin ray thickened although concave posteriorly, tapering distally to a thin ray. Unbranched ray anterior to the last one also thickened in large fish. Lateral line scales 33-38. Flank scales have a wavy anterior margin, very fine and numerous circuli, few anterior and more numerous posterior radii, posterior scale field with tubercles and a subcentral anterior focus. Pelvic axillary scale present. There is a scale between the anus and the anal fin origin. Gill rakers short, just reaching the base of the second raker below when appressed. Barbels 4, thin but moderately long. Snout projecting over inferior mouth but not strongly folded over the upper lip. Upper lip free. Lower lip thicker in the middle but no sucker, posterior edge wavy. Horny edge to lower jaw. Weakly papillose behind the lower lip. Large nasal flap and large first suborbital bone. Pharyngeal teeth in the major row quite massive and obviously hooked with a concave space below. The most anterior tooth is the largest and posterior teeth are slender. Minor row teeth are similar but smaller. Gut complexly coiled. Total vertebrae 46-48.

Meristic values for Iranian specimens are:- dorsal fin branched rays 6(1, ? deformed) or 8(7), anal fin branched rays 5(8), pectoral fin branched rays 16(2) or 17(6), pelvic fin branched rays 8(8); lateral line scales 34(1), 36(5), 37(1) or 38(1) and caudal peduncle scales 12(8); total gill rakers 17(1), 18(5), 19(1) or 20(1); pharyngeal teeth 2,3,5-5,3,2 (5) (sic); and total vertebrae 46(1), 47(5) or 48(2).

Sexual dimorphism

Unknown.

Colour

There is an indistinct silvery stripe along the flank. Dark pigment ends about 1 scale above the lateral line along the flank. Peritoneum black with some silvery patches.

Size

Reaches 10.9 cm total length.

Distribution

Found in the Tigris-Euphrates basin. Initially recorded from Harmaleh on the Dez River of Khuzestan (ZSM 25716, 25717), Abdoli (2000) maps distributions from the upper Karun, upper Dez, and upper Karkheh including the Simarreh rivers.

Zoogeography

The relatives of this species are confined to the Levant, a distribution matched by various other unrelated taxa indicating former connections between these basins (see Krupp (1985c; 1987)).

Habitat

Unknown.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

No parasites reported from Iran. One specimen examined was from the gut of a Silurus triostegus taken at Harmaleh on the Dez River.

Economic importance

None.

Conservation

This species is rare in Iran and is poorly known elsewhere. Assuming this is not the result of inappropriate collecting techniques, then the status of this species should be assessed by field surveys.

Further work

The distribution and biology of this species is very poorly documented and needs attention.

Sources

Type material: See above (BM(NH) 1850.10.21:31-36).

Iranian material: ZSM 21862, 5, 60.5-86.9 mm standard length, Khuzestan, Dez River at Harmaleh (31º57'N, 48º34'E); ZSM 25716, 2, 76.4-79.3 mm standard length, same locality as preceding; ZSM 25717, 1, 78.5 mm standard length, "from the stomach of a wels", same locality as preceding.

Comparative material: BM(NH) 1973.5.21:185, 1, 57.5 mm standard length, Iraq, Tigris River at Jadriyah, Baghdad (no other locality data); FMNH 51235, 1, 43.9 mm standard length, Iraq, Tigris River at Rustam Farm near Baghdad (no other locality data); FMNH 51236, 12, 38.7-59.5 mm standard length, same locality as preceding.

Genus Hypophthalmichthys
Bleeker, 1859

The silver carp genus contains 3 species with a native distribution in eastern Asia. Two species have been widely introduced for food in aquaculture and for phytoplankton control.

The genus is characterised by an elongate and compressed body, very small scales (usually over 100 in the lateral line), eyes low on the head with their lower margin below the mouth corner level, a terminal mouth, no barbels, gill rakers long and thin, a very long gut, branchiostegal membranes joined and free of the isthmus, a short dorsal and elongate anal fin, both spineless, pharyngeal teeth in 1 row, and a ventral keel from the throat or pelvic fins to the anus.

Hypophthalmichthys molitrix
(Valenciennes, 1844)

Common names

فيتوفاگ (= fitofag or phytophag, i.e. phytophage or plant eater), كپور نقره اي (= kopur-e noqrehi or kapoor-e-noghreie, meaning silver carp), آزاد پرورشي (= azad-e parvareshi meaning free farmed).

[silver carp, thickforehead, white thickforehead; belyi tolstolobik, tolpyga and maksun in Russian].

Systematics

Leuciscus molitrix was originally described from China. Populations in the Anzali Mordab are hybrids with Hypophthalmichthys nobilis (J. Holčík, in. litt., 1989).

Key characters

The abdomen has a compressed keel extending from the breast to the vent, the eyes are positioned low such that they are visible from the underside of the head, and scales are minute. The similar bighead (H. nobilis) can be distinguished by the long pectoral fins which extend past the origin of the pelvic fins, a shorter keel (pelvic fins to anus), and gill raker structure (free, no spongy root mass).

Morphology

Dorsal fin with 2-3 unbranched rays followed by 6-7 branched rays, anal fin with 2-3, usually 3, unbranched rays and 11-15 branched rays, pectoral fin branched rays 17, and pelvic fin branched rays 7. Lateral line scales 85-125 Scales are rounded to oval with a posterior focus and very few posterior radii. Total vertebrae 36-40. Pharyngeal teeth 4-4, well-developed, compressed and with striated grinding surfaces. Gill rakers exceed 650 and are longer than the gill filaments. The left and right sides of the gill arches are united by a mucous membrane to form a continuous band; the gill rakers are distinguishable distally but the roots form a spongy mass. The gut is long and complexly coiled. The diploid chromosome number is 48 (Klinkhardt et al., 1995; Varasteh et al., no date). There were 6 pairs of metacentric, 14 pairs of submetacentric and 4 pairs of acrocentric chromosomes in the latter study.

Sexual dimorphism

Unknown.

Colour

Overall colour is silvery, the back bluish to grey-black with upper flanks olivaceous but silver laterally and ventrally, and the fins red or immaculate.

Size

Reaches 1.50 m and 56 kg.

Distribution

The natural distribution is from the Amur River in the former U.S.S.R. southward to southern China. In Iran, it has been introduced to the Sistan basin including the Hamun Sabari and the canal flowing into Chahnimeh (Ahmadi and Wossughi, 1988; Mansoori, 1994; J. Holčík, in litt., 1996), the Voshmgir Reservoir on the Gorgan River (Petr, 1987) as well as widely stocked in Mazandaran and Gorgan provinces in reservoirs and lakes by both governmental agencies and private companies. It is also introduced to the Anzali Mordab and its outlets (Holčík and Oláh, 1992), the Gorgan, Tajan, Aras, and Safid rivers, and the Anzali Mordab (Kiabi et al., 1999; Abbasi et al., 1999), Lake Zaribar, Kordestan (Abzeeyan, 5(5):III, 1994), reservoirs and fish farms in Khuzestan, Mahabad Dam (Abdi, 1999; www.mondialvet99.com, downloaded 31 May 2000; Shamsi et al., 2009).

Abdoli (2000) records this species generally from the Dasht-e Kavir, Kerman-Na'in, Sistan and Hamun-e Jaz Murian basins; from the middle and lower Kor River and the Pulvar River in the Kor River basin; from the middle and lower Zayandeh River in the Esfahan basin; from the Khersan River, lower Karun River, the lower Jarrahi river, the lower Dez River the upper Karkheh River and the lower Kashkan River in the Tigris River basin; the middle and lower Qareh Su and Qom rivers, the lower Karaj and Shur rivers in the Namak Lake basin; the lower Zarrineh river and the middle and lower Talkheh River in the Orumiyeh basin; the middle Kashaf River in the Tedzhen basin; the middle and lower Atrek and Gorgan rivers, the lower Babol, Heraz, Chalus, Tonekabon and Safid rivers, the Anzali Talab and the Caspian coast generally in the Caspian Sea basin.

This species is also recorded from the Karakum Canal and Kopetdag Reservoir in Turkmenistan (Shakirova and Sukhanova, 1994; Sal'nikov, 1995) and may eventually reach Iranian waters in the Tedzhen (= Hari) River and Caspian Sea basins. It is pond-cultured in Iraq and is known from open waters such as the Shatt al Arab and Tigris River (Al-Hassan, 1994).

Zoogeography

This species is an exotic, introduced to Iran from a variety of sources. It may become established in the large river systems of southern Iran and Iraq from escapees (Al-Hassan, 1994).

Habitat

This species is a riverine fish in its native habitat, or is found in water bodies connected to rivers, but is extensively cultivated in ponds for food throughout Asia. Silver carp can live in the Caspian Sea at salinities of 5-8‰ although a few are found at 10-12‰. It enters rivers to spawn (Abdusamodov, 1986). Temperatures in the range 0-40ºC are tolerated although 26-30ºC or 30-34ºC is preferred in different studies. It is more cold-tolerant than bighead carp. This species can be difficult to catch as it will jump over nets, to a height of about 2 m, and when frightened by noise has been known to jump into boats. Malek Nedjad and Parivar (1993) consider that the level of lead pollution in the Anzali Mordab (average 0.124 p.p.m. in surface waters, 0.1956 p.p.m. in deeper waters) caused 8% mortality in fertilised eggs and nearly 18% of eggs are useless for fisheries work.

Age and growth

Terek River silver carp first mature at 4 years for males and 5 years for females. About 15% of females mature at 4 years but 87% of the females and 85% of the males are in the 5-7 age groups (Abdusamadov, 1986). Maturity varies with locality, at 2-8 years, with males maturing a year earlier than females. Silver carp can reach 18-23 kg in 4-5 years. Life span is at least 20 years. Abdolmalaki (2004) examined the fishery for this carp in the Mahabad Reservoir, Iran and found mean fork length was 51.15 cm and mean weight was 2272.1 g. The length-weight relationship was W = 0.013L^3.04. The von Bertalanffy growth parameters were L_∞ = 150 cm, K = 0.128, instantaneous rate of total (Z), natural (M) and fishing (F) mortality were 1.68/year, 0.22/year and 1.46/year. The exploitation rate (E) was calculated to be 0.82. The biomass was 158.5 t with a maximum sustainable yield estimated at 68 t. A decreased fishing effort was recommended as the annual catch in 1998-1999 was 101,123.5 kg.

Food

The gill rakers form a very fine, sponge-like mesh used to filter small planktonic food, aided by the epibranchial organ that produces mucus to trap very small particles. This species is a pump filter feeder, taking smaller particles than bighead carp. Food in Lake Kinneret, Israel is phytoplankton from February to August and predominately zooplankton from September to January, a response to a decrease in phytoplankton biomass in summer-fall (Spataru and Gophen, 1985). Cladocerans and cyclopoid copepods dominate the biomass of zooplankton taken. The ability to take cyclopoids is due to the large mouth, strong sucking power and the high filtration rate when feeding. Food is taken passively rather than selectively.

Reproduction

Silver carp require cool, flowing water to breed. The spawning migration begins at the end of April in the Terek River of Dagestan at 16-17°C, with a peak between the middle of May and the beginning of June. Generally spawning occurs between 18 and 26°C. Fecundity reaches 1,340,500 large, greyish eggs (and in the Kara-kum Canal 1,525,000 eggs; elsewhere to 5.4 million eggs). Water hardened eggs are 4.9-5.6 mm, smaller than those of bighead carp. Spawning takes place after a sharp rise in water level and current speed. Spawners chase each other near the water surface where eggs and sperm are shed. Eggs are first found in the drift in the second week of June and hatch 34-70 hours later depending on temperature. Some larvae reach rice fields and live there until autumn when the fields dry up where some are lost, others migrating. Other larvae are carried into the Caspian Sea where they are sensitive to the prevailing salinity at 1-1.5 days old (Abdusamadov, 1986). It does not breed in the wild in Iran.

Parasites and predators

Mokhayer (1989) reports metacercariae of the eye fluke, Diplostomum spathaceum from this species in Iran, which can cause complete blindness and death in commercially important species, as well as black-spot, Posthodiplostomum cuticola. Jalali and Molnár (1990b) record the monogenean Dactylogyrus hypophthalmichthys from this species at fish farms in Iran. Masoumian and Pazooki (1998) surveyed myxosporeans in this species in Gilan and Mazandaran provinces, finding Myxobolus pavlovskii. Akhlagi (1999) reports that high temperatures (up to 32°C) stresses this species and leaves it open to infection with Aeromonas hydrophila. Safari and Khandagi (1999) record Clostridium botulinum from 1.1% of fresh and smoked samples of this species in Mazandaran Province. Ebrahimzadeh Mousavi and Khosravi (1999; www.mondialvet99.com, downloaded 31 May 2000) record the toxigenic fungi Aspergillus flavus, Alternaria, Penicillium and Fusarium from this species and the pond water at a fish farm in northern Iran. Akhondzadeh et al. (2002) and Akhondzadeh Basti and Zahrae Salehi (2003) show that the psychotropic pathogen Listeria monocytogenes is found in market and fish farm samples of this species. Naem et al. (2002) found the following parasites on the gills of this species from the western branch of the Safid River, namely the protozoans Ichthyophthirius multifilis and a Trichodina species and the monogenean trematode Dactylogyrus hypophthalmichthys. Jalali et al. (2002) and Jalali and Barzegar (2006) record several parasites from this species in Lake Zarivar, namely Dactylogyrus hypophthalmichthys, D. suchengiaii, Diplostomum spathaceum, two species of Argulus, and myxosporean plasmodia. Jalali et al. (2005) summarise the occurrence of Gyrodactylus species in Iran and record G. sprostonae from Safid River fish. Araghi Soureh and Jalali Jafari (2006) recorded Dactylogyrus hypophthalmichthys and D. suchengtaii from this species in the Mahabad River of the Lake Orumiyeh basin, the latter species being a new record for Iran. Shamsi et al. (2009) found Dactylogyrus hypophthalmichthys and D. suchengtaii in this species from fish farms, and Zarivar Lake, and Mahabad reservoir, respectively.

Economic importance

This species is the most productive freshwater fish in the world, with 3.1 million metric tons produced in 1997 (versus 2.2 million mt for Cyprinus carpio).

Holčík and Oláh (1992) report a total catch of 6585 kg in the Anzali Mordab in 1990, at 8.8% of the total fish catch being the third largest catch. This species is used in polyculture with common and grass carps and comprises 50-63% of the fishes. They are fed through pond fertilization without supplementary feeding (Emadi, 1993b). Silver carp in oil are packaged in northern Iran (http://www.netiran.com/business.html, downloaded 31 October 2003). They are also found in fish stores in Ahvaz, Khuzestan (personal observations, September 1995). Iran shares with Uzbekistan the most production of this species among North African and Near Eastern aquaculture; the Iranian catch increasing from nothing in 1989 to 24,720 tonnes in 1994 (Food and Agriculture Organization, Fisheries Department, 1996). Bartley and Rana (1998b) however give a production of 15,228 t for 1995. Market price in 1995 was about U.S.$1.00/kg in 1995, lower than for grass carp at about $2, but silver carp have the higher stocking ratio (Rana and Bartley, 1998a). Kals and Bartels (2004) give some recommendations for improving silver carp farming in Iran.

Abdolmalaki (2004) reported that 67.3% of the 150,261 kg of fishes caught in Mahabad Reservoir in the1998-1999 fishing season were this species. The fishing effort was 69 beach seine hauls and 2530.7 fishing effort units with 7-15 cm gill nets, each unit being 100 m of gill net for 24 hours.

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in aquaculture and food. The food of this species being phytoplankton it has been introduced to areas lacking filter feeders such as Europe and Israel in efforts to control algal blooms associated with eutrophication. The results in Israel were controversial and difficult to interpret because of chemical treatments for nuisance dinoflagellates and introductions of other fish species (Opuszynski and Shireman, 1995). In some cases the effects of the introduction were negative because the silver carp fed from September to January on zooplankton which themselves reduce algal blooms. There was also competition with a more valuable native species of cichlid for the same zooplankton resource and the native cichlid was a more efficient consumer of the nuisance dinoflagellate. Elsewhere, the consumption of plankton may remove food for native, larval fishes.

This species has been studied extensively as a commercial species in various parts of the world. Studies specific to Iran include Azari Takami et al. (2000) on female broodstock selection; Nazifi et al. (2000) on the effects of the organophosphate trichorofon on serum biochemistry; Nazifi et al. (2001) on haematological parameters after experimental intoxication with trichlorofen; Javadian et al. (2003) on the effect of ice storage on lipid and other chemical changes, moisture and texture and appearance; Moradian et al. (2003) on the effects of the hormone thyroxine on the survival of early growth stages (the ratio of hatched eggs in 0.5 p.p.m. thyroxine was highest); Kashani Sabet et al. (2004) on inducing ovulation with hormones combined with dopamine antagonists; Alavi Talab (2007) on use of skins and fins for gelatin extraction; Roshan and Moini (2009) on washing minced flesh with cold water and brine to increase quality of surimi (1.5% brine for 10 minutes was best); etc.

This fish can be dangerous to boaters as it leaps out of the water (Kolar and Lodge, 2002). Akhondzadeh Basteh et al. (2006) found the bacterial pathogens Listeria monocytogenes in fresh and smoked H. molitrix, Staphylococcus aureus in smoked H. molitrix, Escherichia coli and Salmonella dublin in fresh H. molitrix and Vibrio haemolyticus in smoked H. molitrix.

The Iranian Fishery Research Institute has made ice cream using this fish. The product is deoderised so there is no fishy flavour and a protein in the fish blood is reported to lower the temperature at which ice crystals form meaning less cream or fat is needed in the product (Iran Daily, 11 February 2009).

Conservation

Krasznai (1987) and Petr (1987) give details of fish farms propagating this species in Iran. For example, 30 million fish were produced by the Safid Rud Fish Farm in 1986. 20 million carp, silver carp and grass carp fingerlings were produced in the Shahid Rajaae Hatchery in Sari for release across Iran in reservoirs and dams (Abzeeyan, Tehran, 4(7):VII, 1993). Piri and Ordog (1999) describe the effects of herbicides and insecticides on this species. These chemicals are used extensively on rice fields in Gilan where aquaculture is widely developed.

Hybridisation (presumably with H. nobilis) has led to poor growth and a decline in the fishery (Shehadeh, 1997).

Further work

The biology of this species in Iran needs investigation in relation to its effects on native fishes and its distribution monitored.

Sources

Opuszynski and Shireman (1995) summarise the biology and culture of this species.

Comparative material: CMNFI 1977-0590, 3, 120.3-135.5 mm standard length, Israel, Kibbutz `En Hamifraz (32º59'N, 35º05'E).

Hypophthalmichthys nobilis
(Richardson, 1844)

Common names

sar gondeh or sargundeh (= bighead) in the Caspian basin, كپور سرگنده (= kopur-e sargondeh, meaning bighead carp), mahi kopur sar gondeh, fitofag.

[pestryi tolstolobik in Russian; bighead, painted thickforehead].

Systematics

Leuciscus nobilis was originally described from Canton, China. Eschmeyer et al. (1996) give the date of publication as 1845, Reshetnikov et al. (1997) as 1846.

Howes (1981) reaffirms the placement of this species in Hypophthalmichthys Bleeker, 1860, considering that the characters of abdominal keel length, pharyngeal dentition and gill raker form are insufficient to place this species in the distinct genus Aristichthys Oshima, 1919. Other authors disagree (see Eschmeyer, 1990; Reshetnikov et al., 1997).

Populations in the Anzali Mordab are hybrids with Hypophthalmichthys molitrix (J. Holčík, in litt., 1989). Artificial hybrids with Ctenopharyngodon molitrix have been developed in Iran and their karyology studied (Dorafshan and Kalbasi, 2007). The F1 hybrids were triploids with 3n=72 and NF=132.

Key characters

The similar silver carp (H. molitrix) can be distinguished by the short pectoral fins which do not extend past the origin of the pelvic fins, a longer keel (throat to anus), and gill raker structure (continuous band uniting both sides, roots fused into a spongy mass).

Morphology

Dorsal fin unbranched rays 3 followed by 7 unbranched rays, anal fin unbranched rays 3 with 11-14 branched rays, pectoral fin branched rays 16-19, and pelvic fin branched rays 7-9. Lateral line scales 92-115. Scales are a rounded oval with a slightly posterior focus, very few posterior radii and numerous fine circuli. Total vertebrae number 36-41. Pharyngeal teeth 4-4, with smooth grinding surfaces. Gill rakers are very numerous. The gut is elongate and convoluted. The diploid chromosome number is 48 (Klinkhardt et al., 1995).

Sexual dimorphism

Males have a sharp edge along the dorsal surface of several anterior pectoral fin rays.

Colour

Overall colour is silvery and the body has numerous scattered small black spots which makes some fish speckled or darker in overall colour. A few larger blotches may be present. The head is often darker and contrasts with the silvery body. Fins are greyish and similar to the adjacent body colour with the caudal fin darkest, sometimes reddish.

Size

Attains 1.57 m in length and 77.5 kg, possibly to 1.95 m.

Distribution

This species is a native of China and was first introduced to Iran in 1966 from a hatchery in the Krasnodar region of the former U.S.S.R. (Anonymous, 1970b). It is reported from fish farms in the Caspian Sea basin and is stocked in reservoirs throughout Iran (Coad and Abdoli, 1993b). Introduced to the Kor River in Fars (A. Alamdari, in litt., 1997) and reported from the Gorgan, Tajan, Aras, and Safid rivers, and the Anzali Mordab (Abbasi et al., 1999; Kiabi et al., 1999; Jalali et al., 2005). Abdoli (2000) records it generally from the Sistan, Hormuz, Kor, Kerman-Na'in, Dasht-e Kavir, Esfahan, Namak, Tigris, and Orumiyeh basins and from the lower Gorgan, Neka, Babol, Haraz, Chalus and Tonekabon rivers, the Gorgan and Anzali mordabs and along the Caspian coast. Also in Lake Zarivar (Shamsi et al., 2009).

Also reproducing naturally in the Karakum Canal and recorded from the Kopetdag Reservoir in Turkmenistan (Shakirova and Sukhanova, 1994; Sal'nikov, 1995) and may eventually reach Iranian waters in the Tedzhen (= Hari) River basin. Introduced to Iraq for fish farming.

Zoogeography

An exotic introduced to Iran.

Habitat

In their natural habitat, bigheads are found in large rivers and associated floodplain lakes. They migrate upstream to spawning grounds when water levels rise, moving to flooded land afterwards, and returning to the river channel as water levels fall. Bigheads can live in the Caspian Sea at salinities of 5-8 p.p.t. although a few are found at 10-12 p.p.t.. They can adjust gradually to salinities of 15-20 p.p.t. They enter rivers to spawn (Abdusamodov, 1986) but are known to spawn in the Karakum Canal of Turkmenistan. Preferred temperatures for feeding and reproduction are within the general range 20-30°C. Activity almost ceases at 10°C and the critical thermal maximum is 38.8°C.

Age and growth

Males achieve first maturity at age 5 and males of 5-7 years and 81-90 cm make up 90% of the run in the Terek River. Most females mature at 6 years and 81% of the females on the spawning run are 6-7 years old and 75-100 cm (Abdusamodov, 1986). Males mature at 2-4 years and females at 3-5 years (and 10 kg) in Turkmenistan. Sexual maturity varies widely with environmental conditions, 2-6 years for males and a year later for females. Life span is up to 16 years. Growth is rapid, attaining 18-23 kg in 4-5 years.

Food

Zooplankton is almost exclusively the food of this species. Phytoplankton and detritus may be taken when zooplankton biomass is low. Most feeding occurs during summer and peaks daily in the range 1200 to 2000 hours. This species is both a pump feeder, using the buccal pump to push food-laden water through the gill rakers, and a ram feeder, swimming with the mouth open to force water through the gills, with intermittent gulps. Feeding often occurs at the water surface, in contrast to silver carp, as well as in the water column and on the bottom.

Reproduction

A spawning migration of this species enters the Terek River in the second week of May at water temperatures of 18-19°C, numbers increasing until the end of June. Spawning takes place after a sharp rise in water level and current speed. Males actively chase females near the water surface, occasionally butting the female's belly, and sometimes leaping out of the water. Eggs and sperm may be cast into the air. Fecundity attains 1,860,800 eggs. Unswollen eggs are 1.4-1.5 mm in diameter and water hardened eggs are 5.7-6.2 mm. Eggs are first found in the drift in the second week of June and hatch 34-70 hours later depending on temperature. Some larvae reach rice fields and live there until autumn when the fields dry up where some are lost, others migrating. Other larvae are carried into the Caspian Sea where they are sensitive to the prevailing salinity at 1-1.5 days old (Abdusamadov, 1986).

Parasites and predators

Jalali and Molnár (1990b) record the monogeneans Dactylogyrus aristichthys and D. nobilis from this species in Iranian fish farms. Naem et al. (2002) found the following parasites on the gills of this species from the western branch of the Safid River, namely the protozoan Trichodina sp., monogenean trematodes Dactylogyrus nobilis, D. aristichthys, and Gyrodactylus sp.. Jalali et al. (2005) summarise the occurrence of Gyrodactylus species in Iran and record G. sprostonae and G. sp. in fish from the Safid River. Masoumian and Pazooki (1998) surveyed myxosporeans in this species in Gilan and Mazandaran provinces, finding Myxobolus pavlovskii. Shamsi et al. (2009) found Dactylogyrus aristichthys, D. nobilis and D. taihuensis in this species from fish farms and Zarivar Lake.

Economic importance

Holčík and Oláh (1992) report a catch of 466 kg in the Anzali Mordab in 1990. Aquaculture production in 1995 was 1269 tonnes (Bartley and Rana, 1998b). Marjan Iran Company was selling 1500-1800 g fish for U.S.$1.90/kg in August 2003 (http://groups.yahoo.com/groups/hilsa/message/25).

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in aquaculture and food. It ranks fourth on world aquaculture production. This species has been used in Israel to reduce zooplankton populations in reservoirs in an effort to improve water quality (Opuszynski and Shireman, 1995). It has a higher net production in culture systems than H. molitrix and Ctenopharyngodon idella. The consumption of plankton may remove food for native, larval fishes, and affect the diet of piscivorous fishes and birds. This species can also cause habitat alteration, increasing turbidity, and introduce diseases and parasites. In Iran, it has been used in aquaria for investigations on the effect of lead nitrate on blood serum electrolytes (Jamili et al., 2006).

Conservation

Krasznai (1987) and Petr (1987) give some details of propagation of this species in Iran. Some populations are hybrids (see above) and there is a danger of loss of genetic purity in fish farm stocks should breeding adults be captured in the wild. As an exotic, there is no need for conservation.

Further work

The biology of this species in relation to native species should be investigated for Iran.

Sources

Jennings (1988), Opuszynski and Shireman (1995) and Kolar et al. (2005) summarise the biology and culture of this species.

Comparative material: CMNFI 1980-0530, 2, 230.6-255.8 mm standard length, Japan, pond cultured (no other locality data).

Genus Iranocypris
Bruun and Kaiser, 1948

This genus contains a single species found only in Iran and the characters of the species are therefore the characters of the genus. This blind cave species is placed in a world-wide context by Proudlove (1997a; 1997b).

Iranocypris typhlops
Bruun and Kaiser, 1948

Dorsal view of head Ventral view of head

Courtesy of R. Mehrani

Courtesy of Amir Hosin Zalaghi, 11-19 May 2010 (all fish returned to cave alive)

Courtesy of Kiavash Golzarianpour

Iranocypris typhlops movie.3gp, courtesy of Kiavash Golzarianpour

Common names

mahi-ye kureghar or mahi-ye kur-e qar (= blind cave fish), kopur mahi kureghar, ماهي كور (= mahi kur).

[Iran cave barb].

Systematics

The holotype is in the Zoological Museum of Copenhagen (ZMUC P 26475) and measures 46.5 mm total length and 38.5 mm standard length (Nielsen, 1974; personal observations). The paratypes number 5 (in Nielsen (1974)), or 6 (in Bruun and Kaiser (1948)) but only 4 were found in ZMUC in December 1999. Paratypes (ZMUC P 26476, 26477, 26478, 26480) measure 19.5-42.0 mm total length and 16.5-34.5 mm standard length according to Bruun and Kaiser (1948). Two fish (P 26476 and P 26480) were used in histological studies and one consists of the body only. The type locality is given below and the fish were collected by E. Kaiser on 6-5-1937 from "lok 80" (= locality 80; but no field notes by E. Kaiser are available in ZMUC).

The date of authorship for this species is variously listed as 1943 on an official reprint, as 1944-49 in one set of Contents and "ready from the press 1944" in another set of contents. Proudlove (2006) states that is did not appear until 1948 because of World War II.

Bruun and Kaiser (1948) believe this species to be related to the genus "Barbus", members of which also have two pairs of barbels, although Saadati (1977) considers this unlikely since most "Barbus" from the Tigris River basin are large fishes.

Key characters

The only eyeless, depigmented cyprinid species in Iran, it is very distinctive.

Morphology

The body is compressed and the head somewhat flattened. There are two pairs of barbels, one pair at the mouth corners and one about half way along the upper lip. The upper lip has a feebly crenulated edge. The mouth is subterminal and horseshoe-shaped. A mental disc is developed in some fish, absent in others. There are significant differences in some morphometric and meristic characters in fish with, and without, a disc (Sargeran et al., 2008). There is no visible trace of eyes in most fish. The skin is naked except for a few rows of scales behind the pectoral fin base, although some individuals may have more flank scales. There are about 32 myomeres along the flank. A lateral line is present. The dorsal fin has 3 unbranched and 7-8 branched rays, the anal fin 3 unbranched and 4-5 branched rays, the pectoral fin 14-17 branched rays and the pelvic fin 5-7 branched rays. Pharyngeal teeth in 3 rows, 1 to 3 in the outer row, 3 to 4 in the middle row and 3-5 in the inner row. Anterior teeth are very enlarged and conical, appearing as rounded knobs while the posterior teeth in the main row are flattened and slightly hooked. Smaller fish have less conical anterior teeth with a tiny hook at the tip and posterior teeth have a short, flat to slightly concave surface below the tip. Tooth counts are difficult to make with accuracy as it is not always clear to which row a tooth belongs. Smaller fish can be interpreted as 2,3,5-5,3,2 while larger fish may possibly lose a tooth and have a 2,3,4-4,3,2 count. Gill rakers very short, not reaching the adjacent raker when appressed and numbering 10-13 total. Total vertebrae 34-36. Gut s-shaped.

Meristics in specimens examined, including the holotype and 2 paratypes: dorsal fin branched rays 7(30) or 8(29); anal fin branched rays 4(1) or 5(49); pectoral fin branched 12(1), 13(9), 14(8), 15(23) or 16(8); pelvic fin branched rays 6(18), 7(31) or 8(1); total gill rakers 10(2), 12(4) or 13(1); pharyngeal teeth 2,3,5-5,3,2(1), 3,4,5-5,4,2(1), 2,3,4-5,3,2(1) or 2,3,4-4,3,1(1); total vertebrae 34(3), 35(8) or 36(1).

Sexual dimorphism

None reported.

Colour

This species is almost entirely unpigmented although live fish are pinkish to red from the blood showing through the skin. The gill filament area is bright red and some fish give an overall impression of red like a goldfish. Small, black pigment cells were visible in two small fish over the brain and just behind it and in these two fish and three others a very small, black pigment spot deep in the tissues on the side of the head may indicate a rudimentary but non-functional eye. Gut contents are visible through a semi-transparent body wall. Preserved fish are yellowish-white.

Size

Reaches 55.0 mm total length (Kiavash Golzarian, pers comm., 6 April 2008).

Distribution

Found only at "Kaaje-ru" above the garden "Bagh-e Loveh", "Lowa" or "Levan" (probably Loven at 33°04'N, 48°37'E) which is about 4 km from kilometre 382 on the railway from Bandar Shapur to Tehran and approximately 12 km north of the railway station Tang-e Haft. The stream below the cave locality is the "Ab-e Serum" which runs into the "Ab-e Zezar" which is a tributary of the Dez River, in Lorestan Province. Further locality details are given in Bruun and Kaiser (1948). The locality is at 744 m and 33°04'38.6"N, 48°35'33.1"E according to the Iranian Fisheries Research and Training Organization Newsletter, 21:3, 1998 and Kiavash Golzarian, pers comm., 6 April 2008).

Zoogeography

The relationships of cave species, with their reduced characters, are problematical but the three rows of pharyngeal teeth and mouth structures indicate a possible relationship with Garra.

Habitat

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

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

The fish may be seen swimming freely in the well, up to a 20 at a time may be counted. They can be caught with a dip-net.

Sampling in December 2000 recorded a water temperature of 18.5°C, pH 7.5 and a conductivity of 334 µS. Aquarium specimens have been maintained at 5-28°C and were very resistant to changes in oxygen levels (R. Mehrani, (pers. comm., 2000). Amir Hosin Zalaghi recorded the following parameters on 19 May 2010:-

pH 7.3, 18.0ºC, conductivity 506.0 µS/cm, TDS 255.0 mg/l, CO₃ 0.0 mg/l, HCO₃ 152.0 mg/l, Cl 35.0 mg/l, SO₄ 65.0 mg/l, Ca 59.1 mg/l, Mg 27.8 mg/l, K 0.0 mg/l, Th 250.0 mg/l, turbidity 0.65 NTU, COD 3.0 mg/l. BOD 0.0 mg/l, total alkalinity 101.0 , alkalinity-f 0.0, DO 7.9 mg/l, TSS 0.5 mg/l, NO₃ 0.5 mg/l, NO₂ 0.0 mg/l and Na 12.0 mg/l.

Cave locality with R. Mehrani, 4 December 2000

Cave locality showing friable rock surrounds

Age and growth

Unknown although R. Mehrani (per. comm., December 2000) kept fish in aquaria for 18-24 months.

Food

Unknown but the aquarium specimens referred to above were fed Artemia, dried and fine-ground Gammarus, zooplankton and phytoplankton. Faecal contents were phytoplankton and one fish was observed to scrape the aquarium wall. Occasionally aquarium fish will swim upside down with the snout at the water surface and may be feeding on an algal film. Fish with a disc can attach to and graze on the substrate; a significantly longer intestine in such fish may be indicative of a detrital feeding habit (Sargeran et al., 2008).

Reproduction

Unknown.

Parasites and predators

None reported from Iran.

Economic importance

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

Conservation

A fine of 10,000 rials (U.S.$139.94, 15 March 1978) was imposed specifically for illegal fishing of this species (Anonymous, 1977-1978), now 100,000 rials (U.S.$11.04, 7 April 2008). It is on the 1994 IUCN Red List of Threatened Animals as one of two rare fish species from Iran (see also Paracobitis smithi) and is on the 2000 IUCN Red List and subsequent ones as VU D2 (Vulnerable, acute restriction in its area of occupancy; see also Proudlove (2001)). The habitat is of unusual importance for studies on evolution in unique environments. Coad (2000a), using 18 criteria, found this species to be one of the top 4 threatened species of freshwater fishes in Iran.

B. Sandford (in litt., 1979) considered this fish to be endangered. The cave appeared to be a recently collapsed system and the network of fissures could be quite small. The main pool is at the end of a narrow cleft, overhung by a cliff of friable shale. Shale fragments fall spontaneously and the nearer end of the pool has a floor of shale fragments. Coupled with recent collecting the number of extant specimens may be quite low but this is impossible to confirm.

Local informants in December 2000 estimated that 5-6 parties visit the site each year. The number of specimens taken is unknown but an estimated 66+, possibly more than 100, have been collected in recent years (from 2000 to 2008). Eight specimens are referred to in the literature, 4 specimens were caught in 1998 (R. Mehrani, pers. comm., 2000), in the two years 1999-2000 13 specimens were collected by one party, 18 by another in December 2000 (R. Mehrani and IFRO staff, N. Najafpour, IFRO, F. Razi, Darabad Museum, Tehran and B. W. Coad), 10 specimens by Ali Ebrahimi (pers. comm, 25 January 2006), 11 by Kiavash Golzarian (pers comm., 6 April 2008), and more than 10 by others.

Four fish collected in 1998 survived 2 years in an aquarium (R. Mehrani, pers. comm., December 2000). They were fed on Artemia, zooplankton, phytoplankton and fine-ground Gammarus. Water temperature ranged from 5 to 28°C and resistance to changes in oxygen levels was high. Fish were sometimes observed to swim upside down at the water surface.

The establishment of a small park or reserve around the site and education of the local people to maintain a watch on the cave would be most useful to protect this species, and the other cave species at this site, from unauthorised collectors. A survey of the local people and the Department of the Environment files should be made to determine the numbers of visitors to this remote site.

Further work This is a small species of fish of unusual appearance and provenance and could be bred and sold as an aquarium and experimental species, providing that numbers at the site warrant removal of breeding stock. If successful, this would ensure survival of the species. Captures at present appear to be fortuitous and give no real picture of the population size; removal of more specimens would have to be carefully planned and monitored. Surveys of groundwater recharge in the area and a more thorough investigation of the cave system should be undertaken to assess the status of the habitat.

Sources

Movaghar (1973) is an additional reference, in Farsi, on this species.

Type material: See above (ZMUC P 26475, P 26476, 26477, 26478, 26480).

Iranian material: CMNFI 2007-0124, 8, 27.3-42.2 mm standard length, Lorestan, type locality as above.

Genus Kosswigobarbus
Karaman 1971

Much of the past literature on this genus appeared under Barbus (q.v.)

Kosswigobarbus kosswigi
(Ladiges, 1960)

Common names

ابوحنج (abu henej or abu hanaj= father of the hook or spine; possibly abu hanash or abu henesh, father of the snake), shebeh shirbot.

[Kosswig's barbel].

Systematics

This species was described as Cyclocheilichthys kosswigi Ladiges, 1960 from the "Batman suyu" (the holotype is possibly a female, 162.7 mm, Turkey, Siirt Province, Batman suyu (the Batman stream enters the Tigris River at 37°47.30"N, 41°00'E near Batman), April 1939, C. Kosswig). The holotype is in the Zoologischen Instituts und Zoologischen Museums der Universität Hamburg (ZMH H1148). The genus Cyclocheilichthys Bleeker, 1860 is found only in Southeast Asia.

A new and monotypic genus, Kosswigobarbus, was erected for this species by Karaman (1971) but this was synonymised with Barbus by Coad (1982f). However Bănărescu (1997) and Ekmekçi and Banarescu (1998) consider Kosswigobarbus to be valid. Borkenhagen (2005) considers that kosswigi and sublimus should be placed in Barbus (Carasobarbus) or Carasobarbus, favouring treating Carasobarbus as a subgenus until Barbus is revised thoroughly. Karaman (1971) distinguished the genus on the basis of the fin ray characters, a well-developed rostral flap, numerous fine pores on the head, and a large lachrymal bone. These characters are found in other "Barbus" species in Southwest Asia and the whole complex of large-scaled Barbus requires a detailed revision (see also under Carasobarbus luteus and K. sublimus).

The relationships of this species appear to lie with other Southwest Asian species formerly in the catchall genus "Barbus" which possess a compressed body, large scales with counts of 38 or less in the lateral line, a smooth dorsal fin spine, 9 or more branched dorsal fin rays and 6 anal fin branched rays such as Barbus apoensis Banister and Clarke, 1977, B. exulatus Banister and Clarke, 1977 (both in Southwest Arabia), B. chantrei (Sauvage, 1882) and B. canis Valenciennes, 1842 (both from the Levant), B. luteus (Heckel, 1843) from the Tigris-Euphrates and neighbouring basins, and B. sublimus Coad and Najafpour, 1997 from Khuzestan in the Tigris-Euphrates basin of Iran. Borkenhagen (2005) regards these species as a monophyletic group characterised by 6 branched anal fin rays, smooth and ossified last unbranched dorsal fin ray, modally 10 branched dorsal fin rays, less than 40 scales in the lateral line, medium body size, large, round to shield-shaped scales with numerous parallel radii, pharyngeal teeth usually 2,3,5-5,3,2, gill rakers short, stout and slightly curved, and barbels short.

Key characters

This species is characterised by having two pairs of thin barbels, 6 branched anal fin rays, the last unbranched dorsal fin ray strong and sharp-edged but smooth, 9-11 branched dorsal fin rays, large scales (38 or less in the lateral line), and a deep and compressed body.

Morphology

The rostral flap shows variable development, sometimes overlapping the upper lip to become visible in ventral view and other times not so well developed. Lips are thick, continuous and fleshy and there is a large median lobe to the lower lip. The mouth is small, ventral and u-shaped. The dorsal fin origin lies over or slightly in advance of the pelvic fin origin. The dorsal fin margin is strongly concave and the last unbranched dorsal fin ray is a strong spine without teeth. The caudal fin is deeply forked. The anal fin is long and may overlap the caudal fin base.

Scales are regularly arranged over the body. A low sheath of scales is found at the base of the anal and dorsal fins, being most evident anteriorly, and enclosing the anal papilla. There is a pelvic axillary scale. Anterior scale radii are few (5-11 in five scales from one specimen 126.6 mm SL) while posterior radii are numerous (35-40). There is a scaled keel or ridge before the dorsal fin as the back narrows dorsally. Pharyngeal tooth formulae 2,3,5-5,3,2, 2,3,5-4,3,2, 2,3,4-5,3,2 and 2,3,4-4,3,2. The teeth are quite small even in the largest specimens. Teeth are hooked at the tip and strongly recurved there, teeth are conical and have a small, concave to irregular or even rounded grinding surface below the tip. The fifth and most anterior tooth in the main row is small to minute in most fish and may be absent but this is not size related as both large and small specimens have or lack this tooth. The gut is elongate and coiled.

Meristic data from Iranian and other Tigris-Euphrates specimens: dorsal fin branched rays 9(5), 10(34), 11(1), mean 9.9 after 4 unbranched rays; anal fin branched rays 6(40) (not 7 as in the original description) after 3 unbranched rays; pectoral fin branched rays 15(2), 16(6), 17(4), 16.1; and pelvic fin branched rays 7(1), 8(11). Lateral line scales 29(1), 31(2), 32(1), 33(2), 34(6), 35(3), 36(1), 37(1), 38(1), 41(1) mean 34.2 (Kuru's (1975) range is 32-36); scales above the lateral line 6(7), 7(10), 8(1), mean 6.7; scales below the lateral line 5(5), 6(13), mean 5.7; scales between lateral line and pelvic fin 4(11), 5(1), mean 4.1; predorsal scale rows 11(1), 12(2), 13(5), 14(3), 15(1), mean 13.1; and caudal peduncle scales 13(2), 14(2), 15(6), 16(2), mean 14.7. Total gill rakers 10(2), 11(2), 12(3), 13(3), 14(1), mean 11.9. Pharyngeal teeth 2,3,5-5,3,2(4), 2,3,4-4,3,2(3) or 2,3,4-5,3,2(1). Total vertebrae 39(5), 40(4), mean 39.4.

Caudal peduncle length in head length 1.2-1.3, mean 1.3; caudal peduncle depth in caudal peduncle length 1.5-1.7, mean 1.6; pelvic fin length in standard length 4.6-5.4, mean 5.0; pelvic fin length in pelvic fin origin to anal fin origin distance 1.2-1.6, mean 1.3; dorsal fin spine length in head length 0.8-0.9, mean 0.9; and longest dorsal fin ray in head length 0.9-1.0, mean 0.9.

Sexual dimorphism

Sample sizes are too small to investigate accurately.

Colour

Upper flank scales are outlined by pigment, most evidently anteriorly on each scale. Fins are lightly pigmented with scattered melanophores on both rays and membranes with some concentration on dorsal fin membranes although the extent varies individually. The peritoneum is black.

Size

Reaches 19.4 cm total length (the holotype).

Distribution

Found in the Tigris-Euphrates basin of Turkey and Iran (Coad, 1982f; Coad and Najafpoiur, 1997; Abdoli, 2000). It may also occur in the Zohreh River (Gh. Izadi, pers. comm., 2001).

Zoogeography

Karaman (1971) considers that the closest relatives of this species are to be found in the Indo-Malayan region.

Habitat

This species is found in large rivers in Iran which, however, in mid-summer are more stream-like in water flow. Collections are from the plains of Khuzestan and from altitudes in excess of 1600 m in the Zagros Mountains. Temperatures in early July range from 21 to 23°C. One locality was polluted and others were cloudy or muddy. The river beds are composed of stones.

Age and growth

Unknown.

Food

The elongate gut and black peritoneum suggest a plant component to the diet but examination of two gut contents reveal insect remains including chironomid larvae.

Reproduction

Unknown.

Parasites and predators

Sohrabi and Jalali (2002) report the nematode Schulmanella petruchewskii from the liver of this species caught in the Dez River.

Economic importance

This species is too rare in Iran to be of any economic importance.

Conservation

Recommendations are difficult to make since the ecological requirements of this species are unknown. It appears to be rare but this may only be inadequate sampling techniques. Further collections in addition to the holotype have been made in southern Anatolian Turkey (Kuru, 1978-1979) but it does not seem to be common. Endangered in Turkey (Fricke et al., 2007).

Further work

Intensive field work utilising a wide variety of techniques should be directed to determining the abundance and distribution of this species. An adequate material base would then enable ecological studies to be carried out and conservation measures determined.

Sources

Some counts from Kuru (1975) on Turkish material.

Type material: See above, Cyclocheilichthys kosswigi (ZMH H1148).

Iranian material: CMNFI 1979-0275, 1, 126.6 mm standard length, Lorestan, Kashkan River drainage (33º25'N, 47º58'E); CMNFI 1979-0277, 1, 116.5 mm standard length, Lorestan, Kashkan River drainage (33º30'N, 47º59'E); CMNFI 1979-0289, 1, 103.5 mm standard length, Kermanshahan, Diyala River drainage (34º28'N, 45º52'E); CMNFI 1979-0290, 2, 120.1-122.1 mm standard length, Kermanshahan, Diyala River drainage (34º31'N, 45º35'E); CMNFI 1979-0368, ?, ? mm standard length, Khuzestan, Karkheh River (32º24'30"N, 48º09'E); uncatalogued, 1, 173.3 mm standard length, Khuzestan, Karkheh River near Shush (no other locality data); uncatalogued, 3, 140.1-179.0 mm standard length, Khuzestan (no other locality data); uncatalogued, 1, 160.0 mm standard length, Khuzestan, Karun River basin near Izeh (no other locality data).

Comparative material: BM(NH) 1974.2.22:1281, 1, 31.2 mm standard length, Iraq, Al Hadithah (34º07'N, 42º23'E); BM(NH) 1974.2.22:1292-1296, 4, 35.5-98.5 mm standard length, Iraq, Al Hadithah (34º07'N, 42º23'E).

Kosswigobarbus sublimus
(Coad and Najafpour, 1997)

Common names

None.

Systematics

The holotype is CMNFI 1995-0009, female, 113.5 mm, Iran, Khuzestan, A'la River at Pol-e Tighen, 31°23.5'N 49°53'E, 20 September 1995, B. W. Coad, N. Najafpour and party. Paratypes are CMNFI 1995-0009A, 41.9 mm, same locality as the holotype (lost in the mail while on, loan September 2005), CMNFI 1995-0010, female, 115.3 mm, A'la River, 2 km above Pol-e Tighen, 31°23.5'N 49°54'E, 20 September 1995, B. W. Coad, N. Najafpour and party, and CMNFI 1995-0011, 3 females, 90.5-98.6 mm, same locality as holotype, early December 1994, Gh. Eskanderi (one specimen lost in the mail while on loan, September 2005).

The species was named after its river of capture, the only known locality for this species. A`la means "most high" or "exalted".

Key characters

A member of the genus Barbus sensu lato characterised by the unique combination of the following characters: large scales (24-27 in the lateral line), 37-38 total vertebrae, 10-11 branched dorsal fin rays, 6 branched anal fin rays, a relatively short and smooth dorsal fin spine (spine length in head length 1.0-1.1), lower lip with a rounded median lobe and a posterior free flap, a compressed body (depth 3.3-3.5 in standard length), a short caudal peduncle (length in head length 1.5), long pelvic fins (length in standard length 4.1-4.5), and a short dorsal fin (longest dorsal fin ray in head length 1.1-1.2).

Morphology

Dorsal fin branched rays 10(5), 11(1), mean 10.2; anal fin branched rays 6(6); pectoral fin branched rays 14(1), 15(5), mean 14.8; and pelvic fin branched rays 8(6). Lateral line scales 24(1), 25(2), 26(2), 27(1), mean 25.5; scales above the lateral line 4(1), 5(5), mean 4.8; scales below the lateral line 4(3), 5(3), mean 4.5; scales between lateral line and pelvic fin 3(6); predorsal scale rows 9(5), 10(1), mean 9.2; and caudal peduncle scales 12(5). Total gill rakers 10(1), 11(1), 12(2), 15(1), mean 12.0. Total vertebrae 37(2), 38(4), mean 37.7. A specimen from the Khersan River had 39 total vertebrae and one from the Ardal River had 40 total vertebrae; both these fish being unusual in other counts too. Esmaeili et al. (2006) give the following characters for their 6 specimens from Fars: 11 branched dorsal fin rays, 6-8 anal fin branched rays, 16-18 branched pectoral fin rays, 24-28 lateral line scales, and 10-12 total gill rakers.

The body is relatively deep (depth 3.3-3.5 times in standard length) and compressed. The snout is rounded and overhangs the upper part of the thick upper lip. The extent of overlap varies individually. The lower lip is also thick but has a rounded protuberance at its centre, visible in lateral view. The protuberance is variably developed as a flap which is free posteriorly and at the rearmost sides. The posterior barbel is longer and thicker than the anterior barbel. The anus lies just anterior to the anal fin origin.

Scales are regularly arranged over the whole body, there is a pelvic axillary scale, and scales at the anterior base of the anal fin form a small sheath around the bases of the anal rays. Radii are found on the anterior and posterior fields of each scale, being most numerous posteriorly, about three times as many. Some radii extend into the lateral fields. Circuli are numerous and on the posterior field break up into bubble-like shapes.

The dorsal fin is slightly to strongly concave on its margin. The spine tapers and is thin and flexible at the tip. The dorsal fin origin lies over the pelvic fin origin. The caudal fin is deeply forked with the lower lobe more developed and with longer rays than upper lobe. The anal fin reaches or obviously passes the base of the caudal fin rays. This variation in length does not appear to be size or sex related. The posterior margin of the anal fin is straight to concave. The pelvic fin has a straight to rounded posterior margin. The pectoral fin margin is concave and in some fishes is falcate.

The gut is elongate with anterior and posterior loops. Gill rakers are short and reach to the adjacent raker when appressed. Pharyngeal teeth are rounded with a hooked tip and a flattened area below the tip. On three specimens counts were 2,3,5-4,3,2, 1,3,5-4,3,2 and 3,3,4-4,3,2.

Sexual dimorphism

Sample size of the type series is too small to document sexual dimorphism.

Colour

The overall live colour of the species is silvery with the back olive-green. Scales are outlined with dark pigment. The pectoral, pelvic, anal and caudal fins are a faintly pigmented with orange to yellow hues, most apparent when the fin is collapsed. Much of these fins is grey to hyaline. The dorsal fin is grey to hyaline. The eye is silvery with grey-brown pigment at the upper margin. The peritoneum is silvery with numerous melanophores merging to give an overall dark appearance.

In 70% ethanol the pigmentation pattern is as follows. Upper to mid-flank scales have the margins and bases pigmented with melanophores, outlining the scales. Most pigment is concentrated at the scale base giving a slight appearance of rows of spots. Larger fish are more fully pigmented so the back and upper flank then appear dark. The dorsal surface of the head is finely speckled black. The dorsal fin has dark pigment on the membranes, on the distal half or the whole fin, with less pigment on the rays. The caudal fin is mostly hyaline with dark pigment lining the rays. The pectoral and anal fins have some dark pigment lining or on the anterior rays and, in larger fish, on the membranes. The pelvic fin is hyaline. The smallest specimen has a distinct mid-caudal base spot and another spot on the back at the anterior dorsal fin base. Fins are more hyaline than in larger fish.

Size

Reaches 115.3 mm standard length. The maximum size is 15.5 cm (Iranian Fisheries Research and Training Organization Newsletter, Tehran, 18:5, 1997).

Distribution

Known from the A'la River in Khuzestan Province in the Tigris River basin and the Fahlian River in Fars (Esmaeili et al., 2006).

Zoogeography

This species is known from the A`la River, which joins with the Rud Zard (rud = river), and emerges from the foothills of the Zagros Mountains onto the Khuzestan plains where it is tributary to the Jarrahi River. The Jarrahi feeds the Shadegan Marshes and is mostly lost there. In flood times, there may be a connection through the marshes to the Karun River and thence to other large river systems in the Tigris-Euphrates basin. However, it is suspected that the ecological requirements of this species limit it to fast flowing rivers over hard substrates and the marsh system isolates it from other river systems. Collections in the Rud Zard at Rud Zard village and Bagh-e Malek on several occasions have not included the new species although the Rud Zard would appear to be a suitable habitat.

The range extension of 380 km southwest of the A`la River to the Fahlian River near Noorabad in Fars places this species in the headwaters of the Zohreh River which drains to the northern Persian Gulf. This may indicate headwater captures or possibly former interdigitating drainages on the Khuzestan plain.

Habitat

The type habitat is a cloudy river in a wide flood plain at about 800 m. The river bed is stones and pebbles. Water is led off from the river at intervals to irrigate the rice fields of the villages of Meydavud-e `Olya (31°24'N. 49°52'E) and Meydavud Pa'in (31°23'N. 49°49'E) which extend along the bank of the A`la River. This water abstraction is a potential threat to the well-being of fishes in this river system. The water demands of rice growing are large and there is little or no rain through the summer months in this area. Air temperatures in September can exceed 40°C and evaporation from the fields and the river is commensurate.

The fish were caught at the type locality in relatively fast water (0.9 m.s^-1) over a one hour fishing period. In September 1995, the river was at the seasonal low water and the type locality was 10 m wide, 40 cm deep and had a discharge of ca. 2.9 m³.s^-1. The water was also cloudy for the collection in December 1994 at the type locality but the river was wider and had more flow after rain. The second locality had more flow and was deeper and wider than the type locality, to about 30 m and 80 cm. Fishes were caught by electroshocker and cast-net and were difficult to catch and few in number. Other species captured were the cyprinids Luciobarbus barbulus, Tor grypus, Barilius mesopotamicus, Capoeta trutta, Cyprinion macrostomus and Garra rufa, and the sisorid catfish Glyptothorax silviae.

The Fahlian River capture site was shallow, had relatively clear water, a heterogenous bed morphology (sand, gravel, stone, pebbles, rock, etc.), and an absence of aquatic and riparian vegetation (Esmaeili et al., 2006).

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

None reported.

Economic importance

None reported.

Conservation

Known only from the type series, its conservation status is unknown. It appears to be rare and possibly restricted to areas with running water year round.

Further work

The distribution of this species should be investigated within the A'la River and neighbouring drainages and accurate estimates of its numbers and biology procured as part of a process to determine its conservation status.

Sources

Type material: See above, CMNFI 1995-0009, CMNFI 1995-0009A, CMNFI 1995-0010, CMNFI 1995-0011.

Genus Labeo
Cuvier, 1816

Labeo dero
(Hamilton, 1822)

Mashkel (= Mashkid) River basin in Pakistan on the southeastern border of Iran (Zugmayer, 1913; Mirza, 1971; 1972; 1974; 1975; 1992). No Iranian record.

Labeo gedrosicus
Zugmayer, 1912

Mashkel (= Mashkid) River basin in Pakistan on the southeastern border of Iran (Zugmayer, 1912; Mirza, 1971; 1972; 1974; 1975; 1992; Neumann, 2006). No Iranian record.

Labeo macmahoni
Zugmayer, 1912

Makran basin (Dasht River) in Pakistan on the southeastern border of Iran (Zugmayer, 1912; Mirza, 1971; 1972; 1974; 1975; Mirza and Saboohi, 1990; Neumann, 2006). Placed in a new subgenus, Tariqilabeo by Mirza and Saboohi (1990). Kullander et al. (1999) and Mirza and Arshad (2008) consider this taxon to be, questionably, a synonym of Crossocheilus diplocheilus (Heckel 1838). A syntype is in the Naturhistorisches Museum Vienna (NMW 81256). No Iranian record.

Genus Leucaspius
Heckel and Kner, 1858

The genus Leucaspius has not been recently revised in detail and its composition remains uncertain. There may be several species in Europe but only one has a wide distribution and this is found in Iran.

The genus is characterised by a moderately compressed and elongate body; an incomplete lateral line on up to about 13 scales; moderately large, easily detached scales; short dorsal and somewhat longer anal fin; belly without a keel but somewhat compressed; terminal mouth with lower jaw entering the depression of the upper; pharyngeal teeth usually in 2 rows; and gill rakers of moderate size and density.

Leucaspius delineatus
(Heckel, 1843)

Common names

mahi-ye riz-e noqrei or mahi-e-rize-noghreie (= small silvery fish).

[gafgaz ustuzani in Azerbaijan; ovsyanka, verkhovka, Kavkazskaya verkhovka or Caucasian verkhovka in Russian; sunbleak, white aspe, rain bleak; belica; Moderlieschen in German].

Systematics

Squalius delineatus was originally described from Wien and Mähren, Austria. The Caspian Sea basin taxon is given by Berg (1948-1949) as Leucaspius delineatus delineatus natio caucasicus Berg, 1949, described from Transcaucasia, which is distinguished by a lower average dorsal fin branched ray count (7-8 rather than 8 or rarely 9 for the typical form of Europe). This natio has no taxonomic standing but has been applied as a subspecies by some authors (Arnold and Längert, 1995).

Key characters

The large, rounded papillae around the genital opening are distinctive in females, and for both sexes the combination of an incomplete lateral line with moderately large scales is distinctive.

Morphology

Dorsal fin with 2-3, usually 3, unbranched rays followed by 7-10 branched rays (usually 8 in Europe but counts of 7 and 8 are about equally frequent in the Caucasian populations according to Berg (1948-1949) but Abdurakhmanov (1962) gives a frequency of 94% for 8 rays and only 6% for 7 rays in fish from Azerbaijan), anal fin with 3-4, usually 3, unbranched rays followed by 9-17 branched rays (10-12 in the Caucasian subspecies), pectoral fin branched rays 11-16 and pelvic fin branched rays 7-8. Lateral series scales 36-53; lateral line incomplete with 0-13 pored scales anteriorly. Scales bear few anterior and posterior radii, have few circuli, a subcentral anterior focus and are a vertical oval in shape. Gill rakers 10-17 (rarely 20, usually 13-16), reaching the second raker below when appressed. Vertebrae 36-40. Pharyngeal teeth very variable 5-5, 5-4, 4-4, 4-5, 1,5-5, 5-5,1, 1,5-4, 5-4,1, 1,4-4, 4-4,1, 1,5-5,1, 1,5-5,2, 1,4-4,1, 1,4-5,1, 1,5-4,1, 1,5-4,2, 1,4-5,2, 2,5-4,1, 2,5-4,2, 2,5-5,2 and even 2,5-6,1. The frequency of various counts varies with locality, and even whether single row counts dominate over two-rowed counts (Arnold and Längert, 1995). Teeth are hooked at the tip and slightly to strongly serrated. The belly is compressed in the mid-line between the pelvic fins and the vent but does not form a strong keel. The gut is an elongate s-shape. The chromosome number is 2n=50 (Klinkhardt et al., 1995).

Meristic values for Iranian specimens are:- branched dorsal fin rays 8(10) or 9(1); branched anal fin rays 10(4) or 11(7); branched pectoral fin rays 11(7), 12(3) or 13(1); branched pelvic fin rays 7(10) or 8(1); scales in lateral series 39(3), 40(5), 41(1), 43(1) or 45(1); total gill rakers 13 (4) or 14(7); and total vertebrae 36(1) or 37(9). One specimen showed fusions of abdominal vertebrae.

Sexual dimorphism

Females have a unique fold of skin in the shape of two, large, rounded papillae around the genital opening. The male is a little smaller than the female. Males develop prominent nuptial tubercles on the dorsal head surface, snout, on the lower jaw in three pairs and on the upper jaw in two pairs for a total of about 60 tubercles. The male genital opening is depressed.

Colour

The back is olive-green to brown and the flanks and belly silvery-white. A steel blue or bluish-green stripe begins at the rear third of the body and extends back, broadening, to the tail base. Fins are hyaline or slightly yellowish. The peritoneum is light.

Size

Attains 12 cm total length although only up to 5.6 cm total length in the Caucasian form.

Distribution

Found in western and central Europe from the Rhine and north of the Alps east to northern drainages of the Black Sea and the western and northern drainages of the Caspian Sea. Leucaspius delineatus caucasicus is found in the north Caucasus including the Black Sea parts and in Transcaucasia. In the southern Caspian Sea basin, it is found in the lower reaches of the Kura River, Imeni Kirova Bay and the Lenkoran region of Azerbaijan (Kuliev, 1989). A single specimen from Iran was collected by Mr. Akbar Nasrollazadeh near Siah Darvishan (which is at 37°22'N, 49°26'E) in Gilan on 27 May 1993. In June 1996 over 50 specimens were caught in the Anzali Mordab (= Talab) by K. Abbasi and A. Sarpanah of the Gilan Fisheries Research Centre (Iranian Fisheries Research and Training Organization Newsletter, 15:4, 1997). Also reported from the Anzali Talab by Abbasi et al. (1999) and present in the Safid River and Amirkelayeh Wetland (Nasrollazadeh, 1999; K. Abbasi, pers. comm., 2001).

Zoogeography

The Caspian shore of Iran has been surveyed in some detail during the 20th century and it is curious that this species was only discovered towards its end. It may simply have been confused with other small, silvery minnows although it should be noted that some of the surveys were carried out by Russian workers familiar with this species. It may be a recent introduction with other, commercial exotics, and therefore may not be from a Caucasian population.

Habitat

Found in still or slowly flowing water with vegetated shores in large schools. It can be found in fish ponds, ditches, gravel pits and quarries as well as natural habitats. Still water is required for reproduction. It is tolerant of a wide range of temperatures, pH and salinity depending on adaptation, e.g. temperature range of 3-32.8°C (Arnold and Längert, 1995). This small fish is found in large schools near the water surface. It may appear in small ponds without any apparent connection to other water bodies, hence the German name that has been interpreted as "Moderlieschen" or motherless. However, the German name may more correctly mean mud lover (G. H. Copp, in litt., 16 June 2004).

Age and growth

Life span is about 4-6 years with growth fairly continuous over this period.

Food

Diet comprises plankton such as cladocerans, copepods and rotifers, benthic chironomids, flying insects which land on the water surface, and also some algae and detritus.

Reproduction

There is often a spawning migration against the water flow (up to 2-3 m/sec) to new waters. Eggs are laid in strings which are wound spirally around plants by the female, aided by the fold of skin around the genital opening. They may also be laid in a disc-shaped patch on any flat surface. Several spawnings occur over a few weeks in March to September in Europe. The eggs are guarded and fanned by the male who covers them with a bacteriostatic dermal mucus. Up to 485 eggs are found in females and have diameters up to 0.5 mm in Azerbaijan, up to 3500 eggs and 1.5 mm in Europe. Maximum egg production over two seasons is about 500-600 (Abdurakhmanov, 1962). Clutch sizes are about 50-350 eggs (Arnold and Längert, 1995). A minimum temperature of 18°C is required for reproduction.

Parasites and predators

None reported from Iran but eaten by a wide variety of other fishes in Europe and numerous parasites reported (Arnold and Längert, 1995).

Economic importance

The scales have been used in the production of artificial pearls as with Alburnus alburnus (a relative of A. hohenackeri). It has also been used in aquaria and garden ponds and as bait by anglers.

Conservation

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

Further work

More specimens from Iranian waters need to be examined to determine if this species belongs to a distinct subspecies or is an exotic population of the European type subspecies. Biology of the Caucasian subspecies is unknown.

Sources

Arnold and Längert (1995) summarise biology of European populations in detail.

Iranian material: Uncatalogued material, 11, 31.2-37.5 mm standard length, Gilan, swamp near Hendeh Khaleh (37º23'N, 49º28'E).

Genus Luciobarbus
Heckel, 1843

Much of the past literature on this genus appeared under Barbus (q.v.)

Luciobarbus barbulus
(Heckel, 1849)

Common names

lab pahn (= broad lip), برزم (berzem or barzam); berzem lab pahn in Khuzestan and Iraq to distinguish it from B. pectoralis; boz mahi (= goat fish) or سس ماهي (= sos or sas mahi in the Dalaki and Shapur river basins); dolenj.

[abu-barattum (= owner or father of lips), abu baratem, abu bratum or nabbash in Arabic; Orontes barbel].

Systematics

Howes (1987) places this species in Barbus sensu stricto. Karaman (1971) places this species in the synonymy of Barbus rajanorum but other authorities consider it to be Luciobarbus pectoralis (q.v.). Almaça (1983) placed this species as a subspecies of Barbus mystaceus but later (1984a, 1984b, 1986, 1991) retained barbulus as a full species, known only from the Levant, despite Heckel's record from both the Qarah Aqaj (= Mand) of Fars, Iran and the Quwayq (= Kueik) River of the Levant. I retain it as a species under Heckel's name for this taxon until the systematics of this and related species can be worked out as indicated above. It is separated from mystaceus according to Almaça (1983) by having thinner lips, shorter barbels, the last unbranched dorsal ray weaker and shorter, more dense denticles spread over a shorter length of ray, higher anal fin, gill rakers less numerous and the upper dorsal profile is rectilinear and oblique to the back.

The type locality of Barbus Barbulus is the "Fluss Kara-Agatsch....bei dem Dorfe Geré" (= Qarah Aqaj or Mand River, Fars; possibly near Kereft, 29°01'N, 52°52'E) and presumably the "Kueik bei Aleppo" (Heckel, 1846-1849b). J. Valiallahi, pers. comm., 2001 and Edmondson and Lack (2006) suggest Jereh at 29°15'N, 51°58'E but this is in the Hilleh River drainage, a Dalaki River tributary. In addition, "Geré" takes a hard G in German, not a J. There may be some confusion of names and rivers here.

A possible syntype of barbulus from the Qarah Aqaj was located by Almaça (1983, 1986) in the Naturhistorisches Museum Wien (NMW 53957) and seen by me but is in too poor condition to be of much value, being mostly bones. Another syntype is listed as NMW 6596 and measures 119.3 mm standard length. In 1997, this was the only syntype recognised and is possibly the same as NMW 53957 re-numbered as the latter was not located in 2002. The catalogue in Vienna lists only 1 fish, while Heckel's description refers to several fish. NMW 6596 is mostly bones and is dried. The fleshy lip fold of the original description could not be discerned, teeth are missing and the dorsal fin is broken off short.

"Syntypes" of mystaceus are in the Naturhistorisches Museum Wien from Mosul on the Tigris River (NMW 16472 (1 specimen), NMW 50394 (2), NMW 54384 (2)) and NMW 54385 (2) but note that authors such as Karaman (1971) and Almaça (1983, 1991) refer the species description to Heckel (1843) in error. These were not marked as being syntypes as observed on a 1997 visit to Vienna.

Key characters

This species is characterised by having two pairs of barbels, a serrated and very strong dorsal fin spine similar to that in Capoeta trutta in its proportions relative to the body, usually 8 dorsal fin rays (never 10), fleshy lips, and 47 or more lateral line scales.

Morphology

The inferior mouth is moderate in size, with moderate to thick lips and with or without a median lower lip lobe. Some fish have very thick lips so a central lobe is apparent. Some show such a degree of lip development as to appear almost abnormal while fish of similar size or larger lack this hypertrophy. In the latter case, the anterior head may be bluntly rounded and foreshortened rather than having an almost straight upper margin tapering to a pointed end. Barbels are relatively thin, occasionally quite thick. The anterior barbel does not extend past the anterior eye margin level and the posterior one not past the posterior eye margin in all sizes of fish. Rarely the anterior barbel extends to mid-eye level and the posterior one almost to the anterior operculum margin.

Dorsal fin with 4 unbranched and 8-9, usually 8, branched rays, anal fin with 3 unbranched and 5 branched rays. The last unbranched dorsal fin ray is usually very strong with a moderate density of denticles extending along much of the ray but its strength is variably developed. Pectoral fin branched rays 17-19, pelvic fin branched rays 8-9, usually 8. Lateral line scales 47-59. Scale focus subcentral anterior, many fine circuli, and numerous radii on all fields, curved in the lateral fields. A pelvic axillary scale is present but not strongly developed or apparent. Gill rakers 14-24, reaching the second raker when appressed. The interior raker surface may be covered with spinules, the internal base is heavily tubercular and the tips may become club-shaped. Pharyngeal teeth 1 or 2,3,4 or 5-5 or 4,3,2 or 1, hooked at the tip but spoon-like below with the fourth tooth of the inner row molariform, with or without a blunt projection (hooked in small fish) and much larger than the third, and the fifth tooth very small and rounded and sometimes absent apparently independent of size. The gut is elongate and complexly coiled with one anterior and 3 posterior loops. Total vertebrae 44 (Howes, 1987).

Meristics in Iranian fish are as follows: dorsal fin branched rays 8(26); anal fin branched rays 5(26); pectoral fin branched rays 17(6), 18(15), or 19(5); pelvic fin branched rays 8(25) or 9(1); lateral line scales 47(1), 48(3), 49(4), 50(4), 51(2), 52(5), 53(4), 54(1), 56(1) or 57(1); total gill rakers 15(1), 17(1), 18(4), 19(3), 20(6), 21(5), 22(2), 23(3) or 24(1); pharyngeal teeth 2,3,5-5,3,2(18), 2,3,5-4,3,2(3), 2,3,4-5,3,2(3), 2,3,5-5,3,1(1) or 2,3,4-4,3,2(1); total vertebrae ?.

Sexual dimorphism

Unknown.

Colour

The back and upper flank are brownish, the lower flank yellowish and the belly whitish. Upper flank scales are outlined with pigment, and the anterior edge of the dorsal fin and the caudal fin margin are black in preserved fish. Small fish have a few spots on the upper to mid-flank or may be profusely speckled in preservative.

Small live fish are silvery overall and have anal and caudal fins orange to bright red, especially the lower caudal fin lobe. The dorsal fin is grey and the pectoral and pelvic fins yellowish. The operculum is greenish. The lower flank is greenish-golden and the upper flank brown to grey. Large specimens are silvery with clear fins. The belly in small and large fish is white and the back grey or green to brown. The iris is silvery. The peritoneum is black.

Size

Reaches 62 cm total length (Atabak Mahjoor Azad, pers. comm., 16 June 2008); J. Valiallahi (pers. comm., 2001) believes this species reaches 1.5 m and 90 kg in the Zagros rivers of western Iran.

Distribution

Found in the Tigris-Euphrates basin, the Orontes River and the Quwayq River. In Iran it is found in the Tigris River basin (Abdoli, 2000; Ghorbani Chafi, 2000), in the Gulf basin from the Zohreh River and from the Shapur and Dalaki rivers (Gh. Izadpanahi, pers. comm., 1995), the lower Mand River (M. Rabbaniha, pers. comm., 1995), the Helleh, Dozgah, Dasht-e Palang (and its tributary the Shur) (Abdoli, 2000), in the Kor River basin (Abdoli, 2000), although not confirmed by specimens seen by me, and possibly in the Hormuz basin.

Zoogeography

Almaça (1991) believes that this species originated from a colonisation wave from South Europe.

Habitat

van den Eelaart (1954) records this species from rivers in Iraq, moving into lakes and marshes on the floods but never far from rivers.

Age and growth

Unknown.

Food

Diet is benthic organisms including insects. Large plant remains and detritus are also present in gut contents of Iranian fish.

Reproduction

Al-Habbib et al. (1986) report spawning during July and August in fish from the Tigris River at Mosul, Iraq. Reproduction in Iran has not been studied.

Parasites and predators

None reported from Iran.

Economic importance

This species is a preferred catch of anglers at Ahvaz in Khuzestan, second only to shirbot (Tor grypus).

Conservation

The population numbers of this species have not been well-studied nor has its distribution been well-documented. Since it does appear on fish markets in Khuzestan, is a large species and its habitats are under threat, it may require protection. Endangered in Turkey (Fricke et al., 2007).

Further work

The biology, distribution and population numbers of this species need investigation in Iranian waters.

Sources

Type material: See above, Barbus barbulus (NMW 6596), and note comments.

Iranian material: CMNFI 1979-0024, 1, 128.7 mm standard length, Fars, neighbourhood of Shiraz (no other locality data); CMNFI 1979-0109, 2, 91.1-91.6 mm standard length, Fars, Mand River at Shahr-e Khafr (28º56'N, 53º14'E); CMNFI 1979-0135, 1, 215.4 mm standard length, Fars, tributary to Mand River (28º08'N, 53º10'E); CMNFI 1979-0271, 1, 61.8 mm standard length, Lorestan, Kashkan River drainage (33º39'N, 48º32'30"E); CMNFI 1979-0290, 1, 139.1 mm standard length, Kermanshahan, Diyala River drainage in Qasr-e Shirin (34º31'N, 45º35'E); CMNFI 1979-0293, 1, 210.8 mm standard length, Fars, Mand River at Kavar (29º11'N, 52º41'E); CMNFI 1979-0349, 1, 126.0 mm standard length, Fars, Mand River at Kavar (29º11'N, 52º41'E); CMNFI 1979-0393, 1, 112.1 mm standard length, Khuzestan, Jarrahi River drainage (31º18'N, 49º37'E); CMNFI 1979-0497, 2, 117.4-134.4 mm standard length, Fars, Mand River at Band-e Bahman (29º11'N, 52º40'E); CMNFI 1980-0907, 1, ? mm standard length, Iran (no other locality data); CMNFI 1991-0153, 1, 230.0 mm standard length, Khuzestan, Zohreh River (no other locality data); CMNFI 2007-0109, 3, 85.1-138.7 mm standard length, Kordestan, Qeshlaq River basin south of Sanandaj (ca. 35º16'N, ca. 47º01'E); CMNFI 2007-0110, 1, 191.1 mm standard length, Kordestan, Yuzidar River basin (ca. 35º05'N, ca. 46º56'E); CMNFI 2007-0111, 1, 153.0 mm standard length, Kermanshahan, Alvand River near Sar-e Pol-e Zahab (ca. 34º36'N, ca. 45º56'E); CMNFI 2007-0113, 2, 123.9-139.6 mm standard length, Kermanshahan, Qareh Su tributary northwest of Kermanshah (ca. 34º25'N, ca. 47º01'E); CMNFI 2007-0117, 4, 43.4-155.5 mm standard length, Kermanshahan, Gav Masiab near Sahneh (ca. 34º24'N, ca. 47º40'E); uncatalogued, 1, 60.2 mm standard length, Khuzestan, Rud Zard at Rud Zard (31º22'N, 49º43'E).

Comparative material: BM(NH) 1920.3.3:23-30, 9, 80.2-98.9 mm standard length, (); BM(NH) 1931.12.21:4, 172.5 mm standard length, (); BM(NH) 1971.4.2:5, 1, 140.3 mm standard length, Iraq, Tigris near Mosul (36º20'N, 43º08'E); BM(NH) 1972.3.16:2, 69.4 mm standard length, Iraq, 10 km northwest Qala Dize (); BM(NH) 1974.2.22:1270, 174.6 mm standard length, (); BM(NH) 1974.2.22:1271-1272, 2, 91.9-210.2 mm standard length, (); BM(NH) 1974.2.22:1273-1274, 58.4-62.0 mm standard length, (); BM(NH) 1974.2.22:1275-1277, 3, 182.4-201.0 mm standard length, (); BM(NH) 1974.2.22:1278, 81.9 mm standard length, (); BM(NH) 1974.2.22:1289, 173.3 mm standard length, ();

Luciobarbus brachycephalus
(Kessler, 1872)

Common names

zardek, زرده پر (= zardehpar), سس ماهي (= sos or sas mahi), سس ماهي خزري (sas mahi khazari), sassmahi-ye Daryaye-Khazar.

[xazar sirbiti or shirbit in Azerbaijan; Kaspiiskii usach or Caspian barbel and korotkogolovyi ustach or short-headed barbel in Russian; Aral barbel; short-headed barbel].

Systematics

Barbus brachycephalus was originally described from the Syr Darya in Uzbekistan.

Howes (1987) considers the generic placement of this species to be problematical. It has slender barbels, 7 branched dorsal fin rays and the cranium is broad and flat, all characters at odds with Barbus sensu stricto. Doadrio (1990) places it in the subgenus Luciobarbus Heckel, 1843 based on a series of 4 synapomorphic osteological characters, namely the exoccipital contacts the pterotic "largely" (sic, probably broadly), high medial process of the urohyal, narrow exoccipital apophysis of the pterotic, and wide 4th and 5th infraorbitals.

Barbus obtusirostris (non Valenciennes, 1842) Jakovlev, 1870 (nomen praeoccupatum), described from the Volga River delta, Russia, is a synonym.

A possible syntype of B. brachycephalus from the Aral Sea is in the Naturhistorisches Museum Wien (NMW 53971) (Almaça, 1986). The NMW card index lists this fish plus 2 fish in NMW 53972 and 1 fish in NMW 53973 as syntypes. Syntypes in St. Petersburg, Russia are lost (Bogutskaya in Bănărescu and Bogutskaya, 2003). Syntypes of Barbus brachycephalus caspius are in the Zoological Institute, Russian Academy of Sciences, St. Petersburg under 2892 (8 fish), Transcaucasia, 3895 (8), Lenkoran, 9076 (22), 9085 (10), 9109(2), 9117(11), 9118(1), 9124(8), 9128(9), all from the lower Aras River and Lenkoran, 17042(2), 17043(1), 17044(1), all from the Bank Fishery along the lower Kura River. Syntypes under 10619 are apparently lost and a fish under 9108 is actually a Luciobarbus capito (Bănărescu and Bogutskaya, 2003).

The Caspian Sea basin subspecies is Luciobarbus brachycephalus caspius (Berg, 1914), described originally from the Caspian Sea basin (Eschmeyer et al., 1996). Karaman (1971), however, considers differences with the type subspecies of the Aral Sea basin to be minor and not worthy of subspecific recognition. Differences are in body proportions and the Caspian barbel has a smaller eye, lower dorsal fin, less deep body and head, longer pectoral-pelvic distance, shorter pelvic-anal distance, and dorsal fin further back than in the Aral barbel (Berg, 1948-1949). Fricke et al. (2007) list this taxon as a full species but also have brachycephalus in the same system in Turkey (Kura-Aras).

Key characters

The 7 branched dorsal fin rays and the predorsal distance shorter than the postdorsal distance distinguishes this species from B. capito, and colour pattern distinguishes it from B. lacerta, the other Caspian Sea barbels. ?and mursa

Morphology

The mouth is moderate in size and subterminal. Lips are thin to moderate, without a median lobe on the lower lip, and barbels are of moderate thickness. The anterior barbels can reach the level of the posterior eye margin and the posterior barbels reach or pass the preopercle level but barbel lengths show marked individual variation.

Dorsal fin with 3-5, usually 4, unbranched and 6-8, usually 7, branched rays, anal fin with 2-3, usually 3, unbranched and 5-6, usually 5, branched rays. Pectoral fin branched rays 14-17 and pelvic fin branched rays 7-8, usually 8. The dorsal fin denticles on the last unbranched ray are usually moderate in number, but may be lost in very large adults, are usually well-developed and extend along four-fifths of the ray (Karaman, 1971). This ray is very strong. Lateral line scales 62-90, commonly 65-77. Scales are elongate with a central focus and few anterior and posterior radii in young fish. There is a pelvic axillary scale. Gill rakers 16-25, short and reaching the one below when appressed. Pharyngeal teeth 2,3,5-5,3,2, hooked at the tip with the fourth tooth of the inner row large and blunt and the first three spatulate, rarely 2,3,4-5,3,2 or 2,3,4-4,3,2. Total vertebrae 45-50, usually 46-49, mode 48. The gut is coiled anteriorly. The chromosome number is 2n=100 (Klinkhardt et al., 1995).

Iranian specimens have the following meristics: branched dorsal fin rays 7(3), branched anal rays 5(3), branched pectoral fin rays 16(1) or 17(2), and branched pelvic fin rays 8(3). Lateral line scales 69(2) or 71(1). Total gill rakers 18(1) or 19(2). Pharyngeal teeth 2,3,5-5,3,2(2) or 2,3,4-5,3,2(1). Total vertebrae ?.

Sexual dimorphism

Abdurakhmanov (1962) reports on fish from the Kura River basin where males have a longer dorsal fin base and females have a greater maximum body depth, width and girth. Bogutskaya in Bănărescu and Bogutskaya (2003) report that males have a shorter head and longer unpaired fins; nuptial tubercles and colouration are absent.

Colour

The back is dark green, flanks and belly lighter, and the two areas may contrast as in Luciobarbus capito. No dark spots on the body. Fins greyish. Peritoneum brown.

Size

Reaches 22.5 kg (Robins et al., 1991) and 1.2 m.

Distribution

Found in the Caspian and Aral seas and their tributaries. In Iran, it was formerly known from the Anzali Mordab but is probably no longer present (Holčík and Oláh, 1992; but see below) and it was listed as rare in the Safid Rud (Derzhavin, 1934). Nedoshivin and Iljin (1929) and Nevraev (1929) recorded it from the Gorgan, Astrabad and Enzeli (= Anzali) regions. Recent works place it in rivers from the Astara to the Neka and Gorgan Bay peninsula, in the Anzali Mordab, and along the whole Caspian Sea coast but these are summaries of past and present distributions (Riazi, 1996; Abbasi et al., 1999; Kiabi et al., 1999; Abdoli, 2000). This species is now very rare in the Caspian Sea basin of Iran, with only a couple of specimens found in a recent survey (M. Ramin, pers. comm., 2000).

Zoogeography

Almaça (1984b) considers this species to be a Sarmatian Sea remnant, a Neogene brackish-water basin.

Habitat

Enters rivers to spawn but does not ascend as high as Luciobarbus capito. It prefers deep sections of rivers with stony and gravel bottoms. In the Caspian Sea it may be found at 13-25 m depth. On the Kura River in Azerbaijan there is a spring run and one in August-September. The spring run begins in March and lasts about 50 days; the summer run starts after a short interruption and lasts about 190 days. The water temperature at the start of the spring run is 6.7-11.0°C but the most intensive migration is in summer at 25.2-27.2°C (Bogutskaya in Bănărescu and Bogutskaya, 2003). Spring run fish spawn in the same year. This species has been recorded at depths of 11.0-11.9 m in the Iranian Caspian Sea (Knipovich, 1921). Riazi (1996) reports that this species migrates into the Siah-Keshim Protected Region of the Anzali Mordab. Jolodar and Abdoli (2004) state that it is more abundant in Gilan than in Mazandaran coastal waters.

Young females usually enter the sea immediately but males may remain in fresh water for 3-5 years. Spawners return to the sea.

Age and growth

Most fish examined by Razivi et al. (1972) from commercial catches in Iran were 2-7 years old, 38.0-69.0 cm long and weighed 698-4658 g. Low recruitment is attributed to poor spawning success, a result of water abstraction during its spawning season. Sexual maturity is attained at 6-8 years. Holčík and Oláh (1992) note that the Anzali region catches are dominated by 3-5 year old fish, 38-71 cm fork length, with rapid growth and a weight of 2 kg attained during the fifth year of life. Abdurakhmanov (1962) gives a maximum life span of 13 years in Azerbaijan. Females live longer than males which only reach 10 years (Bogutskaya in Bănărescu and Bogutskaya, 2003).

Food

No detailed literature reports but gut contents of small specimens from Iran contain crustaceans, and insects such as, curiously, ants, thrips and mosquitos. This fish evidently feeds on insects taken at the surface and is reported as leaping out of the water to take flying insects (Bogutskaya in Bănărescu and Bogutskaya, 2003). Mayflies and caddisflies are also taken and gut contents includes detritus. Crustaceans are the main food taken in the Caspian Sea (Abdurakhmanov, 1962) but molluscs are also recorded as well as small fish.

Reproduction

This barbel spawns in swift streams over pebbles or sand during July and August in Iran and the eggs attach to rocks (Razivi et al., 1972). Holčík and Oláh (1992) and Makeeva and Pavlov (2000) state that eggs are semipelagic, hatching as they drift downstream over 2 days at 25°C. Fry are carried downstream. Up to 1,259,000 bright-yellow eggs are produced of 1.4 mm diameter and the spawning season on the Kura River begins at the end of April, peaks in June and ends at the end of August. Favoured temperatures are 20-23°C (Abdurakhmanov, 1962). First spawning is at 5-7 years of age with females taking a year longer to mature than males (Bogutskaya in Bănărescu and Bogutskaya, 2003).

Parasites and predators

Molnár and Jalali (1992) record the monogenean Dactylogyrus affinis from this species in the Safid River.

Economic importance

This species is caught as a food fish in Iran. Nevraev (1929) records catches of 37 to 962 individuals from the Anzali region for the years 1914-1915 to 1917-1918. It was abundant in the Anzali Mordab with total catches for Iran of 54.6 t and 32.9 t in 1969/70 and 1970/71 (28.7 t and 14.4 t for the Anzali region alone) but few fish are captured now (Holčík and Oláh, 1992) (note that these figures were taken from Appendix 11, on page 10 they are reversed). They are caught in rogas (outflowing rivers from the Anzali Mordab) and inflowing rivers of the mordab (lagoon) in late winter and early spring. On the Kura River of Azerbaijan average weight in catches was 5.6 kg for females and 3.5 kg for males and the catch from 1920-1944 varied from 0.2 to 3.6 thousand centners.

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

Conservation

Vulnerable in Turkey (Fricke et al., 2007). Stocks of this species have declined because of poor habitat for spawning and the construction of dams and weirs which restricted access to spawning grounds. Water abstraction for irrigation during the summer spawning season would have to be balanced against the requirements of the fish. Larvae of spring spawners are lost when they enter irrigation channels and become stranded in fields (Razivi et al., 1972).

Once known from the Anzali Mordab, it is now absent to rare there and apparently replaced by Luciobarbus capito (Holčík and Oláh, 1992). Kiabi et al. (1999) consider this species to be critically endangered in the south Caspian Sea basin according to IUCN criteria. Criteria include commercial fishing, sport fishing, few in number, habitat destruction, limited range (less than 25% of water bodies), absent in other water bodies in Iran, and absent outside the Caspian sea basin.

This species is regarded as critically endangered through illegal overfishing, pollutants and the destruction of breeding and nursery grounds. Only 2 specimens were caught in the 3 years prior to 2000 during a study of "Barbus" species in Iran. Additionally, during the 6 month beach seine fishing season (October to April) for the years 1998 and 1999 along the Caspian shore, no specimens were caught in 138 beach seines used 51,000 times (M. Ramin, pers comm., 2000).

Further work

Detailed surveys, perhaps returning captures alive, need to be carried out to monitor the status of this species in Iran.

Sources

Type material: ?

Iranian material: CMNFI 1970-0553, 2, ? mm standard length, Gilan, Sowsar Roga (37º27'N, 49º30'E); CMNFI 1980-0120, 1, 115.3 mm standard length, Mazandaran, Babol River at Babol Sar (36º43'N, 52º39'E);

Luciobarbus capito
(Güldenstaedt, 1773)

L. capito (above) and L. mursa (below) Aras River, 2 October 1994
Courtesy of Asghar Abdoli

Common names

usach bulatmai, usach chanari; zardi, zardek, zardak, زرده پر (= zardehpar), zard pareh, اورنج (oranj, orenj, orenge or ourange, possibly from the yellowish fin colour)سس ماهي (= sos, sas or sass mahi), pulad mahi (= steel fish from body colour), ses mahi bozorg (= big ? fish, ses being a word of unknown meaning).

[zardapar, shirbit, yastibas zardapar for natio platycephalus, all in Azerbaijan; tchanari in Georgian; bulatmai in Turkish; usach (or usatch) bulatmai and usach chanari in Russian]. Bulatmai is derived from Farsi, bulat = pulad or steel, mai = mahi or fish in reference to the colour on the upper flank].

Systematics

Cyprinus capito was originally described from the Kura River, Transcaucasia. No types are extant.

Howes (1987) places this species in Barbus sensu stricto. Doadrio (1990) places it in the subgenus Luciobarbus Heckel, 1843 based on a series of osteological characters (listed under Luciobarbus brachycephalus).

Cyprinus bulatmai Hablizl, 1783 (after Berg (1948-1949; Rainboth (1981) has Gmelin, 1774 as the author while Eschmeyer et al. (1996) have Gmelin, 1784 (originally described from Anzali, Iran), Cyprinus chalybatus Pallas, 1814 (originally described from Anzali, Iran), Cyprinus mystaceus Pallas, 1814 (partim, from Tiflis), Barbus conocephalus Kessler, 1872 described from the Zeravshan River, Uzbekistan, Barbus lacertoides Kessler, 1872 described from the Syr-Darya in the neighbourhood of Khodzhent (= Leninabad), Tajikistan, Barbus capito var. tiflissica Kamenskii, 1899 described from the Kura River at Tiflis (= Tbilisi), Georgia, and Barbus bilkewitschi Bulgakov, 1923 (originally described from the "Atrek", i.e. the Atrak River in Turkmenistan on the northeastern border of Iran; also spelt bilkewitchi on page 236 in Bulgakov but bilkewitschi on the plate), are synonyms. Barbus capito serratus Sokolinskii, 1927 is a subspecies from the southern Caspian Sea and Barbus capito platycephalus Abdurakhmanov, 1960 is a subspecies or a natio in the lower Kura River basin (see Abdurakhmanov (1962) for further details). Berg (1948-1949) and Karaman (1971) consider Barbus capito serratus to be a synonym of B. c. capito.

Bianco and Banarescu (1982) record this species from the Hablehrud and the Kul River basin at Darab in Persian Gulf drainages. The 2 specimens have 52 lateral line scales, 8 branched dorsal fin rays and 18-19 gill rakers. They acknowledge that these 2 fish have fewer scales than L. capito from the Caspian Sea basin but believe they may represent a new subspecies. These fish are presumed to be misidentifications as L. capito is restricted to the Caspian Sea basin.

Laloei et al. (2003) using the mitochondrial cytochrome-b gene found no separable populations of this species in 60 samples from the Iranian Caspian Sea coast and rivers.

Key characters

The 8 branched dorsal fin rays and the predorsal distance considerably longer than the postdorsal distance distinguishes this species from L. brachycephalus, and colour pattern distinguishes it from Barbus lacerta, the other Caspian Sea barbels. ?mursa

Morphology

There is a rounded keel on the back in front of the dorsal fin. The mouth is moderate in size, inferior and horseshoe-shaped. Lips are fleshy and well-developed with tubercles but there is no free median lobe on the lower lip. Barbels can be the most developed in thickness in this species among the Luciobarbus considered here but this can vary. The anterior barbel extends back between the anterior eye margin level and its middle and the posterior barbel extends to the posterior eye margin level or almost to the preopercle in young and some adults.

Dorsal fin with 3-5, usually 4-5, unbranched rays followed by 7-9, usually 8, branched rays and anal fin with 2-4, usually 3, unbranched and 5-6, usually 5, branched rays. Pectoral fin branched rays 15-19 and pelvic fin branched rays 7-9. The dorsal fin denticles on the last unbranched ray may be lost in very large adults but are evident for two-thirds or more of the spine length in most fish (Karaman, 1971; Almaça, 1981). The last unbranched ray is moderately strong and the denticles are of moderate density along it. Lateral line scales 51-72, usually 60-66 (Karaman (1971) gives 36-70 but he includes 8 subspecies over a wide range within his definition of the species). There is no obvious pelvic axillary scale although scales in this region are elongate. The scale focus is slightly subcentral anterior, there are numerous fine circuli, and there are radii on all fields with those on the lateral fields few and often curved. Gill rakers 12-19, rarely to 22, increasing in number with the size of the fish, reaching the one below or slightly further when appressed, rounded and knobbed tip, and a large internal rounded extension. Pharyngeal teeth usually 2,3,5-5,3,2 with minor variants, hooked and spoon-like below with the depression below the crown filled in, the fourth one in the inner row the largest and pointed or blunt and rounded, the fifth smaller and blunt. The gut is long and complexly coiled with several anterior and posterior loops. Total vertebrae 42-45 (Howes, 1987), 45-47 (Elanidze, 1983), 43-49 (Bogutskaya, Bănărescu and Almaça in Bănărescu and Bogutskaya, 2003). Chromosome number 2n=100, NF=172 (Pourali Darestani et al., 2006).

Iranian fish have the following meristics: branched dorsal fin rays 7(1) or 8(49), anal fin branched rays 5(50), pectoral fin branched rays 16(3), 17(27), 18(18) or 19(2), pelvic fin rays 7(2) or 8(48); lateral line scales 53(3), 54(4), 55(7), 56(10), 57(6), 58(7), 59(7), 60(5) or 61(1); total gill rakers 13(3), 14(15), 15(18), 16(9), 17(4) or 18(1); pharyngeal teeth 2,3,5-5,3,2(33), 2,3,5-5,3,1(1), 2,3,4-5,3,2(2), 2,3,5-4,3,2(1), 2,3,5-5,2,1(1), 2,2,5-5,3,2(1); and total vertebrae ?

Sexual dimorphism

Unknown, apparently no spawning colouration or breeding tubercles.

Colour

The upper flank and head are steel-grey (hence bulat mahi) and the lower flank and belly are a strongly contrasting pale yellow or pearly-white. Occasionally fish with a uniform coloration are found and preserved material may be uniform. The steel-grey upper flank may be comprised of dark scale margins surrounding a silvery-grey scale centre. The lateral line may be darkly pigmented. Spots may occur individually on the body. The iris is silvery with a grey exterior ring and a very narrow interior golden ring. Barbels are white with grey on the inner surface. The dorsal fin is greyish and may have some dark grey spots. The caudal fin has a greyish or yellowish or slightly orange upper lobe, sometimes with faint dark grey spots, a more strongly coloured and larger yellow-orange to canary-yellow lower lobe and pink margins. The pectoral fin is whitish with a little or considerable amount of pink or yellow. The pelvic and anal fins are canary-yellow to orange with a white margin. Young fish may be darkly speckled and mottled on the mid and upper flank rather like Barbus lacerta. Peritoneum dark brown.

Size

Reaches 1.05 m and 15 kg in literature reports. A specimen from the Sardabrud was 85 cm and 5.5 kg (A. Abdoli, pers. comm., 1995).

Distribution

Found in the basins of the Black, Caspian and Aral seas. Karaman (1971) gives a distribution from the Iberian Peninsula and North Africa to Southwest Asia but he includes 8 subspecies within his definition of "Barbus" capito.

In Iran, this species is found in the Caspian Sea basin, in rivers from the Aras to the Atrak and the Anzali Mordab, the Qezel Owzan and Shahrud in the upper Safid River basin, and the along the sea coast (Derzhavin, 1934; Bianco and Banarescu, 1982; Almaca 1984a; Aliev et al., 1988; Holčík and Oláh, 1992; Kiabi et al., 1994; Abbasi et al., 1999; Kiabi et al., 1999; Abdoli, 2000).

Luciobarbus capito conocephalus (Kessler, 1872) is reported from the Karakum Canal, Kopetdag Reservoir and Uzboi lakes in Turkmenistan (Shakirova and Sukhanova, 1994; Sal'nikov, 1995) and may eventually be recorded from the Tedzhen River and Caspian Sea basins in Iran.

Zoogeography

Almaça (1984b) considers this species to be a Sarmatian Sea remnant, a Neogene brackish-water basin, and related to Euro-Mediterranean "Barbus".

Habitat

This species avoids muddy bottoms (Solak, 1977) although Bogutskaya, Bănărescu and Almaça in Bănărescu and Bogutskaya (2003) report that it prefers warm, deep, slowly-flowing water above gravel, sand or mud and can be found in lacustrine habitats. Spawning migrations in the Kura River of Azerbaijan go as far up as Aragva and generally it ascends to the uppermost tributaries of rivers it enters. The spawning run in the Kura lasts almost the whole year except for the two coldest months. However the main spawning runs are in September-October and April (Bogutskaya, Bănărescu and Almaça in Bănărescu and Bogutskaya, 2003). The Caspian Sea form is anadromous but there are also resident forms in the rivers there. Knipovich (1921) reports this species at depths of 9.15-14.2 m, possibly deeper, in the Iranian Caspian Sea. There are both resident and andromous populations in the Anzali lagoon (Karimpour, 1998).

Age and growth

Solak (1989c) examined a population of this species in the Aras River in Turkey and found a life span of over 4 years, but over 6 years in the Çoruh River of the Black Sea basin of Turkey. In the Caspian Sea basin fish may live up to 8 years (Abdurakhmanov, 1962). Anadromous fish are heavier than fish of the same length that are river residents. Maturity is attained at 3-5 years with females mature one year later than males. Spring migrants spawn that summer while summer or autumn migrants overwinter to spawn the following spring or summer (Bogutskaya, Bănărescu and Almaça in Bănărescu and Bogutskaya, 2003). Shajiee et al. (2002) found a sex ratio of 3:1 for male:female fish in the Caspian Sea off Gilan and a life span of 8 years. Gonadosomatic and hepatosomatic indices, length-weight relationships and other growth and fecundity indices were given.

Food

Stomach contents consist of insects, crustaceans and worms, and filamentous algae and other plant material with associated invertebrates. Terrestrial insects, small fishes and frogs are also taken. Abdoli (2000) reports Ephemeroptera, Trichoptera and Chironomidae. One specimen from Iran had fish remains, possibly a small Luciobarbus capito. Abdurakhmanov (1962) reports grasshoppers and ants, presumably taken at the surface.

Reproduction

Eggs number up to 193,600 and diameters up to 1.8 mm in Azerbaijan (Abdurakhmanov, 1962). A fish with well-developed testes was caught in the Gorgan River on 7 July, suggesting a spawning season of late spring and summer, agreeing with egg diameters of fish from Azerbaijan which are largest in June.

Parasites and predators

Molnár and Jalali (1992) record the monogenean Dactylogyrus linstowi from this species in the Safid Rud. Masoumian and Pazooki (1998) surveyed myxosporeans in this species in Gilan and Mazandaran provinces, finding Myxobolus musculi. Masoumian et al. (2005) recorded the protozoan parasites Ichthyophthirius multifilis and Trichodina perforata from this species in water bodies in West Azarbayjan. Masoumian et al. (2003) record Myxobolus musculi while Pazooki et al. (2003) and Pazooki (2006) record Rhabdochona hellichi, Paradiplozoon homoion and Pseudocapillaria tomentosa, all reports from fishes captured in the Tajan and Zarem rivers of Mazandaran. Sattari et al. (2002) and Sattari (2004) records the presence of the nematode, Eustrongylides excisus, in the body cavity. This parasite can damage muscles in commercial species and render them unsuitable for sale. Sattari et al. (2004, 2005) surveyed this species in the inshore area of the Caspian Sea, recording Eustrongyloides excisus and Anisakis sp. Pazooki et al. (2007) recorded various parasites from localities in West Azarbayjan Province, including Neoechinorhynchus rutili from this species. Miar et al. (2008) examined fish in Valasht Lake and the Chalus River, Mazandaran and found the metazoan Bothriocephalus gowkongensis.

Economic importance

Holčík and Oláh (1992) report a catch of only 9 kg in the Anzali Mordab for 1990. This species had a catch of 17 tonnes in 1997, 28 t in 1998 and 7 t in 1999 during the 6 month beach seine fishing season (October to April). For the years 1998 and 1999, 138 beach seines were used 51,000 times (M. Ramin, pers. comm., 2000). This species was of minor importance commercially in the former U.S.S.R. and is a sport fish in Georgia (Bogutskaya, Bănărescu and Almaça in Bănărescu and Bogutskaya, 2003). In East Azarbayjan it reaches sizes large enough for sport fishing and as a commercial species (Ghasemi, 2002).

Conservation

Kiabi et al. (1999) consider this species to be conservation dependent in the south Caspian Sea basin according to IUCN criteria. Criteria include commercial fishing, sport fishing, medium in numbers, habitat destruction, widespread range (75% of water bodies), absent in other water bodies in Iran, and present outside the Caspian Sea basin. Mostafavi (2007) lists it as conservation dependent in the Talar River, Mazandaran. Vulnerable in Turkey (Fricke et al., 2007).

Further work

Biology and numbers of this species needs investigation.

Sources

Type material: ?

Iranian material: CMNFI 1970-0521, 7, ?-102.5 mm standard length, Gilan, Safid River near Lulaman (no other locality data); CMNFI 1970-0525, 5, 111.9-133.4 mm standard length, Gilan, Safid River near Mohsenabad (no other locality data); CMNFI 1970-0526, 19, ? mm standard length, Gilan, Safid River 6 km below Astaneh Bridge (37º19'N, 49º57'30"E); CMNFI 1970-0531, 1, 157.3 mm standard length, Mazandaran, Larim River (36º46'N, 52º58'E); CMNFI 1970-0536, 1, 194.4 mm standard length, Gilan, Siah River estuary near Rudbar (36º53'N, 49º32'E); CMNFI 1970-0538, 10, 36.7-188.5 mm standard length, Gilan, Qezel Owzan River near Manjil (36º44'N, 49º24'E); CMNFI 1970-0543A, 1, 170.4 mm standard length, Gilan, Caspian Sea at Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1970-0546, 10, 39.3-61.8 mm standard length, Gilan, Safid River canal (no other locality data); CMNFI 1970-0553, 1, 58.1 mm standard length, Gilan, Sowsar Roga (37º27'N, 49º30'E); CMNFI 1970-0563, 1, 70.1 mm standard length, Gilan, Caspian Sea at Kazian Beach (ca. 37º29'N, ca. 49º29'E); CMNFI 1970-0568, 8, 62.5-132.0 mm standard length, Gilan, Caspian Sea at Kazian Beach (ca. 37º29'N, ca. 49º29'E); CMNFI 1970-0581, 6, 41.3-65.5 mm standard length, Gilan, Caspian Sea near Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1970-0587, 3, 69.0-91.4 mm standard length, Mazandaran, Babol River at Babol Sar (36º43'N, 52º39'E); CMNFI 1979-0431, 2, 240.9-265.5 mm standard length, Mazandaran, bazaar at Now Shahr (no other locality data); CMNFI 1979-0437, 1, ? mm standard length, Gilan, Safid River 2 km west of Astaneh (37º16'30"N, 49º56'E): CMNFI 1979-0452, 2, 53.5-56.5 mm standard length, Azarbaijan-e Khavari, Qezel Owzan River 6 km from Mianeh (37º23'N, 47º45'E); CMNFI 1979-0486, 2, 69.2-78.8 mm standard length, Mazandaran, Atrak River draiange (37º44'N, 56º18'E); CMNFI 1979-0488, 1, 95.8 mm standard length, Mazandaran, Atrak River at Maraveh Tappeh (37º55'N, 55º57'30'E); CMNFI 1979-0491, 1, 191.5 mm standard length, Mazandaran, Gorgan River 15 km northeast of Kalaleh (ca. 37º33'N, ca. 55º44'E); CMNFI 1979-0686, 19, ? mm standard length, Gilan, Safid River (37º24'N, 49º58'E): CMNFI 1979-0695, 4, ? mm standard length, Gilan, Safid River at Manjil Bridge (36º46'N, 49º24'E): CMNFI 1979-0788, 2, 152.0-202.4 mm standard length, Mazandaran, Gorgan River at Khadje Nafas (37º00'N, 54º07'E); CMNFI 1980-0116, 8, ?-62.4 mm standard length, Gilan, Safid River at Astaneh Bridge (37º16'30"N, 49º56'E); CMNFI 1980-0123, 8, ? mm standard length, Gilan, Safid River (ca. 37º22'N, ca. 49º57'E): CMNFI 1980-0127, 1, ? mm standard length, Gilan, Caspian Sea near Hasan Kiadeh (37º24'N, 49º58'E): CMNFI 1980-0132, 8, ? mm standard length, Gilan, Safid River at Kisom (37º12'N, 49º54'E); CMNFI 1980-0138, 2, 132.5-137.6 mm standard length, Gilan, Safid River estuary (ca. 37º28'N, ca. 49º54'E); CMNFI 1980-0905, 1, 188.9 mm standard length, Mazandaran, Gorgan River at Khadje Nafas (37º00'N, 54º07'E); CMNFI 1980-0908, 3, 67.2-91.3 mm standard length, Gilan, Safid River estuary (ca. 37º28'N, ca. 49º54'E); uncatalogued, 2, 245.7-272.8 mm standard length, Markazi, Shah River (no other locality data);

Luciobarbus esocinus
Heckel, 1843

Darreshahr, Simarreh River, April 1987,
photo by N. Atarody; courtesy of B. Kiabi

Kermanshah, Simarreh River,
courtesy of B. Kiabi

more photos

Common names

سونگ (= soong) or بچ ( = bach) in northern Khuzestan and Lorestan, anzeh, anzah, narbach, and anzeh-bach at Ahvaz and in southern Khuzestan (meanings unknown); بل زرد (= balzard).

[bizz (in Iraq), farkh-el-biz (= cheerful one (Heckel, 1843b) or baby of bizz), farch or mangar in Arabic; "Tigris salmon", "Euphrates salmon", pike barb, pike barbel].

Systematics

Howes (1987) places this species in Barbus sensu stricto. Labeobarbus Euphrati Sauvage, 1882 described from "Biredjik (Euphrates)", Turkey (not "Irak" as in Bertin and Estève (1948)) is a synonym.

Karaman (1971) places this species in the synonymy of "Barbus" xanthopterus as he considered the only difference to be scale count and the range of variation for these species is unknown. Almaça (1983, 1986) agrees that several meristic characters are similar while the main differences are a shorter head and barbels in esocinus and dotted coloration in esocinus as opposed to uniform in xanthopterus (isn't the reverse true?). He maintains them as separate species because information on variability in characters is lacking.

Examination of the types of L. esocinus (NMW 54088, 2, 58.5-61.5 mm standard length, 54091, 372.4 mm, 54092, 321.3 mm) and L. xanthopterus (NMW 54841a (a syntype), 216.5 mm, 54786 (not a type), 292.8 mm) in Vienna showed the following differences. Head size differs in the two taxa in that esocinus postorbital length is very elongate and the head tapers anteriorly in a distinctive fashion. Head length in standard length is 3.2-3.6, mean 3.4 for esocinus and 4.0-4.2, mean 4.1 for xanthopterus and postorbital length in standard length is 5.9-7.2, mean 6.5 for esocinus and 7.7-7.8, mean 7.8 for xanthopterus with the higher values for esocinus based on smaller fish which tend to have proportionately larger heads. Total gill raker counts are 8-10, mean 9.3 for esocinus and 12-13, mean 12.5 for xanthopterus. Larger esocinus appear to lose anterior rakers with age but still have fewer than xanthopterus of similar size. Lateral line scale counts are 63-70, mean 67.3 in esocinus and 57-60, mean 58.5 in xanthopterus. On this limited basis I am maintaining the two species as distinct. An Iranian specimen, 284.3 mm standard length (ZSM 21830 from the Dez River), falls within the ranges given above. see xanthopterus?

Almaça (1986) records syntypes of Luciobarbus esocinus in the Naturhistorisches Museum Wien from the type locality as given by (Heckel, 1843b) "bei Mossul in Tigris", Iraq (NMW 54088 (2 specimens), NMW 54091 (1), and NMW 54092 (1) but Heckel (1843b) does not specify the number of types). A syntype is in the Senckenberg Museum Frankfurt (SMF 454, formerly NMW; 281.2 mm standard length) and another syntype is also there but dried (SMF 6785, formerly NMW) (F. Krupp, pers. comm., 1985). The catalogue in Vienna lists 2 fish in spirits and 2 fish stuffed.

The mounted holotype of Labeobarbus euphrati is in the Muséum national d'Histoire naturelle, Paris (MNHN A.6961) and measures 1650 mm total length (Bertin and Estève, 1948). Eschmeyer et al. (1996) indicate that the catalogue number may be A.6971.

Key characters

This species is characterised by large size, a long, tapering and depressed head (rather pike-like in shape), two pairs of barbels, a serrated dorsal fin spine, lateral line scale count high (63-78), moderately developed lips, and no large flank spots. Head length in standard length 3.1-3.7, mean ? and postorbital length in standard length 5.9-7.2, mean ? for ? specimens ?-? mm standard length including the types listed above. (includes SMF454 281.2 mm SL, HL 75.6, postorb 44.1, snout 21.5

Morphology

Dorsal fin with 4 unbranched and 8 branched rays, anal fin with 3 unbranched and 5 branched rays, pectoral fin with 16-18 branched rays and pelvic fin with 8 branched rays. Lateral line scales 62-78. Scales are regularly arranged, the smallest being on the isthmus anterior to the pectoral fin bases. There is a pelvic axillary scale. Scales have a central focus, numerous fine circuli, a wavy or rounded anterior margin, and radii on the anterior and posterior fields with a few widely spaced ones on the lateral fields. Gill rakers 8-12, well spaced and just touching the one below when appressed. Pharyngeal teeth 2,3,5-5,3,2, hooked with the third tooth of the inner row slightly larger than the fourth and the fifth smaller. Heckel (1843b) gives 2,3,4-4,3,2, and teeth from large specimens seen at Ahvaz in 1995 by me had 2,3,4-4,3,2 and 2,3,5-4,3,2, the anteriormost tooth being small or absent. Even small specimens (85.7 mm standard length) may have the anteriormost tooth absent. Total vertebrae 48 (Howes, 1987) or 48-50 based on comparative materials listed below. The last unbranched dorsal fin ray is very strong, with a low density of denticles but with fine denticles extending over much of the ray. The mouth is large, terminal and almost horizontal and extends back to the anterior eye margin. Lips are thin to moderate without a median lobe to the interrupted lower lip, and barbels are thin to very thin. The anterior barbel does not reach past the nostril level and the posterior barbel does not pass the mid-eye to rear eye level. The nostril is elongate and closer to the eye than the snout tip. The cephalic canals on the suborbital series have numerous branches. The gut is an elongate s-shape with several anterior loops.

Meristics for an Iranian specimen:- dorsal fin branched rays 8; anal fin branched rays 5; pectoral fin branched rays 17; pelvic fin branched rays 8; lateral line scales 69; and total gill rakers 9.

Sexual dimorphism

Unknown.

Colour

The back has numerous scattered, black spots on an olivaceous background, the spots extending onto the base of the dorsal fin. Spots may be weak or absent but this is comparatively rare. Overall colour is silvery with the anal and caudal fins dark red. The flanks and belly are lighter. The eye is yellowish in colour. Young fish have a yellow tinge or sulphur yellow colour to the fins.

Size

Frequently up to 3 hundredweights (= 152.4 kg) in the Zab River of Iraq southeast of Mosul (Heckel, 1846-1849a); a fish 6'4" (1.93 m) long with a girth of 3'10" (1.17 m) and a weight of 215 lbs (97.6 kg) from the Euphrates River at Hakika (Light, 1917; wrongly identified as "Barbus" scheich according to the editors in an article by Gudger (1945a)); 69 inches (1.75 m) measured over the curve of a back with a 38 inch (0.97 m) girth and a weight of 123 lbs (55.8 kg) caught in the Diyala River, Iraq on a light 14-foot rod taking 1½ hours to land (Bagnall, 1919); 96 lb (43.6 kg) fish caught near Kizil Robat (= As Sa`diyah) in the Diyala River on a lump of atta (a ball of dough)(MacKay, 1919)(Bagnall, a Major, out-doing MacKay, a Brigadier-General); 140 lbs (63.6 kg) Tigris salmon caught on a 2" spoon at Samarra (Lane, 1920); hundreds of good weight up to 112 lbs (50.8 kg), one caught on a hand-line at 170 lbs (77.2 kg), one netted at 252 lbs (114.4 kg), and reputedly over 300 lbs (136 kg)(Radcliffe, 1926); up to two yards (1.83 m) as evidenced by a photograph of a specimen draped over a donkey in Iraqi Kurdistan (Hamilton, 1937); 2 m and 150 kg in Iraq (van den Eelaart, 1954; Herzog, 1967); a 167 lb (75.8 kg) Tigris specimen and a 213 lb (96.7 kg) specimen at Nassiriyah on the Euphrates, called both gattan and "Euphrates salmon" but it was presumably the latter (Vesey-Fitzgerald and Lamonte (no date)); weights up to 300 lbs (136 kg) and the largest taken on rod-and-line as 220 lb (100 kg) and 7 feet (2.1 m), baits used included atta and dates, and chicken or sheep liver (Mahdi, 1962). Beck (pers. comm., 2000) reports the largest fish seen in the 1990s along the Syrian Euphrates and its tributaries weighed 198 kg. A fish caught in 2001 on the Euphrates River near Birecik in Turkey with a net weighed 111 kg and was 2.4 m long (www.fishing-worldrecords.com, downloaded 16 February 2007).

Iranian records of large specimens include one by Mr. Chabok-Savar, a Game Warden or biologist of the Department of the Environment who caught a specimen about 80 kg in the Simareh River in 1973 and N. Atarody, also a Game Warden or biologist, caught two large specimens in April 1987 from "Tang-e Gheer" on the Simareh near Darreh Shahr (Abzeeyan, Tehran, 3 (August-September):19, 1992). A 1.65 m and 75 kg specimen is reported from the Dez River and a 2.1 m specimen is reported from the market at Ahvaz in 1993 (this last fish may have weighed 150 kg, original report not seen; J. Valiallahi, pers. comm., 2001). The Gav Masiab, a river in Kordestan, is reputedly named for these large fishes ("river with fishes as large as a cow")(J. Valiallahi, www.modares.ac.ir, downloaded 4 July 2000). Floor (2003) gives a photograph of a large specimen from the Karun River.

Distribution

This species is found in the Tigris-Euphrates basin including its Iranian portion and the adjacent northern Gulf basin (Marammzai, 1995; Abdoli, 2000). It is reported as common in the Dez Dam (Gh. Eskandary, pers comm., 2000).

Zoogeography

Almaça (1984b, 1991) considers that the origin of this species lies with a group that migrated southwards in the late Pliocene from the Dacian Lake of the Sarmatian Sea and speciated in Mesopotamia.

Habitat

Found in large rivers and dams but also the more limited environment of palace ponds at Baghdad. Details of environmental requirements unknown.

Age and growth

Life span is at least 10 years (Ahmed, 1982). Fish in Khuzestan were found to have a sex ratio of 4.2:1.0 male:female (Gh. Eskandary, pers. comm., 2000; Eskandari et al., 2004). In the Dez Dam of northern Khuzestan females had a length range of 156-1350 mm and a weight range of 31.7-26,500 g while for males figures were 183-1065 mm and 48-12,208 g. Males matured faster than females, annual growth is slow and asymptotic length is more than 2 m. It appears to have a longer reproductive life compared to pre-maturation life (Eskandari et al., 2004).

Food

This species is a predator on other fishes. In the Dez Dam, all samples had fish in their stomachs although the gut to body length ratio indicates omnivory (Eskandari et al., 2004).

Reproduction

van den Eelaart (1954) reports spawning in Iraq in March. Eggs are laid between large stones in the deep part of rivers in March or April. Some fingerlings drift down into lakes and marshes. Eskandari et al. (2004) report a very short spawning season in the Dez Dam in spring after reservoir water levels rise through spring flooding. The fish is a total spawner with eggs released in upstream areas and shallows of the reservoir over gravel at 24ºC. Ünlü (2006) gives age at first maturity as 4 years in the Turkish Tigris River with spawning between large stones.

Parasites and predators

None reported from Iran.

Economic importance

In Iraqi Kurdistan these fish were caught and tethered by a cord passed through the lips until eaten by the villagers (Elliot, 1977). At Altan Keupri on the Lesser Zab River in Iraq a drugged bait was used to stupefy the fish so it could be netted and dragged to shore (Hamilton, 1937).

This species was being considered for aquaculture during the year 2000 in Khuzestan although fish larger than 1 m are needed to be ripe adults. Anglers and commercial fishermen seek this fish in the Iranian Zagros Mountains using ducklings (!) as bait (J. Valiallahi, pers. comm., 2001).

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

Conservation

This species is under severe threat in the Syria Euphrates and its tributaries. A survey in 1997-1998 caught only a single juvenile and the commercial fisheries had not more than two dozen fish. Blast fishing and poisoning had led to a decline in age of catches since 1993. Large scale water abstraction, dam building and pollution had destroyed habitats (R. Beck, pers. comm., 2000). It is listed by Stone (2007) as one of the world's largest freshwater fishes, presumed to be threatened.

A report of fish kill, presumably of this species, in the "Cham Ghorah" River near Mahabad in July 1999 numbering about half a million fish was owing to desiccation of the habitat (J. Valiallahi, www.modares.ac.ir, downloaded 4 July 2000).

Further work

Distribution and numbers are needed for a conservation assessment.

Sources

Type material: See above, NMW 54088.

Iranian material: ZSM 21830, 1, 284.3 mm standard length, Khuzestan, Dez River at Harmaleh (31º57'N, 48º34'E). Some specimens observed in the IFRTO laboratory at Ahvaz (pharyngeal arches).

Comparative material: BM(NH) 1892.9.1:30, 1, 197.3 mm standard length, Iraq, Al Faw (29º58'N, 48º29'E); BM(NH) 1920.3.3:80-82, 3, 85.7-147.0 mm standard length, Iraq, Basrah (30º30'N, 47º47'E); BM(NH) 1931.8.12:5, 1, 111.8 mm standard length, Iraq, near Mosul (no other locality data); BM(NH) 1974.2.22:1297, 1, 166.5 mm standard length, Iraq, Diyala River (no other locality data); BM(NH) 1974.2.22:1810, 1, 220.1 mm standard length, Iraq (no other locality data).

Luciobarbus kersin
(Heckel, 1843)

Common names

برزم (= berzem).

[shissan, jassan, gassan, djissan, barsam or bunni, kersin at Aleppo, all in Arabic; kersin barbel].

Systematics

Karaman (1971) places this species as a synonym of his Barbus capito pectoralis. Almaça (1983) suggests that kersin may be only subspecifically distinct from Barbus (= Luciobarbus) pectoralis (q.v.) but later (Almaça, 1984b) retains it as a full species until further information becomes available. Krupp (1985c) also synonymises this species with Barbus (= Luciobarbus) pectoralis.

Syntypes of Barbus Kersin from "Aleppo", the type locality given by Heckel (1843b) or "Gewässern von Aleppo" (Heckel, 1846-1849a), are in the Naturhistorisches Museum Wien (NMW 54212 and 54215) (Almaça, 1986). Krupp (1985c) lists the following syntypes of B. kersin all from Aleppo and collected by Th. Kotschy: 1 specimen, 141.2 mm standard length as measured by me (NMW 54212), 4, 89.1-135.1 mm standard length as measured by me (NMW 54213), 1, 166.0 mm standard length as measured by me (NMW 54215) and 1, 152 mm standard length (formerly NMW, now in the Senckenberg Museum Frankfurt as SMF 610). The card catalogue in 1997 listed NMW 54215 as "? lectotype" and NMW 54213 as "? paralectoptypes" (sic). Eschmeyer et al. (1996) list 1 syntype in the Museum für Naturkunde, Universität Humboldt, Berlin (ZMB 3237, formerly NMW). The catalogue in Vienna lists 6 specimens.

Key characters

This species differs from the similar L. pectoralis by the smaller scales and body depth being greater than head length (equal in pectoralis) (Berg, 1949). ?see BWC95-32 for a kersin after Valiallahi, its a dark Barbus without the very large D spine and a different colour pattern

Morphology

Dorsal fin unbranched rays 3-4, usually 4, branched rays 7-8, anal fin with 3 branched and 5-6, usually 5, unbranched rays. Pectoral fin branched rays 17, pelvic fin rays 8. Lateral line scales 49-58. Gill rakers 19. Pharyngeal teeth 2,3,5-5,3,2. The last unbranched dorsal fin ray is strong (as in pectoralis)with a low density of coarse denticles extending over much of the ray. The mouth is moderate in size and subterminal. The highly rounded snout projects a little. Lips are thin to moderate but not fleshy and lack a median lobe. The upper lip is covered partly by the snout. Barbels are thin, the anterior barbel not extending back beyond the anterior eye margin and the posterior barbel not beyond the middle of the eye. Body depth is equal to or greater than head length in the types examined by me.

Sexual dimorphism

Unknown.

Colour

The body lacks distinctive markings and is olive to reddish-brown above, silvery on the flanks and white below. The dorsal and caudal fins have a blackish margin.

Size

Attains 70.1 cm total length (Menon, 1956). Reaches 2 m and over 100 kg (Khalaf, 1961).

Distribution

Found in the Tigris-Euphrates, Quwayq and Orontes River basins. In Iran, it is found in the Tigris River basin in the Karun and Karkheh rivers, and the northern Gulf basin in the Zohreh and Helleh rivers and questionably the southern Gulf basin (Abdoli, 2000).

Zoogeography

Almaça (1984b, 1991) considers that the origin of this species lies with a group that migrated southwards in the late Pliocene from the Dacian Lake of the Sarmatian Sea and speciated in Mesopotamia.

Habitat

The main habitat of Iraqi fish is rivers, entering marshes and lakes during floods but returning to rivers in June (van den Eelaart, 1954).

Age and growth

Unknown.

Food

This species is said to eat a wide range of food items (Beckman, 1962).

Reproduction

Eggs are deposited on clay or gravel bottoms during mid-February to early March (van den Eelaart, 1954).

Parasites and predators

Gussev et al. (1993a) describe new species of monogeneans from this species in the Dez River, Khuzestan, namely Dactylogyrus deziensis, D. deziensioides and D. kersini. Ebrahimzadeh and Kailani (1976) record parasite species in the cestode genera Caryophyllaeus and Isoglaridacris and the protozoan Myxosoma from Barbus (= Luciobarbus) kersin taken in the Karun River.

Economic importance

None.

Conservation

Endangered in Turkey (Fricke et al., 2007) but status in Iran unknown.

Further work

The biology of this species in Iran needs study and a molecular comparison with putative synonyms would be of value in clarifying distinctiveness.

Sources

Type material: See above, NMW 54212, NMW 54213 and NMW 54215.

Iranian material: None.

Comparative material: BM(NH)1974.2.22:1324, 186.2 mm standard length (); BM(NH)1920.3.3:41-50, 12(5 examined), 110.9-165.3 mm standard length (); BM(NH)1920.3.3:31-40, 10, 141.7-310.9 mm standard length ();

Luciobarbus mursa
(Güldenstaedt, 1773)

L. capito (above) and L. mursa (below) Aras River, 2 October 1994
Courtesy of Asghar Abdoli

Common names

سس ماهي (= sos, sas or sass mahi), mahi siah (= black fish), zardek-e qalami (= slender or straight yellow one), ses mahi koloft safid rud (= Safid River thick fish, the meaning of ses, sos or sas being unknown but referring to "Barbus"), sas mahi-ye lab koloft (= thick lip "Barbus" fish), زرده پر (= zardehpar).

[mursa or shchirbit in Azerbaijan; murtsa or mursa in Georgian; murtsa or Araksinskaya murtsa, both in Russian].

Systematics

Cyprinus mursa was originally described from the Kura River at Tbilisi, Georgia. Syntypes are presumed lost (Bogutskaya, Bănărescu and Almaça in Bănărescu and Bogutskaya, 2003)

Barbus miliaris De Filippi, 1863 described from a "fiumicelli presso Teheran" (= a stream near Tehran), Barbus mursoides Kessler, 1877 described from Transcaucasia (presumably the Kura-Araks basin), Barbus microphthalmus Sauvage, 1882 from "Tiflis" (presumably the Kura River at Tbilisi, Georgia) (and Barbus macrophthalmus and Barbus mycrophtalmus in Chantre (1882) which are presumably misspellings of this name; the former is in any case preoccupied by B. macrophthalmus Bleeker, 1855 described from Indonesia), Barbus kessleri Derzhavin, 1929 described in Latin from the "Keredsh flumen" (= Karaj River near Tehran), and Barbus dageti Fowler, 1958 are synonyms. Barbus dageti was coined because Fowler believed Barbus kessleri was preoccupied by Puntius kessleri Steindachner, 1866; Puntius Hamilton, 1822 is not now considered a synonym of Barbus sensu lato (Eschmeyer, 1990) although Eschmeyer et al. (1996) have Barbus kessleri listed as preoccupied by Puntius kessleri. Eschmeyer et al. (1996) record 3 syntypes "whereabouts unknown" for Barbus dageti, i.e. Derzhavin's Barbus kessleri types. Dadikyan (1986) refers Aras River fish from Armenia to Barbus mursa mursoides.

Three specimens (presumably syntypes)(MZUT N.676) of Barbus miliaris are stored in the Istituto e Museo di Zoologia della R. Università di Torino (Tortonese, 1940).

The holotype of Barbus mursoides is in the Zoological Institute, St. Petersburg (ZISP 2863) from the Caucasus collected by Hohenacker in 1838.

Two syntypes of Barbus microphthalmus, measuring 340 mm total length, are in the Muséum national d'Histoire naturelle, Paris (MNHN A.3923, formerly MNHN 1881-1007 and MNHN 1881-1008) (Bertin and Estève, 1948).

Howes (1987) considers the generic placement of Barbus miliaris as problematical. It has a series of preanal scales and a prominent genital papilla similar to schizothoracines, and a lachrymal bone similar to Barbus (= Tor) grypus and B. sharpeyi. Karaman (1971) considers Barbus miliaris from the Namak Lake basin of Iran to be a subspecies of the Caspian Sea basin type subspecies, differentiated by larger scales (78-92 compared to 85-103), less fleshy lips, an undeveloped lower lip lobe, feebly ossified last dorsal fin spine, and shorter pectoral fins. Derzhavin (1929b) in describing his Barbus kessleri on fish 121-154 mm total length with well-developed gonads states that the lower lip is clearly trilobate. Berg (1949) recognises miliaris as distinct from mursa on the basis of a shorter snout, somewhat larger scales, fewer scale rows above the lateral line, smaller dimensions and different colour. Bianco and Banarescu (1982) and Almaça (1984b) retain it as a full species although Bianco and Banarescu (1982) also suggest that this species may be a subspecies of their wide-ranging taxon Barbus cyclolepis Heckel, 1837. Almaça (1984a) points out that his conclusion is based in part on small specimens in poor condition and that there is not enough data to take a sound decision (Almaça notes that gill raker counts are low and the lower lip lobe undeveloped in accordance with Karaman (1971) but these are characters which I believe may be size and age related). Almaça (1992) also distinguishes the two taxa on the shorter barbel in miliaris (not exceeding the middle of the eye as opposed to not exceeding the rear border of the eye), slope of the dorsal fin oblique in miliaris as opposed to oblique to nearly perpendicular, and pharyngeal teeth in miliaris 5,3 (or 4),2 as opposed to 4-5,3,2. These characters too may be size dependent or individually variable, as are those of Berg (1949). I consider that miliaris is at most a subspecies of mursa.

Key characters

The high scale counts are an important character as is the presence, usually, of a fleshy three-lobed lower lip.

Morphology

Dorsal fin unbranched rays 3-5, usually 4, followed by 7-8, usually 8, branched rays, anal fin unbranched rays 3 followed by 5 branched rays. Pectoral fin branched rays 13-17 and pelvic fin branched rays 6-8, usually 7. Lateral line scales 74-103, often 85 or more. Scales are small, horizontally elongated and almost rectangular, with a anterior margin variably indented, a very anterior focus, relatively few and well-spaced circuli, and few radii on all fields. A single pelvic axillary scale is not developed but a series of enlarged scales may be separated from other scales by a fold of skin. Gill rakers 7-18 (10-18 in Bogutskaya, Bănărescu and Almaça in Bănărescu and Bogutskaya (2003), lower counts from the literature perhaps not including rakers on the upper arm of the arch. There may also be differences due to size and, independent of size, the rakers on the lower arch anteriorly are variably developed, sometimes being reduced to bumps which were counted and sometimes not even bumps are present. The larger rakers reach the second adjacent raker when appressed. Vertebrae 41-45. Pharyngeal teeth 2,3,5-5,3,2, rarely 1,2,3,5-5,3,2,1; or with only 4 teeth in the main row (e.g. see Heckel (1843b)). Teeth are hooked and the fourth inner row tooth is slightly larger or smaller than the third. The fifth tooth is smaller (sometimes minute) than teeth 3 and 4 and may be pointed or blunt. The grinding surface below the tip is short, uneven and concave to rounded. The mouth is moderate in size, inferior, horseshoe-shaped with moderate to thick fleshy lips and an undeveloped to strongly developed median lower lip lobe (see above). Barbels are thick, the anterior one not extending back beyond the nostril level and the posterior one not exceeding the middle or posterior eye margin. The last dorsal fin spine is moderate to strong and has many, closely-packed denticles from one-half to four-fifths of the spine length, although denticles are lost in adults. The gut is elongate with 2-3 anterior loops. Chromosome number 2n = 100, NF 140 (Pourali Darestani et al., 2006).

Meristic variation in Iranian specimens:- dorsal fin branched rays 7(1) or 8(18); anal fin branched rays 5(19); pectoral fin branched rays 14(1), 15(2), 16(13) or 17(3); pelvic fin branched rays 7(13) or 8(6); lateral line scales 75(1), 80(2), 82(1), 84(1), 85(1), 87(1), 88(3), 89(3), 90(3), 91(1), 92(1) or 95 (1); total gill rakers 9(2), 10(1), 12(1), 13(5), 14(4), 15(2) or 16(3); pharyngeal teeth 2,3,5-5,3,2(7), 2,3,4-5,3,2(5), 2,3,5-4,3,2(2) or 3,4,5-5,3,2; and total vertebrae 43(1), 44(6) or 45(2).

Sexual dimorphism

Etessami (1982) reports an hermaphrodite in this species in the Namak Lake basin. A female specimen, 112.5 mm standard length, caught on 15 July had tubercles on the top and upper sides of the head. Male tuberculation in large adults has not been reported on.

Colour

Overall colour is a pale grey to olive-grey to brownish, slightly darker over the back, and the belly is white to yellowish-brown. The sides of the head and flanks can have golden tints. The iris is grey with a narrow rim of silver immediately around the pupil or may be yellow-gold. The dorsal and caudal fins are pale grey to dark reddish-brown. The caudal fin bears several series of small dark spots. The pectoral and pelvic fins have pale brown rays and transparent membranes but may be pink. The anal fin may be colourless except for a little grey pigment over the last unbranched and first branched rays to an overall reddish-brown. The margins of the pelvic and anal fins are well-developed and white, while the pectoral fin has a very narrow white margin. Young may have numerous dark spots on the back and upper flank, lost in adults.

Size

Attains 39.5 cm or 43 cm total length (Jolodar and Abdoli, 2004).

Distribution

This species occurs in the Kura River basin of the southwestern Caspian Sea and in southern tributaries of the Caspian from Iran. In Iran, it is reported in the Caspian Sea basin from the Aras to the Gorgan rivers (Abbasi et al., 1999; Kiabi et al., 1994; 1999; Abdoli, 2000), the Namak Lake basin (Wossughi, 1978; Rainboth, 1981; Almaca, 1984a; Bianco and Banarescu, 1982; Abdoli, 2000), and the Lake Orumiyeh basin in the Arnar Chay, Nowruzlu Chay, Tatavi and Zarrineh rivers (Günther, 1899; Abdoli, 2000).

Zoogeography

This species is possibly a Caspian Sea endemic, depending on the status of populations in the Namak and Orumiyeh Lake basins.

Habitat

Avoids muddy bottoms, preferring streams with rapid water, gravel and sand bottoms and a rich benthos. It may also be found in lacustrine habitats (Solak, 1977; Bogutskaya, Bănărescu and Almaça in Bănărescu and Bogutskaya, 2003).

Age and growth

Solak (1989b) reports a life span of 6 years in the Aras basin of Turkey. Maturity is attained at 2-3 years (Bogutskaya, Bănărescu and Almaça in Bănărescu and Bogutskaya, 2003).

Food

Food items include chironomids, as much as 70-100% of the diet at times, crustaceans such as copepods and ostracods, insects, worms, plankton, vegetation and detritus (Abdurakhmanov, 1962; Bogutskaya, Bănărescu and Almaça in Bănărescu and Bogutskaya, 2003). Iranian fish guts contain plant fragments, aquatic insects such as chironomids and Ephemeroptera (mayflies), and crustaceans such as amphipods.

Reproduction

Fecundity is up to 25,000 eggs. The spawning season is probably in May and June as noted for Georgian fish in Abdurakhmanov (1962) but may extend from April to August, the peak depending on locale (Bogutskaya, Bănărescu and Almaça in Bănărescu and Bogutskaya, 2003). Fish caught on 6 July in Mazandaran, 64 km west of Dasht had large, possibly atretic, eggs measuring about 1.5 mm although Bogutskaya, Bănărescu and Almaça in Bănărescu and Bogutskaya (2003) report a maximum egg diameter of 2.5 mm.

Parasites and predators

Masoumian et al. (2003) record Myxobolus azerbajdzanicus, M. kovali, M. squamae, M. tauricus, M. rutili and M. osmaniae while Pazooki et al. (2003) and Pazooki (2006) record Rhabdochona hellichi, Bothriocephalus gowkongensis and Paradiplozoon homoion, all reports from fishes captured in the Tajan and Zarem rivers of Mazandaran.

Economic importance

Said to taste even better than trout (Abdurakhmanov, 1962), it is caught by some anglers but is not commercially important (Bogutskaya, Bănărescu and Almaça in Bănărescu and Bogutskaya, 2003).

Conservation

Kiabi et al. (1999) consider this species to be near threatened in the south Caspian Sea basin according to IUCN criteria. Criteria include sport fishing, medium in numbers, habitat destruction, widespread range (75% of water bodies), present in other water bodies in Iran, and absent outside the Caspian Sea basin. Mostafavi (2007) lists it as near threatened in the Talar River, Mazandaran. Bogutskaya, Bănărescu and Almaça in Bănărescu and Bogutskaya (2003) report that is is extremely rare in Azerbaijan. Endangered in Turkey (Fricke et al., 2007).

Further work

The presence of this species in the Lake Orumiyeh basin and the taxonomic status of Namak Lake basin populations need careful examination.

Sources

Type material: ?

Iranian material: CMNFI 1970-0525, 1, 49.3 mm standard length, Gilan, Safid River near Mohsenabad (ca. 37º22'N, ca. 49º57'E); CMNFI 1970-0538, 5, 42.5-82.7 mm standard length, Gilan, Qezel Owzan River near Manjil (36º44'N, 49º24'E); CMNFI 1970-0545, ?, ? mm standard length, (); CMNFI 1970-0589, 1, 110.0 mm standard length, Gilan, Safid River opposite Kisom (37º12'N, 49º54'E); CMNFI 1979-0084, 2, 92.5-96.8 mm standard length, Mazandaran, Chalus River (no other locality data); CMNFI 1979-0253, ?, ? mm standard length, (); CMNFI 1979-0456, 2, 44.1-50.2 mm standard length, Markazi, Shah River at Lowshan (36º37'30"N, 49º31'E); CMNFI 1979-0481, 1, 142.7 mm standard length, Mazandaran, stream 3 km west of Ghalahleekesh (37º18'30"N, 55º31'E); CMNFI 1980-0132, 1, 112.5 mm standard length, Gilan, Safid River at Kisom (37º12'N, 49º54'E); CMNFI 1991-0158, ?, ? mm standard length, (); CMNFI 1993-0136, 1, 109.9 mm standard length, Mazandaran, Sardabrud (36º39'42'N, 51º22'36'E); CMNFI 2007-0086, 1, 182.2 mm standard length, Azarbaijan-e Khavari, Qareh Su basin near Nir (ca. 38º02'N, ca. 48º00'E); Behnke 8 Behnke 79 CMNFI 2007-00, ?, ? mm standard length, (); FMNH 51245, 2, 108.4-128.5 mm standard length, Markazi, Rayy (35º35'N, 51º25'E); ZMH 2429, 98.1 mm standard length, Markazi, Tehran (no other locality data).

Luciobarbus pectoralis
(Heckel, 1843)

Common names

basan, برزم (berzem or barzam), ? tu'ini.

[nebbash or sheikh san in Arabic; Heckel's Orontes barbel (Fricke et al., 2007)].

Systematics

Howes (1987) places this species in Barbus sensu stricto. Barbus perniciosus Heckel, 1843 described from "Gewässern bei Damascus", Luciobarbus Schejch Heckel, 1843 described from "Mossul" (also spelt schech on p. 1019 and p. 1098 in Heckel, presumably in error, and sometimes emended to scheich), Labeobarbus Orontis Sauvage, 1882 from the "Canal de l'Oronte à Antioche", Turkey and possibly Barbus (= Luciobarbus) kersin Heckel, 1843 (q.v.) and possibly Barbus (= Luciobarbus) barbulus Heckel, 1849 (q.v.) are synonyms (see Krupp (1985c)). Barbus pectoralis was described from the "Orontes" (Heckel, 1843b) but the catalogue in Vienna reads "Damascus" (possibly in confusion as this part of the catalogue has been overwritten).

The holotype of Labeobarbus orontis is in the Muséum national d'Histoire naturelle, Paris (MNHN A.3868), with a length of 600 mm (Bertin and Estève, 1948). The catalogue in the Naturhistorisches Museum Wien appears (the catalogue is overwritten here) to list a single specimen opposite each of the names Barbus perniciosus and Barbus pectoralis, probably the holotypes. Krupp (1985c) records the holotype of L. pectoralis as being 116 mm standard length (NMW 54474 and the holotype of B. perniciosus as being 105 mm standard length (NMW 54472). My examination of NMW 54474 showed a length of 117.8 mm standard length.

Karaman (1971) places pectoralis as a subspecies of Barbus (= Luciobarbus) capito but Almaça (1986) disagrees on several grounds, especially on the number of pharyngeal teeth (5 in capito and 4 in pectoralis in the main row (yet Heckel (1843b) gives 5 main row teeth for pectoralis, 4 for the synonym schejch), see also below). Luciobarbus kersin has 5 main row pharyngeal teeth, an indication that it may be distinct. Almaça (1986) points out that the specimen in the Naturhistorisches Museum Wien (NMW 54475) referred to as the holotype of pectoralis by Karaman (1971) is from the wrong locality and was collected at a later date.

Barbus Rajanorum Heckel, 1843 described from "Aleppo" and later in Heckel (1846-1849a) from "Gewässern von Aleppo" is a hybrid of this species and Capoeta damascina (F. Krupp, in litt., 1986) and Almaça (1991) also believes it to be founded on a hybrid; see also Almaça (1983; 1991), Berg (1949) and Karaman (1971) for conflicting views). Almaça (1983) could not find any specimens attributable to Barbus rajanorum and the holotype housed in the Naturhistorisches Museum Wien is lost. However, the type locality for this taxon is "Aleppo" (Heckel (1843b) and Krupp (1985c) states that the holotype is NMW 54494, 190 mm standard length, Aleppo, 1842, Th. Kotschy. The catalogue in Vienna lists a single specimen and the card catalogue in 1997 lists this fish as the holotype.

Karaman (1971) places Barbus (= Luciobarbus) barbulus, Luciobarbus schejch and fish Heckel (1843) referred to Luciobarbus mystaceus (Pallas, 1814) as synonyms of B. rajanorum. Berg (1949) also places L. mystaceus of Heckel in B. rajanorum. However mystaceus of Pallas would have priority (authors such as Karaman (1971) and Almaça (1983, 1991) refer the species description to Heckel (1843) in error) but, as Berg (1949) points out, Pallas's Cyprinus mystaceus is partly Barbus (= Luciobarbus) mursa and Barbus (= Luciobarbus) capito. Almaça (1983) recognises Barbus mystaceus with two subspecies, mystaceus from Aleppo, Tigris at Mosul and the Euphrates and barbulus (see above under this latter species). Krupp (1985c) places Barbus (= Luciobarbus) barbulus and Heckel's Luciobarbus mystaceus in Barbus pectoralis.

I am uncertain as to the identity of Barbus mystaceus (Pallas, 1814) reported by Heckel (1843b) from the "Tigris bei Mossul", Iraq, in regard to Iranian Luciobarbus species and do not assign any Iranian specimens collected by me to it. F. Krupp (in litt., 1987) considers Heckel's mystaceus to be identical with B. barbulus but that Heckel's mystaceus differs from that of Pallas, as previously noted by Berg (1949). Heckel's B. mystaceus is most probably either B. barbulus or B. pectoralis.

Barbus schejch is recognised as a distinct species by Almaça (1983, 1991) but only one specimen, a syntype from the Tigris (in the Naturhistorisches Museum Wien, NMW 50399), was available to him. It measures 136.5 mm standard length. Two other specimens identified as syntypes of this this species are under NMW 54520 with standard lengths 175.4 and 270.7 mm. The barbels in the 50399 are very short, not reaching the eye and about equal in length while in the other two syntypes the posterior barbel reaches the mid-eye and the barbels are subequal. The lips are fleshy, like Luciobarbus barbulus, but there is no central lobe in 50399, present in the smaller of the two other syntypes and poorly developed in the larger. The complete dorsal fin spine bears 29 teeth in the 50399 and 29 or 35 in broken spines of the other two fish. Gill rakers number 22 in 50399 and 16 or 18 in the other two fish. Lateral line scales number 52 (or 54 to end of scale row on caudal fin) in the syntype and 57(58) or 58(60). Main row pharyngeal teeth are 4-4 in 50399, missing in the other two fish. These data are somewhat contradictory and further data are required to resolve the status of this nominal species. The catalogue in Vienna lists 4 fish in spirits and 4 fish stuffed.

The synonymy of Luciobarbus barbulus with L. pectoralis remains uncertain. The putative holotype of B. pectoralis (NMW 54474) was compared with a specimen of similar size from Iran referred to L. barbulus (CMNFI 1973-0393). The L. pectoralis specimen is partly dried so direct measurement comparisons are not possible. The L. pectoralis specimen has more teeth in the dorsal fin spine (27 teeth even though it is broken off, much more than 30 presumably in the intact spine), barbels in pectoralis are shorter, the posterior one reaching the anterior half of the eye, the anterior one short of the mouth angle, mouths similar in shape but lips appear to be less fleshy, gill rakers number 16, lateral line scales number 44, and 4 main row pharyngeal teeth but there is a trace of a fifth tooth not fully ossified. ? check counts on NMc fish?

Key characters

The dorsal spine is much stronger than in Luciobarbus barbulus and arises from an elevated base that supports the dorsal fin base. The body is deeper than in L. barbulus and the lips usually less fleshy.

Morphology

Dorsal fin with 4 unbranched and 7-9 branched rays (7 in the holotype, usually 8), anal fin with 3 unbranched and 5, rarely 6, branched rays. Pectoral fin branched rays 16, pelvic fin rays 8. Lateral line scales 42-60 (44 in the holotype; 42 in Barbus perniciosus). Gill rakers 14-17 (to 21 if Barbus schejch is included (Almaça, 1986)). Pharyngeal teeth 2,3,4-4,3,2 in 16 fish, 2,3,5-5,3,2 in 9 fish examined by Krupp (1985c), rarely 2,3,5-4,3,2 or 2,3,4-5,3,2 (1 fish each), spoon-shaped or pointed with the fourth tooth of the inner row large and globose. Larger fish usually have 4 teeth in the main row and the fourth tooth is globose. Smaller fish with 5 teeth in the main row have cylindrical teeth. All intermediates stages exist (Krupp, 1985c). The mouth is moderate in size and subterminal. Lips are thin to moderate and the median lobe of the lower lip may be present or absent. Barbels are thin to moderate, the anterior one not extending back beyond the nostril to anterior eye margin level and the posterior one not beyond the middle to the posterior margin of the eye. The last unbranched dorsal fin ray is moderate to very strong with a low density of denticles extending along much of the ray. Larger specimens have a lesser extent of denticles along the ray. The body form is extremely variable.

Sexual dimorphism

Unknown.

Colour

The back is brown to bluish-green and the flanks yellowish to silvery-white.

Size

46.9 cm standard length (Krupp, 1985c).

Distribution

Tigris-Euphrates basin and the Orontes and Quwayq rivers. In Iran, it is found in the Tigris River basin in the Hawr Al Azim and the lower Karkheh, Karun and Jarrahi rivers, in the Kor River basin, and in the Gulf basin in the middle and lower Helleh, middle and lower Mand and Dasht-e Palang rivers (Abdoli, 2000).

Zoogeography

Almaça (1984b) considers that the origin of this species (as Barbus schejch) lies with a group that migrated southwards in the late Pliocene from the Dacian Lake of the Sarmatian Sea and speciated in Mesopotamia and later (Almaça, 1991) that this species (as Barbus (= Luciobarbus) pectoralis) originated from a colonisation wave from South Europe.

Habitat

Unknown.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

None reported from Iran.

Economic importance

Occasionally caught and used for food.

Conservation

Endangered in Turkey (Fricke et al., 2007). Confusion over its identity has made a conservation assessment difficult for Iranian waters.

Further work

The taxonomic status of this species and its presence in Iranian waters should be resolved.

Sources

Type material: See above, Barbus pectoralis (NMW 54474).

Luciobarbus subquincunciatus
(Günther, 1868)

Common names

سليماني (= solimani or soleimani).

[abou khazzama, a'djzan, agzan or adzan, all in Arabic; black spot barb, leopard barbel, Mesopotamian barbel].

Systematics

The type locality of this species is unknown. Günther (1868) gives the following account:- "From the Collection of the East-India Company.- Although no record of the history of this specimen has been preserved, it is probable that it came from Mesopotamia, as other examples from this country are preserved in precisely the same manner". The type specimen is a "Skin, 15 inches long" (= 5.9 cm).

Krupp (1985a) removes this species from Bertinius Fang, 1943 since the enlarged molariform pharyngeal teeth on which this genus was erected are due to convergence and are not evidence of monophyly. Howes (1987) places this species in his Barbus sensu stricto.

Key characters

The numerous, large, dark spots arranged in an almost quincunx pattern are distinctive.

Morphology

Dorsal fin with 3-4 unbranched and 8 branched rays, anal fin with 3 unbranched and 5 branched rays. The last unbranched dorsal fin ray is very strong and bears denticles along almost its whole length or three-quarters of the length. Pectoral fin branched rays 14-18, pelvic fin rays 7. Lateral line scales 75-88. Scales have few radii on all fields, fine circuli and a focus slightly subcentral anterior. Total gill rakers about 10-13, broad based and triangular in shape with highly tubercular distal or foliose margin. The longest raker reaches the one below when appressed. Total vertebrae 45 (Howes, 1987). Pharyngeal teeth 2,3,3,-3,3,2, occasionally 2,3,4-4,3,2, the usual number of teeth in the inner row in large specimens being 3 (Krupp, 1985a). The third inner row tooth is the biggest by far and is molariform. Juveniles have 5 inner row teeth (Krupp, 1985c). Total vertebrae 46. The mouth is horseshoe-shaped, small and inferior. Lips are well-developed and fleshy. The median lobe of the lower lip is undeveloped. The barbels are thick. The gut has many anterior loops, the number increasing with size.

Meristics for Iranian material:- dorsal fin branched rays 8(1), anal fin branched rays 5(1), pectoral fin branched rays 14(1), and pelvic fin branched rays 5(1). Lateral line scales 83(1). Total gill rakers ?. Pharyngeal teeth 2,3,4-4,3,2 (1).

?others

Sexual dimorphism

Unknown.

Colour

The whole body, head, fins, barbels, lips and even eyeball are covered with dark spots about the same size as or larger than the eye. Some larger flank spots are 2-3 times the eye diameter. Spots on fins are elongated along the fin length. These spots are arranged in patterns similar to a quincunx, hence the species name. A quincunx comprises four spots, one at each corner of a square with the fifth spot in the middle of the square. Sometimes a spot runs into an adjacent one. Some spots below the lateral line may be elongate, three times longer than wide, and arranged vertically. Occasional fish lack spots on the mid-flank but are still distinctively spotted elsewhere. The overall colour is greenish to brownish-yellow with the belly white. Peritoneum dark brown to black.

Size

Reaches 33.8 cm total length (Menon, 1956), 45.7 cm (Khalaf, 1961) or 60 cm (Sauvage, 1884).

Distribution

Found in the Tigris-Euphrates basin including its Iranian portion in such rivers as the Jarrahi (Wossughi, 1978; Rainboth, 1981; Abdoli, 2000).

Zoogeography

Almaça (1991) believes that this species originated in Mesopotamia.

Habitat

Unknown but recorded from rivers and artificial reservoirs.

Age and growth

Şen et al. (1992) examined 9 fish in Keban Dam Lake, Turkey and found age groups 3-7, growth rings being best expressed in sectioned dorsal fin rays. The length-weight relationship was logw = - 5.78723 + 3.27533 logl and the mean K_(TL) was 0.8234. The b value indicates the habitat is suitable for the species.

Food

The molariform pharyngeal teeth and evidence from gut contents showed this species is an obligate molluscivore (Krupp, 1985a).

Reproduction

Unknown.

Parasites and predators

None reported from Iran.

Economic importance

This species occasionally occurs in commercial catches in Khuzestan but is not a common food fish compared to other Barbus sensu lato species. It has been investigated for aquaculture in Khuzestan but fish are rare and so adults are caught and released.

Conservation

This species is now very rare in Iran and "critically endangered". Reports of 1 fish taken in the Gav Masiab River in 1991, 4 fish from the Karun River in 1995 and 1 fish from the Karun River at Ahvaz in 1997 were the only records for the 1990s (M. Ramin, pers. comm., 2000). The stock of this species in the Gav Masiab River is severely reduced and during 4 years of collecting in western Iran only one fish was caught (J. Valiallahi, www.modares.ac.ir, downloaded 4 July 2000; pers. comm., 2001; Valeolahy, 2000).

Syrian populations in the Euphrates River and parts of its tributaries are also in a parlous state (R. Beck, pers. comm., 2000).

Further work

The biology of this distinctive species should be investigated.

Sources

Type material: ?

Iranian material:- CMNFI 1993-0133, ?, ? mm standard length, (): ZMH 2506, 1, 308.0 mm standard length, Kermanshahan, Karasu-Gamasiab-Seymarreh, Kermanshah (); FMNH70794,1, ?, Javanarud near Kermanshah (); and market specimens from Khuzestan.

Comparative material:- CMNFI 1980-1036, 1, 177.5 mm standard length, Turkey, Keban Dam on Murat Nehri near Elazig (38º41'N, 39º14'E); CMNFI 1986-0676, 1, 283.0 mm standard length, Turkey, Keban Dam on Murat Nehri (no other locality data); BM(NH) 1874.4.28:15, 1, 415.3 mm standard length, Iraq, Tigris River near Baghdad (33º21'N, 44º25'E); BM(NH) 1875.1.14:3-5, 3, 377.6-468.2 mm standard length, Iraq, Tigris River (no other locality data); BM(NH) 1974.2.22:1353, 1, 253.4 mm standard length, Iraq, Sirwan River, Diyala (no other locality data);

Luciobarbus xanthopterus
Heckel, 1843

Common names

گطان (gatan or gattan).

[gattan or ghattan, nobbash, or thekar, all in Arabic; yellowfin barbel].

Systematics

Howes (1987) places this species in Barbus sensu stricto. Almaça (1983) briefly reviews the placement of this species in synonymy; most ichthyologists now regard it as a distinct species. Luciobarbus xanthopterus has been considered as a variant of L. schejch but differs in gill raker count (10-13 in xanthopterus, 21 in schejch) and main row pharyngeal tooth count (5 in xanthopterus and 4 in schejch) (Almaça, 1983; see also discussion under Luciobarbus pectoralis; Heckel (1843b), however, gives the main row count for xanthopterus as 4). It has also been considered as a synonym of esocinus (q.v.) (Almaça, 1986).

Fayazi et al. (2006) used mtDNA to study differentiation between populations of this species in the Karun, Karkheh and Jarrahi rivers in Iran. Diversity was low although the Karun and Karkheh fish grouped together, leading to the recommendation that fish from the Jarrahi should not be used to stock other river basins.

Almaça (1986) records syntypes of Luciobarbus xanthopterus from the type locality given by Heckel (1843b) "Tigris bei Mossul", Iraq in the Naturhistorisches Museum Wien under NMW 54841 (10 specimens, one large fish at 216.5 mm standard length and 9 smaller fish at 48.6-63.4 mm; one of these was noted as being listed as the lectotype in 1997 (? the largest), and the catalogue number was 54841a) and NMW 54786 (1 specimen, 292.8 mm, not listed as a type in 1997). Material listed under NMW 1843 (? catalogue number or collection date) may also be syntypes. Eschmeyer et al. (1996) list 1 dried syntype under NMW 91215. The catalogue in Vienna lists 4 fish in spirits and 2 fish stuffed.

Key characters

This species is characterised by two pairs of barbels, a serrated dorsal fin spine, small scales (57-68 in lateral line), and a subterminal to terminal and oblique mouth. The elongate postorbital length is also seen in Luciobarbus esocinus but is more marked in the latter (check this?) and scale counts are different (check?).

Morphology

Dorsal fin with 4 unbranched and 7-9, usually 8, branched rays, anal fin with 3 unbranched and 5 branched rays. Pelvic fin branched rays 8. Pectoral fin branched rays 14-18 (Jawad, 1975). Lateral line scales 57-68. Scales have rounded dorsal, ventral and posterior margins and an anterior margin with a central protuberance and indentations above and below. Circuli are fine and radii are found on the anterior and posterior fields and sometimes the lateral fields. The focus is subcentral anterior. There is no distinct pelvic axillary scale. Gill rakers 7-13, short and reaching the adjacent raker when appressed. Pharyngeal teeth 2,3,5-5,3,2, strongly hooked the fourth tooth of the inner row being the largest and anterior teeth being rounded with a small flat or concave grinding surface below the tip. Qasim and Niazi (1975) gave a tooth formula of 4,3,2-2,3,4, i.e. 2,3,4-4,3,2 as does Heckel (1843b) and teeth were molariform. Total vertebrae 44 (Howes, 1987), 40-42 (Qasim and Niazi, 1975), 42 (Wossughi, 1978) or 46 (BM(NH) 1973.5.21:198). The last unbranched dorsal fin ray is moderately to very strong, has a low denticle density and is serrate along much of its length. Barbels are thin, the maxillary barbels are longer than rostral barbels but both are short, the rostral ones not extending beyond the level of the nostrils and the maxillary ones not extending back beyond mid-eye to rear eye level. The gut has one anterior and two posterior loops in an elongate s-shape. The mouth is moderate in size, inferior and an elongate u-shape in young fish and as development progresses becomes terminal in adults (Karaman, 1971; Almaça, 1984b). Lips are thin to moderate and the lower lip has no median lobe.

Iranian specimens had the following meristics:- dorsal fin branched rays 8(2); anal fin branchd rays 5(2); pectoral fin branched rays 18(2); pelvic fin branched rays 8(2); lateral line scales 57(1) or 68(1); total gill rakers 7(1) or 10(1); pharyngeal teeth 2,3,5-5,3,2(2); and total vertebrae ?.

Sexual dimorphism

Unknown.

Colour

The body is without distinctive marks. The back is brownish to bluish-grey, the flanks silvery to silvery-yellow, and the belly white. The scales are outlined by melanophores. The overall colour from a marsh habitat is darker than from a riverine habitat, the pigment outlining scales being thicker for example especially at the scale base. The eye is red in marsh specimens, white to yellowish elsewhere (M. Al-Mukhtar, pers. comm., 1995). All fins are lemon-yellow to orange with some darker melanophores. The unbranched dorsal fin rays and the uppermost caudal fin rays are black.

Two small specimens from Iran have irregular spots and blotches on the flank. The peritoneum is silvery with melanophores developed dorsally.

Size

Al-Hassan et al. (1986) report a specimen 1.5 m total length and 8.6 kg from the Abu Al Khasib area in the Shatt al Arab, Iraq.

Distribution

This species is found in the Tigris-Euphrates basin including its Iranian portion such as the lower Karun River and adjacent lower reaches of the Jarrahi River (Abdoli, 2000), the Qareh Su at Kermanshah, the Karkheh River and Hawr al Azim.

Zoogeography

Almaça (1984b, 1991) considers that the origin of this species lies with a group that migrated southwards in the late Pliocene from the Dacian Lake of the Sarmatian Sea and speciated in Mesopotamia.

Habitat

van den Eelaart (1954) and Al-Hamed (1966b; 1972) describe the habitat for this species in the Tigris River as distributed in the deep, open waters of lakes and vegetated marshes and to a lesser extent in the river and its tributaries. Mature fish move upstream to the spawning grounds in February-March and spent fish descend to their original habitat in lakes and marshes. In summer, beginning in June, under low water level conditions and high temperatures, the smaller fish remain in the deepest depressions of lakes but the large fish (3 kg or more) migrate up rivers and tributaries in search of cooler water, returning in September and October when temperatures fall to fatten over winter.

In Khuzestan, this species is most abundant in the Karkheh River in March and in the Hawr al-Azim in December, migrating from the wetland to the river in spring. Younger fish are more abundant in the wetland and older fish in the river (Iranian Fisheries Research Organization Newsletter, 22:3, 2000; Eskandary et al., 2000). Another study showed this species to be most abundant in the Karkheh River in December, with a migration from wetlands in spring to the main river (Tehran Times, 1 October 2000).

Age and growth

Life span is at least 11 years (Al-Ahmed, 1966a). Al-Hamed (1966b; 1972) working on Tigris River fishes found males to mature at about 43 cm and females at about 48 cm, maturity being attained in the fourth year of life and spawning occurring at the beginning of the fifth. Some fish mature at age group 3 and some as late as age group 5. Males outnumber females on the spawning grounds, comprising 62% of the population. Tigris River and Al-Tharthar reservoir fish in Iraq had 7 age groups with growth good in the first three years and slower thereafter (Ali, 1979). In Keban Dam Lake, Turkey, age determination was best made on sectioned dorsal fin rays (of scales, otoliths, vertebrae and opercula) and up to 9 age groups were detected (Duman and Şen, 1995).

In the Karkheh River, male fish are mature at 151-200 mm (one year old) and females at 501-550 mm (3 years old). The sex ratio is 1:1.31 for males:females but this is not significantly different from 1:1 (Eskandary et al., 2000).

Food

Al-Hassan et al. (1986) report isopods and molluscs. Al-Hamed (1965) considers this species to be an omnivore, consuming filamentous algae, detritus, frogs, molluscs and fishes and even planktonic organisms. Organic matter is obtained in periods of food shortage by engulfing mud from the pond bottom. van den Eelaart (1954) reports food to be plants, epiphytes and plankton. In cold winters they take no food. Ali (1979) for Iraqi waters gives insects and plankton as the principle foods. Al-Shamma'a et al. (2009) found fish from Lake Habbaniyah, Iraq to feed mostly on animal materials (76.25) including molluscs and insects and their larvae. Feeding was most active in spring but high feeding intensities were also observed in autumn.

In Khuzestan it is omnivorous, feeding mainly on insects and vegetation, but also taking secondarily shrimps, snails and ostracods (Iranian Fisheries Research Organization Newsletter, 22:3, 2000; Eskandari et al., 2003). In the Karkheh River food is insects and vegetation mainly, with shrimps, gastropods and ostracods secondary food choices (Tehran Times, 1 October 2000). The intestine fullness is greater in fish in the Hawr al Azim, less in the Karkheh River which is used mainly for spawning (Eskandari et al., 2003).

Reproduction

van den Eelaart (1954) and Al-Hamed (1966b; 1972) studied the reproduction of this species in Iraq. Eggs are deposited on fine gravels overlying a layer of coarse sand in shallow, wide holes excavated by the fish. Water depth varies from 30 to 150 cm. Egg diameter is 1.0 mm and fecundity up to 340,000 grey eggs. Al-Hassan et al. (1986) record up to 350,000 eggs for their large fish from the Shatt-al-Arab. The spawning season on the Tigris River between Beled and Tigrit is April and May. Fish appear on the spawning grounds in schools just before dark and remain there until shortly before midnight, making loud noises by splashing, jumping and chasing.

In Khuzestan, spawning fish are 63.7-80.0 cm total length with a relative fecundity of 18.9-142.5 eggs/g body weight and a minimum and maximum absolute fecundity of 136,924 and 549,211 eggs (Iranian Fisheries Research Organization Newsletter, 22:3, 2000; Eskandari et al., 2003). In the Karkheh River, spawning took place at surface water temperatures of 25.5-28.65°C in turbid water after a spring migration from wetlands into the river (Tehran Times, 1 October 2000; Eskandary et al., 2000). Spawning occurs annually in May and June in the Karkheh River and maximum egg diameter is 2.25 mm (Eskandary et al., 2000).

Parasites and predators

Bykhovski (1949) reports a new species of monogenetic trematode, Dactylogyrus inutilis, from this species in the Karkheh River, Iran. Ebrahimzadeh and Nabawi (1975) list Anisakidae from this species in the Karun River. Moghainemi and Abbasi (1992) record a wide range of parasites from this species in the Hawr al-Azim in Khuzestan.

Economic importance

This species appears regularly in the markets of Ahvaz and Wossughi (1978) states it is of great economic importance. Sharma (1980) reports that gattan is the third most important fish species at Basrah fish market, accounting for 510,503 kg for the period from October 1975 to June 1977. Petr (1987) reports the annual catch for 1976 in Iraq was 2543 t. This species has been studied for pond culture in Khuzestan where over 95% of young survived, using hormones to stimulate reproduction (Iranian Fisheries Research Organization Newsletter, 28:3, 2001). Zadeh et al. (2009) investigated the optimal dietary carbohydrate to lipid ratio for fingerlings of this species. Mortezavizadeh et al. (2009) induced reproduction in this species with carp pituitary extract. Sperm production was increased, 86% of females responded positively, 480 eggs/gram were produced and mean survival rate was 77.95%. Propagation was best at the beginning of March at 19.0-24.5°C.

englers in Iraq catch this fish which will reject any bait showing resistance, requiring a fast strike at the first indication that the fish has taken the bait.

Conservation

Several hundred thousand gattan juveniles have been introduced into the Hawr al Azim in Khuzestan in order to restock and protect this resource. The fish were artificially bred from breeders using hormone treatment (Iranian Fisheries Research Organization Newsletter, 39:3, 2004; Network of Aquaculture Centres in Asia, downloaded 11 January 2007). Endangered in Turkey (Fricke et al., 2007).

Further work

Sources

Type material: ?

Iranian material: ZMH 4071, 1, 151.6 mm standard length, Kermanshahan, Qareh Su at Kermanshah (?). uncatalogued, 2, 93.7-112.8 mm standard length, Kermanshahan, Sarab-e Yavari (34º28'N, 46º56'E);

Comparative material: BM(NH) 1893.6.23:25, 1, 198.0 mm standard length, Iraq, Al Faw (29º58'N, 48º29'E); HL51.5, Postorb 26.6, for body proportions; g.r 13; BM(NH) 1973.5.21:197, 1, 227.8 mm standard length, Iraq, Shatt al Arab (no other locality data) HL 60.3, postorb 28.8, g.r. 21; BM(NH) 1973.5.21:198, 1, 143.0 mm standard length, Iraq, Shatt al Arab (no other locality data). HL 34.6, postorb 15.1, g.r. 18.

Genus Mesopotamichthys
Karaman, 1971

Much of the past literature on this genus appeared under Barbus (q.v.)

Mesopotamichthys sharpeyi
(Günther, 1874)

Common names

بني (benni, benny, beni, binni, bini, binny (meaning possibly nose)), سليماني (= solimani or soleimani, meaning unknown).

[binni, bunni, bunnei, bunia or aradah, all in Arabic].

Systematics

Howes (1987) places this species outside the genus Barbus sensu stricto as defined by him because it has the non-elongate lachrymal bone with a sensory canal running along the antero-dorsal border, a derived condition. Karaman (1971) erected a new genus for this species, Mesopotamichthys, which is not accepted here nor by Krupp (1985c). However Bănărescu (1997) and Ekmekçi and Banarescu (1998) recognise this genus as valid. The large-scaled "Barbus" of Southwest Asia require a detailed revision probably with additional, molecular characters (see also under Carasobarbus luteus and Kosswigobarbus kosswigi).

Barbus faoensis Günther, 1896 described from "Fao (Persian Gulf)", Iraq is a synonym; Karaman (1971) retains it as the subspecies of the lower part of the Tigris-Euphrates basin with the type subspecies in the upper part of the Tigris River basin.

Barbus sharpeyi was described from "Baghdad". The three syntypes are in the Natural History Museum, London (BM(NH) 1874.4.28:20 labelled "R. Tigris nr. Baghdad. Sharpey", well sealed in its jar and not measured accurately, and BM(NH) 1874.4.28:27 and BM(NH) 1875.1.14:16 labelled "R. Tigris. Sharpey" and measuring 147.6-178.0 mm standard length). The large holotype of Barbus faoensis is also there, strongly sealed in its jar, labelled "Persian Gulf. Kurrachee Museum" (BM(NH) 1888.5.17:4).

Al-Mukhtar and Al-Hassan (1999) describe a hybrid of this species and Carassius auratus from Al-Hayei (= Al Ha'i), a seasonal lake between the Karkheh and Dez rivers in Khuzestan.

Key characters

The absence of barbels, the last dorsal fin unbranched ray moderately ossified but lacking teeth, and the low scale count are characteristic.

Morphology

Dorsal fin with 4 unbranched and 7-9, usually 8, branched rays, anal fin with 2-3 unbranched and 4-5, usually 5, branched rays, pectoral rays 13-19, usually 16-17, and pelvic fin branched rays usually 8. The last third or quarter of the last unbranched dorsal fin ray is thin, flexible and tapering. The gut has several loops, two anteriorly and three posteriorly. Pharyngeal tooth formula is 2,3,5-5,3,2, sometimes with only 4 teeth in the main row but the anterior tooth is missing in both small and large fish and so does not appear to be age related, with teeth hooked at the tip but not strongly on the posterior teeth which are spoon-shaped with the hollow of the spoon filled in with bone. Total gill rakers number 13-19, reaching the raker below or just beyond when appressed. Total vertebrae 38-42 (lower values, 38-39, may not include hypural plate). Lateral line scales 29-37. A small pelvic axillary scale may be present or scales in this area may be so weakly developed as not to be an apparent axillary scale. Scales have a slightly anterior focus, fine concentric circuli, many radii on all fields and the posterior, exposed field bears numerous small tubercles. The mouth is slightly subterminal. Lips are well-developed but not fleshy and the lower lip is interrupted in the middle. Microscopic studies of the pharynx and oesophagus have been carried out by Alboghobeish and Moosavi (1998) who confirm that it is adapted for herbivory. Chromosome number 2n=98 (Balasem et al., 1994). Alboghobeish and Hamidian (2006) studied the distribution of alarm cells in the skin of this species.

Iranian fish have the following meristics: branched dorsal fin rays 8(2), branched anal fin rays 5(2), branched pectoral fin rays 16(2) and branched pelvic fin rays 5(2). Lateral line scales 30(1) or 31(1). Total gill rakers 16(1) or 18(1). Pharyngeal teeth 2,3,5-5,3,2(2). Total vertebrae 41(1) or 42(2) based on CMNFI 79-0087 (42) and 87-0017 (41 and 42). 41(3) or 42(2) based on BM(NH) 1920.3.3:71-75, 42 (1973.5.21:195), 41 (1973.5.21:196), 42 (1874.4.28:27), 41 (1875.1.14:16)

Sexual dimorphism

Unknown.

Colour

Overall colour is greenish to light brown or golden brown with the belly white to silvery or yellowish-brown. Scales on the back and uppermost flank have solid dark brown pigment on the exposed part of the scale. The scale edge is thinner and so appears lighter. The eye is brownish orange, golden or silvery. Fins are darker than the adjacent body, a deep reddish-brown, with melanophores on rays and membranes in preserved fish. The peritoneum is black.

Size

Attains 55 cm and 4 kg (van den Eelaart, 1954; Al-Hamed, 1966b; 1972). Reaches at least 3.5 kg in Khuzestan (J. Gh. Marammazi, pers. comm., 1995).

Distribution

This species is found in the Tigris-Euphrates River basin including its Iranian portion in such marshes as the Hawr Al Azim and in rivers and in the northern Gulf basin in the Zohreh River (Marammazi, 1995; Abdoli, 2000).

Zoogeography

Karaman (1971) considers that this species originated from the Indian line of the Torini, a tribe of Cyprinidae, in which Karaman includes such genera as Carasobarbus and Kosswigobarbus, Garra, Hemigarra (recognised as Hemigrammocapoeta here) which have Iranian members.

Habitat

van den Eelaart (1954) and Al-Hamed (1966b; 1972) report some movement from lakes and marshes, from the end of February to the beginning of March, to rivers in the Tigris-Euphrates basin of Iraq during floods for about 3 weeks. There is a return to lakes and marshes for spawning in mid-March to mid-April. However, most fish remain in marshes and lakes for most of the year, in overgrown areas avoiding open water. Low water levels and high temperatures in the lakes and marshes may cause a migration to their deepest parts or into the lower reaches of the main and more permanent rivers. This species is less tolerant of low oxygen than Luciobarbus xanthopterus which probably accounts for them not being caught together in any number.

Marammazi (1994) considers this species to be stenohaline and so restricted in its distribution in the Zohreh River which drains to the northern Persian Gulf. The influence of salinity on growth rate is examined by Orian et al. (1993).

Age and growth

Al-Hakim et al. (1976) studied some aspects of the biology of this species in Razzaza Lake, Iraq. Females are longer and heavier than males at advanced ages. Life span of females is 9 years and for males 8 years. Maturity starts in the third year at 32-35 cm total length. Males mature earlier than females. Al-Hamed (1966a; 1966b; 1972) found Tigris River fish in Iraq to mature at 25 cm for males and 28 cm for females in the second year of life and spawning took place early in the third year. A few matured in age group 1 and some as late as age group 3. Males are somewhat more abundant than females on the spawning grounds, averaging 57.4% of the fish caught. Maximum age is 6 years. Ali (1982b) found this species to mature in the fourth year of life in Iraq, with growth better in the marshes than in Tharthar Reservoir. Epler et al. (1996) found fish up to age 6+ years in fresh and salty Iraqi lakes. Nasir et al. (1989) reports on the biology of this species in the Al-Hammar Marsh, Iraq and found a sex ratio of 1 female:3 males for all months and length groups caught. No explanation for this skewed ratio was found.

Food

Al-Hamed (1965) found this species to be strictly herbivorous, feeding on unicellular Chlorophyceae, diatoms and filamentous algae when young and on higher plants and detritus when older. Nasir et al. (1989) and Epler et al. (1996) confirm that this species in Iraq is completely herbivorous although some copepods and molluscs are taken, most probably incidental to filamentous algae, diatoms and detritus. van den Eelaart (1954) reports feeding even in cold winters. In the Karun River, diet includes such plants as Potamogeton, Salvinia, Nuphar and Phragmites (Annual Bulletin 1993-94, Iranian Fisheries Research and Training Organization, Tehran, p. 91-92, 1995).

Reproduction

van den Eelaart (1954) and Al-Hamed (1966b; 1972) studied reproduction in this species on the Tigris River in Iraq and Al-Nasih (1992) in fish ponds. Spawning occurs chiefly in lakes and marshes, with some spawning in the lower reaches of rivers. Eggs are deposited on submerged, or partially submerged, vegetation, from the surface down to about 1 m depth. Eggs are large, yellow and measure up to 1.7 mm in diameter and number up to 158,000. Epler et al. (1996) give a relative fecundity of 10,021 to 28,471 eggs for fish 4+ to 6+ in age from Iraqi lakes with fish spawning in April in a freshwater lake and February/March in a saline lake. Al-Nasih (1992) gives details of larval development. The spawning season in Lake Saniyah just north of Amara is March and April, with some ripe fish caught in May. Fish appear on the spawning grounds about sunset and left before darkness is complete. They return in the early morning and leave again at about 0800 hours. These fish chase each other, dart about singly or in pairs and sometimes come to the surface and splash. Al Mukhtar et al. (2006) investigated this species in the Hawizah Marsh as a source of spawners for aquaculture. Ripe eggs appeared in January and 25% were running in February and 30% in March. Half of the fish were spent in April. The spawning migration was lead by males in October and December with females increasing rapidly in February. Males disappeared in April. Absolute fecundity reached 236,160 eggs.

Petr (1987) reports spawning in Iran at 15-16°C in February in clean water of rivers with sandy bottoms. In the Karun River this species spawned in March-April in river estuaries (Annual Bulletin 1993-94, Iranian Fisheries Research and Training Organization, Tehran, p. 91-92, 1995). A specimen caught in March had well-developed testes. Spawning in Shadegan Marsh, Khuzestan is in March and in branches of the Karkheh River in March to April (Al Mukhtar et al., 2006). Shadegan Marsh is one of the most important spawning areas in Iran (Mohammadi and Marammazi, 2001).

Parasites and predators

Bykhovski (1949) reports a new species of monogenetic trematode, Dactylogyrus pavlovskyi, from this cyprinid in the Karkheh River, Iran. Ebrahimzadeh and Nabawi (1975) list species of the protozoans Trichodina and Myxosoma and the trematode Dactylogyrus as well as the nematode Camallanus lacustris, from this species in the Karun River. Jalali and Molnár (1990a) record two monogenean species, Dactylogyrus spp., in the Dez River and Molnár and Jalali (1992) a new species of monogenean, Dogielius persicus, from this species in the Dez and Karun rivers of Khuzestan. Gussev et al. (1993b) record Dactylogyrus pavlovskyi in the Dez River.

Masoumian et al. (1994) describe a new species of Myxosporea from the gills of this species in the Karun River, namely Myxobolus persicus, and later (Masoumian et al., 1996) another new species of Myxosporea, Myxobolus nodulointestinalis, in the gut lining of this species and also from rivers of southwestern Iran. Masoumian et al. (1996) describe a new species of Myxosporea, Myxobolus bulbocordis, from the heart of fish caught at various localities in Khuzestan. Molnár et al. (1996) report additional new species from this fish in Khuzestan, namely Myxobolus iranicus in the spleen and Myxobolus sharpeyi in the gill cartilage. Myxosporeans are potentially dangerous to fishes such as Mesopotamichthys sharpeyi which may be used in fish culture in Khuzestan. Masoumian and Pazooki (1999) list Myxobolus persicus, M. karuni, M. sharpeyi, M. nodulointestinalis, M. bulbocordis and M. iranicus from this species in various localities in Khuzestan. Shamsi et al. (2009) found Dactylogyrus nchoratus in this species from fish farms and the Karun River.

Peyghan (1994) reports ichthyophthiriasis in cultured Barbus (= Mesopotamichthys) sharpeyi in Khuzestan. This parasite causes severe skin and gill damage and mortality reaches 80%. A combination of formalin and malachite green with transfer of fish to another pond having a better environment cured the condition.

Molnár and Pazooki (1995) record philometrid nematodes from this species in the Karun River, and these are presumed to be a new species. Pazooki and Molnár (1998) later describe Philometra karunensis as the new species from the swimbladder and adominal cavity of this fish.

Moghainemi and Abbasi (1992) record a wide range of parasites from this species in the Hawr al-Azim in Khuzestan.

The monogeneans Dactylogyrus povlovskyi, D. barbioides, D. anchoratus, D. carassobarbi and Dogielius persicus are recorded from this species in the Karun River with heavier infestations in spring and summer than in autumn and winter. These gill parasites caused no serious injuries but were thought to be important in respect of monitoring infestation levels on fish farms in Khuzestan (www.avz1.8m.com/fulltext.htm, downloaded 28 October 2002). Papahn et al. (2004) record the monogeneans Dactylogyrus pavlovskyi, D. barbioides, D. anchoratus, D. carassobarbi and Dogielius persicus from this species in the Karun River at Ahvaz.

Economic importance

This species is second in importance after sobour (Tenualosa ilisha) at the Basrah fish market in Iraq with a weight from October 1975 to June 1977 of 772,775 kg. Nasir et al. (1989) record a total catch for Iraq of 5000 tonnes per year and Petr (1987) for Iraq in 1976 a catch of 4243 t. Young (1976) noted that this species was regarded as the tastiest fish available from the marshes of Iraq.

Petr (1987) has suggested investigating fish farming of this species in Khuzestan and Al-Nasih (1992) carried out such an investigation for Iraq (see below). The Khuzestan Fisheries Research Centre at Ahvaz has successfully bred this species in pond and pen culture using hormone stimulation of broodstocks (Emadi, 1993a; Iranian Fisheries Research and Training Organization pamphlet; Iranian Fisheries Research and Training Organization Newsletter, 5:2, 1994; Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 49, 1996) and in polyculture with Chinese carps such as Ctenopharyngodon idella (Annual Bulletin 1993-94, Iranian Fisheries Research and Training Organization, Tehran, p. 93-94, 1995; Annual Report, 1995-1996, Iranian Fisheries Research and Training Organization, Tehran, p. 36-37, 1997). Private companies also culture this species in Khuzestan. In Khuzestan, over 95% of young survived, using hormones to stimulate reproduction. Mortezazadeh et al. (2009) studied the effects of propofol as an anaesthetic on cultured fish.

Yazdipour et al. (1991) give a report on propagation of this species in Iran.

Sharifian (2000) gives details of whole body analysis. The highest protein content was in the 30-95 mm and 100-140 mm length groups.

Conservation

Local fishermen in Khuzestan believe numbers of this species declined in the Shadegan marshes after young Hypophthalmichthys molitrix from the Caspian were released. As a food fish, its population biology should be monitored in Khuzestan. Several hundred thousand juveniles have been introduced into the Hawr al Azim in Khuzestan in order to restock and protect this resource (Network of Aquaculture Centres in Asia, downloaded 11 January 2007).

Further work

Al-Nasih (1992) investigated the use of this popular food fish for aquaculture in Iraq. Although its growth rate is slower than in Cyprinus carpio, a popular fish for aquaculture, its plankton feeding makes it adaptable to pond life without competition with Cyprinus carpio, it has tasty flesh, reaches 2 kg, and has a relatively high fecundity. Hormonal injections with hypophysial extract from the more readily available Cyprinus carpio induced breeding in this species. Natural production can be increased to 450-600 kg/ha with the use of mineral fertilizers in ponds to stimulate plankton growth. The biology of this species has been investigated in Khuzestan with a view to aquaculture (Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 6, 1996).

Sources

Type material: See above, Barbus sharpeyi (BM(NH) 1874.4.28:20, BM(NH) 1874.4.28:27, BM(NH) 1875.1.14:16), Barbus faoensis (BM(NH) 1888.5.17:4).

Iranian material:- CMNFI 1979-0087, 1, 228.0 mm standard length, ? (); CMNFI 1991-0154, 277.8 mm standard length, ?; material observed on market stalls in Ahvaz, Khuzestan.

Comparative material:- CMNFI 1987-0017, 146.0-175.4 mm standard length, Iraq, vicinity of Basrah (no other locality data); BM(NH) 1920.3.3:71-75, 17, 58.7-115.0 mm standard length, Iraq, Basrah (30º30'N, 47º47'E); BM(NH) 1920.3.3:76-77, 1, 261.4 mm standard length, Iraq, Basrah (30º30'N, 47º47'E); BM(NH) 1922.5.24:1, 1, 113.5 mm standard length, Iraq, Basrah (30º30'N, 47º47'E); BM(NH) 1973.5.21:195, 1, 185.5 mm standard length, Iraq, Shatt-al-Arab (no other locality data); BM(NH) 1973.5.21:196, 1, 186.1 mm standard length, Iraq, Shatt-al-Arab (no other locality data).

Genus Mylopharyngodon
Peters, 1881

This genus comprises a single species so the characters of of the genus are the same as for the species.

Mylopharyngodon piceus
(Richardson, 1846)

Common names

None.

[black carp, snail carp, black amur, Chinese black carp, Chinese roach, black Chinese roach]

Systematics

This species was originally described from Canton, China.

Key characters

The pharyngeal teeth are strong, grooveless molars in 1-2 series and the overall colour is black. distinguishing it from the similar grass carp, Ctenopharyngodon idella.

Morphology

The mouth is terminal to slightly subterminal with thin lips. Dorsal fin origin slightly in advance of level of pelvic fin origin. Dorsal fin with 3 unbranched and 7-9 branched rays, anal fin with 3 unbranched and 7-10 branched rays, pectoral fin branched rays 16, pelvic fin branched rays 8, and lateral line scales 39-46. Gill rakers short and stout, 14-23. Total vertebrae 36-41. The molar pharyngeal teeth are very strong and in one or two rows, commonly 1,4-4,1 or 4-5. Intestine short, 1-2 times total length. The chromosome number is 2n = 48. Artificial hybrids with Ctenopharyngodon idella, Cyprinus carpio, Hypophthalmichthys molitrix and H. nobilis have been bred.

Meristics on Iranian material from Abbasi (2003): Dorsal fin branched rays 7(2), anal fin branched rays 8(2), lateral line scales 41(1) and 42(1), gill rakers 17(1) and 19(1), and pharyngeal teeth 4-5 (1) and 5-5(1).

Sexual dimorphism

Nuptial tubercles are present on the operculum, interorbitally, pectoral fin rays and on the scales of males.

Colour

The overall colour of the body and fins is black to blackish-brown or blue-grey. Fins are characteristically dark but may be somewhat lighter than the body, a blackish-grey, but darker than fins usually are in cyprinids. Scales have dark edges conferring a cross-hatched appearance. The lower head surface may be whitish and the belly greyish-white.

Size

Reaches 2.0 m and 106.0 kg.

Distribution

The native distribution is in East Asia from Amur River basin to south China and perhaps Vietnam but it has been introduced worldwide in suitable waters as a farm fish. The Food and Agriculture Organization web site "Database on Introductions of Aquatic Species" lists this species as introduced to Iran for aquaculture and research from China in 1992. It is reported from the Karakum Canal and the Kopetdag Resevoir in Turkmenistan (Shakirova and Sukhanova (1994) and Sal'nikov (1995) and may eventually reach the Tedzhen River basin in Iran. It is recorded from the Aras River in Armenia (Gabrielyan, 2001) and may eventually be recorded from the Iranian reach of this river. A large specimen from Gilan in the Caspian Sea basin shown me in Tehran in December 2000 appeared to be this species but was frozen solid so certain key characters could not be examined. The Iranian Fisheries Research and Training Organization Newsletter (27:2, 2001) has an illustration of this specimen which weighed 7.5 kg and was 96 cm long. Abbasi (2003) gives the length of this fish as 97 cm total length and a capture date of October 2000 in a beach seine near Bandar Anzali coastal waters. In March 2001 a fish 80 cm total length and weighing 4.8 kg was caught in the same general area. Shamsi et al. (2009) record it from fish farms in Gilan.

Zoogeography

An exotic species introduced to Iran.

Habitat

The main habitat is floodplain lakes and the lower reaches of rivers, down to about 10 m. It may also be found in canals and reservoirs. Fast flowing water is required for reproduction. Temperature tolerances are similar to the grass carp, Ctenopharyngodon idella. Oxygen levels as low as 2 mg/l are tolerated.

Age and growth

Life span is possibly 20 years or more, with maturity attained at 3-11 years in various habitats. Fish introduced to the Karakum Canal showed a relatively rapid growth attaining 48 cm and 1.47 kg in their first year. At six years old fish were 125 cm and 29 kg.

Food

The principal food item of this species is molluscs, snails and clams, although apparently it does not eat large groups of zebra mussels. A 4-year-old fish can eat 1-2 kg of molluscs per day. The molar pharyngeal teeth crush the mollusc shells, the soft tissues are ingested and the broken shells spat out. Young fish eat zooplankton and aquatic insects and adults may take some crayfishes and other benthic invertebrates.

Reproduction

A stretch of river is required for drifting eggs to mature. before settling in quieter areas of floodplain lakes or the river itself. The length of river varies with temperature and velocity of the particular river. Adults migrate up rivers to spawn. Spawning starts at 26-36ºC, and perhaps as low as 18ºC, when rivers begin to rise in flood in spring and summer. Up to 1,180,000 eggs are produced annually in a single batch, more in hatchery fish where an average of 1.5-2.1 million eggs is cited. Swollen eggs have a diameter of 5.6 mm, swelling 4-5 times during hydration. Eggs hatch in 1-2 days.

Parasites and predators

Shamsi et al. (2009) found Dactylogyrus magnihamotus in this species from fish farms. Various Caspian Sea basin species are reported as predators including Esox lucius, Silurus glanis, Sander lucioperca, Aspius aspius, Perca fluviatilis and Lota lota.

Economic importance

None in Iran as not extensively used. This species is widely used on fish farms elsewhere to control parasite problems occasioned by molluscs acting as hosts for trematodes. Both diploid and triploid forms are used in farming. It is also important as a food fish.

Conservation

This species is an exotic and requires no conservation.

Further work

Additional field work is required to determine distribution and abundance in the wild as it can seriously affect native mollusc populations.

Sources

Iranian material: The specimen noted above.

Nico et al. (2005) summarise the biology of this species on which much of the above is based.

Genus Parabramis
Bleeker, 1865

Parabramis pekinensis
(Basilewsky, 1855)

Reported from the Karakum Canal and Kopetdag Reservoir in Turkmenistan (Shakirova and Sukhanova, 1994; Sal'nikov, 1995) as an exotic from China. May eventually be found in the Tedzhen (= Hari ) River and Caspian Sea basins of Iran. No Iranian record.

Genus Pelecus
Agassiz, 1835

The sabre carp genus contains only a single species found from the Baltic to the Black, Caspian and Aral Sea basins including Iran. The characters of the genus are the same as under the species.

Pelecus cultratus
(Linnaeus, 1758)

Common names

shamshir mahi (= scimitar or swordfish), shamshir mahi ab shirin (= freshwater swordfish), kuli.

[gilincbalig in Azerbaijan; chekhon' in Russian; sichel, ziege, razorfish, sabre carp, sabrefish].

Systematics

Pelecus cultratus was originally described from the Baltic Sea.

Pelecus cultratus kurensis Smirnov, 1943 is the Kura River basin subspecies but Berg (1948-1949) considered that other populations over the range of this species had been insufficiently studied to validate this subspecies.

Key characters

This species is easily recognised by the scaleless keel extending from the throat to the anal fin and the decurved and wavy lateral line.

Morphology

The body is elongate and strongly compressed. The mouth is almost vertical. The pectoral fins are long and curved, used for rapid manoeuvring when swimming normally and folded against the body when swimming rapidly. The lower lobe of the caudal fin is larger than the upper, with more rays and a stiffer ventralmost ray. The lower jaw is hooked in older specimens and has a tubercle which fits into an upper jaw notch. Gill openings are very wide with the branchiostegal membranes attached far forward, under the eye level. Muscles on the back extend forward to reach the anterior eye margin. Dorsal fin unbranched rays 2-3, usually 3, followed by 6-10, usually 7, branched rays, anal fin unbranched rays 2-3, usually 3, and branched rays 23-31, pectoral fin branched rays 13-17, and pelvic fin branched rays 6-8. Lateral line scales 88-120. Scales have extremely fine circuli, a central to subcentral posterior focus and very few posterior radii. Gill rakers number 15-26 (reaching the second or third raker below when appressed) and vertebrae 46-52. Pharyngeal teeth number 2,5-5,2, are narrow and are very strongly hooked at the tip with obviously serrate edges. Variants include 2,5-4,2, 2,4-5,2 and 2,5-5,3. The gut is an elongate s-shape. The chromosome number is 2n=50 (Klinkhardt et al., 1995).

Sexual dimorphism

Unknown.

Colour

The back is greenish and the flanks silvery. Fins are hyaline to grey, although the paired fins and the anal fin can be a bright yellow.

Size

Reaches 60 cm and 3.5 kg but most fish in the Volga-Caspian region are 80-180 g.

Distribution

Found in the drainages of the Baltic, Black, Caspian and Aral seas. This species is reported as rare in the lower Safid River (Derzhavin, 1934) and is found in the Anzali Mordab (Abbasi et al., 1999; Kiabi et al., 1999). Recorded by Abdoli (2000) from the middle to lower Safid River, Anzali Talab and adjacent Caspian coast. Recorded from the Atrak River but not yet from its Iranian reach (Reshetnikov et al. (1997). Also reported from the Karakum Canal and Kopetdag Reservoir of Turkmenistan (Shakirova and Sukhanova, 1994; Sal'nikov, 1995) and so may eventually be found in the Tedzhen (= Hari) River basin of Iran.

Zoogeography

A European and western Asian species with its origins in a Danubian or Sarmatian fauna.

Habitat

This species lives primarily in the sea but is also anadromous and may live permanently in larger tributaries. Jolodar and Abdoli (2004) note two forms in Iran, one resident in fresh water and a migratory form. It inhabits surface waters, aided by fin and mouth modifications for this mode of life (Adamicka, 1984). It migrates to the fresh water of large rivers to spawn. In the sea and larger rivers it occurs in schools.

Age and growth

Sexual maturity in the Volga River is attained at 3-4 years and a minimum length of 20 cm in males, 4-5 years and 22 cm for females. The largest immature males are 24 cm long, females 25 cm and the maximum age of non-spawners of both sexes does not exceed 6 years. The spawning stock is mostly fish 4-10 years old. Males have a slightly smaller maximum size than females and life span is at least 16 years (Sil'chenko, 1976). Spawning takes place first at age 2 in the Kura River of Azerbaijan. Growth is faster than in other populations, the Kura fish being the same size at age 2 as age 3 fish from the Don River of the Black Sea and age 4 fish of the Aral Sea.

Food

Food is taken by the vertical mouth from surface waters and includes insects and spiders. Young fish feed on zooplankton and even adults will do this if crustaceans are abundant. In the sea, various crustaceans are taken and these may be pursued near the bottom. Full grown Pelecus (50-60 cm) will capture fish such as Clupeonella, gobies (Gobiidae), Cyprinidae and even sticklebacks (Gasterosteidae) (Adamicka, 1984).

Reproduction

Spawning takes place in the latter half of May in the Kyubyshev Reservoir of the Volga River at 13.5-14.1°C or as high as 18-22°C. Water depths are 2.0-3.5 m. The main spawning takes place around sunset over a period of 24 days with a peak period of 10-12 days. Fecundity reaches 71,400 eggs. Eggs sink to the bottom but swell to an average diameter of 4.7 mm within an hour of fertilisation. Once swollen, any slight movement of the water will suspend the eggs in the water column. Eggs develop pelagically in floodplains, main rivers, side channels, bays and lakes but all these diverse habitats have a high oxygen content through flowing water or wind mixing. In the Volga flow rates are 0.28-1.1 m/sec on a sand-gravel bottom. Spawning may also take place in brackish water where the eggs float. Spawning in the Kura River of Azerbaijan takes place at the end of April and in May. The larvae are phototropic and active swimmers.

Parasites and predators

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

Economic importance

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in textbooks and for food. The scales contain silvery crystals of guanine which are extracted and used to make essence d'orient for artificial pearls. In the period 1909-1913, the catch in the Volga-Caspian region was more than 14 million fish annually. The flesh is fatty and bony and so this species is best smoked.

Conservation

Lelek (1987) classifies this species as intermediate to rare in Europe but fairly common in the Caspian Sea basin. Kiabi et al. (1999) consider this species to be critically endangered in the south Caspian Sea basin according to IUCN criteria. Criteria include sport fishing, few in numbers, habitat destruction, limited range (less than 25% of water bodies), absent in other water bodies in Iran, and present outside the Caspian Sea basin.

Further work

The distribution and biology of this species in Iranian waters needs study although it is probably too rare to be easily captured in numbers.

Sources

Meristics based partly on Wais (1995).

Iranian material: None available.

Comparative material: CMNFI 1971-0825, 1, 232.0 mm standard length, Czechoslovakia, Lake Ozirna (no other locality data); CMNFI 1987-0220, 2, 96.9-105.7 mm standard length, Rumania, Lake Călăraşi (44º20'N, 27º20'E).

Genus Petroleuciscus
Bogutskaya, 2002

This genus currently comprises 7 species of which 3 are found in Iran. Some species in this genus were formerly placed in the subgenus Squalius of the genus Leuciscus; Squalius is now recognised as a distinct genus too. The genus is distinguished by the small size of adults, the reduced number of vertebrae (modally 34-38 in total, rarely 39 or 40 ?check esfahani), few sensory cephalic pores (7-10 in the supraorbital canal, 12-19 in the infraorbital canal and 12-17 in the preoperculo-mandibular canal), a relatively small supraethmoid-mesethmoid block, narrow infraorbitals, and a deep neurocranium with a normally developed interorbital septum (Bogutskaya, 2002). The genus is found in the basins of the Aegean, Black, Caspian and Aral seas, the upper Tigris River , Lake Orumiyeh, and central and south-central Iran.

Petroleuciscus esfahani
Coad and Bogutskaya, 2010

Petroleuciscus persidis
(Coad, 1981)

Common names

'rus mahi persidis.

[Persian chub].

Systematics

Trewavas (1972) has discussed the generic nomenclature of Pseudophoxinus Bleeker, 1859 in which this species was originally described. It is more closely related to the genus Leuciscus according to N. Bogutskaya (pers.comm., 1994; 1996). Bogutskaya (1996) placed this species in her Leuciscus borysthenicus (Kessler, 1859) group which also includes L. ulanus and deserves subgeneric rank, but later erected a new genus Petroleuciscus.

Saadati (1977) described, but did not name, a new species in the related genus Phoxinellus Heckel, 1843 from the Bid Sorkh River in the Simareh River basin between Sahneh and Kangavar in Kermanshahan (the Bid Sorkh Pass is at 34°26'N, 47°49'E). I was unable to find this species in his collections and no further material has come to light. It had 8-9 dorsal fin branched rays, 8-9 branched anal fin rays, 8 branched pelvic fin rays, 47-56 scales in a complete lateral line, and 20 long and crowded gill rakers.

The type locality of Pseudophoxinus persidis is the "upper Shur River drainage at "Koorsiah" village, near Darab on Darab-Fasa road, 28°45.5'N, 54°24'E, Fars". The holotype is a 54.7 mm standard length male held at the Canadian Museum of Nature, Ottawa under CMNFI 1979-0154A (see figure above). Paratypes comprise 95 fish, 34.7-58.8 mm standard length, from the same locality as the holotype under CMNFI 1979-0154B, and 5 fish, 74.7-92.4 mm standard length, under CMNFI 1979-0499 from an "irrigation ditch at village 32 km west of Kor River bridge on road to Dariush Dam, 30°04.5'N, 52°36'E, Fars". Paratypes were distributed to the following institutions from CMNFI 1979-0154B: BM(NH) (2), ROM (2), UAIC (1), UBC (2) and UMMZ (2).

Key characters

This species is characterised by having a pharyngeal tooth count of 1,5-4,1, modally 7 branched dorsal fin rays, anal fin branched rays 7-9, pelvic fin rays 7-8, pored lateral line scales 35-43 in a complete lateral line, short gill rakers numbering 10-14 on the whole first arch, total vertebrae 34-37, a light coloured peritoneum but with numerous melanophores, and flanks with a lateral stripe evident posteriorly but fading anteriorly and not reaching the head.

Morphology

The mouth is terminal and oblique and the rictus reaches back to a level just anterior to the anterior eye margin. A pelvic axillary scale is present. The subcircular to oval scales bear numerous fine circuli and radii on the anterior and posterior fields (total radii number 23-44). Pharyngeal teeth are strongly hooked, concave below the hook and serrated along the margins of the concave surface, particularly on the anterior edge. In the largest fish, the hook may be much reduced to absent, serrations may be absent, and anterior teeth in the main row are rounded. Gill rakers reach the adjacent raker when appressed. The gut is a short and simple s-shape.

Meristic values are as follows: dorsal fin branched rays 6(2) or 7(189) after 3 unbranched rays, anal fin branched rays 7(3), 8(37) or 9(10) after 3 unbranched rays, pectoral fin branched rays 13(10), 14(20) or 15(20), pelvic fin branched rays 7(31) or 8(19), lateral line scales 35(2), 36(19), 37(4), 38(10), 39(3), 40(6), 41(5) or 43(1), total gill rakers 10(1), 11(9), 12(32), 13(6) or 14(2), pharyngeal teeth 1,5-4,2(1), 1,5-4,1(13) or 1,5-4,0(1), and total vertebrae 34(1), 35(20), 36(27) or 37(2). Karyotype 2n=50 (Esmaeili and Piravar, 2006a).

Sexual dimorphism

Number of anal fin branched rays is significantly higher in females (mean 8.3 versus 8.0 in males). Pectoral and pelvic fin lengths, longest dorsal and anal fin rays and caudal peduncle length are shorter in females than in males while head length, head width and predorsal length are longer in females than in males.

Colour

The back and upper flanks are dark and the belly cream. The straight lateral stripe extends from a diffuse area on the tail base to a level at or in front of the dorsal fin origin but does not reach the head. The stripe overlaps the lateral line on the caudal peduncle but lies above it on the flank, paralleling the back. Dorsal fin membranes are lightly speckled with melanophores which tend to be concentrated along the fin ray margins. The caudal and anal fin membranes are mostly clear with pigment restricted to fin ray margins. The pectoral fin may be pigmented on the membranes and there is often strong pigment along the posterior edge of the first unbranched ray. The pelvic fin has little or no pigmentation. Large fish are much darker overall than small fish, obscuring the stripe and with more pigment on fin rays and membranes.

Size

Reaches 9.2 cm standard length.

Distribution

This species is found in the Kor River and Hormuz basins (Coad, 1981e; M. Hafezieh, pers. comm.). Abdoli (2000) reports this species from the Kor and Pulvar rivers, the Hilleh, middle Mand, Shur (tributary of the Dasht-e Palang) and the Kul rivers.

Zoogeography

Durand et al. (2000) place this species in a Leuciscus (= Squalius) cephalus "complex", i.e. descendents of peripheral isolates of a widespread ancestral species, later re-invaded by Danubian S. cephalus.

Habitat

Habitat knowledge is restricted to field data of collections. This species was collected in large rivers, streams, springs, irrigation ditches and qanats, but all these have in common a stream-like environment for much of the year. They are relatively shallow (20 cm to 2 m), variable width (0.5-75 m), medium to slow current, some submergent and emergent aquatic vegetation and a bottom varying from pebbles and gravel to mud. Water temperatures varied from 15 to 23°C from October to January and presumably would be over 30°C in the summer. Conductivity ranged from 0.3 to 1.0 mS. Altitude ranged from 980 to 1940 m. The lower reaches of some of the capture rivers are salty and may not support this species.

Age and growth

Unknown. Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 10 fish measuring 4.21-8.33 cm standard length. The a-value was 0.0178 and the b-value 3.229 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases).

Food

Gut contents include insect remains and fragments of large plants. Diet presumably consists mostly of aquatic invertebrates.

Reproduction

Reproduction in this species is unknown but egg development in adult fish collected in winter and young of the year collected in October suggest spring and early summer as the spawning season.

Parasites and predators

Jalalai et al. (2000) record Dactylogyrus sphyrna, a monogenean, from the gills of this species in the Kor River basin. Barzegar and Jalali (2006) report parasites in this species from Kaftar Lake as Lernaea cyprinacea, Trichodina sp., Ichthyophthirius multifilis and Dactylogyrus sphyrna.

Economic importance

None.

Conservation

The distribution of this species in various small habitats probably means that it is not in immediate danger. However water is in short supply in this part of Iran and abstraction may threaten its survival.

Further work

The ecology of this species should be studied and detailed distributional information compiled.

Sources

Based on Coad (1981e).

Type material: See above, Pseudophoxinus persidis (CMNFI 1979-0154A, 1979-0154B, and 1979-0499).

Iranian material: CMNFI 1979-0025, 11, 19.4-79.6 mm standard length, Fars, Kor River at Marv Dasht (29º51'N, 52º46'30"E); CMNFI 1979-0114, 1, 47.1 mm standard length, Fars, Mand River (29º41'N, 52º06'E); CMNFI 1979-0156, 6, 43.8-64.3 mm standard length, Fars, qanat in Rashidabad (28º47'N, 54º18'E); CMNFI 1979-0163, 8, 31.1-48.3 mm standard length, Fars (no other locality data); CMNFI 1979-0164, 15, 28.8-52.6 mm standard length, Fars (no other locality data); CMNFI 1979-0194, 3, 48.9-58.2 mm standard length, Fars, upper Shur river drainage (28º45'30"N, 54º24'E); CMNFI 1979-0292, 7, 37.0-54.0 mm standard length, Fars, Lapu'i spring near Zarqan (29º48'N, 52º39'E); CMNFI 1979-0305, 2, 23.8-25.5 mm standard length, Fars, Pulvar River at Pasargad (30º12'N, 53º12'E); CMNFI 1979-0342, 33, 23.6-43.2 mm standard length, Fars, Kor River at Band-e Amir (29º46'N, 52º51'E): CMNFI 1979-0503, 4, 32.2-113.5 mm standard length, Fars (no other locality data); USNM 258445, 18, 15.5-44.5 mm standard length, Fars, Chasht Khvar (ca. 29º44'N, ca. 53º12'E).

Petroleuciscus ulanus
(Günther, 1899)

Common names

None.

Systematics

This species was originally described in the genus Leuciscus. Saadati (1977) considers this species to be in the genus Alburnus but did not examine the types. Leuciscus gaderanus Günther, 1899 is a synonym, even Günther (1899) the describer of the two species indicating that this may be the case.

The 2 syntypes of Leuciscus ulanus, 68.0-84.5 mm standard length, are in the Natural History Museum, London and are "from Ula on the Zola Chai", Günther (BM(NH) 1984.10.10:1-2). The 3 syntypes of Leuciscus gaderanus, 44.6-73.0 mm standard length, are "from the Gader Chai" which is near Ocksa (BM(NH) 1899.9.30:113-115). Additionally there are 5 specimens of Leuciscus gaderanus, 11.0-54.5 mm standard length "from near the mouth of the Nazlu Chai at Superghan" (BM(NH) 1899.9.30:108-112). Günther (1899) refers to 3 young specimens from the latter locality as syntypes but of these 5 fish under this catalogue number and locality given as "Superghan" on the label only 2 fish are small (11.0-12.5 mm SL) and the other 3 fish are larger (45.8-54.5 mm SL). Another collection comprising 1 specimen, 27.0 mm standard length, is from the Urmi River according to the label (BM(NH) 1899.9.30:107). Its type status is unclear since the locality is wrong but the size is small and it could be the third of Günther's "three young specimens" since its catalogue number is in sequence.

Key characters

Distribution and meristic characters identify this species.

Morphology

Dorsal fin unbranched rays 3, branched rays 7-9, anal fin unbranched rays 3, branched rays 7-11, pectoral fin branched rays 12-14, and pelvic fin branched rays 7-8. Lateral line scales 36-45. There is a pelvic axillary scale. Scales with a slightly anterior focus and numerous radii on the anterior and posterior fields. Total gill rakers 12-16, reaching the first or second raker below when appressed. Pharyngeal teeth 2,5-4,2, more rarely 2,4-5,2, 2,5-5,2 or 2,4-4,2, hooked at the tip and strongly serrated below on the larger teeth. Total vertebrae 37-38. The mouth is oblique and extends back to just behind the front margin of the eye and the lower jaw protrudes slightly or hardly at all (Günther (1899) has the upper jaw slightly overlapping the lower). The gut is an elongate s-shape and may have an anterior loop to the left.

Meristic values for Iranian specimens are:- dorsal fin branched rays 7(5) or 8(15); anal fin branched rays 7(1), 8(6), 9(11) or 10(2); pectoral fin branched rays 12(5), 13(10), or 14(5); pelvic fin branched rays 7(19) or 8(1); lateral line scales 38(3), 39(6), 40(6), 41(1), 42(1) or 45(2); total gill rakers 12(2), 13(7), 14(6), 15(3) or 16(2); pharyngeal teeth 2,5-4,2(9), 2,4-5,2(2), 2,5-5,2(1) or 2,4-4,2(1); and total vertebrae 37(4) or 38(1).

Sexual dimorphism

Males bear tubercles on the pectoral fins mostly in a single file, occasionally two together, branching with the rays. Small tubercles are also found thickly on the top and sides of the head but no pattern was discernible in the specimens examined (types of L. ulanus).

Colour

There is a narrow, straight black stripe running from the upper half of the eye to the end of the lateral line separating the bluish back from the silvery flanks (Günther, 1899), best developed in preserved specimens posteriorly. Live fish have a grey-silver flank with some spots, a brown-green back and a light silver abdomen. Lower fins are pale to light yellow and the dorsal and caudal fins are light grey (Abbasi and Sabkara, 2004a; 2004b). The lower half of the operculum below the mid-eye level has few or no spots while the upper half is heavily pigmented. There is also no pigment below the eye or it is restricted to a thin line around this lower margin. The flanks are dotted with minute pigment spots. The back mid-line has a black stripe, most obvious predorsally. The rays and membranes of the dorsal fin, caudal fin and anterior pectoral fin bear melanophores; melanophores weak to absent on the anal and pelvic fins. The peritoneum is silvery brown with scattered melanophores.

Size

Reaches 14.2 cm for length (Abbasi and Sabkara, 2004b).

Distribution

This species is endemic to the Lake Orumiyeh basin (Günther, 1899). This species was found only in the Gadarchai and Mahabad River and was not caught in the Zolachai, Nazluchai, Baranduzchai and sampled areas of the Siminerud and Zarrinerud in recent collections by Abbasi and Sabkara (2004a; 2004b).

Zoogeography

Petroleuciscus kurui (Bogutskaya, 1995) from the upper Tigris River system of Turkey is the closest relative of this species (Bogutskaya, 1995) although other members of the Orumiyeh ichthyofauna are related to fishes from the Caspian Sea basin.

Habitat

Essentially unknown, found in stream or river habitats.

Age and growth

Fish caught by Abbasi and Sabkara (2004b) in the Gadarchai and Mahabad River were 34-142 cm in fork length and 1.4 years old. Males comprised 28.8% and females 71.2% of the total population. Maturity was attained at 2 years.

Food

Generally unknown but traces of insects and large quantities of filamentous algae were found in the gut contents of a few specimens. The algae may be an accidental inclusion as the gut is short and probably cannot digest plant material. Fish caught by Abbasi and Sabkara (2004a; 2004b) however contained a wide range of phytoplankton (19 genera) and zooplankton (7 groups), as well as benthic organisms (4 groups). Daphnia, Chydrus and chironomids were dominant gut contents. It was considered to be an omnivorous fish preferring to feed on zooplankton and other mid-water animals.

Reproduction

Spawning of fish in caught by Abbasi and Sabkara (2004b) took place from 1 April until 1 July with absolute fecundity estimated at 1810-16,115, average 6437 eggs.

Parasites and predators

Jalali et al. (2005) summarise the occurrence of Gyrodactylus species in Iran and record G. sp. in fish from the Baranduz and Halaj rivers.

Economic importance

None.

Conservation

This species is known only from the type series and a few other collections. This suggests that it is quite rare or has a restricted habitat preference.

Further work

The biology, distribution and abundance of this poorly known species should be investigated to see if it requires protection.

Sources

Type material: See above for L. ulanus (BM(NH) 1984.10.10:1-2) and L. gaderanus (BM(NH) 1899.9.30:113-115 and BM(NH) 1899.9.30:108-112).

Iranian material: CMNFI 1970-0560, 35,11.4-48.8 mm standard length, Azarbayjan-e Bakhtari, Mamiyand Chay (ca. 36º59'N, ca. 45º39'E); CMNFI 2007-0096, 6, 38.8-68.5 mm standard length, Azarbayjan-e Bakhtari, Baranduz Chay basin (ca. 37º25'N, ca. 45º10'E); BM(NH) 1899.9.30:107, 1, 27.0 mm standard length, Azarbayjan-e Bakhtari, Urmi River (no other locality data).

Genus Pimephales
Rafinesque, 1820

This genus is endemic to North America where there are 4 species in Arctic, Atlantic and Gulf of Mexico drainages. It occurs in Iran as an exotic.

These fishes are small, less than 12 cm standard length, with a stout to slender body. They are uniquely characterised by the last, unbranched dorsal fin ray in males being short, blunt and separated from the succeeding rays by a membrane. There are usually 7 branched dorsal and 6 branched anal rays. The lateral line is complete to incomplete. The head and back before the dorsal fin is flattened and the predorsal scales are small, irregularly arranged and crowded. The intestine varies from long to short, the peritoneum from black to silvery. Pharyngeal teeth are in one row. The flank has a dark band often terminating in a spot at the caudal fin base.

The species of this genus are of some commercial importance as bait or food for sport fishes.

Pimephales promelas
Rafinesque, 1820

Common names

None.

[fathead minnow].

Systematics

This exotic is unrelated to native Iranian fishes and was described from Lexington, Kentucky, U.S.A..

Key characters

This species is unique in Iran in that the last, unbranched dorsal fin ray is short and separated by a membrane from the next, longer ray in males. Other characters include the absence of a barbel, scales in front of the dorsal fin are much smaller and more crowded than flank scales, the back is flattened, the lateral line is incomplete to nearly complete, the mouth is small and terminal, and by the colour and tubercle patterns described below.

Morphology

Dorsal fin branched rays 7, rarely 8, anal branched rays 6, sometimes 7, pectoral rays 14-18 and pelvic rays 8-9. Scales in lateral series 39-56, lateral line pores usually ending before the dorsal fin origin but rarely may be almost complete. Scales are oval with a markedly anterior focus, few circuli and moderate numbers of radii restricted to the posterior field. Vertebrae number 35-38.

Pharyngeal teeth 4-4, slightly hooked at the tip, compressed and with elongate and concave cutting surfaces. The gut is elongate with several loops. Gill rakers 12-16, short and touching the adjacent raker when appressed.

Sexual dimorphism

Large tubercles in males are found on the snout in 3 (rarely 4) rows with 4-15 in the lower row, and with up to 11 tubercles on the lower jaws. There are also smaller tubercles on top of the head and pectoral rays. Tubercles first develop in early April in Canada and males begin to lose them in late July. By early September only scars remain. Males also develop a blue-black to grey, spongy, wrinkled pad on the back between the head and dorsal fin. Fin membranes swell. Breeding males darken, particularly the head and dorsal fin, and the stripe is not apparent. The body can be completely black with a white band at the head-body region and under the dorsal fin. This occurs only during aggression or sexual activity. The lateral banding enhances the robust appearance of fish which must maintain a territory over several weeks without much opportunity to feed. Weight loss is replaced by water to help maintain the image of a fat and vigorous male. Very frightened fishes blanch.

Females have a protruding ovipositor.

Colour

The back and upper flank are dark olive-green to brown, silvery to golden flanks and a whitish belly. There is a mid-flank stripe and a predorsal stripe. Scales on the upper flank are outlined by pigment. A spot at the tail base is faint to absent. Peritoneum black.

Size

Reaches 10.2 cm total length but usually much less.

Distribution

This species was reported from a reservoir, probably the Yengi Kand where bass (Micropterus salmoides) and bluegills (Lepomis macrochirus) were introduced, south of Tehran at 75 km from Asia-beg (Andersskog, 1970) (presumably Asia Bak at 35°19'N, 50°30'E on the Tehran-Esfahan highway). The MMTT catalogue also has a record for the "Dusadj Reservoir", 90 km west of Saveh, presumably in Markazi Province and the Namak Lake basin too. These reservoirs may be the same locality as "Dusadj" or Duzaj, Yang-e Kand and Yanguikand are all villages in the same general area west of Saveh. The reservoir is deduced from maps to be at about 35°19'N, 49°55'E.

A report for the Golpayegan Reservoir (Golpayegan is at 33°27'N, 50°18'E) in the Namak basin by Saadati (1977) was an error (R. J. Behnke, in litt., 1979).

? Map

Zoogeography

The fathead minnow is native to North America and was introduced to Iran accidentally in shipments of largemouth bass, Micropterus salmoides, and bluegills, Lepomis macrochirus. Fathead minnows are pond-raised in North America as food for bass.

Habitat

Fathead minnows are found in ponds, small lakes and slow-flowing brooks, often associated with vegetation. They tolerate high turbidity, high temperatures, low oxygen and high alkalinity and so are able to survive in desiccating or other conditions unfavourable to most fishes.

Age and growth

Males grow faster and larger than females, typical of nest defending species. Life span is about 3 years with maturity attained as early as 1 year, rarely in the year of birth.

Food

Food is bottom sediment for its organic content including plant material, aquatic insects and zooplankton.

Reproduction

Spawning runs from April to August in North America, once water temperatures reach 14°C and light-dark hours are 16-8. Males choose a spawning site under a log, rock, plant stems or even a lily pad or any solid artificial structure in shallow water. Cavity spawning in mud-bottomed habitats prevents the eggs from being smothered as well as offering protection from predators for the relatively few eggs spawned from a small fish. The male will clean out the cavity, spending up to 10 hours on the task, using his tubercles to scrape, pulling debris with his mouth and sweeping with his tail fin. The spongy pad on the back may serve to test spawning sites and eggs chemically. The pad secretes a mucus which is smeared on the spawning site perhaps to improve it for egg survival since mucus protects against disease and parasites. Diseased eggs are eaten by the male. The mucus may also serve to indicate ownership of a nest site. Spawning male fatheads lose their ability to produce alarm or fright chemicals on skin injury, otherwise the continual pad rubbing would disturb spawning activities by releasing alarm substance and scaring away females.

Females may enter the spawning site casually, be chased there by a male or enticed by a face to face encounter and leading to the nest. The male lifts and presses the female on her side between himself and the roof of the spawning site and the female rapidly undulates through an s-shape. Egg deposition is thought to use the same unusual mechanism as in the bluntnose minnow (Pimephales notatus (Rafinesque, 1820)). As each egg is extruded, it is transferred to the upper side of the female and the undulation rolls it along between her side and the nest cavity roof. When the adhesive egg reaches the tail she presses it against the roof where it sticks. How the female transfers the egg from her papilla to her side is unknown and the question remains why this sideways process is used when other roof spawners simply turn upside down and deposit eggs directly. The process may serve to roll eggs into a vacant roof space, occupying the roof most efficiently. Eggs laid on top of others may not attach well or prevent proper development of the underlying eggs.

Males court females by swimming rapidly up to them and then freezing at 3-5 cm away, and by leading females with a zig-zag or straight-line motion from the female to the nest site. Males will also display to females by erecting their fins for 2-3 seconds and by jump-swims in which a male swims upwards to a female then rolls on his side and swims back down. Butting and lateral quivering also occur, perhaps attempts to assess spawning condition of the female. Males defend the eggs against other fishes, including female fatheads, by using the snout tubercles to butt and tail swipes to intimidate by sending a pressure wave sensed by the lateral line system. Chasing and biting are common and 2 males may carousel (or circle head-to-tail) trying to contact each other. Leeches and turtles are also driven away. Some eggs are lost while the male is distracted chasing away predators. Repeat spawning may be necessary to replace lost eggs. Males also aerate and clean the eggs with fin movements. A nest will contain eggs in various stages of development as the male will spawn with several females. Females will deposit eggs in several nests. Orange, mature eggs are up to 1.6 mm in diameter with 12,000 or more per nest. A female will release up to 10,164 eggs in a season but from 9 to 1136 at a time. Spawning intervals are 2 to 16 days.

Parasites and predators

It is an important food for many other fishes and aquatic birds.

Economic importance

In the U.S.A. fatheads are raised as bait fish and as forage fish for introduction into bass fishing lakes. They are also used extensively as a laboratory animal for tests of toxic compounds. They have even been used to evaluate the biological effects of materials from the moon.

Conservation

This exotic species may have deleterious effects on the native fishes and its conservation is of no utility. It is extremely fecund and could compete with native species for food and habitat. A variety of parasites have been reported from this species including some which can have devastating effects on fish populations such as Ligula intestinalis. The introduction of this fish into Europe led to an outbreak of enteric redmouth disease (Yersinia ruckeri) which spread to native and farmed stocks of commercially important fishes such as trout (Michel et al., 1986; Welcomme, 1988). Native fishes may be less well adapted to withstand the depredations of exotic diseases and parasites. Ideally it should be extirpated but, if this is not possible, its interactions with native fishes should be studied and attempts made to prevent its spread to other water bodies.

Further work

The status and numbers of this exotic should be checked by field work. Any data on this exotic species held in files of the Department of the Environment should be published so that this "grey" literature is not lost to future students of Iranian fishes.

Sources

Based on North American literature summaries such as Scott and Crossman (1973) and Becker (1983) as well as personal observations. The biology of Iranian populations is unknown.

Genus Pseudogobio
Bleeker, 1859

Pseudogobio rivularis
(Basilewsky, 1855)

Reported from the Tedzhen River in Turkmenistan on the northeastern border of Iran by Aliev et al. (1988) in their text but not in their distributional table. This exotic species is from China. Also recorded from the Karakum Canal and Kopetdag Reservoir (Shakirova and Sukhanova, 1994; Sal'nikov, 1995) and may eventually reach the Caspian Sea basin. No Iranian record.

Genus Pseudorasbora
Bleeker, 1859

This genus contains 3 species with a native distribution in eastern Asia including the Amur River basin shared between Russia and China, in Japan, other parts of China, and in Korea. One species is an exotic now found in Europe and accidentally introduced in Iran.

The genus is characterised by a small and transverse mouth positioned at the top of the snout rather than the anterior tip, the lower jaw has a trenchant edge and projects slightly beyond the upper jaw, no barbels, pharyngeal teeth are in a single row, the gut is short, scales are large, gill rakers are rudimentary, dorsal and anal fins are short and spineless, and there is no keel on the abdomen.

Pseudorasbora parva
(Temminck and Schlegel in Siebold, 1842)

Common names

آمورچه (amurcheh or amorcheh), amoornama, parva.

[stone moroko, topmouth gudgeon, topmouth minnow, false rasbora; chebachek or Amur chebachok in Russian].

Systematics

Leuciscus parvus was originally described from Nagasaki, Japan.

Subspecies have been described in China but exotic introductions are usually referred to the type subspecies. Reshetnikov et al. (1997) and Bănărescu in Bănărescu (1999) give the date for this species as 1846. The phenotype of this species is highly influenced by environmental conditions and this is suggested to be one of the attributes that make this fish a successful invasive species (Záhorská et al., 2009).

Key characters

The mouth structure is unique, being very small and lying entirely before the nostril level, almost vertical, opening antero-dorsally with the gape entirely visible in dorsal view. The lower jaw protrudes to form the most anterior part of the head.

Morphology

Dorsal fin unbranched rays 3 followed by 7-8, usually 7, branched rays, anal fin with 2-3, usually 3, unbranched rays and 5-7 branched rays, usually 6, pectoral fin branched rays 11-14, and pelvic fin branched rays 6-8, usually 7. Lateral line scales 30-40, with the lateral line rarely incomplete. A pelvic axillary scale is present. The scale radii are restricted to the posterior field. Gill rakers are rudimentary and are only well-developed at the junction of the upper and lower arms of the gill arch. These rakers are stubby and rounded, reaching the adjacent raker when appressed, and bearing fine, fleshy fimbriae which extend onto the adjacent parts of the gill arch. Anterior rakers are absent and patches of fimbriae are found. Rakers number 6-16, usually 9-13. Total vertebrae 31-38 presumably the result of different counting methods; Naseka (1996) gives 36-38 and fish from Turkey have 34-37, cf. below. Pharyngeal teeth usually 5-5, rarely 6-5, with the tips strongly hooked and the area below the hook flattened and without ridges or only very weakly ridged on some teeth. The gut is an elongate s-shape. The chromosome number is 2n=50 (Klinkhardt et al., 1995).

Meristic values for Iranian specimens are:- dorsal fin branched rays 7(11) or 8(1), anal fin branched rays 6(12), pectoral fin branched rays 13(7) or 14(5), pelvic fin branched rays 7(4) or 8(8), lateral line scales 34(4), 35(5) or 36(3), pharyngeal teeth 5-5(9), 6-5(2) or 4-4(1), and total vertebrae 34(2) or 35(10).

Sexual dimorphism

A horny pad develops on the jaws in males and females during spawning and strong, sharp tubercles in males. One tubercle is found between the eye and the nostril, one below the nostril (this may be absent), one next to the upper lip on a line across from the one below the nostril, 5-8 in a row from the extreme corner of the mouth along the side of the head over the flesh of the cheek, and 2-3 below the lower lip from the tip of the lower jaw to the end of the jaw on the lower head surface. Lower head surface tubercles may coalesce at the base but each tubercle bears a single rounded cusp. Rarely a tubercle may have a single base but two cusps.

Males are larger than females and have larger fins. Spawning males are darker than females and the flank has a metallic violet sheen.

Colour

The head and body has a mid-lateral stripe but this is obscured in adults by crescentic speckles situated posteriorly on each scale. The back is light grey, the flanks silvery and the belly whitish. Dorsal and anal fins are speckled and turn almost black in spawning fish.

Preserved fish have a cream coloured belly with the back much darker. The head is black dorsally and fades to cream ventrally. The scales on the back and flanks, but not the belly, carry a broad band of pigment which follows the scale margin distally. The extreme edge of the scale is hyaline but the arc of pigment effectively defines the posterior scale margin and outlines the scale pattern of the back and flanks. Pigmentation on fins is mostly restricted to the rays and their margins but is found also on fin membranes to varying extents. Pigmentation is strongest distally on all fins. The dorsal fin, particularly in smaller fish, bears patches on the membranes posterior to branched rays 1 or 2 through 5 or 6, starting on ray 1 or 2 below the mid-point of the ray length and descending gradually behind successive rays to lie near the base behind the last ray. These patches are vertically short and do not touch the succeeding ray. The leading edge of the dorsal, anal, pectoral and pelvic fins and the upper and lower edges of the caudal fin are black in large, and some small, fish. Pigment may be concentrated along the mid-line forming a thin stripe, only apparent posteriorly in some fish. There is a dark line along the mid-line of the back. The peritoneum is silvery with some scattered melanophores.

Size

Attains 12.0 cm (Movchan and Kozlov, 1978).

Distribution

The natural range of this species is in eastern Asia as given above under the genus. It has been introduced to Iran by accident (Abdoli, 1992; Coad and Abdoli, 1993b). It is now found in ab-bandans at Avaness, Hasan Tabeeb and Shaeed Ziaee (all about 40-45 km east of Gorgan), Teer Tash and Lemrask (about 20-25 km east of Behshahr), Lapoo (about 4 km east of Babol Sar) on the Caspian Sea coast, and at Gorgan-Aliabad, Mazandaran, from the Safid River estuary and neighbouring waters and the Anzali Talab (Iranian Fisheries Research and Training Organization Newsletter, 6:8, 1994; Anonymous, 1994; Abbasi et al., 1999), from the Atrak, Gorgan, Gharasu, Tajan, Babol, Haraz, Sardab, Pol-e Rud, and Safid rivers (Kiabi et al., 1999), from fish ponds at Arak probably inadvertently carried there with carp fingerlings imported from Gilan on the Caspian shore; at Mashhad in northeastern Iran, in springs near Kermanshah and in Lake Zeribar, Kordestan (Coad, 1996g); in Sistan at Hamun Kushk, and the canal flowing into the Chahnimeh (J. Holčík, in litt., 1996), and from the International Wetlands of Alma-Gol, Adji-Gol and Ala-Gol (Patimar et al., 2002a; 2002b; Patimar, 2008).

Abdoli (2000) records it generally from the Dasht-e Kavir, Dasht-e Lut, Kerman-Na'in and Sistan basins; the lower Kashaf River in the Tedzhen River basin; the middle Atrek, lower Neka, Babol, Heraz, Chalus, Tonekabon, and Safid rivers and the Anzali Talab in the Caspian Sea basin; the middle to lower Talkheh and lower Zarrineh rivers in the Orumiyeh basin; the lower Shur and middle and lower Qareh Chai in the Namak Lake basin; the middle and lower Zayandeh River in the Esfahan basin; and the Simarreh and lower Gav Masiab rivers in the Tigris River basin. Jolodar and Abdoli (2004) record it from most water bodies on the Iranian Caspian coast. Esmaeili et al. (2011?) add the Gulf and Lake Maharlu basins and it is now probably distributed throughout Iran.

It is also recorded in the Karakum Canal, Kopetdag Reservoir and Tedzhen River of Turkmenistan (Shakirova and Sukhanova, 1994; Sal'nikov, 1995) so may well reach the Tedzhen (= Hari) River basin of Iran eventually. Pipoyan (1996a) reports it from the Araks River in Armenia.

Zoogeography

This species was first recorded in western Eurasia in Romania in 1960 as an accidental introduction with Chinese carps from the lower Yangtze River of China. The species is now widespread in Europe and is becoming common in western Asia including Kazakhstan, Uzbekistan, Kyrgyzstan and southern Anatolian Turkey as well as Iran (Wildekamp et al., 1997).

Habitat

This species prefers well-vegetated areas as protection from predators. It may be found in streams, rivers and ponds, and more rarely in the shallows of large lakes. It is apparently quite resistant to pollution (Bănărescu in Bănărescu, 1999) and is found in waters that freeze over and that attain 30ºC in summer (Boltachev et al., 2006).

Age and growth

Life span is about 5 years with maturity attained at 1-2 years, usually at 1 year in Europe or the second year of life in the Crimea (Boltachev et al., 2006). Most fish in a population are 2-3 years old or 1-2 years in the Crimea. Patimar et al. (2002a; 2002b) report 4 age groups from the International Wetlands of Alma-Gol, Adji-Gol and Ala-Gol, with the smallest mature specimens found at 2 years. Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 33 Iranian fish measuring 3.29-5.99 cm standard length. The a-value was 0.0286 and the b-value 2.763 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases).

Food

This species feeds on benthos but also some zooplankton. Food items include various aquatic insects such as stone flies, caddis flies, chironomids, water sawbugs and midge larvae but guts also contain sponges, bryozoans, Spirogyra, detritus, and fragments of higher plants (Movchan and Kozlov, 1978). It may also feed on the eggs and juveniles of native fishes. Young fish take zooplankton (Movchan and Kozlov, 1978). Bănărescu in Bănărescu (1999) reports also isopods and aquatic worms and, in fish ponds, artificial food.

Reproduction

Reproduction begins at 16-18°C and lasts two months in its native Amur River basin. Fecundity is about 5000 elliptical eggs with a diameter of 2.0-2.5 mm, and this species has intermittent spawning with up to 85 eggs per batch in introduced populations in Central Asia (Makeyeva and Mokhamed, 1982). Up to 60 batches may be laid in a spawning season. The spawning site is cleaned of ooze and plant material. Adhesive eggs are deposited on the lower surface of stones, and occasionally sticks or empty mollusc shells, and are protected by the male using the head tubercles to drive away other fishes. The ellipsoidal eggs are laid in strips, usually of 5 eggs but as many as 10. Males clean the eggs and remove dead ones. The spawning season in Central Asia is April to August and spawning takes place in warm, shallow and calm waters in the morning. Spawning in the Crimea is in second half of May or in June, late May to July in the Ukraine and from the end of June to the beginning of August in the Amur (Boltachev et al., 2006). Female specimens from Iran collected in March, April and May are ripe and males have well-developed breeding tubercles and Patimar et al. (2002a; 2002b) report a spawning peak in April in the International Wetlands of Alma-Gol, Adji-Gol and Ala-Gol of Iran.

Parasites and predators

Malek and Mobedi (2001) report Clinostomum complanatum from this species in Mazandaran, in the Shiroud. Sander lucioperca and Silurus glanis are predators in Turkmenistan (Aliev et al. (1988).

Economic importance

Makeyeva and Mokhamed (1982) and Movchan and Kozlov (1978) report competition with commercial species and predatory behaviour on carp larvae if there is insufficient food. Male reproductive aggression may inhibit breeding of native fishes. Boltachev et al. (2006) report that it is a facultative parasite of other fishes in enclosed areas including commercial species such as the silver carp (Hypopthalmichthys molitrix). Areas of the body are attacked such as behind the dorsal fin and above the anal fin out of sight of the affected fish. Skin and muscles are eaten away. Welcomme in Courtenay and Stauffer (1984) regards this species as a pest when introduced. Bănărescu in Bănărescu (1999) reports this species as a competitor for food with native species in Europe.

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

Conservation

None required for an introduced species.

Further work

The distribution of his species as an exotic in Iran should be thoroughly documented and its biology and effects on native species studied.

Sources

Iranian material: CMNFI 1991-0160, 3, 47.8-55.8 mm standard length, Mazandaran, Abgeere Avanes (37º03'N, 54º47'E); CMNFI 1991-0161, 1, 52.5 mm standard length, Mazandaran, Madarso River (37º23'N, 55º47'E); CMNFI 1993-0134, 7, 45.0-59.1 mm standard length, Mazandaran, Gorgan-Aliabad (37º01'30"N, 54º47'36"E); uncatalogued material, 1, 86.9 mm standard length, Kermanshahan, sarabs near Kermanshah (no other locality data).

Comparative material: CMNFI 1983-0204, 7, 51.7-76.6 mm standard length, Turkey, Edirne, Meriç River at Ipsala (40º55'N, 26º23'E); CMNFI 1983-0343, 5, 49.9-83.4 mm standard length, same locality as preceding.

Genus Rhodeus
Agassiz, 1832

The bitterlings comprise about 18 species in Europe, Asia Minor, the Caspian Sea basin, China, Japan and Korea with 1 species in Iran.

They are small fishes with deep, compressed bodies, an incomplete lateral line (about 10 pored scales or less), large to moderate-sized scales, females with an ovipositor, males larger than females (unusual in cyprinid fishes), brightly coloured and tuberculate when spawning, pharyngeal teeth in 1 row and not or only slightly serrated, mouth small, oblique and subterminal, no barbels, dorsal fin short to moderately long and spineless, anal fin of similar length, gill rakers short, intestine long and spirally coiled, and peritoneum black.

Rhodeus amarus
(Bloch, 1782)

Common names

ماهي مخرج لوله اي (= mahi-ye makhraj lulehi or mahi-e-makhraj looleei, meaning tube-like vent fish), rodeus.

[karka in Azerbaijan; gorchak in Russian; bitterling, European bitterling].

Systematics

Holčík and Duyvené de Wit (1964) reviewed the systematic status of Rhodeus sericeus (Pallas, 1776) in Europe and western Asia (but not Iran) in comparison to the Chinese populations of this species. They refer European and western Asian populations to Rhodeus sericeus amarus (Bloch, 1782) and Iranian populations were long regarded as this subspecies. Cyprinus sericeus was described from Dauriya; Cyprinus amarus was described from the Elbe basin, Germany and no types are known (Eschmeyer et al., 1996). Later, Holčík and Jedlička (1994) consider subspecies not to be warranted as the key characters used in distinguishing them (pored lateral line scales, scales in lateral series, gill rakers) showed clinal variation with longitude and the number of segments was also related to latitude, elevation, mean annual air temperature, and fish size. Bohlen et al. (2006) consider Rhodeus from the Vistula to the Volga to belong to the R. amarus eastern clade based on cytochrome b sequences.

Kottelat (1997) considers Rhodeus amarus to be a distinct species since it is diagnosable (although differences with Rhodeus sericeus are slight and largely overlap), and it is separated by 2-4 million years in time and 4000 km in space. However he does admit that the immense distributional gap may be a source of bias. Holčík in Bănărescu (1999) considers amarus to be synonymous with sericeus. Bogutskaya and Komlev (2001) found no characters to clearly confirm the specific status of R. amarus. The name amarus is retained here as an indication that the taxa are geographically isolated. Van Damme et al. (2007) refer the Caspian bitterling to a species as yet undescribed.

This species was described from Lake Müggelsee near Köpenik, Berlin, Germany and 3 syntypes are in the Museum für Naturkunde, Humboldt-Universität zu Berlin (or Zoologisches Museum Berlin, ZMB 3393).

Key characters

The presence of an ovipositor in females, the flank stripe and few pored scales are distinctive.

Morphology

Dorsal fin with 2-4, usually 3, unbranched and 7-11, usually 9, branched rays, anal fin with 2-4, usually 3, unbranched and 6-12, usually 9, branched rays, pectoral fin branched rays 10-13, and pelvic fin branched rays 4-8, usually 6-7. Pored lateral line scales 0-11, usually 4-6, along the flank 28-45, usually about 30-32 in some reports but 32-38 in Abdurakhmanov (1962), 37-40 in Holčík and Jedlička (1994), 30-38 in Pipoyan (1996b) and 35-40 in Holčík in Bănărescu (1999). Gill rakers 9-16, usually 10-14, reaching the raker below when appressed. Vertebrae 33-39. Pharyngeal teeth 5-5, rarely 5-4, 4-5 or 4-4. Teeth are elongate and narrowly compressed with a slight to strong hook at the tip and a very long, narrow and concave grinding surface below the tip. The gut has numerous coils. The chromosome number is 2n=48 (Klinkhardt et al., 1995).

Meristic values for Iranian specimens are:- dorsal fin branched rays 8(4), 9(52), 10(1) or 11(1); anal fin branched rays 8(18), 9(39) or 10(1); pectoral fin branched rays 10(3), 11(24), 12(29) or 13(2); pelvic fin branched rays 4(1), 6(3), 7(53) or 8(1); scales in lateral series 32(1), 33(2), 34(22), 35(29) or 36(4); pored lateral line scales 3(1), 4(7), 5(37), 6(11) or 7(2); total gill rakers 10(3), 11(14), 12(27), 13(10) or 14(4); pharyngeal teeth 5-5(20); and total vertebrae 34(9), 35(36), 36(12) or 38(1). Holčík and Jedlička (1994) give ranges in their Iranian sample of 33-41 for lateral series scales, 0-6 pored scales and 9-14 for gill rakers, in general agreement with data here.

Sexual dimorphism

Males develop a triangular or crescent-shaped patch of 7-20 tubercles on each side of the snout and small tubercles are found above the eyes. The female develops an ovipositor near the genital opening and it may be longer than the body. The flank stripe is longer and about one half wider in males compared to females, and male colouration is distinctive. Males are generally larger than females.

Colour

Males are particularly colourful in the spawning season: the top of the head and back are olive to bright green, reddish or dark violet, the iris is bright red, flanks are iridescent with violet and steel-blue colours most evident, the throat and belly are orange to blood-red, dorsal and anal fins are bright red and margined with black, the caudal fin is green at the base and yellow distally, and pectoral and pelvic fins are yellowish. Females are more yellowish and less iridescent than males in the spawning season. Outside the breeding season both sexes are similar in colour with a grey-green back, silvery flanks and yellowish belly. A grey-green to greenish-blue stripe originates under the dorsal fin and extends back to the tail base, broadening posteriorly. The dorsal fin is blackish and other fins reddish to yellowish. The dorsal, and sometimes the anal fin, has a dark interrupted stripe. The iris is silvery or yellowish. The peritoneum is dark.

Size

Reaches a reputed 18.0 cm but usually not more than 7.0-9.0 cm.

Distribution

Found from western Europe north of the Pyrenees and Alps to the Caspian Sea basin. In Iran, it is recorded from Astara to the Gorgan River including the Anzali Talab (Derzhavin, 1934; Holčík and Oláh, 1992; Abbasi et al., 1999; Kiabi et al., 1999; Abdoli, 2000). Also probably in the Araks River of Armenia (Pipoyan, 1996b). They may have been introduced into the Zarrineh River of Azarbaijan-e Gharbi based on photographs by Saber Shiri (pers. comm., 14 June 2008).

Zoogeography

Bohlen et al. (2006) assumed a continuous distribution of Rhodeus from Europe through Siberia to east Asia during the Pliocene, loss of the Siberian population in the late Pliocene or early Pleistocene, subsequent isolation, and later post-glacial expansion from several refugia in the Euro-mediterranean zoogeographic subregion. The Caspian Sea populations were not examined but seem to belong to R. amarus although the authors suggest a revisionary study is required for European bitterlings. Van Damme et al. (2007) consider this species was restricted to Ponto-Caspian and Aegean regions and its presence in western and central Europe is associated with the spread of Cyprinus carpio cultivation, and more recently, anthropogenic alterations to habitats and temperature changes.

Habitat

This species favours heavily vegetated areas of small lakes, ponds and slow-moving rivers, rarely in faster water. It is found in the lower reaches of rivers on the Caspian coast of Iran (Jolodar and Abdoli, 2004). The bottom is usually a fine sand or a thin layer of mud. Swan mussels (Anodonta) and freshwater clams (Unio) share this kind of habitat and are necessary for reproduction. Other genera of clams include Pseudanodonta, Cristaria, Margaritifera and Dahurinaia (Smith et al., 2004). Spawning occurs at water temperatures of 12-24°C, although 15-21°C is optimal (Holčík in Bănărescu, 1999), Van Damme et al. (2007) giving 23°C as optimal. Its distribution is limited by the 16°C July isotherm (Van Damme et al., 2007).

Age and growth

Maximum life span is 5-8 years, the higher values being uncertain (Holčík in Bănărescu, 1999). Maturity is attained during the second or third year, the life span of most individuals. Rarely some fish may mature as early as before the age of 1 year or as late as the fourth year. Growth is faster in ponds than in rivers. Males outnumber females at a ratio of 1.2-1.5:1, especially in spawning areas, although females apparently outlive males (Holčík in Bănărescu, 1999).

Food

Food consists of diatoms and detritus with aquatic insects and crustaceans being mostly accidental inclusions. They also take eggs of Rutilus rutilus, Cyprinus carpio and Rutilus frisii, and also their own eggs which fail to be deposited in the clam or mussel. Most gut contents in Iranian fish examined were plant fragments, filamentous algae, detritus and sand grains.

Reproduction

Reproduction takes place mainly in April and May, but can run from the end of February (in Azerbaijan) to August. The female develops an ovipositor from the genital opening, up to 6 cm long. In Iranian fish the best developed ovipositor was seen in a fish collected on 12 May when it extended back two-thirds along the caudal fin length. Eggs in this fish were 2.2 mm. Fish taken in September, October, November and January had short ovipositors progressively increasing in length with time. A fish taken on 4 July had an ovipositor extending only half way along the anal fin but no large eggs. Spring spawning is indicated but an ovipositor can be seen in varying degrees of development year round in adult females. Holčík in Bănărescu (1999) reports ovipositor length up to 126.5% of standard length in the Anzali Mordab in April.

The ovipositor lays eggs inside freshwater clams and mussels, using the excurrent siphon as the entry route. Apparently the flow of water and a high oxygen content out of this siphon encourage egg laying. Before egg laying, the female nudges the clam repeatedly to accustom the mollusc to stimulus so that it does not close up its shell. In the absence of clams, the bitterling does not become sexually mature. Clams and mussels with high numbers of bitterling larvae or filled with glochidia are avoided, as is Anodonta cygnaea, a mussel which is able to eject eggs and larvae and has low oxygen levels in its excurrent siphon (Kottelat and Freyhof, 2007). Males select and defend a particular clam against other males. They may headbutt each other or strike flanks, dislodging scales. Sneaking occurs despite this. The female deposits eggs 1-2 at a time and the male sheds sperm which are sucked into the clam on its feeding current. This process may be repeated with the same or different females. The female deposits about 40-100 eggs at each spawning and can spawn at least 5 times a day. The movement of the ovoid eggs through the ovipositor can be clearly seen as they are somewhat larger than the distensible ovipositor. Spawning bouts last 1-3 days with intervals of 5-7 days, variable with feeding conditions and temperature (Smith et al., 2004). Fecundity is up to 22,136 eggs and maximum egg diameter to 3.1 mm with width up to 1.52 mm. Eggs hatch 2-5 weeks later and the young leave the clam after 2 days when the yolk sac has been absorbed. Young fish leave the clam singly or in pairs. The fish gain the advantage that the eggs and young are protected inside the clam, even should the shore area dry out since the clam will move to deeper water. The clam is unharmed and is able to disperse its own young or glochidia by their attachment to the fins of the adult fish. Bitterlings seem however to have an immunological response to glochidia, having far fewer than other fishes that are less intimately associated with clams. Circumstantial evidence laid out by Smith et al. (2004) involving inhibition of free water circulation by the fish embryos, damage to clam gills, and increased consumption of oxygen by clams, indicate the relationships is a parasitic rather than a commensal one.

Smith et al. (2004) give a detailed account of reproduction, behaviour and development in this species.

Parasites and predators

None reported from Iran.

Economic importance

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in textbooks and aquaria and because it has been introduced outside its natural range. The species is used as a model in behavioural studies. Van Damme et al. (2007) regard it as a parasite on freshwater mussel populations in western and central Europe outside its native range, escaping glochidia infection and having low egg ejection rates.

Conservation

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

Further work

Although well-studied elsewhere, the biology of this species in Iran has not been investigated in detail.

Sources

Iranian material: CMNFI 1970-0510, 21, 34.5-47.5 mm standard length, Gilan, Golshan River (37º26'N, 49º40'E); CMNFI 1970-0512, 25, 26.6-43.9 mm standard length, Gilan, Shalman River (37º08'N, 50º15'E); CMNFI 1970-0514, 2, 36.6-38.3 mm standard length, Gilan, Shafa River estuary (37º35'N, 49º09'E); CMNFI 1970-0520, 7, 28.9-45.0 mm standard length, Gilan, Astara River (ca. 38º25'N, ca. 48º52'E); CMNFI 1970-0526, 4, 30.7-43.2 mm standard length, Gilan, Safid River below Astaneh Bridge (37º19'N, 49º57'30"E); CMNFI 1970-0579, 8, 32.6-49.0 mm standard length, Gilan, Old Safid River estuary (37º23'N, 50º11'E); CMNFI 1970-0580, 5, 35.8-53.8 mm standard length, Mazandaran, river near Iz Deh (36º36'N, 52º07'E); CMNFI 1979-0265, 9, 16.1-47.4 mm standard length, Gilan, Anzali Mordab at Abkenar (37º28'N, 49º20'E); CMNFI 1979-0472, 6, 37.7-47.7 mm standard length, Mazandaran, stream west of Mahmudabad (36º37'N, 52º12'E); CMNFI 1980-0117, 3, 42.2-47.8 mm standard length, Gilan, Golshan River (37º26'N, 49º40'E); CMNFI 1980-0127,9, 32.6-43.0 mm standard length, Gilan, Caspian Sea near Hasan Kiadeh (37º24'N, 49º58'E).

Rhodeus ocellatus
(Kner, 1866)

Reported from the Karakum Canal and Kopetdag Reservoir in Turkmenistan (Shakirova and Sukhanova, 1994; Sal'nikov, 1995) as an exotic from China. May eventually be found in the Tedzhen (= Hari) River and Caspian Sea basins of Iran. No Iranian record.

Romanogobio
Bănărescu, 1961

There are about 17 species in the genus found from Europe to China with two species recorded from Iran. Members of the genus have a shallower body than the related Gobio, an elongated, usually cylindrical, caudal peduncle, epithelial keels on dorsal scales, an anus placed closer to ventral fins, absolute or average prevalence of caudal vertebrae over abdominal ones, and a higher number of preanal vertebrae (Naseka, 1996). Bănărescu in Bănărescu and Paepke (2002) notes that he considers Romanogobio as a subgenus and further work is needed to resolve this difference of opinion. Most literature is under the genus Gobio.

Romanogobio macropterus
(Kamensky, 1901)

Common names

گاو ماهي (= gav mahi, probably in error for Neogobius and related gobies).

[Kur gumlagcisi in Azerbaijan; Kurinskii peskar' or Kura gudgeon in Russian].

Systematics

Gobio macropterus Kamenskii, 1901 was originally described from the Caucasus. The 14 specimens in the type series were deposited in the Georgian State Museum, Zoological Section, Tbilisi (ZMT) and in Kharkov University, Ukraine. Naseka et al. in Bănărescu (1999) cites types of Gobio macropterus in the Museum of the Caucasus (Tbilisi, presumably the Georgian State Museum) under numbers 128a (1 fish from Alazan) and 129 (4 fish from Kars-tschai) and 1-2 fish from the Kura (catalogue number unknown) in Khar'kov University. See below under R. persus for its changing status.

Key characters

The 7 branched dorsal fin rays are characteristic and this is the only Romanogobio species in the Caspian Sea basin of Iran (see also below under R. persus). It is separated from the related species in Iran (Gobio gobio) by having the body only slightly compressed and the caudal peduncle being cylindrical (caudal peduncle depth at anal fin insertion less than or about equal to caudal peduncle width) and by faint spots on the dorsal and caudal fins.

Morphology

Sexual dimorphism

Colour

Size

Distribution

Found in the Kura and Aras river basins of Armenia, Azerbaijan and Iran. Abdoli (2000) maps the middle Aras River and its lower Qara Chai tributary.

Zoogeography

Habitat

Age and growth

Food

The chitinous remains of aquatic insects comprised 30% of the gut contents from samples in the Kura River basin, caddisflies 21%, mayflies 14%, chironomids 12%, sevryuga eggs 6%, fish scales 3%, and much of the remainder was detritus at 20% (Abdurakhmanov, 1962).

Reproduction

Spawning takes place in May in Azerbaijan (Abdurakhmanov, 1962) and each female may spawn several times in a season. Eggs number up to 15,840 and are up to 1.62 mm in diameter. Water temperatures of 12-18°C are recorded for Georgia in April to June (Naseka et al. in Bănărescu, 1999).

Parasites and predators

Economic importance

Conservation

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

Further work

Sources ?check

Iranian material: CMNFI 1980-0155, 3, 45.9-69.3 mm standard length, Azarbayjan-e Khavari, Qareh Su near Ardebil (ca. 38º15'N, ca. 48º18'E).

Comparative material: CMNFI 1980-0806, 1, 93.4 mm standard length, Turkey, Kars Çayi, Kars (40º37'N, 43º05'E); CMNFI 1986-0007, 4, 49.4-84.0 mm standard length, Turkey, Kars, Kars Çayi north of Kars (41º00'N, 43º00'E).

Romanogobio persus
(Günther, 1899)

Common names

گاو ماهي (= gav mahi, probably in error for Neogobius and related gobies).

[Persian gudgeon].

Systematics

This species was once thought to belong to the genus Rheogobio Bănărescu, 1961 variously regarded as a synonym of Gobio, a subgenus or a distinct genus. P. M. Bănărescu (in litt., 1984) and A. Naseka (pers. comm., 1994; in litt., 1995) place Gobio persus in the subgenus Romanogobius sensu Bănărescu (1961; 1992a) and Naseka (1996) elevates Romanogobio to a genus.

A. Naseka (pers. comm., 1994; in litt., 1995; Naseka et al. in Bănărescu, 1999) has studied R. persus from the Orumiyeh basin and from the Kars, Kura and Aras rivers of Turkey, Georgia and Azerbaijan. He distinguished two subspecies, R. p. persus from the Orumiyeh basin and R. p. macropterus from the Kura River basin which includes the Kars River in Turkey and the Aras River on the border of Iran and Azerbaijan. Gobio macropterus Kamensky, 1901 is now regarded as a distinct species (Naseka and Freyhof, 2004).

The type series of Gobio persa consists of 7 specimens, 53.1-65.9 mm standard length, in the Natural History Museum, London from "Ocksa in the Gader Chai" in the description and "Ockra. NW Persia. Günther" in the jar, the former being more accurate (BM(NH) 1899:30:90-96).

Key characters

The 7 branched dorsal fin rays are characteristic and this is the only Romanogobio species in the Orumiyeh basin. The number of lateral line scales is 40 to 42 with modes of 40 or 41 (41 to 45 with modes of 42 and 43 in R. macropterus), total vertebrae are 37 to 40 with modes of 38 and 39 (38 to 42 with modes of 40 and 41), and the connection between the supraorbital and infraorbital head canals is usually absent (present). The Orumiyeh fish also have a shorter caudal peduncle, a shorter snout, a shorter barbel, and a longer predorsal distance.

It is separated from the related species in Iran (Gobio gobio) by having the body only slightly compressed and the caudal peduncle being cylindrical (caudal peduncle depth at anal fin insertion less than or about equal to caudal peduncle width) and by faint spots on the dorsal and caudal fins.

Morphology

The barbel is broad and fleshy, the mouth subterminal and horseshoe-shaped with thick, papillose lips. The lower head surface between the jaws and the sides of the head are papillose in a mature female.

Dorsal fin with 3, rarely 4, unbranched and 7, rarely 8, branched rays, anal fin with 3, rarely 2, unbranched and 5-7, usually 6, branched rays, pectoral fin branched rays 11-16, and pelvic fin branched rays 6-8. Lateral line scales 39-45. A pelvic axillary scale is present. The throat, breast and anterior belly are naked. Dorsal scales bear epithelial keels, usually one central keel and one to several lateral ones. Scales have a very anterior focus and few posterior radii. Gill rakers 0-6, small and irregularly spaced. Vertebrae 36-42. Pharyngeal teeth 2,5-5,2 or 3,5-5,3, or more rarely 2,5-5,1, 2,5-5,3, 2,4-5,3, 2,4-6,3, and 3,5-5,2. Teeth are strongly hooked at the tip, broadly concave or flattened below the tip, the surface sloping medially. The gut is s-shaped with a slight anterior loop to the left. The anus lies between the pelvic fins, remote from the anal fin origin. The karyotype is 2n=50.

Meristic counts in Iranian fish are: dorsal fin branched rays 7(22); anal fin branched rays 6(21) or 7(1); pectoral fin branched rays 11(1), 12(1), 13(6), 14(7), 15(6) or 16(1); pelvic fin branched rays 7(22); lateral line scales 39(3), 40(7), 41(3), 42(2) or 43(2); pharyngeal teeth 3,5-5,3(5); and total vertebrae 37(1), 38(7), 39(10), 40(4), 42(1).

Sexual dimorphism

Males and females bear irregular-shaped, elongate tubercles on the head. In a female specimen (and presumably males too), the pectoral and pelvic fin rays and adjacent membranes bear tubercles both dorsally and ventrally although the latter are less well developed. Males have longer pectoral and pelvic fins than females, reaching the pelvic fin and anal fin origin respectively in males. The snout to anus distance is more than half body length in females and about equal in males, head width is less than head depth at nape in females and equal in males, and snout length is longer than postorbital distance in females and about equal in males.

Colour

The upper flank is a light yellow-grey to brown with each scale outlined with, or partially filled in with, dark pigment, fading to a yellowish colour below. The mid-flank has 6-12 elongate to rectangular black spots. The lateral line pores may have small dark spots above and below reminiscent of Alburnoides species. The back also bears vague dark spots. Dorsal, caudal and pectoral fins bear up to 5 thin to broad discontinuous bars, the pigment being on the rays. Scales above the lateral line in mature males have longitudinal streaks. Peritoneum silvery with scattered melanophores.

Size

Reaches ?

Distribution

Found in the Lake Orumiyeh basin (Günther, 1899). Abdoli (2000) maps the middle and lower Talkheh, lower Tatavi and Zarrineh rivers in the Lake Orumiyeh basin.

Zoogeography

This species is part of a gobionine fauna found across Eurasia and this wide distribution may be suggestive of further work that could be done to clarify relationships of these fishes.

Habitat

Presumed to be similar to its relative.

Age and growth

Presumed to be similar to its relative.

Food

The principal food is aquatic insects and crustaceans but detritus and vegetation are also taken and presumably, though rarely, the eggs and fry of other fishes. Abdoli (2000) lists Trichoptera, Ephemeroptera and Chironomidae as food items.

Reproduction

Presumed to be similar to its related species. An Iranian specimen had relatively large eggs (1.2 mm) when captured on 23 June.

Parasites and predators

None reported from Iran.

Economic importance

None, although the recorded habit of feeding on fish eggs by its relative may be true of this species and make it important in conservation of other species.

Conservation

Endemic to the Lake Orumiyeh basin where few specimens have been caught and deposited in museums. The numbers of this species in the wild are unknown.

Further work

The biology of this species, and its conservation status, have not been thoroughly investigated.

Sources check?

Type material: See above, BM(NH) 1899:30:90-96.

Iranian material: CMNFI 2007-0101, 12, 28.8-64.3 mm standard length, Azarbayjan-e Bakhtari, Tat'u River (ca. 36º54'N, ca. 46º07'E).

Genus Rutilus
Rafinesque, 1820

The roaches are found in Europe and western Asia where there are about 15 species (Bogutskaya and Iliadou, 2006). Three species are found in Iran.

The genus is characterised by having pharyngeal teeth in one row, usually 6-5, more rarely 6-6 or 5-5, with conical crowns on the anterior teeth and posterior teeth slightly hooked and truncated, scales are large to moderate in size, numbering 33-68 and with numerous fine circuli and radii on all fields, few and short gill rakers (17 or less), short gut, usually a light peritoneum, few to moderate numbers of dorsal and anal fin rays (7-13), the dorsal fin commonly having 4-5 unbranched rays, abdomen behind the pelvic fins rounded or with a slight but scaled keel, and various osteological characters (Bogutskaya and Iliadou, 2006).

Rutilus caspicus
(Yakovlev, 1870)

Caspian Sea species, R. rutilus freshwater residenth

Rutilus frisii
(Nordmann, 1840)

Common names

ماهي سفيد (= mahi safid or safid mahi, meaning white fish), sifid mahyi and asbalan mahi (in Gilaki), mahi safid daryacheh khazar or mahisephid-e-daryaye khazar (= Caspian Sea white fish), talaji (in Mazanderani).

[kutum, ak-balyk (= white fish) or ziyad in Azerbaijanian; akbalyk or kutum in Turlmenian; kutum in Russian; Southern Caspian roach; pearl roach; Caspian kutum].

Systematics

Leuciscus Frisii was originally described from the market in Odessa and the Danube, Dniester, South Bug, Dnieper and Don rivers, draining to the Black Sea.

Rutilus frisii kutum Kamenskii, 1901 is the generally accepted Caspian Sea basin subspecies although J. Holčík (pers. comm., 1994) considers that this is not a good taxon (see also Holčík and Jedlička, 1994). It was originally described as Leuciscus Frisii var. kutum from the Caspian Sea, essentially in the southern part, spawning in the rivers and streams of Transcaucasia and Persia (Kura, Araxes streams of the Lenkoran district) and in lesser numbers elsewhere. This subspecies is distinguished from the type subspecies from the Black Sea by having fewer lateral line scales (about 55-58 versus about 60-66), shallower body (depth equal to or less than head length versus exceeding head length), anal fin longer than high versus shorter than high, lower lobe of caudal fin usually shorter than head versus longer (young R. f. kutum have this lobe as long as head), and dorsal fin as high as long versus higher than long (young R. f. kutum have dorsal fin higher than long). Kotlík et al. (2008) discuss divergence and gene flow between Black and Caspian Sea populations, the majority of the migrations occurring during the Pleistocene (ca. 10,000-1,800,000 years ago). The refugial populations in the Black and Caspian seas have diverged despite periods of migrations between them.

Leuciscus frisii caspius Lönnberg, 1900 described "from the Volga delta" has priority over Rutilus frisii kutum which may be a nomen nudum as it is listed in Radde (1899) as "Leuciscus Frisii Nordm. var. Kutum Kam." without a description, the description only appearing in Kamenskii (1901). However, the name caspius has not been used while kutum appears widely in the literature as well as being the common Russian name of the fish. Additionally, as mentioned above, there may be no need of a subspecific name. Kottelat (1997) considers kutum to be a distinct species as do Bogutskaya and Iliadou (2006) and Fricke et al. (2007).

Leuciscus friesii Kessler, 1870 from the Volga delta is presumably a misspelling.

A hybrid of this species and Ctenopharyngodon idella has been bred at the Astaneh Ashrafie Fisheries Research Station (Sefidrud Research Station) and named "Samur" (Iranian Fisheries Research and Training Organization Newsletter, Tehran, 11:6, 1996; 18:6, 1997; Khara et al., 2002; Nouruz Fashkhami et al., 2002). Gynogenesis may have occurred. Artificial hybrids with Rutilus rutilus (possibly including R. caspicus) and Abramis brama have been bred in Iran (Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 39-40, 1996).

Key characters

This species is distinguished from the related Rutilus rutilus and R. caspicus by the higher scale count (usually >50) and the posterior part of the swimbladder being elongate and conical or pointed rather than rounded.

Morphology

Dorsal fin with 3 unbranched and 8-10, usually 9, branched rays, anal fin with 3 unbranched and 9-12, usually 10, branched rays, pectoral fin branched rays 16-19 and pelvic branched rays 8-9. Lateral line scales 47-68, mostly 55-58. Scales are regularly arranged over the body. A pelvic axillary scale is present. Scales have numerous circuli, numerous posterior radii, few but distinctively crowded anterior radii (more than in Rutilus rutilus and R. caspicus) and an almost central focus which is broken up into a network of lines. The posterior scale margin is crenulate and the anterior margin is wavy to crenulate but scale margins vary greatly between individual scales. Total gill rakers number 7-12 and are very short, hardly reaching the one below when appressed. Pharyngeal teeth usually 6-5, crowns rounded above a slender stalk, posterior teeth with a weakly hooked tip, posteriormost tooth margin may be serrated. The gut is an elongate s-shape. Chromosome number 2n=50 (Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 43, 1996; Noruz Fashkhani and Kosroshahi, 1995; Klinkhardt et al., 1995). Adibmoradi and Sheibani (2002) carried out a histological study of the brain of this species and concluded, in part, that vision was sharp and the fish were good chasers of food.

Meristic values for Iranian specimens are:- dorsal fin branched rays 8(1), 9(58) or 10(1); anal fin branched rays 9(24), 10(35) or 12(1); pectoral fin branched rays 16(15), 17(30), 18(14) or 19(1); pelvic fin branched rays 8(49) or 9(11); lateral line scales 47(1), 49(1), 50(1), 51(1), 52(7), 53(8), 54(6), 55(5), 56(11), 57(7), 58(7) or 59(5); total gill rakers 8(1), 9(21), 10(25) or 11(13); pharyngeal teeth 6-5(23), 5-5(1) or 5-4(1); and total vertebrae 40(1), 41(7), 42(34), 43(3) or 44(2).

Sexual dimorphism

Females are larger than males. The breeding tubercles are evident in males and may develop as early as the end of summer before the spring spawning season in the following year. Large tubercles are found on the top of the head above the eye level, behind the eye, between the eye and the mouth and on the snout as well as on upper flank scales. Tubercles are white in contrast to the dark head.

Colour

Small specimens up to a year old are silvery on the flanks and belly and the back is steel-grey to pale brown or pale olive. In adults, back scales are circled with black and there is a strong contrast between the back and flank. The anterior part of each flank scale, particularly those of the lateral line, is darkly pigmented. Lateral line scales may have two dots, one above and one below the opening as in some Alburnoides spp. but not as pronounced. The sides of the head are silvery with some yellow and darker pigment, the latter particularly in front of the eye. Adults are a bright silvery on the flank. The belly is pearly-white. The iris is silvery, with some spots, and with a marked dark spot above. The dorsal and caudal fins have some grey and a faint orange tint while the pectoral, pelvic and anal fins are colourless to lightly pigmented with black. The pectoral fin may be orange.

Size

Attains 66 cm body length and 4.065 kg in Iran (Farid-Pak, 1968a). Ouseley (1819-1823) ate one almost 3 feet long (ca. 0.9 m). In Iran during the 1950s, catches were 36-67 cm long (Farid-Pak, no date). Reaches 7.0 kg in weight.

Distribution

Found on the northern shores and drainages of the Black Sea, and the western shores and drainages of the Caspian Sea. Other populations are found in northwestern Turkey, Bulgaria and other parts of Europe, some subspecifically distinct from Rutilus frisii kutum. In Iran, it is found along the whole Caspian coast, entering almost all the rivers to spawn and also the Anzali Mordab and Gorgan Bay (Nedoshivin and Il'in, 1929; Kozhin, 1957; Holčík and Oláh, 1992; Riazi, 1996; Oryan et al., 1998; Abbasi et al., 1999; Kiabi et al., 1999; Abdoli, 2000; Lasheidani et al., 2008).

It is also found in Valasht Lake, Mazandaran (landlocked and introduced about 1850 to provide food for the royal family which had a summer residence there) where it possibly hybridises with native Alburnus chalcoides (Armantrout, 1980).

von den Driesch and Dockner (2002) record this species as faunal remains in an excavation at the Great Mosque in medieval Siraf on the Persian Gulf coast. This may well be a misidentification but, if not, indicates that a fondness for this species has a long history considering that a preserved fish or its remains had to be transported all the way from the Caspian Sea.

Zoogeography

A European and western Asian species with its origins in a Danubian or Sarmatian fauna.

Habitat

This species is migratory, spawning in rivers in March-April and returning to the Caspian Sea. Rezavi (1997) found three populations in Iran, one autumn and two spring populations. Il'in (1927a) records schools of fish entering the Anzali Bay as numbering in the several tens of thousands. On one fishing ground next to the Djifrud in February 1914, 12,000 fish were taken simultaneously. However, while the majority of fish migrate into lowland rivers not far from the sea to spawn among bulrushes and cattails, some migrate far upstream into the mountains of Tavalesh (= Talish) and Gilan where spawning conditions are very different. These fish may well overwinter in the river (Derzhavin, 1934). Such fish reached altitudes of about 1000 m before environmental changes inhibited the migration. Young fish migrate downstream, the migration ending in August, and in Azerbaijan enter the sea within 20-50 days. Holčík (1994; 1995) stated that Iranian young from the Anzali Mordab never entered the sea but remained in fresh or brackish water for 1-2 years. Riazi (1996) reports that this species migrates into the Siah-Keshim Protected Region of the Anzali Mordab. Young fish descend the Atrak River and feed in the sea at depths of 4.0-8.5 m. They winter in Iranian waters (Savenkova, 1985). Some fish are reported at depths of 36.6-53.0 m in the Iranian Caspian Sea (Knipovich, 1921).

Age and growth

Azari Takami et al. (1990) described the biology of this species in Iran. Males normally mature between their third and fourth year, sometimes earlier, females during their fourth year. Life span is at least 9 years in Dagestan (Shikhshabekov, 1979) and 8 years in Iran (Holčík and Oláh, 1992). Spawners are 3-8 years old and the principal age groups are 4-5 years for males and 5-6 years for females in the Anzali Mordab (Holčík and Oláh, 1992; Holčík, 1995), older than the 3-4 years of fish reported in 1970-1971. However, recently males are maturing at age 2 and females at age 4 with most spawners at age 3 and 4 years respectively (Holčík and Oláh, 1992). The average size of mature females (700 g) has been decreasing (Bartley and Rana, 1998b). Males grow faster than females until about the third year of life and then more slowly. Males are smaller and have a shorter life span than females (Holčík and Oláh, 1992). An annual increase in length of 17% over the first four years was observed in tagged fish, decreasing to 4.7% annually in subsequent years (for weight the figures are 61.2% and 25.7%). Sex ratios in spring were 3.08 males:1 female and in autumn 0.78 males:1 female (Annual Report, 1995-1996, Iranian Fisheries Research and Training Organization, Tehran, p. 55, 1997). Afraei Bandpei et al. (2008) surveyed biological characteristics in samples from Iranian coastal waters. Average fish size was 11.73 cm fork length, range 8.9-29.0 cm, average weight was 22.74 g and range 8.8-321.2 g. Females were larger than males (12.27 cm and 32.26 g on average compared to 11.03 cm and17.07 g). Sex ratio was 1.3:1 male:female. Overall condition factor was 1.23.

Fish may spend 1 or 2 years in fresh water after hatching (Holčík and Oláh, 1992) and this affects growth, 1 year fish growing more quickly than those spending 2 years in fresh water, maturing earlier and having a shorter lifespan. Growth in the Anzali Mordab is slower than it was 20 years prior to the report of Holčík and Oláh (1992), probably owing to a higher population density and more competition for food resources. The commercial catch in Iran was 3-7 years old, 39.0-57.1 cm long and weighed 613-2525 g (Razivi et al., 1972).

Growth in Iranian fish farms is up to 1.5 g in 8 weeks after a hatching rate of 75% (Aslaanparviz, 1994).

Food

Zarbalieva (1987) provides data on feeding of this species in the Caspian Sea off Azerbaijan. Fish concentrate on a sandy-shellrock bottom and remain there for most of the year to feed. The crab Rhithropanopeus harrisii dominates in the diet, 67.9-93.7% by weight. Molluscs, mainly Cerastoderma lamarckii, comprise 30% by weight of the food of fish 30-40 cm long. Fish larger than 40 cm seldom take molluscs but occasionally Clupeonella spp. Molluscs used to be the main diet item of this fish. Juveniles in the Anzali Mordab of Iran feed mostly on phytoplankton in contrast to the zooplankton reported for Azerbaijan fish (Holčík, 1995). This is a consequence of the poor productivity of this lagoon. Small Iranian specimens from the Caspian Sea had bivalve shell remains, plant remains and in one case a worm. Oryan et al. (1998) state that Cardium is the main food of this species on the eastern and western coasts of Bandar Anzali. Crabs and Balanus are also important. The hepatosomatic index is highest in February and March, the prespawning period. Afraei Bandpei et al. (2008) found diet in the southern Caspian Sea was Mytilaster (48.46%), Gastropoda (31.84%), Cerastoderma (19.65%) and Balanus (0.05%).

The hybrid with Ctenopharyngodon idella fed principally on macrophytes with phytoplankton as a secondary food in pond sin Gilan (Khara et al., 2002).

Reproduction

Fish spawn annually according to an Iranian study by Azari Takami et al. (1990) and probably return to their river of birth. Temperature, and probably river flow, are the factors determining the entrance of fish into the rivers on the spawning migration. Male fish run before females. Adeli Mosabbab and Piri (2005) report spawning in Iranian rivers from the end of March to the middle of April, beginning at 10°C. A large female, 60 cm long, 3.24 kg and 7 years old contained 124,712 eggs but some large females may have over 250,000 eggs (see also Farid-Pak (1968a) where fish 38-40 cm long have a mean value of 53,900 eggs, ranging up to 174,400 for fish 62-64 cm long; Yousefian and Mosavi (2008) give March to May as the spawning season. Abdurakhmanov (1962) gives a fecundity of 290,000 eggs for fish in Azerbaijan). Maximum egg diameter is 2.0 mm (Farid-Pak, 1968a). There are two spawning migrations, winter and spring, and so two forms or stocks of this species (Azari Takami et al., 1990). There may be three stocks based on electrophoretic studies of blood proteins associated with the two spawning migrations (Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 41, 1996). The winter form enters rivers with emergent or submerged aquatic plants at the end of autumn or the beginning of winter. Eggs are deposited on these plants. The principal rivers were the Nahang Roga, Pir Bazar Roga, Sowsar Roga, and Anzali Roga (all draining the Anzali Mordab) but the stock has declined with overfishing and destruction of the habitat. The Safid River is listed as the main spawning ground by the Caspian Sea Biodiversity Database (www.caspianenvironment.org). The spring form enters rivers at the end of winter or the beginning of spring and spawns on gravel or sand. Adults return downstream after spawning, in the early morning. The run occurs both by day and night but increasing water clarity near the end of the spawning season limits runs to the night.

The migration into the "Dinachal" (? = Denya Chal) River occurred on 1 November in 1975, when sexually immature fish were caught. By 11 February, 5 spawners were caught and the peak migration occurred on 26 March when 3,845 fish were caught at a water temperature of 10.5°C. The migration was continuous until 11 May. Water temperatures between 28 March and 23 April in 1976 and 1977 varied between 9.1°C and 13.5°C when peak migrations occurred in the "Havyg" (= ? Haviq) River. Here the spring form of this species started to migrate at 6°C and the optimum water temperature appears to be between 11 and 13°C. Yousefian and Mosavi (2008) found a spawning temperature range of 5 to 21°C centred on 11°C. Fertilisation rate in 4 rivers was 75-92% and hatchability was high. Rivers with high turbidity had low hatchability because of egg damage. Further details of natural and artificial reproduction are given by these two authors.

The main migration (99%) in the Anzali Mordab began in February and lasted 3 months at 8-10°C with a much smaller run (only about 1%) in November-December. The Mordab is now overgrown with reeds or has a silty, vegetated bottom and is no longer a spawning site (Holčík and Oláh, 1992), an economic and poetic loss. Holčík and Oláh (1992) translate Il'in (1927a) on Anzali Mordab mahi safid "During this time the reeds rustle as if of wind as the fish rubs pushing their flanks to the reed stalks or pushing forward among reeds standing in rows".

Khaval (1998) reports a spawning migration into the Safid River despite construction, sand removal and pollution. The migration peaks in the second half of March at a water temperature of 11°C.

Each female is flanked by two males at spawning in shallow water. Sometimes the backs of the fish protrude from the water. Females sharply rub their lower abdomen and pectoral area on the gravel bottom, males and females convulse, and eggs are shed and fertilised. Eggs are adhesive and hatch in 20 days at 10-14°C. Embryonic development in glass incubators at 14-16°C is described by Parivar et al. (1993). Cleavage and gastrulation begin within 24-30 hours, hatching at 216 hours (10 days after fertilisation) and resorption of yolk is complete at day 16. Some males and females are injured through contact with the gravel stream bed. Embryonic development takes 10-15 days at 8-16°C and 5-6 days at 20°C (Aslaanparviz, 1994). Young fish go down to the sea in summer according to some works although Holčík and Oláh (1992) believe that they stay in fresh water or brackish river mouths for 1-2 years before entering the sea.

Kousha et al. (2007) examined some aspects of reproductive endocrinology in this species, specifically on sex steroid-binding protein in plasma. Najafipour et al. (2008) examined 17-αhydroxy progesterone hormone levels, egg diameters and liver weight in fish caught at sea and in the Chalvand River near Astara. These factors were used to indicate physiological indices of reproduction, hormone levels and egg diameters increasing in fish caught in the river, while liver weight decreased. Egg diameters increased from 1.55 to 2.14 mm.

Parasites and predators

Mokhayer (1976b) records the digenetic trematodes Aspidogaster limacoides and Asymphylodora macracetabulum. Eslami and Kohneshahri (1978) report various helminths from this species in Iranian waters. These are the monogenean Diplozoon paradoxum, the aspidogastrean Aspidogaster limacoides, the digenean Asymphylodora kubanicum and larvae of the nematode Anisakis sp. This fish therefore is potentially a source for human infection with Anisakis. Mokhayer (1989) reports metacercariae of the eye fluke, Diplostomum spathaceum from this species in Iran, which can cause complete blindness and death in commercially important species. Jalali and Molnár (1990a) record the monogeneans Dactylogyrus frisii from this species in the Khosk (= ? Khoshk) River (Caspian basin) and D. rarissimus and two monogenean species, Dactylogyrus spp., from the Safid Rud. Jalali and Molnár (1990b) record the monogeneans Dactylogyrus frisii and D. rarissimus at fish farms in Iran. Molnár and Jalali (1992) report the monogeneans Dactylogyrus suecicus, D. haplogonus and D. turaliensis from this species in the Safid Rud. Gussev et al. (1993b) record the monogenean Dactylogyrus haplogonus from this species in the Safid Rud. Masoumian and Pazooki (1998) surveyed myxosporeans in this species in Gilan and Mazandaran provinces, finding Myxobolus bramae. Safari and Khandagi (1999) record Clostridium botulinum from 2.2% of fresh and smoked samples of this species in Mazandaran Province. Shamsi and Jalali (2001b; 2001c) detail monogenean parasites for Safid Rud and Caspian Sea samples, including 6 species of Dactylogyrus. Jalali et al. (2005) summarise the occurrence of Gyrodactylus species in Iran and record G. prostae and G. sp. in fish from the Safid River.

Amini (2006) records Gyrodactylus sp., nematodes, Diplostomum sp., Trichodina sp., Aeromonas sp. and Hydrophila sp. in fingerlings from hatcheries in Iran. The latter two parasites caused a heavy mortality in the Shahid Rajaii hatchery ponds.

Tavassoli and Moghir (20020 describe a squamous cell carcinoma in the oral cavity of this species, the first record for the Caspian Sea.

Economic importance

Safid mahi is the most popular fish in Iran with the highest economic value (Azari Takami et al., 1990). More than 70% of bony fish caught in the coastal Caspian Sea of Iran are this fish (Yousefian and Mosavi, 2008). It may be available out of season as mahi qachaq or bootleg fish, an indication of its popularity. Batmanglij (1999) notes that this fish is baked and served with herbed rice. A stuffing consists of garlic, parsley, tarragon, scallions, coriander, mint, ground walnuts, barberries, raisins, lime juice, salt and pepper, sautéed in butter (he also incorrectly refers to mahi-e safid as striped bass). In 1996 a news report stated that "In the port city of Anzali, one small white fish sells for $10" (http://www.iran-e-azad.org/english/boi/03400201.96). In Rasht, 60% of the inhabitants consume no other fish and "86% of ladies in Rasht can only cook one kind of seafood" (Khairkhah, 1994). Ouseley (1819-1823) reported that this species "seemed most abundant, and was found in all the great rivers of this country near the sea; for several days it had furnished the principal dish of my dinners and often of my breakfasts". Holmes (1845) recorded catches with cast-nets in the Anzali Mordab over 150 years ago which were even then worth £1400-1500. O'Donovan (1882) reported that a small stream in the Atrak River drainage was so crowded with this species that individuals could only move by floundering and jumping over one another, and the horses in crossing the stream, trod them to death by scores. Lönnberg (1900) reported an annual catch of some millions in Persian waters.

It is caught in rivers and lagoons and by large, mechanically hauled beach nets in the Caspian Sea. Sea nets can be 1500 m long and 18 m deep. The fishing season begins in October and reaches a maximum between 20 February and 10 March, ready for the "Now Ruz" or New Year celebrations when many Iranians eat this fish with rice (Emadi, 1979).

The roe of this species is also eaten, salted or unsalted (the fish are called asbalan mahi and the roe shur-e asbal in Gilaki, the local dialect of Gilan). Whole female fish are soaked for one year in a mixture of salt and madder in special clay jars. The jars are traditionally buried in the ground and hermetically sealed but a modern technique has the roe removed from the fish after 20 days. Esmaeilzadeh et al. (2004) studied the nutrient composition and marinade qualities of this fish and compared them to those for grass carp (Ctenopharyngodon idella), the latter being preferable according to the organoleptic properties. The marinades could be stored for 6 months at 10ºC.

The average weight of fish caught in the Anzali Mordab in the early years of this century was 2 kg, the catch in the 1914-1915 season was 47,000 fish and in 1913-1915 123,000 fish. On 26 February 1914 a single haul in the sea near Anzali took 41,045 fish (Il'in, 1927a; Berg, 1948-1949). Il'in (1927a) estimated a yearly catch in Anzali Bay to be 3-4 million fish, a figure conflicting with these other reports.

Nevraev (1929) gives catches for various fishing regions in Iran in the early twentieth century. For the period 1901-1902 to 1913-1914 the catch in the Astara region was 0 to 29,053 individuals, for 1901-1902 to 1917-1918 the catch in the Anzali region was 2565 to 124,195 individuals, in the Safid River region from 1904-1905 to 1917-1918 the catch was 100 to 31,799 individuals, and in Astrabad (= Gorgan) region from 1900-1901 to 1912-1913 the catch was 4000 to 323,500 individuals. The total catch for Iran in the 1914-1915 season was 443,000 fish. The catch in Iran from 1956/1957 to 1961/1962 varied from 197,884 kg to 2,066,580 kg (Vladykov, 1964), from 1965/66 to 1968/69 it varied between 159 and 1252 tonnes (Andersskog, 1970), from 1963/64 to 1968/69 it varied between 121.3 and 1252 t (RaLonde and Walczak, 1970b; 1972), from 1987 to 1991 it varied between 3500 t and 8855 t (Holčík and Oláh, 1992), and between 1989 and 1998 it varied between 11,792 kg and 14,336 kg (Caspian Environmental Programme, 2001a) - catch figures are at variance with each other. Holčík and Oláh (1992) report a catch of 3107 kg in the Anzali Mordab for 1990 and for 1932-1964 a range of 95.1-3488.9 t. They are also caught in rogas and inflowing rivers of the mordab in late winter and early spring. Moghim et al. (1994) estimate that coastal areas of the southern Caspian Sea have a total biomass of 24,000 t with a maximum sustainable yield of 7000 t. In 1993-1994 the total catch of this species, including the illegal catch, was 11,175 t with the total stock estimated at 25,400 t and a maximum sustainable yield of 9300 t. More than 25% of the catch was young fish indicative of non-standard methods being used (Annual Report, 1995-1996, Iranian Fisheries Research and Training Organization, Tehran, p. 19-20, 1997). In 1994-1995, the biomass of this species in Iran was 241,000 t (sic, probably 24,000 t) and the maximum sustainable yield was 9000 t (Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 37, 1996). About 62% of the bony fish catch in the Caspian Sea of Iran in 1993-1994 was this species with the mullet Liza aurata second at 22% (Annual Bulletin 1993-94, Iranian Fisheries Research and Training Organization, Tehran, p. 83, 1995). The catch from beach seine cooperatives along the Iranian coast was 8477 t in in 2004-2005, a 2500 t decrease over the previous year. The biomass of this species in Iranian coastal waters for 2003-2004 was estimated art about 25,000 t (Abdolmalaki, 2006a).

Bartley and Rana (1998a; 1998b) comment that the fishery collapsed in 1980 but has risen from 500 tonnes in 1981 to around 10,000 t in 1996 after restocking from around 400,000 fingerlings/year in 1981 to around 142 million/year in 1997. Rana and Bartley (1998a) note that 7 million 1 g fingerlings were released into the Caspian Sea in 1997 which contradicts their earlier report. There is only a state supported stocking programme but ERM-Lahmeyer International GmbH, DHI Water & Environment and GOPA Consultants (2001a) note that this fish is being successfully managed in Iran while it has been fished almost to extinction in waters off Daghestan and Azerbaijan. Market price is high at about U.S.$5.00/kg, five times the price for silver carp. Export prices are higher, 700 g of smoked "mahi-sefid" cost £21.00 in 2004 (www.superhormuz.com, downloaded 19 January 2004). However this species is difficult to culture beyond the 40-50 g stage and growth is slower than carp.

The conflicting ranges seen in Andersskog and RaLonde and Walczak are typical of the wide variations in reports; figures can only be taken as general guides for many Iranian fisheries. Abdolmalaki (2006c) states that fluctuations are due to destruction of spawning grounds, overfishing and release of fingerlings. The mean catch sizes were 3110 tonnes in 1937-1947, 990 t in 1967-1977 and 8505 t in 1987-1997. A minimum catch of 121 t was taken in 1964 and a maximum of 11,175 t in 1994. Catch-per-unit-effort also showed high variation,. The calculated stock biomass was 1300 t in 1071 and between 18,489 and 25,400 t between 1990 and 2000. The mean biomass in the past 10 years was 22,750 t, a 17-fold increase over the year 1971. The catch in the previous 10 years was 35-46% of the annual stock. A decrease in stock size during 1998-1999 was attributed to an exploitation rate being more than maximum sustainable yield, a decreased mean weight of released fingerlings and a lowered return rate.

Catches in the Bandar-e Anzali area have been as high as 5,480 t in 1939-1940 but fell to 85 t in 1961-1962 (Vladykov, 1964; RaLonde and Walczak, 1972) but have risen again as indicated above. This is attributed to a massive stocking effort with 170 million fingerlings released in Iran in 1991 (cf. Emadi (1993a) below), about a quarter in the Anzali Mordab (Holčík and Oláh, 1992). The Sari hatchery produced 400 million "white fish" over the previous 10 years (Tehran Times, 30 May 1998). Fingerling production in 1996 was 142.1 million (Bartley and Sana, 1998a). Sea ranching increased the yearly catch to 8500 t in 1991, the highest recorded catch in the past being 5850 t in 1940 (Emadi, 1993a). Natural stocks in the past were very high. Migrating fish were so dense at the Mordab mouth that they were caught in buckets and jumping fish literally fell into boats.

Savenkova (1990) comments on the marketing of this species in southwestern Turkmenistan. Salehi (2003) analysed the economics of production of finglerings in Iran. The cost of labour was 50%, feed and fertiliser 20%, maintenance 10%, and harvesting, handling and releasing 6%. The average cost per fingerlings was 37 rials in 2001 (rising to 123 rials in 2003 (Salehi, 2008)). As the average rate of fingerling return was 8.3% and the average weight and age of commercially caught fish was 815 g and 3.7 years, it was expected that 19,257,494 individuals weighing 16,000 t would be harvested over the Iranian year 2004-2005. The value would be an estimated 345 billion rials at the 2001 price (8050 rials to a U.S. dollar in 2001; $42.86 million). Salehi (2008) estimated the wholesale price of the catch as 505 billion rials in 2004 and reviewed the comparative economics of fingerling production for 2001-2003.

Experimental culture of triploid mahi safid has been carried out at the Gilan Fisheries Research Centre (Iranian Fisheries Research and Training Organization Newsletter, 2:2, 1993) to increase fish weight for exploitation but apparently was unsuccessful although the techniques worked with grass carp, Ctenopharyngodon idella (Annual Bulletin 1993-94, Iranian Fisheries Research and Training Organization, Tehran, p. 70-71, 1995). In addition, monoculture and polyculture of this species has been investigated, the latter with grass carp (Ctenopharyngodon idella) and silver carp (Hypophthalmichthys molitrix) (Iranian Fisheries Research and Training Organization Annual Report, 1992-93; Danesh Khoshashi, 1997). From an average initial weight of 7 g, monocultured fish weighed 158 or 177 g on average at the end of each of two one-year periods, polycultured fish weighed 158 or 168 g, and maximum weight attained was 250 or 300 g (Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 38, 1996). Danesh Khoshasi (1997) gives figures of 177 g for mono-cultured fish after 6 months in ponds, poly-cultured fish weighed 185 g (with a maximum of 300 g) while stocking with 7 g fish gave 168 g and 192 g for mono- and poly-culture fish respectively (maximum weight 250 g)(these two reports on the same experiment have confusing figures). The fish were fed on pellets especially made for this species, nitrate and phosphate fertiliser and cow and chicken manure were added to the ponds, and water temperature was 15.0-27.64°C.

This species has been used in Iran for experimental studies, e.g. on the acute LC₅₀ and bioconcentration of mercuric chloride (Gharaei et al., 2006; Gharaei and Esmaili-Sari, 2008); on the toxicity and LC₅₀ of phenol and 1-naphthol (Shariati et al., 2004); the effects of anionic detergents and diazinon on the LC₅₀, both separately and in combination (Tehranifard et al., 2002; Tehranifard et al., 2007); etc.

The hybrid with grass carp, Ctenopharyngodon idella, reached an average weight of 100 g and a length of 22 cm after 5 months in the first report cited above and 6.7 g and 9.17 cm after 4 months in the second report. Larvae were fed live food twice a day and hybrids larger than fingerling size ate grass. The hybrid phenotype resembled the safid mahi but was a herbivore. It was expected that this hybrid would be used as a new culturable "species".

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

Conservation

Azari Takami et al. (1990) list the following reasons for a decline in the commercial catch of this species:- a) regression of the Caspian Sea which decreased the surface area of the Anzali Lagoon and increased the growth rate of aquatic vegetation, b) mechanisation of farming and a consequent increased demand for irrigation water leading to reduced river flows during the spawning migration, c) use of fertilisers and pesticides in rivers draining into the Anzali Mordab (and pesticides and herbicides such as Diazinon, Malathion, Machete and Saturn have a highly toxic effect on fingerlings of this species (Piri et al., 1999); and the herbicide butachlor reduces sperm volume and increases abnormal sperm (Lasheidani et al., 2008)), d) pumping of river water for irrigation causing mass mortalities of fry, e) industrial development increasing the pollution load, and f) excessive catches of adults to the extent that all spawners in a river were taken. Emadi (1979) added such factors as road construction and the removal of sand from banks and river beds, erosion caused by felling trees and shrubs along river banks and in the mountains, construction of bridges and raising of their substructures which formed barriers to migration, and climatic changes. Illegal fishing and non-standard nets threaten the stocks while fingerling release (120-140 million) and improvement of natural spawning areas through rises in water level have contributed to stock increases (Annual Report, 1995-1996, Iranian Fisheries Research and Training Organization, Tehran, p. 19-20, 1997). Three million fingerlings weighing 3-5 g were released into the Anzali Lagoon (Iranian Fisheries Research Organization Newsletter, 49:4, 2006).). Poaching is rampant, even on the spawning grounds, and is also a significant factor in decline of this species (RaLonde and Walczak, 1972).

In the 1970s, rivers were rented to fishermen to exploit to an unlimited degree such that all spawning fish were taken and there was no recruitment for 4 years (Carl Bond Archives, Oregon State University, Corvallis).

The catch declined from 5854 t in 1918 to 172 t in 1937. Actual catches are about 20-30% larger because of local sale and consumption (Emadi, 1979). RaLonde and Walczak (1970b) note a decline from 1556 t in 1957 to 162.1 t in 1967 so the catch does not decline evenly. Emadi (1979) also pointed out that intensive sea fishing and fishing in rivers hindered spawning of the winter form, and led to a situation where only the spring form spawned. Leasing of rivers to fishermen was discontinued in an attempt to alleviate the decline in this species. The migration distance up rivers has decreased from about 25 km to about 8 km in recent years because of water abstraction and dams for agriculture (Bartley and Rana, 1998b). However, Yousefian and Mosavi (2008) report that artificial stocking has increased catches from 1000 t in 1981 to 10,000 t in 2002.

A further problem is the restocking programme only takes spring-run fish; the fall-run stock may no longer exist (but see below). In addition, stocks from various rivers are mixed and may result in outbreeding depression, the loss of adaptations to specific rivers in terms of migration patterns, spawning time, behaviour and other factors (Bartley and Rana, 1998b).

Measures to combat loss of this valued fish included a ban on fishing in the Anzali Mordab and its tributary rivers (catches here were up to 1000 t per year (Emadi, 1979)), effective control of illegal fishing and artificial spawning experiments (although the latter was insufficient to replace stocks). For several years starting in 1925, artificial breeding raised larvae for release in the ten most important rivers. In 1976-1977, fingerlings were raised to increase the stock in the Caspian Sea. Over 5 million fry were produced by the "Havyg" (? = Haviq) Hatchery alone in 1977.

Emadi (1979) recorded releases of 28-44 million fingerlings and 250 million larvae "in recent years". A farm in the Siah Kal region of Gilan Province near Rasht was expected to produce 40 million white fish "roe", presumably fry, in 1985 (Kayhan International, 20 May 1984). The Dr. Beheshti Hatchery near Rasht expected to release more than 60 million fingerlings in the Iranian year 1993-1994 (Abzeeyan, Tehran, 4(5):VI, 1993). This hatchery had a peak production for the period 1973-1993 of over 140 million fingerlings in 1989, rising from the low period of 1973-1982 with less than 10 million fingerlings, and with subsequent decreases to about 50 million fingerlings in 1993 (Abzeeyan, Tehran, 5(3 & 4):IX-X, 1994). The number of fingerlings released from 1986 to 1991 climbed from 38 million to 170 million (Holčík and Oláh, 1992). Krasznai (1987) also refers to propagation of this species at "Sad-e Sangar" (Dr. Beheshti) and Siah Kal Fish Farms near Rasht in Gilan. Emadi (1993a) gives fingerling production at government hatcheries as follows: 25.3 million in 1983, 28.3 million in 1984, 38.0 million 1985, 51.7 million in 1986, 72.0 million in 1987, 84.3 million in 1988, 140.2 million in 1989, 156.3 million in 1990, 110.0 million in 1991, and 145.0 million in 1992. The Shahid Rajaee Hatchery in Sari released 358 million fingerlings, possibly in a single year (Abzeeyan, Tehran, 4(7):VII, 1993), although 70 million are reported as released annually to Mazandaran rivers in 1995 (Abzeeyan, Tehran, 6(8):III, 1995) and production of 70 million kutum fingerlings annually is reported in 2001 (Iranian Fisheries Research Organization Newsletter, 28:3, 2001). In 1997, 142 million fingerlings were produced for restocking (Bartley and Rana, 1998b). In 1999-2000, 150 million juveniles were released into the Caspian Sea (Iranian Fisheries Research Organization Newsletter, 23:4, 2000). From October to March 2000, 80 million juveniles raised in the Shahid Ansari aquaculture and breeding centre in Gilan were released into the Caspian Sea and neighbouring water bodies (Iranian Fisheries Research Organization Newsletter, 26:2, 2001). Billard and Cosson (2002) cite a mean of 100 million alevins released per year, reaching 140 million in some years, and give a brief overview of production facilities. Mosabbab and Piri (2005) record the release of fry into the Gorgan River of the southeast Caspian Sea as 16,663 million in 2000, 19,119 in 2001, 12,263 million in 2002, 11,931 in 2003 and 10,413 in 2004, the decrease being due to a fall in capture of brood fish. Amini (2006) reports release of fingerlings into the Larim, Goharbaran, Shirood, Tonekabon, Sardabrud, Mirud, Babol, Asbuchin and Sorkhrud rivers in 2000-2001 numbering 53.7 million and 62.5 million. Fork length was 36.7 mm and 43.3 mm and condition factor 1.13 and 1.2 0 respectively by year. In 2006, a report has more than 150 million juveniles being released into the Caspian Sea every year (www.iranfisheries.net, downloaded 28 July 2006). Sources obviously conflict on exact numbers, nevertheless marked variations in production are evident over short periods. The South Caspian Fisheries of Azerbaijan also released larvae in Soviet waters, more than 150 million in 1983 for example (Zarbalieva, 1987).

The Inland Water Aquaculture Research centre in Anzali has artificially propagated the autumn or fall-run stock (www.iranfisheries.net, downloaded 28 July 2006). Three million fish were released in July 2006, probably at 3-4 g with more to be released at a heavier weight. Paykhan Heyrati and Dorafshan (2007) compared the effectiveness of two kinds of dopamine antagonist combined with a gonadotropin releasing hormone analogue on ovulation and fertilisation success under hatchery conditions. Spawning induction techniques using hormones provide high quality gametes in quantity for aquaculture programmes.

Azari Takami et al. (1990), Woynarovich (1985) and Bartley and Rana (1998b) detail the technique for artificial spawning, incubation and raising of fingerling mahi safid. Adults are caught as they enter rivers on the spawning migration by blocking the river with wooden tripods interlaced with twigs as screens and using a meshed net. The Shaeed Ansari Fish Farm (of Shilat, the Iranian Fisheries Company) takes fish from 5 main rivers, forming a brood stock of 10,000 females and 20,000 males (Bartley and Rana, 1998b). Almost 100% of migrating fish are caught. Ripe fish are stripped and the eggs fertilised with sperm. Eggs from 2 females are pooled and mixed with milt from 2 males. The adhesive layer is washed away with water and continuous stirring. Once the eggs are completely separated from each other they are placed in incubator jars. Eggs may be placed in net-bottomed trays in the river for 2 days before being sent to a hatchery. Egg development takes a minimum of 7 days but larval development only takes 1-2 days. Larvae are fed with a mixture of milk and eggs until they are 5 days old and then put in the earthern rearing ponds. In the Shaeed Ansari Fish Farm, fingerlings are released in a river mouth at 1 g and most enter the Caspian Sea within 3 days (Bartley and Rana, 1998b). Fingerlings are grown in ponds to a weight of 2-3 g and then released in the Safid River which carries them down to the sea (Petr, 1987). Production of 1-2 g fingerlings attains 3 tonnes per hectare in fish farms (Emadi, 1993a). Farabi et al. (2007) studied brood stocks and fingerlings in the Shirod, Tonekabon, Tajan and Goharbaran rivers from March 2004 to March 2005. The mean length, weight and condition factor for broodstock females and males were 43.75 and 36.5 cm, 1189.5 and 678.13 g and 1.42 and 1.38 respectively. Eggs collected over a 62 day period weighed 4931 kg, mostly at the end of March and beginning of April. Percentage of survival of eggs in the four rivers listed above was 94.5, 95.1, 87.7 and 96.9 respectively. Newly hatched larvae averaged a total length of 6±2 mm and a weight of 2±0.2 g. The number of fish produced at less than 1g was 16,942,454 representing 19.9% of the total released in Mazandaran in 2004. Fish in the 1.0-1.5 g weight class, suitable for release, numbered 62,905,247. Parasites of fry were Diplostomum, Dactylogyrus, Butriocephalus, nematodes, Trichodina and Epistelis.

Studies on the best methods of rearing fingerlings and fry have been carried out in Iranian hatcheries. The use of rotifers (Brachionis plicatilis) as food is more effective in larval growth than concentrated food preparations; after 40 days the rotifer-fed fish reached 440 mg and 31.36 mm on average and those fed concentrated food reached 17.2 mg and 14.75 mm (Fallahi et al., 2006; Kapourchali, 2006). Neverian et al. (2006) studied variation in protein levels fed to advanced fry at an isocaloric digestable energy of 3400 kcal/kg. The highest treatment at protein level 35% showed increased body protein, decreased fat and improved flesh quality in the carcass. Trichlorfon, an organophosphate pesticide is used to protect zooplankton such as rotiferans, food for larval safid mahi, against predators. However it does slow growth of the fish and the larvae must only be added to treated water 72 hours after the last application of the chemical (N. Chookbar, www.netiran.com, downloaded 18 April 2005; Chobkar et al., 2005; 2008). The sex steroid binding protein has been investigated as a biomarker for sexual behaviour in this species (Kousha et al., 2006). Spawning induction studies have been carried out on broodfish in Iran using carp pituitary extract and GnRH analogue, the latter alone or combined with metoclopramide. The latter treatment gave 90% ovulated females, 71.3% ovulation index and 68.4% fertilisation success, significantly higher than values for carp pituitary extract. However, the latency period was longer. Paykhan Heyrati et al. (2006) and Paykhan Heyrati and Dorafshan (2007) induced spawning with a gonadotropin releasing hormone analogue, both alone and in combination with the dopamine antagonists domperidone and metoclopramide, the combination with both antagonists being equally successful. Noverian et al. (2008) studied the effect of dietary digestible energy level on growth indices of advanced fry. Body protein and fat improved when the dietary energy was raised from 2500 to 2800 kcal kg^-1. Haghighi et al. (2009) investigated the effects of dietary lipid levels on fry, growth, survival, protein and energy retention decreasing with an increase of dietary lipid level; a lipid level of 8% and 4314 kcal/kg bringing about the best growth performance. Akbarian et al. (2009) found that mid to high light intensities and long photoperiods could be useful in hatcheries where no particular light regime had been used.

Savenkova (1990) lists several reasons for reductions in catches in the Atrak River stock of Turkmenistan on the border with Iran. These include ineffective fish passes, poor marketing strategies, and the low quality of fish reaching the spawning grounds.

A fine of 1,500 rials is imposed specifically for illegal angling of this species (Anonymous, 1977-1978).

Kiabi et al. (1999) consider this species to be of least concern in the south Caspian Sea basin according to IUCN criteria. Criteria include commercial fishing, sport fishing, abundant in numbers, habitat destruction, widespread range (75% of water bodies), absent in other water bodies in Iran, and present outside the Caspian Sea basin. The 2000 IUCN Red List lists this species as DD (Data Deficient). Coad (2000a), using 18 criteria, found this species to be one of the top 4 threatened species of freshwater fishes in Iran. Extinct in Turkey (Fricke et al., 2007).

Further work

The heavy demand for this fish as a traditional food warrants extensive management and conservation of stocks.

Sources

Iranian material: CMNFI 1970-0513, 9, 51.1-66.1 mm standard length, Gilan, Shafa River estuary (37º35'N, 49º09'E); CMNFI 1970-0516, 27, 42.3-61.0 mm standard length, Gilan, Lomir River (38º14'N, 48º52'30"); CMNFI 1970-0518, 9, 43.4-60.7 mm standard length, Gilan, Haviq River estuary (38º10'N, 48º54'E); CMNFI 1970-0520, 19, 46.7-73.9 mm standard length, Gilan, Astara River (ca. 38º25'N, ca. 48º52'E); CMNFI 1970-0563, 24, 40.8-58.1 mm standard length, Gilan Caspian Sea, Kazian Beach (ca. 37º29'N, ca. 49º29'E): CMNFI 1979-0088, 1, 104.3 mm standard length, Gilan, Safid River (no other locality data); CMNFI 1979-0471, 1, 106.0 mm standard length, Mazandaran, Caspian Sea west of Alamdeh (36º35'N, 51º48'E); CMNFI 1979-0686, 30, 42.8-52.2 mm standard length, Gilan, Safid River above ferry (37º24'N, 49º58'E); CMNFI 1980-0159, 1, 117.0 mm standard length, Gilan, Caspian Sea at Kazian Bridge (37º28'30"N, 49º28'E); CMNFI 1980-0160, 1, 88.5 mm standard length, Iran, Caspian Sea basin (no other locality data); CMNFI 1993-0140, 2, 64.5-71.5 mm standard length, Mazandaran, Tirom River, Ramsar (36º51'48"N, 50º48'E).

Comparative material: BM(NH) 1879.11.14:31-32, 2, 410.0-430.0 mm standard length, Russia, Astrakhan (ca. 46º24'N, ca. 48º05'E).

Rutilus kutum
Kamenskii, 1901

?separate out frisii data

Rutilus rutilus
(Linnaeus, 1758)

need to separate out caspicus information?

Common names

كلمه (= kolme, kollme, kolmeh or koolmeh, meaning unknown), mahi kolme, kolme Gorgan, telagi or talaji (in Gilaki and Mazandarani, meaning unknown, latter three used for R. r. caspius natio knipowitschi), mahi cheshm qermez (= redeye fish), kolme Kura or kolme Anzali for R. r. caspius natio kurensis, kolme Gorgan or kolme Turmenistan for R. r. caspius natio knipowitschi

[kulma, kyumen, xazar kulmasi and Kur kulmasi in Azerbaijan; kasli akcapagy in Turkmenian; Armyanskaya plotva or Armenian roach, karmraki and last for schelkovnikovi, all in Armenia; vobla, severo-kaspiiskaya vobla or North Caspian vobla, Astrakhanskaya vobla or Astrakhan vobla, Kurinskaya vobla or Kura vobla, Astrabadskaya vobla or Astrabad vobla, Turkmenskaya vobla or Turkmenian vobla, all in Russian; roach being the English equivalent of vobla].

Systematics

Cyprinus Rutilus was originally described from lakes in Europe, the type being from Sweden.

Holčík and Skořepa (1971) revised the roach and found no reason to maintain subspecies, of which they list 12. Rutilus rutilus caspicus (Yakovlev, 1870) (North Caspian or Astrakhan vobla) was the subspecies found in the Caspian Sea basin with natio knipowitschi Pravdin, 1927 (Astrabad or Turkmenian vobla) in Gorgan (= Astrabad) and Gasan-kuli bays and the Gorgan, Atrak and Qareh Su rivers and natio kurensis Berg, 1932 (Kura vobla) in Kyzylagach Bay, the Kura River, Astara and rarely the Anzali Mordab. The Kura vobla was said to differ from the Astrabad vobla by a greater body depth, smaller eye and more rapid growth. R. r. caspicus was distinguished from other subspecies by having modally 9 branched dorsal fin rays, darker fins, more inferior mouth, higher dorsal and anal fins, longer pectoral and pelvic fins, a deeper head, and larger eyes (see Berg (1948-1949) for more details). R. r. caspicus was originally described as Leuciscus rutilus var. caspicus from the Volga River delta, Russia. The types are unknown (Eschmeyer et al., 1996). Eschmeyer et al. (1996) also list Leuciscus rutilus var. wobla Grimm, 1896 described from the North Caspian Sea and mouths of rivers (Volga, Ural, Emba, Terek, Kura, Astara) entering the Caspian Sea. This is presumably a synonym of Rutilus rutilus judging from the subspecific name which is the Russian word for this species (wobla = vobla or roach). Kottelat (1997) lists it as a nomen nudum.

Holčík and Skořepa (1971) recognize the Caspian Sea populations as the morph or morpha migratorius, perhaps a first step towards differentiation and speciation. Mironovskii and Kas'yanov (1986; 1987) however retained Rutilus rutilus caspicus as a distinct subspecies in the Caspian Sea based on a multivariate analysis of 12 meristic and 7 morphometric characters. Mironovskii (1991; 1992) also demonstrated differences between Turkmen and Azerbaijan samples. Kuliyev (1984) considered these variations to be responses to different and changing conditions of the environment. Naddafi et al. (2002) also demonstrated differences in meristic (anal fin rays, predorsal scale number and total body vertebrae of 12 characters studied) and morphometric characters (7 measurements of 28 studied) for fish from the Anzali Mordab and the Gorgan River estuary of Iran. Keyvanshokooh and Kalbassi (2006) however found the value of Nei's genetic distance (d = 0.04) to be small between these two populations using DNA. The populations had similar levels of polymorphism. Keyvanshokooh et al. (2007) compared populations in the Anzali Wetland and Gorgan Bay using microsatellite markers and found differences between both populations were not significantly different for average number of alleles per locus nor for observed heterozygosities. Parafkandeh Haghighi and Rezvani (2005) and Parafkandeh Haghighi (2006) used the trace element content in otoliths to demonstrate the presence of two different populations in the southern Caspian Sea, the Anzali-Kura and the Gorgan-Turkmen, as noted above on meristic and morphometric values. Patimar et al. (2005) found morphological differences between fish from the Gomishan Wetland and those from the Adji-gol and Alma-gol wetlands in southeast Iran, and between the latter two wetlands. Inter- and intra-population variation was higher in the Gomishan fish. Rezvani et al. (2006) used mitochondrial DNA to compare Anzali and Turkmen roach and found the former to be more genetically variable, attributing this to there being more rivers, and therefore more spawning populations, in that area. Reyhani et al. (2010) used microsatellite markers and found higher heterozygosity in Anzali Wetland fish compared to those from Gorgan Bay. However, gene flow was high and population differentiation was non-significant.

Various body forms of this species have been given names as morpha, sometimes taken from species names regarded as synonyms, although as morpha these have no taxonomic significance, e.g. prasinus from Leuciscus prasinus Agassiz, 1835 (= elongata from Leuciscus rutilus var. elongata Fatio, 1882) for an elongate body and rutiloides from Leuciscus rutiloides Selys-Lonchamps, 1842 (= elata from Leuciscus rutilus var. elata Fatio, 1882) for a wide-backed body. A golden-orange or red aberration was known as aurata from Leuciscus rutilus var. aurata Fatio, 1882.

Bogutskaya and Naseka (2004) and Kottelat and Freyhof (2007) recognise Rutilus caspicus (Yakovlev, 1870) as the semi-anadromous species in the Caspian Sea with R. rutilus in rivers and lakes. Curiously, R. caspicus and R. frisii showed low genetic divergence in Ketmaier et al. (2008), a study using DNA but sample sizes were low.

Three "types" of Rutilus rutilus caspicus have been reported from Iran (Annual Report, 1995-1996, Iranian Fisheries Research and Training Organization, Tehran, p. 54-55, 1997). One type lives in the Anzali Mordab and is relatively small, the second is larger and migrates between Anzali and the Kura River of Azerbaijan and the largest migrates between the northern and southern coasts of the Caspian Sea.

Rutilus rutilus schelkovnikovi Derzhavin, 1926 was described from the Karasu which falls into the Aras River 5 km above the mouth of the Zanga River (Razdan River), Echmiadzin District, Armenia, and Rutilus rutilus uzboicus Berg, 1932 from lakes in the Uzboi River Valley (with the type from Lake Yaskhan) in Turkmenistan north of the Iranian border. Rutilus rutilus var. fluviatilis (Jakovlev, 1870) described from the delta of the Volga River is another synonym.

Hybrids with Blicca bjoerkna are reported from the Aras River basin in Armenia, which is shared with Iran. Artificial hybrids with Rutilus frisii kutum and Abramis brama have been bred in Iran (Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 39-40, 1996).

Key characters

This species is distinguished from the related Rutilus frisii by the lower scale count (< 50) and the posterior part of the swimbladder being rounded rather than pointed.

Morphology

Dwarf and large forms are recognised in Dagestan by Shikhshabekov (1969) with different life histories and distinct spawning stocks are known from the north Caspian Sea.

Lateral line scales 39-48, mostly 42-47. Scales have fine circuli around a central focus. Radii may be found on all fields but fish often lack radii on lateral fields. Usually there are very few anterior and posterior radii (e.g. in a fish 150.7 mm standard length there are 2 anterior and 4 posterior radii; but counts are quite variable, at least 7-14 radii total). Scales have a wavy anterior margin and a finely crenulated posterior margin. There is a pelvic axillary scale and a scaled keel behind the pelvic fins. Dorsal fin branched rays 8-12, usually 9 in the Caspian Sea (but see below), after 2-5 unbranched rays, anal rays 8-12, mostly 9-10, after 2-4 unbranched rays, pectoral fin with 14-18 branched rays, and pelvic fin with 7-9 branched rays. Gill rakers 9-17, usually 10-14, short and stubby and reaching the adjacent raker when appressed. Vertebrae 38-43. Lake populations generally have more vertebrae than river populations and the number of vertebrae is genetically fixed, generally following the maternal genotype (Izyumov and Kas'yanov, 1995). Pharyngeal teeth 6-5 or 5-5, rarely 6-6, 4-5, 5-4, or 5-6 with the grinding surface slightly folded to form a long, hollow crown for the largest teeth. There is a rounded and hooked tip to teeth 1-4. The tooth margin in the posteriormost 2 teeth is serrated. The anteriormost 2 teeth are rounded. Teeth are more serrated and hooked in smaller fish. The gut is s-shaped with a small anterior loop. The chromosome number is 2n=50 (Klinkhardt et al., 1995).

Meristics for Iranian specimens:- dorsal fin branched rays 9(22) or 10(18); anal fin branched rays 9(16), 10(23) or 11(1); pectoral fin branched rays 14(1), 15(4), 16(22), 17(12) or 18(1); pelvic fin branched rays 7(4), 8(33) or 9(3); lateral line scales 40(1), 41(4), 42(16), 43(11), 44(5) or 45(3); total gill rakers 11(4), 12(15), 13(18) or 14(3); pharyngeal teeth 6-5(22), 6-6(1), 6-4(1) or 5-5(1); and total vertebrae 37(1), 39(2), 40(9), 41(64) or 42(19).

Sexual dimorphism

In a male caught on 26 April, moderate-sized tubercles are evenly distributed over the top and sides of the head, with fewer tubercles on the undersurface. Scales bear one strong tubercle in the exposed mid-scale, anterior and dorsal scales have smaller tubercles on the scale margin and belly scales have minute tubercles on the margin and exposed scale base. Fine tubercles follow the branching fin rays in single file on all fins, both dorsally and ventrally on the pectoral and pelvic fins, and in 1-2 rows on the last unbranched ray of each fin. Some fish have some scales with 1-3 tubercles, usually 2, one above the other. When there are 3 tubercles these are variably arranged, e.g. in an L-shape or a reversed L. Females also bear tubercles on the head and scales but these are less well-developed and are not present on fins.

Colour

Overall colour is silvery. The back is grey and the sides of the head golden. The belly is pearly-white. The iris is silvery with a black spot above the pupil, some gold on the upper part, and may be orange to red. Pectoral, pelvic and anal fins light grey with a darker band on the edge. The middle of the pelvic and anal fins may be transparent, orange to pink or red. The peritoneum is silvery with moderately dispersed but evident melanophores.

Size

Attains 56.0 cm and 3.13 kg. Larger fish may be hybrids with Rutilus frisii. In Iran during the 1950s, catches were 16-35 cm long (Farid-Pak, no date).

Distribution

Found from the British Isles and France eastwards to Siberia, north of the Alps; in the Black Sea basin but not in Turkey, except the northwestern corner and the Yesilirmak River basin, and on all shores of the Caspian Sea basin including that of Iran, its rivers and bays (Nedoshivin and Il'in, 1929; Berg, 1948-1949; Kozhin, 1957; Riazi, 1996; Abbasi et al., 1999; Kiabi et al., 1999).

Abdoli (2000) has natio knipowitschi in Gorgan Bay and the neighbouring Caspian coast and natio kurensis in the lower Safid River, Anzali Talab and neighbouring Caspian coast to Astara and the middle Aras River.

Rutilus rutilus aralensis Berg, 1916 (but see above on subspecies status) is reported from the Karakum Canal and Kopetdag Reservoir in Turkmenistan (Shakirova nd Sukhanova (1994; Sal'nikov, 1995) and may eventually be found in the Tedzhen (= Hari) River basin of Iran.

Zoogeography

A European and western Asian species with its origins in a Danubian or Sarmatian fauna.

Habitat

This species is eurytopic, living in rivers, streams, lakes, reservoirs, and fresher parts of seas. It occurs in schools close to vegetation in fresh waters.

This species gathers in front of the Volga River delta in the north of the Caspian Sea in large shoals in autumn when water levels decrease. At water temperatures of 5°C they descend to the sea floor and "lie in winter sleep" (Holčík and Skořepa, 1971). It is not known if this occurs off the warmer Iranian shore. Generally the vobla lives in the sea itself and migrates to rivers for spawning. Although most spawning takes place in fresh water, some occurs at a salinity of 6.5‰. This species is the most tolerant of the semi-anadromous fishes, the lethal salinity level being 15-16‰. In the north Caspian Sea, this fish are found mostly shallower than 7.5 m but Knipovich (1921) reports this species at 36.6-53.0 m in the Iranian Caspian Sea.

A mass migration of the Kura vobla (natio kurensis) into the Anzali Mordab occurred between 21 and 28 February 1915, only to disappear after 4 April. Riazi (1996) reports that this species migrates into the Siah-Keshim Protected Region of the Anzali Mordab. Males migrate into the Anzali and Gomishan wetlands of Iran earlier than females and the larger fish enter first (Naddafi et al., 2005).

Natio knipowitschi lives in the sea in Gorgan and Gasan-kuli bays and enters rivers there. Savenkova (1994) reported as many as 2000 million juveniles descending the Atrak River in May and June in high productivity years and as few as 83 million in low productivity years. The sea in July and August is as warm as 28-30°C off the Iran-Turkmenistan border and the juvenile fish from the Atrak descend to 11-12 m. Savenkova (1994) gives further details of movements in relation to temperature, e.g. when waters cool to 8-12°C off Turkmenistan, young fish move to warmer Iranian waters to overwinter. ERM-Lahmeyer International GmbH, DHI Water & Environment and GOPA Consultants (2001a) report that the Gomishan Lagoon at 37º11N, 53º57'E is an important staging area for this species.

Age and growth

Generally growth is a function of type of feeding (a diet favouring molluscs gives higher growth than herbivory) and temperature. High growth rate is shown by semi-migratory fish but is also shared by some populations resident in lakes and rivers. Kas'yanov et al. ( 1995) review the complex relationships of environmental factors affecting growth in this species.

A life span up to 19 years has been reported although most fish are 10 years or less. Growth is fastest in the second and third years of life when most fish mature sexually (Strubalina and Chernyavskiy, 1992).

Some fish may mature at one year of age and a length of 9-11 cm in the Atrak River population (Petr, 1987). Growth in the Atrak population juveniles is heavily dependent on temperature regimes in the sea, the timing of descent to it, and stock abundance. Fish which descend earlier because of low flow and have a longer feeding season, find a favourable, warm regime and few competitors will have faster growth (Savenkova, 1994).

Young-of-the-year Turkmen fish reach 4.9-5.8 cm in July while in the northern Caspian they only attain 3.3-4.1 cm, although the latter have a higher condition factor, possibly because of higher benthic biomass and heterogeneity (Savenkova, 1994). The dwarf form in Dagestan is 11 cm and 31 g at 3 years of age while the large form is 20.3 cm and 130 g. Both are sexually mature (Shikhshabekov, 1969).

The commercial catch in Iran during 1971/72 was 3-6 years old, 20.4-26.0 cm long and weighed 146-300 g (Razivi et al., 1972). The average weight in 1995 was 200-300 g (Abzeeyan, Tehran, 5(9):V, 1995).

In Gorgan Bay, the age composition is from 1+ to 6+ with age group 3+ and 4+ each accounting for about 30%. Growth rate for all south Caspian populations is higher than the north Caspian populations, two-year-olds being the same length as three-year-olds. Average weight is 247 g with larger fish at 30 cm weighing 583 g. Numerous fish entered Gorgan Bay in mid-January in the early years of this century and by the end of that month ascended all the tributary rivers, in particular the Qareh Su (= Karasu).

Growth rate in the Anzali Mordab was higher than in the Gomishan Wetland of the southeastern Caspian Sea, limited by higher salinity in the latter (Naddafi et al., 2002a, 2002b, 2005; Paghe et al., 2005). The population density was higher in Gomishan as Anzali suffers more from pollution, overfishing, illegal fishing and other human impacts. Mean condition factors for the Gomishan population were 1.84 for males and 2.09 for females while in the Anzali population these were 2.03 and 2.2 respectively. The most abundant age groups in the commercial fishery of both areas were 3+, 4+ and 5+. Maximum age at Gomishan was 8+ years. Growth was rapid in the first year with a sharp decline in the second year and a steady, less rapid decline in subsequent years. Females were longer than males at in each age class. Sex ratios were did not differ significantly from 1:1. The Anzali population had longer body lengths than fish from other studies, perhaps due to a longer growing season and higher water temperatures in the south Caspian Sea, better food supply and genetic factors.

Food

The young feed first on phytoplankton and then switch to zooplankton and benthic insect larvae with growth. Adults eat benthic animals and plants with molluscs an important diet item where present as in the southern Caspian Sea. A mollusc diet occasions significant wear on the pharyngeal teeth (Holčík and Skořepa, 1971). There has been a long-term variation in the diet of roach in the Caspian Sea, molluscs declining and being replaced by crustaceans and plants (Strubalina and Chernyavskiy, 1992). Plant foods in Europe include water thyme, milfoil, duckweed, stonewart and algae scraped from reeds and rocks (Muus and Dahlstrøm, 1999). There is little feeding during spawning in rivers. Young-of-the-year in the Atrak delta feed on the polychaete worm Nereis (up to 89.5%) and with growth the mollusc Cerastoderma lamarckii (up to 70.8%), and to a lesser extent the exotic mollusc Abra ovata (5.2%) since this abundant species is partially buried and is not readily available. Polychaetes and molluscs have high biomasses in the silty sediments of around 30-50 g/sq m. Sometimes protozoa and crustaceans figure prominently in the summer diet but diet varies considerably between months and years (Savenkova, 1994).

Reproduction

A spawning migration runs up rivers in the Caspian Sea basin in spring, the larger fish first. Spawning grounds are mostly flooded meadows that warm up easily (Caspian Sea Biodiversity Database, www.caspianenvironment.org). Immature fish only migrate in autumn after their gonads have matured, spending the summer in deeper water. The fish return to the sea after spawning in the same sequence as the upriver migration. Young remain in the river until early summer when they too go down to the sea. However some populations of this species are exclusively freshwater and do not migrate.

Spawning is continuous and takes about 5-6 hours. Fecundity reaches about 202,000 eggs and egg diameters 1.6 mm. In the Volga delta this species spawns in the same areas as Abramis brama. However, it spawns earlier at temperatures of 7-16°C on last years dead vegetation while the bream spawns later at 15-23°C on new growth. Spawning can occur on stony bottoms in the absence of vegetation. Spawning is a noisy process which can be heard from some distance. Each female is accompanied by 2-3 males who push on her belly, sometimes lifting her part way out of the water, as they swim in circles. With many fish engaged in this process, the water foams. The fish gradually approach the shore where the eggs are shed in water only 5-20 cm deep as the female is squeezed by two males. Other males take part in the process, as many as 10 at a time. Incubation takes 4 days at 17-20°C. Young actively move downstream once they reach 20-25 mm.

The breeding migration in the Atrak River starts in January-February and continues until April, the fish traveling 70-80 km upriver (Petr, 1987). Nümann (1969) states that the spawning migration takes place from November to December in Iranian waters. The migration in the Atrak peaks at 10-12°C and lasts 20-25 days. Spawning peaks at 15-18°C in March on the lower Atrak floodplains (in 1908 spawning in the lower Atrak was observed in mid-April). In the absence of flooding, no spawning occurs. After spawning, females migrate back to the sea first. In the Qareh Su of Gorgan Bay, spawning took place 20-25 km upriver from the mouth in 1907 and 1908. It started in mid-February, peaked in the second half of February and the first half of March, and ended between 20 and 30 March. Males with running milt first appeared between 14 and 20 February in 1915 and ripe females a week later. Mass spawning took place at the beginning of March (Nedoshivin and Il'in, 1929). The main spawning rivers in Iran are the Qareh Su, Goharbaran, Larim, Siah-Darvishan and Shafa River (Iranian Fisheries Research and Training Organization Newsletter, 9:5, 1995).

Fecundity in the Gomishan Wetland (4262-98,804 eggs) was significantly higher than in the Anzali Mordab (6035-32,141) for a given body size. Egg diameters ranged from 0.9 to 1.45 mm and were not significantly different between the two wetlands. The peaks for the gonadosomatic index (GSI) curves were early March for males and mid-March to early April for females. The GSI was higher in Gomishan (Naddafi et al., 2005).

Parasites and predators

The main predator of this species in the north Caspian Sea is Sander lucioperca, accounting for 65% of its food. Silurus glanis and Esox lucius and various birds such as the pelican are also important predators (Kushnarenko, 1978). The Caspian seal, Pusa caspica, is a significant predator on this species (Krylov, 1984). On the spawning grounds, this fish can be picked up by hand and falls easy prey to birds and other predators.

Mokhayer (1976b) records the cestode larva Diagramma. Jalali and Molnár (1990a) record the monogenean Dactylogyrus turaliensis from this species in the Safid River. Masoumian et al. (2002) recorded parasites from fish captured form the coast of the southeast Caspian Sea as Anisakis larvae, Aspidogaster limacoides, Bothriocephalus gowkongensis, Dactylogyrus turaliensis and Diplostomum spathaceum. Youssefi et al. (2005) record the pleroceroid larvae of Ligula intestinalis in fish from the Aras Dam. A toxin produced by this parasite can cause infertility and weight loss in the fish and can be harmful to humans. Khara et al. (2006a) record the eye fluke Diplostomum spathaceum for this fish in the Amirkalayeh Wetland in Gilan, having the highest infection rate in 7 species examined. Khara et al. (2006b) record the cestode Caryophyllaeus fimbriceps from this species in the Boojagh Wetland of the Caspian coast.

Economic importance

This species is one of the most important commercial species in the Caspian Sea with a catch in the north Caspian and lower reaches of rivers in 1930 being 2,591,000 centners (15,000,000,000 fish) or 45.5% of the total catch of fishes in the Caspian Sea or nearly 20% of the whole Soviet fish catch. Archaeological studies on the eastern Caspian shore in the former Soviet Union has shown this species to have been fished for 5000 years ago (Tsepkin, 1986). It is a significant food for the sturgeon Huso huso (Keyvanshokooh and Kalbassi, 2006).

The main fishing season in Iran is February and March (Farid-Pak, no date). Nevraev (1929) gives catches for various fishing regions in Iran in the early twentieth century. For the Anzali region from 1914-1915 to 1917-1918 the catch was 6000 to 244,800 individuals (the catch declining steadily over this period), and in Astrabad (= Gorgan) region from 1909-1910 to 1912-1913 the catch was 8,348,800 to 21,790,000 individuals. The catch in Iran from 1956/1957 to 1961/1962 varied from 2989 kg to 1,092,719 kg (Vladykov, 1964), from 1965/66 to 1968/69 it varied from 32 to 74 tonnes (36, 74, 32 and 35 t respectively) (Andersskog, 1970) and from 1963 to 1967 from 2.4 to 47.0 t (23.1, 2.4, 3.7, 30.7, 47.0 t respectively) (RaLonde and Walczak, 1970b). Vladykov (1964) reports catches from the Anzali region for the period 1933/1934 to 1961/1962 to vary from 57 to 716,974 kg, with none reported in some years. Holčík and Oláh (1992) report an annual catch in the Anzali Mordab from 1932-1964 of 0.8-449.7 t. The total catch in 1989 (or 1990) was only 26 t but increased to 120 t in 1995 because of the rising Caspian Sea level which provided more spawning grounds (Abzeeyan, Tehran, 5(9):V, 1995; Iranian Fisheries Research and Training Organization Newsletter, 9:5, 1995). Conflicting values from reports where they overlap are typical of fisheries statistics from Iran which can only be taken as general trends rather than absolute values (see below also). The Food and Agriculture Organization, Rome gave the catch for Rutilus spp., presumably this species, for the years 1980-1985 as 0, 0, 0, 121, 347, and 350 t respectively. The catch in Gorgan Bay 20-30 years ago according to Petr (1987) was about 4000 tonnes per year but is now negligible. This figure conflicts with the ones cited above. In the 1914-1915 fishing season in Gorgan (= Astrabad) Bay, 24.2 million fish were taken, of which 21.9 million were taken in the Qareh Su. Petr (1987) cites catches of 830 t in 1978 for the southeastern Caspian Sea and for 1979 only 120 t, probably including Soviet catches. Recruitment falls when the Atrak River, a major spawning ground, fails to flood adequately. The dwarf form in Dagestan is of no commercial value and may have arisen with changing environmental conditions (Shikhshabekov, 1969).

Moini et al. (2005) have investigated the production of marinades using this fish using various recipes, examining their chemical composition over time and their comparative tastes.

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in aquaculture and as food, in sport and in textbooks and because it has been introduced outside its natural range. Everard (2006) is one book of many describing the biology and importance of the roach.

Conservation

Holčík and Oláh (1992) report a catch of only 5 kg in the Anzali Mordab for 1990. This species has been cultured at "Sad-e Sangar" Fish Farm near Rasht (3 million larvae a year), at Astara and at Anzali on the Caspian among other localities (Mokhayer, 1972b). In 1999-2000, 30 million juveniles were released into the Caspian Sea (Iranian Fisheries Research Organization Newsletter, 23:4, 2000). Noroozi et al. (2006) detail capture of brood stock from the Gorgan River estuary for release into 2 ha earthen ponds enriched with manure and fertiliser. The optimum temperature for spawning brood stocks is 12-17ºC, found in mid-March to late April. Pine branches were placed in the ponds as spawning sites. Ponds were stocked with 700 females averaging 150 g and 350 males averaging 100 g. Eggs hatched on the sixth day. Larvae were fed on natural zooplankton (Rotatoria and Daphnia) and artificial food.

The Atrak River population can be conserved by regulating the discharge so that the spawning grounds flood at the appropriate time (Petr, 1987). Beach seines with mesh sizes of 28 mm, 33 mm and 36 mm caught fish of ages 2.8 years, 2.9 years and 3.3 years respectively. This data indicates that 58.3%, 50.5% and 3.6% of the total catch is non-standard respectively. Seines with a cod end of at least 36 mm are recommended for stock protection (Iranian Fisheries Research and Training Organization Newsletter, 6:8, 1994). Catches are increasing with the rise of the Caspian Sea water level as noted above, indicating how natural events beyond human control can have a significant effect on stocks.

There have been few studies on the effects of environmental factors on this species in Iran. Gerve'ei et al. (2008) determined the LC₅₀ and gill tissue lesions caused by aluminium sulphate under laboratory conditions and Keramati et al. (2010) studied the effect of the agricultural

organophosphate diazinon on enzyme activity, also under laboratory conditions.

Further work

As an important food fish, this species should be effectively managed and conserved.

Sources

Wossugh-Zamani (1991d) and Akbari-Pasand (1996) give accounts of this species in Farsi.

Iranian material: CMNFI 1970-0509, 3, 73.4-82.9 mm standard length, Gilan, Safid River at Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1970-0510, 4, 58.6-83.8 mm standard length, Gilan, Golshan River (37º26'N, 49º40'E); CMNFI 1970-0531, 14, 65.5-115.8 mm standard length, Mazandaran, Larim River (36º46'N, 52º56'E); CMNFI 1970-0532, 1, 74.9 mm standard length, Gilan, Caspian Sea near Bandar-e Anzali (37º28'N, 49º27'E); CMNFI 1970-0543A, 1, 119.4 mm standard length, Gilan, Caspian Sea at Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1970-0549, 1, 109.7 mm standard length, Mazandaran, Qareh Su near Alm Imamzadeh (no other locality data); CMNFI 1970-0563, 5, 40.3-112.2 mm standard length, Gilan Caspian Sea, Kazian Beach (ca. 37º29'N, ca. 49º29'E): CMNFI 1970-0583, 6, 35.7-84.9 mm standard length, Gilan, Nahang Roga River (37º28'N, 49º28'E); CMNFI 1979-0431, 1, 157.0 mm standard length, Mazandaran, fish bazaar at Now Shahr (no other locality data); CMNFI 1979-0788, 2, 64.9-66.6 mm standard length, Mazandaran, Gorgan River at Khadje Nafas (37º00'N, 54º07'E); CMNFI 1980-0126, 4, 162.2-178.3 mm standard length, Gilan, Caspian Sea near Bandar-e Anzali (37º28'N, 49º27'E); CMNFI 1980-0136, 4, 72.1-109.7 mm standard length, Mazandaran, Fereydun Kenar River estuary (36º41'N, 52º29'E); CMNFI 1980-0148, 1, 85.4 mm standard length, Gilan, Pir Bazar Roga River (37º21'N, 49º33'E); CMNFI 1980-0157, 2, 73.1-132.0 mm standard length, Mazandaran, Gorgan River estuary (36º59'N, 53º59'30"E); CMNFI 1980-0905, 9, 98.5-203.0 mm standard length, Mazandaran, Gorgan River at Khadje Nafas (37º00'N, 54º07'E).

Genus Scardinius
Bonaparte, 1837

Howes (1981) placed this genus in Rutilus Rafinesque, 1820 on osteological grounds. Bogutskaya (1988) disagrees. Scardinius is usually separated from Rutilus by having 2, as opposed to 1, rows of pharyngeal teeth and a ventral keel on the body. Howes (1981) considers pharyngeal teeth to be of only species importance and the keel is variously developed in Rutilus.

The genus contains perhaps 6 species and is found from the British Isles and the Iberian Peninsula throughout Europe to the Caspian and Aral Sea basins, with one species in Iran.

The genus is characterised by a pharyngeal tooth count of 3,5-5,3 with major row crowns laterally compressed and bearing 5-8 serrations, scales moderate in size in a complete lateral line, few, short gill rakers, a keel on the belly behind the pelvic fins covered with scales, a short gut and light peritoneum, dorsal and anal fins of moderate length, dorsal fin origin well behind the origin of the pelvic fins, terminal and oblique mouth,

Scardinius erythrophthalmus
(Linnaeus, 1758)

Common names

sorkh pareh or sorkh par (= red fin), mahi sorkh baleh, ماهي چشم قرمز (= mahi chesm ghermez).

[giziluzkac in Azerbaijan; krasnoperka in Russian; rudd, redeye, redfin, pearl roach].

Systematics

See above under the genus. Cyprinus Erythrophthalmus was originally described from northern Europe. This species is widely known to spawn with other cyprinid fishes making hybrids a common occurrence. Some Iranian material appears to be hybrids of this species and another, unknown parental species but this has not been investigated.

Key characters

This species is often confused with Rutilus rutilus and R. caspicus but can be distinguished by the posterior position of the dorsal fin (in relation to the pelvic fins), the belly keel, the upturned mouth, and the serrated pharyngeal teeth in 2 rows.

Morphology

Dorsal fin with 2-4 unbranched and 7-10, usually 8 branched rays, anal fin with 3-4 unbranched followed by 9-13, usually 11, branched rays (Abdurakhmanov (1962) initially gives 8-9 anal fin branched rays for Azerbaijan fish but this may be a misprint as a subsequent table lists 9-11 rays), pectoral fin with 13-16 branched rays, and pelvic fin with 7-9 branched rays. Lateral line scales 36-45. There is a pelvic axillary scale. Scales are squarish in shape, with sharp dorsal and ventral anterior corners, a wavy anterior margin, central focus, fine circuli which are coarser on the posterior field, and very few anterior and posterior radii (e.g. 2 anterior and 3 posterior primary radii reaching the focus from the margin). The scale margin is indented where radii terminate and the thick posterior radii are visible on the flank. Gill rakers short and widely spaced, touching the adjacent one when appressed, and numbering 8-13. Vertebrae 37-42. Pharyngeal teeth mostly 3,5-5,3 with variants 3,5-5,2, 3,5-5,1, 3,5-4,3, 2,5-5,3 and 2,5-5,2, narrow and elongate, slightly hooked and with about 5-8 strong serrations on each tooth. There is a strongly-developed, scaled keel between the vent and the pelvic fin base. The gut is s-shaped with an anterior loop. The chromosome number is 2n=48-50 (Klinkhardt et al., 1995).

A single Iranian specimen had the following meristics:- dorsal fin branched rays 8; anal fin branched rays 10; pectoral fin branched rays 16; pelvic fin branched rays 8; lateral line scales 37; total gill rakers 10; pharyngeal teeth 3,5-5,3, and total vertebrae 39..

Sexual dimorphism

Males develop breeding tubercles on the head and body.

Colour

The back is blue-black to greenish- or olive-brown, the flanks are brassy and the belly silvery-white. Upper flank scales have dark bases. The tips of the caudal, anal and pelvic fins are a bright, blood red in the spawning season and the dorsal fin is black proximally and red distally. The iris is yellow to orange, or gold, with a red spot at the top. Peritoneum silvery with scattered melanophores. Young are much less brightly coloured than adults.

Size

Attains 62.0 cm and 3.01 kg.

Distribution

Found from the British Isles and north of the Pyrenees east to the Caspian and Aral sea basins. It is recorded from the Lenkoran in Azerbaijan and, in Iran, from the Anzali Mordab, Bojaagh wetland, Safid River, Haraz River and Babol River (Derzhavin, 1934; Holčík and Oláh, 1992; Abbasi et al., 1999; Abdoli, 2000; Jolodar and Abdoli, 2004; K. Abbasi, pers. comm., 21 February 2005).

Zoogeography

This species is part of a European and West Asian fauna whose origins may lie in a Danubian or Sarmatian fauna.

Habitat

Rudd can favour heavily overgrown areas (Shikhshabekov, 1979) and are generally found in shallow warm lakes or slow moving rivers. They are usually inhabitants of midwater or near the surface but they overwinter in deep water. They are regarded as fairly hardy and are adapted to eutrophic, and presumably therefore, polluted waters. In Iranian waters its density is highest in the Anzali Mordab (K. Abbasi, pers. comm., 21 February 2005).

Age and growth

Sexual maturity is attained at 3-4 years in Dagestan at lengths of 17-29 cm and 80-530 g. A stunted form is found in rice paddies at an age of 2 years, 7.5-11.0 cm and 10-23 g (Shikhshabekov, 1979). Elsewhere life span is at least 17 years.

Food

Food is aquatic macrophytes as well as insect larvae, crustaceans, molluscs and more rarely fish eggs and fry. The young feed on zooplankton.

Reproduction

Spawning takes place at water temperatures of at least 18-20°C in June-July in Dagestan. Each female can be accompanied by two males, one on each side. Two batches of eggs may be spawned in this period (Shikhshabekov, 1979). In the Volga Delta, spawning takes place from April until the end of June. Eggs attach to water plants. The young remain attached to vegetation until the yolk-sac is absorbed. Fecundity is up to 232,000 eggs with a diameter of 1.5 mm. Hatching takes 3 days at 20-22°C.

Parasites and predators

Masoumian and Pazooki (1998) surveyed myxosporeans in this species in Gilan and Mazandaran provinces, finding Myxobolus pfeifferi. Sattari et al. (2004) record the nematode Raphidascaroides acus larvae from this species in Gilan. Khara et al. (2006b) record the nematode Raphidascaris acus from this species in the Boojagh Wetland of the Caspian coast. The Caspian seal, Pusa caspica, is a predator on this species (Krylov, 1984).

Economic importance

Holčík and Oláh (1992) report a catch of 98 kg in the Anzali Mordab in 1990.

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in aquaria and aquaculture, as food, in sport and in textbooks and because it has been introduced outside its natural range.

Conservation

Lelek (1987) classifies this species as vulnerable in Europe. Kiabi et al. (1999) consider this species to be conservation dependent in the south Caspian Sea basin according to IUCN criteria. Criteria include sport fishing, medium numbers, habitat destruction, limited range (less than 25% of water bodies), absent in other water bodies in Iran, present outside the Caspian Sea basin. Near threatened in Turkey (Fricke et al., 2007).

Further work

The distribution and biology of this species and its potential for hybridisation with other cyprinids requires study in Iran.

Sources

Iranian material: Uncatalogued, 1, 103.7 mm standard length, Gilan, swamp near Hendeh Khaleh (37°23'N, 49°28'E).

Genus Schizocypris
Regan, 1914

This genus of medium-sized snow trouts contains only 2 species found in Pakistan, Afghanistan and Iran. Coad and Keyzer-de Ville (2005) revised the genus.

It is characterised by a rounded and moderately elongate body, scales small but larger near the shoulder region, belly scaleless, a wide and transverse mouth with the snout projecting, no barbels or barbels vestigial, pharyngeal teeth with a flat tip unlike Schizothorax and a formula of 2,3,4-4,3,2 rather than 2,3,5-5,3,2 as in Schizothorax, dorsal and anal fins short but 6 branched anal fin rays not 5 as in related genera in the same area, dorsal fin with a strongly serrated spine, scales in the vent region are split and enlarged to flank the urogenital region, and radii are on all scale fields.

Rainboth (1981) includes Capoeta trutta, Capoeta fusca and Capoeta nudiventris (= C. fusca) in this genus but this is incorrect. These species show some enlargement of scales around the anus and anal fin region but it is not as marked and definitive as in true Schizocypris and other characters of the genus are absent.

Schizocypris altidorsalis
Bianco and Banarescu, 1982

Common names

gorgak (= small wolf), anjak or khaju (A. A. Pasand, pers. comm., 5 November 2000 but see under Schizothorax zarudnyi and S. pelzami).

Systematics

Schizocypris brucei, non Regan, 1914 (Annandale and Hora, 1920) is a synonym. Schizocypris brucei Regan, 1914 is thus restricted to the Gomal River drainage in the Indus River basin of Pakistan (originally described from the Wana Toi, a Gomal River tributary). A third species in the genus Schizocypris is S. ladigesi Karaman, 1969 from the Kankai River, also in the Indus River basin.

Kähsbauer (1964) reports a hybrid between Schizothorax schumacheri and Schizopygopsis stoliczkae from Sistan which may in fact be this species.

The holotype of Schizocypris altidorsalis is in the Istituto di Zoologia dell'Universitá di L'Aquila, Italy under IZA 8169 and is 73.7 mm standard length (my measurment) (Bianco and Banarescu, 1982). The type locality is "Nahr-Taheri near Zabol, Seistan". Paratypes include 5 fish from the type locality under IZA 7841, 35-65 mm standard length (4 fish seen by me, 35.4-62.0 mm standard length) with further specimens in the Institutul de Stiinte Biologice, Bucuresti, Romania (ISBB 3136). Three paratypes, 68-73 mm standard length, from "Rud-Sistan, 8 km from Zabol, Seistan" are under IZA 7844 (69.9-74.3 mm standard length measured by me) with further specimens under ISSB 3137. Two paratypes from the Nahr Taheri are in the Zoologischen Instituts und Zoologischen Museums der Universität Hamburg (ZMH 6091, 77.2-81.9 mm standard length) (Wilkens and Dohse, 1993; examined by me), 2 paratypes are in the American Museum of Natural History, New York (AMNH 40952), 1 paratype is in the Muséum national d'Histoire naturelle, Paris (1982-1018), 1 paratype is in the United States National Museum, Washington (USNM 227928), 2 paratypes are in the Academy of Natural Sciences, Philadelphia (ANSP 150977), and 2 paratypes are in CMNFI 1982-0368 (formerly IZA 7841).

Syntypes of Schizocypris brucei are in the Natural History Museum, London under BM(NH) 1913.4.15:100-109 (10 fish) and in the Zoological Survey of India, Calcutta under ZSI F9832/1 (1) (Menon and Yazdani, 1968; Eschmeyer et al., 1996).

Berg (1949) advanced the possibility that this species is the juvenile of Schizothorax zarudnyi since barbels and scale cover develop with age. However the tooth formula is very distinctive as is the anal fin branched ray count and lack of barbel development at all sizes in S. altidorsalis (Coad and Keyzer-de Ville, 2005).

Key characters

This species is characterised by a very high dorsal fin with a strongly denticulated spine. The spine is longer than the head and the denticles easily catch the skin when the fish are handled. This is particularly true of small fish, larger ones are not so snaggly. Bianco and Banarescu (1982) give values for spine length of 24.4-29.8% of standard length in altidorsalis, 19.4-20.4% in brucei, and 14.7-19.5% in ladigesi. For 35 altidorsalis 66.1-175.1 mm standard length examined here, spine length is 23.3-31.3% of standard length and 14.0-18.5% for 20 brucei 102.3-170.8 mm standard length. Dorsal spine length in head length is 0.7-0.9, mean 0.9 for altidorsalis, 1.2-1.6, mean 1.4 for brucei (see also below). Scales in the lateral line are 87-96 (brucei has 74-81 and ladigesi 78-88) according to Bianco and Banarescu (1982). Specimens examined by me have lateral line counts of 81-95, mean 87.4 for 60 altidorsalis and 73-91, mean 79.6 for 56 brucei, showing some overlap but scales are definitely smaller on average in altidorsalis.

Bianco and Banarescu (1982) describe the body as mostly scaled except on the anterior part of the breast (scaled on the mid-line of the back in front of the dorsal fin as in ladigesi, naked in brucei), and scales embedded on most of the body except the caudal peduncle and the lateral line; but see below.

Morphology

Dorsal fin unbranched rays 4, branched rays 7-8, anal fin unbranched rays 3, branched rays 5-6, pectoral fin rays 14-20, pelvic fin rays 7-9, lateral line scales 82-96, total gill rakers 24-30, reaching the third raker below when appressed in large fish but only one raker below in small fish, pharyngeal teeth usually 2,3,4-4,3,2(16) with variants 2,3,4-4,3,3(1), 2,3,5-4,3,2(2) and 2,3,5-4,4,2(1), and total vertebrae 43-45. Since spines are often broken off, the height of the dorsal fin can be measured as the longest branched ray. For this species it is 16.5-28.8% standard length (mean 24.3, 60 fish) while in S. brucei, the taxon fish in Sistan were formerly assigned to, it is 11.6-20.0% (mean 16.8, 56 fish), clearly distinguishing these species.

Scales are regularly arranged over most of the body. In some fish, scales near the tail are difficult to distinguish. Shoulder scales are moderately large anteriorly above the lateral line and decrease in size posteriorly. The back is naked in a narrow band for a short distance anterior to the dorsal fin (not so according to Bianco and Banarescu (1982) but visible in fish examined by me). Flank scales are small but those at the dorsal fin base are a little larger. Lateral line scales are larger than those on the flank but only on the anterior lateral line. The breast is scaleless according to Bianco and Banarescu (1982) but is scaled on the breast in large specimens and some small ones too. There is a pelvic axillary scale. The scale focus is anterior with radii on all fields and in overall shape is oval to rounded. Radii are found on all fields but are few in number (15).

The mouth is inferior, transverse, and slightly arched. The lower lip is developed only laterally. The lower jaw is covered by a horny sheath in some specimens, lost in others. There are usually no barbels (n = 105) although one fish, 70.1 mm standard length, had a minute pair of barbels hidden in the lip grooves and two other fish of similar size had respectively a single right and a single left minute barbel. S. brucei has small but protruding barbels on both sides in 15 fish, a left barbel only in 3 fish, a right barbel only in 5 fish and no barbels in 8 fish.

The gut is very elongate and complexly coiled. The anterior main row pharyngeal tooth is peg-like while the rest are spatulate with a deep central groove and crowns flared on each side of the groove.

Meristics for Iranian specimens are:- dorsal fin branched rays 7(2) or 8(38); branched anal fin rays 5(1) or 6(39); pectoral fin branched rays 14(1), 16(11), 17(23), 18(4) or 20(1); pelvic fin branched rays 7(1), 8(35) or 9(4); lateral line scales 81(1), 82(2), 83(3), 84(4), 85(6), 86(8), 87(10), 88(7), 89(5), 90(4), 91(3), 92(3), 93(3) or 95(1); total gill rakers 24(1), 25(4), 26(5), 27(10), 28(12), 29(6) or 30(2); pharyngeal teeth 2,3,4-4,3,2(16), 2,3,4-4,3,3(1), 2,3,5-4,3,2(2), or 2,3,5-4,4,2(1); and total vertebrae 43(16), 44(33) or 45(11).

Sexual dimorphism

Unknown.

Colour

The back is bluish and the flanks and belly are silvery. The flanks may have a few small to numerous black spots.

Size

Reaches 17.5 cm standard length.

Distribution

This species is endemic to the Sistan basin of Iran, and presumably adjacent Afghanistan (Bianco and Banarescu, 1982; J. Holčík, in litt., 1996). The distribution of this species (as S. brucei) in Khorasan and Gorgan reported by Wossughi (1978) is incorrect.

Zoogeography

A relative of other schizothroacine species found along the mountain chain from Iran to China. See also the genus Schizothorax.

Habitat

Occurs in a wide range of habitats from ditches to hamuns. Reported from pools in dry river beds and still, reedy channels in Sistan. E. Penning (pers. comm., 28 July 2005) states that this was the dominant fish species in Hamun-e Puzak and Hamun-e Saberi in April 2005 after a dry period when the hamuns flooded at the end of February. The fish enter the flooding hamuns from the upstream parts of rivers. In July, water levels fell from1-2 m to less than 1 m and this species was absent, presumably having returned to the more permanent rivers. They were also observed swimming up a fish staircase at the Sistan Dam.

Age and growth

Unknown.

Food

The principal food is aufwuchs and detritus as evidenced by the sectorial mouth and elongate gut. Gut contents are a fine mush. E. Penning (pers. comm., 28 July 2005) observed filamentous algae in the gut.

Reproduction

Most fish examined by me were young and no data on reproduction is available. E. Penning (pers. comm., 28 July 2005) noted that fish at 20-30 cm caught in April 2005 had eggs 1 mm in diameter and were approaching or ready for spawning.

Parasites and predators

Jalali et al. (1995) describe a new species of monogenean, Dactylogyrus schizocypris, from fish taken in the Chahnimeh water reservoir near the Hamun Lake in Sistan.

Economic importance

Individual fishermen caught 5-10 kg per day of this species in 2005 although they considered individuals as small and catches very low. Fish were 10-20 cm long with some larger ones at 20-30 cm (E. Penning, pers. comm., 28 July 2005).

Conservation

Further knowledge of biology is needed to assess conservation status.

Further work

The biology of this species needs study.

Sources

Type material: See above, Schizocypris altidorsalis (IZA 8169, IZA 7841, IZA 7844, ZMH 6091 and CMNFI 1982-0368).

Iranian material: CMNFI 1979-0072, 2, 120.8-162.9 mm standard length, Sistan, river near Zabol (ca. 30º58'N, ca. 61º28'E); CMNFI 1979-0223, 1, 19.9 mm standard length, Sistan, ditch south of Lutak (30º45'N, 61º24'E); CMNFI 1979-0224, 13, 48.2-77.5 mm standard length, Sistan, Hirmand River effluent (30º53'N, 61º27'E); CMNFI 1979-0225, 1, 147.7 mm standard length, Sistan, Hirmand River effluent (30º58'N, 61º28'E); CMNFI 1979-0226, 107, 60.0-82.5 mm standard length, Sistan, pool east of Kuh-e Khajeh (30º57'N, 61º17'E); CMNFI 1979-0228, 16, 18.1-73.8 mm standard length, Sistan, ditch 1 km from Zabol (31º02'N, 61º31'E); CMNFI 1979-0229, 11, 61.0-84.9 mm standard length, Sistan, ditch 5 km from Zabol (31º03'N, 61º33'E); CMNFI 1979-0231, 2, 17.5-19.9 mm standard length, Sistan, irrigation jube 3 km from Zabol (31º01'N, 61º32'E); CMNFI 1979-0232, 23, 41.8-78.8 mm standard length, Sistan, irrigation ditch 11 km from Zabol (30º58'N, 61º36'E); CMNFI 1979-0233, 2, 66.0-71.5 mm standard length, Sistan, irrigation ditch 15 km from Zabol (ca. 30º57'N, ca. 61º38'E); CMNFI 1979-0234, 15, 13.9-82.4 mm standard length, Sistan effluent of the Hirmand River near Zahak (30º54'N, 61º40'E); CMNFI 1979-0236, 1, 14.4 mm standard length, Sistan, irrigation ditch 27 km from Zabol (ca. 30º52'N, ca. 61º22'E); CMNFI 1979-0237, 5, 21.4-72.1 mm standard length, Sistan, irrigation ditch 18 km south of Zabol (30º53'N, 61º27'E).

Comparative material: USNM 182276, 5, 95.7-153.1 mm standard length, Afghanistan, Arghandab Reservoir (ca. 31º51'N, 65º55'E); USNM 182277, 1, 159.7 mm standard length, Afghanistan, Arghandab River at Kandahar (ca. 31º35'N, 65º45'E); USNM 182281, 3, 71.2-79.9 mm standard length, Afghanistan, Lashka-dah, Helmand River (ca. 31º35'N, ca. 64º21'E); USNM 182282, 5, 142.1-159.4 mm standard length, Afghanistan, Laskha-dah area (ca. 31º35'N, ca. 64º21'E); ZMUC 261629-34, 6, 131.4-203.7 mm standard length, Afghanistan, Sistan, Feyzabad (31º28'N, 61º31'E).

Material of Schizocypris brucei and S. ladigesi, nominal species not found in Iran, are listed in Coad and Keyzer-de Ville (2005)

Genus Schizopygopsis
Steindachner, 1866

This genus contains about 8 species distributed from Iran along the Himalayas to China. There is a single species in Iran.

The osmans are characterised by an elongate and almost cylindrical body, almost scaleless, scales being restricted to the complete lateral line, the flank above the pectoral fin and as enlarged scales around the vent and anal fin base, the mouth is terminal to inferior, the lower jaw is sometimes covered by a horny sheath, lips are present but may only be developed at the mouth corners, no barbels, pharyngeal teeth in 2 rows, spatulate or cochleariform, short dorsal and anal fins, dorsal fin last unbranched ray thickened (but not strongly) and serrated or denticulate (denticles lost with age), peritoneum black, and gut very elongate.

Schizopygopsis stoliczkai
Steindachner, 1866

Common names

kopur-e barfi (= snow trout).

[lozhnyi osman or false osman in Russian; Pamir snowcarp].

Systematics

The type locality of this species is a stream near Hanle Monastery, Ladakh, India. Syntypes are in the Naturhistorisches Museum Wien under NMW 9255 (1), NMW 9256 (1), NMW 51472 (9) and NMW 51473 (2) (Eschmeyer et al., 1996; 1997 Vienna card catalogue).

The species name is spelt Stoličkai by Steindachner (1866) which becomes correctly stoliczkai since accents are not used in Latin nor capitals for the scientific species name. It is often spelt stoliczkae in the general literature.

Schizopygopsis stoliczkae infraspecies sewerzowi Herzenstein, 1890, originally described from the Bulun Kul and Karasu, Amu Darya basin in the Pamir Mountains has no taxonomic validity (it occurs together with the typical form in the upper Amu Darya) but it was used to characterise the form in the Sistan basin of Iran (Berg, 1948-1949). However, Annandale and Hora (1920) report both this and the stoliczkae form, without intermediates, in Sistan. The sewerzowi form differs from the typical form by having a larger eye (1.2-1.3 times in interorbital width as opposed to 1.5-1.7 times and 3.8-4.4 times in head length as opposed to 4.8-5.7 times), a spotted rather than monotone body, smaller size (much less than half that of the typical form), somewhat deeper body, and more oblique mouth with apex at the lower eye level. Data presented below under Food indicates that this species can be very plastic in its characters so subspecies designations would require extensive study of both characters and ecology. The Iranian fish are poorly represented by specimens and are referred here simply to the species.

Syntypes of sewerzowi (dated 1891 in Eschmeyer et al. (1996)) are in the Zoological Institute, St. Petersburg under ZISP 8747 (2), ZISP 8748 (2), ZISP 8749 (2), ZISP 8780 (1), ZISP 8901 (2) (Eschmeyer et al., 1996).

Kähsbauer (1964) reports a hybrid between Schizothorax schumacheri and Schizopygopsis stoliczkae from Sistan which may in fact be Schizocypris altidorsalis.

Key characters

The anal fin base has a sheath of enlarged scales and there are a few scales in the shoulder region and along the lateral line but the body is nearly naked, the anal fin has 5 branched rays, barbels are absent, and pharyngeal teeth are in 2 rows.

Morphology

Dorsal fin branched rays 7-9, usually 8, after 3-4 unbranched rays, anal fin branched rays 5-6, usually 5, after 2-3 unbranched rays, pectoral fin branched rays 12-20, and pelvic fin branched rays 8-10. Total vertebrae 48 (Howes, 1987). The lateral line is complete, a pelvic axillary scale is present and the anus and anal fin base are sheathed in enlarged scales. Scales are also present on the anterior lateral line and between there and the anterior pectoral fin base. A double row of scales extends forward from the anal sheath to the pelvic fin bases. Lateral line scales 96-120.

The dorsal fin spine is weakly developed and lacks denticles in large fish while young have many well-developed denticles. The gut is very elongate and coiled. The lower jaw is strong, with a dark brown, horny plate. The pharyngeal teeth number 3,4-4,3 and have a rounded base becoming spatulate distally with a rounded, hooked tip. Some teeth have a weak, flat cusp with a bump posteriorly below a rounded tip.

Meristic values for Iranian specimens are:- dorsal fin branched rays 8(3); anal fin branched rays 5(3); pectoral fin branched rays 17(1), 18(1) or 19(1); pelvic fin branched rays 9(3); lateral line scales 91(1) or 92(1); total gill rakers 11(3), reaching the raker below when appressed; and total vertebrae 48(1) or 50(2).

Sexual dimorphism

Tubercles are present on the anal fin of two male specimens from Sistan in a single file on the last unbranched and first 3 branched rays. They number up to 8 and are often widely spaced. Tubercles are found low on the flank above the anal fin and anus and on the upper flank behind the dorsal fin level. They also line the posterior lateral line. There are also a very few tubercles on the top of the head, side of the snout, below the eye and on the operculum, widely scattered and small.

Colour

Dark or yellowish with small dark brown or blackish spots extending onto the fins, or olive with large grey spots, or bluish-grey. Spots can be so numerous as to give the appearance of an overall blackish colour. Tilak (1987) however states that blackish spots are absent in both old and freshly preserved material. Both spotted and non-spotted forms occur in Sistan. The belly is whitish. Fins are pink. Peritoneum dark brown to black.

Size

Reaches 75.4 cm but the Sistan form is only up to 22.0 cm and is regarded as a dwarf form by Annandale and Hora (1920).

Distribution

Found in the mountainous areas of Afghanistan, Pakistan, India and western China. In Iran, restricted to the lowland Sistan basin where reported from the Hirmand River delta and 8 miles east of Lab-e Baring (Annandale, 1921; Vijayalakshmanan, 1950).

Zoogeography

The presence of this species in Sistan is an example of a riverine highway enabling species usually found at higher altitudes to penetrate into lowlands. See also the genus Schizothorax.

Habitat

Outside Iran the adults favour the main river while young are found in shallow streams and pools.

Age and growth

Savvaitova et al. (1989) cite a life span up to 22 years. Maturity in the Sistan form is attained at 18.0 cm. Berg (1949) reports a male fish 14 cm long with fairly well-developed testes and tubercles on the anal fin which bears out the dwarf nature of the Sistan fish.

Food

Savvaitova et al. (1989) examined feeding in certain lakes of the Pamirs at altitudes over 3220 m. This is much higher than the Sistan populations and the data may not be relevant. However the false osman is a poorly studied species so such information gives a basis for comparison and future research. The false osman is a very plastic species and can adapt to a variety of conditions. The osmans of the Pamirs were divisible into four groups: herbivores, detritivores, molluscivores and predators reflected in the structure of the gut and its length, the number of gill rakers, eye diameter, length and position of fins, and the shape of the horny plate on the lower jaw. Food includes higher aquatic plants, aquatic insects, diatoms and blue-green algae, detritus, molluscs and fish. In herbivores consuming periphyton, the horny plate on the lower jaw is rasp-shaped with the pointed end aimed anteriorly. In detritivores, the plate is sharper, and in predators, it is larger with the pointed edge directed upwards to grasp prey. Predatory behaviour only develops in fish over 30 cm, until which they eat plants.

Chaudhary et al. (1991) indicate that in a Pakistani population gut length increases with age and diet changes gradually from an omnivorous to a herbivorous one.

Reproduction

Spawning in Asia occurs generally in June and July. Eggs are large at 2.0 mm diameter.

Parasites and predators

None reported from Iran.

Economic importance

None.

Conservation

This species is rarely collected in Sistan and may require conservation measures.

Further work

Detailed surveys are needed to document the occurrence of this species in Sistan and to study its biology in a lowland habitat. Comparative studies with samples of the species from other, remote localities may demonstrate some level of taxonomic distinction as is seen in other schizothoracines of Sistan.

Sources

Iranian material: BM(NH) 1905.4.7:1-2, 2, 113.0-116.5 mm standard length, Sistan (no other locality data); ZISP 25854, 1, 111.5 mm SL, Sistan (no other locality data).

Comparative material: BM(NH) 1931.10.26:2-4, 2, 85.5-138.6 mm standard length, India, Ladkah, stream into Pangong Lake (no other locality data).

Genus Schizothorax
Heckel, 1838

The snow trouts, mountain barbels or Indian trouts are found from Iran to China, favouring mountainous areas but occasionally in lowlands. There are about 56 species (more if some other genera are included), with 3 in Iran.

The genera Racoma McClelland and Griffith in McClelland, 1842 and Aspiostoma Nikol'skii, 1897 are synonyms of Schizothorax Heckel, 1838 (Eschmeyer, 1990).

Schizothorax intermedius, and other species, have been placed in the genus Schizothoraichthys Misra, 1962 (e.g. in Tilak (1987). However Schizothoraichthys is regarded as a synonym of Schizopyge Heckel, 1847 by Jayaram (1981) or of Schizothorax (see Talwar, 1978; Eschmeyer, 1990). Schizopyge is itself regarded as a synonym of Schizothorax by some authors, e.g. Talwar (1978). Talwar (1978) separates the genus Oreinus McClelland, 1839 from Schizothorax by the margin of the lower jaw having a firm and hard horny covering which is thickest internally and a thick lower lip with a free posterior edge forming a sucker. Schizothorax has a non-suctorial lower lip and a lower labial fold interrupted or entire in the middle. However Talwar and Jhingran (1991) contradict this view and use Schizothoraichthys for Schizothorax and Schizothorax for Oreinus. Tilak (1987) recognises the name Schizothorax for fishes with strip of papillated tissue on the chin and Oreinus as a synonym; Schizothoraichthys is used then for fishes without the papillated chin. Oreinus is regarded as a synonym of Schizothorax by Jayaram (1981) and Eschmeyer (1990). Mirza (1991a; 1991b) recognises a tribe Schizothoracini, with the genera Schizothorax, Schizopyge, Racoma McClelland (and Schizocypris). I have retained Schizothorax here as the oldest name in view of these conflicting opinions.

There are various records of nominal Schizothorax species from the Helmand River basin in Afghanistan summarised in Coad (1981d); they have not been reported from the Sistan lowlands of Iran.

This genus is characterised by an elongate and almost cylindrical body, very small scales, over 100 in the series next to the lateral line, scales in complete lateral line somewhat larger, the vent and anal fin base are sheathed in enlarged scales and there may be enlarged scales near the pectoral fin and edge of the gill opening, dorsal and anal fins are short, dorsal fin with a thickened last unbranched ray bearing denticles (denticles lost with age), pharyngeal teeth in 3 rows and hooked at the tip, 4 barbels (rostral and maxillary), mouth inferior or subterminal, lower jaw may have a horny sheath, a papillated area on the chin may be present or absent, the lower labial fold may be interrupted or not in the middle, elongate gut and black peritoneum, and poisonous eggs. Members of the Schizothoracinae tribe are of polyploid origin with 2n=98 and 3n=148.

The ancestors of the schizothoracines in general were barbinines in the eastern part of the Qinghai-Xizang Plateau as it rose and water temperatures decreased in the late Miocene to early Pliocene (Sizhong, 1995). Primitive genera like Schizothorax migrated westwards earlier and further than more specialised genera such as Schizopygopsis (although both reach their westernmost distribution in Iran).

These fishes generally prefer rapids and pools of the larger streams at temperatures of 8-22°C although some occur in lakes with inflowing streams (Sharma, 1988). They are found in streams above 3000 m. Food varies from detritus to insects, plankton and fish depending on the species. The spawning season may be in late summer and early fall or in spring. Egg counts vary from a few hundred to over 50,000 and egg diameters may attain 3.6 mm. Some species show a spawning migration from warm lakes to cold streams.

Deaths have occurred from eating poisonous eggs of members of this genus but none are reported from Iran (see under S. zarudnyi). Symptoms include abdominal pain, nausea, vomiting, diarrhoea, dizziness, headache, fever, bitter taste, dryness of the mouth, intense thirst, sensation of chest constriction, cold sweats, rapid irregular weak pulse, low blood pressure, cyanosis, pupillary dilatation, syncope, chills, dysphagia and tinnitus. Severe cases show muscular cramps, paralysis, convulsions, coma, and death. Victims generally recover within 3-5 days with supportive treatment but it may take longer. Treatment is symptomatic and there is no known antidote or therapeutic data available. The patients' stomach should be evacuated as soon as possible after ingestion of eggs (Halstead, 1967-1970; Coad 1979b). Fish eaten during the breeding season should be cleaned with care to remove all traces of the eggs to avoid contamination of the flesh as cooking does not destroy the toxin.

Schizothorax esocinus
(Heckel, 1838)

Reported from the Helmand River drainage of Afghanistan; records summarised in Coad (1981d). No Iranian record.

Schizothorax intermedius
McClelland, 1842

Common names

mawda (in Sistan), marinka.

[marinka obyknovennaya or common marinka in Russian.

Systematics

Schizothorax intermedius was described from the "Cabul river at Jullalabad. Tarnuck River" in the Indus River basin.

Oreinus plagiostomus McClelland, 1842 described from the "Helmund river at Girdun Dewar" in Afghanistan, Racoma brevis McClelland, 1842 described from the "Helmund River", Racoma labiatus McClelland, 1842 described from "Pushut, Koonar river near Jullalabad" in the Indus River basin but also reported from the Helmand River basin by Annandale and Hora (1920) and Schizothorax ritchieana McClelland, 1842 described from "Affghanistan. In the Helmund there is a variety of this species ..... which will probably prove to be distinct." are probably synonyms (Berg, 1949).

Berg (1948-1949) reviews three morphae or forms of this species which indicate the great variation in this taxon. He also notes how barbel length varies independently of morpha and how the lips may be very strongly developed. The forms are as follows and the latter two were originally described as distinct species and are recognised as such by some authors: typica, eurystomus Kessler, 1872 and fedtschenkoi Kessler, 1872. Eurystomus has a transverse jaw covered by horny padding and a strong or weak dorsal fin spine. Fedtschenkoi has a lower jaw without the horny pad and the lower lip continuous or interrupted (the form with an interrupted lip is called morpha irregularis Day, 1896; the date is incorrect and should be 1877) and the dorsal fin spine is weak with 10-22 denticles extending to the mid-point or two-thirds along the ray. The forma typica has a crescent-shaped lower jaw without horny padding, the dorsal fin spine is well-developed with 12-32 denticles extending distally beyond the mid-point, and the lower lip is interrupted.

Schizothorax schumacheri Fowler and Steinitz, 1956 described from "Zabol, Eastern Iran" is also a synonym (Saadati, 1977). The holotype of Schizothorax schumacheri (ANSP 71950) at 244 mm total length and a paratype (ANSP 71951) at 130 mm to the end of the broken caudal fin, are stored in the Academy of Natural Sciences of Philadelphia (Böhlke, 1984).

Kähsbauer (1964) reports a hybrid between Schizothorax schumacheri and Schizopygopsis stoliczkae from Sistan which may in fact be Schizocypris altidorsalis.

Tilak (1987) reports Schizothorax richardsonii (Gray, 1832) from Sistan based on 2 fish in the Zoological Survey of India, Calcutta (F. 1226-1227/1), which I have not seen, and this may be the correct name for this fish.

Key characters

Gill raker counts and distribution separate this Schizothorax from others in Iran.

Morphology This is a very variable species, depending on habitat (Mirzaev, 1998). Dorsal fin with 2-4 unbranched and 5-9, usually 8 (Kullander et al. (1999) give 6-7 for their Kashmir specimens of S. curvifrons), branched rays, anal fin with 1-3 unbranched and 4-7, usually 5, branched rays, pectoral fin branched rays 14-19 and pelvic fin branched rays 7-10, usually 8. Lateral line scales 85-121, scale series next to the lateral line 115-165. Pharyngeal teeth 2,3,5-5,3,2. Gill rakers 10-17 (Kullander et al. (1999) give 21-28 for their Kashmir specimens of S. curvifrons). Total vertebrae 48 (Howes, 1987) or 40-43 (2000a) – presumably excluding 4 Weberian vertebrae). There is considerable variation in lower jaw form in specimens attributed to this species. The lower jaw may be crescent-shaped with or without a sharp horny sheath, or covered with a deciduous horny layer, or transverse and covered by a horny sheath. Lips may be interrupted medially or continuous, and can be very strongly developed. The dorsal fin spine may be well-developed with numerous denticles or weakly-developed with denticles not beyond the middle of the spine. Various morpha or infraspecies have been described to refer to these forms (see Berg, 1948-1949). Barbel length is highly variable. Young about 30 mm long have a naked body and no barbels. The karyotype is 2n=98-100. Usually silvery and occasionally with minute black spots on the upper half of the body but usually without spots. The head is olive-green. In preservative a pale greyish brown above, flanks and lower surfaces brilliant silvery or light yellow. Fins greyish to pale olive with lower ones whitish. Iris bright silvery white.

SizeReaches 60.0 cm (Solijonov, 2007).

Sexual dimorphism

Unknown.

Colour

Usually silvery and occasionally with minute black spots on the upper half of the body. In preservative a pale greyish brown above, flanks and lower surfaces brilliant silvery. Fins greyish to pale olive with lower ones whitish. Iris bright silvery white.

Size

Reaches 50 cm.

Distribution

Found in the basins of the Indus, the Amu Darya and Syr Darya rivers, the Tarim basin and the Helmand (= Hirmand) of Afghanistan and Iran.

Zoogeography

See under genus description.

Habitat

Reported from both lotic and lentic environments but little is known of its environmental requirements.

Age and growth

Life span is at least 8 years. Solijonov (2007), in a study of fishes in the Pamir-Alai Transboundary Conservation Area, found this species as being most active in the evening and so is seldom caught in the daytime when it hides in refuges among rocks. It maintains station in fast water of Pamir-Alai mountain streams behind rocks and in whirlpools. Males mature here at 2-3 years and females at 3-4 years and spawning takes place in mid-May.

Food

Food is small aquatic fauna, vegetation and detritus. Akhrorov and Kondur (1981) found macrophytes, detritus and molluscs to be important foods in a Pamir lake, varying with the year of sampling such that molluscs dominated in one year and macrophtyes in another.

Reproduction

Spawning takes place between May and September, depending on locality, and up to 8678-59,895 eggs are produced in fish 21.5-37.1 cm and 211-913 g (Mitrofanov et al., 1988). Spawning is probably non-annual in some areas (Maksunov, 1971).

Parasites and predators

None reported from Iran.

Economic importance

None.

Conservation

This species is not well-documented in Iran and no assessment of conversation needs can be made. It may be a stray from higher latitudes in Afghanistan.

Further work

Surveys should be done to confirm its presence in Iran.

Sources

Iranian material: None.

Schizothorax labiatus
(McClelland, 1842)

Reported from the Helmand River drainage of Afghanistan; records summarised in Coad (1981d). May be a synonym of Schizothorax intermedius (McClelland, 1842). No Iranian record.

Schizothorax pelzami
Kessler, 1870

Common names

shir mahi.

[Zakaspiiskaya marinka or Transcaspian marinka, and forel, in Russian].

Systematics

Schizothorax raulinsii Günther, 1889 described from a skin from the "Hari-rud River, near Khusan" and Schizothorax pelzami iranicus Karaman, 1969 are synonyms. Wossughi (1978) considers Schizothorax pelzami iranicus to be only a large specimen of S. p. pelzami and Coad and Keyzer de Ville (2004) concur.

The holotype of Schizothorax Poelzami is in the Zoological Institute, St. Petersburg (ZISP 8036, 265 mm total length) and is from "Fl. Schach-rud. accursus fl. Sefid-rud in Persia. 1889. Univ. Petropol.". The Shah Rud or Shah River is a tributary of the Safid Rud of the Caspian Sea basin but this species does not occur there. Berg (1948-1949) cites A. N. Derzhavin who suggests that this Shah-rud is south of Astrabad (= Gorgan). There is a Shahr Now River in the Tedzhen or Hari Rud basin where this species is found (shahr is the Farsi for city and may have been a general term for a major river of northeastern Iran as it flows through a city). However, the type probably came from the environs of the city of Shahrud (= Emamshahr) in the Damghan basin, a sub-basin of the Dasht-e Kavir basin (see below).

Another specimen listed as a type from the "Schah-Roude. Persia. Pelzame. St. Petersburg University" measuring 78.6 mm standard length is in the Natural History Museum, London (BM(NH) 1897.7.5:24). Kessler (1870) and Eschmeyer (1998) list 4 syntypes so two appear to be lost but Coad and Keyzer de Ville (2004) point out a disparity in size range (120-180 mm for the syntypes according to Kessler (1870)) while the London fish is too small and may not be a type despite its label.

The type of Schizothorax raulinsii is from the Hari-rud River, near Khusan, Afghanistan. BM(NH) 1886.9.21:181, is a skin marked as a syntype, 312.8 mm standard length, Hari-rud River, near Khusan, presumably the skin referred to by Eschmeyer et al. (1996). The skin is also marked Schizothorax “aitchisonii” (= S. raulinsii, a synonym of S. pelzami); Albert Günther of the British Museum apparently confused the names Aitchison and Raulins, collectors of specimens on the Afghan Delimitation Commission; aitchisonii was never used and the skin is a syntype of raulinsii. Eschmeyer lists a skin and 2 other syntypes in the Natural History Museum, London and 2 syntypes in the Zoological Survey of India, Calcutta (ZSI F11477-78). These 4 additional specimens are presumably the 4 smaller fishes mentioned by Günther (1889) in his original description as being collected at Bezd on the Jam River in Iran. BM(NH) 1886.9.21:171-172, 2, 87.8-100.9 mm standard length, Bezd, are not currently listed as syntypes in the British Museum however.

Schizothorax pelzami iranicus Karaman, 1969 is described from "Teheran in Quelle" (= Tehran in a spring) based on a single specimen. Schizothorax pelzami does not occur in the Namak Lake basin in which Tehran lies and the subspecies may have come from the Damghan part of the Dasht-e Kavir basin. The subspecies differs from the type subspecies by having a weakly ossified spiny ray in the dorsal fin (only the first half with small teeth), smaller eyes, longer snout and an overall brown to blackish-grey body colour with all fins, lips and barbels dark-coloured as opposed to the sharp boundary between the dark brown dorsal side and the light ventral side (Karaman, 1969). The Damghan part of the Dasht-e Kavir basin is the type locality of the species and S. p. iranicus is a synonym (Coad and Keyzer de Ville, 2004).

The holotype of S. p. iranicus is in the Zoologischen Instituts und Zoologischen Museums der Universität Hamburg (ZMH 4116, 327.5 mm standard length).

Starostin (1936) reports a hybrid of this species and Capoeta heratensis (= Capoeta capoeta) from Turkmenistan.

Key characters

This is the only schizothoracine species in northeast Iran and is easily recognised by its high lateral line scale count and the enlarged scales around the anus and anal fin.

Morphology

Dorsal fin branched rays 7-8 after 3-4 unbranched rays, anal fin branched rays 5-6, usually 5, after 3 unbranched rays. Pectoral fin branched rays 16-21, pelvic fin branched rays 7-9. The dorsal fin spine is very strong and thick with well-developed and widely-spaced teeth. Lateral line scales 84-108, lateral series scales 155-170, about 32 between the dorsal fin spine and the lateral line and about 27 between the lateral line and pelvic fin. The belly is scaled up to the isthmus. There is a scaled pelvic axillary process. The anal papilla and anal fin lie in a groove formed by enlarged scales, the groove extending about one third to half way between the anal fin origin and pelvic base. Scales are oval and obliquely inserted into scale pockets on mid-flank, sloping backwards postero-dorsally. The focus is subcentral anterior and radii are present on all fields. Circuli are few in these small scales. Pharyngeal teeth usually 2,3,5-5,3,2. Teeth are rounded with an evident hooked tip and posterior teeth have a short to medium flat grinding surface below the tip. Teeth may also be spatulate or have a spatulate shape with the hollow filled in. Gill rakers 9-15, relatively short and reaching the adjacent raker or slightly beyond when appressed. Occasional rakers are forked. The mouth is inferior. The lower lip is interrupted medially. The lower jaw may have a sharp horny sheath but this is mostly lacking. Mouth shape varies from a u-shape to a sector mouth (a gentle arch), the latter with a horny edge. The anterior barbels extend back to the anterior eye margin or the mid-eye while the posterior barbels extend to the rear eye margin or beyond. Barbel size is variable and not obviously related to size. The gut is very elongate and coiled.

Meristic values for Iranian specimens are:- dorsal fin branched rays 7(11) or 8(22); anal fin branched rays 5(33); pectoral fin branched rays 16(2), 17(4), 18(12), 19(6), 20(8) or 21(1); pelvic fin branched rays 7(2), 8(30) or 9(1); lateral line scales 85(2), 86(2), 88(2), 89(1), 90(2), 91(2), 92(2), 93(2), 95(2), 98(2), 99(7), 100(3), 102(1), 104(1), 105(1) or 108(1); total gill rakers 9(1), 10(5), 11(6), 12(7), 13(8), 14(1), 15(2), 16(1), 17(1) or 18(1); pharyngeal teeth 2,3,5-5,3,2(15), 2,3,4-5,3,2(2), 2,3,5-4,3,2(1), 2,2,5-5,3,2(1), 1,3,5-5,3,2(1); and total vertebrae 43(6), 44(10), 45(1), 46(2), 47(2), or 49(1).

Sexual dimorphism

A male specimen, 123.9 mm standard length, caught on 6 April had small to moderate sized tubercles on the top and sides of the head but these were not fully developed. The largest tubercles are found between the nostrils and the upper lip on the snout. No tubercles were noted on the fins. A fish taken on 5 November also had small but distinctive tubercles.

Colour

The overall coloration is silvery without any pattern but the back and upper flank are blackish to olive or brassy and the belly is whitish in small fish to a strong yellow in large fish. The back may be iridescent blue-green. The lateral line may be lighter than the surrounding flank, appearing as a thin, whitish line. The lips, pectoral, pelvic and anal fins are yellow. Fins bases are bright orange, the gill slit has a bright orange streak and the isthmus is bright orange. All fins may be flushed with red in freshly caught material and the lower flank and belly can have pinkish tinges. The iris is red dorsally. The peritoneum is a dark brown.

Preserved fish have a uniform brown colour with faint to dark speckles arranged irregularly on the flank. There are no obvious patterns on the fins although they are darkened by melanophores on both rays and membranes.

Günther (1889) reports the caudal fin as black, but this is possibly a dried or otherwise abnormal specimen. The colouration of the iranicus nominal subspecies in the original description (cited above) is not borne out by specimens from Damghan, the presumed locality of the type specimen, and again may be an artefact of preservation or simply a variation.

Size

Attains 54 cm (Muhomedieva, 1967) and reputedly 3 kg in qanat specimens (R. J. Behnke, in litt., 1981).

Distribution

Found in the Tedzhen and Murgab rivers of Afghanistan and Turkmenistan including Iranian drainages of the former known as the Hari River in its Iranian reach (Aliev et al., 1988). It is recorded from the Jam River, the Sharak River, the Akhland River near Mashhad, the Kashaf Riverand various smaller water bodies in Khorasan, the upper Kal Shur, Jajarm and Jovein rivers in the Dasht-e Kavir basin, as well as Cheshmeh Ali at Damghan and Cheshmah Badash near Shahrud further west, the westernmost distribution of the schizothoracine fishes (Günther, 1889; Nikol'skii, 1897; 1899; Abdoli, 2000; Coad and Abdoli, 2000b).

Wossughi (1978) records this species from the Hamun-See (= Sistan) but this is an error.

Zoogeography

Saadati (1977) found slight differences between fish from the Dasht-e Kavir basin and from the Hari River basin, in raker counts and caudal peduncle depth. He concludes that isolation in the Dasht-e Kavir basin is relatively recent and migration has occurred westwards in the past 15-25,000 years. The occurrence of this species in the western Dasht-e Kavir basin (the Damghan basin) is the westernmost distribution of the schizothoracine fishes. See also the genus Schizothorax.

Habitat

Found in springs, streams, rivers and qanats, the principal habitats of northeast Iran but environmental requirements are unknown.

Age and growth

Life span exceeds 7 years (Muhomedieva, 1967). Abdoli et al. (2007) found males reached 23.3 cm and 145.6 g and females 34.0 cm and 428 g in the Laiinsoo River. The sex ratio was male:female 2.5:1.

Food

The diet is 93-99% fishes including Cyprinus carpio (Muhomedieva, 1967). Other foods include small Capoeta capoeta, Cyprinus carpio, chironomids, caddis flies, dragonflies, other aquatic insects, and plant material (Aliev et al., 1988). Crustaceans, plant fragments and filamentous algae, and possibly fish eggs, may also be found in gut contents. Abdoli (2000) lists Plecoptera, Ceratopogonidae, Trichoptera, Ephemeroptera, Chironomidae and Simuliidae. Abdoli et al. (2007) found fish in the Laiinsoo River had a diet dominated by chrironomid pupae and simuliid larvae, with 9 other benthic invertebrate food items taken.

Reproduction

Egg diameter reaches 2.0 mm and numbers reach 36,300 eggs (Aliev et al., 1988). Iranian fish caught on 6 April had developing eggs suggesting a spring or early summer spawning period.

Parasites and predators

None reported from Iran.

Economic importance

None.

Conservation

The numbers and details of distribution need to be examined before an assessment can be made.

Further work

The biology of this species in Iran is unknown in detail.

Sources

Type material: See above, S. pelzami (ZISP 8036) (and possibly (BM(NH) 1897.7.5:24)) and S. p. iranicus (ZMH 4116).

Iranian material: CMNFI 1993-0124A, 1, 118.7 mm standard length, Semnan, Cheshmeh Ali-Damghan (36º17'N, 54º05'E); CMNFI 2007-0003, 8, 94.3-188.1 mm standard length, Semnan, Cheshmeh Ali (ca. 36º17'N, ca. 54º46'E); CMNFI 2007-0004, 6, 75.6-109.5 mm standard length, Semnan, Cheshmeh Bedasht (ca. 36º35'N, ca. 55º03'E); CMNFI 2007-0012, 2, 77.8-127.6 mm standard length, Khorasan, qanat at Bagh-e Jan (ca. 36º00'N, 58º38'E); CMNFI 2007-0013, 2, 55.9-80.4 mm standard length, Khorasan, qanat 5 km north of Boghai (ca. 36º02'N, ca. 59º31'E); CMNFI 2007-0014, 3, 88.7-98.1 mm standard length, Khorasan, Kuh-e Sang Park, Mashhad (ca. 36º18'N, ca. 59º36'E); BM(NH) 1914.1.1:16-17, 2, 98.0-163.9 mm standard length, Khorasan, Kashaf River, Mashhad (no other locality data); BM(NH) 1914.1.1:18-20, 3, 138.5-242.0 mm standard length, Khorasan, small stream near Mashhad (no other locality data); BM(NH) 1914.1.1:21-23, 2, 151.2-165.7 mm standard length, Khorasan, Cheshmeh-e Saby (no other locality data); BM(NH) 1914.1.1:24-29, 5, 147.6-188.8 mm standard length, Khorasan, Langar, Jam River (35º23'N. ca. 60º25'E).

Schizothorax plagiostomus
Heckel, 1838

Described from Kashmir; records summarised in Coad (1981d) for the Helmand River basin in Afghanistan but these may refer to Oreinus plagiostomus, a probable synonym of Schizothorax intermedius (q.v.). No Iranian record.

Schizothorax zarudnyi
(Nikol'skii, 1897

Common names

hamun mahi (= hamun or lake fish), ماهي خواجو (= mahi khvaju or khaju, after the historic island in Lake Hamun), sefidak or vatani (A. A. Pasand, pers. comm., 5 November 2000), shir mahi (= milk fish), anjak (Fowler and Steinitz (1956) report three kinds of fish being caught by fishermen in Sistan named anjaq (hence Oreinus anjac, see below), mawda (? unknown) and mahrmahé (= mar mahi, snake fish, probably a nemacheilid)).

Systematics

The holotype of Aspiostoma zarudnyi is in poor condition with the tail detached and the body impaled on a wooden spike when examined by me. Nikol'skii (1897) in his original description states in Latin "Specimen valde destructum". It is in the Zoological Institute, St. Petersburg (ZISP 11195a) and measures about 265 mm standard length. Nikol'skii (1897) gives the catalogue number as 11115 (sic, incorrectly according to Berg (1949)) and the type locality as "Palus Neizar in Seistan. 3.VI.96.". Berg (1949) gives the collection locality as "Neizar near the southern tip of Lake Hamun-i-Farah, western edge of the Helmand delta in northwestern Seistan" based on Zarudnyi (1901).

This species was originally described in the genus Aspiostoma Nikol'skii, 1897, a synonym of Schizothorax Heckel, 1838 (Eschmeyer, 1990). Bianco and Banarescu (1982) place this species as Schizopyge zarudny, the species name being a mis-spelling. Schizopyge Heckel, 1847 is regarded by these authors as the correct name for snow trouts without a suctorial disc on the chin (synonym Schizothoraichthys Misra, 1962 (see discussion above under the genus Schizothorax).

Barbus microlepis Keyserling, 1861 described from "Flüsschen bei Anardareh, zwischen Herat und Lasch" is a synonym and is pre-occupied by Barbus microlepis Bleeker, 1851. Oreinus anjac Fowler and Steinitz, 1956 from "Zabol, Eastern Iran" is also a synonym as suggested by Saadati (1977) although it may be a hybrid.

The holotype of Oreinus anjac (ANSP 71949) at 281 mm total length is stored in the Academy of Natural Sciences of Philadelphia (Böhlke, 1984).

This species is closely related to Schizothorax intermedius but is distinguished by much smaller paired fins, longer and narrower branchial isthmus, and the scales slightly enlarged at the base of all fins, especially the dorsal and anal (Annandale and Hora, 1920).

Key characters

The only common Schizothorax species in Sistan, is is recognised by the large barbels and enlarged scales around the anal fin.

Morphology

Dorsal fin with 4 unbranched and 7-8 branched rays, anal fin with 3 unbranched and 5 branched rays. The last unbranched dorsal fin ray is moderately strong and serrated, the serrations being proportionately longer and spinier in young. Pectoral fin branched rays 18 and pelvic fin branched rays 9. Lateral line scales 97-114, at least 190 in the scale series immediately above the lateral line. The breast is naked or sparsely scaled. There is a pelvic axillary process. The scale sheath around the anal papilla and anal fin extends about one third to half way between the anal fin origin and the pelvic fin base. Scales are very small, horizontally ovoid and have an almost central focus. Scales are obliquely inserted in the scale pockets on the mid-flank above the lateral line and below the dorsal fin. Scales on the nape are none to minimally imnbricate. Radii are found on all fields and are numerous. Gill rakers on the lower arm 30-39 (Nikol'skii's (1897) count of 25 is incorrect (Berg, 1949); but see below for wider variation). Gill rakers are long, reaching the third to the sixth adjacent raker when appressed. The interior margin of each raker is serrated. Pharyngeal teeth usually 2,3,5-5,3,2, spoon-shaped with a slightly hooked tip. Anterior teeth are more rounded and thicker. There are 2 pairs of barbels, the anterior ones long to rudimentary in literature sources. The barbels are subequal in length, the anterior ones not reaching the eye and the poterior ones not reaching beyond the eye. The mouth is usually slightly subterminal but can be terminal or have the lower jaw projecting slightly. The gut is elongate and coiled. The chromosome number is 2n=96, NF=142, comprising 9 pairs of metacentric, 14 pairs of submetacentric and 25 pairs of acrotelocentric chromosomes, and the fish is a tetraploid (Hosseini and Kalbasi, 2003; Kalbassi et al., 2008).

Meristics for Iranian specimens: dorsal fin branched rays 7(2) or 8(34); anal fin branched rays 5(35); pectoral fin branched rays 16(1), 17(7), 18(10) or 19(16); pelvic fin branched rays 9(30) or 10(5); lateral line scales 93(1), 97(1), 98(2), 99(3), 100(4), 101(5), 103(4), 105(2), 106(5), 107(2), 108(1), 109(2), 113(1) or 114(1); total gill rakers 24(1), 25(1), 26(1), 27(1), 28(1), 29(1), 31(5), 34(6), 35(2), 36(8), 37(1), 38(2), 40(1) or 41(1); pharyngeal teeth 2,3,5-5,3,2(14), 2,3,5-5,3,3 (1), 2,3,5-5,2,1(1), 2,3,5-5,3,1(1), 2,2,5-5,3,2 (1), 2,2,5-4,2,2(1), 2,3,4-5,3,2(2), 2,3,4-4,3,2(1) or 1,2,5-5,3,2(1); and total vertebrae 47(11), 48(11) or 49(4).

Sexual dimorphism

Males develop prominent nuptial tubercles on the snout and on the scales. Females have a soft and distended belly during the breeding season (CIRSPE, 2006b).

Colour

Overall colour is silvery, the back and head darker with indistinct fine dots. The flanks may be spotted with black and some small areas may be more lightly pigmented and appear as indistinct spots or blotches. There are melanophores on the fin rays and membranes. Adult males may have reddish fins and dull red specks on the dorsal surface. Young are more silvery than adults. Generally there is no distinctive pattern on the body and fins.

Colour varies with the environment. In muddy water, the back and fins are pale olive-green, the flanks tinged with green and the belly pure white while in the yellow water of the reed beds the back and flanks are much darker, almost black, and even the belly is darkish. The peritoneum is brown to black.

Size

Reaches 62.1 cm total length and and over 2.2 kg. Ahmadi and Wossughi (1988) give average weights of 300 to 2000 g in commercial catches while fish more than 12 kg are reported recently (Iranian Fisheries Research Organization Newsletter, 30-31:5, 2002).

Distribution

This species is restricted to the Sistan basin including the Chahnimeh (Nikol'skii, 1897; Bianco and Banarescu, 1982; J. Holčík, in litt., 1996.

Zoogeography

See under the genus description.

Habitat

Found in the open lake, in reed beds and in pools in Sistan (Annandale, 1921). It is the only species in Sistan common in the open lake in winter. Young probably make their way up upstream in the flood season as only adults are found in the lake in winter. The species is extremely abundant in pools left in stream beds when the floods recede. Spawning may occur in rivers as fry have not been found in the lakes (Iranian Fisheries Research Organization Newsletter, 30-31:5, 2002). Zabihi (2006) characterises it as a potamodromous species and notes that in March and April, if there is no flow in the rivers and thus no migration from the lake is possible, female gonads are reabsorbed.

Age and growth

Sexual maturity may only be attained after 4 years. Zabihi (2006) examined 697 specimens with a length and weight range of 24.5-62.1 cm and 137-2204 g. Half the male fishes were mature at 29-31 cm and for females at 38-40 cm.

Food

The diet comprises almost exclusively small fishes.

Reproduction

Eggs in fish caught in spring by me were developing but very small. Specimens with mature, yellow eggs have been caught in December. However this species is mature in Esfand (20 February-20 March), incubation is 6-7 days at 16.20-17.75ºC, maximum egg diameter is 3.8 mm when washed and the yolk sac is absorbed at 6-7 days (M. Abedi, Islamic Azad University of Savad Kooh, abstract). Zabihy et al. (2004) and Zabihi (2006) found maximum oocyte size in March and April at 14-18ºC when the gonadosomatic index was highest at 7.9-9.6. The mean absolute and relative fecundities for fish 460-1380 g was 26,256 and 34,418 eggs respectively. The species was a total spawner showing a synchronous ovary. Eggs are adhesive to prevent them being washed away by strong currents (CIRSPE, 2006b).

Parasites and predators

Datta (1937) describes the male of the acanthocephalan Eosentis rigidus from the intestine of this species.

Economic importance

Annandale in Annandale and Hora (1920) described the fishery for this species in the early years of the 20th century. Annandale commented that the flesh tasted like trout but was bony. The bones can be softened by cooking in vinegar. This fish is still caught and eaten and appears in local markets. Ahmadi and Wossughi (1988) cite an annual catch of 300-500 t while Iran Daily (24 August 2006) gives 700-1000 t before the drought.

Annandale and his assistants became sick from eating the eggs of this species but he maintained that the local fishermen ate it without any deleterious effects. Fish were caught in a net about 4 feet (1.22 m) deep and 100 feet (30.5 m) or more long anchored at each end by a tamarisk stick stuck into the lake bottom and with the bottom of the net on the lake bed and the top of the net slightly above the surface. The net was positioned in relation to the wind, and therefore the prevailing currents, so it formed a semi-circle. The net was arranged in a gap in the reed beds or just outside the reeds in the open lake if the weather was exceptionally calm. Pools in the reeds were kept open to facilitate the fishing. The fishermen riding their tutins (reed boats) would drive the fish into the net by beating the water with poles and ululating. The two ends of the net were then lifted out of the water by the men in the two end boats such that the net formed a bag. The net was drawn into the two boats as rapidly as possible. A similar but shorter net was used to drag small channels while the men using it waded. A small-meshed bag net attached to one horizontal and two upright poles was also used in pools of dry stream-beds. The net was dragged by ropes, the men wading through the water. Some large fish were killed in the flooded swamps by striking at them with swords. Another net consisted of a bag about 7 feet (2.14 m) long and 6 feet (1.83 m) by 2 feet (0.61 m) at the mouth. The mouth was held open by poles tied together at one end to make a fork. The fork pivoted on a post on the bank. The mouth of the net had fine lines across it, the lead string of which was held by the fishermen to warn him that a fish had entered the net so he could pull the net out of the water to retrieve the fish. The net was placed along a bank where the current swirled forming a backwater, at the mouth of a small canal, or as the focus of a line of stakes blocking a channel. The season for this type of net began as early as August or not until October. It lasted several weeks. March and April could also be a favourable time if the river was not very full but the fish did not move as actively. The large fish caught were attracted to feed on the numerous small fish which migrated up river along the shore and were checked where backwaters met the main flow.

A more recent description of fishing in Sistan is found in Fowler and Steinitz (1956). Fishing takes place in fall on rivers and in the lake, preferably the early evening or morning when it is cool. River fishing is preferred to lake fishing and some fishermen never go out onto the lake. Lake fishing using boats may last up to 3-5 days at a time. River fishing is carried out with a cotton-thread seine weighted by stones at the bottom and with gourds as floats. The seine is tied to sticks at each end and the sticks to ropes leading to shore. Four to ten men can own a seine and the catch is divided among those fishing that day. Fishing is done in teams of 4-7 men or often in two teams with two nets. Half the men are on one side of the river and half on the other, pulling each other's nets in and out of the water. Only large, fat fish are kept, the others being thrown back. One net catches 30-40 fish in a day which are then sold in Zabol. Women never fish nor are present during fishing as they bring bad luck. Fish are always cooked before eating as eating raw fish is reputedly fatal.

Conservation

Ahmadi and Wossughi (1988) state that introductions of various fishes such as Cyprinus carpio, Carassius auratus, Ctenopharyngodon idella and Hypophthalmichtys molitrix are a possible source of competition for native species such as this schizothoracine since they are voracious, take spawning sites and carry diseases and parasites. Native catches have decreased in favour of introduced species. M. Abedi, Islamic Azad University of Savad Kooh, has studied artificial reproduction in this species including larval development. CIRSPE (2006b) and Iran Daily (24 August 2006) give details on artificial reproduction of this species. Males over 600 g and females over 900 g were found to be suitable for breeding, ideal water temperature range was 18-22ºC, fecundity range was 24,300-37,640 eggs for fish 0.8-1.1 kg, egg size was 1.54 mm (presumably not water hardened; see above), and survival from egg to 11 mm fingerling was about 10%.

Further work

The molecular relationships of this species to other Schizothorax species and related genera could be revealing.

Sources

Type material: See above, Aspiostoma zarudnyi (ZISP 11195a).

Iranian material:- CMNFI 1979-0072, 1, 199.1 mm standard length, Sistan, Hirmand River near Zabol (30º58'N, 61º28'E); CMNFI 1979-0223, 1, 34.4 mm standard length, Sistan, irrigation jube 1 km south of Lutak (30º45'N, 61º24'E); CMNFI 1979-0225, 11, 182.2-225.3 mm standard length, Sistan, effluent of Hirmand River (30º58'N, 61º28'E); CMNFI 1979-0226, 2, 146.8-167.2 mm standard length, Sistan, pool near Kuh-e Khajeh (30º57'N, 61º17'E); CMNFI 1979-0231, 2, 23.1-25.8 mm standard length, Sistan, irrigation jube 3 km from Zabol (31º01'N, 61º32'E); CMNFI 1979-0232, 2, 26.3-26.5 mm standard length, Sistan, irrigation jube 11 km from Zabol (ca. 30º58'30"N, ca. 61º36'E); CMNFI 1979-0235, 13, 148.4-193.7 mm standard length, Sistan, effluent of Hirmand (30º54'30"N, 61º41'E); CMNFI 1979-0237, 8, 24.4-45.3 mm standard length, Sistan, irrigation jube 18 km south of Zabol (30º53'N, 61º27'30"E); BM(NH) 1920.1.20:35, 1, 232.5 mm standard length, Sistan, near Lab-e Baring (ca. 31º07'N, ca. 61º12'E); ZMH 5902, 2, 372.3-369.7 mm standard length, Sistan, Hamun See (no other locality data); ZMH 5903, 3, 274.2-298.7 mm standard length, Sistan, Hamun See (no other locality data); ZMH 6088, 2, 48.3-136.1 mm standard length, Sistan, Rud Sistan (no other locality data).

Genus Squalius
Bonaparte, 1837

The members of this genus were formerly placed in the dace genus Leuciscus Cuvier, 1816. Some subgenera have been elevated to generic rank and vice versa. The genus Leuciscus is not monophyletic based on allozyme data for a limited number of European taxa (Hänfling and Brandl, 2000). There may be about 22 species in the genus Squalius with 3 species reported from Iran. It has been suggested that species formerly considered to belong to the subgenus Squalius should be simply regarded as part of a Leuciscus cephalus complex characterised by serrated pharyngeal teeth in 2 rows (2,5-5,2) and an almost straight or convex anal fin margin. This complex would include L. cephalus, L. gaderanus (= L. ulanus) and L. lepidus among Iranian species (Bogutskaya, 1994). Bogutskaya (2002), however, places L. persidis and L. ulanus in a new genus Petroleuciscus, q.v. and places L. cephalus and L. lepidus in the genus Squalius, and this is followed here.

Bogutskaya (2002) gives the characters of Squalius as numerous total vertebrae (commonly more than 40, up to 48); increased number of sensory cephalic pores (up to 12-20 in the supraorbital canal) in most species; often fused and very expanded fourth and fifth infraorbitals; and depressed neurocranium with a reduced interorbital septum. Other characters are a somewhat compressed body, moderate to large scales, a complete lateral line, no barbels, mouth terminal or subterminal, no notch in the upper jaw accommodating a tubercle on the lower jaw, thin lips with the lower one interrupted medially, a short dorsal fin without a thickened ray, a moderately long anal fin, long and hooked pharyngeal teeth in 2 rows (2,5-4,2, 2,5-5,2 or 3,5-5,3 modally) usually with hooked tips and spoon-shaped crowns, short gut, no keel on the belly, and short and relatively few gill rakers.

Rückert-Ülkümen (2000) and Rückert-Ülkümen and Matzke-Karasz (2000) document Lower Miocene fossils of Leuciscus, presumably Squalius, from western Turkey.

Squalius cephalus
(Linnaeus, 1758)

Common names

mahi safid (= white fish) in the Caspian basin and in central Iran; mahi-ye safid rudkhanehi or mahi-e-sephid-e-roodkhaneie (= literally white river, presumably in the sense of white river fish) in Khuzestan; kuli; 'rus mahi or aroos mahi.

[enlibas or gafgaz enlibasi, nour enlibasi for natio kaznakovi, all in Azerbaijan; tepug in Armenia; bir-aan siphaloos, baeaan, barayan, or berak (= breast perhaps in allusion to the broad and fleshy chin (Heckel, 1843b)) at Aleppo, all in Arabic; Kavkazskii golavl' or Caucasian chub in Russian; European chub].

Systematics

Cyprinus Cephalus was originally described from southern Europe.

Leuciscus orientalis Nordmann, 1840 described from Abkhazia, Georgia, Squalius Berak Heckel, 1843 described from "Aleppo" (earlier in the same work - p. 1041 - spelled Berag without a description, presumably an error), Squalius cephalopsis Heckel, 1843 described from "Aleppo", Squalius orientalis Heckel, 1849 described from "Flusse Kueik bei Aleppo", Squalius turcicus De Filippi, 1865 described from "Dell' Arasse presso Erzerum" (Aras River at Erzerum in Turkey), and Squalius agdamicus Kamenskii, 1901 from Agdam in the Kura River basin, are synonyms.

Several natio and varieties within what is now Squalius cephalus have been described from Iran or contiguous drainages and are listed as follows. They have no taxonomic validity but the names may reoccur. Leuciscus cephalus orientalis natio kaznakovi Berg, 1912 was described from Lake Nour near Vandam, Nukha District, Azerbaijan (in the Tur'yan-chai basin of the Kura River basin but not connected to it). Squalius turcicus var. platycephala Kamenskii, 1897 was described from Lake Taparavani (= Lake Paravani at 41°26'N, 43°48'E) and the Kyrchbulach River in the upper Kura River basin, Georgia. Leuciscus cephalus orientalis natio aralychensis Barach, 1934 was described in Latin from "Turcia, Aralych. fl. Kara-su", and in Russian "Reka Kara-su u vblizi Aralykha podnoshchiya Ararata" (= Kara-su River at the foot of Ararat in the vicinity of Aralykha). Leuciscus cephalus orientalis natio zangicus Barach, 1934 was described in Russian from "R. Zanga v Armenii" (i.e. at Erivan). Leuciscus cephalus orientalis natio ardebilicus Barach, 1934 was described in Latin from "Ardebil, fl. Balyk-tchai in systemate fl. Arax. inf." and in Russian "R. Balyk-chaya, vblizi Ardebilya v Persii" (i.e. Balyk River or Balyk-chai) in the upper Aras River basin of Iran.

Iranian populations were usually recognised as Leusciscus cephalus orientalis (Nordmann, 1840). Bianco and Banarescu (1982) correctly point out that there has been no critical revision of the subspecies and that differences are slight. Recognition of subspecies is disputable according to Reshetnikov et al. (1997).

Turan et al. (2007) examined morphological variation in Turkish populations from the Black, Caspian, Aegean, Mediterranean and Tigris-Euphrates basins and found three groups, but no discrete taxa.

The nominal Iranian subspecies, L. c. orientalis, differs from the European subspecies, L. c. cephalus, by having a more elongate body, a dark stripe behind the operculum and on average fewer scales and anal fin branched rays (Berg, 1948-1949; 1949).

The syntypes of Cyprinus cephalus are in the Naturhistoriska Riksmuseet, Stockholm under LP 81 (1 fish) and in the Zoologiska Museet, Uppsala Universitet, Uppsala under ZMUU Linnaeus Collection 213 (1) (Eschmeyer et al., 1996).

Three syntypes of Squalius orientalis, 111-146 mm standard length from Aleppo are in the Naturhistorisches Museum Wien under NMW 49438 (Krupp, 1985c). Two more fish from Aleppo are under NMW 49440 and measure 130.0-168.3 mm standard length.

Six syntypes of Squalius berak are in the Naturhistorisches Museum Wien under NMW 48915 and 3 syntypes are in the Senckenberg Museum Frankfurt under SMF 469, formerly in NMW, and 3 syntypes of Squalius cephalopsis are under NMW 49438 and 2 other syntypes are under NMW 49440 (Eschmeyer et al., 1996). The Vienna catalogue lists 6 specimens as Squalius berak and 2 specimens as Squalius cephalopsis but the Vienna card index in 1997 agrees with Eschmeyer et al. (1996) for the NMW specimens.

Natio ardebilicus syntypes are in the Georgian State Museum, Zoological Section, Tbilisi under ZMT 136h, natio aralychensis syntypes are under ZMT 22-11 (10), and natio kaznakovi syntypes are in the Zoological Institute, St. Petersburg under ZISP 12089 (5) (Eschmeyer et al., 1996).

A hybrid between Squalius cephalus and Chalcalburnus (= Alburnus) chalcoides is reported from Turkey (Ünver and Erk'akan, 2005).

Key characters

The relatively large scales outlined by pigment give this fish a distinctive appearance and, combined with large body size, fin ray counts, rounded belly and the absence of barbels, separates it from other Iranian cyprinids.

Morphology

Dorsal fin branched rays 7-9, usually 8, after 2-3, usually 3, unbranched rays, anal fin branched rays 7-10, usually 8 after 3 unbranched rays, pectoral fin branched rays 14-19, and pelvic fin branched rays 6-9, usually 8. Lateral line scales 38-48. Scales have few to moderate numbers of radii on the anterior and posterior fields. The focus is central to subcentral anterior and circuli are fine. On the posterior field, circuli break up into "bubbles" and are coarser than on other fields. The anterior scale margin is wavy, sometimes irregular and others with a small central protuberance and indentations above and below. Gill rakers short, 7-12, touching the adjacent raker when appressed. Vertebrae 40-46. Pharyngeal teeth 2,5-5,2, with variants 2,5-4,2, 2,4-4,2, 2,5-5,1, 1,5-5,2, 1,5-5,1, 2,5-5,3, 2,6-5,2, or even 1,5-5,1,2 and 1,2,5-5,2,2. Teeth are very narrow, strongly hooked at the tip and strongly serrated so that there is no obvious flat surface. The serrations on the teeth of orientalis are stronger than in the typical subspecies. The gut is an elongate s-shape. Chromosome number is 2n=50 (Al-Sabti, 1986; Klinkhardt et al., 1995).

Meristics for Iranian specimens: dorsal fin branched rays 7(3), 8(103) or 9(1); anal fin branched rays 7(3), 8(60) or 9(44); pectoral fin branched rays 14(3), 15(13), 16(39), 17(46), 18(5) or 19(1); pelvic fin branched rays 6(1), 7(1), 6(100) or 9(5); lateral line scales 40(2), 41(20), 42(45), 43(30), 44(9) or 46(1); total gill rakers 7(2), 8(14), 9(58), 10(31) or 11(2); pharyngeal teeth ?; and total vertebrae 40(6), 41(29), 42(45) or 43(6).

Sexual dimorphism

Abdurakhmanov (1962) reports that head length, eye diameter, caudal peduncle length, dorsal and anal fin heights, pectoral and pelvic fin lengths and lower caudal fin lobe length are all greater in males while postorbital length, interorbital width, head depth and pectoral-pelvic fin distance are greater in females.

Colour

Overall colour is silvery to grey. Scales of the lateral line and upper back have a strong dark pigmentation along the posterior margin and a distinct dark spot anteriorly. Scales may have some gold in their centre. The back is dark brown or reddish-brown to blue-grey and the belly and lower head are pearly-white to a silvery yellow. The operculum is a strong copper-yellow colour and the opercular opening is dark, nearly black. The iris is silvery and has very little gold, or is golden with a lot of dark grey pigment. There may be an upper spot of dark grey on the iris. The dorsal and anal fins are grey and may have some pink. The dorsal fin is blackish distally. The pectoral fin has a little pale grey pigment and there may be a yellow or yellow-pink spot at its upper base with this colour extending onto the first 2-3 rays. The pelvic and anal fins are pink to red-pink with somewhat colourless posterior margins, especially in spawning males. The caudal fin is a pale pink to dirty reddish with some dark pigmentation to the posterior margin.

Size

Possibly as long as 80 cm although 45 cm is a more likely maximum (Berg, 1948-1949) and a weight of 6-8 kg. Reaches 43 cm total length in the Madar Su of Golestan National Park (A. Abdoli, pers. comm., 1995). Reaches 39.2 cm and 870 g in Taham Dam in northern Iran (Iranian Fisheries Research Organization Newsletter, 54 & 55:4, 2008). Attains 39.0 cm fork length in Turkey (Yerli et al., 1999). Maximum size is possibly 85.0 cm and 10.0 kg.

Distribution

Found from the British Isles and the Iberian Peninsula eastwards in its southern distribution to Turkey and Iraq, the northern half of Iran, and the whole Caspian and Aral seas drainages. In Iran, it is recorded from rivers along the Caspian Sea coast including the Anzali Mordab (Holčík and Oláh, 1992; Roshan Tabari, 1997; Shamsi et al., 1997; Abbasi et al., 1999; Kiabi et al., 1999) but also the Aras River, the Namak Lake basin (Wossughi, 1978; Bianco and Banarescu, 1982), the Lake Orumiyeh basin (Günther, 1899); Lake Zaribar, Kordestan (Abzeeyan, 5(5):III, 1994), and the Esfahan basin and the Tigris River basin (Berg, 1949). Abdoli (2000) maps this species from the middle Atrek and Gorgan, middle and upper Neka, middle and lower Babol and Heraz, Chalus and Tonekabon, Safid, Shahrud and Qezel Owzan, middle Aras and its tributary Qareh Chai, all in the the Caspian Sea basin, the middle and lower Talkheh, Zarrineh and Tatavi in the Lake Orumiyeh basin, the Karaj, Shur, Abhar, Qareh Chai and Qom rivers in the Namak Lake basin, and the Marun, upper Karun and middle Khersan, Dez, middle and upper Karkheh, Simarreh, Kashkan and lower Gav Masiab rivers in the Tigris River basin.

Zoogeography

This widely distributed species has been split into a number of subspecies but the principal one from the Middle East or Southwest Asia is recognised as S. c. orientalis. This has not been examined in detail recently using classical techniques and the relationships between isolated populations are unknown. Durand et al. (1999), working on the cytochome b of European populations, found that a lineage from a Ponto-Caspian refugium recolonised the Baltic area in the Holocene after glaciation. Durand et al. (2000), again examining cytochrome b, consider that S. cephalus may have originated from Mesopotamia and, in the late Pliocene, used the large inland lake of Anatolia existing at that time for dispersion. Uplift of the Anatolian Plateau, climatic changes and river isolation was probably the main vicariant event leading to a quick radiation in these chubs.

Habitat

In the Caspian Sea basin, it is found mainly in upstream waters (Jolodar and Abdoli, 2004).

Age and growth

Öztaş (1988; 1989), Öztaş and Solak (1988) and Türkmen et al. (1999) studied the chub in the Aras River of Turkey and found its condition factor to be higher in summer and autumn and to vary between age groups. Life span there is over 8 years, compared with 7 years in Lake Aksehir, Anatolia (Altindag, 1996) and in the Savur stream in the Tigris River basin of southeast Anatolia (Ünlü and Balcı, 1993a; 1993b), over 8 years in the Euphrates River in Turkey (Özdemir and Şen, 1986), and 10 years in Sariyar Dam Lake near Ankara (Ekmekci, 1996a). In the Aras, females grow faster than males and there are more sexually mature females in the older age groups. Sexual maturity is attained at age 2-3 in males and ages 3-4 for females. Mean condition factor for Aras males was 1.326 and for females 1.333. In the Savur stream in the Tigris River basin, 80% of females and 75% of males reaching sexual maturity at their third (14.6 cm fork length) and second years (13.0 cm) respectively. Females grow larger than males and live about 2 years longer as they do in Lake Aksehir. Sexual maturity is attained at ages 3-4 in Sariyar Dam Lake and these fish at age 1 have a mean fork length of 90 mm, mean weight of 9.4 g and a condition factor of 1.263 which by age 10 can reach 370 mm, 910 g and 1.805. Aksehir Lake fish reached 44 cm forked length and 1766 g for females and 31 cm and 557 g for males. In Lake Tödürge, Turkey, Ünver (1998) found that males reached maturity in their second or third year of life while females were in their third or fourth year. Over 68% of the catch was female and life span attained 7 years. Yerli et al. (1999) invetsigated growth in Lake Çıldır, Turkey and also found 8 age groups, dominated by those in age group 3 at 67.3% and by males at 73% and with maximum fork length at 390 mm and maximum weight at 720 g. Maximum life span is up to 15 years. The weight class 150-450 g dominated in Taham Dam in Iran (Iranian Fisheries Research Organization Newsletter, 54 & 55:4, 2008).

Food

Food items include mayfly and caddisfly larvae, other small organisms such as molluscs, and crayfishes, small fishes and frogs. Large fish feed mainly on other fish. Reputedly even fruit fallen in the water will be eaten. Trout eggs and fry are also eaten. Guts of Iranian specimens contained a wide variety of organisms including ants (presumably taken at the water surface), aquatic insects such as chironomids among others, crustaceans, filamentous algae, higher plant fragments, scales of cyprinids and the remains of a Paracobitis malapterura. Abdoli (2000) reports Ephemeroptera, Chironomidae and Trichoptera. In Taham Dam in Iran algae, higher plants, bivalve and insects were found (Iranian Fisheries Research Organization Newsletter, 54 & 55:4, 2008).

Reproduction

Öztaş (1989) examined reproduction of this species in a stream tributary to the Aras River in Turkey. Spawning begins at the end of May although most fish spawn in June. Water temperatures at this time are 12-18°C. Fecundity is up to 61,808 eggs and maximum egg diameter is 1.39 mm. Türkmen et al . (1999) found spawning between May and July in the Aras River proper with fecundity up to 17,187 eggs. Ünlü and Balcı (1993a; 1993b) found spawning to take place from May to late June in southeast Anatolia in a tributary of the Tigris River. Fecundity reaches 20,140 eggs and egg diameter 1.5 mm. Equations for the relationship between fecundity (F) and length (FL), weight (W) and ovary weight (GW) were given as F = 0.0458 FL^2.3680 (r = 0.745), F = 270.96 W^0.7912 (r = 0.761) and F = 1888.86 GW^0.8163(r = 0.775). In Sariyar Dam Lake near Ankara, spawning takes place between April and June (Ekmekci, 1996a). Ünver (1998) found fish in Lake Tödürge, Turkey spawning between May and July with a mean egg diameter of 0.65 mm (highest 1.04 mm) and a mean fecundity of 14,500 eggs (highest 28,664 eggs).

In Azerbaijan, Abdurakhmanov (1962) gives spawning temperatures as 12-21°C, maximum fecundity as 118,000 eggs and maximum egg diameter as 1.8 mm. Spawning in Iran appears to take place in spring judging from egg development.

Around 40% of the fish in Taham Dam in Iran spawned in early June and around 30% did not spawn at all (Iranian Fisheries Research Organization Newsletter, 54 & 55:4, 2008).

Parasites and predators

Ergens and Gusev (1965) report the monogenean helminth Dactylogyrus prostae in this species from Bandar-e Shah (= Bandar-e Torkeman) on the Caspian Sea coast. The monogeneans Diplozoon paradoxum and D. megan are recorded from this species in the Tajan River, Mazandaran (Iranian Fisheries Research and Training Organization Newsletter, 6:7, 1994). Shamsi et al. (1997) report Clinostomum complanatum, a parasite causing laryngo-pharyngitis in humans, from this species. Mirhasheminasab and Pazooki (2003) list Ergasilus peregrinus, Tracheliastes polycolpus and Lernaea cyprinacea from this species in Mahabad Reservoir, the latter being the most dangerous parasite. Jalali et al. (2005) summarise the occurrence of Gyrodactylus species in Iran and record G. sp. from fish in the Safid River. Masoumian and Pazooki (1998) surveyed myxosporeans in this species in Gilan and Mazandaran provinces, finding Myxobolus minutus and M. muelleri. Pazooki et al. (2005) record Lamprolegna compacta, Ergasilus peregrinus and Lernaea cyprinacea from this species in waterbodies of Zanjan Province. Miar et al. (2008) examined fish in Valasht Lake and the Chalus River, Mazandaran and found the protozoan Chilodonella hexastica.

Economic importance

Listed as economically important in the Turkish Euphrates River (Özdemir and Şen, 1986).

Conservation

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in textbooks and for sport. Kiabi et al. (1999) consider this species to be of least concern in the south Caspian Sea basin according to IUCN criteria. Criteria include sport fishing, medium numbers, habitat destruction, widespread range (75% of water bodies), present in other water bodies in Iran, and present outside the Caspian Sea basin.

Ekmekci (1996a) recommends a minimum fishing size of 28 cm and a prohibition of fishing during the April-June spawning season for this Anatolian population.

Further work

The distribution and biology of this species in Iran needs study, as does the systematics of isolated taxa.

Sources

Iranian material: CMNFI 1970-0506, 6, 42.5-116.3 mm standard length, Gilan, Shalman River (37º08'N, 50º15'E); CMNFI 1970-0536, 3, 89.0-223.0 mm standard length, Gilan, Siah River estuary near Rudbar (36º53'N, 49º32'E); CMNFI 1970-0538, 7, 40.8-137.2 mm standard length, Gilan, Qezel Owzan River (36º44'N, 49º24'E); CMNFI 1970-0583, 6, 41.4-123.8 mm standard length, Gilan, Nahang Roga River (37º28'N, 49º28'E); CMNFI 1970-0589, 1, 142.0 mm standard length, Gilan, Safid River (37º12'N, 49º54'E); CMNFI 1979-0253, 33, 46.4-100.4 mm standard length, Markazi, Qareh Su River drainage (34º52'N, 50º49'E); CMNFI 1979-0255, 2, 90.7-90.8 mm standard length, Markazi, Bar River drainage (33º51'30"N, 50º23'E); CMNFI 1979-0276, 1, 59.7 mm standard length, Lorestan, Chamesk River (ca. 33º19'N, ca. 47º53'30"E); CMNFI 1979-0462, 12, 50.7-104.5 mm standard length, Markazi, Mazdaqan River (35º06'30"N, 49º40'30"E); CMNFI 1979-0469, 6, 86.0-127.5 mm standard length, Mazandaran, river west of Alamdeh (36º37'30"N, 51º35'E); CMNFI 1979-0474, 6, 71.3-80.6 mm standard length, Mazandaran, Tajan River (36º34'N, 53º05'E); CMNFI 1979-0482, 2, 112.2-160.1 mm standard length, Mazandaran, river between Minudasht and Dowlatabad (37º19'30"N, 55º31'E); CMNFI 1979-0493, 7, 85.2-114.5 mm standard length, Mazandaran, Tajan River drainage (36º19'N, 53º23'E); CMNFI 1979-0494, 12, 9.1-92.6 mm standard length, Mazandaran, Talar River tributray (36º21'N, 52º51'30"E); CMNFI 1980-0120, 2, 41.8-118.1 mm standard length, Mazandaran, Babol River at Babol Sar (36º43'N, 52º39'E); CMNFI 2007-0092, 5, 25.5-45.9 mm standard length, Azarbayjan-e Khavari, Zilber Chay (38º42'N, 45º16'E); CMNFI 2007-0100, 4, 41.2-171.8 mm standard length, Azarbayjan-e Bakhtari, Kalwi Chay near Piranshahr (ca. 36º44'N, ca. 45º10'E); CMNFI 2007-0101, 1, 162.0 mm standard length, Azarbayjan-e Bahktari, Tata'u River (ca. 36º54'N, ca. 46º07'E); CMNFI 2007-0105, 1, 96.9 mm standard length, Kordestan, Zarineh River basin (ca. 36º06'N, ca. 46º20'E); CMNFI 2007-0109, 1, 101.1mm standard length, Kordestan, Qeshlaq River basin (ca. 35º16'N, ca. 47º01'E); CMNFI 2007-0113, 1, 99.5 mm standard length, Kermanshahan, Qareh Su River tributary (ca. 34º25'N, ca. 47º01'E).

Comparative material: BM(NH) 1974.2.22:71-72, 2, 68.0-93.1 mm standard length, Iraq, Baghdad (33º21'N, 44º25'E); BM(NH) 1974.2.22:74, 1, 152.1 mm standard length, Iraq, Greater Zab near Aski Kalak (ca. 36º16'N, 43º39'E); BM(NH) 1974.2.22:75, 1, 154.0 mm standard length, Iraq, Qizilja River, Lesser Zab (no other locality data).

Squalius latus
Keyserling, 1861

Flank scale

Common names

None.

[Zakaspiiskii elets or Transcaspian dace in Russian; Murgab dace].

Systematics

Squalius latus was described from the "Fluss. Heri-Rud bei Herat" (now in Afghanistan, then in Persia; Herat is at °'N, °'E).

Squalius transcaspiensis Berg, 1898 from "Habitat in flum. Tedschent, prope Aschabat in provincia Transcaspica" is a synonym. The type locality is presumably the Tedzhen River in Turkmenistan (not Iran as given in Eschmeyer et al. (1996)) although Ashkhabad is not on the Tedzhen River. Many syntypes are in the Zoological Museum of Moscow State University (MMSU) according to Eschmeyer et al. (1996). Possibly a subspecies of Squalius lehmanni (Brandt, 1852) according to Nikol'skii (1938) and Svetovidova (1967) or of Leuciscus leuciscus (Linnaeus, 1758) according to V. V. Kafanova (cited in Bogutskaya, 1994). Bogutskaya (1994) considers S. latus to be distinct.

Key characters

Dorsal and anal fin ray counts, both 7-9, lateral line scale counts under 48, and distribution identify this species in northeastern Iran.

Morphology

Dorsal fin branched rays 7-9, after 3 unbranched rays, anal fin branched rays 7-9 after 3 unbranched rays, pectoral fin branched rays 14-16, and pelvic fin branched rays 7-8. Pharyngeal teeth number 2,5-5,2; 2,5-5,1; 1,5-5,1; 3,4-4,2; 3,5-5,2; or 2,5-4,2. In specimens from the Tedzhen (= Hari) River, part of which is the Iran-Afghanistan border:- dorsal fin branched rays 7(12), anal fin branched rays 7(1), 8(6) or 9(5), pectoral fin branched rays 15(6) or 16(6), and pelvic fin branched rays 7(1) or 8(11). Lateral line scales 39(1), 40(1), 41(2), 42(2), 43(1), 44(2), 45(1), 46(1), or 47(1). A pelvic axillary scale is present. Scales bear a few anterior and posterior radii. Total gill rakers number 8(2), 9(1), 10(5) or 11(4) and reach the raker below when appressed. Total vertebrae 39(4), 40(6) or 41(3) (includes data from x-rays of ZISP 18361 and 11047). Pharyngeal teeth number 2,5-5,2(3) and have narrow cutting edges on each side of a shallow groove below a hooked tip and in larger fish some teeth are serrated below the tip. The mouth is oblique and extends back to the level of the nostrils, the lower jaw protruding slightly.

Sexual dimorphism

Unknown.

Colour

The back and upper flank are reddish-brown, the flanks yellowish and the belly silver. The dorsal and caudal fins are yellowish-brown and the other fins are reddish. The base of all fins becomes bright orange during spawning.

Size

Reaches 26.7 cm.

Distribution

This species is found in the Tedzhen River (= Harirud in Iran) and the Murgab River to the east in Afghanistan and Turkmenistan.

Zoogeography

This species is part of a complex of species in the genus Squalius and the related Leuciscus found from western Europe to Siberia. Its origins may lie in a Ponto-Caspian-Aralian refugium, becoming isolated and speciating in the eastern part of this area.

Habitat

Unknown in detail.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

None reported from Iran.

Economic importance

None.

Conservation

Numbers and habitat requirements are unknown and so a conservation assessment cannot be made.

Further work

The biology of this species needs study as well as it relationships to other Squalius.

Sources

Type material: There are no type specimens.

Iranian material: None.

Comparative material: ZISP 10357a, 1, 63.2 mm standard length, Turkmenistan or Afghanistan, Tedzhen River (no other locality data); ZISP 10359, 4, 93.4-130.5 mm standard length, Turkmenistan or Afghanistan, Tedzhen River (no other locality data); ZISP 10360, 6, 58.4-122.0 mm standard length, Turkmenistan or Afghanistan, Tedzhen River (no other locality data); ZISP 10419, 1, 107.6 mm standard length, Turkmenistan Iolotan near Merv (37º18'N, 62º21'E).

Squalius lepidus
Heckel, 1843

Common names

kavar or kawar.

[bara'an or bir-aan abiadh in Arabic; Tigris dace].

Systematics

Squalius lepidus was originally described from the "Tigris bei Mossul" (Heckel, 1843b).

Alburnus doriae De Filippi, 1865 and Alburnus maculatus Keyserling, 1861 are synonyms according to Coad (1982d; 1985h) but these are errors. Note also that Alburnus maculatus Kessler, 1859 from the Salghir River at Simferopol, Crimea, Ukraine preoccupies Alburnus maculatus Keyserling, 1861 and is a distinct species of Alburnoides (Bogutskaya and Coad, 2009).

The syntypes of Squalius lepidus are in the Naturhistorisches Museum Wien according to Krupp (1985c) under NMW 49342, 2 specimens, 229.6-245.5 mm standard length as measured by me and NMW 49343, 2, 88.0-107.8 mm standard length, collected by Th. Kotschy in 1843. Bogutskaya (1994), however, lists a lectotype (237.0 mm standard length) and 5 paralectotypes under NMW 49342, 2 paralectotypes under NMW 49343, and 1 paralectotype under SMF 847 (the paralectotypes measure 90-8-233.8 mm standard length). Eschmeyer et al. (1996) also lists 1 syntype under NMW 49344 and this specimen (164.8 mm standard length) appears in the 1997 Vienna card catalogue as a syntype along with the other 4 fish mentioned by Krupp (1985c). One syntype is in the Senckenberg Museum Frankfurt (SMF 847, formerly NMW) (F. Krupp, pers. comm., 1985). The catalogue in Vienna lists 6 fish in spirits and 1 fish stuffed.

Key characters

This species is distinguished from other Squalius by the elongate and pointed head with a projecting lower jaw, an adaptation for piscivory which develops early and is not evident in other Squalius even when these feed on fish as adults.

Morphology

Dorsal fin branched rays 8-10, usually 8 (Krupp, 1985c) or 9 (Bogutskaya, 1994)(8 in four syntypes, 9 in one), after 3 unbranched rays, anal fin branched rays 9-11, usually 9, after 3 unbranched rays, pectoral fin branched rays 14-18, and pelvic fin branched rays 8. Lateral line scales 42-50. Scales have few radii on the anterior and posterior fields. Gill rakers 7-12, reaching the one below when appressed (near junction of upper and lower arches) and sometimes curled. Pharyngeal teeth 2,5-5,2 or 2,5-4,2, strongly hooked and serrated. Total vertebrae 42-46. The projecting lower jaw fits into a notch in the upper jaw.

Sexual dimorphism

Unknown.

Colour

The back and upper flank are brown to bluish-brown or blackish, the flanks generally silvery, and the belly silvery-yellow. The upper flank, head and fins may be sprinkled with black spots. Flank scales have a spot at their base. All fins are reddish, with the pectoral and anal fins the brightest. The dorsal fin has a black margin. The caudal fin is blue-grey and its margin is blackish.

Size

Reaches 54.2 cm (Karabatak, 1997).

Distribution

Found in the Orontes (= Asi), Quwayq and Tigris-Euphrates basins as well as some lakes in central Anatolia (Beyşehir, Akşehir) and the rivers Ceyhan and Şeyhan in southern Anatolia.

Records from Iran are based on literature and include Lake Zaribar in Kordestan, Kermanshah, the Marakeh River, Luristan, and Dez River below Dez Dam (= formerly Mohammad Reza Shah Pahlavi Dam) (Wossughi, 1978). Also recorded from Mendeli on the Iran-Iraq border (Berg, 1949). Abdoli (2000) records this species from rivers of Khuzestan such as the Jarrahi, Marun, Karun, middle and lower Dez and middle Karkheh; and the Qom, Shur and Zayandeh rivers probably in error. A record from a canal near Gaz a few miles from Esfahan is probably an error (Keyserling, 1861).

Zoogeography

Durand et al. (2000) using cytochrome b suggest that this species is fully introgressed with L. cephalus mtDNA (= S. cephalus) and so question the taxonomic validity of this species. Morphological data contradicts this conclusion. Durand et al. (2000) conclude that their data does indicate that "L. lepidus and L. cephalus might have had different dispersion histories over the same geographical range" and that "L. lepidus introgression by the chub (L. cephalus) is ancient, explaining the complete sorting of the lepidus lineage".

Habitat

Generally unknown but found in rivers and lakes.

Age and growth

Karabatak (1997) studied fishes identified as this species from Lake Beyşehir in Turkey and recorded condition coefficients of 1.30-1.45 pre-spawning, 1.13-1.24 spawning and 1.24-130 post-spawning, with maximum values in the pre-spawning period coincident with maximum gonad development. Males and females showed similar length-weight relationships although females were slightly heavier because of their eggs. Çolak (1983) studied the age of this species in the Keban Dam, Turkey and found a maximum age of 8 years, that females grew faster and were longer than males, and growth was rapid until five years of age when it fell by almost half in each succeeding year. von Bertalanffy formulae, sexes combined, for two years were L_t = 87.92 (1-e^{-0.112(t+1.464)}) and L_t = 65.64 (1-e^{-0.214(t+0.878)}).

Food

Plant remains and fish scales have been found in gut contents. Fish appear to be the main diet item even in young from about 10 cm in length (Bogutskaya, 1994).

Reproduction

Karabatak (1997) studied fishes identified as this species from Lake Beyşehir in Turkey and found a peak spawning in mid to late June at 20.0-21.5°C with the highest gonadosomatic index on 15 May at 16.0-17.2°C. The spawning season could extend from late May to early July. Ünlü (2006) gives age at first maturity as 2-3 years in the Turkish Tigris River with spawning over sand, stone and gravel.

Parasites and predators

Williams et al. (1980) report the digenean helminth Allocreadium isoporum from this species in the Zayandeh River at Esfahan but the fish species was most probably misidentified.

Economic importance

Economically important in Turkey in Lake Beyşehir (Karabatak, 1997).

Conservation

The biology and distribution of this species in Iran need elucidation so a conservation assessment can be made.

Further work

The biology of this species needs study along with its systematic relationships with other Squalius species as morphology and molecular analyses are contradictory.

Sources

Some counts are taken from Bogutskaya (1994).

Type material: See above, Squalius lepidus (NMW 49342, NMW 49343 and NMW 49344).

Iranian material: None.

Comparative material: NMW 87786, 2, 329.0-381.5 mm standard length, Turkey, Karaviran at Konya (ca. 37º52'N, ca. 32º31'E); NMW 91149, 11, 69.6-115.9 mm standard length, Syria, Euphrates River at Jarabulus (36º49'N, 38º01'E); NMW 91620, 2, 91.6-95.6 mm standard length, Syria, Euphrates River at Jarabulus (36º49'N, 38º01'E); NMW 94441, 1, 175.7 mm standard length, Turkey, Palu, Murat River (38º42'N, 39º57'E).

Genus Tinca
Cuvier, 1816

The tench genus contains a single species found from Europe to Siberia including the Caspian Sea basin.

The characters of this genus include very small, elongate scales deeply embedded in slimy skin, pharyngeal teeth in a single row, all fins rounded and without spines, a short barbel at the terminal mouth corner, no keel on the belly, moderately long gill rakers, and short dorsal and anal fins.

Tinca tinca
(Linnaeus, 1758)

CMNFI 1979-0439, Gilan, Anzali Mordab

Common names

لاي ماهي (= lie, lai, lay or laay mahi, meaning bottom-dwelling or mud fish); nazi; tilkhos; saboni (= soapy?); hashtarkhan kapur.

[lil baligi in Azerbaijan; sew-zukgná in Armenia; lin' in Russian; tench, common tench, green tench].

Systematics

Cyprinus Tinca was described originally from European lakes. Ljabner et al. (2010) found the western and eastern phylogroups of tench were a single species under the biological species concept, although they were separate phylogenetic species. Tench from a western European and a Ponto-Caspian refugium came into contact by postglacial expansion after a separation of 750,000 years, and showed free interbreeding.

Key characters

The dark coloration, rounded fins and slimy body are characteristic of this species.

Morphology

Dorsal fin unbranched rays 2-4, usually 3-4 followed by 6-9, usually 8, branched rays, anal fin unbranched rays 3-4 and branched rays 5-9, usually 6-7, pectoral fin branched rays 13-18, and pelvic fin branched rays 7-10. Lateral line scales 70-120, gill rakers 10-16, usually 11-15, and vertebrae 35-44, usually 39-43. Anterior gill rakers may be difficult to distinguish from throat tubercles. Central rakers touch the one below when appresssed. Scales are very elongate ovals with a very anterior focus near the anterior scale margin. Radii are numerous on all fields. Pharyngeal teeth 4-5, 5-4, 5-3 or 4-4, more rarely 5-5, slightly to strongly hooked and expanded at the tip into a head supported on a narrow stalk. The oblique head has rounded edges and is concave in the middle. The gut is s-shaped with a small anterior loop. The chromosome number is 2n=48 (Klinkhardt et al., 1995).

Meristic values for Iranian specimens are:- dorsal fin branched rays 8(28); anal fin branched rays 5(1), 6(11), 7(15) or 8(1); pectoral fin branched rays 15(10), 16(13) or 17(5); pelvic fin branched rays 9(27) or 10(1); lateral line scales very embedded and obscured by skin and mucus, range about 82-95; total gill rakers 11(4), 12(7), 13(8), 14(7) or 15(2); pharyngeal teeth 5-4(5), 4-5(2) or 4-4(1); and total vertebrae 37(1), 38(7), 39(14) or 40(1).

Sexual dimorphism

The second pelvic fin ray in males is thickened and is accompanied by a muscular protuberance from the flank. Pelvic fins reach the anal fin in males.

Colour

The overall colour is a blackish-green, green or dark brown to bronze. Colour varies with habitat and is lighter in areas with less vegetation. The iris is red and the lips are yellowish-orange. Fins are grey to greenish or blackish. Golden, red and orange forms, some with black spots, may be bred in garden ponds and aquaria. Preserved fish have a dark body and fins with a lighter belly.

Size

Reaches 84.0 cm total length and about 8.5kg, possibly 10.0 kg.

Distribution

Found from the British Isles and the Iberian Peninsula across Europe to Siberia including southern drainages of the Black and Caspian seas. In Iran it is reported from the Anzali Mordab and neighbouring rivers to the Gorgan River (Holčík and Oláh, 1992; Riazi, 1996; Abbasi et al., 1999; Kiabi et al., 1999; Abdoli, 2000; Daghigh Roohi and Mokhayer, no date). Khara et al. (2005) record it from the Amirkelayeh Wetland near Lahijan.

Zoogeography

This widespread species has no close relatives among Eurasian cyprinids.

Habitat

Tench inhabit bays, small and shallow lakes, slow rivers and estuarine areas rich in vegetation. They are confined to the lower reaches of rivers in Iran and do not penetrate upstream (Berg, 1948-1949). Riazi (1996) reports that this species is native (resident) to the Siah-Keshim Protected Region of the Anzali Mordab. They are essentially inactive in winter. Tench are reported to bury themselves in mud during severe winters, lying dormant until spring. They can tolerate weak brackish water (to 12‰), acidic waters, low oxygen conditions and temperatures as high as 37°C for short periods. Preferred temperatures are 15-23.5°C (Brylińska, Bryliński and Bănărescu in Bănărescu, 1999). They remain alive for long periods when removed from the water. In Iran, adults have been caught in gill nets at 1-2 m (Annual Bulletin 1993-94, Iranian Fisheries Research and Training Organization, Tehran, p. 78-79, 1995).

Age and growth

Anonymous (2006) studied of tench in the Amirkelayeh Lagoon and found age groups 1+ to 7+ years. Vetlugina (1992) studied the biology of tench in the Volga Delta using sections of dorsal fin rays for aging. Life span is 13 years or more. Females grow faster than males and comprise 90-100% of older fish. Growth rate is similar to fish from Dagestan, nearer Iranian waters. Tench begin to mature at 3 years (30-40% of fish are sexually mature) and by age 4-5, 60% are mature. In Dagestan they mature at 3-4 years (Shikhshabekov, 1977). Life span is up to 30 years elsewhere. Turkish populations in lakes formed by dams range up to 5 or 6 years of age (Altindağ et al., 1998; 2002). These authors give age-length and age-weight relationships and condition factors.

Food

Diet comprises insect larvae, crustaceans, small molluscs, aquatic worms, vegetation and detritus. Chironomids may constitute as much as 91.8% of the diet in the second year of life while older fish favour snails and algae and macrophytes. This fish is probably a carnivore, switching to vegetation when animals are not available. Food may be picked out of vegetation or nosed out of mud as much as 13 cm deep using the snout (Brylińska, Bryliński and Bănărescu in Bănărescu, 1999). Anonymous (2006) studied the diet of tench in the Amirkelayeh Lagoon and found 17 food groups including Odonata, snails, water plants, Trichoptera, Chironomidae, Hemiptera, Ephemeroptera, Perca fluviatilis, Diptera, Gammarus, Tubifex, plant seeds, Simulium, water bugs, water ticks, zooplankton and phytoplankton. Phytoplankton, snails and Hemiptera had the highest frequenceis at 68.5%, 65.7% and 34.0% respectively. This fish is an omnivore and diet varied with season age and sex.

Reproduction

Spawning occurs in shallow water with little current and abundant vegetation. Spawning aggregations can be seen in April in the Volga Delta and spawning takes place in May and June (Vetlugina, 1992). Tench in the Anzali Mordab have well-developed, 1.0 mm eggs in early June. Water temperatures at peak spawning are 20-26°C. In Dagestan, spawning takes place in June-July at water temperatures no lower than 19-20°C (Shikhshabekov, 1977). Each female is accompanied by 2-3 males and different males will fertilise the egg batches as they are released. Tench spawn 2 or 3 times at intervals of about 20-30 days, shedding eggs onto surface vegetation. Eggs are greenish, adhesive and up to 1.57 mm in diameter. Fecundity is up to 863,000 eggs (or 124,850 per kg of body weight) and increases with age, length and weight. Larvae have attachment organs which enable them to hang onto plants for the first few days of life. Maximum fecundity is 1,241,200 eggs in eastern Europe.

Parasites and predators

Daghigh Roohi and Mokhayer (no date) record the trematode Asymphylodora tincae from this fish in the Anzali Lagoon, the first record of this parasite in Iran. Khara et al. (2005) found parasites of this species in the Amirkelayeh Wetland were Raphidascaris acus, Camallanus lacustris, Asymphylodora tincae, Diplostomum spathaceum, Dactylogyrus sp., Caryophyllaeus fimbriceps, Lernaea sp. and Trichodina sp. Khara et al. (2006a) record the eye fluke Diplostomum spathaceum for this fish in the Amirkelayeh Wetland in Gilan, although it had the lowest abundance of 7 species examined. Sattari et al. (2004, 2005) surveyed this species in the Anzali and Amirkelayeh wetlands, recording Raphidascaris acus larvae and Camallanus lacustris. Khara et al. (2006b) record the trematode Asymphylodora tincae from this species in the Boojagh Wetland of the Caspian coast.

Economic importance

Commercial catches are made in the Volga Delta where there are spring (April) and fall (September-October) fishing seasons. Up to 6,300 tonnes are taken using fyke nets, trap nets and seines. Holčík and Oláh (1992) report a catch of 540 kg in the Anzali Mordab in 1990.

The tench is a sought-after sport fish in Europe, putting on a strong fight for its weight (personal experience). There is an ornamental form, orange-yellow or reddish, which is kept in park ponds.

The roe or eggs of this species are particularly poisonous and it should be cleaned with care to avoid contamination of the flesh although it is not a common market fish in Iran (Halstead, 1967-1970; Coad, 1979b). Probable symptoms and treatment are given under the genus Schizothorax.

The tench is reputedly a "doctor fish" and other species are said to rub against its slimy body as a cure for injuries. Rubbed on humans, it is said to cure fever, headache, toothache and jaundice. In Iran, this species has been proposed as a component of polyculture as it feeds on snails, a host for diplostomiasis-causing parasites. A decrease in snail frequency varied from 43 to 82% in experiments using adult and juvenile tench (Iranian Fisheries Research Organization Newsletter, 37:3, 2003).

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use as food, in textbooks and because it has been introduced outside its natural range.

Conservation

Lelek (1987) classifies this species as intermediate in Europe (liable to be transferred to vulnerable or rare categories if their habitat deteriorates further). Kiabi et al. (1999) consider this species to be of least concern in the south Caspian Sea basin according to IUCN criteria. Criteria include commercial fishing, sport fishing, abundant in numbers, habitat destruction, widespread range (75% of water bodies), absent in other water bodies in Iran, and present outside the Caspian Sea basin.

There have been attempts to breed this species artificially in Iran to increase the recruitment rate and control loss (Annual Bulletin 1993-94, Iranian Fisheries Research and Training Organization, Tehran, p. 78-79, 1995; Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 39, 1996; Sirang, 1997). Brood fish weighed 0.2-0.45 kg and were kept in earthen ponds. Carp culture techniques were used with injection of gonadotropic hormones from carp pituitary at 3-8 mg per kg body weight applied in two doses. Water temperatures were 20-24°C, eggs were placed in saline carbamide solution after fertilisation to remove stickiness and washed in pure water. Swelled eggs measured 0.65-0.8 mm and were bright yellow to greenish-yellow. The number of eggs per spawner was 10,848-17,710 and 80-350 eggs per 1 g body weight. Fertilisation rate was 70-85%. Eggs were incubated for 2.0-2.5 days with a survival rate of hatched fry at 85-95% and size of newly hatched larvae at 4.5-5.0 mm.

Further work

The biology of this species has been well-studied elsewhere in its extensive range but not under Iranian conditions.

Sources

Iranian material: CMNFI 1970-0514, 2, 63.9-106.9 mm standard length, Gilan, Shafa River estuary (37º35'N, 49º09'E); CMNFI 1970-0553, 1, 70.9 mm standard length, Gilan, Sowsar Roga River (37º27'N, 49º30'E); CMNFI 1970-0580, 3, 28.0-36.2 mm standard length, Mazandaran, river near Iz Deh (36º36'N, 52º07'E); CMNFI 1979-0439, 1, 170.9 mm standard length, Gilan, Anzali Mordab (ca. 3727'N, ca. 4925'E); CMNFI 1980-0148, 14, 33.9-49.7 mm standard length, Gilan, Pir Bazar Roga River (37º21'N, 49º33'E); CMNFI 1980-0916, 6, 38.2-51.1 mm standard length, Gilan, Nahang Roga River (no other locality data); CMNFI 1993-0137, 1, 150.4 mm standard length, Mazandaran, Sari (36º34'N, 53º04'E).

Genus Tor
Gray1834

Much of the past literature on this genus appeared under Barbus (q.v.)

Tor grypus
(Heckel, 1843)

Above photographs showing fleshy lips courtesy of Atabak Mahjoor Azad

Common names

شيربت (= shirbot, shirbod or shilbot), shaboot, سس ماهي (= sos or sas mahi), rumi, shebhe shirbot, سرخه (sorkheh, meaning= reddish, a local name in the Zohreh River - J. Gh. Marammazi, pers. comm., 1995), rumi (not heard of in Khuzestan).

[shabout or hamrawi in Arabic; large scaled barb, Tigris barbel].

Systematics

Howes (1987) places this species outside the genus Barbus sensu stricto as defined by him because it has the non-elongate lachrymal bone with a sensory canal running along the antero-dorsal border, a derived condition. Karaman (1971) and Ekmekçi and Banarescu (1998) placed it in the genus Tor (see Barbus) and it may belong in Naziritor Mirza and Javed, 1985 (M. R. Mirza, pers. comm., 6 December 2003). Al-Hassan (1984) looked at several "Barbus" species and found the electropherogram of this species to be distinctive, perhaps indicating that molecular studies could resolve the relationships of this species. This distinction was reiterated by Jawad (2003).

The type locality of Barbus Grypus is "Tigris bei Mossul" (Heckel, 1843b) and Krupp (1985c) records a syntype (dried) from the Naturhistorisches Museum Wien (formerly NMW) now in the Senckenberg Museum Frankfurt under SMF 2613, 375 mm standard length. One syntype is in the Museum für Naturkunde, Universität Humboldt, Berlin (ZMB 8788, not located February 2006). One syntype is in Vienna under NMW 54160, 2 are under NMW 54161 (280.9-318.9 mm total length as measured by me), and 1 is under NMW 91023 (Eschmeyer et al., 1996). The catalogue in Vienna lists 3 fish in spirits and 2 fish stuffed.

Labeobarbus Kotschyi Heckel, 1843 described from the "Tigris bei Mossul" is a synonym. Krupp (1985c) records a syntype from the Naturhistorisches Museum Wien under NMW 49729, Th. Kotschy (188.8 mm standard length as measured by me). A dried specimen (NMW 59462) is also a syntype. Eschmeyer et al. (1996) also lists another syntype, NMW 91022. The catalogue in Vienna lists 1 fish in spirits and 2 fish stuffed.

Key characters

This species is identified by having two pairs of barbels, a strong, smooth spine in the dorsal fin, and less than 44 scales in the lateral line.

Morphology

The forehead is more rounded than in type material of kotschyi, although kotschyi types are smaller than grypus types which may account for this distinction. The mouth is inferior, horseshoe-shaped and has fleshy lips. The median lobe of the lower lip is well-developed in some individuals (such specimens were described as kotschyi - this form is rare in Khuzestan according to N. Najafpour, pers. comm., 1995) and intermediates can be seen) but not in others (grypus) (Karaman, 1971). The median lobe may extend back almost as far as the level of the rear margin of the lower lip or be distinctive with free lateral and rear margins but only extend back one third of this distance. The much fleshier lip structure in kotschyi (the upper lip can be reflexed for example) may be a form of hypertrophy seen in other cyprinid fishes (see Roberts and Khaironizam (2008) for further discussions on this feature)). The last unbranched dorsal fin ray is smooth and spine-like, with sharp edges but no serrations although serrations are weakly developed in young fish. Barbels are about equal in length. The gut has two anterior and two posterior loops.

Dorsal fin unbranched rays 4, branched rays 7-9, usually 8, anal fin unbranched rays 3, branched rays 5, pectoral fin branched rays 14-18 (including counts from Jawad (1975)), and pelvic fin branched rays 7-8, usually 8. Total gill rakers 16-22 (including counts by Jawad (1975)). Krupp (1985c) cites 13-17 gill rakers, presumably lower arch ones only. Gill rakers reach the second raker below or beyond when appressed, with large tubercles or branches on the inner surface in 2 rows alternating left and right. Lateral line scales 32-43. A pelvic axillary scale is present. Scales have a subcentral anterior, almost central, focus, numerous fine circuli and many radii on all fields with the exposed part of the scale tubercular. Pharyngeal teeth 2,3,4-4,3,2 in the literature, but see below, anterior teeth rounded, the most anterior one small and blunt, posterior ones spatulate with hooked tips. Total vertebrae 44-47 (Howes, 1987), 43-45 (Jawad, 1975) or 47(8), 48(7), 49(5) or 50(1) in fish seen by me (BM(NH) 1974.2.22:1283-1284, 1299-1315, 1317, 1323; 1920.3.3:1-18, 1874.4.28:24-6).

Ali et al. (1981) found differences in morphology for fish from Al-Therthar Reservoir and the Tigris River in Iraq, not by locality but by habitat type.

Iranian specimens have the following meristic data: dorsal fin branched rays 7(1), 8(20) or 9(1); anal fin branched rays 5(22); pectoral fin branched rays 15(2), 16(15), 17(4) or 18(1), and pelvic fin branched rays 7(1) or 8(21). Lateral line scales 32(4), 33(4), 34(4), 35(3), 36(3) or 37(3). Total gill rakers 16(4), 17(2), 18(3), 19(6), 20(4) or 21(2), with some evidence of higher counts in larger fish. Pharyngeal teeth usually 2,3,5-5,3,2(14) with variants 2,3,5-4,3,2(2), 2,3,4-5,3,2(2), 2,2,5-5,3,2(1) and 1,2,5-4,3,2(1), in contrast to literature reports of 4 main row teeth being typical.

Sexual dimorphism

Ali et al. (1981) found no sexual dimorphism in their Iraqi samples.

Colour

Overall colour has a pale rose to light orange effect, usually without other markings. The back is a dark olive-brown to blackish-green with the flanks pale rose, light orange to yellowish to silvery and belly silvery to milk-white. There may be an indistinct stripe along the mid-flank. Large fish have the upper flank darkened from the overall orange colour of the mid-flank and the lower flank scales are rimmed in white so they stand out. Lips are pale red. The operculum is golden. The pectoral, pelvic, anal and caudal fins are bright orange or pink at the base (perhaps white after preservation), distally blackish. Pectoral and pelvic fins may be dark overall with a reddish to reddish-brown tinge, and the leading edge of the pelvic fin pink. The anal fin may be a bluish-black distally. In some fish the caudal fin is black proximally and reddish distally. In large fish the pectoral, pelvic, anal and caudal are progressively darker in this order. The anal and pelvic fins, the pectoral fins less so, may be heavily pigmented with melanophores on rays and membranes so as to appear black in preserved fish. The dorsal fin is hyaline. The eye rim is yellow-green to lime-green. Young fish may have some scales darkened, giving a mottled effect and are more silvery on the flank than large fish. Their pectoral and pelvic fins are more orange and the anal and caudal fins are only slightly tinged with colour. The caudal fin carries a lot of grey. The smallest fish have a very faint fin colouration. Peritoneum black.

Size

Attains 96.0 cm and 9.7 kg in Dukan Reservoir, Iraq and 96.0 cm and 11.0 kg in Atatürk Dam Lake on the Euphrates River in Turkey (Al-Hakim et al., 1981; Oymak et al., 2008). Gruvel (1931) cited 1.5 m and 30 kg for Syria. Banister (1980) gives nearly 2 m and 100 kg but this may be confusion with B. esocinus although Krupp (1992) also cites almost 2 m. Reputedly reaches 60 kg in Lorestan (S. Nazeeri, pers. comm., 2000) and Ghofleh Marammazi (2004) found fish up to 20 kg in Khuzestan. Specimens reach 3 kg even in the small canals of the sugar-cane fields of Khuzestan.

Distribution

This species is found in the Tigris-Euphrates basin and the Orontes River basin. In Iran it is found in the Tigris River basin up to the Simarreh including marshes such as the Hawr Al Azim, in the Gulf basin in the Zohreh, Shapur, Helleh, Dalaki, Dasht-e Palang, Shur, Dozgah and Mand rivers and Lake Famur (= Perishan), although rare in the latter, and in the Hormuz basin in the Hasan Langi and Kul rivers (Berg, 1949; Gh. Izadpanahi, pers. comm., 1995; Marammazi, 1995; M. Rabbaniha, pers. comm., 1995; Maafi, 1996b; H. R. Alizadeh, pers comm., 2000; Abdoli, 2000).

Zoogeography

Karaman (1971) considers this species to have an Indian line of descent, placing it in the genus Tor which most subsequent authors restrict to the Indian subcontinent and southeast Asia.

Habitat

van den Eelaart (1954) and Al-Hamed (1966b; 1972) describe the habitat for this species in the Tigris River as distributed throughout the river and its tributaries. It is a strong swimmer. Mature fish move upstream to the spawning grounds and spent fish descend to their original habitat. In summer under low water level conditions and high temperatures, the smaller fish remain in the lower reaches of rivers but the larger fish migrate up rivers and tributaries, returning in September and October when temperatures fall. This species may enter marshes on floods, favouring areas where there is fresh river water, but returns to rivers as it requires a higher oxygen concentration than most marsh residents. Heydarpour (1978) gives a temperature range of 9-31ºC for this species under culture conditions in Khuzestan.

Marammazi (1994) considers this species to be versatile in its habitats in the Zohreh River which drains to the northern Persian Gulf. It was found throughout the river in contrast to Mesopotamichthys sharpeyi which, being stenohaline, was restricted in its distribution. The form with a well-developed median lobe is said to occur in rocky habitats. This species is considered to be the dominant fish in the Karun and Zohreh rivers (Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 48, 1996; Iranian Fisheries Research and Training Organization Newsletter, 17:1, 1997). Ghofleh Marammazi (2004) found it in almost all water bodies in Khuzestan where it occurred under a wide range of temperatures and salinities. However its presence on the Khuzestan plain was for feeding while for reproduction it required more northerly areas with sandy or gravel substrates, high water flow, low temperatures and high oxygen content (Ghofleh Marammazi, 2000). Ramin (2009) records this species as the most abundant in the Karkheh River out of 37 species and subspecies.

Age and growth

Al-Hakim et al. (1976) studied some aspects of the biology of this species in Razzaza Lake, Iraq. Males are longer than females before maturation and shorter thereafter. Females reach 13 years and males 8 years of age and fish mature at 45-48 cm total length in their fifth year. Males mature earlier than females. Ali et al. (1981) found this species to mature at 3-5 years of age and 40-50 cm in the Al-Therthar Reservoir (about 65 km northwest of Baghdad) and the Tigris River (Kut Dam) in Iraq. Jiad et al. (1984) studied this species in the Al-Hindiya Dam in Iraq and found similar results to the studies cited above.

Al-Hakim et al. (1981) studied this species in the Dukan Reservoir, west of Sulaimaniyah, Iraq. Life span is 17 years for females and 11 years for males. Growth slows with age, and especially after maturity, and is fastest in the first year of life. 30% of males mature at age group 3 (39 cm) and all were mature at age group 6 (48 cm). Al-Hamed (1966a; 1966b; 1972) working with Tigris River populations in Iraq, found males to mature at about 45 cm and females at about 50 cm, with most fish mature in their fourth year and spawning at the beginning of their fifth year of life. Some fish mature in age group 3 and some as late as age group 5. Maximum age observed was 12 years. Males outnumber females, being two thirds of the fish on the spawning grounds. Al-Hakim et al. (1981) found all females are mature at 51 cm (age group 7) but only 10% at 42 cm. Males mature earlier than females and may grow faster and die younger.

Ali et al. (1981) found growth to be better in fish from Al-Therthar Reservoir compared to those from the Tigris River in Iraq. The fishing methods used, both commercially and experimentally, caught mature fish of 40-50 cm and 3-5 years of age.

Oymak et al. (2008) examined age and growth in the Atatürk Dam Lake on the Euphrates River in Turkey. Fourteen age classes were found with age classes 4-6 for females and 2-4 for males dominant. von Bertalanffy growth equations were given for males and females.

Growth in a polluted section of the Diyala River, Iraq is poor compared to other populations (Khalaf et al., 1984; Khalaf et al., 1985).

Food

Al-Hamed (1965) found this species to be a herbivore taking filamentous algae and higher plant parts. Incidental food items taken while feeding on plants include fish tissue and scales. Fallen ripe fruits from trees overhanging the water are also consumed as are cereal grains from loading docks. It may also take some small fishes. Iranian specimens contain filamentous algae, plant fragments and associated invertebrates. Ghofleh Marammazi (2000) considers it to be an omnivore.

Reproduction

In Dukan Reservoir, Iraq spawning takes place from the beginning of May until the end of June. van den Eelaart (1954) and Al-Hamed (1966b; 1972) studied the reproduction of this species in Iraq. Eggs are deposited on fine gravels overlying a layer of coarse sand in shallow, wide holes. Water depth varies from 30 to 150 cm. Egg diameter is 1.5 mm and fecundity up to 147,000. The spawning season on the Tigris River between Beled and Tigrit is late May to late June after an upriver migration in April. Fish appear on the spawning grounds in schools just before dark and remain there until shortly before midnight, making loud noises by splashing, jumping and chasing. After spawning, the fish return downriver but do not enter marshes as these are now too warm. Oymak et al. (2008) examined reproduction in the Atatürk Dam Lake on the Euphrates River in Turkey and found a sex ratio of 1:1.34 (females/males), with a spawning period in May to July, a fecundity up to 235,764 eggs and a mean egg diameter of 2.183 mm.

Its presence in areas of the Khuzestan plain is mainly for feeding while reproduction occurs in the northern parts of this province where there are sandy and gravel substrates, fast current, low temperatures and high oxygen content (Ghofleh Marammazi, 2004).

Parasites and predators

Bykhovski (1949) report a new species of monogenetic trematode, Dactylogyrus pavlovskyi, from this species in the Karkheh River, Iran. Molnár and Jalali (1992) describe a new species of monogenean, Dogielius persicus, from this species in the Dez and Karun rivers of Khuzestan. Masoumian et al. (1994) describe two new species of Myxosporea from the gills of this species in the Karun River, Khuzestan, namely Myxobolus karuni and Myxobolus persicus. Molnár et al. (1996) report additional new species from this fish in Khuzestan, namely Myxobolus iranicus in the spleen and Myxobolus mesopotamiae in connective tissue of the caudal and pectoral fins. The latter myxosporean is also reported from Barbus rajanorum as is a new species Myxobolus shadgani infecting the gills - the identity of the host fish is unknown as Barbus rajanorum is not a distinct species (see under Luciobarbus pectoralis). Myxosporeans are potentially dangerous to fishes such as Tor grypus which may be used in fish culture in Khuzestan.

The monogeneans Dactylogyrus povlovskyi, D. barbioides, Dogielius persicus, Gyrodactylus sprostonae and Paradiplozoon sp. are recorded from this species in the Karun River with heavier infestations in spring and summer than in autumn and winter. These gill parasites caused no serious injuries but were thought to be important in respect of monitoring infestation levels on fish farms in Khuzestan (www.avz1.8m.com/fulltext.htm, downloaded 28 October 2002).

Ebrahimzadeh and Nabawi (1975) list a nematode species Philometra and Ascaridae from this species in the Karun River. Ebrahimzadeh and Kailani (1976) record parasites in the genera Myxosoma (protozoan) and Isoglaridacris (cestode) and also a nematode from Barbus (= Tor) grypus taken in the Karun River.

Masoumian and Pazooki (1999) list Myxobolus persicus, M. karuni, M. mesopotamiae and M. iranicus from this species in various localities in Khuzestan. Moghainemi and Abbasi (1992) record a wide range of parasites from this species in the Hawr al-Azim in Khuzestan. Farahnak et al. (2002) record Contracaecum sp. and Anisakis sp. from this fish in Khuzestan Province.

Papahn et al. (2004) record the monogeneans Dactylogyrus pavlovskyi, D. barbioides, Dogielius persicus, Gyrodactylus sprostonae and Paradiplozoon sp. from this species in the Karun River at Ahvaz, the latter two being first records for the host in Iran.

Economic importance

An important food fish, comprising 23% of the total fish production in Iraq for example and forming the most important commercial fish there (Al-Hakim et al., 1981). Petr (1987) reports the catch for all Iraq in 1976 as 519 t. The weight at the Basrah fish market from October 1975 to June 1977 was only 3,330 kg however (Sharma, 1980) and Khalaf et al. (1984) rank it third in the inland wholesale trade of Iraq for the period 1967-1970.

This species is the preferred catch of anglers at Ahvaz in Khuzestan, with bread or potato as bait. There is a good demand for this species in local markets of Khuzestan (Ghofleh Marammazi, 2004).

Petr (1987) has suggested that this species be investigated for fish farming in Khuzestan. The Khuzestan Fisheries Research Centre at Ahvaz has experimented with this species in pond culture (Emadi, 1993a; Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 6, 1996).

Conservation

The stock of this species in the Gav Masiab River is severely reduced and only 3 fish were caught in western Iran in the Zagros rivers during a 4-year survey (J. Valiallahi, www.modares.ac.ir, downloaded 4 July 2000; pers. comm., 2001). Shirbot were considered to be on the verge of extinction in the Gav Masiab River of the Tigris River basin, through pollution, overfishing, dam building, aquaculture, and introduction of exotics (IranMania.com, 29 December 2006). Probably in decline in Turkey (Fricke et al., 2007).

Further work

The biology of this species in Iran needs investigation as does its putative relationship to South Asian fishes in the genus Tor.

Sources

Type material: See above, Barbus grypus (NMW 54161) and Labeobarbus kotschyi (NMW 49729).

Iranian material: CMNFI 1979-0155, 1, 42.8 mm standard length, Fars, spring at Gavanoo (28º47'N, 54º22'E); CMNFI 1979-0291, 1, 31.4 mm standard length, Kermanshahan, Diyala River drainage (34º24'N, 45º37'E); CMNFI 1979-0356, 5, 22.5-40.7 mm standard length, Khuzestan, Karkheh River drainage at Hoveyzeh (31º27'N, 48º04'E); CMNFI 1979-0360, 1, 375.5 mm standard length, Khuzestan, canal branch of Karkheh River (31º40'N, 48º35'E); CMNFI 1979-0364, 1, 22.3 mm standard length, Khuzestan, river at Abdolkhan (31º52'30"N, 48º20'30"E); CMNFI 1979-0384, 2, 202.2-222.2 mm standard length, Khuzestan, Ab-e Shur drainage (32º00'N, 49º07'E); CMNFI 1979-0391, 1, 220.5 mm standard length, Khuzestan, stream in Marun River drainage (31º28'N, 49º51'E); CMNFI 1979-0392, 1, 62.4 mm standard length, Khuzestan, Zard River (ca. 31º32'N, ca. 49º48'E); CMNFI 1979-0395, 2, 32.7-38.0 mm standard length, Khuzestan, stream in Marun River drainage (ca. 30º57'N, ca. 49º51'E); CMNFI 1979-0402, 1, 80.7 mm standard length, Bushehr, Mand River 12 km north of Kaki (ca. 28º25'N, ca. 51º32'E); CMNFI 1991-0153, 1, 253.5 mm standard length, Khuzestan, Zohreh River (no other locality data); CMNFI 1993-0141, 1, 80.2 mm standard length, Bushehr, Dalaki River (29º28'N, 51º15'E); CMNFI 1995-009A, 4, Khuzestan, A'la River at Pol-e Tighen (31º23'30"N, 49º53'E); BM(NH) 1980.8.28:7, 1, 72.0 mm standard length, Khuzestan, Dezful (32º23'N, 48º24'E); ZMH 2508, 1, 343.3 mm standard length, Khuzestan, Karun River at Ahvaz (31º19'N, 48º42'E); ZSM 21864, 1, 157.7 mm standard length, Khuzestan, Dez River at Harmaleh (31º57'N, 48º34'E);

Comparative material:- BM(NH) 1874.4.28:24-26, 3, 231.2-254.7 mm standard length, Iraq, Tigris River at Baghdad (33º21'N, 44º25'E); BM(NH) 1920.3.3:1-18, 5, 104.8-196.0 mm standard length, Iraq, Basrah (30º30'N, 47º47'E); BM(NH) 1973.5.21:191, 1, 205.4 mm standard length, Iraq, Shatt-al-Arab; BM(NH) 1973.5.21:192, 1, 139.5 mm standard length, Iraq, Shatt-al-Arab; BM(NH) 1974.2.22:1283-1284, 2, 121.0-164.5 mm standard length, Iraq, Khalis (no other locality data); BM(NH) 1974.2.22:1299-1315, 9, 64.9-101.5 mm standard length, Iraq, branch of Khalis River (no other locality data); BM(NH) 1974.2.22:1317, 93.9 mm standard length, Iraq, branch of Khalis River (no other locality data); BM(NH) 1974.2.22:1323, 1, 160.1 mm standard length, Iraq, Basrah (30º30'N, 47º47'E); BM(NH) 1974.2.22:1328, 1, 161.9 mm standard length, Iraq, Basrah (30º30'N, 47º47'E); KU 10516, 1, 124.1 mm standard length, Iraq, Basrah (30º30'N, 47º47'E).

Genus Vimba
Fitzinger, 1873

This genus is found in the basins of the Baltic, Black and Caspian seas and has a single species. It is characterised by a compressed, moderately deep body with an inferior, crescentic mouth, a scaleless keel between the pelvic and anal fins, a scaleless groove in front of the dorsal fin and an evident keel behind it, pharyngeal teeth in a single row, short dorsal and long anal fin, gill rakers short, and scales moderate in size. Bogutskaya (1986) using skull morphology reaffirms the generic separation of Vimba Fitzinger, 1873 from Abramis Cuvier, 1816 although Howes (1981) considers it to be a synonym.

Vimba persa
(Pallas, 1814)

CMNFI 1979-0435, Gilan, stream 10 km west of Ramsar

Common names

سياه كولي (= siah kuli or siahkooli, meaning black fish, cooli, couli, coli, kooli or kuli being any small fish; in Gilaki), mahi siah kuli.

[garasol in Azerbaijan; chernospinka, Kaspiiskii rybets or Caspian vimba, both in Russian; southern white-eye, vimba].

Systematics

Cyprinus persa Gmelin, 1774 is a nomen nudum - see Kottelat (1997) - and was later made available by Pallas. It was described originally from "Persa; in lacubus ad Cyrum", i.e. the southern coast of the Caspian Sea in lakes of the Kura River system in Azerbaijan. It is distinguished by larger scales and usually fewer anal rays from V. vimba (Linnaeus, 1758) which it was formerly synonymised with or recognised as a subspecies of. Cyprinus Vimba was originally described from lakes of Sweden. Hänfling et al. (2009) found phylogenetically distinct mtDNA sequences for Caspian Sea basin samples, the Caspian clade having diverged from a western or Pontic clade 1-2 MYA at the beginning of the Pleistocene. Caspian populations could then rank as a separate species or subspecies although they consider further work involving western Caucasian populations is needed to support one conclusion over the other. Naseka and Bogutskaya (2009) recognise V. persa as a species.

A hybrid with Alburnus chalcoides was reported from the Safid River (Petrov, 1926).

Key characters

The snout projects over the lower jaw and in large fish is quite bulbous, there is a keel on the belly and on the back, and fin ray counts are distinctive.

Morphology

Western and southeastern populations in the Caspian Sea can be distinguished morphometrically, and represent different stocks.

Dorsal fin with 2-3 unbranched rays (always 2 in the subspecies persa (Berg, 1948-1949) but the first unbranched ray is minute and visible in x-rays in Iranian specimens) and 7-9, usually 8, branched rays, anal fin with 3 unbranched and 12-22 branched rays (16-18 in persa but see below), pectoral fin branched rays 11-18 and pelvic fin branched rays 7-10. Lateral line scales 47-64, 48-54 in persa (but see below). The lateral line runs below the midline of the caudal peduncle. Predorsal scales are small and crowded. A pelvic axillary scale is present. The naked ventral keel begins 0-3 scales behind the pelvic fin bases. Scales at the anal fin base form a sheath. The anterior scale margin is wavy and the posterior margin is crenualte. There is a central focus, numerous fine circuli and few anterior and posterior radii. Gill rakers 12-20, small and reaching the raker below when appressed. Pharyngeal teeth usually 5-5, with the largest teeth having long and narrow, flat to slightly concave crowns, and tips recurved or very slightly hooked. Vertebrae 38-45. Gill rakers 14-19. The gut is s-shaped. The chromosome number is 2n=50 (Klinkhardt et al., 1995) or 2n=52 (Reshetnikov, 2002).

Meristic values for Iranian specimens are:- branched dorsal fin rays 7(1) or 8(39); branched anal fin rays 16(3), 17(13), 18(18) or 19(6); branched pectoral fin rays 14(8), 15(22), 16(7) or 17(3); branched pelvic fin rays 8(10) or 9(30); lateral line scales 47(1), 48(4), 49(7), 50(10), 51(14), 52(1), 53(1), 54(1) or 55(1); total gill rakers 15(1), 16(2), 17(12), 18(14), 19(9) or 20(2); pharyngeal teeth 5-5(15), 5-4(4) or 4-5(1); and total vertebrae 41(3), 42(9), 43(24) or 44(3). Abbasi et al. (2004) found 149 Safid River fish to have mean values of 50.83 lateral line scales, branched dorsal fin rays 7.96 and branched anal fin rays 17.58.

Sexual dimorphism

Females are slightly larger than males of the same age and differ morphometrically on account of the eggs distorting body shape. The males become black on the back, reddish on the belly, their fins become red and the tips of the dorsal and caudal fins become dark, and they develop minute tubercles on the body during the spawning season (Kuliev, 1988; Abbasi et al., 2004). Females may also develop tubercles but to a lesser extent. Safid River fish showed differences in 2 meristic and 16 morphometric characters, especially body depth and lengths of dorsal, pectoral, pelvic and anal fins (Abbasi et al., 2004).

Iranian specimens have small tubercles lining the scale margins and larger tubercles over the whole head but particularly on the dorsal surface and upper sides. Fin rays bear small tubercles in files following the branching of the rays. The pelvic fin has weakly developed tubercles on its ventral surface as well as dorsally. The unbranched pectoral and pelvic rays bear several rows of tubercles.

Colour

The back is a reddish-brown to grey-blue, flanks are silvery and the belly yellowish. Paired fins are red at the base, pink distally. The anal fin base is red while other fins are grey to hyaline. Spawning fish develop black stripes along the dorsal and ventral body.

Size

Reaches 60.0 cm and 3 kg. The subspecies persa is smaller, to 30 cm.

Distribution

Found from central Europe to the Caspian Sea basin. In Iran it is recorded from the Aras to the Atrak rivers in the Caspian Sea basin including the Manjil Reservoir on the Safid River, the Anzali Mordab and Gorgan Bay, and the southeast Caspian Sea, southwest Caspian Sea and south-central Caspian Sea (Kozhin, 1957; Nümann, 1966; Holčík and Oláh, 1992; Riazi, 1996; Abbasi et al., 1999; Kiabi et al., 1999; Abdoli, 2000). It has also been introduced to Sistan.

Zoogeography

This species reaches its most south-easterly occurrence in Iran and has been given subspecific status. Its relationships lie with European taxa (see Abramis).

Habitat

Caspian vimba have a sparse distribution in the sea and are not fished there commercially. It is more common in Gilan than Mazandaran and Golestan coastal waters (Jolodar and Abdoli, 2004). The semi-migratory form enters fresh water or brackish water only for reproduction in spring. After spawning, it migrates to river mouths to feed until the next reproductive season (Kuliev, 1988). Riazi (1996) reports that this species is native (resident) to the Siah-Keshim Protected Region of the Anzali Mordab. S. Bazari Moghaddam (www.meeresschule.com/cgi-bin/abstracts/gastbuch.asp, downloaded 17 January 2005) records a migration into the Safid River in spring for reproduction. Feeding continues on this migration. Knipovich (1921) reports this species from depths of 36.6-53.0 m in the Iranian Caspian Sea. In fresh water it occurs in schools in the lower reaches of rivers, in deep water over stone and gravel bottoms. It may also occur in lakes over mud bottoms.

Age and growth

Maturity is attained in the second or third year of life, males maturing at age 2 in the Anzali region. In the Safid River migrating fish were 2-4 years old, predominately 3-year-old fish (S. Bazari Moghaddam, www.meeresschule.com/cgi-bin/abstracts/gastbuch.asp, downloaded 17 January2005). Four-year-olds predominate in the spawning population in Kyzylagach or Imeni Kirova Bay, Azerbaijan. Most spawning females are 16-23 cm (46%) and males 13-19 (42%). Large fish spawn first and the number of smaller fish spawning increases towards the end of the reproductive season (Kuliev, 1988; Shikhshabekov, 1979). Most fish on the spawning migration into the Anzali Mordab are 170-250 mm and ages 3-4 years (Holčik and Oláh, 1992). Fish on the spawning migration of the Safid River had a fork length of 116-208 mm and a weight of 21.1-116.1 g in males and 122-222 mm and 23.1-170.0 g in females. Spawning males were 2-6 years old and females 3-7 years (Abbasi et al., 2005). Maximum life span is about 15 years.

Food

Diet is aquatic insects, crustaceans, snails, worms and algae on muddy bottoms. Iranian specimens had zebra mussels and insect remains. S. Bazari Moghaddam (www.meeresschule.com/cgi-bin/abstracts/gastbuch.asp, downloaded 17 January 2005) reports oligochaetes, chironomids and Odonata in fish from the Safid River.

Reproduction

Kuliev (1988) and Shikhshabekov (1979) studied reproduction in the Kyzylagach Bay of the southwestern Caspian Sea and the waters of Dagestan respectively. The spawning migration begins in March or April at 10-13°C and spawning takes place at the end of April at 16-20°C, continuing until the end of May or into June.

Fish enter the Anzali Mordab of Iran in mid-January at a water temperature of 8-9°C, peaking from 21 April-10 May at 19-21°C (Holčík and Oláh, 1992). Khaval (1998) reports a spawning migration into the Safid River despite construction, sand removal and pollution. Fish from the Shafa River estuary in Iran caught on 10 April had highly developed eggs measuring 1.3 mm. Fecundity is up to 89,200 eggs per female, increasing with age and body size (elsewhere in Europe to 200,000 eggs with a diameter of 1.4 mm). Spawning is non-intermittent, in contrast to Black Sea vimbas. Eggs are deposited on gravel or stones where there is a current of 0.6-0.9 m/second. Concrete structures and flooded fields may be used as long as there is some current (Holčík and Oláh, 1992). The eggs may form a layer up to 10 cm thick. Initially attached to plants or stones, the eggs are later washed down between the plants or stones. Other fishes eat these eggs and mortality is high. Some fish deposit eggs in sandy shallows of bays or on the roots of reeds and bulrushes. The young migrate to the coastal zone of the Caspian Sea for the summer, moving to greater depths as winter approaches. At temperatures of 17-22°C, eggs incubate for 70-77 hours (Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 37-38, 1996).

Abbasi et al. (2005) found that the Safid River population started the spawning migration in March and this continued until July, peaking in mid-April to late May. Gonad weight for females increased with distance from the estuary. Spawning occurred from late May to late June, peaking in May at 18-29ºC water temperatures. Eggs were shed on pebble and gravel grounds 25-75 km from the estuary. The Disaam tributary was the major spawning site.

Parasites and predators

Jalali and Molnár (1990a) record the monogeneans Dactylogyrus cornoides and D. haplogonus from this species in the Safid River and Pazooki and Aghlmani (no date) record the trematode Asymphylodora kubanicum from Iranian specimens. Sattari et al. (2007) record the cestodes Caryophyllaeus laticeps and Caryophyllaeus fimbriceps, the digenean Diplostomum spathaceum and the monogenean Dactylogyrus extensus in this species in the Anzali wetland of the Caspian shore.

Economic importance

The vimba catch over the whole Caspian Sea basin was less than 100 tonnes per year in the 1980s (Kuliev, 1988). The catch by local fishermen in the Anzali Mordab region in 1990-1991 was 823 kg or about 8400 fish (Holčík and Oláh, 1992). They are caught in rogas and inflowing rivers of the mordab in late winter and early spring. In 1994-1995, the population of this species was noted as declining in recent years (Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 37-38, 1996).

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

Conservation

Weirs are a problem for this species in Iran as they block the spawning migration, the fish massing below the obstruction, and causing re-absorption of eggs and sperm (Holčík and Oláh, 1992). Aquaculture of this species has been investigated in Iran; it can be bred semi-artificially using methods similar to that for Rutilus species (Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 38, 1996).

Lelek (1987) classifies this species as intermediate to rare in northwestern Europe.

Kiabi et al. (1999) consider this species to be near threatened in the south Caspian Sea basin according to IUCN criteria. Criteria include commercial fishing, sport fishing, abundant in numbers, habitat destruction, widespread range (75% of water bodies), absent in other water bodies in Iran, and absent outside the Caspian Sea basin. Mostafavi (2007) lists it as near threatened in the Talar River, Mazandaran.

Further work

The relationships of the Caspian Sea taxon to other Vimba needs investigation as do certain aspects of its biology in Iranian waters as it is under some threat.

Sources

Iranian material: CMNFI 1970-0531, 8, 44.4-70.5 mm standard length, Mazandaran, Larim River (36º46'N, 52º56'E); CMNFI 1970-0543A, 9, 38.9-87.4 mm standard length, Gilan, Caspian Sea at Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1970-0544, 1, 162.2 mm standard length, Gilan, Caspian Sea near Bandar-e Anzali (37º28'N, 49º27'E); CMNFI 1979-0431, 1, 145.0 mm standard length, Mazandaran, fish bazaar at Now Shahr (no other locality data); CMNFI 1979-0435, 1, 141.1 mm standard length, Gilan, stream 10 km west of Ramsar (36º57'N, 50º37'E); CMNFI 1979-0436, 5, 115.1-156.5 mm standard length, Gilan, stream 26 km west of Ramsar (37º02'30"N, 50º27'E); CMNFI 1979-0437, 1, 155.2 mm standard length, Gilan, Safid River, 2km west of Astaneh (37º16'30"N, 49º56'E); CMNFI 1979-0438, 2, 139.5-152.6 mm standard length, Gilan, Gholab Ghir River (37º27'N, 49º37'E); CMNFI 1979-0788, 2, 28.5-50.7 mm standard length, Mazandaran, Gorgan River (37º00'N, 54º07'E); CMNFI 1980-0120, 7, 47.8-65.5 mm standard length, Mazandaran, Babol River at Babol Sar (36º43'N, 52º39'E); CMNFI 1980-0121, 3, 139.5-152.2 mm standard length, Gilan, Shafa River estuary (37º35'N, 49º09'E); CMNFI 1980-0126, 1, 170.8 mm standard length, Gilan, Caspian Sea near Bandar-e Anzali (37º28'N, 49º27'E); CMNFI 1980-0127, 2, 180.3-182.1 mm standard length, Gilan, Caspian Sea near Hasan Kiadeh (37º24'N, 49º58'E); CMNFI 1980-0129, 1, 51.9 mm standard length, Mazandaran, Tajan River (36º49'N, 53º06'30"E); CMNFI 1980-0138, 10, 28.1-105.0 mm standard length, Gilan, Safid River estuary (ca. 37º28'N, ca. 49º54'E); CMNFI 1980-0142, 1, 180.1 mm standard length, Gilan, Nahang Roga River (no other locality data); CMNFI 1980-0908, 4, 87.8-135.7 mm standard length, Gilan, Safid River estuary (ca. 37º28'N, ca. 49º54'E).

Basin/Dorsal fin branched rays	7	8	9	x	S.D.
Hormozgan	27	6		7.2	0.39
Lake Maharlu		90	2	8.0	0.15
Sa`di's Tomb	5	20		7.8	0.41
Gulf	16	173	1	7.9	0.29
Tigris River	15	176	3	7.9	0.30

Basin/Pectoral fin branched rays	10	11	12	13	14	15	16	18	x	S.D.
Hormozgan			1	3	18	10	1		14.2	0.78
Lake Maharlu		1	2	15	50	23	1		14.1	0.78
Sa`di's Tomb			5	16	2	2		1	13.2	1.24
Gulf		1	4	57	84	42	1		13.9	0.81
Tigris River	1	1	10	54	93	33	2		13.8	0.88

Basin/Pelvic fin branched rays	6	7	8	9	x	S.D.
Hormozgan		20	12		7.4	0.49
Lake Maharlu		8	84		7.9	0.28
Sa`di's Tomb		9	16		7.9	0.27
Gulf	1	110	79		7.4	0.50
Tigris River	2	51	138	3	7.7	0.50

Basin/Lateral line scales	25	26	27	28	29	30	31	32	33	34	35	36	37	38	40	x	SD
Hormuz							2	7	19	10	1					33.0	0.87
Lake Maharlu					1	4	13	21	32	30	9					31.2	1.54
Sa`di's Tomb						2		6	6	7	4					33.1	1.39
Gulf	1	1	5	16	25	24	25	42	36	19	5					31.2	2.06
Tigris River						2	2	24	44	59	45	9	2	1	1	33.9	1.34

Basin/Total vertebrae	32	33	34	35	36	37	x	S.D.
Hormozgan		7	24	7	3		34.2	0.78
Lake Maharlu		6	38	21	3		34.3	0.70
Sa`di's Tomb			14	9			34.4	0.50
Gulf	2	15	36	34	9		34.3	0.93
Tigris River		23	52	76	17	2	34.5	0.92