Freshwater Fishes of Iran

Species Accounts

Cyprinidae

Capoeta
to Mylopharyngodon

 

Revised: 07 November 2010

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Genus Capoeta
Valenciennes, 1842

The genus Capoeta has a wide distribution in Southwest Asia and contains about 20 species of which 7 occur in Iran. Its affinities are uncertain and may lie with the European Barbus/Aulopyge group or with Cyprinion and its southern and east Asian relatives (Karaman, 1971; Howes, 1982; Krupp, 1985c; Bănărescu, 1992b).

Varicorhinus Rüppell, 1836 (as used for Southwest Asian cyprinids) is a synonym of Capoeta Valenciennes, 1842 (see Karaman (1969) for further details: Capoeta is distinguished from Varicorhinus of Africa since it has a denticulate last unbranched dorsal fin ray (as opposed to smooth), very small to medium-sized scales (large), lachrymal bone narrow and covering only a small part of the upper side of the rostrum (large and covering most of the rostrum), suborbital bones narrow and long (short and wide), posterior maxillary process not extending back to a level with the centre of the jugal (extends back to a level of the centre of the suborbitals), lower jaw long (short). Scaphiodon Heckel, 1843 has been used for Capoeta and Cyprinion species in Southwest Asia. The nomenclatural status of this genus is reviewed by Bănărescu in Bănărescu (1999).

This genus Capoeta is characterised by a compressed to rounded and moderately elongate body, small to moderately large scales (lateral line counts 37-99), scales at the anal fin base and anus not usually enlarged (sometimes variably enlarged as is the case with certain cyprinids), an inferior, transverse mouth, the lower jaw with a sharp, horny sheath, barbels absent or in 1 or 2 pairs, dorsal fin short (usually 7-9 branched rays) with the last unbranched ray thickened and bearing serrations (serrations sometimes reduced to absent), anal fin short (usually 5 branched rays), gill rakers short, moderate to numerous, pharyngeal teeth in 3 rows with spoon-shaped and truncate tips, a very long and coiled gut (ca. 7-10 times body length), mostly of uniform colour, and a black peritoneum.

The general name for the members of this genus in northern Iran is سياه ماهي (= siah mahi, meaning black fish) while in the south they are called twiny or touyeni and even gel cheragh (= mud-eater, mud-grazer). The name Capoeta is derived from the Armenian and Georgian name for female Capoeta capoeta packed with eggs, namely "Kapwaeti". Other general names for members of this genus shol khar, ghel khar or choul khar, all variant spoken intonations meaning mud eater.

The origin of Capoeta in Southwest Asia follows the same route as the genus Barbus (q.v.).

CMNFI 1977-0510A, 4, mm standard length, Fars, qanat at Naqsh-e Rostam (29º59'30"N, 52º54'E); CMNFI 1979-0026, , mm standard length, Fars, Shapur River at Shapur (29º47'N, 51º35'E); CMNFI 1979-0027, , mm standard length, Fars, Chehel Chashmeh (ca. 29º43'N, ca. 52º02'E); CMNFI 1979-0036, 2, 83.9-118.3 mm standard length, Fars, Shapur River at Shapur (29º47'N, 51º35'E); CMNFI 1979-0043, , mm standard length, Fars, qanat behind Sarvestan (29º16'N, 53º14'E); CMNFI 1979-0044, , mm standard length, Fars, qanat at Mian Jangal (29º09'N, 53º27'E); CMNFI 1979-0053, 6, 47.3-79.5 mm standard length, Fars, Shur River tributary (ca. 28-29º58-03'N, ca. 52º34-35'E); CMNFI 1979-0054, 16, 35.8-127.9 mm standard length, Fars, Shur River tributary (ca. 28-29º58-03'N, ca. 52º34-35'E); CMNFI 1979-0057, , mm standard length, Fars, Shapur River 4 km from Shapur (29º49'N, 51º34'E); CMNFI 1979-0058, 6, 75.6-115.3 mm standard length, Fars, jube over Shapur River at Shapur (29º47'N, 51º35'E); CMNFI 1979-0059, 2, 45.0-50.4 mm standard length, ID, more than one species? Fars, Pulver River 8km south of Sivand (30º01'30"N, 52º57'E); CMNFI 1979-0061, , mm standard length, ID, more than one species? Fars, stream tributary to Pulvar River (30º04'N, 53º01'E); CMNFI 1979-0063, 2, 201.0-206.7 mm standard length, Fars, qanat under Sa'adi's Tomb, Shiraz (29º37'N, 52º35'E); CMNFI 1979-0067, , mm standard length, Fars, qanat at Zarqan (ca. 29º46'N, ca. 52º43'E); CMNFI 1979-0068, , mm standard length, Fars, qanat 12 km from Shiraz on Esfahan road (ca. 29º43'N, ca. 52º34'30"E); CMNFI 1979-0073, 5, 28.9-86.6 mm standard length, Fars, Mand River beyond Chehel Chashhmeh (ca. 29º42'30"N, ca. 52º01'30"E); CMNFI 1979-0074, , mm standard length, Fars, Mand River backwater (29º41'N, 52º06'E); CMNFI 1979-0075, , mm standard length, Fars, Mand River at Pol-e Kavar (29º11'N, 52º41'E); CMNFI 1979-0079, 2, 120.7-149.9 mm standard length, Fars, Mand River 5 km above Band-e Bahman (ca. 29º12'N, ca. 52º38'E); CMNFI 1979-0079, 1, 159.7 mm standard length, Fars, Mand River 5 km above Band-e Bahman (ca. 29º12'N, ca. 52º38'E); CMNFI 1979-0090, , mm standard length, Esfahan, Gav Khuni (ca. 32º21'N, ca. 52º49'E); CMNFI 1979-0093, 1, 73.9 mm standard length, (); BWC 76-77 check CMNFI # CMNFI 1979-0109, 1, 91.1 mm standard length, Fars, Mand River ar Shahr-e Khafr (28º56'N, 53º14'E); CMNFI 1979-0109, 1, 103.4 mm standard length, Fars, Mand River ar Shahr-e Khafr (28º56'N, 53º14'E); CMNFI 1979-0111, 10, 8.7-54.6 mm standard length, Fars, stream 21-22 km from Shiraz (29º37'30"N, 52º21'E); CMNFI 1979-0113, , mm standard length, Fars, qanat under Sa'adi's Tomb (29º37'N, 52º35'E); CMNFI 1979-0114, , mm standard length, Fars, Mand River at road bridge (29º41'N, 52º06'E); CMNFI 1979-0115, 4, 154.4-172.6 mm standard length, qanat under Sa'adi's Tomb (29º37'N, 52º35'E); CMNFI 1979-0125, 1, 137.8 mm standard length, Bushehr, Dalaki River near Dalaki (ca. 29º28'N, ca. 51º21'E); CMNFI 1979-0128, 16, 34.6-108.6 mm standard length, Fars, Shur River (28º51'N, 52º31'E); CMNFI 1979-0128, 18, 17.2-135.3 mm standard length, Fars, Shur River (28º51'N, 52º31'E); CMNFI 1979-0129, , mm standard length, Fars, spring 2 km north of Farrashband (28º54'N, 52º04'E); CMNFI 1979-0130, 5, 44.4-93.3 mm standard length, Fars, Shur River 4 km west of Firuzabad (28º51'N, 52º32'E); CMNFI 1979-0131, 58, 25.5-140.0 mm standard length, Fars, Mand River tributary (28º38'N, 52º49'E); CMNFI 1979-0132, 23, 51.1-74.4 mm standard length, Fars, Mand River tributary (28º35'N, 52º58'E); CMNFI 1979-0154B, 6, mm standard length, Fars, upper Shur River drainage near Darab (28º45'30"N, 52º24'E); CMNFI 1979-0155, 7, 36.2-80.5 mm standard length, Fars, spring at Gavanoo (28º47'N, 54º22'E); CMNFI 1979-0156, 3, 54.6-122.9 mm standard length, Fars, qanat at Rashidabad (28º47'N, 54º18'E); CMNFI 1979-0157, , mm standard length, Fars, qanat at Hadiabad (28º52'N, 54º13'E); CMNFI 1979-0158, , mm standard length, Fars, qanat over Qasook River (28º54'N, 53º53'30"E); CMNFI 1979-0159, 87, 23.1-167.3 mm standard length, Fars, qanat at Qaziabad (ca. 28º54'N, ca. 53º43'E); CMNFI 1979-0160, 4, 66.3-138.4 mm standard length, Fars, Arteshkadeh Pomp spring (29º09'N, 53º37'E); CMNFI 1979-0161, 29, 33.2-88.3 mm standard length, Fars, qanat on Neyriz to Shiraz road (29º10'30"N, 53º41'E); CMNFI 1979-0162, 9, ?-88.3 mm standard length, Fars, qanat behind Sarvestan (29º16'30"N, 53º14'E); CMNFI 1979-0163, 1, 73.8 mm standard length, ?Fars, neighbourhood of Shiraz (no other locality data); CMNFI 1979-0164, 1, 49.4 mm standard length, ?Fars, neighbourhood of Shiraz (no other locality data); CMNFI 1979-0165, 7, 30.0-96.6 mm standard length, Kerman, qanat at Ahmadabad (30º32'N, 55º38'E); CMNFI 1979-0166, 67, 37.1-123.1 mm standard length, Kerman, qanat at Hassanabad-e Nuq (30º43'N, 55º50'E); CMNFI 1979-0168, , mm standard length, Kerman, qanat at Shahabad (29º07'N, 58º16'E); CMNFI 1979-0169, , mm standard length, Kerman, qanat 10 km from Mahan (30º08'30"N, 57º17'E); CMNFI 1979-0170, , mm standard length, Kerman, qanat at Baghin (30º12'N, 56º48'E); CMNFI 1979-0171, , mm standard length, Kerman, qanat at Bardesir (29º56'N, 56º34'E); CMNFI 1979-0187, , mm standard length, Hormozgan, stream and pools at Sar Khun (27º23'30"N, 56º26'E); CMNFI 1979-0191, , mm standard length, Fars, stream 10 km east of Furg (ca. 28º16'N, ca. 55º18'E); CMNFI 1979-0192, , mm standard length, Fars, qanat 2 km east of Rostaq (28º26'30"N, 55º04'E); CMNFI 1979-0195, , mm standard length, Fars, jube on road to Fasa (ca. 28º54'N, ca. 53º53'30"E); CMNFI 1979-0198, , mm standard length, Fars, stream at Tadovan (28º47'N, 53º24'30"E); CMNFI 1979-0199, 6, 70.8-102.1 mm standard length, Fars, qanat 18 km from Jahrom (ca. 28º23-25'N, ca. 53º31-40'E); CMNFI 1979-0202, , mm standard length, Fars, Mand River (29º01'N, 53º00'E); CMNFI 1979-0203, , mm standard length, Fars, qanat at Dudej (29º33'N, 52º59'E); CMNFI 1979-0204, , mm standard length, Fars, qanat on road to Kharameh (29º33'N, 52º59'E); CMNFI 1979-0205, 12, 45.9-200.5 mm standard length, Fars, jube at Runiz-e Pa'in (29º12'N, 53º42'E); CMNFI 1979-0206, , mm standard length, Fars, qanat on road to Kharameh (29º12'N, 53º40'E); CMNFI 1979-0207, 12, 24.2-83.7 mm standard length, Fars, jube 22 km from Neyriz (29º16'N, 54º28'E); CMNFI 1979-0208, 6, 39.9-130.4 mm standard length, Fars, qanat 47 km from Neyriz (ca. 29º11'N, ca. 54º40'E); CMNFI 1979-0209, 60, 43.6-138.9 mm standard length, Kerman, qanat at Kuch Kuluh (29º25'N, 56º03'E); CMNFI 1979-0211, 63, 33.2-94.3 mm standard length, Kerman, river on road to Baft (29º19'N, 56º12'E); CMNFI 1979-0212, 73, 26.0-99.1 mm standard length, Kerman, qanat on road to Baft (29º14'N, 56º17'E); CMNFI 1979-0213, 5, 51.4-60.2 mm standard length, Kerman, stream in Kharan River drainage (29º15'N, 56º25'E); CMNFI 1979-0214, , mm standard length, Kerman, qanat pool on road to Baft (ca. 29º15'N, ca. 56º28'E); CMNFI 1979-0215, 15, 39.7-125.9 mm standard length, Kerman, Kharan River drainage (29º14'N, 56º37'E); CMNFI 1979-0216, 11?, 51.1-65.8 mm standard length, Kerman, qanat 9 km from Baft (ca. 29º13'N, ca. 56º42'E); CMNFI 1979-0217, 15, 39.7-125.9 mm standard length, Kerman, Kharan River drainage (ca. 28º59'30"N, ca. 56º51'30"E); CMNFI 1979-0221, , mm standard length, Kerman, Halil River drainage (28º51'N, 57º52'E); CMNFI 1979-0241, , mm standard length, Fars, Shapur River at Shapur (29º47'N, 51º35'E); CMNFI 1979-0243, , mm standard length, Esfahan, Zayandeh River at Falavarjan (32º33'N, 51º31'E); CMNFI 1979-0246, , mm standard length, Shahrestan-e Bakhtiari va Chahar Mahall, upper Karun River drainage (31º57'30"N, 50º59'E); CMNFI 1979-0251, , mm standard length, Esfahan, stream 1 km east of Daran (32º59'N, 50º26'E); CMNFI 1979-0251, , mm standard length, Esfahan, stream 1 km east of Daran (32º59'N, 50º26'E); CMNFI 1979-0255, , mm standard length, Markazi, Bar River drainage 2 km west of Shahabiyeh (33º51'30"N, 50º23'E); CMNFI 1979-0269, 1, 125.0 mm standard length, Lorestan, Dez or Karkheh River drainage (no other locality data); CMNFI 1979-0271, , mm standard length, Lorestan, Kashkan River drainage (33º39'N, 48º32'30"E); CMNFI 1979-0272, , mm standard length, Lorestan, river at Nokhor (33º40-47'N, 48º28-45'E); CMNFI 1979-0273, 7, 66.7-137.6 mm standard length, Lorestan, Kashkan River drainage (33º26'N, 48º19'E); CMNFI 1979-0274, 3, 28.9-141.8 mm standard length, Lorestan, Kashkan River drainage (33º27'N, 48º11'E); CMNFI 1979-0276, , mm standard length, Lorestan, Kashkan River drainage (ca. 33º19'N, ca. 47º53'30"E); CMNFI 1979-0277, 2, 116.2-133.4 mm standard length, Lorestan, Kashkan River drainage (33º30'N, 47º59'30"E); CMNFI 1979-0278, 3, 93.5-114.7 mm standard length, Lorestan, Kashkan River drainage (33º34'N, 48º01'E); CMNFI 1979-0279, 1, 126.0 mm standard length, Lorestan, Khorramabad River 16 km from Nurabad (33º37'N, 48º18'E); CMNFI 1979-0279, 5, 115.6-155.8 mm standard length, Lorestan, Khorramabad River 16 km from Nurabad (33º37'N, 48º18'E); CMNFI 1979-0280, 3, 104.7-107.7 mm standard length, Lorestan, Kashkan River drainage (33º43-47'N, 48º12-15'E); CMNFI 1979-0282, 6, 110.3-130.3 mm standard length, Lorestan, Seymareh River drainage at Nurabad (34º05'N, 47º58'E); CMNFI 1979-0283, 2, 113.7-125.0 mm standard length, Kermanshahan, Qareh Su drainage (34º21'N, 47º07'E); CMNFI 1979-0285, 3, 125.5-148.0 mm standard length, Kermanshahan, Qareh Su drainage (34º26'N, 46º37'E); CMNFI 1979-0286, , mm standard length, Kermanshahan, Ravansar River at Ravansar (34º43'N, 46º40'E); CMNFI 1979-0287, 2, 128.2-136.1 mm standard length, Kermanshahan, Chashmeh Javari 2 km from Ravansar (ca. 34º42'N, ca. 46º40'E); CMNFI 1979-0288, 62, 37.6-153.7 mm standard length, Ilam and Poshtkuh, Gangir River at Sarab Ewan (33º50'N, 46º18'E); CMNFI 1979-0289, , mm standard length, Kermanshahan, Diyala River drainage (34º28'N, 45º52'E); CMNFI 1979-0290, , mm standard length, Kermanshahan, Diyala River drainage at Qasr-e Shirin (34º31'N, 45º35'E); CMNFI 1979-0291, , mm standard length, Kermanshahan, Diyala River drainage (34º24'N, 45º37'E); CMNFI 1979-0306, , mm standard length, Kerman, qanat on road to Baft (29º13'N, 54º33'E); CMNFI 1979-0307, 5, 50.9-73.4 mm standard length, Kerman, river at Sartal 6 km from Baft (ca. 29º17'N, ca. 56º38'E); CMNFI 1979-0308, 67, 20.5-246.9 mm standard length, Kerman, river 44 km from Baft (29º02'N, 56º50'E); CMNFI 1979-0309, , mm standard length, Kerman, Fahraj River at Azizabad (28º57'N, 58º42'E); CMNFI 1979-0315, 2, 53.5-65.5 mm standard length, Baluchestan, Bampur River 2 km from Karevandar (27º51'N, 60º46'E); CMNFI 1979-0315, 34?, 53.7-85.1 mm standard length, (); note two collections? CMNFI 1979-0337, , mm standard length, Baluchestan, stream near Kanowak (ca. 28º40'N, ca. 60º48'E); CMNFI 1979-0341, 14, 27.2-75.9 mm standard length, Kerman, Tahrud west of Bam (29º23'N, 57º52'E); CMNFI 1979-0343, , mm standard length, Fars, lake near Deh Bid (ca. 30º32'N, ca. 52º49'E); CMNFI 1979-0411, 7, 42.2-76.5 mm standard length, Hormozgan, Minab River past Rudan (27º24'N, 57º12'E); CMNFI 1979-0419, 1, 62.2 mm standard length, Fars, stream 7 km from Rostaq (28º29'N, 55º01'E); CMNFI 1979-0420, 6, 57.1-150.6 mm standard length, Fars, Rudbar River at Bahregan (30º11'N, 52º03'E); CMNFI 1979-0422, , mm standard length, Boyer Ahmadi-ye Sardsir va Kohkiluyeh, stream in Yasuj valley (30º36'N, 51º34'E); CMNFI 1979-0424, , mm standard length, Fars, stream on Yasuj to Nurabad road (30º18'N, 51º30'30"E); CMNFI 1979-0425, , mm standard length, Fars, Haft Barm-e Kudian (29º49'N, 52º02'E); CMNFI 1979-0426, , mm standard length, Esfahan, qanat at Abbasabad-Natanz (33º36'N, 51º49'E); CMNFI 1979-0458, 2, 90.7-108.4 mm standard length, Markazi, Khar River 6 km north of Ab-Garm (35º47'N, 49º20'E); CMNFI 1979-0460, 3, 54.4-65.0 mm standard length, Hamadan, stream 16 km south of Asadabad (34º39'N, 48º05'E); CMNFI 1979-0462, , mm standard length, Markazi, Mazdaqan River (35º06'30"N, 49º40'30"E); CMNFI 1979-0466, , mm standard length, Esfahan, qanat at Meymeh (33º27'N, 51º10'E); CMNFI 1979-0484, , mm standard length, Khorasan, stream 22 km west from Bojnurd (37º28'N, 56º44'E); CMNFI 1979-0497, 3, 49.8-113.0 mm standard length, Fars, Mand River at Band-e Bahman (29º11'N, 52º40'E); CMNFI 1979-0497, 7, 102.2-132.0 mm standard length, Fars, Mand River at Band-e Bahman (29º11'N, 52º40'E); CMNFI 1979-0499, , mm standard length, Fars, ditch 32 km from Kor River bridge (30º04'30"N, 52º36'E); CMNFI 1979-0501, 6, 34.1-110.9 mm standard length, Fars, Mand River at Kavar (29º11'N, 52º41'E); CMNFI 1979-0502, , mm standard length, Fars, Haft Barm-e Kudian (29º49'N, 52º02'E); CMNFI 1993-0126, , mm standard length, Kermanshahan, Sarab-e Yavari (34º28'N, 46º56'E); CMNFI 2007-0030, , mm standard length, Baluchestan, stream near Eskelabad (28º35'N, 60º48'E); CMNFI 2007-0031, , mm standard length, Baluchestan, headwaters of Bampur River (27º51'N, 60º46'E); CMNFI 2007-0037, , mm standard length, Kerman, Hosseinabad and Gamatabad qanats at Bam (29º06'N, 58º21'E); CMNFI 2007-0038, , mm standard length, Kerman, Mehtiabad qanat (29º06'N, 58º21'E); CMNFI 2007-0039, , mm standard length, Kerman, Tahrud River (ca. 29º23'N, ca. 57º53'E); CMNFI 2007-0040, , mm standard length, Kerman, Qahariz qanat at Jupar (30º04'N, 57º08'E); CMNFI 2007-0041, , mm standard length, Kerman, qanat at Baghin (30º12'N, 56º48'E); CMNFI 2007-0042, , mm standard length, Kerman, qanat at Negar (29º52'N, 56º50'E); CMNFI 2007-0043, , mm standard length, Kerman, qanat at Emamzadeh Sultan (ca. 29º40'N, ca. 56º45'E); CMNFI 2007-0044, , mm standard length, Kerman, Qal'eh-ye Askar stream (ca. 29º28'N, ca. 56º38'E); CMNFI 2007-0045, , mm standard length, Kerman, Kharan River drainage at Baft (29º14'N, 56º38'E); CMNFI 2007-0047, , mm standard length, Kerman, qanat at Hoshun (29º14'N, 56º19'E); CMNFI 2007-0048, , mm standard length, Kerman, qanat at Hasanabad (ca. 28º50'N, ca. 55º50'E); CMNFI 2007-0049, , mm standard length, Hormozgan, upper Kol River basin at Hajjiabad (ca. 28º19'N, ca. 55º55'E); CMNFI 2007-0063, , mm standard length, Fars Mand River tributary outside Jahrom (28º36'N, 53º37'E); CMNFI 2007-0065, , mm standard length, Fars, Barm-e Dalak (ca. 29º35'N, ca. 52º38'E); CMNFI 2007-0066, , mm standard length, Fars, qanat under Sa'adi's Tomb, Shiraz (29º37'N, 52º35'E); CMNFI 2007-0067, , mm standard length, Fars, Sivan River (ca. 30º02'N, ca. 52º57'E); CMNFI 2007-0068, 5, 59.0-89.6 mm standard length, Fars, qanat 4 km south of Abarqu (ca. 31º07'N, ca. 53º14'E); CMNFI 2007-0069, , mm standard length, Yazd, qanat at Zarej (ca. 31º58'N, ca. 54º17'E); CMNFI 2007-0070, , mm standard length, Yazd, qanat at Ardakan, (32º19'N, 53º59'E); CMNFI 2007-0073, , mm standard length, Esfahan, Zayandeh River at Tanderan (32º47'N, 51º02'E); CMNFI 2007-0075, , mm standard length, Hamadan, Malayer River south of Malayer (ca. 34º17'N, ca. 48º47'E); CMNFI 2007-0076, , mm standard length, Markazi, Malekabad qanat (34º05'N, 49º53'E); CMNFI 2007-0083, , mm standard length, Azarbayjan-e Khavari, Qaranqu River basin west of Sar Eskand Khan (ca. 37º25'N, ca. 46º55'E); CMNFI 2007-0084, , mm standard length, Azarbayjan-e Khavari, Talkheh River basin west of Sarab (ca. 37º56'N, ca. 47º19'E); CMNFI 2007-0091, , mm standard length, Azarbayjan-e Khavari, Zilber Chay basin west of Marand (38º30'N, 45º23'E); CMNFI 2007-0100, , mm standard length, Azarbayjan-e Bakhtari, Kalwi Chay near Piranshahr (ca. 36º44'N, ca. 45º10'E); CMNFI 2007-0108, , mm standard length, Kordestan, Qeshlaq River basin north of Sanandaj (ca. 35º33'N, ca. 47º08'E); CMNFI 2007-0109, , mm standard length, Kordestan, Qeshlaq River basin south of Sanandaj (ca. 35º16'N, ca. 47º01'E); CMNFI 2007-0110, , mm standard length, Kordestan, Yuzidar River basin (ca. 35º05'N, ca. 46º56'E); CMNFI 2007-0115, , mm standard length, Kermanshahan, Qareh Su basin north of Kermanshah (ca. 34º34'N, ca. 46º47'E); CMNFI 2007-0116, , mm standard length, Kermanshahan, Gav Masiab River basin west of Sahneh (ca. 34º28'N, ca. 47º36'E); CMNFI 2007-0117, , mm standard length, Kermnashahan, Gav Masiab River basin near Sahneh (ca. 34º24'N, ca. 47º40'E); CMNFI 2007-0117, , mm standard length, Kermnashahan, Gav Masiab River basin near Sahneh (ca. 34º24'N, ca. 47º40'E); CMNFI 2007-0122, , mm standard length, Markazi, Khar River basin south of Takestan (ca. 35º56'N, ca. 49º30'E); USNM 200308, 2, 37.5-47.3 mm standard length, Lorestan, Ab-e Khorramabad (33º30'N, 48º13'E); ? damascina USNM 205933, 5, 97.5-142.4 mm standard length, Baluchestan, Karavandar Creek (no other locality data); ? damscina USNM acc. 303854, 14, 29.1-44.6 mm standard length, Fars, Lake Arzhan (29º36'N, 51º59'E). ? damscina USNM acc. 303854, 4, 55.7-99.0 mm standard length, Fars, pool east of Sangkar (29º48'N, 53º29'E); ?damascina

Capoeta aculeata
(Valenciennes, 1844)

 

Flank scale                                                            Left pharyngeal arch

Common names

shum (= unlucky or inauspicious, possible meaning), سياه ماهي (= siah mahi), زرده پر (= zardehpar), siah mahi aculeata.

Systematics

Chondrostoma aculeatum was originally described from "eaux douces de la Perse".

Scaphiodon macrolepis Heckel, 1849 described from the "Confluenten des Araxes bei Persepolis" (probably the Pulvar (= Sivan) River, Fars near Persepolis) and Varicorhinus bergi Derzhavin, 1929 described in Latin from "Keredsh flumen propea Teherane, Persia septentrionalis" (Karaj River near Tehran, northern Iran) are synonyms.

Six syntypes (MNHN 2357) of Chondrostoma aculeatum in poor condition are stored in the Muséum national d'Histoire naturelle, Paris (Bertin and Estève, 1948; Coad and Krupp, 1994). They measure 86-179 mm standard length (Coad and Krupp, 1994) or 105-210 mm total length (Bertin and Estève, 1948). The largest specimen is designated as the lectotype.

Two syntypes of Scaphiodon macrolepis are in the Naturhistorisches Museum Wien under NMW 55896 and measure 100-175 mm standard length (Kähsbauer, 1964). Two other fish are marked as syntypes under NMW 51653 and are from Persepolis collected by Th. Kotschy. The catalogue in Vienna lists 4 fish and the 1997 card index agrees these 4 fish are the syntypes.

Types of Varicorhinus bergi are unknown (Eschmeyer et al. (1996)).

Berg (1949) considers both aculeata and macrolepis as distinct species although very close, the latter distinguished from the former by a deeper body and a shorter head. Karaman (1969) and Bianco and Banarescu (1982) place both aculeata and macrolepis in Capoeta capoeta; Karaman does suggest that macrolepis could belong in aculeata. Saadati (1977) considers aculeatus not more than subspecifically distinct from macrolepis, not realising the former has priority.

Key characters

This species differs from all others in the genus Capoeta in the lower number of lateral line (93% of 314 fish had range of 39-48) and caudal peduncle scales (90% of 303 fish had a range of 16-20). Capoeta capoeta, a related species, usually has 54 or more lateral line scales and 20 or more caudal peduncle scales.

Morphology

Dorsal fin with 3-5, modally 4, unbranched and 7-9, modally 8, branched rays. The last dorsal fin unbranched ray is thickened and serrated, the denticles being long and narrowly spaced but not strongly developed. Distally this spiny ray is flexible. Smaller fish have proportionately larger and more extensive denticles than larger fish. The extent of denticles from the base distally varies between about two-thirds and three-quarters. Anal fin with 3 unbranched and 5-6, modally 5, branched rays, pectoral fin with 14-21 branched rays, and pelvic fin with 7-10 branched rays.

Lateral line scales 36-52. Caudal peduncle scales 13-23. Scale shape is squarish with shallowly rounded to straight dorsal and ventral margins, sharp corners anteriorly, and a large to moderate central protuberance on the anterior margin. Radii are most numerous on the posterior field but even there are few, relatively few laterally and few anteriorly. Circuli are very fine but break into coarser "bubbles" on the posterior field. The focus is subcentral anterior. The pelvic fin axillary scale varies greatly in size.

The mouth is slightly arched or even straight in ventral view. The horny edge to the lower jaw is usually well-developed but may be lost in preserved specimens. Gill rakers number 16-25 and are short, reaching past the first or second raker when appressed. Rakers are thick and usually hooked at their tips. Pharyngeal teeth are modally 2,3,4-4,3,2 (in 10 fish). Major row teeth are spatulate with a wide crown in large fish. Total vertebrae number 39-44. The gut is extremely elongate with numerous anterior and posterior coils.

Meristic characters in Iranian fish are: dorsal fin branched rays 7(50), 8(255) or 9(4); anal fin branched rays 5(177) or 6(1); pectoral fin branched rays 14(2), 15(3), 16(12), 17(52), 18(123), 19(79), 20(27) or 21(6); pelvic fin branched rays 7(23), 8(183), 9(102) or 10(6); lateral line scales 36(1), 37(5), 38(8), 39(15), 40(25), 41(48), 42(56), 43(38), 44(40), 45(23), 46(22), 47(15), 48(10), 49(1), 50(4), 51(2) or 52(1); scales around the caudal peduncle 13(1), 15(5), 16(48), 17(52), 18(73), 19(64), 20(37), 21(12), 22(5) or 23(6); total gill rakers 16(3), 17(13), 18(40), 19(49), 20(62), 21(43), 22(43), 23(27), 24(16) or 25(6); and total vertebrae 39(1), 40(26), 41(90), 42(103) or 44(16).

Sexual dimorphism

Males have moderately large tubercles on the anal fin rays following the ray branching (2-4 tubercles on last 4 branched anal rays), small tubercles on the lowest caudal fin ray, very fine tubercles on top of the head, larger tubercles on the side of the head, largest on the snout below the eye and nostril as far as the mouth, connecting across the snout, and numbering 1-5 moderately large tubercles on flank scales variously arranged on each scale and best developed on the posterior part of the body.

Colour

The back is almost entirely black to green-brown or olive-green, the upper flank is brownish, the belly and lower flank are yellow up to the lateral line, only the belly centre being white. The flanks are generally silvery in live fish. Some fish have small black spots on the sides and fins. Preserved fish have pigment on the posterior, exposed margin and so are outlined on the flank. The sides of the head are golden-brown. Flank spots may be in 5 longitudinal rows above, and 2 rows below, the lateral line. Some populations have fish with spots and mottles on the body and fins but these are probably occasioned by a parasitic infestation. Fins are often reddish-brown to pink although pelvic and anal fins may be yellowish-green and the dorsal and caudal fins very light to hyaline. Preserved fish have pigment on the rays and membranes of fins without any distinctive pattern. The dorsal and caudal fins are darker than the lower fins. The iris is golden to orange. The peritoneum is black.

Size

Reaches 23.4 cm standard length.

Distribution

This species is found in the Tigris River, Namak Lake, Dasht-e Kavir, Kerman-Na'in, Esfahan, Kor River basins (Rainboth, 1981; Bianco and Banarescu, 1982; Ghorbani Chafi, 2000). Abdoli (2000) maps this species from theKerman-Na'in basin generally; the upper Kal Shur, Jajarm and Jovein rivers in the Dasht-e Kavir basin; the middle and upper Shur and Abhar, Qareh Chai and Qom rivers in the Namak Lake basin; the Zayandeh and Shur rivers in the Esfahan basin; the Jarrahi and Marun, upper Karun and Khersan, Dez, Karkheh, Simarreh and Kashkan rivers in the Tigris River basin.

Zoogeography

Saadati (1977) suggests that this species moved eastward to basins on the plateau during more pluvial periods from the Tigris River basin. See also above under genus.

Habitat

Unknown in detail.

Age and growth

Unknown.

Food

Gut contents include filamentous algae, plant fragments and diatoms with large amounts of sand. This species has been seen turning belly up to feed (field notes for specimens from Jajarm, Khorasan).

Reproduction

Reproduction has not been studied in this species. Specimens from the Khorramabad River contained eggs 1.5 mm in diameter on 6 July and some seemed to be reabsorbing eggs. Spawning presumably takes place in late spring and summer.

Parasites and predators

Barzegar et al. (2004) examined this species for parasites in fish from the Beheshtabad river in Chahar Mahall va Bakhtiari Province and found Dactylogyrus lenkorani, Gyrodactylus sp. and Myxobolus sp. Masoumian et al. (2007) record the myxosporean parasite Myxobolus cristatus from this species in the Zayandeh River. Mehdipoor et al. (2004) record the monogeneans Dactylogyrus chramuli, D. lenkorani and D. gracilis in the Zayandeh River. Barzegar and Jalali (2006) report parasites in this species from Kaftar Lake as Lernaea cyprinacea and Trichodina sp.

Economic importance

None.

Conservation

This species is widely distributed in Iran and does not appear to be in need of conservation but its biology and habitat requirements are unknown. Kamali-Far et al.(2009) have used carp pituitary extract in an attempt to induce spawning in this species. Hatchery production could then be used to supplement natural stocks. However, the attempt was unsuccessful. Note that the identity of the species used in this study needs verification judging from the photograph in the paper.

Further work

The biology of this species needs study as does its habitat requirements and conservation needs.

Sources

Type material: See above, Chondrostoma aculeatum (MNHN 2357).

Iranian material: CMNFI 1979-0025, 2, 65.3-68.1 mm standard length, Fars, Kor River near Marv Dasht (29º51'N, 52º46'30"E); CMNFI 1979-0059, 155, 22.9-67.4 mm standard length, Fars, Pulvar River 8 km south of Sivand (30º01'30"N, 52º57'E); CMNFI 1979-0061, 6, 28.6-64.9 mm standard length, Fars, stream tributary to Pulvar River (30º04'N, 53º01'E); CMNFI 1979-0069, 1, 28.7 mm standard length, Fars, qanat at Naqsh-e Rostam (29º59'30"N, 52º54'E); CMNFI 1979-0070, 16, 25.9-60.0 mm standard length, Fars, Pulvar River near Naqsh-e Rostam (29º59'N, 52º54'E); CMNFI 1979-0090, 2, 153.6-160.5 mm standard length, Esfahan, Gav Khuni (ca. 32º21'N, ca. 52º49'E); CMNFI 1979-0116, 49, 24.3-52.1 mm standard length, Fars, Kor River near Marv Dasht (29º51'N, 52º46'30"E); CMNFI 1979-0117, 14, 34.4-44.1 mm standard length, Fars, Pulvar River at Naqsh-e Rostam (29º59'N, 52º54'E); CMNFI 1979-0252, 3, ?, mm standard length, Markazi, jube at Baqerabad (34º55'N, 50º50'E); CMNFI 1979-0253, 5, 40.4-103.7 mm standard length, Markazi, stream in Qareh Chay drainage (34º52'N, 50º49'E); CMNFI 1979-0270, 1, 121.8 mm standard length, Lorestan, Kashkan River draiangae outside Khorramabad (33º26'N, 48º19'E); CMNFI 1979-0271, 1, 52.1 mm standard length, Lorestan, stream in Kashkan River drainage (33º39'N, 48º32'30"E); CMNFI 1979-0273, 28, 51.4-104.5 mm standard length, Lorestan, stream in Kashkan River drainage near Khorramabad (33º26'N, 48º19'E); CMNFI 1979-0274, 6, 20.6-59.2 mm standard length, Lorestan, stream in Kashkan River drainage (33º27'N, 48º11'E); CMNFI 1979-0275, 1, 50.9 mm standard length, Lorestan, Kashkan River 2 km from Ma'mulan (33º25'N, 47º58'E); CMNFI 1979-0279, 18, 41.1-129.9 mm standard length, Lorestan, Khorramabad River (33º37'N, 48º18'E); CMNFI 1979-0282, 7, 99.2-130.8 mm standard length, Lorestan, river at Nurabad (34º05'N, 47º58'E); CMNFI 1979-0283, 2, 125.2-186.3 mm standard length, Kermanshahan, Qareh Su near Kermanshah (34º21'N, 47º07'E); CMNFI 1979-0343, 1, 146.6 mm standard length, Fars, lake near Deh Bid (ca. 30º32'N, ca. 52º49'E); check ID? CMNFI 1979-0365, 1, 25.0 mm standard length, Khuzestan, stream in Doveyrich River drainage (32º25'N, 47º36'30"E); CMNFI 1979-0396, 9, 32.5-58.7 mm standard length, Khuzestan, Kheyrabad River 20 km from Behbehan (30º32'N, 50º23'30"E); CMNFI 1979-0427, 2, 100.5-112.2 mm standard length, Markazi, Cheshmeh Fin at Fin (33º57'N, 51º24'E); checkID? CMNFI 1979-0428, 17, 25.9-104.5 mm standard length, Markazi, stream 3 km south of Sen Sen (34º13'N, 51º16'E); checkID? CMNFI 1979-0458, 9, 48.5-117.8 mm standard length, Markazi, Khar River 6 km north of Ab-garm (35º47'N, 49º20'E); CMNFI 1979-0460, 1, 77.6 mm standard length, Hamadan, stream 16 km south of Asadabad (34º38'N, 48º03'E); checkID? CMNFI 1979-0463, 8, 97.9-135.3 mm standard length, Markazi, Qareh Chay (34º53'N, 50º24'E); checkID? CMNFI 1979-0464, 1, 74.2 mm standard length, Markazi, qanat at Kheyrabad (34º08'N, 50º00'E); CMNFI 1979-0465, 18, 35.7-58.3 mm standard length, Markazi, Qom River (34º18'30"N, 50º32'E); CMNFI 1979-0500, 2, 92.4-98.6 mm standard length, Fars, Pulvar River near Naqsh-e Rostam (29º59'N, 52º54'E); checkID? CMNFI 1980-0156, 27, ? mm standard length, Markazi, Karaj River (35º47'N, 50º58'E); CMNFI 1993-0154, 1, mm standard length, Markazi, Sharra River near Far (34º03'N, 49º20'E); checkID? CMNFI 1993-0156, 1, mm standard length, Markazi, Sharra River (34º03'N, 49º21'E); checkID? CMNFI 2007-0006, 9, 59.9-127.2 mm standard length, Khorasan, spring in Qareh Su basin south of Garmeh (ca. 36º58'N, ca. 56º15'E); CMNFI 2007-0007, 8, 59.4-79.3 mm standard length, Khorasan, stream supplemented by qanats, Kal-e Tangeh (ca. 36º59'N, ca. 56º29'E); CMNFI 2007-0008, 2, 72.1-84.3 mm standard length, Khorasan, qanat at Jajarm (36º57'N, 56º23'E); CMNFI 2007-0009, 18, 35.9-108.1 mm standard length, Khorasan, qanat at Amirabad (ca. 36º31'N, ca. 56º45'E); CMNFI 2007-0010, 11, 80.8-123.1 mm standard length, Khorasan, qanat at Haresabad (36º07'N, 57º37'E); CMNFI 2007-0011, 12, 34.1-85.4 mm standard length, Khorasan, Kalshur River south of Neyshabur (36º05'N, 58º43'E); CMNFI 2007-0071, 10, 70.4-156.9 mm standard length, Esfahan, qanat at Mohammadiyeh, Na'in (32º51'N, 53º06'E); CMNFI 2007-0074, 29, 50.6-100.7 mm standard length, Markazi, Qareh Chai west of Arak (34º03'N, 49º21'E); CMNFI 2007-0075, 16, 29.3-152.5 mm standard length, Hamadan, Hamadan, Malayer River 5 km from Malayer (ca. 34º17'N, ca. 48º47'E); CMNFI 2007-0076, 5, 56.1-97.4 mm standard length, Markazi, Malekabad qanat east of Arak (34º05'N, 49º53'E); CMNFI 2007-0078, 8, 37.6-102.8 mm standard length, Markazi, Qom River (ca. 34º18'N, ca. 50º32'E); CMNFI 2007-0117, ?, mm standard length, Kermanshahan, Gav Masiab River basin near Sahneh (ca. 34º24'N, ca. 47º40'E); CMNFI 2007-0119, ?, mm standard length, Kermanshahan, Gav Masiab River basin near Kangavar (ca. 34º31'N, ca. 48º03'E); CMNFI 2007-0120, 15, 29.0-165.5 mm standard length, Hamadan, Ab Chay near Hamadan (ca. 34º49'N, ca. 48º29'E); CMNFI 2007-0122, 12, 35.0-77.6 mm standard length, Markazi, Khar River basin south of Takestan (ca. 35º56'N, ca. 49º30'E); BM(NH) 1934.10.29:2, 1, 84.0 mm standard length, Markazi, Tehran (no other locality data); BM(NH) 1958.11.7:1-6, 6, 25.6-89.9, Khorasan, Jajarm (36º57'N, 56º23'E); BM(NH) 1975.1.17:255-258, 4, 103.0-161.0 mm standard length, Esfahan, Esfahan (no other locality data); MNHN 1960-611, 2, 127.0-144.0 mm standard length, Markazi, Jajrud east of Tehran (ca. 35º45'N, ca. 51º42'E) USNM 205932, 3, 78.5-159.4 mm standard length, Markazi, stream southwest of Tehran (35º34'N, 51º03'E); ZMH 5905, 2, 57.0-70.0 mm standard length, ?, Jafar Abad qanat (?); ZSM 25703, 1, 76.3 mm standard length, ?, Khorramabad River (no other locality data).

Capoeta barroisi
Lortet, 1894

Common names

siah mahi-ye Dasht-e Arzhani (Arzhan Plain black fish).

[spotted barb, Tigris barb].

Systematics

Subspecies are Capoeta barroisi persica Karaman, 1969 described from "See Zariwar, Mariwan, 120 km westlich v. Sannadaj" (Lake Zaribar near Marivan, Kordestan in the Tigris River basin) and Capoeta barroisi mandica Bianco and Banarescu, 1982 from the "Mand River near Dasht-e-Arzhan" of Fars Province. Krupp (1985c) considers both these to be synonyms of the nominal subspecies, C. b. barroisi.

The subspecies persica is distinguished from the type subspecies by having a more horseshoe-shaped mouth, 8 branched dorsal fin rays, 18 gill rakers, blackish pectoral, pelvic and anal fins, few but very large black spots on the body, a shorter anal fin and a longer pectoral fin, and a deep body, based on a single specimen. Krupp (1985c) considers the characters of mouth form and colour to fall within the range of the nominal subspecies (and by implication the other characters too). Özuluğ and Freyhof (2008) found it difficult to reach a conclusion on the taxonomic status of this subspecies on the basis of a single specimen which could be abnormal.

C. barroisi mandica differs from the type subspecies (C. barroisi barroisi) and C. b. persica in number of scales (61-68 in mandica (58-68 in types examined by me), 69-82 in barroisi, 78-79 in persica), number of gill rakers (21-24 in mandica (22-27 in types examined by me and apparently number is related to size of fish), 27-31 in barroisi, 18 in persica), from barroisi in having usually 8 branched dorsal fin rays (barroisi has 9 but persica also has 8), and from persica by a straight mouth (also straight or transverse in barroisi, arched in persica). Krupp (1985c) considers the scale counts to be within the lower range of the nominal subspecies, gill raker counts and mouth position do not differ from the nominal subspecies, and the dorsal fin ray count of 8 is seen in the subspecies mandica. Krupp observes that meristic and morphometric characters are extremely variable in widely distributed Capoeta species. Özuluğ and Freyhof (2008) examined 5 juvenile specimens from the Mand River and consider the subspecies mandica to be a valid species. Widespread taxa like Capoeta species are prime candidates for  molecular analyses which might help resolve conflicting views on - a single widespread, variable species versus several distinct species.

Berg (1949) considers this species to be close to C. damascina, differing by having a stronger spine in the dorsal fin, hardly an invariant character. Saadati (1977) considers that C. barroisi of Karaman is in fact C. damascina. Turan (2008) using mt 16S rDNA concluded on this evidence that C. barroisi was a subspecies of C. damascina although other genetic markers should be used for a more reliable assessment.

Syntypes of Capoeta barroisi are in the Musée Guimet d'Histoire Naturelle, Lyon (MGHN 3492, 316 mm standard length, from the Orontes near Antakya in Turkey collected by E. Chantre and MGHN 3493, 278 mm standard length, from Buhairat Hims in Syria collected by Th. Barrois) (Krupp, 1985c).

The holotype of C. barroisi persica is in the Zoologischen Instituts und Zoologischen Museums der Universität Hamburg (ZMH H4119, 185.2 mm standard length, Daryacheh-ye Zaribar, 35°32'N, 46°08'E, IV. 1968, W. Nümann (Bianco and Banarescu, 1982; Krupp, 1985c; examined and measured by me).

The holotype of C. b. mandica, 106.9 mm standard length, is in the Istituto di Zoologia dell'Universitá di L'Aquila, Italy (IZA 7890), with 95 paratypes from the same locality in IZA 7891 (now numbering 84 fish measuring 34.2-84.9 mm standard length) and 5 paratypes in the Institutul de Stiinte Biologice, Bucuresti, Romania (ISBB 3123), these 100 specimens having a standard length of 34-86 mm. Six paratypes of mandica are in the Canadian Museum of Nature, Ottawa under CMNFI 1982-0366 (from IZA 7891).?lengths

Key characters

The dorsal fin branched ray count of usually 9 rays is characteristic for the type subspecies but not nominal Iranian ones. Gill raker counts, a head length greater than the dorsal fin spine length and the irregular brownish markings on a silvery-white body are also distinctive.

Morphology

Dorsal fin with 3-4 unbranched and 8-10 branched rays, usually 9 in the type subspecies but 8 in the subspecies mandica, anal fin with 3 unbranched and 5 branched rays. Pectoral fin branched rays 15-16, pelvic fin rays 7-8. Lateral line scales 61-82, 58-68 in the subspecies mandica. Scales on the belly in front of the pelvic fins are small and may not be imbricate. Pharyngeal teeth 2,3,4-4,3,2. Gill rakers 18-31, reaching the second adjacent raker when appressed. The last dorsal fin unbranched ray is very strong, but narrows distally, and bears large denticles or serrations on three-quarters of its length. The snout in the holotype of the mandica subspecies has a depression in front of the nostrils.

ZMH6086 D8, A5 P15, V7, ll 58, gr 24, flanks speckled dorsally, mouth a gentle arch almost straight, large denticles in damaged D spine.

Sexual dimorphism

Tubercles in males are found from eye to eye around the snout with fine tubercles sparse on the top of the head. Most flank scales have a single, centrally-placed tubercle as do scales on the caudal peduncle. There is a single row of tubercles on the last three anal fin rays. There are some weak tubercles on the side of the head.

Colour

There are numerous, small, distinctive brown to black spots on the head, flank and dorsal and anal fins. The back and upper flank are silvery-white with slate to violet overtones and the belly is yellowish. The holotype of the subspecies mandica has light specking and mottling on the upper flank and back. All fins have some speckling on the rays and membranes but no clear rows of spots. Smaller fish (paratypes) have darker and bigger speckles than the holotype which extend lower on the flank.

Size

Reaches 31.6 cm standard length.

Distribution

The subspecies C. barroisi barroisi is found in the Ceyhan, Orontes, Quwayq and Tigris-Euphrates basins, C. b. persica only in Lake Marivan or Zaribar of the Tigris River basin of Iran, and C. b. mandica in the Mand River of the Gulf basin. Krupp (1985c) includes Iranian Gulf drainages for the type subspecies. Abdoli (2000) has Jarrahi and lower Karun in the Tigris River basin and the middle and lower Hilleh, lower Mand, and lower Dasht-e Palang rivers in the Gulf basin.

Zoogeography

Taxa in Iran are remote from the type locality of this species and may be indicative of the links between the Levantine fauna and the former tributaries of the Tigris-Euphrates basin in southern Iran. See also above under genus.

Habitat

Unknown.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

None reported from Iran.

Economic importance

None.

Conservation

Nothing is known of biology and a conservation assessment cannot be made. Endangered in Turkey (Fricke et al., 2007).

Further work

The biology of this species needs study as a prerequisite for a conservation assessment. The relationships of nominal Iranian taxa to material from the type locality need further study using molecular techniques.

Sources

Type material: Capoeta barroisi mandica (IZA 7890, 7891, CMNFI 1982-0366 (from IZA 7891)) and C. barroisi persica (ZMH H4119).

Iranian material: ZMH 6086, 1, 73.6 mm standard length, Fars, Shur Fluβ, zufluβ von Mand-Fluβ (= C. b. mandica) P. Bianco ?check this against his paper.

Comparative material: BMNH 1974.2.22:1853-1856, 3, 48.9-60.2 mm standard length, Iraq, Kaliasan near Sulaymaniyah

Capoeta buhsei
Kessler, 1877

Common names

shamshiri (= sword-like), mahi sibili (= moustached fish, from Karaj Lake).

Systematics

Varicorhinus nikolskii Derzhavin, 1929 described in Latin from the "Keredsh flumen" (= Karaj River near Tehran) is a synonym. Saadati (1977) places Capoeta buhsei in Capoeta damascina.

The 2 syntypes of Capoeta Buhsei, 200.7-211.4 mm standard length, are in the Zoological Institute, St. Petersburg (ZISP 2330) and were collected "iz Persii" (= from Persia) by Dr. Buhse in 1849. The 11 syntypes of Varicorhinus nikolskii have not been located (Eschmeyer et al., 1996).

Key characters

This species is distinguished by its low total gill raker count of 9-17, mean 12.4, modes 12 and 13, lower arch rakers 7-10 (cf. C. damascina which has 17-25 total rakers, mean based on ? specimens), the absence of a keel in front of the dorsal fin, the mouth structure, and by a very weak, unserrated or barely serrated dorsal fin spine in large fish (cf. C. damascina).

Morphology

Dorsal fin unbranched rays 3-4, branched rays 7-9, anal fin unbranched rays 3, branched rays 5., pectoral fin branched rays 14-19, and pelvic fin branched rays 7-9. Lateral line scales 72-99. Scales are found on the back and on the belly. A pelvic axillary scale is present. Scales have parallel dorsal and ventral margins, a rounded posterior margin and an anterior margin with a rounded central protuberance. Radii are found on all fields including a few long and curved ones on the lateral fields. The focus is subcentral anterior and circuli are numerous and fine. Gill rakers 9-19 in literature (but see below), including some counts probably for the lower arm only, and reach the second raker below when appressed but only the next raker in small fish. Pharyngeal teeth in the main row are spatulate, the crowns flat, narrow and curved. Tooth counts are 2,3,4 or 5-5 or 4,3,2. The fifth tooth in either row is small and variably present. This may be size related although the fish examined here were all relatively small and showed no clear trend. The gut is elongate with several long coils.

The mouth is large and a shallow horseshoe-shape with the horny lower jaw layer weakly developed but the lower lip corners in particular fleshy and well-developed. The last unbranched dorsal fin ray is weak with the distal half thin and flexible. Denticles are found on the basal third to two-thirds or more of the ray, their extent and size variable but usually small, weak and less extensive in large fish, while larger and more extensive in the smallest fish. For fish 48.9-174.0 mm standard length extent of dorsal fin spine serrations in spine length is 0.3-0.8, mean 0.6. The lower lip is apparent and finely ridged. The upper lip and snout are covered with unculi which occur also over the head but more widely spaced out. The upper lip unculi are densely concentrated and are broader than other head ones. Unculi are also on the lower head surface and belly scales back to the pelvic fins, and on the anal, pectoral and pelvic fin rays and membranes.

Meristics are as follows: dorsal fin branched rays 8(35) or 9(3); anal fin branched rays 5(38); pectoral fin branched rays 14(1), 17(20), 18(11) or 19(6); pelvic fin branched rays 7(1), 8(5), 9(31); lateral line scales 72(2), 73(1), 75(3), 76(4), 77(2), 78(1), 79(7), 80(4), 81(5), 82(2), 83(2), 84(2), 86(2) or 91(1); total gill rakers 9(2), 10(2), 11(4), 12(12), 13(11), 14(4), 15(2) or 17(1); pharyngeal teeth 2,3,4,-4,3,2(7), 2,3,4-5,3,2(6), 2,3,5-4,3,2(5), 2,3,5-5,3,2(1), 1,3,4-4,3,2 (1) or 2,3,4-4,3,1(1); and total vertebrae 43(1) and 44(6) (USNM 20593 and the syntypes).

Sexual dimorphism

One male specimen measuring 94.6 mm standard length bears large tubercles on anal fin rays, fine tubercles scattered on the head, on the back and upper flanks one tubercle per scale at the scale centre but not on every scale, all along the lateral line at one tubercle per scale, and below the lateral line only in the area above the anal fin.

Colour

Overall colour is brownish in preservative without spots or any distinctive markings. The back is dark. The peritoneum is dark brown to black in preserved fish.

Size

Reaches 25.7 cm.

Distribution

This species is endemic to the Namak Lake basin of Iran (Derzhavin, 1929; Wossughi, 1978; Abdoli, 2000). Abdoli (2000) questionably maps it from the Esfahan basin. A report from Lake Zaribar, Kordestan (Abzeeyan, 5(5):III, 1994) is presumably a mis-identification and records from springs of Kul River basin near Darab in the Hormuz basin (Bianco and Banarescu (1982) and the Hamun-e Jaz Murian basin (Vossoughi, 1998) are also questionable.

Zoogeography

An endemic of an interior Iranian basin, its zoogeographical relationships to other Capoeta have not been resolved. See also above under genus.

Habitat

Unknown.

Age and growth

Unknown.

Food

Gut contents include aquatic insect larvae and masses of filamentous algae, suggesting that aufwuchs is an important diet item.

Reproduction

Generally unknown but fish caught on 5 June measuring 121.3-132.6 mm standard length have small eggs, perhaps because this size of fish is not mature. A 174.0 mm standard length caught in January has larger eggs than those from the June fish. A male fish caught on 5 May and measuring 146.6 mm standard length has mature testes.

Parasites and predators

Williams et al. (1980) report the helminths Khawia armeniaca (a cestode) and Acanthocephalorhynchoides cholodkowskyi (an acanthocephalan) from this species in the Zayandeh River at Esfahan.

Economic importance

Unknown.

Conservation

The conservation status of this species has not been determined by field studies and assessments can only be done from museum collections.

Further work

The distribution and numbers of this species in the Namak Lake basin should be examined by field studies to determine the population status. This basin is mostly in Markazi (= Central) Province which contains Tehran and a very large human population with great demands on limited water resources. It is probably not under any immediate threat but is an Iranian endemic.

Sources

Type material: See above, Capoeta buhsei (ZISP 2330).

Iranian material: CMNFI 1970-0588, 19, 42.4-128.9 mm standard length, Markazi, Karaj Lake (35º57'N, 51º06'E); CMNFI 1979-0094, 2, 143.1-174.0 mm standard length, Markazi, Karaj Lake (35º57'N, 51º06'E); CMNFI 1979-0266, 2, 52.4-54.3 mm standard length, Esfahan, spring at Nowqan (ca. 33º10'N, ca. 50º05'E); CMNFI 1979-0458, 1, 94.2 mm standard length, Markazi, Khar River (35º47'N, 49º20'E); CMNFI 1979-0459, 2 ?check fish, only 1 in catalogue, 27.0-31.6 mm standard length, Hamadan, stream 2 km south of Razan (35º22'N, 49º02'E); CMNFI 1979-0461, 1, 54.1 mm standard length, Hamadan, qanat at Taveh (35º07'N, 49º02'E); CMNFI 1979-0495, 1, 42.5 mm standard length, Markzai, Nam River west of Firuzkuh (35º43'N, 52º40'E); CMNFI 1980-0154, 71, 12.0-34.9 mm standard length, Markazi, Karaj River below village (35º47'N, 50º58'E); CMNFI 1980-0156, 27, 32.4-54.3 mm standard length, Markazi, Karaj River below village (35º47'N, 50º58'E); CMNFI 1993-0151, 1, 146.4 mm standard length, Markazi, Sharra River near Far (34º03'N, 49º19'E); CMNFI 1993-0152, 2, 121.3-132.6 mm standard length, Markazi, Sharra River near Khosbijan (34º07'N, 49º23'E); CMNFI 1993-0153, 2, 104.3-138.9 mm standard length, Markazi, Sharra River near Emarat (33º52'N, 49º36'E); CMNFI 1993-0154, 1, 124.0 mm standard length, Markazi, Sharra River near Far (34º03'N, 49º20'E); CMNFI 2007-0074, 3, ? mm standard length, Markazi, Qareh Chay (34º03'N, 49º21'E); CMNFI 2007-0078, 5, ? mm standard length, Markazi, Qom River (ca. 34º18'N, ca. 50º32'E): check ID? CMNFI 2007-0079, 14, ? mm standard length, Zanjan, Abhar River basin (ca. 36º16'N, ca. 49º08'E); CMNFI 2007-0120, , mm standard length, Hamadan, Ab Chay (ca. 34º49'N, ca. 48º29'E); CMNFI 2007-0121, 3, 82.5-141.5 mm standard length, Hamadan, Qareh Su basin north of Razan (ca. 35º25'N, ca. 49º02'E); CMNFI 2007-0122, , mm standard length, Markazi, Khar River basin south of Takestan (ca. 35º56'N, ca. 49º30'E); USNM 20593, ?, ? mm standard length, (); ZMH 2632, 1, 148.2 mm standard length, Dojodje (); ZMH 2633, ?, ? mm standard length, above Latian ().

Capoeta capoeta
(Güldenstaedt, 1773)

Common names

tilkhos, سياه ماهي (= siah mahi, meaning black fish), sang lisak (= rock snail?); soru (= slippery) in the Dalaki and Shapur river basins.

[gara balig or Lankaran xramulyasi for C. c. gracilis, Kur xramulyasi for C. capoeta, both in Azerbaijan; khramulya, capoeta, kapuit, kaput (all apparently derived from local names in Georgia and Armenia, namely khramuli and kapweti); Lenkoranskaya khramulya or Lenkoran khramulya, Kurinskaya khramulya or Kura khramulya, Zakaspiiskaya khramulya or Transcaspian khramulya (also marinka is used locally for the Transcaspian khramulya subspecies but this is an error), Araksinskaya khramulya or Araks khramulya, all in Russian; Transcaucasian barb; khramulia; kersin handscherli at Aleppo, in Arabic].

Systematics

Cyprinus capoeta was originally described from Tbilisi, Georgia.

Cyprinus fundulus Güldenstaedt, 1787 from the Caspian Sea, Cyrus River (and Capoeta fundulus Valenciennes, 1842), Scaphiodon asmussii Keyserling, 1861 from "Warme Quelle bei Sultan Karaul, 8 Meilen nordöstlich von Herat" (now in Afghanistan, formerly in Persia), Scaphiodon gracilis Keyserling, 1861 from "Wasserleitung bei Gaes, einige Meilen von Isphahan", Scaphiodon heratensis Keyserling, 1861 described from the "Heri-Rud, ein Fluss bei Herat" (now in Afghanistan, formerly in Persia), Capoeta Hohenackeri Kessler, 1877 from Caucasia (probably lower Kura and Araks rivers, Azerbaijan), Capoeta (Scaphiodon) Steindachneri Kessler, 1872 and Capoeta Steindachneri var. platylepida Kessler, 1872 both from the Zeravshan River, Uzbekistan, and probably Capoeta gibbosa Nikol'skii, 1897 described in Latin as from "Bochsani in Persia orientale" are synonyms. Capöeta Guldenstädtii De Filippi in Tortonese, 1940 from "F. Arasse, Erzerum (Anatolia)" is Capoeta capoeta but it is a manuscript name and is not available (Tortonese, 1940; Eschmeyer et al., 1966); 2 syntypes are in the Istituto e Museo di Zoologia della R. Università di Torino (MZUT N.729).

Eschmeyer et al. (1996) have the date of Cyprinus capoeta as 1772; the type locality is Tiflis, Caspian Sea: they also have Cyprinus fundulus authored by Pallas, 1814 although Berg (1948-1949) has Güldenstädt as the author.

A hybrid of Capoeta capoeta heratensis and Schizothorax pelzami is reported from the northern Kopetdag in Turkmenistan (Starostin, 1936).

Capoeta hohenackeri Kessler, 1877 described from tributaries of the Kura and Aras rivers has a high lateral line scale count in the original description (78) and might be a mislabelled Capoeta tinca (Heckel, 1843) from Black Sea drainages of Georgia and Turkey rather than the Caspian Sea basin.

Capoeta capoeta gracilis is the subspecies of much of Iran and Capoeta capoeta heratensis (figure above) is the subspecies from the Tedzhen River basin (Berg, 1949). The former usually has one pair of barbels, the latter two pairs (but see below). Bianco and Banarescu (1982) limit C. c. gracilis to basins between the Safid River and the Atrak while C. c. capoeta is found in the Kura-Aras basin. Holčík and Jedlička (1994) consider that the two subspecies gracilis and heratensis do not exist but that the taxon C. capoeta exhibits clinal variation.

Bănărescu in Bănărescu (1999) limits C. capoeta gracilis to the Lake Orumiyeh basin and the Safid River in Iran (and the lower Kura River of Azerbaijan) while his C. capoeta aff. gracilis (an unnamed subspecies related to C. capoeta gracilis) is found along the rest of the Iranian Caspian shore. However his material was limited (and did not include any from Esfahan, the type locality of gracilis) and the analysis is based on lateral line scale counts only. Bănărescu in Bănărescu (1999) also states that C. capoeta sevangi de Filippi, 1865 is the subspecies of the Araxes River basin, presumably including Iran, distinguished from the type subspecies, C. capoeta capoeta of the Kura River basin, by having the dorsal fin margin straight or slightly convex as opposed to slightly to moderately notched. This character does not seem to be significant for such wide ranging and variable populations, which he admits in one case at least (Kura River at Mingechaur), show differences between samples from the same locality at different times.

Abdurakhmanov (1962) compares fish from the Kura River basin (presumably C. c. capoeta) with fish from the Lenkoranchai and Bilyashchai in Azerbaijan (C. c. gracilis) and finds that the latter have fewer dorsal fin rays on average, greater head length and depth, smaller eye, longer snout and postorbital distance, greater body depth and caudal peduncle depth, a shorter postdorsal distance, a shorter dorsal fin base, lesser dorsal fin height, a longer anal fin base, a greater pectoral-pelvic distance and a shorter pelvic-anal fin distance.

Dadikyan (1986) refers to Varicorhinus capoeta araxensis subsp. nov. from the Aras River basin in Armenia.

Günther (1899) points out that the considerable morphological variation shown by these fishes has resulted in numerous specific names and that it is difficult to assess them without a large comparative series and better information on localities. Berg (1948-1949) also indicates that the various subspecies are very close to each other and that their distributions are not clearly isolated.

C. c. heratensis shows major variations in body form, sometimes called morpha elata with a deep body and morpha elongata with a shallow and elongate body. These are not taxonomically significant but simply ecomorphs and all intermediates between the two extremes can be found. The deep-bodied form probably formed part of the fishes described as asmussii (Berg, 1948-1949). Reshetnikov and Shakirova (1993) list Capoeta heratensis as a full species.

Samaee et al. (2006) showed differences in morphometry between fish from six rivers along the Iranian Caspian shore with an overall assignment of individuals to a group of 88.6%. The morphometric data were mirrored by molecular data. Differences in morphometry were attributed to environmental and habitat conditions (temperature, turbidity, food availability and water depth and flow) but molecular data indicated a genetic basis, presumably through lack of gene flow between the river populations. Samaee et al. (2009) examined morphological variation with this species in the Shirud of the south Caspian Sea basin. There were no significant differences in meristic characters but morphometric characters varied and could be used to distinguish 5 groups.

Records of Capoeta capoeta from the Tigris River basin at least are probably Capoeta damascina with low scale counts (F. Krupp, in litt., 1986).

Wossughi (1978) described, in a dissertation, a subspecies from the Namak Lake basin (from "Tschmeh Jafar Abad bei Araq") but this work may not be published in the sense of the International Code of Zoological Nomenclature (Ride et al., 1985). In any case, the holotype is Capoeta aculeata and the other material comprises 21 Leuciscus (= Squalius) cephalus orientalis and 4 Capoeta aculeata (F. Krupp, in litt., 1984). The type material, all female, is stored in the Zoologischen Instituts und Zoologischen Museums der Universität Hamburg (holotype, 132 mm standard length under ZMH 5946, and 2 paratypes, 115-121 mm standard length, under ZMH 5947).

Bianco and Banarescu (1982) described C. capoeta intermedia from the Mand River in Fars but this is referred to C. damascina here (q.v.).

The types of Capoeta gibbosa are in the Zoological Institute, St. Petersburg (ZISP 11104) but have dried at some point. Their locality is given by Nikol'skii (1897) in Latin as "Bochsani in Persia orientali. 4.VII.96 (2)". This may be Bozjani at 35°48'N, 59°36'E. Berg (1949) considers that this nominal species is close to C. capoeta gracilis but is distinguished by body proportions (longer caudal peduncle and a longer head) but it is founded on only 2 specimens, hardly an adequate sample.

Types of Scaphiodon asmussii, S. gracilis and S. heratensis were not kept ? phrasing

Key characters

Berg (1948-1949) and Abdurakhmanov (1962) separate C. c. capoeta from C. capoeta gracilis, both of which may occur in Iran, by the following key:-

1(2). Dorsal fin emarginate above; lateral line scales usually 55-59; dorsal fin spine strong with numerous denticles; back behind occiput and particularly in front of the dorsal fin strongly compressed.....C. c. capoeta

2(1). Dorsal fin truncated in adults; lateral line scales usually 47-58; dorsal fin weak; back behind occiput not or only weakly compressed; radii on scales with minute recesses .....C. c. gracilis

Morphology

Dorsal fin unbranched rays 3-5, branched rays 7-10, usually 8-9; anal fin unbranched rays 2-4, branched rays 5; pectoral fin branched rays 15-20; and pelvic branched rays 7-9. Lateral line scales 46-70. In the subspecies gracilis the scales are said to be somewhat larger than in the type form, 47-58, mostly 48-50, and in heratensis the range is given as 50-60 by Berg (1948-1949). Scales are regularly arranged over the body. There is a pelvic axillary scale. Scales have a wavy anterior edge, few anterior and posterior radii and an almost central focus. Gill rakers 16-30, lower counts may refer to lower arm rakers only and total counts in the range 25-30 are probably more typical. Vertebrae 42-47. Pharyngeal teeth 2,3,4-4,3,2 or 2,3,5-4,3,2 with a hooked tip, spatulate below on posterior teeth while anterior teeth are conical. The last unbranched dorsal fin ray is strong with denticles along one half to two-thirds of its length (less strong in gracilis than in the type form). The number of barbels is variable - in fish from Uzbekistan 2 barbels (126 fish or 58.6%), 3 barbels (15 or 7.0%) or 4 barbels (74 or 34.4%)(Amanov, 1970). The subspecies heratensis is characterised by having 4 barbels but this is probably variable in Iran as in Uzbekistan. Six specimens from the Hari River basin of Iran all had 4 barbels.

Levin et al. (2005) found gracilis and heratensis (and steindachneri) to be oligovertebrate with 41-45 vertebrae, modes 42 to 44, compared to the multivertebrate type subspecies capoeta and sevangi with 45-48 vertebrae, mode 46. Morphometry and longevity also differ between these two groups and it was assumed they belong to different phyletic lines.

The Lake Sevan, Armenia subspecies (sevangi) has 2n=150 and is closer to "Barbus" than to African Varicorhinus, a genus in which Southwest Asian Capoeta were once placed (Krysanov, 1999). C. capoeta from the Safid River, the Shahrud in Rudbar and the Madarso River in Golestan National Park also have 2n=150, NF=230-234 (Pourali et al., 2000; Pourali Darestani et al., 2006). C. c. umbla from the Tigris River of Turkey had 2n=150, possibly hexaploid, with 43 meta-submetacentric chromosomes, 32 pairs of subtelo-acrocentric chromosomes with NF=236 (Kiliç Demirok and Ünlü, 2001).

Günther (1899) points out that this species shows considerable morphological variation, even in fish caught at the same place and time. The mouth can vary from straight to a gentle crescent to a distinct crescent, e.g. in three fish from the Nazlu Chai. There are also variations in dorsal fin spine development and the crown of the head can be flattened or convex.

Meristics for Iranian material: ?

Sexual dimorphism

The snout in males has 2-4 rows of tubercles and tubercles are present on scales and the rays of the anal fin. Abdurakhamanov (1962) reports that caudal peduncle length and lower caudal lobe length are longer in males while anal fin height, pelvic-anal fin distance, postorbital distance and interorbital width are greater in females.

Colour

The back is dark grey or green to brownish and the flanks light, silvery or silvery-grey, or yellowish. There may be several large black spots or blotches on the flank. The belly and lower head surface are pearly-white to dirty yellow. Scales are darkly pigmented. The operculum has a broad, yellow-gold spot. The iris is silvery, somewhat darker or yellow-golden above, or golden overall with traces of grey. The front of the dorsal fin and the margin of the caudal fin are black, and the rest of these fins are grey or yellowish-grey with some pink. The black margin to the caudal fin may be best developed on the upper and lower lobes compared to the posterior margin. The pectoral, pelvic and anal fins are grey with some pink or may be an overall pale pinkish. The peritoneum is black.

Size

Reaches 38 cm standard length and 3.5 kg (Amanov, 1970). This species reaches 43 cm in the Aras River basin of Iran (possibly C. capoeta sevangi, see above; A. Abdoli, pers. comm., 1995), 55 cm in Lake Sevan as C. capoeta sevangi (Bănărescu in Bănărescu, 1999) and 43.5 cm fork length and 1.23 kg in Çıır Lake, Turkey as C. capoeta capoeta.

Distribution

The type species is found in the Kura River basin of Azerbaijan with some Aras River basin fishes very similar. It is not known if the fishes from the Aras River basin in Iran belong to the type species or to gracilis. The subspecies Capoeta capoeta gracilis (known to Russian ichthyologists as the Lenkoran khramulya) was described from near Esfahan and is recognised as the one found over much of Iran including the Caspian Sea basin from the Astara to the Atrak including the Anzali Mordab and Gorgan Bay (Derzhavin, 1934; Bianco and Banarescu, 1982; Holčík and Oláh, 1992; Kiabi et al., 1994; Roshan Tabari, 1997; Shamsi et al., 1997; Abbasi et al., 1999; Kiabi et al., 1999; Abdoli, 2000), the Lake Orumiyeh basin including the Urmi River or Shaher Chai, Nazlu Chai and Talkheh, Tatavi and Zarrineh rivers (Günther, 1899; Abdoli, 2000) and Mahabad Dam (Abdi, 1999; www.mondialvet99.com, downloaded 31 May 2000), the Dasht-e Kavir basin (Saadati, 1977), the Gulf basin in the Zohreh, Shapur, Dalaki, Helleh, Shur rivers and the upper and lower Mand River as Capoeta capoeta intermedia ((Gh. Izadpanahi, pers. comm., 1995; M. Rabbaniha (pers. comm., 1995; Abdoli, 2000), the Esfahan basin including the Dopolan River, Gav Khuni marsh (Keyserling, 1861), the Tigris River basin in the Regab River near Kermanshah, Nashad River in the Divadarreh region, Kordestan and near Borujerd, Lorestan (records need verification by specimens), the Tedzhen River basin including the Jam and Kashaf rivers (Berg, 1949). This variable species is also recorded from the Dasht-e Kavir, Bejestan. Sirjan and  Namak Lake basins (Esmaeili et al., 2011?).

The subspecies Capoeta capoeta heratensis (the Transcaspian khramulya) is found in the Tedzhen or Harirud basin of Iran and eastwards including the Kashaf River (Abdoli, 2000). This subspecies is also recorded from the Karakum Canal and Kopetdag Reservoir in Turkmenistan (Shakirova and Sukhanova, 1994; Sal'nikov, 1995) and may eventually reach Iranian waters in the Caspian Sea basin.

Zoogeography

Saadati (1977) suggests that this species entered the Dasht-e Kavir basin from either the Tigris River basin, the Hari River basin or the Caspian Sea basin. See also above under genus.

Habitat

The habitat of this species in the Surkhandar'ya of Uzbekistan is backwaters and channels with weak current and silt beds as well as reservoirs (Amanov, 1970). In Iran, it is one of two most abundant species in Caspian rivers along with Alburnoides bipunctatus (= eichwaldii) (Iranian Fisheries Research and Training Organization Newsletter, 19:4, 1998). Günther (1899) found that Capoeta capoeta placed in saline Lake Orumiyeh water died in 3.5 minutes.

Age and growth

Life span is over 8 years and catches in Uzbekistan are dominated by fish 3-4 or 4-5 years old. Growth is fastest in the first two years of life (Amanov, 1970). Life span in Azerbaijan is over 6 years (Abdurakhmanov, 1962), in northern Anatolia 6 years where vertebrae followed by scales were the best structures for aging (Polat and Işik, 1995), while in Georgia life span exceeds 9 years (Elanidze, 1983), and in Lake Gotchka, Armenia 10 years but only 4+ years in Lenkoran (Bănărescu in Bănărescu, 1999). Günther (1899) reports on a male fish from Ula in the Lake Orumiyeh basin which was only 12.5 cm long yet a sexually mature male, perhaps an instance of a dwarf form. Canbolat et al. (1999) found life span to be over 9 years in Çıldır Lake, Turkey for Capoeta capoeta capoeta. Fish aged 6 years dominated at 31.5% and 61.7% of the sample was female.

In Madarsoo Stream of Golestan National Park, this species had age groups 0-10 years and growth parameters were Lt = 229.67 mm and K = 0.54 for males, 327.95 mm and 0.18 for females (Koohestan Eskandari, 2003). Maku Dam lake in West Azarbayjan has an estimated 9.4-10.7 tonnes of this species with a maximum sustainable yield of 4.5-4.8 t (Saiad Bourani and Ghaninejad, 2004). Average length of this population was 23.9 cm, weight was 1626.8 g and age was 2.6 years. Most fish were 3+ years old and 5+ fish were at a minimum. Infinite length and the growth coefficient were computed as 35.6 cm and 0.39 per year. Total mortality was 0.74, natural mortality 0.37 and fishing mortality 0.37. The Yasalegh Stream in the Gorgan River basin had a male to female ratio of 1:0.54, a maximum weight of 71.2 g for males and 119.4 g for females, and age range of 0-3 years, von Bertalanffy growth equations of Lt = 190(1-exp{-0.462 [t+1]} for males and Lt = 230(1-exp{-0.472 [t+0.742]} for females, and weight growth was isometric (b = 3.052 for males and 3.050 for females). Tilabad River fish had an age structure of 0-4 years and the Talar River 2-4 years, similar to Yasalegh Stream but differing from the Madarsoo Stream. The fish in the latter stream had better living conditions in a national park, no pollution, no fishing, no competition from exotic carps, no other human disturbances, no environmental stress and no food shortages. Patimar et al. (2009) found b values ranging from 2.647 (male at Chekchai) to 2.964 (females at Madarsu) indicating negative allometric growth for fish in the Gorgan River basin. They interpreted this variation to the species' response to different habitat conditions.

Food

Food is mainly detritus and ooze, with some higher plants and small amounts of blue-green algae, which is digested in an intestine almost 7 times longer than the body (Amanov, 1970). Small benthic invertebrates may also be included, such as chironomids and molluscs (Bănărescu in Bănărescu, 1999). In Maku Dam lake, this species is a detritivore consuming Chrysophyta from the phytoplankton and Cyclotella a diatom, from the benthos as well as Chironomidae and Ephemeroptera (Valipour, 2004). In the Talar and Yasalegh rivers of the eastern Caspian Sea basin, 27 genera of phytoplankton were identified in the diet, with Chrysophyta being dominant, but with some differences between older and younger fish in the species consumed (Mostafavi and Abdoli, 2005).

Reproduction

Sexual maturity in Uzbekistan is at ages 2-4 years and lengths of 15-20 cm or in some populations at 10-14 cm. Some fish mature as dwarfs before age 1 and Berg (1948-1949) reports males 8.4 cm long of the subspecies gracilis can be mature. Spawning may take place at any time during the period from March to September (Berg,1948-1949) and is intermittent with the first spawning accounting for up to 85% of the eggs and the subsequent two spawnings for the remainder. The eggs have a diameter up to 2.2 mm in the first spawning, up to 0.75 mm in the second and 0.65 mm in the third. Fecundity is up to 86,800 eggs. Eggs are laid at 50-60 cm on sand and stone beds and in water temperatures of 16-23°C (Amanov, 1970). Fecundity in the Kura River may reach 93,861 at 36-40 cm but this is for C. capoeta capoeta and fecundity for C. capoeta sevangi is less (Bănărescu in Bănărescu, 1999). Eggs are shed in running water and on lake shores, and eggs are covered by sand or small stones.

Parasites and predators

Williams et al. (1980) report the helminths Khawia armeniaca (a cestode) and Acanthocephalorhynchoides cholodkowskyi (an acanthocephalan) from this species in the Zayandeh River at Esfahan. Molnár and Jalali (1992) record for this species the monogeneans Dactylogyrus chramulii, D. gracilis and D. lenkorani in the Safid Rud, D. chramulii and D. lenkorani in the Beshar River of the Persian Gulf drainage, D. gracilis and D. lenkorani in the Zayandeh Rud, D. lenkorani in the Tonekabon and Tajan rivers of the Caspian Sea drainage and the Kor River drainage of Fars, and D. pulcher from the Safid, Tajan, Tonekabon and "Ghasemlu" rivers of the Caspian Sea basin and the Jajrud of the Namak Lake basin. Shamsi et al. (1997) report Clinostomum complanatum, a parasite causing laryngo-pharyngitis in humans, from this species, the highest rate in 9 species examined. Malek (1993) and Malek and Mobedi (2001) report Clinostomum complanatum from this species in Mazandaran, the Shiroud. Up to 60 parasites per fish are recorded, with female fish having the highest infestation (the later study showing no difference between male and female fish), infestation decreasing with increase in body length, and parasites being concentrated in the gill cavity and pharynx.

Masoumian and Pazooki (1998) surveyed myxosporeans in this species in Gilan and Mazandaran provinces, finding Myxobolus musayevi and M. samgoricus. The crustacean parasite Tracheliastes polycolpus is reported from the fins of this species in the Mahabad Dam reservoir (Abdi, 1999; www.mondialvet99.com, downloaded 31 May 2000). Masoumian et al. (2002) investigated parasites from this fish in the Aras and Mahabad dams in northwest Iran and found the protozoan Myxobolus musayevi which is also recorded from this fish in the Tajan River in Mazandaran. Mokhayer et al. (2002) report Acanthocephalorhynchoides cholodkowskyi (Quadrigyridae) from the midgut and Tracheliastes polycolpus (Lernaeopodidae) on the fins of this fish in Golestan National Park, with more parasites on male fish and differences by season and station. Naem et al. (2002) found the following parasites on the gills of this species from the western branch of the Safid River, namely the protozoan Trichodina sp. and the monogenean trematode Dactylogyrus lenkorani. Mirhasheminasab and Pazooki (2003) list Ergasilus peregrinus, Tracheliastes polycolpus and Lernaea cyprinacea from this species in Mahabad Reservoir, the latter being the most dangerous parasite. Rohei Aminjan and Malek (2004) found 9 parasite species in fish from the Shiroud, namely the trematodes Clinostomum complanatum, Diplostomum spathaceum, Posthodiplostomum cuticola, Allocreadium sp., the monogeneans Dactylogyrus pulcher, D. lenkorani, Gyrodactylus mutabilitas and the nematodes Rhabdochona fortunatowi and Capillaria sp. Masoumian et al. (2005) recorded the protozoan parasites Ichthyophthirius multifilis, Trichodina perforata, Chilodonella, sp., Amphileptus branchiarum, Tetrahymena pyriformis, Apiosoma sp., and Vorticella sp. from this species in water bodies in West Azarbayjan. Araghi Soureh and Jalali Jafari (2006) recorded Dactylogyrus gracilis, D. charmulii, D. lenkorani and D. kendalanicus from this species in the Mahabad River of the Lake Orumiyeh basin, the latter species being a new record for Iran. Pazooki et al. (2007) recorded various parasites from localities in West Azarbayjan Province, namely Diplostomum spathaceum, Ligula intestinalis, Digrama sp., Rhabdochona hellichi, Argulus foliaceus, Allocreadium isoporum, Lamprolegna compacta, Myxobolus cristatus and M. musajevi. Pazooki et al. (2005) record Tracheliastes longicollis, Lamprolegna compacta, Neoechinorhynchus rutili, Capillaria sp., Myxobolus musajevi, M. cristatus, Trichodina perforata, Chilodonella piscicola, Ichthyophthirius multifilis and Ichthyobodo necatrix from this species in waterbodies of Zanjan Province. Pazooki et al. (2006) record the monogeneans Dactylogyrus chramuli, D. gracilis, D. lamellatus, D. lenkorani, D. pulcher and Gyrodactylus sp. from this fish in Zanjan Province. Masoumian et al. (2007) record the myxosporean parasite Myxobolus musajevi from this species in the Zayandeh River. Miar et al. (2008) examined fish in Valasht Lake and the Chalus River, Mazandaran and found the metazoan Myxobolus saidovi.

Economic importan

The subspecies Capoeta capoeta heratensis is a food fish in Uzbekistan (Amanov, 1970) and C. capoeta sevangi and C. capoeta capoeta are commercially important in Lake Gotchka, Armenia and eastern Georgia and Azerbaijan respectively (Bănărescu in Bănărescu, 1999). It is also used in sport fishing in Iran (Samaee et al., 2006).

Conservation

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

Further work

The relationships of the various subspecies need study to determine if they are in fact good species.

Sources

Type material: See above, Capoeta gibbosa (ZISP 11104).

Iranian material: CMNFI 1970-0512, 3, mm standard length, Gilan, Shalman River (37º08'N, 50º15'E); CMNFI 1970-0514, 17, ? mm standard length, Gilan, Shafa River estuary (37º55'N, 49º09'E); CMNFI 1970-0516, 6, ? mm standard length, Gilan, Lemir River (38º14'N, 48º52'30"E); CMNFI 1970-0519, 2, ? mm standard length, Gilan, Chelvand River (ca. 38º18'N, ca. 48º52'E); CMNFI 1970-0520, 2 ?7 on data sheet, 94.0-100.9 mm standard length, Gilan, Astara River (ca. 38º25'N, ca. 48º52'E); CMNFI 1970-0521, 3, ? mm standard length, Gilan, Safid River near Lulaman (no other locality data); CMNFI 1970-0522, 10, ? mm standard length, Gilan, Safid River at Astaneh bridge (37º16'30"N, 49º56'E); CMNFI 1970-0525, 5, 92.8-146.1 mm standard length, Gilan, Safid River near Mohsenabad (ca. 37º22'N, ca. 49º57'E); ?see data sheets 78, 22.1-160.4 CMNFI 1970-0526, 8, ? mm standard length, Gilan, Safid River 6 km below Astaneh bridge (37º19'N, 49º57'30"E); CMNFI 1970-0531, 7, 60.2-84.6 mm standard length, Mazandaran, Larim River (36º46'N, 52º58'E); CMNFI 1970-0536, 2, ?5 on data sheets 101.4-125.4 mm standard length, Gilan, Siah River estuary (36º53'N, 49º32'E); CMNFI 1970-0538, 1, ?5 on data sheet 95.4 mm standard length, Gilan, Qezel Owzan River near Manjil (36º44'N, 49º24'E); CMNFI 1970-0557, 3, ? mm standard length, Azarbayjan-e Bakhtari, Shaher Chay (no other locality data); CMNFI 1970-0559, 6, 83.9-125.4 mm standard length, Azarbayjan-e Bakhtari, Baranduz Chay (ca. 37º25'N, ca. 45º10'E); CMNFI 1970-0568, 9, ? mm standard length, Gilan, Caspian Sea at Kazian beach (ca. 37º29'N, ca. 49º29'E); CMNFI 1970-0577, , mm standard length, Gilan, Caspian Sea at Astara (ca. 38º26'N, ca. 48º53'E); CMNFI 1970-0583, 8, 34.1-93.9 mm standard length, Gilan, Nahang Roga (37º28'N, 49º28'E); CMNFI 1979-0589, , mm standard length, Gilan, Safid River opposite Kisom (37º12'N, 49º54'E); CMNFI 1979-0242, 27, 25.6-107.0 mm standard length, Fars, river at Izadkhvast (31º31'N, 52º07'E); check ID? CMNFI 1979-0249, 33, 66.4-114.2 mm standard length, Esfahan, stream at Dizaj (31º55'N, 51º30'E); check ID? CMNFI 1979-0429, 1, ? mm standard length, Mazandaran, Chalus River (36º34'N, 51º23'E); CMNFI 1979-0432, 1, ? mm standard length, Mazandaran, Sardab River branch (36º41'N, 51º22'E); CMNFI 1979-0434, 1, ? mm standard length, Mazandaran, Shir River (36º51'N, 50º49'E); CMNFI 1979-0435, 1, ? mm standard length, Gilan, stream 10 km west of Ramsar (36º57'N, 50º37'E); CMNFI 1979-0433, 1, 115.2 mm standard length, Mazandaran, stream 18 km west of Chalus (36º42'N, 51º15'E); CMNFI 1979-0438, 2, 142.4-144.8 mm standard length, Gilan, Gholab Ghir River (37º27'N, 49º37'E); CMNFI 1979-0441, 1, 121.9 mm standard length, Gilan, river 14 km south of Hashtpar (37º42'N, 48º58'E); CMNFI 1979-0443, 1, ? mm standard length, Gilan, river 34 km west of Hashtpar (38º06'N, 48º53'E); CMNFI 1979-0444, 1, ? mm standard length, Gilan, Chubar River (38º11'N, 48º52'30"E); CMNFI 1979-0446, 1, ? mm standard length, Gilan, Astara River (38º26'30"N, 48º51'E); CMNFI 1979-0449, 2, ? mm standard length, Azarbayjan-e Khavari, river 18 km from Khalkhal (ca. 37º42'N, ca. 48º27'E); CMNFI 1979-0451, 30, 35.8-97.3 mm standard length, Azarbayjan-e Khavari, Qezel Owzan River (ca. 37º30'N, ca. 47º57'E); CMNFI 1979-0452, 1, 79.7 mm standard length, Azarbayjan-e Khavari, Qezel Owzan River 6 km from Mianeh (37º23'N, 47º45'E); CMNFI 1979-0453, 24, 36.1-111.1 mm standard length, Zanjan, Zanjan River (37º06'N, 47º56'E); CMNFI 1979-0469, 2, 56.6-76.2 mm standard length, Mazandaran, river 36 km west of Alamdeh (36º37'30"N, 51º35'E); CMNFI 1979-0474, 1, ? mm standard length, Mazandaran, Tajan River (36º34'N, 53º05'E); CMNFI 1979-0475, 1, 86.4 mm standard length, Mazandaran, stream on road to Bandar-e Shah (36º46'N, 54º00'E); CMNFI 1979-0480, 2, ? mm standard length, Mazandaran, Gorgan River at Gonbad-e Kavus (37º15'30'N, 55º09'E); CMNFI 1979-0481, 3, 101.9-188.0 mm standard length, Mazandaran, stream 3 km west of Ghalahleekesh (37º18'30"N, 55º31'E); CMNFI 1979-0482, 2, ? mm standard length, Mazandaran, river 2km west of Ghalahleekesh (37º19'30'N, 55º31'E); CMNFI 1979-0483, 4, 121.6-160.5 mm standard length, Mazandaran, river 28 km west of Dasht (37º23'30"N, 55º51'30"E); CMNFI 1979-0485, 3, 71.2-99.1 mm standard length, Khorasan, stream 28 km west of Bojnurd (37º33'N, 57º04'E); CMNFI 1979-0486, 66, 17.5-97.8 mm standard length, Mazandaran, stream in Atrak River draiange (37º44'N, 56º18'E); CMNFI 1979-0487, 20, ? mm standard length, Mazandaran, spring 2 km from Maraveh Tappeh (37º54'N, 55º58'E); CMNFI 1979-0488, 9, 29.7-140.4 mm standard length, Mazandaran, Atrak River at Maraveh Tappeh (37º55'N, 55º57'30"E); CMNFI 1979-0489, 78, ? mm standard length, Mazandaran, stream 13 km from Maraveh Tappeh (37º50'N, 55º53'E); CMNFI 1979-0490, 14, 21.0-108.4 mm standard length, Mazandaran, stream in Gorgan River drainage (ca. 37º39'N, ca. 55º42'E); CMNFI 1979-0491 2, ? mm standard length, Mazandaran, Gorgan River northeast of Kalaleh (ca. 37º33'N, ca. 55º44'E); CMNFI 1979-0492, 25? check jar, 9.3-183.4 mm standard length, Mazandaran, river in Gorgan River drainage (37º05'N, 55º15'E); CMNFI 1979-0695, 13, ? mm standard length, Gilan, Safid River at Manjil Bridge (36º46'N, 49º24'E); CMNFI 1997-0003, , mm standard length, (); CMNFI 1980-0116, 1, ? mm standard length, Gilan, Safid River at Astaneh (37º16'30"N, 49º56'E); CMNFI 1980-0120, , mm standard length, Mazandaran, Babol River at Babol Sar (36º43'N, 52º39'E); CMNFI 1980-0121, , mm standard length, Gilan, Shafa River estuary (37º35'N, 49º09'E); CMNFI 1980-0123, , mm standard length, Gilan, Safid River (ca. 37º22'N, ca. 49º57'E); CMNFI 1980-0141, , mm standard length, Gilan, Lisar River estuary (37º59'N, 48º56'E); CMNFI 1991-0163, , mm standard length, Mazandaran, Ramian River (36º58'N, 55º07'E); CMNFI 1993-0138, 1, mm standard length, Khorasan, Bazangan Lake (36º18'N, 60º27'E); CMNFI 2007-0014, 4, 39.4-99.1 mm standard length, Khorasan, pool in Kuh-e Sang Park, Mashhad (ca. 36º18'N, ca. 59º36'E); CMNFI 2007-0086, 6, ? mm standard length, Azarbayjan-e Khavari, Qareh Su basin near Nir (ca. 38º02'N, ca. 48º00'E); CMNFI 2007-0087, 1, ? mm standard length, Azarbayjan-e Khavari, Qareh Su north of Ardebil (38º22'N, 48º19'E); CMNFI 2007-0088, 5, ? mm standard length, Azarbayjan-e Khavari, Qareh Su east of Lari (38º30'N, 48º03'E); CMNFI 2007-0089, 4, ? mm standard length, Azarbayjan-e Khavari, Ahar Chay at Ahar (38º28'N, 47º03'E); CMNFI 2007-0093, 13, ? mm standard length, Azarbayjan-e Bakhtari, Qotur River south of Khvoy (38º30'N, 44º58'E); CMNFI 2007-0094, 6, ? mm standard length, Azarbayjan-e Bakhtari, Nazlu River north of Reza'iyeh (ca. 37º42'N, ca. 45º04'E); checkID? CMNFI 2007-0095, 2, ? mm standard length, Azarbayjan-e Bakhtari, Shahr Chay southwest of Reza'iyeh (ca. 37º27'N, ca. 44º56'E); checkID? CMNFI 2007-0096, 5, ? mm standard length, Azarbayjan-e Bakhtari, Qasemul River in Baranduz Chay basin (ca. 37º25'N, ca. 45º10'E); checkID? CMNFI 2007-0098, 2, ? mm standard length, Azarbayjan-e Bakhtari, river south of Mahabad (ca. 36º42'N, ca. 45º41'E); CMNFI 2007-0099, 1, ? mm standard length, Azarbayjan-e Bakhtari, Kalwi Chay west of Mahabad (ca. 36º35'N, ca. 45º25'E); checkID? CMNFI 2007-0101, 1, ? mm standard length, Azarbayjan-e Bakhtari, Tata'u River south of Miandow Ab (ca. 36º54'N, ca. 46º07'E); CMNFI 2007-0102, 4, ? mm standard length, Azarbayjan-e Bakhtari, Zarineh River near Miandow Ab (ca. 37º00'N, ca. 46º07'E); CMNFI 2007-0103, 9, ? mm standard length, Kordestan, Zarineh River basin north of Saqqez (ca. 36º18'N, ca. 46º16'E); CMNFI 2007-0104, 4, ? mm standard length, Kordestan, Zarineh River basin south of Saqqez (ca. 36º12'N, ca. 46º18'E); CMNFI 2007-0105, 7, ? mm standard length, Kordestan, Zarineh River basin south of Saqqez (ca. 36º06'N, ca. 46º20'E); CMNFI 2007-0106, 9, ? mm standard length, Kordestan, Qezel Owzan River basin near Divandarreh (ca. 35º52'N, ca. 47º05'E); CMNFI 2007-0107, 10, ? mm standard length, Kordestan, Qezel Owzan River basin near Bijar (ca. 35º54'N, ca. 47º20'E); ZSM 24500, 6, 24.2-31.0 mm standard length, Khorasan, stream near Bojnurd (no other locality data); uncatalogued, 2, 75.8-85.5 mm standard length, Khorasan, Hari River at Sarakhs (36º32'N, 61º11'E).

Capoeta damascina
(Valenciennes, 1842)

Common names

sardeh (= cold one, probably zardeh = yellow one, is more correct and appropriate based on yellow-tinged flank) or سياه ماهي (= siah mahi, meaning black fish) in the Caspian basin; siah mahi damascina; tu'ini (meaning unknown) or gel cheragh (= mud-eater, mud-grazer) or tu'ini gelkhorak (= mud-eater, mud-grazer) in Khuzestan; qezel ala (red spots) in Chahar Mahall but mistakenly.

[twena, toyoueni or toueni, bertin or bartin, tin, zardah masih, tela shami; kollur, kellur, kollur hadjiari (= the pilgrim or migrating kollur), kellur dischileki (= the strawberry-coloured kollur), kollur achmar (= the red kollur) and kollur aschkar (= the brown kollur), all at Aleppo; all in Arabic; Mesopotamian barb].

Systematics

Gobio damascinus was described from the "fleuve de Damas" (= river of Damascus, Syria).

Synonyms are Scaphiodon capoeta Heckel, 1843 (non sensu Güldenstädt, 1773) described from "Aleppo", Scaphiodon fratercula Heckel, 1843 described from "Gewässern von Damascus", possibly Scaphiodon Umbla Heckel, 1843 described from the "Tigris bei Mossul", Scaphiodon socialis Heckel, 1843 described from "Um Damascus" (= around Damascus) (Heckel, 1843b) and later more completely from the "Orontes" (Heckel, 1846-1849a) (placed in Scaphiodon Capoëta of Heckel by Steindachner (1864)), Scaphiodon peregrinorum Heckel, 1843 described from "Aleppo" and later from "Fluss Kueik bei Aleppo", Chondrostoma syriacum Valenciennes, 1844 from Abraham's River at the foot of Mount Sinai, Egypt (the correct locality is probably in the Jordan River basin (Coad and Krupp, 1994)), Scaphiodon Amir Heckel, 1849 described from the "Araxes" (= Kor River, Fars), Scaphiodon niger Heckel, 1849 described from the "Araxes oder Benth-Amir" (= Kor River, also known as the Bandamir River), Scaphiodon Saadii Heckel, 1849 described from the "Quellen des Saadi" (Sa`di at 29°37'N, 52°35'E, now within the city of Shiraz) and the "Nähe von Persepolis" (= probably the Pulvar (= Sivan) River near Persepolis, Fars), Scaphiodon chebisiensis Keyserling, 1861 from "Wasserleitung in Chebis" (= canal in Chebis, probably Khabis or Shahdad at 30°25'N, 57°42'E in Kerman), Scaphiodon rostratus Keyserling, 1861 from "Wasserleitungun in der Umgegend von Jezd. Das abgebildete Exemplar stammte aus Meibut" (= canals in the vicinity of Yazd. The specimen drawn originated from Meibut, probably Meybod at 32°14'N, 54°01'E), Barbus belayewi Menon, 1960 (Menon and Yazdani (1968) date this species as 1960, presumably the 1956 edition of the journal was delayed) from the "Tigris, Baghdad, Iraq", and Capoeta capoeta intermedia Bianco and Banarescu, 1982 (non Capoeta intermedia Temminck and Schlegel, 1846 = Acheilognathus lanceolata (Temminck and Schlegel, 1846) (see Boeseman, 1947)) described from the "Mand River near Akbar, southern Iran".

The synonymy of Barbus belayewi is suggested by F. Krupp (in litt., 1986) and W. Rainboth (pers. comm., 1986). The synonymy of S. fratercula is pointed out by Berg (1949) since the species was founded on low lateral line scale counts, a variable character in C. damascina, and on a larger orbit but Heckel's comparison was between fish of greatly differing size and no allowance was made for allometry.

Karaman (1969) places damascina in Capoeta capoeta as a subspecies and umbla as another subspecies. Berg (1949) and Saadati (1977) recognise umbla as a distinct species. The latter is distinguished from the former by a higher scale count (87-99), higher dorsal fin branched rays (9-10), longer dorsal fin, longer caudal fin (shorter than or equal to head length in C. damascina), a markedly transverse mouth, and a weaker dorsal fin spine. Saadati (1977) considers fratercula to be a distinct species from the Tigris and Mand rivers in Iran based on scale count (58-66), more gill rakers (20-22), and a more serrated dorsal fin spine; or a subspecies of Capoeta capoeta based on a close similarity in scale counts, average number of gill rakers, and the dorsal fin origin being anterior to that of the pelvic fins. He also considers that Scaphiodon niger from the Kor River of Fars is possibly a synonym of fratercula. Krupp (1985c) considers the synonymy of C. damascina and C. capoeta as extremely doubtful after examining topotypic material.

Bianco and Banarescu (1982) recognise Capoeta saadi as a distinct species based on an arched mouth rather than transverse as in most subspecies of Capoeta capoeta, with a lightly developed horny cover on the lip, a feebly ossified dorsal fin spine, 13-17 gill rakers, modally 8 dorsal fin branched rays, 53-76 lateral line scales and 24-28 scales around the caudal peduncle. However they do point out the extreme variability in scale counts, for example, from fish taken in the same locality and even between opposite sides of the same fish (5 more scales on one side than the other!). Designation of subspecies on such variable characters is difficult and would require very large series and multivariate analysis techniques. Bianco and Banarescu (1982) regard C. c. intermedia as intermediate between C. c. umbla and their C. c. macrolepis on the basis of scale counts, gill raker counts, smaller transverse mouth than in umbla and a rather light colouration.

Capoeta damascina, with a wide distribution and wide variation in morphology, must be regarded as a species complex until detailed analyses can be carried out. Final resolution of the species composition of this complex may well require extensive material for molecular analyses, as well as re-examination of types and topotypic material over the whole range of the taxon.

The syntypes of Gobio damascinus are in the Muséum national d'Histoire naturelle, Paris (MNHN 4494, 2 specimens, 169-179 mm standard length, Damascus, Syria, Bové, MNHN 3948, 1, 289 mm standard length, Nahr Barada, Syria and MNHN A.3947, 1, 169 mm standard length, Syria) (Krupp, 1985c). Bertin and Estève (1948) give 200-210 mm total length for MNHN 4494 and 330-390 mm total length for MNHN 3947, 3948 and A.789. Eschmeyer et al. (1996) list MNHN 4494 as the lectotype (as designated by Krupp and Schneider (1989) although this collection comprises two fish) with MNHN 3947 (1, dry) and MNHN 3948 (1, dry) and possibly MNHN A.789 (1) as paralectotypes. The latter is listed as a syntype in Bertin and Estève (1948) although the localities listed in this article "Fl. Jourdain, à Damas (Syrie)" is obviously an error on geographical grounds.

Syntypes of Scaphiodon capoeta are in the Naturhistorisches Museum Wien under NMW 51650 (1 fish), NMW 51831 (1), and NMW 55845-55846 (2). Heckel (1843) lists 2 specimens in his description.

The holotype of Chondrostoma syriacum is in the Muséum national d'Histoire naturelle, Paris under MNHN 1945 (Eschmeyer et al., 1996).

The holotype of Capoeta capoeta intermedia is in the Istituto di Zoologia dell'Universitá di L'Aquila, Italy (IZA 7892) and is 92.5 mm standard length, collected by P. Bianco and S Zerunian, 27/5/1976. There are 62 paratypes (IZA 7893) from the same collection as the holotype measuring 36-87 mm standard length and 13 paratypes uncatalogued in the Institutul de Stiinte Biologice, Bucuresti, Romania (ISBB) measuring 68-86 mm standard length (Bianco and Banarescu, 1982). Another paratype under IZA 7894 measures 105.5 mm standard length was examined by me. A paratype of Capoeta capoeta intermedia from the Mand River in Fars is in the Zoologischen Instituts und Zoologischen Museums der Universität Hamburg (ZMH 6090, 83.2 mm standard length) (Wilkens and Dohse, 1993; examined by me), one paratype from the Mand is in the California Academy of Sciences, San Francisco (CAS 48113), one paratype from the Mand is in the United States National Museum, Washington (USNM 227935), and 6 paratypes are in the Canadian Museum of Nature, Ottawa under CMNFI 1982-0367 (formerly IZA 7893).

The holotype of Scaphiodon fratercula was taken from "Gewässern von Damascus", the syntypes of Scaphiodon umbla from the "Tigris bei Mossul", the types of Scaphiodon socialis from "Um Damascus" (but listed as "Orontes" in the catalogue in Vienna, possibly in confusion as this part of the catalogue has been overwritten), and the types of Scaphiodon peregrinorum from "Um Aleppo" according to Heckel (1843b) and "Fluss Kueik bei Aleppo" according to Heckel (1846-1849a).

Two syntypes of Scaphiodon niger are in the Naturhistorisches Museum Wien under NMW 51655 with standard lengths of 140.4 and 188.5 mm (another syntype is under NMW 51654 (232.7 mm), and a fourth under NMW 51656 as seen by me; all 4 are listed as syntypes in the 1997 Vienna card index). Eight syntypes of Scaphiodon amir are under NMW 61472 and measure 42-59 mm standard length and there are also 6 fish under NMW 46081 (138.1-282.3 mm standard length); however the card index in 1997 lists only NMW 46081 (6) and 16508 (1, dried). Fifteen syntypes in the catalogue (18 seen by me and in the Vienna card index in 1997) of Scaphiodon saadii from Sa`di are under NMW 51666 (Eschmeyer et al. (1996) have 52666, apparently in error) and measure 58-123 mm standard length (18.3-123.8 mm standard length when measured by me) with a further 4 syntypes from Persepolis under NMW 55900 measuring 84-114 mm standard length (Kähsbauer, 1964; not in the 1997 card index). There is also 1 syntype (RMNH 3166) in the Rijksmuseum van Natuurlijke Historie, Leiden from NMW (Eschmeyer et al., 1996).

The catalogue in Vienna lists no fish opposite the name S. niger, 6 and 2 fish in one column and 5 in the adjacent column for Scaphiodon amir (cf. above), 10 fish in one column and 10 in the adjacent column for S. saadii (cf. above).

A dried syntype of C. umbla is in the Senckenberg Museum Frankfurt (SMF 6777, formerly NMW) (F. Krupp, pers. comm., 1985; ca. 262.3 m standard length), 2 syntypes are in Naturhistorisches Museum Wien (NMW 55932-55933) and another syntype is under NMW 55934. Eschmeyer et al. (1996) also lists NMW 79373-74, both dried. The catalogue in Vienna lists 2 fish in spirits and 2 fish stuffed and the card index in 1997 lists as syntypes NMW 55932-33 and 79373-74 (dried).

Eschmeyer et al. (1996) note that there are no types of Scaphiodon fratercula in the Naturhistorisches Museum Wien.

Two fish are labelled as syntypes of Scaphiodon socialis in the Naturhistorisches Museum Wien (NMW 55855) which agrees with Heckel's text although the catalogue lists only 1 specimen. Eschmeyer et al. (1996) state that there are no types at NMW presumably after Krupp and Schneider (1989) who state that NMW 55670 (1 fish), 55843 (2) and 55855 (2) are not types.

The types of Scaphiodon peregrinorum number 6 according to the catalogue in the Naturhistorisches Museum Wien and may comprise all or part of NMW 51658 (1), NMW 51659 (1), NMW 51660 (1), NMW 51661 (1), NMW 51662 (1), NMW 51663 (1) NMW 51664 (3), and NMW 51665 (1), all labelled as from "Kueik" and possibly RMNH 2681 (3) in the Rijksmuseum van Natuurlijke Historie, Leiden from NMW (Eschmeyer et al., 1996).

The types of Barbus belayewi are in the Zoological Survey of India, Calcutta, the holotype being ZSI F1046/2 and a paratype ZSI F1047/2 (Menon, 1960; Menon and Yazdani, 1968).

Types of Scaphiodon chebisiensis and Scaphiodon rostratus were not kept.? phrasing

Key characters

The mode of 9 dorsal fin rays, small scales, and the presence of large black blotches often distinguish this species from other Capoeta in Iran.

Morphology

Dorsal fin with 3-5 unbranched rays and 8-10 branched rays (Krupp (1985c) gives frequency distributions for his material from Turkey, Syria, Lebanon, Israel and Jordan as 8(52), 9(144) and 10(4)), anal fin with 3 unbranched and 5-6 branched rays (5(179), 6(21) after Krupp (1985c)), pectoral fin branched rays 15-20, and pelvic fin branched rays 8-10. Lateral line scales 60-99. Gill rakers 17-25 (Saadati (1977) gives 9-21!; Krupp (1985c) 12-18 for the lower arm of the arch, Berg (1949) up to 23 on the lower arm). Pharyngeal teeth 2,3,4-4,3,2, often 2,3,5-5,3,2, with spoon-shaped crowns. The mouth is usually horseshoe-shaped, seldom transverse. The last unbranched dorsal fin ray is moderate to strong with denticles along two-thirds of its length. Heckel (1846-1849b) distinguished his Scaphiodon amir and S. niger by the dorsal fin denticles being horizontal or perpendicular to the spine, not hooked downward as in related species. Berg (1949) did not attach any significance to this character, finding it in small fish from the Sarhadd of Baluchestan and from Jordan.

The karyotype for fish in the Tigris River basin of Turkey identified as Capoeta capoeta umbla is 2n=150, possibly hexaploid (Kılıç Demirok and Ünlü, 2001) and of fish identified as C. damascina from the Wadi Karak, Jordan 2n=148-150, indicating a hexaploid species (Gorshkova et al., 2002).

Body form is highly variable as are scale counts between populations and even within populations when large series are examined (Krupp, 1985c). Subspecific designations can only be valid if very large series from the whole range of the species are compared.

Meristic values for Iranian specimens are :

IZA7892, 7894 (2 fish) D8, A5, P15-15, V7, ll 62-64, gr 24

Sexual dimorphism

Males develop breeding tubercles around the snout and the posterior body on both sides of the lateral line (Khalaf, 1987).

Colour

The back is dark brown or brownish to olive or blue-grey, the flanks silvery with some yellowish tinges, sometimes golden, or yellow-brown or reddish-brown above the lateral line, silvery below. The belly is white to yellowish. Cheeks are golden. Dark brown or black spots numbering up to 20 may be scattered irregularly on the flanks. Fins are reddish-brown, yellowish or grey and may be hyaline. The caudal and pectoral fins may be very dark compared to other fins. The pectoral and pelvic fins may a light pink tinge. The cartilaginous edge to the lower jaw is bright yellow to red-yellow. The peritoneum is black. Some fish may be very black with only the underside of the head and belly yellowish-white (specimens described by Heckel (1846-1849b) as Scaphiodon niger; however since these fish "decompose quickly in the commonly used ethyl alcohol concentrations", they may have been poorly preserved and the black colouration resulted from partial decomposition).

Size

Attains 35.3 cm standard length, about 45.0 cm total length and 0.5 kg.

Distribution

Found from Turkey, Syria, Lebanon and Israel to Iran. In Iran, it is reported from the Tigris River including the the Regab River in Kurdistan and the Selakhor River near Borujerd, the Jarrahi, Marun, Karun, Kuhrang, Bazoft and Khersan rivers, throughout the Dez and Karkheh basins to their uppermost reaches; Lake Zarivar; Esfahan including the Dopolan and Zayandeh rivers, Dasht-e Kavir including the Jajarm and upper Kal Shur rivers, Namak Lake including the Karaj, Shur, Abhar, Qareh Su and Qom rivers; Kor River, Lake Maharlu, Gulf including the Zohreh River and its Kheirabad tributary and the Mand River near Akbar and its Shur (Dasht-e Palang) River tributary, Kerman-Na'in, Dasht-e Lut, Sirjan, Hormuz including the middle to upper Hasan Langi, Kul and its Shur River tributary, and Hamun-e Jaz Murian basins (Lovett, 1873; Nikol'skii, 1899; Berg, 1949; Kähsbauer, 1964; Spillman, 1972; Armantrout, 1980; Rainboth, 1981; Bianco and Banarescu, 1982; Abdoli, 2000; Ghorbani Chafi, 2000; R. Mehrani, pers. comm., 2000; Jalali et al., 2005; Esmaeili et al., 2011?).

Zoogeography

Its relationships with other Capoeta species is generally unclear, as is the status of isolated populations some of which have been named. The larger zoogeographical relationships of this and other Capoeta species remain uncertain. See also above under genus.

Habitat

Unknown in detail.

Age and growth

All males are mature at 18 cm and all females at 20 cm in Khalaf's (1987) study in the Lebanon. In Lake Kinneret, Israel, Stoumboudi et al. (1993) found that fish longer than 25 cm have developed gonads, occasionally males mature between 16 and 25 cm as did females between 20 and 25 cm. Khalaf et al. (2002) found 6 age classes (1+ to 6+) in the Nahr el Khalb, a Lebanese stream. Maximal growth was in July and August and minimal growth between December and February. Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 40 Iranian fish measuring 5.23-19.87 cm standard length. The a-value was 0.0282 and the b-value 2.890 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases).

Food

Khalaf (1985) and Spataru and Gophen (1986) examined the food of this species in Lebanon and Israel respectively. Benthic diatoms and filamentous algae are the main foods. Some other algal species and some zoobenthic organisms are present along with large quantities of mud. The species is classified as a phytobenthophagous fish, one that takes its food from bottom sediments. Leaf remains have also been found in gut contents. Abdoli (2000) lists variety of insects: Chironomidae, Formicidae, Epididae, Empididae, Tipulidae, Tabanidae, Simuliidae, Hydroptilidae, Grouvellinus, Elmis, Hydropsyche, Heptagenia, Baetis and hydracariens.

Reproduction

Khalaf (1987) examined the reproductive cycle in this species for Lebanese waters. Spawning begins in May and ends in July. Eggs number up to 5,138 and egg diameters are up to 2.2 mm. In marked contrast, Stoumboudi et al. (1993) found that gonad weights are greatest in January in Lake Kinneret, Israel, 4 months earlier. This may be evidence of different temperature regimes or populational variation. Fishelson et al. (1996) confirm that this species migrates in winter, December to February in the upper Jordan River of Israel, the process being initiated by rainfall and flooding and a decrease in temperature to 16-18°C. The gonadosomatic index is highest in February and the final months of reproduction are March to May. Lake dwelling fish aggregate and swim up streams as far as 25 km and altitudes of 400-900 m, fattening and ripening at the spawning site. They can jump rapids on this migration (and in Iran large fish cornered in small streams will jump over seine nets!). The females excavate a shallow nest in which to deposit adhesive eggs, up to 4.5 mm in diameter. Dozens of nests are found close together and sand and gravel stirred up by the excavation covers adjacent nests. After spawning the adults return downstream to the stream mouth and lake.

Parasites and predators

Dollfus (1970) describes a new cestode Coelobothrium monodi from this species at "Nasratabad", possibly from the Dasht-e Lut basin. Jalali et al. (1995) describe two new species of monogeneans, Dactylogyrus rohdeianus and D. capoetae, from fish caught in the "Chaghalnandi" River, a Karkheh River tributary north of Ahvaz. González-Solís et al. (1997) report the nematodes Rhabdochona denudata and Rhabdochona fortunatowi from this species in the Mand River, Fars. O. M. Amin (pers. comm., 1998) has identified the acanthocephalan Acanthocephalorhynchoides cholodkowskyi from specimens collected in the Mand River west of Shiraz, Fars. Jalali et al. (2002) and Jalali and Barzegar (2006) record Trichodina pediculus, Dogielius molnari, Gyrodactylus sp., Dactylogyrus carassobabrbi and D. lenkorani from this species in Lake Zarivar. Barzegar et al. (2004) examined this species for parasites in fish from the Beheshtabad river in Chahar Mahall va Bakhtiari Province and found Dactylogyrus lenkorani, Gyrodactylus pulcher, Dactyolgyrus sp., Allocreadium isoporum and Myxobolus molnari. Mehdipoor et al. (2004) record the monogeneans Dactylogyrus lenkorani and D. pulcher in Zayandeh River fish. Barzegar and Jalali (2006) report a parasite in this species from Kaftar Lake as Dactylogyrus lenkorani. Masoumian et al. (2007) record the myxosporean parasites Myxobolus samgoricus and M. varicorhini from this species in the Zayandeh River. Nazari Chamak et al. (2010) found the following myxozoan parasites in the genus Myxobolus: buckei, cristatus, karelicus, musajevi, samgoricus, suturalis and varicorhini in fish from the Halil River, Kerman.

Economic importance

This species is of no economic importance in Iran although in Israel annual catches in Lake Kinneret have been as high as 29 tonnes or 7% of the total fishery (Spataru and Gophen, 1986). Heckel (1846-1849b) reports that this species was "greatly appreciated as food fish by the local people" in the Kor River basin, Fars (as his Scaphiodon amir).

The eggs are reputedly poisonous and this is said to account for the low population of introduced Oncorhynchus mykiss in Gahaar Lake, Lorestan (R. Mehrani, pers. comm., 2000).

Conservation

A widely distributed species, probably not in need of conservation. Endangered in Turkey (Fricke et al., 2007).

Further work

The relationships of isolated and named taxa under this species, and the relationships of this species to other Capoeta, would benefit from molecular analyses.

Sources

Type material: See above, Capoeta capoeta intermedia (IZA 7892, 7894, CMNFI 1982-0367 (formerly IZA 7893), ZMH 6090) P. G. Bianco. ? check this last speciemne; Scaphiodon amir (NMW 61472, 46081); Scaphiodon niger (NMW 51655, 51654, 51656); Scaphiodon saadii (NMW 51666).

Iranian material: CMNFI 1993-0154, 1, ? mm standard length, Markazi, Sharra River near Far (34º03'N, 49º20'E); CMNFI 1995-0020, , mm standard length, (); CMNFI 1995-0021, , mm standard length, (); CMNFI 1997-0004, , mm standard length, ();

Comparative material: BM(NH) 1974.2.22:1856, 227.1 mm standard length, Iraq, Mosul (36º20'N, 43º08'E); BM(NH) 1934.9.5:3-5, 6, 14.8-45.5 mm standard length, Rawanduz River, Razanok.

Capoeta fusca
Nikol'skii, 1897

Common names

سياه ماهي (= siah mahi, meaning black fish).

Systematics

The 2 syntypes, listed in Latin as from "Mondechi in Persia orientali", are in the Zoological Institute, St. Petersburg (ZISP 11108) and measure 121.9-172.9 mm standard length. Berg (1949) gives the locality in Russian as "Mondekhi, northern periphery of the Bajistan Salt Desert in southeast Khorasan". This locality is possibly Mandehi or Miandehi at 34°53'N, 58°38'E. Nikol'skii (1897) lists a series of specimens in Latin, presumably all of which he regarded as types, sic:- "11108. Mondechi in Persia orientali. 12.IV.96 (2). 11109. Persia orientalis. 1896. (6). 11110. Persia orientalis. 1896. (5). 11111. Persia orientalis. 1896. 11112. Kuss in Persia orientali. 6.IV.96.", the last two lacking number of specimens. Berg (1949) gives 20 specimens for 11109, 6 specimens for 11110, and 1 specimen for each of the last two. Catalogue dates in ZISP for all these are 26.IV.96, presumably new style, while Berg (1949) gives new style dates 24.IV.1896 for the first and 18.IV.1896 for the last (and this last is 26.IV.1896 in the catalogue). Only ZISP 11108 specimens are regarded as syntypes by Berg (1949). Berg (1949) also points out the confusion over the date when Zarudnyi, the collector, was at "Kuss" (= Khusf at 32°46'N, 58°53'E) given by Nikol'skii as 6.IV.96 old style but on this date Zarudnyi was at "Kiaz-khak" near Asadabad (35°38'N, 59°21'E) south of Mashhad and only reached Khusf on 8 (or 20 new style).VI.96. This is not particularly critical in this instance but serves to point out the difficulties of reconciling literature, field notes, catalogues, and jar labels.

Capoeta nudiventris Nikol'skii, 1897 is a synonym. The syntypes are in the Zoological Institute, St. Petersburg (ZISP 11106) according to Berg (1949) and comprise 3 fish 92.4-121.5 mm standard length. Berg (1949) gives the type locality as "Zeride near Bajistan in southeast Khorasan, 30.IV.1896" (the date in the jar is 26.IV.1896). Nikol'skii (1897) lists 3 collections all from "Saride in Persia orientali. 18.IV.96." with numbers 11105, 11105 (presumably an error for 11106), and 11107 and 6 (actually 7 in the jar and according to Berg (1949)), 3, and 5 specimens respectively. Berg (1949) lists the 5 specimens under 11107 as from "Chakhak in the Al'kor region between Bajistan and Birjand. 9.V.1896", presumably at 33°17'N, 58°54'E. These 5 fish are 37.0-55.2 mm standard length, collected on 25.IV.1896 in the ZISP catalogue and not listed as types in the jar, nor in the catalogue, nor in Berg (1949). The 7 fish in ZISP 11105 measure 46.8-75.3 mm standard length, are from the same locality listed under ZISP 11106 in Berg (1949) and are listed as types in the ZISP catalogue, though not in Berg (1949). Judging from the labels and catalogue sheets, the types are probably from Sarideh at 34°22'N, 58°14'E and comprise 11105 and 11106.

Rainboth (1981) places both fusca and nudiventris in the genus Schizocypris on the basis of the enlarged scales around the vent and anal fin base, a condition reported on by Berg (1949) also but not considered by this latter author to warrant inclusion of these fish in Schizocypris.

Key characters

The strong mode of 7 branched dorsal fin rays, distribution, and the relatively low scale count aid in identifying this species.

Morphology

Dorsal fin with 3 unbranched rays and 7-8, strong mode at 7, branched rays, anal fin with 3 unbranched and 5 branched rays, pectoral fin branched rays 14-20 and pelvic fin branched rays 7-9. Lateral line scales 42-62, mostly 47-56. Scales are found regularly arranged over the whole body and are enlarged around the anus and anal fin base. There is a pelvic axillary scale. Scales are oval and have a subcentral, markedly anterior focus, numerous radii on all fields and moderate numbers of circuli. Gill rakers 11-20, short and touching the raker below when appressed. The mouth is horseshoe-shaped. The last unbranched dorsal fin ray is weak with only a few fine denticles along the basal half. The pharyngeal teeth are very spatulate up to the tip but are thick. There is an occasional trace of a fifth tooth in the major row but all the fish examined had only 4 strongly developed main row teeth. The gut is very elongate with several anterior and posterior loops.

Some populations or individuals may show a very light belly extending up onto the lower flank rendering scales hard to see. Capoeta nudiventris was apparently founded on specimens like this. Some scales low on the flank are incompletely imbricate and deeply embedded in the skin. Berg (1949) in examining the types of fusca and nudiventris found the extent of the scales ventrally to be the same and nudiventris is not naked on the lower flank and belly.

Meristics for Iranian specimens:- dorsal fin branched rays 7(77); anal fin branched rays 5(77); pectoral fin branched rays 14(1), 15(1), 16(8), 17(23), 18(26), 19(13) or 20(5); pelvic fin branched rays 7(8), 8(64) or 9(5); lateral line scales 46(4), 47(6), 48(8), 49(10), 50(10), 51(9), 52(9), 53(9), 54(9), 55(2) or 56(1); total gill rakers 13(1), 14(11), 15(25), 16(26), 17(11), 18(1), or 20(1); pharyngeal teeth 2,3,4-4,3,2(20); and total vertebrae 40(9), 41(42), 42(20) or 43(4).

Sexual dimorphism

Unknown but males presumably bear large tubercles in the breeding season.

Colour

The back and flanks are dark while below the lateral line the body can be very light. The dorsal, anal and caudal fin membranes are dark. Young fish may have a mid-lateral stripe as wide as the eye ending in an indistinct dark blotch on the caudal fin base. The peritoneum is dark brown to black.

Size

Reaches 21.1 cm total length.

Distribution

This species is found in eastern Iran in the Tedzhen River (including Kashaf River), Dasht-e Kavir, Bejestan, Dasht-e Lut and Sistan basins in rivers, springs and qanats, some of the latter not easily located on maps (Nikol'skii, 1899; Berg, 1949; Abdoli, 2000). A record from the "Schalman Rud" presumably in the Caspian Sea basin is most probably an error (Wossughi, 1978).

--- Zirkhuch may be Zir-e Kuh at 32°48'N, 59°50'E ? check this and if reasonable search for Zir and add in all text - Zirkhuch is in eastern or southeastern Khorasan for sure

Zoogeography

Saadati (1977) considers that this species entered eastern Iran from the west via the Namak Lake basin. See also above under genus.

Habitat

Karaman (1969) considers that this species shows the greatest adaptation among Capoeta species to desert life: an elongate and low body, scaleless belly in many individuals, weak spiny dorsal fin ray, reduced number of dorsal fin rays, short dorsal fin which can easily lie flat against the body, and the mouth structure.

Age and growth

Unknown.

Food

Gut contents of the few fish examined contained fragments of large plants including large seeds, filamentous algae and sand grains.

Reproduction

Fish caught in April and May have mature eggs along with some immature eggs, indicating that spawning may occur in stages. Fish caught in November have small but obvious and developing eggs.

Parasites and predators

None reported from Iran except for black spots on the head and fins (syntypes of nudiventris as noted by Nikol'skii (1897)) which are probably encysted larvae of trematodes (Berg, 1949).

Economic importance

This species has been studied in aquaria for the toxicity of lead acetate (Omidi et al., 2009). Toxicity decreased with increase in water hardness, qanat water with a high water hardness (310 mg L-1) showing low toxicity.

Conservation

A widely distributed species apparently able to survive in a wide range of desert habitats, it may not be in need of conservation.

Further work

Biology in mostly unknown and would help confirm the impression that it is not in need of conservation.

Sources

Type material: See above, Capoeta fusca (ZISP 11108) and Capoeta nudiventris (ZISP 11105, 11106).

Iranian material: CMNFI 2007-0005, 7, 27.8-84.2 mm standard length, Semnan, spring at Nardin (ca. 37º03'N, ca. 55º47'E); check ID? CMNFI 2007-0015, 8, 60.1-85.6 mm standard length, Khorasan, qanat at Khalaj (ca. 34º54'N, ca. 58º52'E; CMNFI 2007-0016, 8, 85.5-171.4 mm standard length, Khorasan, qanat and jube at Bidokht (ca. 34º21'N, ca. 58º46'E); CMNFI 2007-0017, ?, ? mm standard length, Khorasan, qanat at Dasht-e Bayaz (ca. 34º02'N, ca. 58º47'E); CMNFI 2007-0018, 15, 21.7-92.4 mm standard length, Khorasan, Shur River (ca. 33º52'N, ca. 59º41'E); CMNFI 2007-0019, 9, 32.7-141.3 mm standard length, Khorasan, qanat between Esfideh and Abbasabad (ca. 33º29-39'N, ca. 59º38-46'E); CMNFI 2007-0020, 23, 43.7-115.1 mm standard length, Khorasan, qanats at Marak and Rabi'an (ca. 32º55-58'N, ca. 59º26-27'E); CMNFI 2007-0021, 16, 24.8-56.3 mm standard length, Khorasan, Shah Abbas qanat in Asadabad (32º55'N, 60º01'E); CMNFI 2007-0022, 6, 56.7-112.1 mm standard length, Khorasan, qanat pool at Mud-e Dahanab (32º43'N, 59º31'E); CMNFI 2007-0023, 6, 82.5-113.1 mm standard length, Khorasan, qanat at Sarbisheh (32º34'N, 59º48'E); BM(NH) 1958.11.7:1-6, 6, 26.1-90.9 mm standard length, Khorasan, near Jajarm (no other locality data).

Capoeta trutta
(Heckel, 1843)

Common names

tu'ini (and variant spellings in transliteration such as touyeni, tuyeni, tuini or too'ini) in Khuzestan (meaning unknown); tu'ini gelkhorak in Khuzestan (see C. damascina for meaning); shir mahi (= milk fish), barg bidy or barg-e bidi (= willow leaf, perhaps from shape and colour), berzem.

[twena, hemira, tela morqat, ethra at Mosul (Heckel (1843b), or takal handscherli (takal = soft or flexible presumably from its small scales, handscherli = armed with a dagger or knife from the dorsal fin spine) at Aleppo (Heckel, 1843b), all in Arabic; trout barb].

Systematics

Rainboth (1981) places this species in Schizocypris on the basis of enlarged scales forming a split to encompass the urogenital region and a bare to partially bare mid-dorsal strip anterior to the dorsal fin. However the schizothoracine fishes are quite different (see accounts for Schizothorax, Schizopygopsis and Schizocypris) and this placement is not accepted here.

The type localities of Capoeta Trutta as given by Heckel (1843b) are "Gewässern bei Aleppo" and the "Tigris bei Mossul". The syntypes are in the Naturhistorisches Museum Wien according to Krupp (1985c) as follows: NMW 55935-37, 55942, 6 specimens 94-274 mm standard length from Mosul, NMW 55926, 55928, 55940-41, 7, 68-192 mm standard length from Aleppo, and in the Senckenberg Museum Frankfurt (SMF 2567 (formerly NMW), 1, 407 mm standard length, from Mosul and SMF 923 (formerly NMW), 1, 175 mm standard length, from Aleppo. Four other syntypes are under NMW 55939, 1 other syntype under NMW 55938 and a dried syntype under NMW 58875. Eschmeyer et al. (1996) lists similar material with the numbers of fish under each catalogue number detailed thus: NMW 55926 (1), NMW 55928 (2), NMW 55935-37 (2, 2, 1), NMW 55939-42 (4, 1, 3, 1), possibly 1 fish in the Rijksmuseum van Natuurlijke Historie, Leiden (RMNH 3164, formerly NMW), 1 syntype in the Senckenberg Museum Frankfurt (SMF 923, formerly NMW) and 1 syntype SMF 2567 (formerly NMW), and 1 dried syntype from the Museum für Naturkunde, Universität Humboldt, Berlin (ZMB 8789; not located in February 2006). The catalogue in Vienna lists only 5 specimens although the card index in 1997 lists NMW fish as syntypes in agreement with Eschmeyer et al. (1996).

Key characters

The combination of small scales, transverse mouth, dorsal and anal fin branched ray counts, the very strong last unbranched dorsal fin ray (longer than head length - usually strong but rarely weak), and the colour pattern identifies this species.

Morphology

Dorsal fin with 3-5 unbranched rays followed by 7-9, usually 8, branched rays, anal fin with 2-3 unbranched rays followed by 5 branched rays, pectoral fin branched rays 14-18, and branched pelvic fin rays 5. Hanel et al. (1992) found 23-31 denticles or teeth on the serrated dorsal fin ray, the largest near the centre of the ray length. Scales in lateral line 68-90, scales above lateral line 15-18 and scales below lateral line 10-17. The back anterior to the dorsal fin is compressed and lacks scales except near the occiput. Scales have a protruding anterior margin but are otherwise rounded, anterior and posterior radii, fine circuli and a subcentral anterior focus. There is a pelvic axillary scale. Gill rakers 23-33, on the lower arm 18-25 (with lowest counts in smallest fish). The rakers reach the second raker below when appressed. Pharyngeal teeth 2,3,4-4,3,2. Teeth are broadly spoon-shaped at the tip, with narrow cusps and stems such that they are quite fragile. A frequency distribution of counts was not taken because of this fragility. Total vertebrae 43-46. The mouth is inferior and transverse with a strong horny cover to the lower jaw. The gut is very elongate with numerous anterior and posterior loops. The karyotype of fish from the Tigris River of Turkey is 2n=150, possibly hexaploid, with 35 meta-submetacentric chromosomes, 40 pairs of subtelo-acrocentric chromosomes with NF=220 (Kılıç Demirok and Ünlü, 2001).

Meristics for Iranian specimens:- branched dorsal fin rays 8(34); branched anal fin rays 5(34); branched pectoral fin rays 14(1), 15(8), 16(18), 17(6) or 18(1); branched pelvic fin rays 5(34); lateral line scales 68(2), 69(1), 70(1), 71(4), 72(5), 73(3), 74(5), 75(2), 76(3), 78(3), 79(1), 80(1), 81(1), 83(1) or 84(1); total gill rakers 22(1), 24(5), 25(4), 26(3), 27(7), 28(8), 29(3), 30(2) or 31(1); and total vertebrae ?more 43(1), 44(6), 45(3) or 46(2) - NMC 79-269, 367, 384, 269, 268, Behnke 231 done.

Sexual dimorphism

Males bear a single tubercle on each flank scale, sometimes 2 tubercles, positioned about the middle of the exposed scale or nearer the posterior edge. The head has small and widely scattered tubercles on the top and sides and large tubercles around the snout from eye to eye below the nostril level. Large tubercles occur in single files on the anal and dorsal fin rays, particularly the posterior rays, becoming apparent on the more anterior rays as tuberculation develops more highly.

Colour

The head and body and the dorsal fin (and sometimes the caudal fin) are covered with small, distinctive black spots, often c- or x-shaped. Spots are apparent through the silver flank colour. Some fish in Khuzestan lack spots but transitional specimens from fully spotted through weakly spotted to immaculate are found. Colour is brownish to yellowish or olive-green on the back with silvery-white flanks and the belly lighter, white with silvery tints. Some fish are very pale almost whitish. Upper flank scales in particular are outlined with dark pigment. The eye is orange above or mostly silvery. Lower fins are orange to yellow at the base and blackish distally, or may be orange to yellow overall. The dorsal and caudal fins are grey or hyaline. The lower rays of the caudal fin have a slight orange-yellow tint. The peritoneum is dark brown to black.

Size

Attains at least 45.8 cm total length. Heckel (1843b) gives 1 Schuh 8 Zoll, or 52.7 cm.

Distribution

Found in the Quwayq, Orontes and Tigris-Euphrates basins including the Iranian portion of the latter (Berg, 1949; Marammazi, 1995) and the Gulf basin in the Zohreh River.

upper Mand ? to check on maps

Zoogeography

Its relationships with other Capoeta species is generally unclear, as are the larger zoogeographical relationships of this and other Capoeta species. See also above under genus.

Habitat

Marammazi (1994) considers this species to be stenohaline but nonetheless more widely distributed than stenohaline Barbus (= Mesopotamichthys) sharpeyi in the Zohreh River which drains to the northern Persian Gulf.

Age and growth

The majority of the population studied by Ünlü (1991) in the Tigris River in Turkey are in age groups 2 and 3 although males live to age 7 and females age 10. Females are usually longer and heavier than males of the same age. Males comprise 41.26% and females 58.74% of this population. In a stream in the Euphrates River drainage of Turkey, Gul et al. (1996) found fish to live for 8 years with 60-90% of the fish in age groups 1 to 3. Females comprised 53.3% and males 46.7% of the population. Kalkan (2008) studied a population in the Karakaya Dam lake on the Turkish Euphrates River. Maximum age was 7 years, age groups 4 and 6 were mostly females whereas age group 3 was mostly male, age-length, age-weight and length-weight formulae were given, and the average growth condition factor was 1.30 for females and 1.28 for males.

Food

Gut contents include diatoms, green algae and large amounts of sand.

Reproduction

Spawning in both the Tigris and Euphrates rivers in Turkey took place in May-June. Males mature at age 2 and females at age 3 in both rivers. Ripe egg size in the Tigris varied between 1.33 and 2.11 mm and egg numbers between 4713 and 18240. Ripe eggs in the Euphrates attained 1.04 mm and the maximum number of eggs per gramme of gonads was 666. Fish from Khuzestan had well-developed eggs on 30 January while adult fish taken on 7 July were not in reproductive condition.

Parasites and predators

Molnár and Jalali (1992) report the monogenean Dactylogyrus pulcher from this species in the Dez River of Khuzestan. Gussev et al. (1993a) describe a new species of monogenean from this species in the Dez River, Dactylogyrus microcirrus. Baska and Masoumian (1996) describe two new species of Myxosporea from fish caught in the Karun River at Ahvaz, Myxobolus molnari taken from the gills and Myxobolus mokhayeri taken from between the soft rays of the fins. The latter species is named after Dr. Baba Mokhayer, an internationally renowned Iranian professor. The new species are of minor pathological importance as the infections are of low intensity and prevalence. Masoumian and Pazooki (1999) list Myxobolus molnari and M. mokhayeri from this species from localities in Khuzestan.

Economic importance

Duman and Duman (1996) give the nutritional value of Capoeta trutta from Keban Dam Lake in Turkey but this fish is little used in Iran.

Conservation

This species does not appear in need of conservation but its biology is too poorly known in Iran to be certain. Kalkan (2008) recommended prohibition of fishing in Turkey during March-August and fish under 22.62 cm should not be retained.

Further work

The biology of this species and its relationships to other Capoeta species needs work.

Sources

Type material: ?

Iranian material:- CMNFI 1979-0020, ?, ? mm standard length, (); CMNFI 1979-0268, 3, 115.7-141.2 mm standard length, Lorestan, between Nowqan and Khorramabad (no other locality data); CMNFI 1979-0269, 2, 114.1-144.1 mm standard length, Lorestan, between Nowqan and Khorramabad (no other locality data); CMNFI 1979-0367, 2, 29.7-54.1 mm standard length, Khuzestan, Meymeh River 11 km north of Dehloran (32º44'30"N, 47º09'30"E); CMNFI 1979-0368, 8, 36.3-67.9 mm standard length Khuzestan, Karkheh River (32º24'30"N, 48º09'E); CMNFI 1979-0376, 1, 55.2 mm standard length, Khuzestan, river tributary to Karkheh River (32º48'30"N, 48º04'30"E); CMNFI 1979-0384, 1, 218.4 mm standard length, Khuzestan, Ab-e Shur drainage (32º00'N, 49º07'E); CMNFI 1991-0153, 2, 153.8-217.2 mm standard length, Khuzestan, Zohreh River (no other locality data); CMNFI 1995-0020, ?, ? mm standard length, (); CMNFI 1995-0021, ?, ? mm standard length, (); CMNFI 1995-0030, ?, ? mm standard length, (); CMNFI 2007-0100, 1, 136.7 mm standard length, Azarbayjan-e Gharbi, Kalwi Chay near Piranshahr (ca. 36º44'N, ca. 45º10'E); CMNFI 2007-0109, 11, 61.3-167.4 mm standard length, Kordestan, Qeshlaq River basin north of Sanandaj (ca. 35º33'N, ca. 47º08'E); CMNFI 2007-0110, 3, 96.6-160.3 mm standard length, Kordestan, Yuzidar River basin (ca. 35º05'N, ca. 46º56'E); CMNFI 2007-0113, 1, 74.8 mm standard length, Kermanshahan, Razavar River 35 km northwest of Kermanahah (ca. 34º25'N, ca. 47º01'E); CMNFI 2007-0116, 1, 95.9 mm standard length, Kermanshahan, Gav Masiab west of Sahneh (ca. 34º28'N, ca. 47º36'E); CMNFI 2007-0117, 2, 153.8-217.2 mm standard length, Kermanshahan, Gav Masiab near Sahneh (ca. 34º24'N, ca. 47º40'E); ZMH 2511, 1, 319.0 mm standard length, Kermanshahan, Karasu-Gamasiab-Seymarreh (no other locality data).

Comparative material: BM(NH) 1931.12.21:8, 1, 113.5 mm standard length, Iraq, Mosul (36º20'N, 43º08'E); BM(NH) 1968.12.13:376-390, 15, 35.6-123.3 mm standard length, Syria, Euphrates River at Mayadine (?); BM(NH) 1974.2.22:1374-1377, 4, 66.3-91.2 mm standard length, Iraq, Baghdad (33º21'N, 44º25'E); BM(NH) 1974.2.22:1382, 1, 86.1 mm standard length, Iraq, Baghdad (33º21'N, 44º25'E); BM(NH) 1974.2.22:1388-1389, 2, 259.4-273.3 mm standard length, Iraq, Tigris River at Samarra (?).

Genus Capoetobrama
Berg, 1916

Capoetobrama kuschakewitschi
(Kessler, 1872)

This species is reported from the Karakum Canal of Turkmenistan (Shakirova and Sukhanova, 1994; Sal'nikov, 1995) and may eventually be found in the Tedzhen River and Caspian Sea basins of Iran. No Iranian record.

Genus Carasobarbus
Karaman, 1971

?

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

 

Carasobarbus luteus
(Heckel, 1843)

Common names

حمري (= hemri), himri; sangal or zangol (= blackish, used at Kermanshah, J. Valiallahi, pers. comm., 2001); lab matiki (= from lipstick by professional fishermen at Kermanshah in reference to red lips, from J. Valiallahi, pers. comm., 2001).

[himri, hamria, hamra, binni hamour, binni hamri, bunni himri, binni, binni shifatha, beni asphar (= yellow son), beni abjad (= white son), beni hamra (= red or yellow son), zuri or bartema, all in Arabic; golden barb, yellow barbel].

Systematics

Heckel (1843b) gives localities for the types of Systomus luteus as "Orontes", and "Tigris", and in the next sentence at "Aleppo" and "Mossul". Two syntypes were examined in the Naturhistorisches Museum Wien under NMW 54250 (but see below). Krupp (1985c) records a 301 mm standard length syntype from Aleppo formerly in the Naturhistorisches Museum Wien, now in the Senckenberg Museum Frankfurt as SMF 6784. Eschmeyer et al. (1996) list the following syntypes: NMW 10827 (1 fish), NMW 54247 (2), NMW 54248 (1), NMW 54249 (1), NMW 54253 (2), NMW 54254 (3), NMW 54255 (2), NMW 54520 (2), NMW 80043 (2) and possibly 2 syntypes in the Rijksmuseum van Natuurlijke Historie, Leiden (RMNH 2463, formerly NMW) as well as the syntype in Frankfurt. The catalogue in Vienna seems to list 5 specimens but this part of the catalogue is overwritten and difficult to interpret. The card index in 1997 lists NMW 53680a (1 fish, the lectotype), 53674 (1), 53675 (1) and 53676 (1) as the syntype series.

Systomus albus Heckel, 1843 from the "Tigris" and "Orontes" and Systomus albus var. alpina Heckel, 1849 are synonyms.

Systomus albus var. alpina was described from the "Flusse Kara-Agatsch und den Alpenseen Pire-San und Deria Kaserun" (= Qarah Aqaj River and Lake Famur, Fars; Pire-San being Parishan and Deria Kaserun being Lake Kazerun, both other names for Lake Famur) (Heckel, 1846-1849b). Krupp (1985c) records 4 syntypes of alpina from Shiraz (sic), Th. Kotschy as NMW 53679 (2 fish) and NMW 53681 (2). NMW 53678 (5 fish, 27.6-60.8 mm standard length), NMW 53679 (2 fish, 63.8-70.5 mm standard length), and NMW 53681 (2 fish, 79.6-93.3 mm standard length) are from the "Kara Agatsch bei Schiraz"; and NMW 53682 (2 fish, 201.7-203.7 mm standard length) are from the "Alpenseen Pire-san und Deria Kaserun": all are possibly syntypes of Systomus albus var. alpina although the catalogue in Vienna lists 5 fish under this name in one column and 4 fish in smaller writing in the adjacent column. The card index in 1997 lists syntypes under NMW 53678 (5), 53679 (2), 53681 (2) and 53682 (2, one of which is the lectotype). Eschmeyer et al. (1996) list 2 fish in the Rijksmuseum van Natuurlijke Historie, Leiden (RMNH 2464) as possible former NMW types of this taxon.

A dried specimen of Systomus albus from Mosul collected by Th. Kotschy may be a syntype (NMW 59485). Eschmeyer et al. (1996) gives the syntypes of this species as NMW 53674 (1), NMW 53675 (1), NMW 53676 (1), NMW 53677 (1), NMW 53680 (1), NMW 91400 (1, dry) and SMF 812 (1), formerly NMW. Krupp (1985c) records the syntype of albus in the Senckenberg Museum Frankfurt under SMF 812 as being 84 mm standard length. The Vienna catalogue lists 4 fish under Systomus albus but the card index in 1997 lists the same NMW fish as Eschmeyer et al. (1996) as above with NMW 53680 as lectotype.

Barbus parieschanica Wossughi, Khoshzahmat and Etemadfar, 1982 is presumably also from Lake Famur or Parishan judging by the name and is a synonym (note that the species name is first spelt parschanica on page 23 in the abstract in Farsi and on page 44 in the English abstract but in the text species description (page 34) and in the table (page 37) it appears as parieschanica, and this is presumably the intended correct spelling). The species locality in the text is "Noorabad of Mamasany". ?

Saadati (1977) refers to a new and undescribed Cyprinion species from Lar in southern Iran but the fish are Carasobarbus luteus.

Günther (1874) placed this species in Barynotus Günther, 1868, a genus with the type species from West Africa. Barynotus is preoccupied in Coleoptera and was replaced by Barbellion Whitley, 1931 (Eschmeyer, 1990). Most authors place the species in Barbus although Karaman (1971) erected a new genus for it, Carasobarbus; and Krupp (1985c) also synonymises Carasobarbus with Barbus. Bănărescu (1997) and Ekmekçi and Banarescu (1998) recognise Carasobarbus as a valid genus however. A group of related species share characters with this species (see also under Kosswigobarbus kosswigi). Carasobarbus may be the generic names for certain members of the group.

Key characters

This species is characterised by a low scale count, smooth last unbranched dorsal fin ray, one or two pairs of barbels, and 10 branched dorsal and 6 branched anal fin rays.

Morphology

Dorsal fin with 4 unbranched rays followed by 9-11, usually and modally 10, branched rays. The last unbranched dorsal fin ray is smooth, thickened, sharp-edged and spine-like. Anal fin with 3 unbranched rays followed by 5-7, usually and modally 6, branched rays. Pectoral fin branched rays 13-17 and pelvic fin branched rays 7-9, usually 8. Lateral line scales 23-36. There is a pelvic axillary scale. There are moderate to many anterior field radii and many posterior field radii and occasionally few lateral radii. The focus is central to subcentral anterior, the anterior scale margin is wavy and the exposed part of the scale is coarse. The concealed part of the scale has numerous fine circuli. Total gill rakers 7-14, reaching the adjacent raker when appressed, sometimes forked at the tip and with spinules on the anterior side. Pharyngeal teeth usually 2,3,5-5,3,2, with the anterior 2-3 teeth rounded and heavier than the posterior teeth. Variants may have 2,3,4 or 1,3,5 (Borkenhagen, 2005). Posterior teeth are hooked at the tip and the grinding surface below the tip is irregular with a protuberant knob which may be striated. The gut is elongate with both posterior and anterior loops.

The mouth is terminal to subterminal and lips are weakly developed. There is one pair of short and thin barbels at the corner in most descriptions. Number and frequencies for 130 fish are 2 barbels (47 or 36.2%), 3 barbels with left anterior present (7 or 5.4%), 3 barbels with right anterior present (5 or 3.8%), or 4 barbels (71 or 54.6%). However, this sample is 112 fish or 86.2% from Fars and Hormozgan. Fish from these provinces, at such localities as the lower Mand River and the Sar Khun oasis north of Bandar Abbas consistently have a high frequency of 4 barbels (58.9%), and with 3 barbel counts included 68.8%, than fish from the Tigris River basin. Even the 18 fish from the Tigris River basin had 5 fish with 4 barbels so, at least in the eastern part of this species range, 4-barbelled fish are not rare.

Body form varies with habitat (Ali, 1982a), there being lake and river forms as with many other cyprinid species.

Iranian specimens have the following meristics: dorsal fin branched rays 9(7), 10(102) or 11(7); anal fin branched rays 5(3) or 6(114); pectoral fin branched rays 14(12), 15(44), 16(48) or 17(13); pelvic fin branched rays 7(9), 8(107) or 9(1); lateral line scales 23(2), 24(10), 25(20), 26(22), 27(28), 28(16), 29(14), 30(4) or 31(1); total gill rakers 8(6), 9(24), 10(40), 11(28), 12(12), 13(3) or 14(2); pharyngeal teeth 2,3,5-5,3,2(19), 2,3,4-5,3,2(4) or 2,3,5-4,3,2(2); and total vertebrae 36(8), 37(53), 38(70), 39(25) or 40(1).

Sexual dimorphism

A 12.7 cm specimen from the Mand River has tubercles on the dorsal, anal, caudal, pectoral and pelvic fins, most strongly on the anal fin rays. Fine tubercles cover the top and sides of the head. A 20 cm fish from the same collection lacked tubercles. Another fish from the lower Mand River (128.5 mm standard length) also has fine tubercles on the upper flank scales as well as the head and fin rays. Ali (1982) reports no sexual dimorphism for Iraqi fish.

Colour

The back and upper flank is dark brown, greenish black or grey-green fading to a whitish or silvery belly all overlain by an orange to yellowish tinge. On the upper flank, scale bases are black-brown with a light blue-grey margin. There is a dark stripe along the mid-line of the back and a dark mid-lateral stripe. Fins are greyish to lime-green, reddish-yellow or orange, becoming blackish distally. The pectoral and pelvic fins tend to be more orange than the anal and caudal fins which are more a faint lime-green. The lips are orange. The eye rim is yellow-green. The peritoneum is black. Small fish have a collection of melanophores at the mid-base of the caudal fin forming a spot-like structure.

The fish described by Heckel (1846-1849b) as Systomus albus var. alpina were also painted live and had a lead-grey body, light brown at the head and reddish-white on the belly. Each scale was black-brown at the base and light blue-grey at the margin, particularly on the upper flank. All fins were blackish and the eyes orange-red.

Size

Attains 38 cm calculated maximum length and 501 g (Ahmed, 1982) or 750 g (Borkenhagen, 2005). Heckel (1843b) gives 17 Zoll for Systomus albus (= 44.8 cm).

Distribution

This species is found in the Orontes and Quwayq rivers and the Tigris-Euphrates basin. In Iran, it is found in the Tigris River basin including the Hawr Al Azim marsh, the Gulf basin including the Helleh, Dalaki, Shapur, Mand and Dasht-e Palang rivers and Lake Famur, the Lake Maharlu basin, the Hormuz basin and the Kor River basin (Wossughi, 1978; Bianco and Banarescu, 1982; Gh. Izadpanahi, pers. comm., 1995; M. Rabbaniha, pers. comm., 1995; Abdoli, 2000). The record from the Kor River basin (Abdoli, 2000) needs confirmation with specimens.

Zoogeography

Karaman (1971) considers that the closest relatives of this species were to be found in India and southern Asia.

Habitat

van den Eelaart (1954) reports that this species in Iraq is a resident in still water and the slower sections of rivers and is the main fish in canals. In summer it goes to the deeper basins of marshes and remains in the shade of plants. It tolerates warm water but does not go into open waters. Al-Hassan and Muhsin (1986) record this species from the Khor al Zubair in southern Iraq where annual temperature range is 12-30°C and annual salinity change is 28-47‰. The fish appear unaffected by these conditions while Heteropneustes fossilis is moribund. Mohamed et al. (1993) report Barbus (= Carasobarbus) luteus from 2 km southward of Fao, Iraq in a pure marine habitat (temperature 13-35°C and salinity 30-47‰). The fish were caught in April which is the flood season.

Age and growth

Ahmed et al. (1984) studied the reproductive cycle of this species in the Hawr al Hammar in southern Iraq near Basrah. Maturity is attained at a minimum of 11.2 cm for females and 12.2 cm for males, at age 1+. The largest fish are 26.0 cm and age 6. Barak and Mohamed (1983) also found 6 age groups for fish from the Garma Marshes, Iraq. Ahmed (1982) studied a population in Tharthar Reservoir about 65 km northwest of Baghdad and found 7 age groups. This study has the fastest growth of Iraqi populations. Khalaf et al. (1988) worked on a population in a flooded gravel pit about 50 km north of Baghdad in Iraq and found fish up to age group 7+. Growth is greatest in the first year (67 mm) and averaged only 22.5 mm in the following years. Growth is slow in consequence of high salinity (3-6% (sic)) and poor food resources. Mohamed et al. (1993) report fish up to 7 years of age in a marine setting in Iraq, Epler et al. (1996) up to 5+ years in fresh and salty Iraqi lakes. Biro et al. (1988) found fish up to age group 8+ in the Diyala River, Iraq. Al Hazzaa and Hussein (2007) describe larval development and growth in the laboratory using fish from a Syrian hatchery. Gökçek and Akyurt (2008) found fish up to 9 years of age in the Turkish Orontes River and give growth parameters for this population. Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 34 Iranian fish measuring 3.20-16.80 cm standard length. The a-value was 0.0232 and the b-value 3.036 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases).

Food

Naama and Muhsen (1986) examined feeding periodicities in this species in the Hawr al Hammar, Iraq. Food is mainly detritus, aquatic plants and algae taken throughout the night and day. Barak and Mohamed (1982) studied food habits in the Garma Marshes, near Basrah, Iraq and found this fish to contain principally aquatic plants, the broken and fragmented leaves and stems of Vallisneria in particular. Diatoms and other algae as well as shrimps, chironomid larvae, gastropods and cladocerans are important foods. Invertebrates are about eight times more important in fish smaller than 30 cm than in larger fish. Plant parts are more important, almost twice as much, in larger fish than smaller. Mohamed et al. (1993) report plant remains to be dominant and fish eggs in lesser quantities in a marine setting in Iraq. Epler et al. (1996) found plants to dominate in fish from fresh and salty Iraqi lakes, although not to the same extent as in Barbus (= Mesopotamichthys) sharpeyi where 95.7-100% of the diet was plants. Tendipedids, worms, detritus and fish were also found in B. luteus.

Khoshzahmat et al. (1981) found that this species did not eat molluscs in Lake Perishan (= Famur), near Kazerun in Iran and assume its diet is aquatic plants.

Reproduction

Spawning in the Hawr al Hammar starts in April and after July no fish are found in a partially spent phase. Eggs are yellow to orange in colour and testes white. The eggs attain 1.86 mm in diameter and number up to 38,433 for the oldest fish. Bhatti and Al-Daham (1978) and Al-Daham and Bhatti (1979) report a spawning season of May-July (peak June-July) for a lower Euphrates River, Iraq population, perhaps as a result of cooler temperatures outside the shallow marshes where warmer temperatures cause an earlier development of gonads. Epler et al. (1996) report spawning in June/July in freshwater Iraqi lakes, earlier in a saline lake. Iranian fish have well-developed eggs in May.

Parasites and predators

Bykhovski (1949) reports a new species of monogenetic trematode, Dactylogyrus persis, from this species in the Karkheh River, Iran. Ebrahimzadeh and Nabawi (1975) list species in the nematode genus Philometra, the protozoan genera Myxosoma and Trypanosoma, the trematode genera Dactylogyrus and Gyrodactylus and the nematode species Camallanus lacustris as well as various unidentified cestodes, trematodes, acanthocephalans and hookworms, from this species in the Karun River. Jalali and Molnár (1990a) records two monogenean species, Dactylogyrus spp., from this species in the Dez River. Molnár and Jalali (1992) describe a new species of monogenean, Dogielius persicus, from this species in the Dez and Karun rivers of Khuzestan. Gussev et al. (1993b) describe a new species, Dactylogyrus carassobarbi, from this species in the Dez River, Khuzestan, the specific name being founded on a misspelling of the genus name Carasobarbus. Masoumian et al. (1994) describe a new species of Myxosporea from the gills of this species in the Karun River, Khuzestan, namely Myxobolus persicus, and later (Masoumian et al., 1996) another new species of Myxosporea, Myxobolus nodulointestinalis, in the gut lining of this species and also from rivers of southwestern Iran. Molnár et al. (1996) report additional new species from this fish in Khuzestan, namely Myxobolus iranicus in the spleen and Myxobolus mesopotamiae in connective tissue of the caudal and pectoral fins. Molnár and Pazooki (1995) record philometrid nematodes from this species in the Karun River, and these are presumed to be a new species.

Masoumian and Pazooki (1999) list Myxobolus persicus, M. karuni, M. sharpeyi, M. nodulointestinalis, M. mesopotamiae and M. iranicus from this species in various localities in Khuzestan. Jalali et al. (2005) summarise the occurrence of Gyrodactylus species in Iran and record G. sp. from Dez River fish. Farahnak et al. (2002) record Anisakis sp. from this fish in Khuzestan Province.

González-Solís et al. (1997) report Proleptinae larvae (Nematoda) from this species in the drainage of Lake Maharlu, Fars. The definitive host is a predatory fish, possibly Mastacembelus mastacembelus, not yet recorded from this basin.

Moghainemi and Abbasi (1992) record a wide range of parasites from this species in the Hawr al-Azim in Khuzestan. Mortazaei et al. (2000) record an infection rate of 1.6% with the worm Bothriocephalus opsariichthydis in Khuzestan marshes.

Economic importance

An important food fish in southern Iraq and Iran (Al-Daham and Bhatti, 1979; Ahmed, 1982). Sharma (1980) reports that hamri were the fourth most important fish species at Basrah fish market, accounting for 267,570 kg from October 1975 to June 1977. Heckel (1846-1849b) reports that they "reach a good size and are very tasty" in Lake Famur, Fars.

In some parts of Southwest Asia this species is regarded as &qut;sacred" kept and bred in special pools where fishing is forbidden (Tortonese, 1934).

The eggs of this species are poisonous (Najafpour and Coad, 2002). A kebab made of about one-quarter of an ovary was eaten. Toxic effects were dizziness, abdominal pain, vomiting, diarrhoea, bitter taste, dryness of mouth, intense thirst, and faintness. One victim as hospitalised for two days and his stomach pumped while a second victim recovered after one day's rest.

Conservation

Vulnerable in Turkey (Fricke et al., 2007). It is a common species in Iranian freshwaters but no detailed conservation assessment has been made.

Further work

The biology of this species in Iran needs study along with its conservation status. Specimens from Fars show differences in body form from those in Khuzestan and this could be investigated.

Sources

Type material: ?

Iranian material: CMNFI 1979-0023, 17, 58.3-161.4 mm standard length, Fars, neighbourhood of Shiraz (no other locality data); CMNFI 1979-0024, 1, 61.5 mm standard length, Fars, neighbourhood of Shiraz (no other locality data); CMNFI 1979-0026, 2, ? mm standard length, Fars, Shapur River (29º47'N, 51º35'E); CMNFI 1979-0047, 1, ? mm standard length, Fars, Ab-e Paravan (ca. 29º34'N, ca. 52º42'E); CMNFI 1979-0076, 1, ? mm standard length, Fars, Barm-e Shur (29º28'N, 52º41'30"E); CMNFI 1979-0087, 1, ? mm standard length, Khuzestan, Karun River at Ahvaz (31º19'N, 48º42'E); CMNFI 1979-0125, 1, ? mm standard length, Bushehr, Dalaki River near Dalaki (ca. 29º28'N, ca. 51º21'E); CMNFI 1979-0129, 26, ? mm standard length, Fars, spring about 2 km from Farrashband (28º54'N, 52º04'E); CMNFI 1979-0135, 19, ? mm standard length, Fars, Mand River tributary (28º08'N, 53º10'E); CMNFI 1979-0154B, 3, 160.7-258.6 mm standard length, Fars, stream channels at Koorsiah (28º45'30"N, 54º24'E); CMNFI 1979-0155, 2, ? mm standard length, Fars, spring at Gavanoo (28º47'N, 54º22'E); CMNFI 1979-0156, 6, ? mm standard length, Fars, qanat at Rashidabad (28º47'N, 54º18'E); CMNFI 1979-0157, 1, ? mm standard length, Fars, qanat at Hadiabad (28º52'N, 54º13'E); CMNFI 1979-0160, 2, ? mm standard length, Fars, spring at Arteshkkadeh Pomp (29º09'N, 53º37'E); CMNFI 1979-0163, 1, 84.9 mm standard length, Fars, neighbourhood of Shiraz (no other locality data); CMNFI 1979-0164, 6, 56.6-91.1 mm standard length, Fars, neighbourhood of Shiraz (no other locality data); CMNFI 1979-0187, 31, ? mm standard length, Hormozgan, stream and pools at Sar Khun oasis (27º23'30"N, 56º26'E); CMNFI 1979-0206, 3, 24.4-25.1 mm standard length, Fars, qanat near Runiz-e Pa'in (29º12'N, 53º40'E); CMNFI 1979-0240, 3, ? mm standard length, Fars, Parishan Lake (ca. 29º31'N, ca. 51º50'E); CMNFI 1979-0304, 5, ? mm standard length, Fars, Parishan Lake (ca. 29º31'N, ca. 51º50'E); CMNFI 1979-0347, 2, ? mm standard length, Fars, Pol-e Berengie (29º27'30"N, 52º32'E); CMNFI 1979-0352, 7, ? mm standard length, Khuzestan, marsh in Jarrahi River drainage (30º33'30"N, 48º48'E); CMNFI 1979-0358, 1, 23.7 mm standard length, Khuzestan, pond southeast of Bostan (31º37'N, 48º07'E); CMNFI 1979-0360, 8, ? mm standard length, Khuzestan, canal branch of Karkheh River (31º40'N, 48º35'E); CMNFI 1979-0364, 6, ? mm standard length, Khuzestan, river at Abdolkhan (31º52'30"N, 48º20'30"E); CMNFI 1979-0371, 7, ? mm standard length, Khuzestan, stream in Karkheh River drainage (32º05'N, 48º19'E); CMNFI 1979-0687, 7, 124.8-154.1 mm standard length, Fars, Shiraz bazar (no other locality data); CMNFI 1979-0789, 4, ? mm standard length, Fars, Lake Parishan (29º31'N, 51º48'E); CMNFI 1991-0154, 1, ? mm standard length, Khuzestan, Hawr al Azim (ca. 31º45'N, ca. 47º55'E); CMNFI 1993-0126, 1, ? mm standard length, Kermanshahan, Sarab-e Yavari (34º28'N, 46º56'E); CMNFI 1993-0127, 1, ? mm standard length, Kermanshahan, Sarab-e Maran (34º44'N, 46º51'E); CMNFI 2007-0060, 2, ? mm standard length, Fars, Chashmeh Ab-e Shirin near Lar (ca. 27º41'N, ca. 54º17'E); CMNFI 2007-0111, 1, ? mm standard length, Kermanshahan, Alvand River near Sar-e Pol-e Zahab (ca. 34º36'N, ca. 45º56'E); ZSM 21861, 5, 172.0-217.2 mm standard length, Khuzestan, Dez River at Harmaleh (31º57'N, 48º34'E).

Comparative material: CMNFI 1987-0017, 3, 97.3-143.9 mm standard length, (); BM(NH) 1934.9.5:6, 1, 117.3 mm standard length, Kurdistan, Ain al Hamra, Shithatha (); BM(NH) 1973.6.21:194, 1, 203.4 mm standard length, Iraq, Shatt al Arab (); BM(NH) 1974.2.22:1338, 1, 134.9 mm standard length, Iraq, Najab Bazar (); BM(NH) 1974.2.22:1346, 1, 108.7 mm standard length, Iraq, Tigris River near Faish Khabour (); BM(NH) 1986.2.14:4-7, 4, 98.6-146.6 mm standard length, Iraq, Baghdad (33º21'N, 44º25'E).

Genus Carassius
Nilsson, 1832

The goldfishes comprise 2-3 species found in Europe, northern Asia and the Far East. Eschmeyer (1990) and Kottelat (1997) comment on the authorship of Carassius. One species is now common in Iran.

These fishes are characterised by a stout and compressed body, last unbranched dorsal and anal fin rays finely serrated, long dorsal and short anal fin, mouth small and terminal, lips thick and fleshy, no barbels, pharyngeal teeth in 1 row and molariform but compressed, numerous gill rakers, and scales large.

Carassius auratus
(Linnaeus, 1758)

Common names

mahi-ye talaee or mahi-ye talai (= gold fish) or اوشين (ooshin or oushein) in Khuzestan; kapur safid by anglers in Khuzestan at Ahvaz; kopur-cheh (= small carp) or كاراس (= karas, karass or karaz) in Mazandaran; kopur cheky (= by the job carp?), kopur chekeh (= drop carp?); ماهي حوض (= mahi-ye howz or mahi-e-hoz, meaning pond or pool fish), mahi-ye howz-e noqrehi (= silvery pond fish, for silvery form), mahi-ye howz-e talaee (= golden pond fish for orange form).

[samak zahabi, buj-buj in Nasiriyah; samti; yayabash in Basrah; karseen in Baghdad; all in Iraqi Arabic; serebryanyi karas or silver crucian carp in Russian; goldfish for auratus, Prussian carp for gibelio].

Systematics

Cyprinus auratus was originally described from China and Japanese rivers.

Pelz (1987) discusses the scientific name of the goldfish and its confusion with Carassius carassius. All diploid goldfish of western Europe are Carassius auratus auratus (from introductions, presumably including releases and escapes in Iran) and all triploid goldfish are C. auratus gibelio from eastern areas. Goldfish do not appear to be native to Iran but Iranian specimens are sometimes referred to Carassius auratus gibelio (Bloch, 1782) known as the Prussian carp, European goldfish or silver crucian carp. Berg (1948-1949) considers the familiar pet "goldfish" to be a domesticated form of the Prussian carp. However these fish probably have a number of origins - from aquarium stock and from China. Kottelat (1997) tentatively recognises Carassius gibelio (Bloch, 1782) as a species native to eastern Central Europe, and Kottelat and Freyhof (2007) map gibelio as the introduced species in the Caspian Sea basin of Iran. Vasil'eva and Vasil'ev (2000) state that fish named in the literature as Carassius auratus gibelio from Europe, Siberia and eastern Asia are triploids and are not a valid subspecies of C. auratus s.s. They consider C. gibelio to be a distinct species as long as it has a unique and ancient origin rather than arising de novo, and as long as the type specimens are triploids. Szczerbowski in Bănărescu and Paepke (2002) recognises C. a. auratus and C. a. gibelio.

Additionally C. auratus may be a tetraploid derivative of Carassius carassius. The native distribution of C. carassius is in Europe and western Asia, reaching northern drainages of the Caspian Sea in the southern limits of its distribution (Libosvárskı, 1962). It differs from C. auratus in having a slightly convex margin to the dorsal fin (straight or slightly concave in C. auratus), caudal fin slightly emarginate (deeply emarginate), usually 6 branched anal rays (always 5), 23-33 gill rakers (37-53), 31-34 vertebrae, usually 32-33 (28-31, usually 29-30), 28-29 fin denticles posteriorly on the dorsal fin spine (10-11), peritoneum light (dark), black spot at the caudal fin base in young and some adults (absent), and a coppery gold body (silvery, pinkish gold, gold or red) (Szczerbowski in Bănărescu and Paepke, 2002). Berg (1948-1949) also cites the characters body rounded, back thick (body angular, back compressed) and scales weakly sculptured (rough), although his comparison is with C. a. gibelio.

Goldfish commonly hybridise with Cyprinus carpio to further confuse the identity of these fishes (L. Nico, http://nas.er.usgs.gov/fishes/accounts/cyprinid/ca_aurat.html, downloaded 24 May 2000). The identity of "goldfish" in Iran has not been thoroughly surveyed and, along with conflicting views on species and widespread introductions from many sources, make it simpler to refer to this taxon as C. auratus for now.

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

Key characters

The combination of spines in both the dorsal and anal fins and the absence of barbels is unique to this species. Szczerbowski in Bănărescu and Paepke (2002) distinguishes the subspecies auratus from gibelio by 21-36 lateral line scales (27-35 in gibelio) and a pink or gold colour (yellowish silver), not very diagnostic. Ilhan et al. (2005) give gill raker numbers of 34-40 for auratus, 42-56 for gibelio and 25-32 for C. carassius in Turkish waters (however note below that counts can increase with growth and see also under C. carassius for somewhat different counts and other distinguishing characters).

Morphology

Dorsal fin with 3-4 unbranched rays followed by 12-20 branched rays, anal fin with 2-4, usually 3, unbranched rays followed by 5-6, usually 5, branched rays, pectoral fin branched rays 11-18, and pelvic fin branched rays 6-9, usually 8. Dorsal and anal fin spine denticles coarse and few (about 10-15).

Lateral line scales 21-36. The anterior scale margin is wavy and there are very few anterior and posterior radii, as few as 3-4. The focus is slightly subcentral posterior. Circuli on the exposed part of the scale are more coarse and widely spaced than on the concealed part of the scale. Gill rakers long with serrated interior margins, reaching the fifth to eighth raker below when appressed with younger fish having longer rakers proportionately. Counts are size dependent in the range 34-54. Total vertebrae 25-34. Pharyngeal teeth 4-4, with very elongate, narrow, flattened and horizontal cusps arising from a much narrower stem. The gut is coiled with several loops. This species is variously reported as only diploid or as a tetraploid (2n=100-104); see above.

There are elongate specimens (morpha humilis, where fish density is high) and deep-bodied specimens (morpha vovki, where fish density is low) but these names have no taxonomic significance.

Meristic values for Iranian specimens are:- dorsal fin branched rays 16(4), 17(3), 18(3), 19(5) or 20(2); anal fin branched rays 5(17); pectoral fin branched rays 11(1), 14(1), 15(4), 16(10) or 17(1); pelvic fin branched rays 7(2) or 8(15); lateral line scales 28(6), 29(9) or 30(2); pharyngeal teeth 4-4(10); and total vertebrae 32(2).

Sexual dimorphism

Breeding males have small nuptial tubercles on the operculum, back and pectoral fin rays.

Colour

The golden or orange colour of artificially bred aquarium goldfish is distinctive. However populations in the wild, if they breed successfully, gradually revert to a wild-type of colour, without the appropriate diet supplement of aquarium fish and, as golden fish, are readily seen and eaten by birds and other fishes. Yanar and Tekelioğlu (1999) found that pigmentation increased with fish weight when specimens were fed the carotenoid zeaxanthin. Wild-type colour is an overall olive-green fading to a white belly. Flanks can be silvery to almost black. Fins are a dark olive-bronze, the membranes in particular being heavily pigmented. Young goldfish are usually green, brown or bronze to almost black and only after about 1 year do they take on the colour of adult auratus or gibelio. Peritoneum dusky to black.

Young fish at Ahvaz, Khuzestan, however, are a bright silvery overall (more so than Cyprinus carpio of similar size), the back is grey, the caudal fin is grey on the proximal half and hyaline distally, and the anal fin rays are white (and thus partly resemble gibelio).

Prussian carp (subspecies gibelio) is a dark steel colour with dark blue or greenish dorsally, silver-grey laterally and white ventrally, dorsal and caudal fins are dark grey and the paired fins and anal fin are light pinkish (Szczerbowski in Bănărescu and Paepke (2002).

Size

Attains 62.0 cm and about 5.0 kg, the subspecies gibelio being smaller, up to 45.0 cm and 1.24 kg.

Distribution

The native distribution is in northern Asia and China, reaching northern drainages of the Caspian Sea in the western limits of its distribution (Libosvárskı, 1962; Plez, 1987). The goldfish has been widely introduced to garden ponds and released from aquaria in temperate to warm waters world-wide. In Iran it has been introduced throughout the Caspian Sea basin where it is reported from the Atrak, Gorgan, Gharasu, Tajan, Babol, Haraz, Sardab, Aras (including the middle Aras and lower reaches of its tributary the Qareh Chai), Tonekabon, Pol-e Rud, and Safid rivers, the Anzali Mordab where it is now the most abundant fish, Gorgan Bay and Alma- and Ala-Gol (Holčík and Oláh, 1992; Shamsi et al., 1997; Roshan Tabari, 1997; Abbasi et al., 1999; Kiabi et al., 1999; Abdoli, 2000; Gasmi and Mirzaei, 2004; Patimar, 2008); the lower Talkheh and lower Zarrineh rivers in the Orumiyeh basin (Abdoli, 2000); the lower Shur, lower Qareh Chai and the Latian Reservoir in the Namak Lake basin (Armantrout, 1980; Hosseini, 1987; Abdoli, 2000); the Hamun Kushk and the Sistan Dam as well as throughout the hamuns in the Sistan basin (Ahmadi and Wossughi, 1988; Mansoori, 1994; J. Holčík, in litt., 1996; field work in the 1970s); Kerman-Na'in and Dasht-e Lut basins generally (Abdoli, 2000); lower Kashaf River in the Tedzhen basin (Abdoli, 2000); throughout Khuzestan where now common (N. Najafpour and M. Al-Mukhtar, pers. comm., 1995; field work 2000, absent in 1970s); middle and lower Hilleh and lower Mand rivers in the Gulf basin; middle Halil and middle to lower Bampur River (Abdoli, 2000); Dalaki and Shapour rivers (Pazira et al., 2005), Safid River, Zayandeh River, Zarivar Lake and the Hamun Lake (Shamsi et al., 2009), and found in garden and park ponds throughout Iran. Some introductions are probably discarded aquarium fish as goldfish are sold as pets and for the Now Ruz (= New Year) festivities. They may also have been introduced accidentally with the commercially important Chinese carps.

This species is also recorded from the Karakum Canal and Kopetdag Reservoir in Turkmenistan (Shakirova and Sukhanova, 1994; Sal'nikov, 1995) and may eventually reach Iranian waters from this source in the Tedzhen (= Hari) River basin.

The Prussian carp (subspecies gibelio) is less widely distributed and its presence and distribution in Iran are not known.

Zoogeography

This species has been introduced to Iran by man. Some are undoubtedly aquaculture pond escapees or aquarium releases. Goldfish are kept in aquaria as part of the Now Ruz (New Year) celebrations in March each year. Tehran television (and the Green Front of Iran, see below) urged people to release them into local waters rather than killing them after the New Year (J. Valiallahi, pers. comm., 2000).

Habitat

Goldfish are hardy and can live in winterkill water bodies with much aquatic vegetation, low oxygen, and high pollution (Gudkov, 1985). They can also survive several hours out of water (Pelz, 1987) and may bury themselves in mud, albeit temporarily when scared (Szczerbowski in Bănărescu and Paepke, 2002). Goldfish appear to favour ponds or pools in streams with aquatic vegetation but are often introduced into small bodies of water as ornamental fish. They are tolerant of turbidity, e.g. clay at 225,000 mg/l, pH from 4.5 to 10.5, very high temperatures (upper lethal limit 41.4°C), and high salinity (17‰). This species was killed under experimental conditions, when gradually acclimated to increasing salinity at 28,200 μmho and, by sudden exposure, at 19,200 μmho (Jassim, 1988). This is a greater tolerance than that shown by Cyprinus carpio, another exotic introduced to Iran. However, Carassius auratus appeared in the Basrah fish market when an increase in the Tigris River discharge reduced the salinity of the Shatt al Arab (N. A. Hussain, in litt., 1994).

In Iran it is one of two most abundant species in Caspian wetland areas along with Gambusia holbrooki (Iranian Fisheries Research and Training Organization Newsletter, 19:4, 1998).

Age and growth

Maturity is attained at 3-4 years in the Volga Delta for goldfish. Life span is 13 years with most growth in the first 2-4 years to a size of 15-20 cm (Gudkov, 1985; Kizina, 1986). Life span in captivity in China may exceed 50 years. Population numbers in confined areas are limited by a chemical released by the goldfish which represses more spawning. Prussian carp live up to 11 years.

In the Anzali Mordab, Holčík and Oláh (1992) found only 6 age groups (as did Bagirova et al. (1990) in reservoirs of Azerbaijan while Pipoyan and Rukhkyan (1998) found 9 age groups in Armenia) with the largest fish 32 cm standard length owing to intense fishing pressure. Growth in mm increments was successively 93, 47, 50, 42, 28, and 37. The population is entirely female (see below). Individual life span is greater in Armenia where males are scarce or absent than in bisexual populations (Pipoyan and Rukhkyan, 1998). Sayad Borani et al. (2001) studied this species (as C. auratus gibelio) in the Anzali Mordab at four localities and found a mean fork length of 19.5 cm (range 2.5-31.5 cm) and a mean weight of 196.8 g. The mean age was 2.6 years. The mean length, weight and age were higher in the Sia-Keshim area of the lagoon. The exploitation rate was 0.47, L was 36.0 cm and K was 0.23 per year. Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 41 Iranian fish measuring 5.65-8.17 cm standard length. The a-value was 0.0419 and the b-value 2.911 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases).

Fish in Buldan Dam Lake, Gediz River basin, Turkey referred to C. gibelio had a maximum age of 6 years and attained 25.5 cm and 269.1 g (Sarı et al., 2008). von Bertalanffy growth parameters were L = 31.66 cm, W = 635.91 g, k = 0.146 year-1 and t0 = -2.166 year. Ratios of total, natural and fishing mortality were calculated as 0.632 year-1, 0.456 year-1 and 0.176 year-1.

Food

Food is predominately zooplankton but also includes aquatic insects, crustaceans, molluscs, worms, detritus, filamentous algae, macrophytes and young fish, switching from one kind of food to another as circumstances warrant. Goldfish have a palatal organ on the roof of the mouth used to taste and touch food and their dense gill rakers aids in feeding on smaller food items. In the recovering Hawr al Hammar, Iraq, diet is 46.1% algae and 25.5% diatoms, with amounts of plants, crustaceans, insects, snails and fish being less than 10% each, in the Hawr al Hawizah 36.3% algae, 21.3% diatoms and 17.5% copepods, with amounts of plants, cladocerans, ostracods and insects being less than 10% each, in the Al Kaba'ish (= Chabaish) Marsh 45.5% algae, 25.2% diatoms, with plants, various crustaceans, insects and snails at less than 10% each (Hussain et al., 2006).

Reproduction

The fish in the Anzali Mordab are all female, reproducing through gynogenesis. Sayad Borani et al. (2001) found fish in Anzali Mordab to have a sex ratio of 99.3 females:0.7 males. Egg development is stimulated by sperm probably from Cyprinus carpio, Tinca tinca, Blicca bjoerkna or Scardinius erythrophthalmus. Here fish may mature at 1 year of age, and coupled with polycyclic ripening of eggs and intermittent spawning, this has led to the dominance of this species in the fresh waters of the lagoon (Holčík and Oláh, 1992). In Armenia, maturity appears to be linked with average annual temperature - at 12.0-13.1°C it occurs at the end of the first year of life while at 8.4-9.0°C it occurs at the end of third and fourth years (Pipoyan and Rukhkyan, 1998). Turkish populations in Topçam Dam Lake, Aydın (Şaşı, 2008) and Buldan Dam Lake, Gediz River basin (Sarı et al., 2008) referred to C. gibelio were 98.84% and 99.44% female. Spawning in the former locality was from March to August, suggesting multiple spawnings with mean fecundity ranging from 37,823 in August to 85,159 in March. Egg diameter reached 1.099 mm in June.

Spawning begins in late April to mid-May in the Volga Delta and occurs in May-June in the Anzali Mordab (Sayad Borani et al., 2001). Eggs are laid in 2-5 batches over a spawning period extending into July. Up to 10 batches are laid elsewhere at 8-10 day intervals with up to 4000 greenish-yellow eggs in each batch. Fecundity reaches 253,200 eggs (elsewhere to 685,700 with absolute fecundity reaching 860,000 eggs). The largest eggs are 1.6 mm in diameter (Gudkov, 1985; Kizina, 1986; Szczerbowski in Bănărescu and Paepke, 2002). Each female is accompanied by 2 or more males and chases are reported with splashing and shooting through the water near the surface. The eggs are adhesive and attach to water plants and hatch in 5-8 days.

Parasites and predators

Mokhayer (1976b) records infectious dropsy and swimbladder inflammation in Iranian goldfish. Saprolegniosis has been reported from goldfish in Iran (Rahbari and Razavilar, 1982). Growths of the fungus Saprolegnia parasitica resembled tufts of cotton wool. Mokhayer (1989) reports metacercariae of the eye fluke, Diplostomum spathaceum from this species in Iran, which can cause complete blindness and death in commercially important species. Jalali and Molnár (1990a) record the monogeneans Dactylogyrus baueri, D. extensus, D. formosus and D. vastator from this species in the Safid Rud. Jalali and Molnár (1990b) report a variety of monogeneans from this species variously in fish farms throughout Iran, namely Dactylogyrus baueri, D. dulkeiti, D. formosus, D. vastator and D. vastator forma minor. Molnár and Jalali (1992) record the monogenean Dactylogyrus intermedius from this species in a petfish farm near Tehran. Gussev et al. (1993a) describe a new species of monogenean from goldfish on a fish farm near Tehran, Dactylogyrus intermedioides. Shamsi et al. (1997) report Clinostomum complanatum, a parasite causing laryngo-pharyngitis in humans, from this species. The helminth Anisakis sp. is recorded from the guts of this species in the Anzali wetland (Ataee and Eslami, 1999; www.mondialvet99.com, downloaded 31 May 2000). Mousavi (2003) records the monogeneans Gyrodactylus sp., G. kabayashi, D. extensus, D. baueri, Trichodina sp., the ciliates Ichthyophthirius multifilis and Ichthyoboda sp. and the copepods Lernaea cyprinacea and Argulus foliaceus from this species in ornamental fish in Iran. Aquarium specimens are often released in the wild at New Year (Now Ruz). Naem et al. (2002) found the following parasites on the gills of this species from the western branch of the Safid River, namely the protozoans Ichthyophthirius multifilis and a Trichodina species, monogenean trematodes Dactylogyrus anchoratum, and Gyrodactylus sp.. Jalali et al. (2002) and Jalali and Barzegar (2006) record Diplostomum spathaceum from this species in Lake Zarivar. Naem (2002) records the monogenean Dactylogyrus anchoratus from fish in Safid River. Mehdipoor et al. (2004) record the monogenean Dactylogyrus baueri in this fish in the Zayandeh River. Masoumian et al. (2005) recorded the protozoan parasite Ichthyophthirius multifilis from this species in the Aras Dam in West Azarbayjan (species identified as C. carassius, presumably goldfish). Jalali et al. (2005) summarise the occurrence of Gyrodactylus species in Iran and record G. kobayashii and G. sp. in fish from the Safid River. Khara et al. (2006a) record the eye fluke Diplostomum spathaceum for this fish in the Amirkalayeh Wetland in Gilan. Sattari et al. (2004, 2005) surveyed this species (as C. carassius) in the Anzali wetland, recording Raphidascaris acus (and larvae) Eustrongyloides excisus and Camallanus lacustris. Pazooki et al. (2007) recorded various parasites from localities in West Azarbayjan Province, and found Eustrongylides excisus. Sattari et al. (2007) record the nematode Raphidascaris acus, the digenean Diplostomum spathaceum and the monogeneans Dactylogyrus extensus,and Gyrodactylus sp. in this species in the Anzali wetland of the Caspian shore. Shamsi et al. (2009) found Dactylogyrus baueri, D. dulikeity, D. extensus, D. intermedius, D. intermedioides and D. wegeneri in this species from localities such as fish farms, the Safid River, Zayandeh River, Zarivar Lake and the Hamun Lake.

Economic importance

This species is raised on Tehran fish farms for the pet trade (Molnár and Jalali, 1992). Ebrahimzadeh Mousavi and Khosravi (2004) report suspected epizootic ulcerative syndrome from ulcerated goldfish.

In the Anzali Mordab, 62% of the total catch is goldfish, an accidental introduction (Petr, 1987). The catch in the mordab in 1990 was 46,472 kg (Holčík and Oláh, 1992). As the salinity of this lagoon increases, the density of goldfish will decrease. Valeipour and Haghighy (2000) record the catch for 1992-1996 at 40% of the species taken. Safaei (2005) gives a goldfish catch figure of 45% of the 313 ton fishery there in 1992. The presence of goldfish in the Anzali Mordab led to a decline in the native fishery there.

This species is caught by anglers at Ahvaz in Khuzestan using bread or potato as bait.

It is known to control mosquito larvae in Bengal (Chandra et al., 2008).

The peculiar type of reproduction is very successful and affects the catches of other cyprinid species, being equivalent to a predatory effect (Holčík and Oláh, 1992).

There is some evidence that this fish disturbs the habitat of native species, muddying waters, and it may compete for food and space. Goldfish have destroyed some amphibian populations in other parts of the world by consuming frog eggs (Coad and Abdoli, 1993b). The Green Front of Iran recommended the release into pools of mosques, parks or natural lakes of the estimated 20 million goldfish kept in aquaria for the Iranian New Year celebrations in March each year. This would have a deleterious effect on habitats not yet colonised by this exotic species. A news report in 2005 cites the death of 5 million fish in transit from the store to the Iranian home at New Year, indicating perhaps that the numbers that do make it are much higher (www.politicalgateway.com, downloaded 5 August 2005). Newspaper articles suggested that goldfish should only be released into "pools" rather than rivers because of all the attendant dangers of this exotic. They are known to prevent reproduction of native species in Sistan (Iran Daily, 17 March 2005, p. 5).

This species is used in Iran as an experimental organism, e.g. in studying the effects of anionic detergents (shampoos, a common water pollutant) on blood parameters, on hepatic and renal pathology and serum biochemical parameters (Shahsavani et al., 2003; Shahsavani et al., 2005; Shahsavani et al., 2006; Shahsavani and Movassaghi, 2006); the use of phenytoin sodium on skin wounds (5mg/l showed best healing improvement)(Shahsavani et al., 2002); the effects of cortisol on testicular apoptosis (Bahmani et al., 2007).

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in textbooks, in aquaria and in aquaculture, as bait, as an experimental species and because it has been introduced outside its natural range. There are numerous, commercial aquarium forms with particular morphologies and colours that are assigned common names, e.g. common, veiltail, comet, fans, calicoe, black-moor, telescope-veiltail, lionhead, egg-fish, shubunkin. Balon (2006) reviews the origin of the species.

Conservation

This species is a successful exotic, in no need of conservation.

Further work

The Carassius species in Iran is generally regarded as C. auratus, the goldfish of aquaria, as it is used extensively in Now Ruz (New Year) celebrations and often released into natural waters. This needs confirmation for all major populations.

Sources

Iranian material: CMNFI 1979-0230, 41, 14.7-38.6 mm standard length, Sistan, Hamun-e Puzak (ca. 31º15'N, ca. 61º42'E); CMNFI 1991-0162, 1, 40.5 mm standard length, Mazandaran, Bagher Tangeh (36º42'N, 52º43'E); CMNFI 1993-0136, 64.0 mm standard length, uncatalogued material, 1, 93.5 mm standard length, Gilan, near Hendeh Khaleh (ca. 37º23'N, ca. 49º28'E); 1, 52.8 mm standard length, Gilan, near Hendeh Khaleh (ca. 37º23'N, ca. 49º28'E); 4, 16.4-50.3 mm standard length, Gilan, near Khoshk Bijar (ca. 37º22'N, ca. 49º47'E).

Carassius carassius
(Linnaeus, 1758)

The crucian carp has been reported as introduced to Iran in the Karun River basin as aquarium releases by Armantrout (1980) without further details and there are other reports such as in the Gorgan River (Y. Keivany, in litt., 1992) and Mahabad Dam (Abdi, 1999; www.mondialvet99.com, downloaded 31 May 2000) but these may be confusion with Carassius auratus. Specimens are needed to confirm the presence of this species in Iran. The native distribution is in Europe and western Asia, reaching northern drainages of the Caspian Sea in the southern limits of its distribution (Libosvárskı, 1962). It differs from C. auratus in having a slightly convex margin to the dorsal fin (straight or slightly concave in C. auratus), caudal fin slightly emarginate (deeply emarginate), usually 6 branched anal rays (always 5), 23-33 gill rakers (37-53), 31-34 vertebrae, usually 32-33 (28-31, usually 29-30), 28-29 fin denticles posteriorly on the dorsal fin spine (10-11), peritoneum light (dark), black spot at the caudal fin base in young and some adults (absent), and a coppery gold body (silvery, pinkish gold, gold or red) (Szczerbowski in Bănărescu and Paepke, 2002). Berg (1948-1949) also cites the characters body rounded, back thick (body angular, back compressed) and scales weakly sculptured (rough), although his comparison is with C. a. gibelio.

Genus Chondrostoma
Agassiz, 1832

The nases are found from the Iberian Peninsula and France to the Caspian and Tigris-Euphrates basins. There are about 26 species of which 2 are known for Iran (Elvira, 1997). Chondrochylus Heckel, 1843 and Chondrochilus Heckel, 1843 are synonyms. Eschmeyer (1990) gives the year of publication for the genus as 1832 as opposed to other authors who give 1835 (e.g. Berg, 1948-1949; Reshetnikov et al., 1997). Doadrio and Carmona (2004) confirm the monophyly of the genus based on the cytochrome b gene with vicariant events accounting for distribution of taxa better than a dispersalist model. Middle East taxa belong to a single lineage with the more differentiated and basal species in the Caucasus and Mesopotamia, having been isolated in the Upper Miocene-Pliocene.

This genus is characterised by being of moderate size, with a somewhat compressed body, scales of moderate to small size (44-106 in the lateral line (Robalo et al. (2007) give a range of 52-78 for their more restricted genus)), scales squarish with radii in the anterior and posterior fields and a subcentral anterior focus, no barbels, an inferior and transverse or crescentic mouth with a cutting edge to the lower jaw, thin upper lip and no lower lip, pharyngeal teeth knife-like and in 1 row with a high count (5, 6 or 7, the same number on each arch or one more on the left), gill rakers short and moderately numerous (up to 40), short dorsal fin without a thickened ray opposite the pelvic fins, 7-10 dorsal fin branched rays, a moderately elongate anal fin with 8-12 branched rays, deeply forked caudal fin and usually concave dorsal and anal fins, a pelvic axillary process always present, 42-49 vertebrae, a black peritoneum, and a long, coiled gut. Elvira (1997) and Robalo et al. (2007) give osteological characters.

Bogutskaya (1997a) places the nominal Iranian species, C. regium and C. orientalis, in a group characterised by a straight or only slightly arched mouth cleft, high vertebral counts (total vertebrae modes 45-47 and abdominal modes 26-28) and often or commonly 4 unbranched rays in the dorsal fin.

Chondrostoma cyri
Kessler, 1877

Common names

shekamsiah-e Aras.

[Kur altagizi in Azerbaijan; chernobryushka or blackbelly, Kurinskii podust or Kura nase, uzkotelii Kurinskii podust, all in Russian; Kura undermouth, Kura nase].

Systematics

Earlier works by Elvira (1986; 1988; 1991) placed this species as a subspecies of C. oxyrhynchum but in Elvira (1997), using the phylogenetic species concept and following the studies of Smirnov (1992), this taxon is recognised as a species. C. oxyrhynchum is then found in more northerly rivers of the western Caspian Sea basin remote from Iranian waters. C. cyri orientalis (Bianco and Banarescu, 1982 is described from Fars (see below under C. regium).

Chondrostoma cyri Kessler, 1877 was described from the Kura River, Tiflis (= Tbilisi), Georgia and Chondrostoma oxyrhynchum from the Kuma River near Georgiyevsk, Russia in the Caspian Sea basin.

Alburnus alasanicus Kamenskii, 1901 described in part from the Alasan, Alazan' or Alazani River, a left bank Kura River tributary in Georgia, Chondrostoma schmidti, Berg, 1910 from the Alazan' River at Naporiri, and Chondrostoma leptosoma Berg, 1914 from the Kars-tchai, a tributary of the Aras River in Turkey, the Aras by Kopri-kei, near Erzurum, Turkey, and the lower Aras at Karadonly and Dzhulfa in the former U.S.S.R., are synonyms. Subspecies are not recognised (Elvira, 1991; 1997). C. leptosoma was founded on an elongate form from the Karasu in the Aras River basin.

Two syntypes of Chondrostoma cyri are also in ZISP (10919) from "Tiflis" collected by Kessler in September 1875. A syntype of Chondrostoma oxyrhynchum is in the Zoological Institute, St. Petersburg (ZISP 2881) from "Fl. Sunsha" collected in 1830 by Ménétries. According to Elvira (1988), the type locality is Kuma R. at Georgijewsk and 2 syntypes are under ZISP 10922. Another syntype of Chondrostoma oxyrhynchum is in the Natural History Museum, London (BM(NH) 1897.7.5:28 (184.8 mm standard length), formerly in ZISP, as is other syntype of Chondrostoma cyri (BM(NH) 1897.7.5:27 (correctly numbered 27, 78.4 mm standard length), formerly in ZISP)(Elvira, 1988; personal observations).

Five syntypes of Chondrostoma leptosoma are in the Zoological Institute, St. Petersburg (ZISP 9098) according to (Elvira, 1988) but there are 15 syntypes under this number from the "Reka Araks", 1888, Warpochowsky as well as additional material listed as syntypes with numbers ZISP 9107 ("Fl. Araxes", 1888, Warpochowsky, 12 fish), ZISP 5180 ("Kars-tschai", 1879, Dr. A. Brandt, 3 fish), ZISP 9099 ("Reka Araks", 1888, Warpochowsky, 4 fish), ZISP 15264 ("Reka Araks", 20.III.1911, 2 fish), and ZISP 15516 ("Reka Araks near settlement Djulfa", 17.VI.1911, 13 fish).

Key characters

This species is the only one in its genus in northern Iran and can be recognised by generic characters.

Morphology

Kuru (1981) gives the following meristic characters for 103 specimens from the Aras and Kura river basins in Turkey:- 10-12 dorsal fin rays, 10-11 anal fin rays, 9-10 pelvic fin rays, 9-15 pectoral fin rays, 52-62 lateral line scales, 13-18 scales around the caudal peduncle, 17-32 gill rakers, and 5-6 pharyngeal teeth on each arch (note that the statistical treatment in this paper is in error and the conclusion that species of Chondrostoma in Turkey are not distinct is therefore incorrect). There is clinal variation in scale numbers, the number increasing from south to north and Elvira (1988; 1991) gives the total range for characters of this species as dorsal fin branched rays 7-9, usually 8, anal fin branched rays 8-10, usually 9-10, pectoral fin branched rays 13-18, usually 14-16, pelvic fin branched rays 7-8, usually 8, lateral line scales 50-68 (to 73 in Kazancheev (1981) and from 48 in Chikova (1967)), scales above the lateral line 7-10, usually 8-10, scales below the lateral line 3-6, usually 4-6, pharyngeal teeth 6-5 or 5-5, more rarely 6-6 and mode 6-5, and gill rakers 21-29. Vertebrae number 43-45.

The mouth is arched with a thin horny layer on the lower jaw. Scales are rounded in overall shape with indentations above and below a central, rounded protuberance on the anterior margin. The anterior margin may be wavy. There are few anterior and posterior radii, few circuli and a subcentral anterior focus. There is a pelvic axillary scale. The gill rakers are short and reach the one below or just past it when appressed. Pharyngeal teeth are compressed and thin but deep with a long, thin and concave grinding surface. Teeth tips may be slightly hooked. The gut has numerous anterior loops.

Thirteen specimens from Djulfa (presumably in Azerbaijan opposite the Iranian town across the Aras River) have dorsal fin branched rays 8(12) or 9(1), anal fin branched rays 9(9) or 10(4), and pharyngeal teeth 6-5(5) or 6-6(1).

Sexual dimorphism

Unknown.

Colour

The flanks are silvery but may have dark pigment spots which may, or may not, form a stripe. Paired fins are orange to reddish and median fins grey. The dorsal and caudal fins have dark margins. The peritoneum is black.

Size

Reaches 80.0 cm and about 5.0 kg.

Distribution

Found in the rivers draining to the western coast of the Caspian Sea from the Kuma River in the north southward to the Kura and Aras river basins in the south. Recorded from the Aras River basin of Iran (Abdoli, 2000).

Zoogeography

This genus has a European and Middle Eastern distribution. Its relationships to other taxa are poorly known.

Habitat

Unknown. Found principally in streams and rivers.

Age and growth

Fish are mature at 2 years of age and life span is at least 5 years.

Food

Diet is assumed to consist of bottom organisms including aquatic insect larvae, detritus and vegetation scraped from the substrate.

Reproduction

Up to 16,217 eggs are produced and maximum diameter is 1.69 mm. The spawning season is in the spring, peaking in April in the Kura River basin (Abdurakhmanov, 1962).

Parasites and predators

None reported from Iran.

Economic importance

None.

Conservation

Kiabi et al. (1999) consider this species to be conservation dependent, in the south Caspian Sea basin according to IUCN criteria. Criteria include sport fishing, possibly few in numbers, limited range (less than 25% of water bodies), absent in other water bodies in Iran, absent outside the Caspian Sea basin.

Further work

Biology in Iranian waters needs study.

Sources

Morphology based on Bianco and Banarescu (1982), Elvira (1986; 1988), Nelva et al. (1988).

Type material: See above, Chondrostoma cyri (BM(NH) 1897.7.5:25, formerly in ZISP), Chondrostoma oxyrhynchum (BM(NH) 1897.7.5:28, formerly in ZISP), and Chondrostoma leptosoma (ZISP 15516).

Iranian material: None.

Comparative material: CMNFI 1980-0812, 2, 101.9-107.9 mm standard length, Turkey, Kars, Selim Çayi (40º28'N, 42º47'E).

Chondrostoma orientale
Bianco and Banarescu, 1982

Chondrostoma cyri orientalis Bianco and Banarescu, 1982 was originally described from the "Pulwar River near Persepolis".

The holotype (IZA 8170, 93.7 mm standard length, examined by me) and 19 paratypes (IZA 7833, 51 specimens under this number, 35.4-90.1 mm standard length) of Chondrostoma cyri orientalis are in the Istituto di Zoologia dell'Universitá di L'Aquila, Italy (Elvira, 1988). Two paratypes of Chondrostoma cyri orientalis are stored in the Field Museum of Natural History, Chicago (FMNH 94519)(Ibarra and Stewart, 1987), 1 paratype is in the Muséum national d'Histoire naturelle, Paris (1982-1014), 1 paratype is in the United States National Museum, Washington (USNM 227934), 2 paratypes are in the Academy of Natural Sciences, Philadelphia (ANSP 150985), and 6 paratypes are in the Canadian Museum of Nature, Ottawa (CMNFI 1982-0365, formerly IZA 7833, 37.8-88.7 mm standard length). The total number of paratypes is 75, originally under IZA 7833 but some dispersed as noted above, with 10 further fish in the Institutul de Stiinte Biologice, Bucurešti, Romania (ISBB) but uncatalogued (Bianco and Banarescu, 1982).

 

Barzegar and Jalali (2006) report parasites in this species from Kaftar Lake as Unio sp., Lernaea cyprinacea, Ichthyophthirius multifilis and Diplostomum spathaceum.

Chondrostoma regium
(Heckel, 1843)

Common names

jokhorak, nazok, nazi; heif-e nan (= waste of bread, i.e. valueless) in Khuzestan; سياه ديم (= siah deem in Behbehan); سياه دم (= siah dom, meaning blacktail); كپور پوزه دار (= kapur puzeh dar).

[baloot muluki, pangki; zurri (= the harmful one) at Mosul (also used for Alburnus mossulensis, Aphanius spp., Gambusia and any small fishes or large fishes when young); terris or terris achmar meleki (= royal red terris) at Aleppo (= Haleb, Syria), all in Arabic; based on Heckel (1843b) for zurri and terris; king nase].

Systematics

Chondrochilus regius Heckel, 1843 was described from the "Orontes" (= Asi) (but see below) and "Tigris". Elvira (1988; 1991; 1997) considers Chondrostoma orientale to be a valid species while Nelva et al. (1988) retain it as a subspecies of C. cyri. Bianco and Banarescu (1982) placed orientalis in C. cyri on the basis of similar dorsal and anal fin ray counts, scale counts and to a certain degree pharyngeal tooth formula.

Banarescu (1960) regarded C. regium as only a race of a widespread species, C. nasus (Linnaeus, 1758). C. nasus has larger scales on average and 6-6 pharyngeal teeth (Berg, 1949); Heckel (1846-1849c) found 47 C. nasus from the Danube River had 6-6 teeth, 2 had 6-7 and 2 had 5-6 while in 13 C. regium the count was 7-6 in 12 fish and 6-6 in 1 fish. Krupp (1985c) considers C. regium to be distinct while recognising the small degree of morphological variation between species in this genus. Data gathered for Iran show a wide range in scale and teeth counts (see below). Ladiges (1960) identified specimens from the same bodies of water in Turkey as members of both species. The earlier literature on the systematics of this genus remain confused (see Elvira (1988) for comments on Ladiges (1966) and Kuru (1981)) and the morphology summarised here for this species does not adequately resolve the problem. There may well be significant variation of a clinal nature, altitude and temperature may be important, and habitat types (lentic or lotic) may affect body form. Most samples examined previously are too small in numbers and differences due to size and sex could not be adequately assessed.

Twelve syntypes of Chondrostoma regium are in the Naturhistorisches Museum Wien (7 fish as NMW 52532-52535 from the Quwayq (= Kueik) River near Aleppo and 5 fish as NMW 52536-52538 from the Tigris River near Mosul)(Elvira, 1988). Krupp (1985c) gives further details. All material was collected by Th. Kotschy in 1842 from the Quwayq and in 1843 from Mosul and the range in standard length for the fish from the Quwayq is 102-166 mm and from Mosul 11.9-24.1 cm. The Vienna catalogue lists only 6 fish but the card catalogue in 1997 lists NMW 52532 (2 fish), 52533 (2), 52534 (2), 52535 (1), 52536 (2), 52537 (1) and 52538 (2) as syntypes. The type locality "Orontes" (= Asi) in Heckel (1843b) seems to be an error.

Key characters

This species is the only one in its genus in southern Iran and can be recognised by generic characters.

Morphology

?re-work

The following counts are from literature sources; my counts in the table below often show a wider range: lateral line scales 56-72 (47-55 for orientalis), scales above the lateral line 9-13 (8-9 in orientalis), and scales below the lateral line 5-6 (4-5 in orientalis). Lateral line scale counts for Iranian fish are as follows: Tigris - 50(1), 51(4), 52(3), 53(12), 54(5), 55(7), 56(4), 57(4), 58(4), 59(8), 60(7), 61(6), 62(5), 63(3), 64(3), 65(1), 66(2), 67(1) or 69(2); Kor (= orientalis) - 49(1), 50(2), 51(3), 52(8), 53(3), 54(7), 55(3) or 57(4). Despite a lower range, the counts for the Kor River basin are matched by a sample from Cheshmeh Javari near Ravansar, Kermanshahan (CMNFI 1979-0287) which have a range of 50-58, leading to a supposition of altitudinal or habitat variation :-

Dorsal fin branched rays 8-11, mode 9 (note Bogutskaya (1997a) gives a mode of 10) (7-9, mostly 8 for orientalis), anal fin branched rays 9-12, mode 11 (note that Bogutskaya (1997a) gives modes of 11 or 12) (9-10, mode 9 for orientalis), pectoral fin rays 14-18, mostly 15-17 (13-15, mostly 14 in orientalis) and pelvic fin rays 6-9, mostly 8 (7-8, mostly 8 in orientalis). Gill rakers 18-36 (probably some lower counts are of rakers on the lower arch only and ranges in single studies, presumably to a consistent technique, are 22-34, 24-31, 25-34 and 25-36) (22-28, 22-30 or 28-32 by different authors for orientalis). Counts for the whole arch on Iranian fish give a wide range of 19-34, highly correlated with size, larger fish having more (or more discernible) rakers than smaller fish (r = 0.5049, p<0.001, n = 90).

Scale radii are few and restricted to the posterior field. Total vertebrae 46-48. Pharyngeal teeth 6-5, 6-6, 6-7, 7-5, 7-6 and 7-7, mode 6-6 or 7-6 (6-5, 6-6, 5-6 and 7-5, mostly 6-6 for orientalis) but see above. The mouth is straight (= transverse) with a thick horny layer on the lower jaw.

Meristics for southern Iranian specimens of Chondrostoma:


Locality/Dorsal Fin Branched Rays

7

8

9

10

x

S.D.

Tigris River Basin

35

46

1

8.6

0.520

Kor River Basin

1

30

8.0

0.180

Locality/Anal Fin Branched Rays

8

9

10

11

12

x

S.D.

Tigris River Basin

4

45

21

9

3

9.5

0.892

Kor River Basin

23

8

9.3

0.445

Locality/Pelvic Fin Rays

7

8

9

x

S.D.

Tigris River Basin

3

77

2

8.0

0.248

Kor River Basin

1

29

1

8.0

0.258

Locality/Lateral Line Scales

Range

x

S.D.

Tigris River Basin

50-69

57.8

4.553

Kor River Basin

49-57

53.2

2.131

 

Locality/Total Vertebrae

42

43

44

45

46

47

48

49

x

S.D.

Tigris River Basin

1

20

17

9

14

10

9

2

45.1

1.833

Kor River Basin

5

20

5

1

43.1

0.680

Sexual dimorphism

Unknown.

Colour

The back is olive-brown with bluish tinges and the flanks and belly are silvery-white. The dorsal and caudal fins are greyish and the other fins hyaline. Some fish have bright orange fins, the pectorals paler, the pelvics and anal fins fringed by white. The dorsal and caudal fins have a black margin, wide on the caudal. These fin colours give them a flag-like effect (Heckel, 1843b). The caudal fin can be orange, distally black, with the extreme edge white in freshly dead fish.

Size

Attains 40 cm and 1 kg.

Distribution

Found in the Tigris-Euphrates basin and the Mediterranean basins of southeastern Turkey and the northern Levant. In Iran found in the Tigris River basin. Additional localities are springs (sarabs) near Kermanshah, the Marun River, the Hawr al Azim marsh (Wossughi, 1978; Abdoli, 2000). Ghorbani Chafi (2000) lists the Bazoft and Kuhrang rivers in the upper Karun River basin and also possibly the Zayandeh River of the Esfahan basin.

Zoogeography

This genus has a European and Middle Eastern distribution. Its relationships to other taxa are poorly known.

Habitat

Found in both rivers and lakes (and reservoirs) but habitat requirements have not been studied in Iran. Ünlü (2006) reports that this species prefers stone grounds and still waters in rivers and lakes in Turkey.

Age and growth

Khalaf et al. (1986) studied this species in the Diyala River, Iraq. Maximum age group is 7+ years, males and females show no difference in weight at the same length and samples from three adjacent areas show no major differences in growth rates. Length-weight relationship was W = 0.0480 L2.49 (n = 255, r = 0.88). Males mature at 15.0 cm and females at 19.0 cm in the Diyala River at Rustamiyah in Iraq (Allouse et al., 1986). A population at Al Kadhmia north of Baghdad in the Tigris River had four age classes dominated by the three year age class, with all fish being sexually mature during the second year. Fish smaller than 15 cm for males and 17 cm for females were immature. The disparity in age structure with the Diyala River population was attributed to pollution in the Diyala (Daoud and Qasim, 1999).

Polat and Gümüş (1995) aged a population of this species in the Bafra Altınkaya Dam lake in Turkey using vertebrae, otoliths, scales, opercle and subopercle. Age reached 5, perhaps 6, years and scales were found to be the best structure to use. Polat et al. (1999) found a similar age range in the Suat Uğurlu Dam, Turkey with annulus (hyaline ring) formation in October to February. Oymak (2000) examined growth characteristics of this species in the Atatürk Dam on the Turkish Euphrates River. Eight age groups were found and age-length and age-weight equations given for females and males were Lt = 38.67[1-e-0.136126(t+3.073799)], Wt = 527.52[1-e-0.136126(t+3.073799)]3.1986 and Lt = 35.01[1-e-0.168137(t+2.754214)], Wt= 724.73[1-e-0.168137(t+2.754214)]3.2779 respectively. The length-weight relationships were obtained as Log W = -5.4153 + 3.1986 Log FL in females and Log W = -5.6212 + 3.2779 Log FL in males. The condition factor was high in age group 7 and high in April and May, lowest in December and January. Gümüş et al. (2002) found deposition of hyaline rings was synchronous with decrease in food diversity in autumn in the Suat Uğurlu Dam, Turkey. Aydin et al. (2004) demonstrated a positive linear relationship between otolith length and fish length for this species in Keban Dam Lake, Turkey.

Food

This species is omnivorous taking insect larvae and eggs and fry of other fishes. Gut contents also include diatoms and algae as well as large quantities of sand. However, Gümüş et al. (2002) examined diet in the Suat Uğurlu Dam, Turkey and found Navicula, Cymbella and Synedra were the most frequently consumed organisms. This species feeds mostly on Bacillariophyta in this dam but also Chlorophyta, Cyanophyta, Xanthophyta, Euglenophyta and Rotifera. Diet varied with seasonable availability of food items.

Reproduction

Studies on the Diyala River population in Iraq found fish to be mature in December and by January females lacked eggs. Each female produces up to 6900 eggs and number of eggs increases linearly with length (Allouse et al. (1986). The breeding season at Al Kadhmia in the Tigris River near Baghdad was March-May (Daoud and Qasim, 1999). Ünlü (2006) reports up to 13,280 eggs for fish in the Tigris River of Turkey. Beckman (1962) states that this species probably spawns in May or June in Syria and Oymak (2000) found that condition factors were highest in April and May in the Atatürk Dam, Turkey.

Parasites and predators

Barzegar et al. (2004) examined this species for parasites in fish from the Beheshtabad river in Chahar Mahall va Bakhtiari Province and found Lernaea cyprinacea, Dactylogyrus ergensi, Ichthyophthirius multifilis and Myxobolus sp. Jalali et al. (2005) summarise the occurrence of Gyrodactylus species in Iran and record G. sp. from the Dez and Karun rivers in Chondrostoma nasus, presumably this species. Riassy et al. (2009) found the digenean eye parasite Tylodelphys clavata in fish from Choghakhor Lagoon.

Economic importance

This species has been caught and used for food in Khuzestan.

Conservation

This species is relatively common and is not widely used as food; it may not need conservation. However it is listed as endangered in Turkey (Fricke et al., 2007).

Further work

Its biology in Iran needs study and this may reveal conservation needs.

Sources

?re-work

Type material: See above, Chondrostoma cyri orientalis (IZA 8170, IZA 7833, CMNFI 1982-0365, formerly IZA 7833).

Iranian material: CMNFI 1979-0025, 16, 22.1-119.0 mm standard length, Fars, Kor River at Marv Dasht (29º51'N, 52º46'30"E); CMNFI 1979-0028, 14, 32.2-139.1 mm standard length, Fars, Kor River drainage (no other locality data); CMNFI 1979-0059, 1, 72.2 mm standard length, Fars, Pulvar River (30º01'30"N, 52º57'E); CMNFI 1979-0061, 14, 9.5-56.5 mm standard length, Fars, stream tributary to Pulvar River (30º04'N, 53º01'E); CMNFI 1979-0245, 5, 35.3-47.1 mm standard length, Sharestan-e Bahktiari va Chahar Mahall, stream in Ab-e Shalamzar drainage (32º08'N, 50º51'E); CMNFI 1979-0247A, 4, 57.2-65.3 mm standard length, Sharestan-e Bakhtiari va Chahar Mahall (31º57'N, 51º01'E); CMNFI 1979-0248, 2, 39.2-65.2 mm standard length, Sharestan-e Bakhtiari va Chahar Mahall, stream 3 km east of Boldaji (31º55'N, 51º05'E); CMNFI 1979-0271, 11, 60.0-131.3 mm standard length, Lorestan, Kashkan River drainage (33º39'N, 48º32'30"E); CMNFI 1979-0272, 1, 58.5 mm standard length, Lorestan, river at Nokhor (ca. 33º40-47'N, ca. 48º28-45'E); CMNFI 1979-0279, 2, 61.8-134.0 mm standard length, Lorestan, Khorramabad River (33º37'N, 48º18'E); CMNFI 1979-0280, 1, 114.5 mm standard length, Lorestan, Kashkan River drainage (ca. 33º43-47'N, 48º12-15'E); CMNFI 1979-0283, 1, 137.0 mm standard length, Kermanshahan, river 15 km before Kermanshah (34º21'N, 47º07'E); CMNFI 1979-0287, 22, 56.6-112.5 mm standard length, Kermanshahan, Chashmeh Javari near Ravansar (ca. 34º42'N, ca. 46º40'E); CMNFI 1979-0286, 11, 77.4-100.4 mm standard length, Kermanshahan, Ravansar River at Ravansar (34º43'N, 46º40'E); CMNFI 1979-0289, 1, 131.5 mm standard length, Kermanshahan, Diyala River drainage (34º28'N, 45º52'E); CMNFI 1979-0368, 4, 54.0-84.5 mm standard length, Khuzestan, Karkheh River (32º24'30"N, 48º09'E); CMNFI 1979-0370, 6, 187.3-221.6 mm standard length, Khuzestan, Karkheh River (32º12'N, 48º14'30"E); CMNFI 1979-0382, 2, 37.7-62.5 mm standard length, Khuzestan, Karun River at Shushtar (32º03'N, 48º51'E); CMNFI 1979-0392, 1, 53.7 mm standard length, Khuzestan, Zard River (ca. 31º32'N, ca. 49º48'E); CMNFI 1979-0421, 5, 114.0-122.0 mm standard length, Boyer Ahmadi-ye Sardsir va Kohkiluyeh, stream in Khersan River drainage (30º24'N, 51º47'E); CMNFI 1979-0499, 1, 113.0 mm standard length, Fars, irrigation ditch 32 km from Kor River bridge (30º04'30"N, 52º36'E); CMNFI 1979-0500, 7, 94.8-110.5 mm standard length, Fars, Pulvar River at Naqsh-e Rostam (29º59'N, 52º54'E); CMNFI 2007-0100, 2, 165.4-165.7 mm standard length, Azarbayjan-e Bakhtari, Kalwi Chay near Piranshar (ca. 36º44'N, ca. 45º10'E); CMNFI 2007-0111, 2, 183.3-191.7 mm standard length, Kermanshahan, Alvand River near Sar-e Pol-e Zahab (ca. 34º36'N, ca. 45º56'E); CMNFI 2007-0113, 2, 106.7-145.0 mm standard length, Kermanshahan, Razavar River, Qareh Su tributary (ca. 34º25'N, ca. 47º01'E); CMNFI 2007-0115, 3, 72.5-96.5 mm standard length, Kermanshahan, Qareh Su basin north of Kermanshah (ca, 34º34'N, ca. 46º47'E).

Comparative material: BM(NH) 1931.8.12:1-3, 2, 136.0-172.2 mm standard length, Iraq, near Mosul (36º20'N, 43º08'E); BM(NH) 1971.4.2:6, 1, 147.7 mm standard length, Iraq, River Tigris near Mosul (36º20'N, 43º08'E); BM(NH) 1974.2.22:81-82, 1, 197.5 mm standard length, Iraq, Great Zab near Eski Kelek and near Bekhne Dam (no other locality data).

Genus Crossocheilus
Kuhl and van Hasselt, 1823

Kottelat (1987) retains the spelling Crossocheilus Kuhl and van Hasselt in van Hasselt, 1823 as first reviser. The name was spelt Crostocheilus early in the text but this has never been used again and Crossocheilus appeared with the description. Crossochilus Günther, 1868 is an incorrect emendation (Eschmeyer, 1990).

The genus is found chiefly in the Oriental Region but extends into Iran with one species. There are about 18 species.

The genus is characterised by an elongate body with a rounded belly; the mouth is inferior and transverse, the crenulated or fringed upper lip being continuous with the snout (not separated by a groove); the lower jaw has a horny covering and behind this are several rows of lobate papillae; 1-2 pairs of barbels; gill membranes attached to isthmus; dorsal and anal fins are short and spineless; the lateral line is complete; scales are large to moderate in size; the intestine is very long; and the peritoneum is black.

The lower surface of the head bears an "adhesive apparatus", the mechanism of which has been investigated by Singh (1993) for Crossocheilus latius latius, a subspecies not found in Iran. The fringed upper lip acts as a food strainer as well as part of the adhesive apparatus. Both this lip and the area behind the lower lip are heavily tuberculate with glandular openings and irregularly arranged hard ridges. Mucus from the glands in conjunction with the ridges holds the fish to the substrate.

Crossocheilus latius
(Hamilton, 1822)

Common names

None.

[ispigoar or dogra in Pakistan].

Systematics

Cyprinus latius was described from the Tista River in India/Bangladesh and types are unknown (Eschmeyer et al., 1996).

Bianco and Banarescu (1982) and Bănărescu (1986) consider Discognathus adiscus Annandale, 1919 described from Sistan (type locality given below) to be a synonym of this species which is represented in Sistan and Baluchestan by Crossocheilus latius diplocheilus (Heckel, 1838), originally described from Kashmir with syntypes in the Naturhistorisches Museum Wien under NMW 48820 (7 fish). Bănărescu (1986) cites 1 fish under NMW 48420 as possibly the holotype but this specimen is dated 1839 which is anachronistic. Berg (1949) considers Discognathus adiscus to be a distinct species. I concur with Bianco and Banarescu (1982) and Bănărescu (1986).

Characters advanced by Berg (1949) for separating the two species are number of barbels (4 in adiscus, 2 in latius diplocheilus where mouth angle barbels are absent or rudimentary, not the rostral ones as implied by Bianco and Banarescu (1982)), upper lip fringe (barely developed in adiscus, distinctly developed in latius diplocheilus), the posterior swimbladder (conical in adiscus, elongate cylinder in latius diplocheilus), and papillae on the lower lip and chin (rudimental in adiscus and latius diplocheilus but the latter has almost free lateral edges and an attached posterior end - this condition is not specified for adiscus). Bianco and Banarescu (1982) and Bănărescu (1986) found some latius diplocheilus specimens to have 4 barbels (and this is given too as a character of C. latius latius), and no difference in development of lip papillae in specimens from the Indus River basin (actually my reading of Berg (1949) cited above does not indicate that papillae development differs but that the sucker area has almost free lateral edges and an attached posterior end; this occurs in Sistan fish but not in 4 fish from the Hamun-e Mashkid and Makran basins of Iran which have a fold in the flesh behind the tuberculate area - these latter fish are very small however, 20.8-27.8 mm standard length, and I lack extensive comparative adult material from outside the Sistan basin in Iran and from neighbouring Pakistan to make an adequate analysis of nominal adiscus and latius diplocheilus in this and other characters). My observations of the posterior swimbladder development indicate a great individual variation in form for Sistan fish: the swimbladder may be conical, elongate and tapering, rounded posteriorly, expanded posteriorly, rounded posteriorly after a constriction, or even a narrow elongate cylinder supposedly characteristic of latius. Fringe development of the upper lip is also quite variable and seems to be relatively well-developed in larger Sistan fish.

Karaman (1971) described a new genus, Hemigarra, for Tylognathus elegans Günther, 1868 and Discognathus adiscus Annandale, 1919. He places Crossocheilus adiscus as the Sistan subspecies of his Hemigarra elegans (= Hemigrammocapoeta elegans here, q.v.) which is found in Mesopotamia. Karaman (1971) distinguishes the two subspecies by the former having densely arranged papillae on the chin as opposed to sparse papillae. Bianco and Banarescu (1982) and Bănărescu (1986) state that it is not related to Hemigrammocapoeta elegans but is a typical Crossocheilus species.

The type locality of Discognathus adiscus is Sistan by implication, as no locality is given for the holotype in Annandale (1919b). Menon and Yazdani (1968) concur. Distribution is given as "small watercourses and pools in the plains of Seistan" and "Nasratabad, irrigation channel in Consulate garden; pool in the desert 5 miles south of Nasratabad; pools in stream-bed 12 miles north of Nasratabad; channels in the reed-beds of the Hamun-i-Helmand near Lab-i-Baring, and channel leading out of the Hamun 12 miles east of Lab-i-Baring; small watercourse, Lutak, southern Seistan", and one of these is presumably the type locality.

Twenty syntypes of Discognathus adiscus are in the Zoological Survey of India, Calcutta (ZSI F9758/1) (Menon and Yazdani, 1968). Annandale (1919b) cites ZSI 9763/1 as the holotype catalogue number. Three syntypes are in the Zoological Institute, St. Petersburg (ZISP 25411) from "Nasratabad, Seistan, Indian Museum, Dr. Hora" and measure 38.0-43.4 mm standard length. Two syntypes (listed as cotypes) measuring 44.8-45.5 mm standard length from "Jellalabad" with the annotation "Ind. Mus. Ex. F 9762/1" are in the Natural History Museum, London (BM(NH) 1919.8.16:7-8; the outside has 1919.3.16:7-8, incorrectly).

Key characters

The characters of the genus, particularly in the mouth region, serve to identify the only species in Iran.

Morphology

Four short barbels are present, the rostral ones longer than those at the mouth corner. The upper lip covers the upper jaw, is granular or tuberculate and has a marginal fringe, variably developed and most apparent in larger fish. The lower lip is only apparent at the sides and the exposed lower jaw has a granular or tuberculate pad without a free posterior margin but with almost completely free edges.

Dorsal fin with 2-3 unbranched and 8-9 branched rays, anal fin with 2-3 unbranched and 5 branched rays, pectoral fin branched rays 14-17, and pelvic fin branched rays 7-9. Lateral line scales 33-39. Scales may have short dorsal and ventral projections from the margin at about one-third of the scale length from the posterior edge. There is a pelvic axillary scale. Scales have 9-10 radii on the posterior field and are elongate with a notably anterior focus. Radii in large fish are parallel rather than divergent. The anus is 4-5 scales in advance of the anal fin origin. Gill rakers 17-25, small reaching the adjacent or second raker when appressed. Pharyngeal teeth usually 3,3,5-5,3,3 or 2,4,5-5,4,2, depending on how the crowded teeth are counted; major row teeth are usually 5 but may be 4 or 6, middle row teeth are 3 or 4, and minor row teeth 2 or 3, more rarely 1 (this difficulty in assigning teeth to rows is the reason for omitting frequency distributions below). Supernumerary teeth may be present to further confuse counts. The crown of major row teeth are flattened, the anterior tooth may be rounded and some teeth may have a small hooked tip. The gut is very long and complexly coiled. The chromosome number is probably 2n=48 (Klinkhardt et al., 1995).

Iranian fish from Sistan and Baluchestan have the following meristic characters: dorsal fin branched rays 8(81) or 9(1), anal fin branched rays 5(81), pectoral fin branched rays 14(28), 15(37), 16(16), or 17(1), and pelvic fin branched rays 7(3), 8(76), or 9(3). Lateral line scales 33(1), 34(2), 35(12), 36(31), 37(32) or 38(4). Total gill rakers 20-25, but not countable with great accuracy since the smallest rakers are difficult to detect at the ends of the arch. Total vertebrae 34(6), 35(20), 36(9) or 37(1).

Sexual dimorphism

Unknown.

Colour

The back is bluish-grey in Sistan fish or brownish to greenish with irregular spots in other populations and the belly light pink to yellowish-white or silvery-white. Fins are pink and the dorsal and caudal fins have a grey tinge. The flank has a bluish, mid-lateral stripe in Sistan fish and in preserved ones scattered melanophores, or small blotches of less than scale size, or clumps of melanophores centred on upper flank scales and more dispersed on the lower flank. There is a broad stripe along the back mid-line. Fins in preserved fish from Sistan are mostly immaculate except in the larger fish with some melanophores lining rays basally. The caudal fin is distinctive in larger fish from Sistan in having the rays of the lower half of the fin heavily pigmented while the upper half rays are only lightly pigmented. Peritoneum is dark brown to black.

Size

Attains 14.6 cm although the largest fish recorded from Sistan was 93.2 mm standard length.

Distribution

Found in submontane areas of Afghanistan, Pakistan and India as well as eastern and southeastern Iran. The main areas of distribution are Sistan, the Hamun-e Mashkid basin including the Simish River and coastal streams of Makran from the Jagin to middle and upper Nikshahr rivers and the middle and upper Bahu Kalat River including its Sarbaz River reach. (Nikol'skii, 1899; Annandale, 1919b; Berg, 1949; Bianco and Banarescu, 1982; J. Holčík, in litt., 1996; Abdoli, 2000).

Zoogeography

This distribution in Iran marks the western limit for the genus and the relationships of the species lie to the east.

Habitat

Very abundant in small streams, including those with rocky or muddy beds, irrigation ditches, channels in reed beds and pools in Sistan, less common in Baluchestan streams. This species is found in large schools in Sistan in still or slow-flowing water, on the bottom during the day but it may swim at the surface in the evenings. It is common in the smallest permanent water channels but Annandale and Hora (1920) reported it to be in small numbers in the reed beds in winter and these were dead or dying, perhaps because of low oxygen conditions associated with vegetation decay. Large numbers die each year in drying stream beds as salt content increases and the water is fouled by sheep and goats. Tekrival and Rao (1999) report its aquarium preferences as 18-22°C, pH 6.5-7.2, algae as food, not too bright lighting, bottom dwelling with stones, roots and crevices preferred and cave brooding reproduction.

Age and growth

Unknown.

Food

Diet is algae on muddy bottoms. The type subspecies is a bottom feeding herbivore taking more than 90% plant food such as algae, diatoms and macrophytes as well as detritus (Sharma, 1984; Singh and Bahuguna, 1984). Iranian fish contain detritus and some insect remains, possibly as accidental inclusions.

Reproduction

Iranian adult specimens were caught in May in Sistan and show signs of developing reproductive organs suggestive of summer spawning.

Parasites and predators

Jalali et al. (2000) describe two new species of monogenean, Dactylogyrus faridpaki and D. eslamii, from this species in the Bahu Kalat River of Baluchestan.

Economic importance

This species is of no economic importance although Butt (1995) suggests it could be cultured as food and as a forage fish in Pakistan.

Conservation

This species does not appear to be under any major threat as it can survive drying of the Sistan lakes in small ditches and streams.

Further work

The biology of this species needs investigation as does the taxonomic status of Sistan populations.

Sources

Mirza (1972) for colour.

Type material: See above, Discognathus adiscus (ZISP 25411, BM(NH) 1919.8.16:7-8).

Iranian material: CMNFI 1979-0224, 8, 43.6-55.4 mm standard length, Sistan, effluent of Hirmand River (30º53'30"N, 61º27'E); CMNFI 1979-0226, 277, 29.7-78.8 mm standard length, Sistan, pool near Kuh-e Khajeh (30º57'N, 61º17'E); CMNFI 1979-0227, 4, 37.0-48.9 mm standard length, Sistan, naizar at Kuh-e Khajeh (30º57'N, 61º16'E); CMNFI 1979-0228, 1, 42.9 mm standard length, Sistan, ditch 1 km from Zabol (31º02'30"N, 61º31'E); CMNFI 1979-0229, 5, 52.3-93.2 mm standard length, Sistan, ditch 5 km from Zabol (31º03'N, 61º33'E); CMNFI 1979-0230, 1, 48.3 mm standard length, Sistan, Hamun-e Puzak (ca. 31º15'N, ca. 61º42'E); CMNFI 1979-0232, 9, 44.0-65.9 mm standard length, Sistan, ditch 11 km from Zabol (ca. 30º58'30"N, ca, 61º36'E); CMNFI 1979-0234, 17, 40.4-49.3 mm standard length, Sistan, effluent of Hirmand River (30º54'N, 61º40'E); CMNFI 1979-0318, 2, 24.0-27.8 mm standard length, Baluchestan, Sarbaz River at Huvar (26º09'N, 61º27'E); CMNFI 1979-0333, 2, 20.8-21.2 mm standard length, Baluchestan, Mashkid River west of Kuhak (ca. 27º05'N, ca. 63º12'E).

Genus Ctenopharyngodon
Steindachner, 1866

The grass carp genus contains only a single species found in East Asia but widely introduced for food and its ability to digest macrophytes.

This genus is characterised by a rounded body and broad head, the eyes are large and positioned at or above the body axis and often visible from the underside of the head, mouth wide and terminal, no barbels, moderate-sized scales, a complete lateral line, dorsal and anal fins short and lacking spines, branchial membranes attached to the isthmus, short unfused gill rakers, brown to black peritoneum, and pharyngeal teeth in 2 rows with the crowns strongly compressed and serrate and with a longitudinal groove on the grinding surface.

Ctenopharyngodon idella
(Valenciennes, 1844)

Common names

كپور علفخوار (= kapour-e alaf khaar or alaf khoar or kopur 'laf khoar, carp grass-eater or grass-eater), آمور (= amur), سفيد پرورشي (safid parvareshi or mahid safid parvareshi meaning cultured white fish, from a resemblance to mahi safid, i.e. Rutilus frisii kutum).

[grass carp, white amur].

Systematics

Leuciscus idella was originally described from China. A hybrid of this carp and Rutilus frisii has been bred at the Astaneh Ashrafie Fisheries Research Station and named "Samur" (Iranian Fisheries Research and Training Organization Newsletter, 11:6, 1996). See under Rutilus frisii for more information.

Key characters

This species is identified by the eyes being low on the side of the head, the anal fin is far back on the body close to the caudal fin, and pharyngeal teeth have large, parallel grooves on the grinding surface.

Morphology

Lateral line scales 34-47. Scales have a wavy anterior edge, central focus and moderate numbers of anterior and posterior radii. Dorsal fin branched rays 6-8, usually 7, after 3 unbranched rays, anal fin branched rays 7-9, usually 8, after 3 unbranched rays, pectoral fin branched rays 13-20 and pelvic fin branched rays 7-8. Gill rakers number 15-18 and touch the adjacent raker when appressed. Vertebrae 40-47. Pharyngeal teeth are 2,5-5,2, 2,4-5,2, 2,4-4,2, or 1,4-5,2 and are obviously serrated with a longitudinal grooves. The gut is long and complexly coiled. The diploid chromosome number is 48, the triploid 72 (Klinkhardt et al., 1995; Nowruzfashkhami et al., no date). Serum immunoglobulins have been characterised by Soltani et al. (2003).

Sexual dimorphism

Nuptial tubercles are evident on the male head, upper caudal peduncle, dorsal and caudal fins and in particular on the pectoral fins, the first ray of which is thickened, while the female has a distended belly and a swollen and pinkish vent.

Colour

The back is dark, olive to greenish-brown, the flanks are silvery but scales are marked with darker pigment on their posterior margin giving the appearance of a row of spots, and the belly is white to cream-yellow. Scale centres may reflect golden or yellowish tints. Upper scales are outlined with dark pigment to give a cross-hatching effect. The fins are grey-green, or grey to black, except the pelvics which resemble the belly colour. Peritoneum brownish black.

Size

Reputed to attain 1.6 m and about 50 kg in its native range; reports of weights up to 180 kg probably being exaggerations. Reaches 80 cm in the Tadjan River near Sari (A. Abdoli, pers. comm., 1995).

Distribution

The native distribution is in East Asia but it has been introduced to Iranian waters. Also introduced to Afghanistan, Pakistan, and Iraq (Shireman and Smith, 1983).

This species was first introduced in the 1950s according to Armantrout (1980) in the Anzali Mordab for vegetation control, adults surviving to the 1960s but no breeding population was established. Also introduced in 1966 from a hatchery in the Krasnodar region of the former U.S.S.R. and stocked in the Anzali Mordab (Anonymous, 1970b) and in October 1970 50,000 fingerlings from the U.S.S.R. were introduced to the Caspian Sea and Anzali Mordab (Griffiths et al., 1972). Three large fish (80 cm) were caught in January 1971 and believed to be from the October introduction and evidence of good growth although they may have been from an earlier stocking. It is reported from the Siah-Keshim Protected Region of the Anzali Mordab (Riazi, 1996), presumably recently stocked, and is stocked in a variety of reservoirs in the provinces of Gilan and Mazandaran but not as widely as silver carp (Hypophthalmichthys molitrix). It is pen cultured in Gomishan Reservoir, Mazandaran (Madbaygi, 1993b). Grass carp were introduced to Khuzestan in the 1970s to control vegetation in irrigation ditches. In April 1974, 1150 fish were released in the Dez Irrigation Project (Saadati, 1974). It is reported from Mahabad Dam (Abdi, 1999; www.mondialvet99.com, downloaded 31 May 2000), from the Safid River and Anzali Talab (Abbasi et al., 1999), from Lake Zaribar, Kordestan (Abzeeyan, 5(5):III, 1994), the Kor River in Fars (A. Alamdari, in litt., 1997), from the Sistan basin in Hamun Sabari, Hamun Kushk and the canal flowing into Chahnimeh (Ahmadi and Wossughi, 1988; Mansoori, 1994; J. Holčík, in litt., 1996), from the Haft Barm lakes near Shiraz in 1984 although these later disappeared, possibly eaten by introduced Sander lucioperca (Petr, 1987). As escapees from a fish farm, they have been found in Lake Famur. Also recorded from the Gorgan, Tajan and Safid rivers, and the Anzali Mordab (Kiabi et al., 1999), and it is mapped from the Kor, Kerman-Na'in, Hormuz, Dasht-e Lut and Sistan basins without exact localities; the Kashaf River in the Tedzhen River basin; middle reaches of the Atrak River, lower reaches of the Gorgan, Neka, Babol, Heraz and Safid rivers and in the Anzali Mordab, all in the Caspian Sea basin, the middle to lower Talkheh and lower Zarrineh rivers in the Lake Orumiyeh basin, the middle to lower Abhar-Shur and Qom River in the Namak Lake basin, the middle to lower Zayandeh River in the Esfahan basin; the lower Karun and Jarrahi rivers in the Tigris River basin, and the lower Jovein and middle Kal Shur rivers in the Dasht-e Kavir basin (Abdoli, 2000).

It was introduced to the Soviet Caspian Sea in 1970-1974 where small populations became established in the Terek River and the Volga delta and to the Karakum Canal and Kopetdag Reservoir of Turkmenistan near the Iranian border (Baltz, 1991; Shakirova and Sukhanova, 1994; Sal'nikov, 1995; Opuszynski and Shireman, 1995).

Grass carp could establish breeding populations in the large rivers of southern Iran and Iraq if the environment proves favourable and there is enough uninterrupted river flow for eggs to hatch.

Zoogeography

This species is an exotic in Iran and has a native range from the Amur River basin of Siberia south to southern China. It has been widely introduced around the world for vegetation control.

Habitat

The natural habitat is large rivers but this species adapts easily to pond culture. Grass carp can live in the Caspian Sea at salinities of 5-8‰ although a few are found at 10-12‰. They enter rivers to spawn (Abdusamodov, 1986). Temperatures in the range 0-41°C and low oxygen concentrations (0.2 mg/l) are tolerated by this species as is high turbidity. Fry have an upper lethal temperature range of 33-41°C and temperatures greater than 38°C are lethal for adults. pH range is 5.0-9.0. Adults prefer densely vegetated inshore areas with depths of 1-3 m. Adults leave the river after spawning and feed in lakes, reservoirs and on floodplains, returning to the river in autumn to overwinter in deep holes separate from the juveniles. Young hide in vegetation of lakes, reservoirs and floodplains. Juveniles may migrate as much as 1000 km up- or downstream from the original spawning site in their native habitat. Young fish overwinter in deep holes in river beds.

Age and growth

Growth rate in Khuzestan canals was 1.8 g per day while in ponds growth was 6.6 g per day when fed alfalfa during a 5-month growing season from April to September (Saadati, 1974; Behnke, 1975a). Males begin to mature at 4 years and females at 5 years in the Terek River of Dagestan (Abdusamadov, 1986). Maturity is attained at 6-10 years in the Amur River, the native habitat, and as early as 10 months in Malaysia. Life span is over 33 years. Growth rate in this species is perhaps greater than in any other fish. Growth to 1 kg in the first year of life and 2-3 kg per year thereafter in temperate areas is very high; in tropical areas a 20 g fingerling can reach 8.5 kg in 1 year. Rates of 10-22 g per day have been reported in various areas of the world depending on local conditions.

Food

Grass carp are herbivores, except for quite small fish (20 mm total length or less) which consume zooplankton. In Khuzestan, the grass carp prefers to eat Potamogeton spp. and Alisma gramineum to Chara and Cladophora (Saadati, 1974). The grass carp can consume 100-150% of its body weight per day of aquatic vegetation. Peak feeding occurs at 25-30°C but food is taken in the range 15-35°C. Grass carp stocked in the Anzali Mordab and fish farms of the Caspian Sea basin consume fresh Azolla, an introduced fern. Grass carp stocked at 800/ha consume 400-500 kg of Azolla daily gaining 800-1200 g in 5 months. In China this species is known to eat grass, leaves, small fishes, insects and other items in addition to aquatic vegetation or when such vegetation is in low supply. About half the plant food passing through the digestive system is undigested and large quantities of plant material must be eaten to sustain life. This consumption rate is the reason for its success at aquatic vegetation control (Greenfield, 1973). Grass carp overwinter without feeding.

Reproduction

A spawning migration to a large river takes place at about 15-17°C water temperature. The female swims in the centre of the river at the surface accompanied by 2-3 males, they roll and rub their bodies together and often jump out of the water. A male prods the female's body to stimulate egg release and leans closely to one side. Eggs are semi-buoyant and require a slow and steady current to keep them off the bottom (minimum water velocity of 0.23 m/second or more to support them and allow hatching; this is found in large rivers where the eggs hatch as they drift downstream; at 20°C and a not unusual velocity of 1.2 m/second, hatching requires 180 km of river). Temperatures should be above 20°C and preferably 21-25°C, or 26-30°C in another source. At these temperatures hatching takes about 40 hours. Flow rates should be 0.7-1.8 m/sec. Spawning occurs after heavy rain in rising rivers, when turbidity may reduce predatory attacks on the semi-pelagic eggs (Greenfield, 1973). This regime is also required for newly hatched fry and such conditions are rare outside their native habitat. In the Terek River of the Caspian Sea basin, the spawning migration begins in mid-April at water temperatures of 15-17°C and continues until August although numbers begin to decrease from the end of May. Spawning takes place after a sharp rise in water level and current speed. Eggs are first found in the drift in the second week of June and hatch 34-70 hours later depending on temperature. Some larvae reach rice fields and live there until autumn when the fields dry up, some being lost, others migrating. Other larvae are carried into the Caspian Sea where they are sensitive to the prevailing salinity at 1-1.5 days old (Abdusamadov, 1986). Up to 100,000 eggs are laid at one time (Greenfield, 1973) and in the Terek River fecundity reaches 1,230,700 eggs (Abdusamadov, 1986). Absolute fecundity may reach 2 million eggs. Eggs are up to 2.5 mm in diameter before fertilisation and are greyish-blue to bright orange. In water they swell to over 5.3 mm in 2 hours, becoming buoyant in flowing water.

Parasites and predators

Mokhayer (1976b) reports the cestode Bothriocephalus gowkongensis and the acanthocephalan Pomphorhynchus perforator. Red-sore disease is reported from fish pond grass carp in Iran by Razavilar et al. (1981). It is caused by a bacterium Aeromonas hydrophila and treatment was unsuccessful. Mokhayer (1989) records metacercariae of the eye fluke, Diplostomum spathaceum from this species in Iran, which can cause complete blindness and death in commercially important species, as well as shedder scales (sic), Echinochasmus perfoliatus. Jalali and Molnár (1990b) record the monogenean Dactylogyrus lamellatus from this species at fish farms in Iran. Viral haemorrhagic disease has been reported from grass carp in Iran (Iranian Fisheries Research and Training Organization Newsletter, 6:6, 1994; 9:6, 1995). Pond-cultured grass carp were found to be infected by the tapeworm Bothriocephalus, with 70-80 parasites causing intestinal obstruction and lowered haemoglobin, haematocrit and erythrocyte values (Esmaeli and Abbasi, 1996). Esmaeli and Peighan (1997) record an Aeromonas-like bacteria from grass carp in Khuzestan Province. Ebrahimzadeh Mousavi and Khosravi (1999; www.mondialvet99.com, downloaded 31 May 2000) record the toxigenic fungi Aspergillus flavus, Alternaria, Penicillium and Fusarium from this species and the pond water at a fish farm in northern Iran. The crustacean parasite Lernaea elegans is reported from this species in the Mahabad Dam reservoir (Abdi, 1999; www.mondialvet99.com, downloaded 31 May 2000). The intestinal helminth Bothriocephalus gowkongensis was recorded from this species on fish farms in West Azarbayjan Province (Azarvandi et al., 1999). Naem et al. (2002) found the following parasites on the gills of this species from the western branch of the Safid River, namely the protozoan Ichthyophthirius multifilis, a copepod crustacean Lernaea sp., monogenean trematodes Dactylogyrus lamellatus, D. ctenopharyngodonis, and Gyrodactylus sp.. Jalali et al. (2002) and Jalali and Barzegar (2006) record Diplostomum spathaceum and Dactylogyrus lamellatus from this species in Lake Zarivar. Esmaeili et al. (2005) found a Flavobacterium columnaris-like bacterium on grass carp form Khuzestan fish ponds, suspected of either causing a 40% mortality or being a secondary factor in the fish kill. Pazooki et al. (2005) record Ergasilus peregrinus from this species in waterbodies of Zanjan Province. Araghi Soureh and Jalali Jafari (2006) recorded Dactylogyrus lamellatus from this species in the Mahabad River of the Lake Orumiyeh basin. Barzegar and Jalali (2006) report parasites in this species from Kaftar Lake as Trichodina sp., Dactylogyrus lamellatus, Lernaea cyprinacea and Diplostomum spathaceum.

Any piscivore will take this species.

Economic importance

This species has been introduced to Iran to control aquatic weeds in drainage and irrigation canals as an alternative to using polluting chemicals or mechanical removal. In some countries at is sought after by anglers. Grass carp may also help to control the snail-carried, human disease schistosomiasis, since the vegetation on which the snails live is severely reduced. They are also a food fish which relies on food sources not available to native fish (few fish consume whole plants). Grass carp consume vegetation at a rate of 100:1, i.e. for every 1 kg increase in grass carp biomass 100 kg of vegetation is consumed. Removal rates may exceed this figure since leaves are bitten off and branches clipped with not all of it being consumed. In the Dez Irrigation Project large amounts of this vegetation were removed daily from screens in test sections. Stocking in the Dez Irrigation Project in Khuzestan showed a removal rate approximately the same as mechanical control (Saadati, 1974; Behnke, 1975a). During a 5 month period the grass carp controlled 250 tons of aquatic vegetation per hectare. The fish preferred plant species which blocked the canals (Potamogeton spp. and Alisma gramineum) rather than those which grew close to the substrate (Chara and Cladophora) and did not interfere with water flow. Shireman and Smith (1983) give details on artificial propagation of this species.

Esmaeilzadeh et al. (2004) studied the nutrient composition and marinade qualities of this fish in Iran and compared them to those for safid mahi (Rutilus frisii) and found them to be preferable according to the organoleptic properties. The marinades could be stored for 6 months at 10ºC.

Fish farming of this species in Sistan was discontinued as its consumption of vegetation was reducing food for other species (www.netiran.com, downloaded 28 February 2005).

Holčík and Oláh (1992) report a catch of 315 kg in the Anzali Mordab in 1990. However Iran acounts for almost all the production of grass carp in the Near East and North Africa (4378 tonnes in 1994) (Food and Agriculture Organization, Fisheries Department, 1996). The aquaculture production in 1995 was 3942 tonnes (Bartley and Rana, 1998b). Grass carp sold for about U.S.$2.00/kg in 1995 (Rana and Bartley, 1998a). Marjan Iran Company was selling 1500-2000 g fish for U.S.$2.10/kg in August 2003 (http://groups.yahoo.com/groups/hilsa/message/25).

The inland waters of Turkmenistan had catches of 23 to 29.7 tonnes for the years 1971-1974 and a catch of 76 tonnes in 1970 when a ban on taking phytophagous fish was lifted.

Greenfield (1973) reviews the advantages and disadvantages of using this species as a weed control agent in the U.S.A. and Charyev (1984) in the Kara-kum Canal in Turkmenistan. Destruction of habitat for fishes and waterfowl, competition with native species and introduction of exotic diseases and parasites are all problems once this fish escapes into a main river suitable for reproduction. Their destruction of plants may interfere with waterfowl management, destroy breeding grounds for other species and facilitate the attacks of predators. Ideally triploids, produced by cold or warm shocks or by hydrostatic pressure on fertilised eggs, should be used initially as they cannot reproduce (Clugston and Shireman, 1987). However the chromosome number of each fish must be checked (by electronically measuring the volume of a red blood cell nucleus) as the process is not 100% effective. Grass carp are reproducing naturally in the Kara-kum Canal, vegetation is controlled, fish stocks have increased and some reduction of mosquitos has been obtained. However the ecosystem has been changed, spawning grounds of commercial species threatened, undesirable species have been introduced accidentally, and reduction in vegetation affects water quality. Grass carp are best used in restricted areas where improved flow and reduced mosquito populations are required but where there is no commercial fishery (Charyev, 1984).

The grass carp has a short gut and about half the plant material eaten is released to enrich the water and promote algal blooms. Oxygen levels and water clarity are reduced. The removal of plants can remove food sources for other fishes, shelter and spawning substrate. Additionally, as noted, the triploid treatment is not always effective and the species can become established.

Iran has had problems with disease outbreaks and poor survival of fingerlings which has led to production problems (Shehadeh, 1997).

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in aquaculture, as food and in textbooks. There are numerous studies on this species as an experimental fish and in relation to aquaculture. Some Iranian studies include Morovvati et al. (2006) on seasonal changes of pronephros lymphoid tissue; Nahavandi et al. (2006) on the chemiluminescent response to determine the effect of various concentrations of diazinon, an organophosphate, on phagocytosis in order to measure immunity after exposure to this toxin; Pourgholam et al. (2006) and Sharifpour et al. (2006) on the toxicity and histopathological effects of diazinon; Sharifpour et al. (2006) on the sub-lethal effects  of diazinon on various organs; Pourgholam et al. (2006) on the toxicity of diazinon and the effects of sub-lethal concentrations on haematological and biochemical indices; etc.

Conservation

No conservation is required for this exotic species. Krasznai (1987) and Petr (1987) give details of fish farms propagating this species in Iran. For example, 10 million were produced in the Safid Rud Fish Farm in 1986. 20 million carp, silver carp and grass carp fingerlings were produced in the Shahid Rajaae Hatchery in Sari for release across Iran in reservoirs and dams (Abzeeyan, Tehran, 4(7):VII, 1993). Feeding and growth studies on this species have also been carried out on this species in the Shaid Rajaee Hatchery (Ahmadi and Rezai, 1998). Experiments on induction of triploidy have been carried out in Iran using cold and heat shocks (M. Hassanzadehsaber, M. Pourkazemi, M. R. Nowruzfashkhami and A. Ghanaatparast (www.meeresschule.com/cgi-bin/abstracts/gastbuch.asp, downloaded 17 January2005).

Further work

Studies on the interactions of this species and native Iranian taxa should be carried out and introductions carefully controlled and monitored.

Sources

Shireman and Smith (1983) give a summary of the biology of this species. There is an extensive literature on herbivorous fishes, a recent book being Opuszynski and Shireman (1995), which has sections on grass carp. Gholipour (1996) has an account in Farsi.

Comparative material: BC65-381, 2, 95.4-98.8 mm standard length, Singapore, fish ponds (no other locality data).

Genus Cyprinion
Heckel, 1843

Scaphiodon Heckel, 1843 has been used for Cyprinion and Capoeta species in Southwest Asia.

Taki (1975) related members of this genus to a common ancestor with Onychostoma Günther, 1896, a Chinese and southeast Asian genus although Li et al. (2008) found this lineage to be unsupported on DNA evidence. Howes (1982) synonymises Semiplotus Bleeker, 1859, a genus found from Nepal to Viet Nam, and Scaphiodonichthys Vinciguerra, 1890, a genus from Indochina, with Cyprinion and refuted Taki's (1975) view using osteological characters, particularly of the jaws. Howes (1982) considers that Cyprinion cannot be defined on any uniquely derived characters. Krupp (1983) considers Howes' revision as unsatisfactory for the reasons that type specimens were not examined, relationships are based on jaw anatomy and other characters are largely excluded, variability of osteological characters within a species are largely unknown, and synapomorphies are not unequivocal. Bănărescu (1992b) and Banarescu and Herzig-Straschil (1995) regard Semiplotus as a distinct genus but probably related to Cyprinion. They comment that Semiplotus differs sharply from Cyprinion s.s. in the absence of barbels, a higher number of branched dorsal fin rays (20 or more), and in a lower number of branched anal fin rays (5 as in most related genera rather than the unusual 7 in Cyprinion). Scaphiodonichthys has 2 pairs of barbels (only 1 in Cyprinion), and 5 branched anal fin rays as well as differing from both Cyprinion and Semiplotus by having the lateral line closer to the ventral margin of the caudal peduncle and divergent rather than parallel striae on the scales. These latter 2 characters justify generic separation of Scaphiodonichthys. Bănărescu (1997) considers Scaphiodonichthys as valid and not a synonym of Cyprinion. Characters used by others to define Cyprinion such as expansion of the proximal part of the pelvic fin rays, interpelvic papillate flaps (Banister and Clarke, 1977) and a naked predorsal ridge (Mirza, 1969) do not occur in all species in this genus. If Semiplotus is included in Cyprinion then several osteological structures, particularly a synarthritic dentary joint, are uniquely derived or synapomorphic.

In the absence of a detailed revision, I have retained species within Cyprinion as the most familiar name in use in Southwest Asia for these fishes. Cyprinion s.s. is found from the Indus River basin west to the Arabian Peninsula and the Tigris-Euphrates basin but excluding northern drainages such as the Lake Orumiyeh, Caspian Sea and Hari River basins and excluding the westernmost edge of Southwest Asia such as the Jordan River basin and coastal drainages of Israel.

The genus Cyprinion is currently under revision by Florian Wicker at the Senckenberg Museum, Frankfurt and the status of the following species may undergo some changes.

Saadati (1977:45) refers to a new and undescribed Cyprinion species from Lar in southern Iran but the fish are Carasobarbus luteus.

A thorough study of the systematics of this genus in Iran depends to some degree on material from other areas which is not readily available, on large series of well-preserved adult specimens, and analyses which demonstrate consistency in characters used to define species. These conditions have not been achieved thus far in any studies undertaken and given the wide distribution and individual variation shown by Cyprinion species an adequate understanding of the species composition is not entirely possible.

This genus is characterised by a moderate sized, compressed body, a thick and blunt snout, an inferior mouth with a straight, crescentic or arched shape and a sharp horny edge to the lower jaw (which may fall off in preserved specimens), 1 pair of small barbels at the mouth corner, the last dorsal fin unbranched ray is thickened and bears weak to strong serrations (highly variable between individuals within a species and not a good character in species definitions), the dorsal fin is long (up to 16 branched rays) and the anal fin short (typically 7 branched rays), a ridge in front of the dorsal fin is formed internally from fused pterygiophores and lacks scales externally, pharyngeal teeth are in 3 rows and are compressed and spoon-shaped, scales large to moderate in size (lateral line counts (31-45), breast and belly scales may be absent (individually variable and not a good character), scale radii are restricted to the posterior field, peritoneum black, and gut very long and coiled (several times body length).

Cyprinion kais
Heckel, 1843


Dez River at Dez Wildlife Refuge, 24 April 2008, courtesy A. Mahjoor Azad

Common names

butak-e dehan kuchek, بوتك (= botak), butak dahan kuchek, butok, لوتك (= lotak), zanbour, زنبور دهان كوچك (= zanbour dahan kuchek).

[bunni saghir, bnaini; kais at Aleppo (= Haleb, Syria), hence the scientific name, all in Arabic; kais kingfish].

Systematics

Cyprinion Cypris Heckel, 1843 is a synonym, being a juvenile with keratinization of the lower jaw incomplete according to Howes (1982), although he did not examine the types. Krupp (1985c) and Banarescu and Herzig-Straschil (1995) agree with this synonymy. Berg (1949) placed C. kais (and C. cypris) in C. macrostomum, as the position of the dorsal fin in relation to the pelvic fins was variable in these fishes and not sufficient to warrant species status as Heckel (1843) stated in describing these species.

The type localities for Cyprinion Kais are "Aleppo" and "Mossul" and for Cyprinion Cypris the "Tigris bei Mossul" (Heckel, 1843b).

The syntypes of C. kais are in the Naturhistorisches Museum Wien comprising 3 fish in NMW 52801 (paralectotypes) and measuring 68.5-97.3 mm standard length, 2 fish in NMW 52802 measuring 120.6-164.3 mm standard length, and 2 fish in NMW 52803 (paralectotypes) measuring 153.4-154.2 mm standard length, the smaller of these being designated as the lectotype by F. Krupp in 1984. Eschmeyer et al. (1996) list possible syntypes in the Rijksmuseum van Natuurlijke Historie, Leiden under RMNH 2485 (2 fish, formerly NMW) and RMNH 2489 (1), and 1 syntype in the Senckenberg Museum Frankfurt (SMF 134, formerly NMW). The catalogue in Vienna lists 5 specimens.

A syntype of C. cypris is in the Senckenberg Museum Frankfurt (SMF 849, formerly NMW) (F. Krupp, pers. comm., 1985). Two syntypes, 63.5-106.2 mm standard length are under SMF 849, the larger one designated as a paralectotype (March 2007). Ten syntypes are in the Naturhistorisches Museum Wien (NMW 52804) measuring 51.2-115.1 mm standard length, the largest being designated as the lectotype (however Banarescu and Herzig-Straschil (1995) give 44.1-110.0 mm standard length for these 10 fish with one at 99.8 mm standard length as lectotype as selected by F. Krupp in 1984). Another specimen, 110.5 mm standard length, may also be a syntype (NMW 52800); and also NMW 59508, a dried specimen (Eschmeyer et al., 1996). The catalogue in Vienna lists 6 fish in alcohol and 1 fish stuffed.

Key characters

Mouth shape is distinctive. It is small and semicircular with a width about the size of the eye diameter and has large lateral lobes (= lower lips)(Kafuku, 1969). The cartilaginous sheath is thickened between the corners of the mouth and is rounded posteriorly with a distinct margin. The cartilage can form a tooth-like structure protruding anteriorly from the lower lip. The mouth in C. macrostomum is wider, arched and lacks the lateral lobes (see also illustrations in Kafuku (1969), Krupp (1985c) and Banarescu and Herzig-Straschil (1995)). These latter authors have the width of the mouth opening as only 13.5-22.0% of the head length (22.0-27.0% in C. macrostomum) for adult fish and the height of the arch or mouth opening (a line perpendicular from a line between the mouth corners to the tip of the lower jaw) 48-80% of the mouth width (29-47% in C. macrostomum), i.e. the mouth is narrower and more arched in C. kais. On this character, therefore, the two species can be distinguished as adults but there is potential for confusion in young fish. A single specimen identified as C. kais on the basis of mouth shape from the Dalaki River of Iran had values of 23.2% and 47.4% which are arguably C. macrostomum values. This specimen has a protruding tooth-like edge to the lower jaw in a u-shaped mouth with well-developed lips posterior to the "tooth".

The intestine is shorter and less complexly coiled in this species and the mean number of gill rakers is less in contrast to C. macrostomum (Kafuku, 1969). The back is higher and more curved, the eyes are larger and the anal fin is more posterior, in addition to the mouth shape (Heckel, 1843b). The dorsal fin origin arises over that of the pelvic fins (Heckel, 1846-1849a). The edge of the dorsal fin is more notched in C. kais than in C. macrostomum (the length of the fourth branched ray is 48-62% of the length of the first ray as opposed to 55-79% in C. macrostomum, with extreme values overlapping, according to Banarescu and Herzig-Straschil (1995)).

The form of the pharyngeal teeth is different from C. macrostomum (see Krupp (1985c) for illustrations where kais has hooked tips and macrostomum does not), there are fewer gill rakers (8-12 on the lower arch in kais, 12-16 in macrostomum), on average there are fewer dorsal fin rays, the last unbranched dorsal fin ray is longer, and interorbital width is smaller. However sample sizes in some studies are small (in Kafuku (1969) only 5 fish of each species were examined), morphometric characters are notoriously size-dependent, gill raker counts are also size dependent, and even pharyngeal tooth form varies with age (small macrostomum have hooked tips). C. kais may well be a good species but a wide-ranging comparison of adults and young and of localities is needed and material from Iran is scarce or equivocal. Further discussion is under C. macrostomum.

Morphology

Dorsal fin with 4 unbranched and 12-16 branched rays, anal fin with 3 unbranched and 7 branched rays. The dorsal fin has the last unbranched developed as a spine with strong teeth except at the extreme tip which is thin and flexible. Pectoral fin with 14-18 branched rays, and pelvic fin with 8-9 branched rays. Lateral line with 36-43 scales. The belly is scaled. There is a well-developed pelvic axillary scale. Scales have a subcentral anterior focus, fine circuli, few posterior radii and no or very few anterior radii. Total gill rakers 10-15, short and reaching the raker below when appressed. Rakers are absent on the anterior arch where there are only tubercles. Pharyngeal teeth 2,3,4-4,3,2, with variants 2,3,5-5,3,2 and 2,3,5-4,3,2, spoon-shaped with a small hook at the tip.

Meristics for Iranian material: dorsal fin branched rays 12(2), 13(2), 14(1) or 15(1); branched anal fin rays 7 (6); branched pectoral fin rays 14(1), 15(2), 16(2) or 17(1); branched pelvic rays 8(4) or 9(2); lateral line scales 38(5) or 39(1); and total gill rakers 12(2), 13(3) or 14(1).

Sexual dimorphism

Tuberculation in a 103.5 mm standard length specimen consisted of ca. 20 tubercles restricted to the area over the lachrymal bone. A specimen 147.5 mm standard length had small to minute tubercles in front of the eye, under the eye, on the mid-preoperculum and on the mid-operculum. Curiously the individual small tubercles on the operculum were connected by thin lines of horny tissue.

Colour

Overall colour is silvery to yellowish-white with the back grey-brown and the lower surfaces a lemon yellow. The lower jaw margin is a glossy yellow. The fish shown above may represent a spawning colouration, not seen in all specimens. The pelvic fins are a bright orange-red, the pectorals paler. Some fish have a less strong colour in the pelvic than in the anal fin. The anal fin is yellow, to orange or greenish, distally black and anteriorly most orange. The caudal fin has light orange to greenish tints. The dorsal fin is black with a yellow-tinged base becoming anteriorly reddish. In preserved fish, there is some concentration of pigment above and below each lateral line pore, scales on the back and upper flank are outlined with pigment, and there is some concentration of pigment into a few to moderate number of diffuse spots on the uppermost flank and back midline. The leading edge of the dorsal fin is very dark (but may be light), dorsal fin membranes are dark, anal fin membranes also dark but to a lesser extent, and the caudal, pectoral and pelvic fins have pigment lining the rays. Peritoneum black.

Size

Attains 21.5 cm total length.

Distribution

This species is found in the Tigris-Euphrates and Quwaiq basins. Abdoli (2000) maps the Jarrahi, Karun, middle to lower Dez, and Karkheh up to the Simarreh rivers of the Tigris River basin. It is also found in the Gulf basin, although rare, and specimens from the sugar cane fields of Khuzestan were seen in 2000 (personal observations, B. W. Coad).

Zoogeography

Zoogeographical comments are under the genus above.

Habitat

This species is recorded from a variety of habitats as listed above and is also known to inhabit canals but nothing is known of its environmental requirements.

Age and growth

Unknown.

Food

Gut contents are filamentous algae in the one specimen examined. Diet may be similar to Cyprinion macrostomus. Curiously, the mouth structure resembles that of the unrelated cutlips minnow, Exoglossum maxillingua (Le Sueur, 1817), from North America. This species feeds on insect larvae, with some molluscs and worms. Food is scraped from the bottom or poked out of crevices using the shovel-like lower jaw. Sand is also taken in and spat out, presumably after food items are extracted. The cutlips also picks out the eyes of other fishes in confined areas (Coad et al., 1995).

Reproduction

Generally unknown. Ünlü (2006) gives age at first maturity as 2 years in the Turkish Tigris River with spawning over sand, stones and gravel in May-June.

Parasites and predators

None reported from Iran.

Economic importance

None.

Conservation

This minnow appears to be rare, or at least is rarely collected, in Iran. Cyprinion macrostomum is much more common and is taken in most seine hauls in streams and rivers. The distribution and population numbers are unknown. Endangered in Turkey (Fricke et al., 2007).

Further work

The biology of this species needs to be investigated and the use of the peculiar jaw structure ascertained. Its great rarity, at least in Iran, leads to the suspicion that it may be a developmental anomaly of Cyprinion macrostomum - the few specimens at hand don't permit a detailed study of characters other than the strikingly different jaw (see comments under C. macrostomum and also above). Development of pharyngeal teeth, gill raker numbers, complexity of gut coils and morphometric characters are all size dependent and show individual and populational variations not analysable here.

Sources

Type material: See above, Cyprinion kais (NMW 52801, 52802 and 52803) and C. cypris (NMW 52804).

Iranian material: CMNFI 1993-0141, 1, 66.3 mm standard length, Bushehr, Dalaki River (29º28'N, 51º15'E); CMNFI 2008-0169, 5, 80.4-98.2 mm standard length, Khuzestan, irrigation ditch in sugar cane fields (31º58'42"N, 48º31'07"E); ZSM 25715, 2, 34.1-65.3 mm standard length, Khuzestan, Dez River at Harmaleh (31º57'N, 48º34'E).

Comparative material: BM(NH) 1920.3.3:50, 1, 83.6 mm standard length, Iraq, Basrah (30º30'N, 47º47'E); BM(NH) 1920.3.3:94-115, 40, 65.3-92.4 mm standard length, Iraq, Basrah (30º30'N, 47º47'E); BM(NH) 1931.12.21:3, 1, 129.8 mm standard length, Iraq, Mosul (36º20'N, 43º08'E); BM(NH) 1974.2.22:115-120, 5, 90.6-147.9 mm standard length, Iraq, Mosul (36º20'N, 43º08'E); BM(NH) 1974.2.22:1105, 1, 115.6 mm standard length, Iraq, Mosul (36º20'N, 43º08'E); BM(NH) 1974.2.22:1106, 1, 101.4 mm standard length, Iraq, Fao (29º58'N, 48º29'E); BM(NH) 1974.2.22:1214-1255 (in part), Iraq, Khalis (33º49'N, 44º32'E); BM(NH) 1984.4.18:30, 63.4 mm standard length, Iraq, Kut Hiwa (no other locality data); FMNH 51229, 1, 103.5 mm standard length, Iraq, Diyala River, 12 miles east of Baghdad (no other locality data); FMNH 51230, 6, 42.9-60.5 mm standard length, Iraq, Diyala River, 12 miles east of Baghdad (no other locality data); FMNH 51231, 2, 64.0-64.8 mm standard length, Iraq, Diyala River, 12 miles east of Baghdad (no other locality data); uncatalogued, 5, 49.1-66.7 mm standard length, Iraq, Shatt al Arab (no other locality data); uncatalogued, 1, 107.2 mm standard length, Turkey, Euphrates River 20 km west of Erzurum (ca. 41º03'N, ca. 39º55'E).

Cyprinion macrostomum
Heckel, 1843

Common names

بوتك (= botak); butok; لوتك (= lotak); butak-e dehan (or dahan) buzorg in Khuzestan; galuk (Mokhayer (1981c); kapour; zanbour (= bee) in Khuzestan and Boyer Ahmadi-ye Sardsir va Kohkiluyeh provinces; زنبور دهان بزرگ (= zanbour dahan bozorg); ?tumbuek (= hunting horn, possible name from Heckel (1843b)).

[hmarriya sefra or himriya sefra, surrah masih, kais at Aleppo (= Haleb, Syria) but see above species (Heckel, 1843b); dombok or dumbek at Mosul (= solid or compact flesh, a good source of food, according to Heckel (1843b)); dunbuk kabir al-fam, benayne, all preceding in Arabic; large-mouthed barb, Tigris kingfish].

Systematics

Originally spelt macrostomus but correctly macrostomum (Berg, 1949). Cyprinion neglectus Heckel, 1849 from the "Tigris bei Mossul" is a synonym (Krupp, 1985c; Banarescu and Herzig-Straschil, 1995). Howes (1982) considered that Cyprinion tenuiradius (q.v.) was only a "variant" of this species but did not examine any material. Berg (1949) places C. kais (q.v.) in the synonymy of this species along with C. cypris (see C. kais).

The type locality of Cyprinion macrostomus is given by Heckel (1843b) as "Aleppo" and "Mossul". Krupp (1985c) lists 5 syntypes from Aleppo, 81-133 mm standard length in the Naturhistorisches Museum Wien (NMW 52805), the largest being selected as the lectotype (hence Aleppo is the type locality as designated by the publication of Banarescu and Herzig-Straschil (1995)). One syntype from Aleppo, 83 mm standard length, is in the Senckenberg Museum Frankfurt (SMF 70, formerly NMW; Eschmeyer et al. (1996) give SMF 870) and 4 syntypes from Mosul, 58-124 mm standard length are in the Naturhistorisches Museum Wien (NMW 52806). My measurements are 82.1-135.0 mm standard length for NMW 52805 and 59.1-126.2 mm standard length for NMW 52806. Another syntype is a dried specimen NMW 52503, and the Rijksmuseum van Natuurlijke Historie, Leiden has 1 syntype under RMNH 2487, formerly NMW) and 1 syntype under RMNH 2488, formerly NMW). The catalogue in Vienna lists 4 specimens.

Seven syntypes of Cyprinion neglectus from Mosul measure 54-131 mm standard length (NMW 52807), the largest being selected as the lectotype (Krupp, 1985c). My measurements are 53.3-131.9 mm standard length (Banarescu and Herzig-Straschil (1995) have 53.1-128.2 mm standard length). All material was collected by Th. Kotschy in 1842 for Aleppo and 1843 for Mosul. The catalogue in Vienna lists only 2 specimens under this name.

?Check lengths against data sheets

Key characters

Distinguished from C. kais by mouth and dorsal fin ray characters as described under that species, by having more gill rakers and a longer and more coiled intestine (Kafuku, 1969). The dorsal fin origin is in front of that of the pelvic fins (Heckel, 1846-1849a). See discussion under C. tenuiradius for distinction from that taxon.

Morphology

Dorsal fin with 4 unbranched and 12-17 branched rays (usually 14-15 according to Banarescu and Herzig-Straschil (1995) but 77% of fish in Iran are 13-14, see below). The last dorsal fin unbranched ray is strong and serrated to the tip. The anal fin has 3 unbranched and 6-7, usually 7, branched rays. In Iranian specimens, 96.1% of 127 fish have 7 rays, the remainder 6 rays. Pectoral fin branched rays are 12-17 and pelvic fin branched rays 7-9, usually 8. Lateral line scales 33-45 (usually 41-44 according to Banarescu and Herzig-Straschil, 1995) but a broader range in Iran, see below). The breast is covered with scales. The pelvic axillary scale is very elongate. Scales are squarish, being deeper than long, often with parallel dorsal and ventral margins (or rounded margins). The anterior margin has a marked central protuberance and the posterior margin is rounded. Radii are numerous on the posterior field and circuli are fine and numerous. The posterior field circuli break into "bubbles". The focus is subcentral anterior. Gill rakers 16-17, on the lower arm 12-16, in the literature but a much wider range in total rakers in Iran (see below). Rakers are short and only touch the raker below or a little further when appressed. Pharyngeal teeth 2,3,5-5,3,2, 2,3,4-4,3,2, and variations on 4 or 5 main row teeth. Teeth are spatulate with broad, flattened crowns. The tips of teeth are slightly hooked in small fish. The most anterior tooth in the main row may be very small or absent (or incompletely ossified and hard to distinguish). The gut is very elongate with complex coils. In small fish, the upper lip is not covered with a fold of the snout as in large fish. Also the gut is not as coiled in young fish as in adults. Chromosome number is 2n=48 (Ünlü et al., 1997).

Meristics for Iranian fish from the Tigris River basin: branched dorsal fin rays 12(4), 13(43), 14(52), 15(26) or 16(3)(mean = 13.9, S.D. = 0.861); branched pectoral fin rays 14(3), 15(44), 16(57) or 17(25)(mean = 15.8, S.D. = 0.771); branched pelvic fin rays 7(7), 8(121) or 9(1)( mean = 8.0, S.D. = 0.246); lateral line scales 33(3), 34(1), 35(12), 36(11), 37(3), 38(11), 39(29), 40(31), 41(25), 42(2) or 45(1)(mean = 38.8, S.D. = 2.211); total gill rakers 13(3), 14(8), 15(15), 16(23), 17(15), 18(24), 19(17), 20(14) or 21(6) (mean = 17.3, S.D. = 2.022); pharyngeal teeth 2,3,5-5,3,2(17), 2,3,4-5,3,2(8), 2,3,5-4,3,2(3), or 2,3,4-4,3,2(2); and total vertebrae ?.

The mouth is usually transverse or slightly arched and usually has a horny covering. Small fish have a crescentic mouth. A wide range of mouth arching is seen in fish of varying sizes and even in fish of the same size and locality of capture. Banarescu and Herzig-Straschil (1995) note that the syntypes of Cyprinion neglectus have a mouth arch which is more curved and not as wide, somewhat intermediate between C. macrostomum and C. kais, being closer to the former. This variation is attributed to the material possibly being from some tributary of the Tigris River, or from isolated ponds, where introgression with C. kais took place. It may well be that variation in mouth shape is more marked than limited sample sizes would indicate. Certainly in smaller fish, e.g. in 20 specimens of C. macrostomum (38.5-54.0 mm SL) examined by me from Iran, values for mouth width and depth as measured in Banarescu and Herzig-Straschil (1995) are not as clear cut and there is a variable developmental gradient in mouth shape. Mouth "height" as a % of width was 29.2-53.8 and width as % of head length was 22.1-36.6. Banarescu and Herzig-Straschil (1995) give "height" as 19-31% of width and width as 26-44% head length for macrostomum and 48-80% and 13.5-22.0% respectively for kais. Large macrostomum and kais (>100 mm SL) can be distinguished on mouth shape but not smaller specimens which bridge the gap between the two species. The possibility that kais is a developmental anomaly of macrostomum, retaining juvenile features, should be investigated.

Sexual dimorphism

Mature males have large tubercles on the snout in a broad band below the nostril level, extending back under the eye and breaking up into a few tubercles on the operculum. There is a large tubercle between the nostril and the eye. Fine tubercles are scattered over the top of the head. Three tubercles are found in rows on the first branched pectoral fin ray and very strong tubercles line each anal fin branched ray in single file. The anterior pelvic fin rays have the occasional 1-2 tubercles or a row of tubercles. Dorsal and caudal fin rays have fine tubercles, much smaller than those on the anal fin. Mid and posterior flank scales have 1-3 small tubercles, variably arranged on the exposed scale.

Colour

The back is bluish-grey to bluish-black or brown, flanks silvery or silvery-yellow and the belly whitish with silvery tints. The upper head is light brown. Scales are outlined with dark pigment and the anterior exposed scale base is darkened. The cleithrum area is pink or orange in some fish with pink or orange spots on up to 5 rows of flank scales but mostly along the anterior lateral line. Fish from a saline stream in Khuzestan had a pale-pink cleithrum and lateral line spots. There is a reddish-yellow spot at the base of the pectoral and pelvic fins. The pectoral, pelvic, anal and caudal fins are yellowish to pinkish or orange proximally and blackish distally. The dorsal fin has a narrow, yellow stripe at the base and the rest of the fin is black. The cartilaginous lower jaw is reddish-yellow to orange. The eye is slightly yellow. Small live fish are silvery overall with a white belly and olive back, the pectoral and pelvic fins slightly orange-yellow and other fins greyish although all fins may be hyaline. The peritoneum is black.

Small preserved fish have an indistinct blotch at the caudal fin base and a similar blotch on the back at the base of the spine in the dorsal fin. In very small fish these blotches are more distinct and there are 4-7 irregular blotches on the mid-flank above the lateral line and 3 blotches at the dorsal fin base. Development of blotches is individually variable, some fish being almost immaculate while in others the blotches extend vertically as bars as far as the back.

Size

Reaches 19.3 cm standard length (Krupp, 1985c).

Distribution

Found in the Orontes, (= Asi), Quwayq and Tigris-Euphrates basins. In Iran, it is found in the Tigris River basin including the Hawr Al Azim, Khersan, Jarrahi and Marun rivers (Berg, 1949; Abdoli, 2000) and the northern Gulf basin in the Shapur, Dalaki and Helleh rivers (Gh. Izadpanahi, pers. comm., 1995), the Zohreh River and possibly Lake Famur - some may be C. tenuiradius. Vossoughi (1998) reports this species from the western Hamun-e Jaz Murian basin based on a fishes with 13-15 branched dorsal fin rays, much higher than for C. watsoni, the taxon to be expected in this area.

Zoogeography

Zoogeographical comments are under the genus above.

Habitat

Known from a variety of habitats such as rivers, streams, reservoirs and ponds, as well as canals and gravel pits. Al-Habbib and Al-Habbib (1979) have demonstrated experimentally for a sample from "Nawaran Spring" north of Mosul, Iraq that this species can survive temperatures up to about 37°C. Akpinar and Aksoylar (1989) and Akpinar (1999) report this species from the Kangal Thermal Spring, Sivas, Turkey at a constant temperature of 35°C. This is the commonest species in catches in southwestern Iran, followed by Garra rufa. In areas under human influence in Lorestan, such as the lower reaches of rivers and near cities, it exceeds 80% in numbers in catches.

Age and growth

Maximum age reported for a population in the "Al-Nibaey" Lakes near Baghdad is 7+ years. Growth is slow and there is no difference in growth between males and females, although the habitat is not considered ideal for these fishes. Females tend to be slightly heavier than males of the same length especially in older fish. The length-weight relationship was W = 0.027 L2.67 (r = 0.78) for both sexes, W = 0.028 L2.65 (r = 0.90) for males and W = 0.020 L2.78 (r = 0.93) for females. Maturity is attained at 10.0-11.1 cm, corresponding to age group 2 (Allouse et al., 1989). The length-weight equation for commercially caught fish in the Tigris River was log W = 2.884 log L-4.623, condition factor was 1.15-1.47 (mean 1.28) and fish were immature up to age 2+ (Al-Nasiri, 1991). Haematology of this species from Sarao Subhana Agha near Sulaymaniyah was examined by Al-Mehdi and Khan (1984).

Food

Major food items in the Baghdad study are of plant origin with occasionally some chironomid larvae, copepods and cladocerans. Khan (1988) found for fish from near Sulaimaniyah, Iraq that diatoms and decayed organic matter are the main foods, with some green algae. Zooplankton are thought to be accidental food items. Guts contain mud and sand, evidence of a bottom feeding habit. Feeding increases at the start of the breeding season. The horny lower jaw covering is used to scrape algal food off hard bottom objects.

Reproduction

Near Baghdad, most fish are mature by April, the gonads occupying about one-third of the body cavity. Ovaries are orange to yellowish and testes milky white. Spawning occurs principally in May and June, with some in early July, but by July most fish are spent.

Iranian material shows minute but developing eggs in a 71.3 mm standard length fish caught on 31 January and specimens caught on 5 July have eggs 1.4 mm in diameter. The 31 January fish has tubercles on the snout and anal rays so tubercles develop quite early and in small fish. A fish caught on 20 September also shows tubercles around the snout. Small fish caught in January about 20 mm SL are presumably the young from the previous season and so show slow growth or are evidence of a prolonged or late spawning season.

Parasites and predators

Gussev et al. (1993a) describe a new species of monogenean from C. macrostomum in the Karun River, Dactylogyrus cyprinioni, and Jalali (1992) a new species of monogenean, Dogielius molnari, in the Dez River, both in Khuzestan. Jalali et al. (1995) describe a new species of monogenean, Dactylogyrus pallicirrus, from fish taken in the Dez River near Ahvaz.

Economic importance

Al-Mehdi and Khan (1984) report this species to be important in riverine and culture fisheries in northern Iraq. Ündar et al. (1990) identify this species and Garra rufa as the "doctor fish" of the Kangal hot spring in Turkey (Timur et al., 1983; Warwick and Warwick, 1989; Kürkçüoğlu and Öz, 1989; and various newspaper and television reports). High water temperatures reduce the amount of plankton available as fish food and the fish nibble away infected skin of humans who bathe in these waters. The fish is known as "striker" (and Garra rufa as "licker") from its behaviour in the spa pools. The healing properties are linked to the high level of selenium (1.3 p.p.m.) in the water, selenium being beneficial in some skin diseases, and possibly to UV light. The fish facilitate the action of the selenium and UV light by softening and clearing away psoriatic plaque and scale, exposing the lesions to the water and sunlight. However, some lesions are made worse and the fish can cause some new ones.

Conservation

This species is widely distributed in southern areas, particularly Khuzestan, and does not appear to be under threat other than that suffered by all species by pollution and water abstraction. Endangered in Turkey (Fricke et al., 2007).

Further work

See comments above on the need for further work to distinguish this species from C. kais, especially when young and below for distinction from C. tenuiradius.

Sources

Type material: See above, Cyprinion macrostomum (NMW 52805, 52806), C. neglectus (NMW 52807).

Iranian material: Tigris basin: and presumably macrostomum CMNFI 1979-0268, 13, 92.2-122.4 mm standard length, Lorestan, Dez or Karkheh drainage between Nowqan and Khorramabad (no other locality data); CMNFI 1979-0269, 4, 104.7-110.6 mm standard length, Lorestan, Dez or Karkheh drainage between Nowqan and Khorramabad (no other locality data); CMNFI 1979-0270, 10, 85.5-122.4 mm standard length, Lorestan, Kashkan River drainage (33º26'N, 48º19'E); CMNFI 1979-0271, 3, 100.7-144.8 mm standard length, Lorestan, Kashkan River drainage (33º39'N, 48º32'30"E); CMNFI 1979-0273, 9, ? mm standard length, Lorestan, Kashkan River drainage (33º26'N, 48º19'E); CMNFI 1979-0274, 14, ? mm standard length, Lorestan, Kashkan River drainage (33º27'N, 48º11'E); CMNFI 1979-0275, 2, 142.4-165.0 mm standard length, Lorestan, Kashkan River drainage (33º25'N, 47º58'E); CMNFI 1979-0278, 4, 93.5-114.1 mm standard length, Lorestan, Kashkan River drainage (33º34'N, 48º01'E); CMNFI 1979-0279, 9, 100.3-149.4 mm standard length, Lorestan, Khorramabad River (33º37'N, 48º18'E); CMNFI 1979-0283, 5, 93.0-144.0 mm standard length, Kermanshahan, Qareh Su drainage (34º21'N, 47º07'E); CMNFI 1979-0287, 1, 112.6 mm standard length, Kermanshahan, Chashmeh Javari 2 km from Ravansar (ca. 34º42'N, ca. 46º40'E); CMNFI 1979-0288, 1, 94.3 mm standard length, Ilam and Poshtkuh, Gangir River at Juy Zar (33º50'N, 46º18'E); CMNFI 1979-0289, 4, ? mm standard length, Kermanshahan, Diyala River drainage (34º28'N, 45º52'E); CMNFI 1979-0290, 11, 49.3-133.0 mm standard length, Kermanshahan, Diyala River drainage at Qasr-e Shirin (34º31'N, 45º35'E); CMNFI 1979-0291, 15, ? mm standard length, Kermanshahan, Diyala River drainage (34º24'N, 45º37'E); CMNFI 1979-0350, 18, ? mm standard length, Khuzestan, Marun River near Marun (30º39'30"N, 50º02'E); CMNFI 1979-0355, 1, ? mm standard length, Khuzestan, stream tributary to Karun River at Salmaneh (30º35'N, 48º22'E); CMNFI 1979-0356, 1, ? mm standard length, Khuzestan, stream at Hoveyzeh (31º27'N, 48º04'E); CMNFI 1979-0360, 2, ? mm standard length, Khuzestan, canal branch of Karkheh River (31º40'N, 48º35'E); CMNFI 1979-0361, 3, ? mm standard length, Khuzestan, jube in Karkheh River drainage (31º42'N, 48º33'E); CMNFI 1979-0363, 1, ? mm standard length, Khuzestan, Karkheh River (31º52'N, 48º20'E); CMNFI 1979-0364, 2, ? mm standard length, Khuzestan, river at Abdolkhan (31º52'30"N< 48º20'30"E); CMNFI 1979-0365, 24, ? mm standard length, Khuzestan, stream in Doveyrich River drainage (32º25'N, 47º36'30'E); CMNFI 1979-0366, 16, ? mm standard length, Khuzestan, stream west of Dehloran (32º45'30"N, 47º05'30"E); ID? CMNFI 1979-0367, 2, ? mm standard length, Khuzestan, Meymeh River 11 km north of Dehloran (32º44'30"N, 47º09'30"E) ID? CMNFI 1979-0368, 12, ? mm standard length, Khuzestan, Karkheh River (32º24'30"N, 48º09'E); CMNFI 1979-0371, 1, ? mm standard length, Khuzestan, stream in Karkheh River drainage (32º05'N, 48º19'E); CMNFI 1979-0373, 12, ? mm standard length, Khuzestan, Bala River north of Andimeshk (32º35'N, 48º17'E); CMNFI 1979-0374, 46, ? mm standard length, Khuzestan, stream tributary to Bala River (32º40'N, 48º15'E); CMNFI 1979-0376, 9, ? mm standard length, Khuzestan, river tributary to Karkheh River (32º48'30"N, 48º04'30"E); CMNFI 1979-0378, 10, ? mm standard length, Khuzestan, stream tributary to Karkheh River (ca. 32º48'N, ca. 48º04'E); CMNFI 1979-0379, 11, ? mm standard length, Khuzestan, Dez River (32º12'N, 48º27'E); CMNFI 1979-0380, 5, ? mm standard length, Khuzestan, stream tributary to Dez River (ca. 32º10'N, ca. 48º35'E); CMNFI 1979-0381, 28, ? mm standard length, Khuzestan, stream 40 km west of Shushtar (ca. 32º10'N, ca. 48º35'E); CMNFI 1979-0382, 67, ? mm standard length, Khuzestan, Karun River at Shushtar (32º03'N, 48º51'E); CMNFI 1979-0383, 1, ? mm standard length, Khuzestan, stream in Ab-e Shur drainage (31º59'30"N, 49º06'E); CMNFI 1979-0384, 7, 86.3-152.2 mm standard length, Khuzestan, Ab-e Shur drainage (32º00'N, 49º07'E); CMNFI 1979-0386, 4, ? mm standard length, Khuzestan, stream 21 km from Haft Gel (ca. 31º34'N, ca. 49º23'E); CMNFI 1979-0387, 6, ? mm standard length, Khuzestan, stream 12 km from Haft Gel, Jarrahi River drainage (31º25'N, 49º38'E); CMNFI 1979-0388, 2, ? mm standard length, Khuzestan, Zard River (31º19'N, 49º44'E); CMNFI 1979-0390B, 23, 36.2-156.2 mm standard length, Khuzestan, stream 3km south of Bagh-e Malek (31º29'N, 49º54'30"E); CMNFI 1979-0391, 1, 154.5 mm standard length, Khuzestan, stream in Marun River drainage (31º28'N, 49º51'E); CMNFI 1979-0392, 5, ? mm standard length, Khuzestan, Zard River (ca. 31º32'N, ca. 49º48'E); CMNFI 1979-0393, 2, 96.9-116.6 mm standard length, Khuzestan, Jarrahi River drainage (31º18'N, 49º37'E); CMNFI 1979-0394, 1, 130.2 mm standard length, Khuzestan, stream in Marun River drainage (31º01'N, 49º45'E); CMNFI 1979-0395, 4, ? mm standard length, Khuzestan, stream in Marun River drainage (ca. 30º57'N, ca. 49º51'E); CMNFI 1979-0396, 1, ? mm standard length, Khuzestan, Kheyrabad River (30º32'N, 50º23'30"E); ID? CMNFI 1979-0398, 23, ? mm standard length, Boyer Ahmadi-ye Sardsir va Kohkiluyeh, stream in Zohreh River drainage (30º24'30"N, 50º37'30"E); ID? CMNFI 1979-0399, 7, ? mm standard length, Fars, stream in Zohreh River drainage (30º19'30"N, 51º15'E); CMNFI 1991-0153, 1, 171.3 mm standard length, Khuzestan, Zohreh River (no other locality data); CMNFI 1991-0154, 1, 109.9 mm standard length, Khuzestan, Hawr al-Azim (ca. 31º45'N, ca. 47º55'E); CMNFI 1993-0128, 1, 110.7 mm standard length, Kermanshahan, Sarab-e Sabz 'Ali Khan (34º25'N, 46º32'E); CMNFI 1993-0149, 1, 121.7 mm standard length, Khuzestan, Karun River (no other locality data); CMNFI 2007-0111, 6, 24.7-173.8 mm standard length, Kermanshahan, Alvand River near Sar-e Pol-e Zahab (ca. 34º36'N, ca. 45º56'E); CMNFI 2007-0112, 6, 46.5-118.8 mm standard length, Kermanshahan, Kerend River basin near Shahabad-e Gharb (ca. 34º06'N, ca. 46º30'E; CMNFI 2007-0113, 1, 122.1 mm standard length, Kermanshahan, Razavar River, Qareh Su tributary (ca. 34º25'N, ca. 47º01'E); CMNFI 2007-0115, 6, 59.7-154.8 mm standard length, Kermanshahan, Qareh Su basin (ca. 34º34'N, ca. 46º47'E); CMNFI 2007-0116, 12, ?-93.0 mm standard length, Kermanshahan, Gav Masiab basin west of Sahneh (ca. 34º28'N, ca. 47º36'E); CMNFI 2007-0117, 1, ? mm standard length, Kermanshahan, Gav Masiab basin near Sahneh (ca. 34º24'N, ca. 47º40'E); BM(NH) 1980.8.28:1, 1, 90.3 mm standard length, Khuzestan, Dezful (32º23'N, 48º24'E); BWC95-20, 14, ? mm standard length, Khuzestan, Rud Zard at Rud Zard (31º22'N, 49º43'E); Gulf fish:- ? tenuiradius CMNFI 1979-0020, 56, ?, mm standard length, Fars, Mand River outside Kavar (29º11'N, 52º41'E); CMNFI 1979-0054, 14, 37.4-64.1 mm standard length, Fars, Shur River tributary (ca. 28º58-29º03'N, ca, 52º34-35'E); CMNFI 1979-0075, 123, 21.3-142.4 mm standard length, Fars, Mand River at Pol-e Kavar (29º11'N, 52º41'E); CMNFI 1979-0109, 5, 63.2-100.2 mm standard length, Fars, Mand River at Shahr-e Khafr (28º56'N, 53º14'E); CMNFI 1979-0128, 7, 19.2-103.8 mm standard length, Shur River (28º51'N, 52º31'E); CMNFI 1979-0131, 19, 16.4-41.7 mm standard length, Fars, Ab-Arak River (28º38'N, 52º49'E); CMNFI 1979-0132, 72?, 15.2-100.1 mm standard length, Fars, Ab-Arak River (28º35'N, 52º58'E); CMNFI 1979-0133, 50, 45.6-95.5 mm standard length, qanat stream near Qir (28º27'30"N, 53º03'E); CMNFI 1979-0135, 18, 21.8-49.2 mm standard length, Mand River tributary (28º08'N, 53º10'E); CMNFI 1979-0157, 4, 23.6-85.4 mm standard length, Fars, qanat stream at Hadiabad (28º52'N, 54º13'E); macrostomum? CMNFI 1979-0193, 1, 36.3 mm standard length, Fars, river 8 km from Darab (28º45'N, 54º27'30"E); macrostomum? CMNFI 1979-0195, 1, ? mm standard length, Fars, jube west of Darab (ca. 28º54'N, ca. 53º53'30"E); CMNFI 1979-0196, 1, 59.9 mm standard length, Fasrs, qanat and pool at Khanehnehrin (28º50'N, 53º31'30"E); not on data sheet check jar? CMNFI 1979-0197, 1, 51.3 mm standard length, Fars, spring nd stream 33 km from Fasa (28º45'N, 53º25'E); CMNFI 1979-0198, 23, 22.3-57.7 mm standard length, Fars, stream at Tadovan (28º47'N, 53º24'30"E); CMNFI 1979-0200, 8, 29.0-46.1 mm standard length, Fars, Mand River tributary (28º36'N, 53º36'30"E); CMNFI 1979-0202, 12, ? mm standard length, Fars, Mand River (29º01'N, 53º00'E); CMNFI 1979-0241, 18, 43.8-72.6 mm standard length, Fars, Shapur River at Shapur (29º47'N, 51º35'E); CMNFI 1979-0347, 2, 105.2-106.7 mm standard length, Fasr, Pol-e Berengie (29º27'30"N, 52º32'E); CMNFI 1979-0348, 4, 52.9-79.1 mm standard length, Fars, stream at Somduldul (ca. 29º28'N, ca. 52º32'E); CMNFI 1979-0404, 25, 20.2-127.9 mm standard length, Bushehr, stream 33 km south of Kaki (28º08'N, 51º47'E); CMNFI 1979-0405, 4, 33.5-36.7 mm standard length, Hormozgan, stream about 13 km north of Rostaq (28º29"N, 54º59'E); ID? CMNFI 1979-0497, 1, 85.6 mm standard length, Fars, Mand River at Band-e Bahman (29º11'N, 52º40'E); CMNFI 1979-0501, 17, 18.7-91.0 mm standard length, Fars, Mand River at Kavar (29º11'N, 52º41'E); CMNFI 1979-0504, 6, ?-93.0 mm standard length, Fars, stream at Pol-e Gaz in Lake Maharlu basin (no other locality data); CMNFI 1979-0789, 1, 164.6 mm standard length, Fars, Lake Parishan (29º45'N, 53º40'E); CMNFI 1993-0141, 1, 64.4 mm standard length, Bushehr, Dalaki River (29º28'N, 51º15'E); ID? CMNFI 2007-0061, 2, ? mm standard length, Fars, qanat pool at Ab-e Barik (ca. 27º52'N, ca. 54º09'E); CMNFI 2007-0063, 6, ? mm standard length, Fars, Mand River outside Jahrom (28º36'N, 53º37'E); USNM 205890, 2, 46.0-48.7 mm standard length, Fars, Lake Parishan (29º45'N, 53º40'E); ZSM 25705, 1, 107.0 mm standard length, Fars, Lake Parishan (29º45'N, 53º40'E).

Comparative material:- CMNFI 1980-0811, 2, 82.6-112.4 mm, Turkey, Akziyaret Deresi, Tigris River system (no other locality data);

BM(NH) 1931.12.21:1-2, 2, 69.5-78.5 mm standard length, Iraq, Mosul (36º20'N, 43º08'E); BM(NH) 1974.2.22:1184, 1, 130.2 mm standard length, Iraq, Sulaimaniyah (); BM(NH) 1974.2.22:1196, 1, 53.0 mm standard length, Hawiya Canal, Lesser Zab (); BM(NH) 1974.2.22:1214-1255 (in part), Khalis (33º49'N, 44º32'E).

Cyprinion milesi
(Day, 1880)

Common names

None.

[sabzug in Pakistan].

Systematics

Barbus milesi was described from "a spring at Tràl", Pakistan.

Berg (1949), Mirza (1969), Mirza et al. (1991) and Howes (1982) recognise this species as valid. If so, synonyms according to Berg (1949), would be Barbus bampurensis Nikol'skii, 1899 described from "Flum. Bampur", Scaphiodon daukesi Zugmayer, 1912 from "Irrigation channels and pools near Panjgur, Baluchistan, Pakistan", and Barbus baschakirdi Holly, 1929 from "Ein Bach bei Guadjik am Wege von Sarzeh in Biabun nach Darpahan in den Bergen von Baschakird, Südostpersien" (= a brook at Guadjik on the way from Sarzeh in Biabun to Darpahan in the Baschakird Mountains, southeast Persia).

Much of my material from southeastern Iran was assigned by me to C. watsoni. Specimens that resemble C. milesi (lacking a shallowly arched or sector mouth with a horny edge but having an oblique u-shaped mouth) are found at the same sample localities as typical C. watsoni. The mouth structure of the putative C. milesi resembles that of juvenile C. watsoni, possibly retained in the adult (paedomorphosis). A Principal Components Analysis does not separate these two forms when the mouth characters are not included in the analysis.

A specimen in the Naturhistorisches Museum Wien under NMW 52736, 34.4 mm standard length, is listed as a syntype under the name Cirrhina milesi but its locality is Gwadur, Hubb River and the type status may be an error.

Five syntypes of Barbus bampurensis, 32.0-64.8 mm standard length, are in the Zoological Institute, St. Petersburg (ZISP 11715) from "Flum. Bampur, 15-23.VII.1898, Zarudnyi". The jar label gives a date of 15-19.VII.1898.

The holotype of Barbus baschakirdi, 52.2 mm standard length, is in the Naturhistorisches Museum Wien under NMW 13798 and a cotype (syntype) of Scaphiodon daukesi, 102.8 mm standard length, is under NMW 19784.

Scaphiodon daukesi types in Munich were destroyed in World War II but one syntype is in the Naturhistorisches Museum Wien under NMW 19784, and two syntypes are in the Zoological Survey of India, Calcutta under ZSI F8028/1 and 8032/1 ((Menon and Yazdani, 1968; Eschmeyer et al., 1996; Neumann, 2006).

Key characters

The mouth is characteristically oblique, longer in lateral view than C. watsoni.

Morphology

The oblique mouth reaches back to the anterior eye margin in small fish and to the rear of the nostril in larger fish. Dorsal fin with 3 unbranched and 10-13 branched rays, anal fin with 2 unbranched and 7 branched rays, pectoral fin with 14-16 branched rays and pelvic fin with 7-8 branched rays. Total gill rakers 11-12. The following description is based mostly on Barbus bampurensis types. Dorsal fin spine strong and serrated, with large teeth in small fish. Lateral line scales 34-39. The scaleless groove before the dorsal fin is weakly expressed. Scales are present on the belly of large fish, almost absent on small fish. Upper flank scales may be regularly or irregularly arranged. Scales have few to no anterior radii, numerous posterior radii, numerous fine circuli, a subcentral anterior focus, and an anterior scale margin indented above and below the mid-line. A pelvic axillary scale is present. The head is more massive in relation to the body than for similar size C. watsoni/kirmanense specimens. The barbel is quite stubby at the base but tapers rapidly to the tip in larger fish. The type series of Barbus bampurensis (= C. milesi) has dorsal fin branched rays 10(4) or 11 (1), anal fin branched rays 7(5), pectoral fin branched rays 14(1) or 15(3) (one unclear), pelvic fin branched rays 7(1) or 8(4), lateral line scales 34(1), 36(1) and 37(3), and total gill rakers 11(3) or 12 (2). Two fish from Sib (see below) had dorsal fin branched rays 9(1) or 10(1), anal fin branched rays 7(2), pectoral fin branched rays 15(2), pelvic fin branched rays 6(1) or 7(1), lateral line scales 35(1) or 37(1), pharyngeal teeth 4,3,2 on the left side, total gill rakers 13(1) or 14 (1), and total vertebrae 38(1) or 39(1). Pharyngeal teeth have a slight hook on the anteriormost tooth with the rest in the main row with scooped-out crowns.

Sexual dimorphism

Tubercles line the anal fin rays and are apparent on the snout in males.

Colour

Copper-brown on the back and upper flank fading to a pinkish belly. Fins are pink and the lateral line has a bright orange streak along it. The preopercle also has orange-golden spots as does the base of the pectoral fins. There is a dark blotch at the base of the caudal fin. The caudal fin base bears a spot in small specimens and there are some much smaller, irregular spots on the caudal peduncle. Peritoneum brown to black.

Size

Attains about 19.0 cm.

Distribution

In Iran, it is recorded from the Sarbaz River of the Makran according to Saadati (1977), the Bampur River of the Hamun-e Jaz Murian basin according to Berg (1949) and the Dozdan River of the Hormuz basin (H. R. Esmaeili). Also in the Mashkid River basin in Pakistan and in rivers draining to the Indian Ocean.

Zoogeography

See under the genus.

Habitat

Unknown.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

None reported from Iran.

Economic importance

None.

Conservation

The distribution, abundance and biology of this species in Iran is poorly known and an assessment for conservation status cannot be given.

Further work

See above.

Sources

Type material: See above, Barbus bampurensis (ZISP 11715),Barbus baschakirdi (NMW 13798) and Scaphiodon daukesi (NMW 19784).

Iranian material: BM(NH)1883.8.2:2-3, 2, 72.2-130.9 mm standard length, Baluchestan, Sib near Dizak (27º15'N, 62º05'E). BWC97-4 no fish on cat sheet?

Comparative material: BM(NH) 1889.2.1:263-264, 2, 89.3-108.7 mm standard length, Afghanistan (no other locality data).

Cyprinion tenuiradius
Heckel, 1849

Common names

[Araxes kingfish (Fricke et al., 2007)]

Systematics

The type locality is the "Kara-Agatsch als aus dem Araxes" (= Qarah Aqaj River and the Kor River, Fars). Sometimes spelt tenuiradiatus (e.g. in Rainboth (1981) but this is incorrect). Syntypes of Cyprinion tenuiradius are in the Naturhistorisches Museum Wien according to Kähsbauer (1964) under NMW 52808 (1 specimen, 116.7 mm standard length), 52809 (2, 52.3-58.0 mm standard length), 52811 (4, 42.7-47.4 mm standard length), 52815 (1, 77.0 mm standard length) and 52816 (2, 75.5-80.8, although Kähsbauer lists only 1 while Banarescu and Herzig-Straschil (1995) list 2 as also found by me). Other material marked as syntypes from the "Kara-Agatsch. Th. Kotschy" includes NMW 52810 (2 , 103.7-110.0 mm standard length), NMW 52812 (2, 103.5-104.8 mm standard length), NMW 52813 (2, 97.7-103.1 mm standard length), NMW 52814 (1, 114.9 mm standard length), and 52817 (1, not examined). The catalogue in Vienna lists 8 specimens in one column and 26 in the adjacent column. Eschmeyer et al. (1996) add 2 fish from the Araxes River, formerly in NMW, now at the Rijksmuseum van Natuurlijke Historie, Leiden under RMNH 2486. The lectotype as selected by F. Krupp in 1984 is NMW 52814 and is published by Banarescu and Herzig-Straschil (1995) with NMW 52808, 52809, 52810, 52811, 52812, 52813, 52815 and 52816 as paralectotypes.

Karaman (1971) assigns this taxon as a subspecies of Cyprinion macrostomum and Bianco and Banarescu (1982) suggest it may be a subspecies in a polytypic species. Berg (1949) records it from the Tigris River where it may be sympatric with C. macrostomum. He considers it to be close to that species, perhaps its southeastern subspecies. Howes (1982) considers tenuiradius to be a variant of C. macrostomum.

Heckel (1846-1849b) distinguishes this species from C. macrostomum by a lower scale count (35-36 as opposed to 42; Berg (1949) gives 35-38 as opposed to 37-43); Krupp (1985c) gives 34-38 compared to 39-43 in macrostomum; Banarescu and Herzig-Straschil (1995) give 36-38, rarely 35 or 39 in C. tenuiradius compared to 41-44, rarely 40 or 45 in C. macrostomum), slenderer body, and a much thinner dorsal spine which is soft in its distal third. The mouth is arched and there is some lower lip development at the mouth corner as in C. kais (see illustrations in Krupp (1985c)). In addition, Berg (1949) gives a branched dorsal fin ray count of 12-13 in C. tenuiradius, 13-15 in C. macrostomum, although Banarescu and Herzig-Straschil (1995) give (12)13-15 for C. tenuiradius from the type locality of Kara-Agasch (sic). Krupp (1985c) states that tenuiradius has a smaller number of scale radii than macrostomum, radii are divergent and the posterior scale margin is curved. However, data for specimens examined by me show overlaps in meristic characters; although means differ, individual fish would be difficult to distinguish on counts alone.

The question then arises as to whether tenuiradius is distinct from macrostomum or merely a variant of a wide-ranging, variable species. The only absolute character is a weaker dorsal fin spine based on examination of type material; other, meristic characters overlap and minor variations in body form are difficult to quantify given a wide range of habitats (lowland rivers and marshes versus highland streams) which may affect shape. The species tenuiradius is retained here as distinct but would benefit from further analyses using new characters, if available, from molecular data.

Key characters

Distribution and a weak spine distinguish this taxon. The dorsal fin spine in macrostomum has teeth extending further along the spine, teeth are more well-developed even near the tip. Spine teeth in tenuiradius are more graded in size as they near the tip and are finer than in macrostomum.

Morphology

Dorsal fin with 4 unbranched and 11-15 branched rays (Berg (1949) has 12-13). The anal fin has 3 unbranched and 6-8 branched rays, usually 7. In 199 Iranian fish, 96.5% have 7 anal fin rays with the rest having 6 rays and 1, presumably anomalous fish, with 9 rays. Pectoral fin branched rays 13-18, pelvic fin branched rays 7-9. Lateral line scales 32-39. Gill rakers 10-21. Scales on the belly may be small and skin covered. There is a naked dorsal keel in front of the dorsal fin, although the area behind the occiput may be scaled and the groove begins nearer the dorsal fin. The mouth is transverse to more or less curved. The dorsal fin spine is weak and serrated only half way or two-thirds of its length. The chromosome number is 2n=50, comprising 13 metacentric, 5 submetacentric and 7 subtelocentric chromosomes pairs. Arm number is NF=86 (Esmaeili and Piravar, 2006).

Meristics for fish from Persian Gulf drainages of Fars, Bushehr and Hormozgan provinces including the Lake Maharlu endorheic basin:- dorsal fin branched rays 11(4), 12(51), 13(175), 14(74) or 15(9) (mean = 13.1, S.D. = 0.746); pectoral fin branched rays 13(3), 14(38), 15(117), 16(41), 17(2) or 18(1)(mean = 15.0, S.D. = 0.733); pelvic fin branched rays 7(23), 8(177) or 9(3)(mean = 7.9, S.D. = 0.345); total gill rakers 10(2), 11(16), 12(27), 13(24), 14(49), 15(35), 16(20), 17(14), 18(8), 19(3) or 21(1)(many counts are based on small specimens and may be low accordingly in comparison with Tigris River basin fishes; mean = 14.2, S.D. = 2.003); and lateral line scales 32(1), 33(15), 34(28), 35(41), 36(47), 37(56), 38(13) or 39(2)(mean = 35.7, S.D. = 1.431).

Sexual dimorphism

Unknown.

Colour

Overall colour is yellowish-white with a light grey back. Scale bases on the flank above the lateral line are brown. The pectoral and pelvic fins have an orange-yellow spot at their base.

Size

Reaches 16.3 cm (Berg, 1949).

Distribution

This species is found in the Gulf and Lake Maharlu basins in Iran (Bianco and Banarescu, 1982; M. Rabbaniha, pers. comm., 1995; Abdoli, 2000).

Heckel (1849) records this species as from the "Araxes", the modern Kor River in Fars. However, the catalogue sheets in Vienna for the types only list the "Kara Agatsch" (= Mand River) and no subsequent collections have been made of this species in the internal Kor River basin although Abdoli (2000) also maps it from the middle to lower Kor River, possibly based on Heckel's report. Berg (1949) records it from the Tigris River basin, perhaps in error, and Fricke et al. (2007) have it in Turkey from the Aras River system of eastern Turkey (presumably a confusion of the modern Aras or Araxes River with the classical Araxes or Kor River of Fars).

Zoogeography

See under the genus.

Habitat

Unknown in detail but found in springs, streams and rivers of varying descriptions.

Age and growth

Unknown. Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 40 fish measuring 5.04-13.49 cm fork length. The a-value was 0.0139 and the b-value 3.063 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases).

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

None reported from Iran.

Economic importance

None.

Conservation

The distribution, abundance and biology of this species in Iran is poorly known and an assessment for conservation status cannot be given. Endangered in Turkey (Fricke et al., 2007) but probably does not occur there.

Further work

See above.

Sources

Type material: See above, Cyprinion tenuiradius (NMW 52808, 52809, 52810, 52811, 52812, 52813, 52814, 52815, 52816).

Iranian material: ? see above and ID

Cyprinion watsoni
(Day, 1872)

Common names

None.

[sehrgoar; sabzug = watsoni and microphthalmum - all in Pakistan].

Systematics

Scaphiodon irregularis Day, 1872 described from "rivers in the Sind hills", India, probably Scaphiodon microphthalmus Day, 1880 from "Quetta", Scaphiodon muscatensis Boulenger, 1887 from Muscat, Oman, Cirrhina afghana Günther, 1889 from "Nushki (N. Baluchistan)" and "small river at Kushk (N.W. Afghanistan), Badghis", Cyprinion kirmanense Nikol'skii, 1899 from "Schur-Ab in Kirmano orient.", Cirrhina afghana var. nikolskii Berg, 1905, Scaphiodon macmahoni Regan, 1906, Scaphiodon baluchiorum Jenkins, 1910 (see below for type locality), Scaphiodon watsoni var. belense Zugmayer, 1912 from the "Purali River, near Las Bela" (in Pakistani Baluchistan), Scaphiodon readingi Hora, 1923 from the "Salt Range, Punjab", India, and Cyprinion microphthalmum infraspecies nikolskii Berg, 1949 described originally in part as Cirrhina afghana var. nikolskii Berg, 1905, and Semiplotus dayi Fowler, 1958 are synonyms.

Semiplotus dayi was coined by Fowler to replace Scaphiodon aculeatus, a misidentification by Day (1880) for Chondrostoma aculeatum (= Capoeta aculeata). Fowler thought that Day's fish represented a new species which he named Semiplotus dayi. Howes (1982) considers Semiplotus dayi to be a synonym of Capoeta capoeta (since Karaman (1969a) synonymises Scaphiodon aculeata with C. capoeta. Day's Scaphiodon aculeatus is placed in the synonymy of Cyprinion microphthalmum infraspecies nikolskii by Berg (1949).

Syntypes (or at least specimens examined by Day) of Scaphiodon watsoni described from rivers on the Sind Hills and the Salt Range of the Punjab, India are in the Zoological Survey of India, Calcutta under ZSI 2596 (1), the Natural History Museum, London under BM(NH) 1889.2.1.370-9 (10, but 14 in jar September 2007, 35.6-93.4 mm standard length), the Australian Museum, Sydney under B.7751 (1), the Zoölogisch Museum, Universiteit van Amsterdam under ZMA 115.924 (2) and ZMA 119.255 (1), the Naturhistorisches Museum Wien under NMW 51671 (1), NMW 51672 (1) and NMW 51673 (1), the Museum für Naturkunde, Universität Humboldt, Berlin under ZMB 11042 (1)(132.6 mm standard length), the Rijksmuseum van Natuurlijke Historie, Leiden under RMNH 8704 (1) (or possibly 2552), the Zoological Institute, St. Petersburg under ZISP 8278 (4 but only 2 fish found by me, 63.6-79.6 mm standard length), and the Field Museum of Natural History, Chicago under FMNH 2303 ? 2302(4, 34.0-72.5 mm standard length as examined by me) (Whitehead and Talwar, 1976; Nijssen et al., 1993; Eschmeyer et al., 1996; Ferraris et al., 2000). The 3 fish in the Naturhistorisches Museum Wien measure 86.6, 80.8 and 93.3 mm standard length respectively and are listed there as syntypes.

ZISP 8279 comprising 3 fish, 51.5-52.1 mm standard length, has the same data as ZISP 8278 and may also be types. It is not clear if these are all types, those in ZISP not being marked as types and those in BM(NH) being marked as "possible types"; they may include material simply collected by Francis Day.

A cotype of Scaphiodon watsoni var. belense (NMW 19833) measures 136.9 mm standard length. Eschmeyer et al. (1996) report 2 fish under NMW 19833 although the Vienna card index in 1997 lists only one syntype under this number. In the Zoological Survey of India, Calcutta there are single syntypes under ZSI F827/1 (a misprint for 8027), ZSI F8029/1, ZSI F8030/1 and ZSI F8031/1 (see also Menon and Yazdani (1968)). The remainder of 42 syntypes were in the Munich Museum but were destroyed in World War II (Neumann, 2006).

Types of Scaphiodon microphthalmus are probably lost. The species was described from 2 specimens taken at Quetta in Pakistan. One specimen was sent to the Florence Museum but a recent search failed to locate it and the other specimen has not been located (Whitehead and Talwar, 1976; Banister and Clarke, 1977). A fish measuring 130.1 mm standard length in the Naturhistorisches Museum Wien is listed as a possible syntype (NMW 55897) and in the 1997 card index as "? holotype" (sic).

Note Howes (1982) and Mirza et al. (1991) consider Cyprinion microphthalmum to be a valid species with muscatensis, afghana, afghana var. nikolskii and baluchiorum as synonyms. Howes places macmahoni in watsoni rather than microphthalmum as Berg (1949) and Mirza (1969) do. Howes (1982) also includes irregularis, kirmanense, and readingi in watsoni.

A syntype of Scaphiodon irregularis is in the Australian Museum, Sydney under AMS B.7883 (Ferraris et al., 2000). Syntypes of Scaphiodon muscatensis are in the Natural History Museum, London under BM(NH) 1885.11.7:35-40 (6, 66.4-89.3 mm standard length) and BM(NH) 1887.11.11:289-291 (3, 72.1-79.3 mm standard length) (Eschmeyer et al., 1996; personal observations). Syntypes of Scaphiodon readingi are in the Zoological Survey of India, Calcutta under ZSI F10353/1 and ZSI 10354/1 (27) (sic, although the catalogue numbers seem to indicate only 2 fish) (Menon and Yazdani, 1968) and in the Zoological Museum of Moscow University (ZMMU) (P-1588) (Pavlinov and Borissenko, 2001).

Three syntypes of Scaphiodon baluchiorum (ZSI F9398 to F9400) and one syntype of Scaphiodon macmahoni (ZSI F1239/1) are in the Zoological Survey of India, Calcutta (Menon and Yazdani, 1968). A syntype of Scaphiodon macmahoni measuring 58.6 mm standard length from "Seistan" is in the Natural History Museum, London and was labelled as Cyprinion watsoni (BM(NH) 1905.11.29:27, 58.6 mm standard length). The type locality of Scaphiodon baluchiorum is "Gishtigan (Bampusht); Kalagan, 3,500 feet; Baluchistan". These localities are in Pakistani Baluchistan; Gishtigan being on the Kulushta River which drains into the Nihing River and then the Dashti River (Jenkins, 1910) (these are near the border of Iranian Baluchestan with the upper reaches of the Nihing being in Iran) and Kalagan possibly being the Kalugar River with headwaters in Iran and draining to the Hamun-i Mashkel in Pakistan. The type locality of Scaphiodon macmahoni is "affluents of the Helmand" (Regan, 1906), presumably an error for "effluents" or the delta of the Helmand.

The holotype of Cyprinion kirmanense, 61.6 mm standard length, is in the Zoological Institute, St. Petersburg under ZISP 11712 from "Schur-Ab in Kirmano orient. 27.VI." The 5 syntypes of Cirrhina afghana var. nikolskii are in the Zoological Institute, St. Petersburg (ZISP 11709) and are from the "Bampur River, 27 VII 1898, N. Zarudnyi" according to Berg (1949) but he mentions 2 additional fish with a somewhat deeper body, presumably also part of the type series. ZISP 11709 does have 7 specimens, 43.0-79.1 mm standard length, with a date 15-27.VII.1898. Four syntypes of Cirrhina afghana measuring 74.6-83.0 mm standard length from "Kushk" annotated Afghan. Boundary Comm. are in the Natural History Museum, London (BM(NH) 1886.9.21:150-154 - note that 150-154 indicates there should be 5 fish) with a further 6 syntypes measuring 44.9-99.5 mm standard length labelled "Nushki" and also annotated Afghan. Boundary Comm. (BM(NH) 1886.9.21:155-159 - note this indicates there should be 5 fish in this jar and probably one fish has been mixed up). Additional syntypes are in the Zoological Survey of India, Calcutta under ZSI 11474-11476 (3) and ZSI 11479-11485 (7) (Eschmeyer et al., 1996).

Berg (1949) places Cirrhina afghana var. nikolskii in his Cyprinion microphthalmum infraspecies nikolskii (see also Berg (1933a)). This infraspecies occurs together with Cyprinion microphthalmum but differs by a stronger osseous ray in the dorsal fin which is serrated almost to the summit (Berg (1949) states that transitions exist). The anterior belly region is scaleless also. ZISP 11709 fish mostly have their dorsal spines snapped off but one fish has osseous ray teeth between three-quarters and four-fifths along the spine and a second about three-quarters. ZISP 25406 from a qanat between Kerman and Bandar-e `Abbas comprises 12 fish, 31.0-53.6 mm standard length, belonging to infraspecies nikolskii according to Berg (1949). These fish, of all sizes, have the last quarter to a third of the osseous spine in the dorsal fin unserrated. The mouth form varies. One large fish has a terminal mouth, moderately oblique in lateral view, and no strong horny layer on the lower jaw. Others have a u-shaped or horny jaw positioned on the lower head surface so there is no real gape in lateral view. Some small fish are transitional between the two types. Fin serration, mouth form and development of scales on the anterior belly seem to be widely variable within samples of Cyprinion from a single locality and presumably a single species.

Berg (1949) recognises Cyprinion watsoni belense as a subspecies, rather than a variety as originally described, from Indian Ocean drainages of southeastern Iran and southwestern Pakistan (Baluchistan). It is distinguished by smaller scales (33-36) from the type form (31-34), hardly a sufficient criterion given the wide distribution range and individual variation shown by these fishes.

This species has not been adequately examined in southeastern Iran and most nominal species are referred to Cyprinion watsoni, the earliest available name for the taxon. C. watsoni is distinguished from other Iranian Cyprinion by having 9-11 dorsal fin branched rays (macrostomum and tenuiradius have 12-15; C. milesi also has a low dorsal ray count but has an oblique mouth, not transverse or arched (Berg, 1949)). Bianco and Banarescu (1982) consider that several subspecies may eventually be defined and that some of the names in synonymy here would then be used.

Berg (1949) also recognises C. irregulare as a distinct species with a low dorsal fin branched ray count as in C. watsoni but usually 37 or more scales in the lateral line, a scaleless groove on the back before the dorsal fin, and upper scale rows anteriorly arranged irregularly and not imbricate and C. microphthalmum with a low dorsal fin branched ray count as in C. watsoni but usually 37 or more scales in the lateral line, a scaleless groove on the back before the dorsal fin barely outlined, and upper scale rows anteriorly arranged regularly and imbricate. C. microphthalmum infraspecies nikolskii is described as having a strong dorsal fin spine with obvious teeth extending to the tip while typical C. microphthalmum has a weak ray with weak teeth only visible when the skin covering the fin is peeled away.

Berg (1949) later states that no great importance should be attached to the upper row scale arrangement and the groove development - if the groove is well-developed then the upper row scales are irregular and this phenomenon can be seen in some C. watsoni and C. microphthalmum specimens. Berg then suggests that irregulare could be regarded as an infraspecies of C. microphthalmum as this type of condition occurs in Capoeta fusca and in Garra rossica. Under the heading C. watsoni Berg also gives mouth shape, scale arrangement, dorsal fin spine serrations, and body form as characters which can vary greatly. These observations serve to confirm the great variability in characters for these fishes. Large series of adults and young would be needed to adequately define species and subspecies.

Mirza (1969) reports C. watsoni, C. microphthalmum and C. milesi from western Pakistan and Iran, the former in Makran drainages and the latter two in the Mashkel (= Mashkid) River basin. The characters used to separate these taxa are an oblique mouth and head length contained less than 4.5 times in total length (= C. milesi), an arched mouth, head length more than 4.5 times in total length, scaleless strip on back conspicuous, and 33-36 lateral line scales (= C. watsoni), and a transverse mouth, head length more than 4.5 times in total length, scaleless strip on back hardly visible, and 37-40 lateral line scales (= C. microphthalmum). Sample sizes in this study were small (22 fish) and these characters show considerable variation in larger samples and between fish of different sizes.

Key characters

The arched mouth and and 9-12, usually 10-11, branched dorsal fin rays serve to identify this species.

Morphology

Young fish have a more horseshoe-shaped mouth than larger and older fish where the mouth is a shallow arch, almost straight. The dorsal fin has 3-4 unbranched and 9-12 branched rays, the last unbranched ray of the dorsal fin being variably serrated and thickened. The extent of serrations appears to vary independently of size, from half to three-quarters or more of the spine length. The anal fin has 1-3, usually 3, unbranched and 6-8, usually 7 branched rays. In Iranian specimens, 89.7% of 419 fish had 7 branched anal fin rays, the remainder having 6 branched rays. Pectoral fin branched rays 11-18, usually 15-16, and pelvic fin branched rays 6-9, usually 8. Lateral line scales 31-43. Scales have well-developed anterior radii as well as posterior and some lateral radii. The scale focus is almost central on mid-flank scales. There is a naked median strip on the back in front of the dorsal fin, about one scale wide, in some fish. Some fish may show poor imbrication of scales on the belly and upper anterior flank. Gill rakers 8-18, reaching to or past the adjacent raker when appressed. Total vertebrae 38. Rarely with a tripartite swimbladder, usually bipartite (Mirza, 1971 - for his C. microphthalmum). Pharyngeal teeth 2,3,4-4,3,2 or 2,3,5-5,3,2, with spoon-shaped crowns. The back in front of the dorsal fin is naked in the mid-line. Mouth shape variable, from and arch to a transverse cutting edge. Scales on the belly can be embedded in the skin, obvious or even absent.

Meristics for Iranian specimens:-
 

Locality/Dorsal Fin Rays

9

10

11

12

x

S.D.

Hamun-e Mashkid

16

3

10.2

0.375

Hamun-e Jaz Murian

3

15

5

10.1

0.596

Dasht-e Lut

2

50

4

10.0

0.328

Makran

1

29

3

10.1

0.348

Hormuz

2

144

124

7

10.5

0.562

Sirjan

1

3

7

10.5

0.688

 

Locality/Pelvic Fin Rays

6

7

8

9

x

S.D.

Hamun-e Mashkid

6

13

7.7

0.478

Hamun-e Jaz Murian

1

21

1

8.0

0.302

Dasht-e Lut

1

54

8.0

0.135

Makran

33

8.0

0.000

Hormuz

2

31

237

8

7.9

0.400

Sirjan

2

9

7.8

0.405

 

Locality/Pectoral Fin Rays

11

13

14

15

16

17

18

x

S.D.

Hamun-e Mashkid

9

10

15.5

0.513

Hamun-e Jaz Murian

1

8

11

2

1

15.7

0.864

Dasht-e Lut

1

2

14

31

8

14.8

0.879

Makran

1

14

15

3

15.6

0.704

Hormuz

1

46

111

99

20

1

15.3

0.863

Sirjan

4

4

3

14.9

0.831

 

Locality/Total Gill Rakers

8

9

10

11

12

13

14

15

16

17

18

x

S.D.

Hamun-e Mashkid

1

4

3

4

3

4

13.8

1.718

Hamun-e Jaz Murian

3

4

10

4

1

1

12.0

1.492

Dasht-e Lut

1

7

8

16

12

10

2

12.2

1.427

Makran

1

6

9

4

8

3

1

1

13.0

1.794

Hormuz

3

7

33

45

64

60

41

10

2

1

12.2

1.569

Sirjan

2

2

1

5

1

12.1

1.375

 

 
Locality/Lateral Line Scales 33 34 35 36 37 38 39 40 41 42 43 x S.D.
Hamun-e Mashkid 1 3 9 1 1 1 2 1
Hamun-e Jaz Murian 1 9 11 1 1
Dasht-e Lut 5 12 26 10 2 1
Makran 2 1 17 8 4 1
Hormuz 1 2 17 57 99 72 20 7 3
Sirjan 3 4 3 1

? vertebrae

The holotype of Cyprinion kirmanense has 10 dorsal fin branched rays, 7 anal fin branched rays, 15 pectoral fin branched rays, 8 pelvic fin branched rays, 37 lateral line scales and 13 total gill rakers (not in above tables).

Sexual dimorphism

Males have snout tubercles and tubercles on the anal fin rays (Regan, 1906; Jenkins, 1910; Berg, 1949). Large tubercles are found on the snout in front of the nostrils, the top of the head, and in rows on the rays of the caudal and anal fin, following the ray branching, in a fish not yet fully mature (40.1 mm standard length). ZMB 11042 (132.6 mm SL) has tubercles thickly present on the snout extending back to the nostrils and then to the eyes, scattered all over the sides of the head, absent on top of the head (may be lost in this old specimen) and large tubercles on anal fin rays near the tip. There is a depression in front of the nostrils in adult males.

Colour

The back and upper flank are dark or copper brown, golden or dark olive, light green-brown or brown-grey, sometimes with bluish or orange tinges, fading to a light or yellowish pink on the lower flanks and belly. The flank scales are silvery and may be outlined in black. Black spots may be present along the flank as may be an orange stripe or a series of 7-9 orange spots above the lateral line anteriorly. The orange colour may be deep, almost red. Occasionally, there may be an orange spot below the lateral line. There may be a vertical orange line over the cleithrum or a spot at its postero-ventral corner. In some fish the whole cleithrum area is red-orange. The operculum, preoperculum and cheek can be iridescent blue. The bases of the pectoral and pelvic fins and part of the operculum may also be pink or orange-coloured. Young have a fine black streak above the lateral line. The dorsal and caudal fins are lead-coloured to black and other fins are pink to yellowish. All the fins except the paired fins may be hyaline. There can be a black caudal base spot, quite marked in some fish, particularly small ones. The peritoneum is black or dark brown.

Size

Attains 23 cm (Zugmayer, 1912).

Distribution

From southeastern Iran east to India. In Iran, it is recorded from the Dasht-e Lut, Hormuz, Hamun-e Jaz Murian, Sistan, Hamun-e Mashkid basins and in the Makran from the Jagin to the Bahu Kalat rivers (Berg, 1949; Spillman, 1972; Bianco and Banarescu, 1982; Abdoli, 2000; Esmaeili et al., 2011?).

Zoogeography

The occurrence of this species (as C. microphthalmum in Banister and Clarke (1977)) in Oman across the Straits of Hormuz is a result of the 120 m lowered sea level from 100,000 B.P. to 10,000 B.P. The Tigris-Euphrates then ran down the Gulf and presumably provided ready access across it. Banister and Clarke (1977) comment that it is surprising that only one species made the crossing but nothing is known of the climate during this 90,000 year period nor of the composition of the Iranian fish fauna. It may well have been quite impoverished. Hora (1956) describes fish paintings on pots from Nal in Pakistani Baluchistan dating from the third millenium B.C. One of the species not clearly represented is a Cyprinion, now found in that area (not extinct as Banister (1980) would have it). This potentially shows how the ichthyofauna in Southwest Asia can change over relatively short periods of time; the changes over 100,000 years must have been considerable and not readily traceable.

Dorsal fin branched rays have moderately strong modes of 10 for the Hamun-e Mashkid, Hamun-e Jaz Murian, Dasht-e Lut and Makran basins but modes of 10 and/or 11 for Hormuz and Sirjan, the westernmost basins. C. watsoni may be showing some introgression with Cyprinion species to the west which have higher counts. However, subsamples within the Hormuz basin do not show a clear pattern of higher modes or means towards the west.

Habitat

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

Age and growth

Unknown. Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 23 Iranian fish measuring 8.34-13.38 cm total length. The a-value was 0.0101 and the b-value 2.952 (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 are primarily herbivorous items including filamentous algae such as Cladophora and Spirogyra, and a wide range of diatoms but some insect material is also found (Mirza, 1969; Farooq et al., 1996).

Reproduction

Spawning takes place in Pakistan at Islamabad from mid- to late March to mid-April (Shaikh and Jalali, 1989, 1991) and near Islamabad (33.3°N, 73.0°E) in April and May (Shaikh and Hafeez, 1993). Gonads begin to develop in December as photoperiod and temperature rise but a continuing warm temperature is the predominant factor for spawning to occur; a fall in temperature halts spawning. Eggs are dark yellow when mature, testes creamy when ripe. Spawning occurs once a year. Up to 150 eggs are recorded in fish from the Ab Garm-e Ganow with a diameter of 1.2 mm.

Parasites and predators

Males were reported as having snout tubercles and tubercles on the anal fin rays (Regan, 1906) but these were the encysted glochidia of a unionid mollusc (B. Prashad in Annandale and Hora (1920). Jalali et al. (1995) describe a new species of monogenean, Dactylogyrus pallicirrus, from fish taken in the Shur River, a Halil River tributary in the Hamun-e Jaz Murian basin. Jalali et al. (2005) summarise the occurrence of Gyrodactylus species in Iran and record Gyrodactylus sp. for fish from the Minab and Halil rivers.

Economic importance

This species is of no economic importance although Butt (1995) suggests that it could be a food source in Pakistan, occurring in shoals of considerable size in rivers that otherwise support little in the form of aquatic protein. It could be cultured as food and as a forage fish. This species has been used to study the effects of heavy metals in Pakistan (Shah, 2002). Higher concentrations of copper and zinc caused lethargy and loss of equilibrium.

Conservation

This species is widely distributed in various basins in southeastern Iran and neighbouring areas and does not appear to under any threat.

Further work

The biology of this species requires study and a thorough review of morphology in relation to named taxa should be carried out.

Sources

Type material: See above and note reservations on type status of some, Cirrhina afghana (BM(NH) 1886.9.21:150-154, BM(NH) 1886.9.21:155-159); Cirrhina afghana var. nikolskii (ZISP 11709); Cyprinion kirmanense (ZISP 11712); Scaphiodon macmahoni (BM(NH) 1905.11.29:27); Scaphiodon microphthalmus (NMW 55897); Scaphiodon muscatensis (BM(NH) 1885.11.7:35-40, BM(NH) 1887.11.11:289-291); Scaphiodon watsoni (FMNH 2303, NMW 51671, NMW 51672 and NMW 51673, ZMB 11042, ZISP 8278, ZISP 8279); Scaphiodon watsoni var. belense (NMW 19833).

Iranian material: CMNFI 1979-0138, 49, ?-66.8 mm standard length, Fars-Hormozgan border, stream in Rasul River drainage (ca. 27º32'N, ca. 54º58'30"E); CMNFI 1979-0143, 22, ? mm standard length, Hormozgan, marsh in Hasan Langi River drainage (27º21'N, 56º50'30"E); CMNFI 1979-0144, 76, ? mm standard length, Hormozgan, Minab River at Minab (27º09'30"N, 57º04'E); CMNFI 1979-0145, 139, ? mm standard length, Hormozgan, Geru River south of Minab (26º55'N, 57º01'30"E); CMNFI 1979-0149, 54, ?-93.8 mm standard length, Hormozgan, stream north of Bandar Abbas (27º36'N, 56º14'E); CMNFI 1979-0150, 34, 43.8-101.6 mm standard length, Hormozgan, stream at Gohreh (27º45'N, 56º05'E); CMNFI 1979-0152, 10, ? mm standard length, Homozgan, Shur River drainage (28º09'N, 55º43'E); CMNFI 1979-0153, 31 24.5-84.7 mm standard length, Fars, qanat stream and pool at Qaleh-ye Biabani (28º31'N, 54º53'E); CMNFI 1979-0154B, 49, ? mm standard length, Fars, stream channels at Koorsiah (28º45'30"N, 54º24'E); CMNFI 1979-0155, 26, ? mm standard length, Fars, springa t gavanoo (28º47'N, 54º22'E); CMNFI 1979-0156, 12, ? mm standard length, Fars, qanat at Rashidabad (28º47'N, 54º18'E); CMNFI 1979-0167, 25, ? mm standard length, Kerman, qanat at Bam (29º06'N, 58º20'E); CMNFI 1979-0168, 50, ?-93.8 mm standard length, Kerman, qanat at Shahabad (29º07'N, 58º16'E); CMNFI 1979-0173, 15, 26.5-84.0 mm standard length, Hormozgan, qanat at Hajjiabad (28º19'N, 55º54'E); CMNFI 1979-0176, 1, ? mm standard length, Hormozgan, Sarzeh River (27º30'30"N, 56º15'30"E); CMNFI 1979-0180, 5, ? mm standard length, Hormozgan, stream 3 km east of Essin (27º19'N, 56º17'30"E); CMNFI 1979-0181, 19, ? mm standard length, Hormozgan, Kul River (27º17'30"N, 56º03'30"E); CMNFI 1979-0183, 14, ? mm standard length, Hormozgan, stream in Rasul River drainage (27º11'30"N, 55º42'E); CMNFI 1979-0185, 4, ? mm standard length, Hormozgan, stream in Rasul River drainage (27º06'N, 55º45'E); CMNFI 1979-0186, 11, ? mm standard length, Hormozgan, strean and pools at Sar Khun oasis (ca. 27º24'30"N, ca. 56º25'E); CMNFI 1979-0187, 54, 18.9-73.6 mm standard length, Hormozgan, stream and pools at Sar Khun oasis (27º23'30"N, 56º26'E); CMNFI 1979-0188, 18, ? mm standard length, Hormozgan, jube at Gohreh (27º45'N, 56º05'E); CMNFI 1979-0189, 25, ? mm standard length, Hormozgan, jube and pool on road to Darab (27º08'30"N, 55º42'E); CMNFI 1979-0190, 44, ?-83.3 mm standard length, Fars-Hormozgan border, spring and pool at Galah Tuyeh (ca. 28º32'N, ca. 55º14'E); CMNFI 1979-0191, 35, 36.6-86.5 mm standard length, Fars, stream 10 km east of Furg (ca. 28º16'N, ca. 55º18'E); CMNFI 1979-0219, 19, 19.1-33.0 mm standard length, Kerman, jube 14 km west of Jiroft (28º37'N, 57º41'E); CMNFI 1979-0220, 4, 28.0-65.5 mm standard length, Kerman, jube 2 km south of Jiroft (28º39'N, 57º43'E); CMNFI 1979-0309, 2, 101.2-108.5 mm standard length, Kerman, Fahraj River at Azizabad (28º57'N, 58º42'E); CMNFI 1979-0310, 1, ? mm standard length, Baluchestan, qanat at Bazman (27º49'N, 60º12'E); CMNFI 1979-0311, 10, ? mm standard length, Baluchestan, Bampur River at Malakabad (27º11'N, 60º27'E); CMNFI 1979-0312, 39, ? mm standard length, Baluchestan, dam on Bampur River (27º11'N, 60º36'E); CMNFI 1979-0313, 68, ? mm standard length, Baluchestan, Bampur River at Bangharabad (27º20'N, 60º46'E); CMNFI 1979-0314, 10, 25.5-118.4 mm standard length, Baluchestan, qanat at Karavandar (27º50'N, 60º46'E); CMNFI 1979-0315, 71, ? mm standard length, Baluchestan, Bampur River 2 km north of Karavandar (27º51'N, 60º46'E); CMNFI 1979-0316, 22, 14.5-69.8 mm standard length, Baluchestan, stream 68 km south of Iranshahr (26º48'N, 61º02'E); CMNFI 1979-0317, 11, 16.5-118.6 mm standard length, Baluchestan, Sarbaz River at Bondan (26º35'N, 61º13'E); CMNFI 1979-0318, 11, ? mm standard length, Baluchestan, Sarbaz River at Huvar (26º09'N, 61º27'E); CMNFI 1979-0323, 6, ? mm standard length, Baluchestan, Sarbaz River (ca. 26º26'N, ca. 61º16'E); CMNFI 1984-0324, 4, ?39, ?51.9-117.8 mm standard length, Baluchestan, Bampur River at Sa'idabad (27º11'N, 60º22'E); CMNFI 1979-0325, 7, ? mm standard length, Baluchestan, qanat at Espakeh (26º51'N, 60º14'E); CMNFI 1979-0326, 10, ? mm standard length, Baluchestan, stream south of Pip (ca. 26º35'N, ca. 60º02'E); CMNFI 1979-0327, 10, 24.0-62.4 mm standard length, Baluchestan, stream in Geh River drainage (26º32'N, 59º57'E); CMNFI 1979-0329, 82, ? mm standard length, Baluchestan, stream at Zaminbandan (27º02'N, 61º20'E); CMNFI 1979-0331, 25, 13.1-50.3 mm standard length, Baluchestan, qanat in Saravan (27º22'N, 62º20'E); CMNFI 1979-0332, 9, 20.8-33.3 mm standard length, Baluchestan, qanat at Kalapurkan (27º14'N, 62º33'E); CMNFI 1979-0334, 4, 26.3-38.4 mm standard length, Baluchestan, Mashkid River (27º04'N, 62º54'E); CMNFI 1979-0335, 2, 66.8-72.2 mm standard length, Baluchestan, qanat at Esfandak (27º07'N, 62º50'E); CMNFI 1979-0338, 17, ? mm standard length, Baluchestan, Tahlab River drainage 8 km from Mirjaveh (28º58'N, 61º24'E); CMNFI 1979-0339, 24, 24.5-76.9 mm standard length, Baluchestan, Tahlab River drainage 16 km from Mirjaveh (28º56'30"N, 61º21'E); CMNFI 1979-0411, 8, ? mm standard length, Hormozgan, Minab River (27º24'N, 57º12'E); CMNFI 1979-0412, 22, 22.0-122.2 mm standard length, Hormozzgan, spring at Saras (27º30'N, 57º34'E); CMNFI 1979-0415, 5, ? mm standard length, Hormozgan, stream south of Ab Garm-e Ganow (27º17'30"N, 56º20'E); CMNFI 1979-0416, 2, 40.1-55.9 mm standard length, Hormozgan, Ab Garm-e Ganow (ca. 27º26'N, ca. 56º20'E); CMNFI 1979-0418, 5, 58.2-111.2 mm standard length, Hormozgan, river near Kahkom (28º09'N, 55º43'E); CMNFI 1991-0141, 3, ?, mm standard length, ; see original sheet? CMNFI 2007-0031, 12, ? mm standard length, Baluchestan, headwater of Bampur River (27º51'N, 60º46'E); CMNFI 2007-0033, 15, ? mm standard length, Baluchestan, Rusgay qanat in Iranshahr (27º13'N, 60º41'E); CMNFI 2007-0034, 3, ? mm standard length, Baluchestan, headwater stream on road to Zaboli (ca. 26º58'N, ca. 61º27'E); CMNFI 2007-0036, 8, ? mm standard length, Baluchestan, qanat at Bazman (27º49'N, 60º12'E); CMNFI 2007-0037, 7, 62.4-166.3 mm standard length, Kerman, Hosseinabad and Gamatabad qanats at Bam (29º06'N, 58º21'E); CMNFI 2007-0038, 9, 62.8-101.2 mm standard length, Kerman, Mehtiabad qanat at Bam (29º06'N, 58º21'E); CMNFI ?, 11, 46.5-69.1 mm standard length, ; 44 ? check, Sirjan on data sheet but no fish at this number in catalogue, check shelf CMNFI 2007-0049, 11, ? mm standard length, Hormozgan, ditches in upper Kol River basin at Hajjiabad (ca, 28º19'N, ca. 55º55'E); 45 CMNFI 2007-0050, 4, 61.2-92.4 mm standard length, Hormozgan, ditches in upper Kol River basin at Hajjiabad (ca. 28º19'N, ca. 55º55'E); 46 CMNFI 2007-0051, 7, 54.6-84.3 mm standard length, Hormozgan, upper kol River basin at Hajjiabad (28º19'N, 55º55'E); 47 CMNFI 2007-0052, 2, 70.7-92.3 mm standard length, Hormozgan, ditch at Qotbabad (27º46'N, 56º06'E); 48 CMNFI 2007-0055, 15, 24.5-75.3 mm standard length, Hormozgan, headwtaer stream in Minab River basin (ca. 27º47'N, ca. 57º12'E); 51 CMNFI 2007-0056, 14, 30.2-70.4 mm standard length, Kerman, qanat at Kahnuj (27º58'N, 57º45'E); 52 CMNFI 2007-0059, 9, ? mm standard length, Fars, Chashmeh Barashk (ca. 27º24'N, ca. 54º06'E); CMNFI 2007-0060, 3, 56.2-93.7 mm standard length, Fars, Chashmeh Ab-e Shirin near Lar (ca. 27º41'N, ca. 54º17'E); 57ID? BM(NH) 1883.8.2:4-9, 5, 42.4-114.9 mm standard length, Baluchestan, Jalq (27º36'N, 62º42'E); BM(NH) 1883.8.2:20-25, 6, 23.1-85.2 mm standard length, Baluchestan, Sib near Dizak (27º15'N, 62º05'E); NMW uncatalogued, 19, 18.1-52.9 mm standard length, Hormozgan, Ab Garm-e Ganow (ca. 27º26'N, ca. 56º20'E). BWC97-4, ?, ? mm standard length, Kerman, Halil River (no other locality data); no fish on cat sheet? BWC97-5, 1, ? mm standard length, Hormozgan, Hasan Langi River (no other locality data);

Comparative material: CAS 28722, 1, 117.4 mm standard length, India, Punjab, Salt Range, Katas Nallah (no other locality data);

Genus Cyprinus
Linnaeus, 1758

The carp genus is found in Europe and Asia and comprises several species of which one has been widely introduced as a food fish.

This genus is characterised by a compressed but heavy body, large size, rounded snout, 2 pairs of barbels, large molar pharyngeal teeth in 3 rows, a very long dorsal fin with the last unbranched ray spine-like and serrated, the anal fin short but with the last unbranched ray spine-like and serrated, the gut is moderately long, and the dorsal and lateral skull bones are sculptured.

Cyprinus carpio
Linnaeus, 1758

Common names

كپور (= kopur, kapur or kapoor (in Gilaki)), kapur-e ainehi (= mirror carp), كپور معمولي (kapur-e ma'mouli or mar'mulleh or ma'muli or maamoli (= common carp)), rashti or كپور رشتي (kapur-e Rashti) in Khuzestan (because their origin was Rasht), mahi-ye gul (= flower fish, meaning in this sense a good fish).

[car or carp shaeeh in Arabic; caki in Azerbaijan; geitan-tsatsan or dliter, both in Armenia; sazan for wild and karp for cultured carp in Russian; common carp, European carp, German carp, wild carp; mirror carp, leather carp, line carp, naked carp (last four referring to scalation), koi (aquarium variety)].

Systematics

Cyprinus Carpio was originally described from Europe. Cyprinus carpio var. caspicus Walbaum, 1792 is described from the mouth of the Volga and Don rivers but is infrasubspecific and the name has not been used in Iran nor has Cyprinus carpio fluviatilis Pravdin, 1945 described from floodplain lakes of the Volga River near Saratov.

Key characters

This species is easily identified by the long dorsal fin, the spine in both the dorsal and anal fins, and the two pairs of barbels.

Morphology

Dorsal fin with 2-5 unbranched rays followed by 14-23, usually 18-20, branched rays, anal fin with 2-4, usually 3, unbranched rays followed by 3-7, usually 5, branched rays, pectoral fin branched rays 13-19, and pelvic fin branched rays 5-9, usually 8. The dorsal fin has the last unbranched ray developed as a toothed spine and the anal fin has a similar spine. Lateral line scales 26-41, mostly 36-39. Scales may be absent (leather carp), restricted to a few, enlarged scales (mirror carp), or only a mid-lateral row of scales (line carp), in cultivated varieties. Wild carp are fully scaled. Individual scales have a central focus, wavy anterior margin, few radii on the anterior and posterior fields in young fish, and medium numbers of radii on fish 12-14 cm standard length. There are numerous fine circuli and the posterior scale field breaks up into bubble-like structures. Gill rakers 17-29 (some literature counts may be lower arm of arch only and there may be size-related variation too) and vertebrae 32-39 (lower counts may not include Weberian vertebrae). Rakers touch the second raker below when appressed and have a row of knobs on their medial surface. Pharyngeal teeth 1,1,3-3,1,1 with variants 1,2,3-3,2,1, 1,2,3-3,1,1, 1,1,1,3-3,1,1, 1,1,3-2,1,1 and 1,3-3,1,1. Posterior major row teeth are large with flattened crowns bearing wavy ridges while more anterior teeth are a rounded knob, or even concave on top of the knob. The gut is elongate with several coils. This species is a tetraploid (2n=98-104)(Al-Sabti, 1986; Klinkhardt et al., 1995).

Meristic values for Iranian specimens are:- dorsal fin branched rays 16(1), 17(5), 18(18), 19(16), 20(10); anal fin branched rays 4(1) or 5(49); pectoral fin branched rays 14(1), 15(7), 16(25), 17(16) or 18(1); pelvic fin branched rays 6(1), 7(6) or 8(43); lateral line scales 34(2), 35(6), 36(10), 37(26) or 38(6); total gill rakers 18(1), 20(2), 21(11), 22(11), 23(15), 24(4), 25(4) or 26(2); total vertebrae 36(1), 37(19), 38(26) and 39(6); and pharyngeal teeth ?. Note that these samples may include individuals which are not native but introduced for fish farming from stocks outside Iran.

There are morphometric and meristic differences between carp from the southwestern and southeastern Caspian Sea but these are ecological not taxonomic. Also, carp from the Anzali Mordab differ from those in the sea by having a longer head, snout, eye and postorbital region (although of course some of these are redundant), greater interorbital width, shorter predorsal and preanal distances, shorter dorsal, anal, pectoral and caudal fins, and a lower anal fin (A. M. Shukolyukov in Berg (1948-1949)). Yousefian (2004) found carp from the Caspian Sea in Iran had a dominant genotype different from those in a fish farm. Meristics (scales and fin rays) and morphometrics (head length and body width) also differed.

Sexual dimorphism

Females are deeper bodied than males because of their eggs and the distance between the pectoral and pelvic fins and the pelvic and anal fins is more. Dorsal and anal fins in males are higher, the anal fin is longer at the base, the pectoral fin is longer and the lobes of the caudal fin are longer. This is accounted for by the greater swimming activity of males during spawning (Kuliyev and Agayarova, 1984). Breeding males have fine tubercles on the head, particularly on the anterior operculum and preoperculum and under the eye, above the lateral line and more frequently below it, and on the fin rays.

Colour

In semi-diadromous carp from the Kura region of the southwest Caspian Sea, the overall body colour is dark yellow, the flanks being golden-yellow with dark shading, and the back is black. The belly and fins are light yellow and the caudal fin is reddish. Lake populations are darker. Young fish from Iran are silvery on the flanks (but not as bright as Carassius auratus), greyish on the back, silver-pearl on the belly, the iris is silvery with grey above and below, the dorsal fin and upper caudal lobe are pale grey, the lower caudal lobe and anal fin are orange, the pelvic fin is pale orange, and the pectoral fin has only traces of orange. The caudal fin may be yellow-orange with lobe margins red. The freshwater resident form in the Anzali Mordab is yellowish, the semi-diadromous form dark. The peritoneum is grey to silvery and may be speckled.

Size

Carp resident in fresh waters are smaller than semi-diadromous carp. In the 1950s in Iran, carp catches were 20-41 cm long (Farid-Pak, no date). Fish up to 1.0 m long are caught in the Caspian basin (A. Abdoli, pers. comm., 1995). Maximum size exceeds 1.2 m and 44.9 kg, possibly to 1.5 m and 69.6 kg.

Distribution

Widely introduced in the Middle East for aquaculture. Found naturally in Iran in the whole Caspian Sea drainage, it is also widely stocked in the provinces of Gilan and Mazandaran (Petr, 1987; Abbasi et al., 1999). It is reported from the Atrak, Gorgan, Gharasu, Tajan, Babol, Aras, and Safid rivers, the Anzali Mordab and the Siah-Keshim Protected Region, Gorgan Bay, the southeast Caspian Sea, southwest Caspian Sea and south-central Caspian Sea as an introduced species, and in all these plus the Haraz and Pol-e Rud rivers as a native species (Riazi, 1996; Kiabi et al., 1999).

It is probably native to the Tedzhen River of Turkmenistan (the Hari River or Harirud in Iran) (Aliev et al., 1988). This species is also recorded from the Karakum Canal and Kopetdag Reservoir in Turkmenistan (Shakirova and Sukhanova, 1994; Sal'nikov, 1995) and may eventually reach Iranian waters in the Tedzhen (= Hari) River basin. These latter stocks may be introduced.

It has been introduced to the Sistan basin to a canal flowing into the Chahnimeh, the Sistan Dam, Hamun Lake and the Hamun Kushk (Ahmadi and Wossughi, 1988; J. Holčík, in litt., 1996; Shamsi et al., 2009)), to the Karaj River in the Namak Lake basin, to the Mahabad Dam (Abdi, 1999; www.mondialvet99.com, downloaded 31 May 2000), to the Haft Barm lakes near Shiraz in 1984 (Petr, 1987), to the Manjil Reservoir on the Safid Rud (Nümann, 1966), to Mehr Dasht Lake, Markazi, to the Zayandeh River Dam (Y. Keivany, in litt., 1992; Ghorbani Chafi, 2000), to Lake Zaribar, Kordestan (Abzeeyan, 5(5):III, 1994), to Lake Famur from nearby fish farms (Maafi, 1996b), to the Kor River basin in Fars (A. Alamdari, in litt., 1997; M. Hafezieh, in litt., 1997), throughout Khuzestan Province (not caught in the 1970s), and to numerous other water bodies and ponds throughout Iran where it reproduces in the wild. Mirror carp found in the Shadegan marshes of Khuzestan are escapees from fish farms. Carp in the Bahu Kalat and Sarbaz rivers of Baluchestan are also escapees from fish farms (A. Mobaraki, pers. comm., June 1999).

Abdoli (2000) records this species as present generally in the Dasht-e Lut, Sistan, Dasht-e Kavir and Kerman-Na'in basins; in lower reaches of rivers from the Gorgan to the Astara and along the Caspian coast, the middle Aras River; the Kashaf River in Khorasan; the middle and lower Talkheh River and Tatavi River in the Orumiyeh basin; the Qareh Chai and Qom rivers in the Namak Lake basin; the middle to lower Zayandeh River in the Esfahan basin; the Aravand, lower Karun, Jarrahi, lower Karkheh rivers in the Tigris River basin; the middle Halil River in the Hamun-e Jaz Murian basin; and the middle and upper Kul River including the Shur River tributary in the Hormuz basin. Jalali et al. (2005) records this species from the Dez, Karun and Zayandeh rivers, the Vahdat reservoir, and the Zarivar and Kaftar lakes.

Zoogeography

The natural distribution of the carp is supposed to be Asia Minor and the Caspian Sea basin where its origins lie in the late Pliocene. From this area, modern wild carp spread east and west, perhaps as late as the last postglacial thermal optimum, and latterly aided by man (Balon, 1974; 1995; Van Damme et al., 2007). Iranian carp may be a mixture of native and introduced stocks (or species - see Kottelat (1997) - but this remains unresolved). It is probably not now possible to distinguish the native stocks morphologically because of admixtures of farmed stocks. All domestic forms probably originated from native Danube River stocks (Bănărescu, Barus and Peňáz in Bănărescu and Paepke, 2002). Khalili and Amirkolaie (2010) compared fish electrophoretically, meristically and morphologically from the Anzali lagoon, Qareh Su and Bandar Gaz. Morphometric differences between east and west were attributed to a relatively smaller body size in the east and the influence of domestic stocks. Electrophoretic differences, particularly obvious at Anzali, were attributed to the larger number of fish farms in this latter area.

Habitat

This species favours an abundance of soft vegetation in shallow water, necessary for successful reproduction. Still waters are preferred but they are found in the lower courses of lowland rivers with moderately flowing water, and occasionally in water exceeding 2 m/sec. They can often be seen basking at the surface or feeding on algae and their dorsal fins break the water surface. Large fish often move into shallows in the afternoon and evening. Carp also leap from the water but the reason is unknown. They rarely descend below 30 m in lakes and avoid fast water in streams. Carp overwinter in depressions in the lower reaches of rivers in the Caspian basin. There are both freshwater resident populations in the Caspian basin and diadromous (or semi-diadromous) ones, the latter living in brackish water near river mouths and only entering fresh water to spawn. The migration up the Ural River may reach 60 km from the river mouth. Some fish apparently spawn in the brackish waters of shallow coastal areas in the Caspian. Knipovich (1921) reports fish in the Iranian Caspian Sea down to 11.9-12.3 m. Riazi (1996) and Karimpour (1998) report that this species is both native (resident) to, and migrates to, the Siah-Keshim Protected Region of the Anzali Mordab.

Carp have a salinity tolerance under experimental conditions of up to 8‰, and for short periods 18.6‰ with acclimation, and this has significance for survival of carp in the Caspian Sea and in waters of southern Iran and Iraq where this species is farmed. Hafezamini and Oryan (2002) and Hafezamini et al. (2003) however found that, under experimental conditions, all fish in their study died at 18‰ in less than 12 hours. Carp eggs hatch in water up to 10‰, with the favourable level being up to 6.6‰ (Al-Hamed, 1971).

Low dissolved oxygen concentrations of 3 mg/l are tolerated and levels as low as 0.5 mg/l can be withstood for 2-3 hours. Normal growth has occurred in fish kept at 35°C. The immune response of carp following exposure to the organophosphate malathion has been studied by Soltani et al. (2003).

This species is considered to be a dominant species in the Karun River along with Barbus (= Tor) grypus (Iranian Fisheries Research and Training Organization Newsletter, 17:1, 1997).

Introduced to Iraqi waters in 1960 as juveniles, this species rapidly became established (Ahmed and Taher, 1988). Cyprinus carpio was caught in large numbers in the Shatt al Arab of Iraq down to the estuary after an increase in the discharge of the Tigris River reduced salinity (N. A. Hussain, in litt., 1994).

Age and growth

Females are larger and mature a year later than males. Sexual maturity is attained in the second year of life, and in a few individuals even by the end of the first year, in the southeastern Caspian Sea; but in the southwestern Caspian Sea this occurs in the third and fourth years (Kuliyev and Agayarova, 1984). Resident carp in Dagestan mature in their third year at about 30 cm and have an average life span of 6 years whereas the semi-anadromous or semi-diadromous form matures in its fourth year at 35-36 cm and has an average life span of 8 years (Shikhshabekov, 1969). Growth is faster in the Kura River of Azerbaijan than in other populations in the Caspian but maturity is later at age 4 or more usually at 5 years. Freshwater residents in the Anzali Mordab are slow-growing compared to the semi-diadromous form and are less common. Fish taken in the commercial operations in Iran were 3-7 years old, 31.0-63.0 cm long and weighed 539-3375 g (Razivi et al., 1972).

This species was stocked in the Dukan and Derbendikhan dams of Iraq in the 1960s where fish up to 3 years of age were reported by Ciepielewski et al. (2001). Decreasing growth rates indicate conditions are not too favourable although growth in the first two years is comparable with that in lakes of central Iraq. The condition coefficient (K) was higher among smaller fish, e.g. fish from Dukan at 230 g had a K of 2.32, at 1 kg K was 1.75. Ahmed and Taher (1989) examined the growth of 0+ carp in Hawr al Hammar, Iraq found the length-weight relationship to be W = 0.00004627 L2.8022 and derived the growth equation Lt = 189.87 (1-e-0.0158 (t + 25.662)) with a calculated length at the end of the first year of life of 189.87 mm. This is relatively larger than for other parts of the world, indicating a successful introduction of this exotic. Al-Nasiri and Dawood (1991) found the smallest mature male was 182 mm and the smallest female was 184 mm in Hawr al Hammar, with sexual maturity achieved in the first year of life. Maximum life span was 8+ years. Epler et al. (2001) found the oldest age groups in Iraqi lakes to be 5+ in Lake Habbaniyah and 3+ in Lake Razzazah. The mean condition factor was 1.47 for Lake Habbaniyah and 1.50 for Lake Razzazah.

In Sariyar Dam Lake near Ankara in central Anatolia, ages range from 0 to 18 years (Ekmekci, 1996b). In their first year, fish have an average fork length of 103 mm and weigh 24 g, in 5 years they average 357 mm and 822 g, and in 10 years 580 mm and 3365 g. In Gölhisar Lake by contrast, a small water body in western Turkey, age composition was from 1 to 6 years and fish attained a maximum of 494 mm and 1922 g (Alp and Balik, 2000).

Maximum life span for this species is reported as 47 years for domestic fish.

Food

Food is derived from browsing on the substrate at all hours, if the temperature is favourable. Browsing muddies the water and can inhibit other species and uproot plants. Mouthfuls of bottom ooze are taken up, spat out and the food items selected. These include aquatic insects, crustaceans, worms and molluscs, and more rarely, fish. Plant material is ground up by the molar pharyngeal teeth and includes algae, seeds, wild rice, leaves and various aquatic plants. Organic sewage is also eaten. Some surface feeding on algal mats or insects will also occur. Feeding almost completely stops in winter and the fish go into a form of hibernation.

Feeding in the Hawr al Hammar, Iraq is related to temperature, the peak intensities being July and the minimum in January with peak activity in September and minimum in January. Feeding does occur year round and smaller fish (<200 mm) have highest feeding activity in spring while adults have this in summer (Hussein et al., 2000a). In a study of the recovering Hawr al Hammar, diet was 25.45% algae, 18.18% snails, 12.73% diatoms and 12.73% copepods, 10.91% insects with plants, cladocerans and shrimps at less than 10% each, in the Hawr al Hawizah diet was 27.3% snails, 18.2% insects and 12.1% for algae, plants and cladocerans with fish, diatoms and copepods at less than 10% each, and in the Al Kaba'ish (= Chabaish) Marsh 33.3% algae, 20.4% insects, 11.1% snails and diatoms and plants at 10.2, with various crustaceans at less than 10% each (Hussain et al., 2006). Hussein et al. (1991) examined diet in the Garma Marshes, Iraq and found crustaceans, molluscs, aquatic plants and seeds, aquatic insects, oligochaetes and fish to be dietary items, selection and numbers varying with carp size and season. Some fish were found to have fed exclusively on only a single, different mollusc species, presumably as opportunity presented. Al-Shamma'a et al. (1996) examined the food of this species in Al Qadisiyah Reservoir, Iraq and found plants, their seeds, molluscs and aquatic insects, all bottom foods. Mangalo and Akbar (1988a, 1988b) studied the food of carp in a farm pond at Al-Latifiyah, Baghdad where zooplankton was the principal diet. The gill rakers show an efficient structure for filtration, indicative of a phytoplanktivorous and omnivorous feeding (Salman et al., 1993). Salman et al. (1994) report a mixture of animal and plant foods, with zooplankton a dominant component of all length groups. The gut is coiled and 3.42 times standard length, indicating omnivory with plant food being important. Epler et al. (2001) gave the diet in Lake Habbaniyah, Iraq as 51.7% plants, 15.7% oligochaetes, 15.2% tendipedids, 7.2% molluscs, 5.2% detritus and 4.1% cladocerans. They also found that where there is significant competition between autochthonous species, as here, carp become another strong competitor for food. Dietary coincidence between carp and shabbout, gattan and himri was 58.5, 68.5 and 54.2 % respectively. Hussein et al. (2000b; 2000c) examined dietary overlap between this species and three native carps in the Hawr al Hammar. Overlap with Barbus (= Mesopotamichthys) sharpeyi was the weakest as this species is a herbivore but small B. luteus (<200 mm) and B. xanthopterus showed strong overlaps. This overlap may explain the decline in some native carps. Hussain and Ali (2006) also examined feeding relationships among fishes in the Hawr al Hammar and found this species to be a carnivore, 26.4% of the diet being crustaceans, 12.7% insects and 30.5% molluscs. Dietary overlap of 84% was found between this species and Barbus (= Luciobarbus) xanthopterus but the availability of food resources offset possible competition, contrasting with the conclusions above. Ciepielewski et al. (2001) found the diet of this species in the Dukan and Derbendikhan reservoirs in Iraq to be mainly algae, copepods and chironomids.

Reproduction

Under natural conditions, males spend more time on the spawning grounds than females and spawn several times. More than 7 million eggs up to 1.71 mm in diameter may be present in a female but only about 500 are laid at a time. Wild carp in the Atrek River had a fecundity range of 16,000 to 543,000 eggs (Bănărescu, Barus and Peňáz in Bănărescu and Paepke, 2002). The resident form is less fecund by about half than the semi-anadromous form (Shikhshabekov, 1969). In the Anzali Mordab a mass spawning run takes place in April with spawning in April-May. The first migratory fish are seen as early as January. Shallow weedy areas or the mouths of rivers are used as spawning sites and adhesive eggs are laid on plants.

Resident populations in Dagestan spawn earlier, by about a month, than the semi-anadromous population which spawn in early May (Shikhshabekov, 1969). Spawning time variations are governed by temperature and the most favourable temperature is 18-20°C.

Carp ascend the Kura River of Azerbaijan in spring and autumn. The spring run begins in mid-March and peaks in April while the weak autumn run lasts from August to mid-October.

Carp have feeding grounds in the coastal waters of the southeastern Caspian Sea and enter the Atrak River in winter to spawn between February and April. Young migrate downstream, this movement ending in July when the river flow is minimal or ceases. When there is no flood, spawning does not occur (Petr, 1987).

Fish in Iraqi ponds grew 25-30 cm in the first year of life and matured in 1-2 years. At 16-26ºC they spawned from late February to late April and again in the autumn (Al-Hamed, 1960). Palm tree fibres were used for egg deposition and eggs hatched in 4-8 days. Al-Nasiri and Dawood (1991) found a fecundity range of 14,150-1,492,500 eggs in Hawr al Hammar with a mean relative fecundity of 182 eggs/g of body weight, and a egg diameters of 0.90-1.02 mm. The gonadosomatic index indicated spawning in March and possibly October-November. Epler et al. (2001) studied reproduction in lakes Tharthar and Habbaniyah and found both sexes to achieve maturity in the first year of life at 13.5 cm for males and 12.6 cm for females. Spawning occurred in May and fecundity was 186-531 thousand eggs/kg body mass.

The spawning behaviour involves stimulation of a female while moving over vegetation and being accompanied by 2-3 males, active movement and spawning being induced by blows from the male(s). The eggs adhere to the vegetation or are lost. Most eggs are shed at night or in the early morning.

Parasites and predators

Eslami and Anwar (1971) record the cestode Caryophyllaeus fimbriceps from this species on the Caspian coast of Iran. Mokhayer (1976b) records the cestodes Bothriocephalus gowkongensis, the nematode larva Anisakis and the acanthocephalan Pomphorhynchus perforator. 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 (1990b) variously record the monogeneans Dactylogyrus anchoratus, D. extensus, D. sahuensis and D. vastator from carp on fish farms throughout Iran and Naem (2002) records D. anchoratus from fish in the Safid River.

Moghainemi and Abbasi (1992) record a wide range of parasites from this species in the Hawr al-Azim in Khuzestan. Mortazaei et al. (2000) record an infection rate of 66% (2 of 3 fish) with the worm Bothriocephalus opsariichthydis in Khuzestan marshes. Sattari and Faramarzi (1997) record Caryophyllaeus fimbriceps, C. laticeps and C. brachycollis from 38% of carp in the Anzali lagoon. Akhlagi (1999; 2000) 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 3.8% of fresh and smoked samples of this species in Mazandaran Province. 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. Farahnak (2000) records Anisakidae from this species in Khuzestan. 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. Masoumian et al. (2002) investigated parasites from this fish in the Aras and Mahabad dams in northwest Iran and found the protozoan Goussia carpelli, also known from carp in the Safid River. Ebrahimzadeh Mousavi et al. (2005) isolated the fungus Branchiomyces spp. from gill lesions of farmed carp in northern Iran. Branchiomycosis or gill rot is a major problem in commercial fish production.

The intestinal helminth Bothriocephalus gowkongensis was recorded from this species on fish farms in West Azarbayjan Province (Azarvandi et al., 1999). Naem et al. (2002) found the following parasites on the gills of this species from the western branch of the Safid River, namely the protozoans Ichthyophthirius multifilis and a Trichodina species, a copepod crustacean Lernaea sp., monogenean trematodes Dactylogyrus anchoratum, and D. achmerowi. Jalali et al. (2002) and Jalali and Barzegar (2006) record parasites from this species in Lake Zarivar, namely Trichodina pediculus, Dactylogyrus extensus, Gyrodactylus stankovici, Diplostomum spathaceum two species of Argulus, Pseudocapillaria tomentosa and Lernaea cyprinacea. Jalali et al. (2005) summarise the occurrence of Gyrodactylus species in Iran and record G. cyprini, G. elegans, G. shulmani, G. sprostonae, G. stankovici and G. sp. from various localities for the carp. Farahnak et al. (2002) record Contracaecum sp. and Anisakis sp. from this fish in Khuzestan Province. Mehdipoor et al. (2004) record the monogenean Dactylogyrus extensus in Zayandeh River fish. Araghi Soureh and Jalali Jafari (2006) recorded Dactylogyrus extensus from this species in the Mahabad River of the Lake Orumiyeh basin. Jalali and Barzegar (2005c) record five species of monogeneans in the genus Dactylogyrus from both farmed and native Cyprinus carpio in Iran. These are D. extensus, D. anchoratus, D. achmerovi, D. vastator and D. sahuensis. Fry and fingerlings are more sensitive to these parasites and this sensitivity is increased with crowding in ponds. The paper also deals with gill histopathology and distribution of the parasites in Iran. Barzegar and Jalali (2006) report parasites in this species from Kaftar Lake as Lernaea cyprinacea,  Trichodina sp., Dermocystidium sp., Dactylogyrus extensus, D. anchoratus, Gyrodactylus sp. and Diplostomum spathaceum. Khara et al. (2006b) record the cestode Caryophyllaeus fimbriceps from this species in the Boojagh Wetland of the Caspian coast. Pazooki et al. (2007) recorded various parasites from localities in West Azarbayjan Province, namely Diplostomum spathaceum, Ligula intestinalis, Digrama sp. and Argulus foliaceus. Sattari et al. (2007) record the cestode Caryophyllaeus fimbriceps, the digenean Diplostomum spathaceum and the monogeneans Dactylogyrus extensus, Gyrodactylus sp. and Diplozoon sp. in this species in the Anzali wetland of the Caspian shore and also mention that the Caryophyllaeus laticeps is also known from this species in the Iranian Caspian Sea. Shamsi et al. (2009) found Dactylogyrus achmerovi, D. anchoratus, D. extensus, D. sahuensis and D. vastator in this species in fish farms, Safid River and Hamun Lake.

The Caspian seal, Pusa caspica, is a significant predator on this species (Krylov, 1984). Esox lucius, Sander lucioperca and Silurus glanis are all predators on small carp. A wide variety of other fishes and birds eat smaller carp as do predatory aquatic insects, frogs and toads. Carp eggs are eaten by many fishes. Adult carp are too large for most predators to take.

Economic importance

Balon (1974; 2006) details domestication of the carp which has been a cultivated fish for over 2000 years. The carp is an important table fish in Iraq where details of farming techniques are given by Ahmed and Taher (1988) and by Mangalo and Akbar (1988a; 1988b). Catches in former Soviet waters of the Caspian Sea were described by Kuliyev and Agayarova (1984). In the southwestern part, size range was 21-64 cm in 1960-1961 (92% of fish were 29-41 cm) but in 1972 95% of fish were 31-55 cm. The smallest carp are caught in the southeastern Caspian where maximum weight was 3.97 kg as opposed to 10.0 kg in the central Caspian. Spawning fish in the southeastern Caspian were only 0.58-1.0 kg while in the southwestern Caspian carp were 0.25-5.75 kg. About 17-30% of commercial catches in the Volga region are estimated to be comprised of hatchery production (Petr, 1987).

Nevraev (1929) gives catches for various fishing regions in Iran in the early twentieth century. For the Safid River region from 1899-1900 to 1917-1918 the catch was 30 to 30,500 individuals with no fish reported in some years, and in Astrabad (= Gorgan) region from 1900-1901 to 1912-1913 the catch was 14,200 to 851,800 individuals. The commercial catch in Iran from 1956/1957 to 1961/1962 varied from 3443 kg to 175,295 kg (Vladykov, 1964), from 1965/66 to 1968/69 varied from 184 to 333 tonnes (Andersskog, 1970) and from 1963 to 1967 from 7.0 to 108.8 t (on a yearly basis 41.1, 9.8, 7.0, 108.8, and 48.1 t respectively) (RaLonde and Walczak, 1970b). Catches from 1933/34 to 1961/62 varied from 34 kg to 1113 t in the Bandar Anzali region and the total catch of the Northern Shilot (Fisheries Company) varied from 9.8 t to 333 t from 1963/64 to 1968/69 (RaLonde and Walczak, 1972). The Food and Agriculture Organization, Rome gave the catches from Iran for the 6 years 1980 to 1985 as 1032, 2000, 1000, 52, 83, and 100 t respectively. The catch in the Anzali Mordab in 1990 was 6855 kg and from 1932 to 1964 the annual catch varied from 1.2-638.6 t (Holčík and Oláh, 1992). These wide fluctuations in reporting and in nominal catches are indicative of the poor fisheries statistics as well as the inter-year variability of catches in Iran.

March and April are the main fishing months in Iran (Farid-Pak, no date). It is the main fish in Iranian carp farms, when raised in polyculture with Chinese carps (Jalali and Molnár, 1990b). Salehi (2004) gives a figure of 25% as the share of common carp in Iranian polyculture (63% being silver carp) and gives a review of the economics of carp polyculture. Kohnehchahry and Heydarpur (1973) outline methods of raising carp using submerged cages which they believe would be suitable for Iranian waters. There were about 4000 ha of carp fish ponds (which presumably included Chinese carps such as Hypophthalmichthys spp.) with an annual production of 2-3 tonnes/ha (White, 1988). Plans were made in the 1980s to increase total pond area (including trout which was at 60 ha and produced 1000 t/year) to 35,000 ha over 10 years to yield an annual harvest of 100,000 t/year. However Bartley and Rana (1998b) report an aquaculture production of 6561 tonnes in 1995 and Salehi (2004) gives 28,060 t from warm-water fish farming, mostly Chinese carps. More than 700,000 carp fingerlings were released in the small and remote province of Chahar Mahall va Bakhtiari alone and 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). Salehi (1999) states that the marketable size of cultured carp is 1 kg with most harvested once annually and almost 90% supplied to market in October-March with a peak in March (the Iranian New Year when fish is a traditional food).

Marjan Iran Company was selling 1500-1800 g fish for U.S.$1.90/kg in August 2003 compared to $2.10 for Ctenopharyngodon idella (http://groups.yahoo.com/groups/hilsa/message/25). In Golestan Province, carp cost 1500-1700 tomans/kg in the early 2000s. Salehi (2006a) analysed the consumer market for this species and its products in Iran.

An exotic species in some parts of the world, carp are a nuisance because they uproot vegetation used by native species for cover, food and spawning. This activity also increases water turbidity to levels which many native species cannot tolerate. Stirred up silt may smother eggs of native species. Carp also compete with other species for food and eat the eggs of other species.

Various studies have been carried out on its culture in Iran and neighbouring countries, e.g. see Freshwater Fishes of Iraq, and its use as an experimental organism, e.g. in Iran, see Mohagheghi and Hedayatifard (Farsi translation of Horváth et al. (1985a and 1985b); Khazraiinia et al. (2000) on acute ammonia toxicity; Rostami Bashman et al (2001) on histopathological lesions after exposure to copper, zinc, mercury and cadmium compounds; Soltani et al. (2001) on the anti-fungal effectiveness of formalin on hatch rate of eggs; Arabi and Heydarnejad (2002) on the deleterious effects of copper and mercury, used in combating algal blooms and weeds, on the gills of carp; Abtahi et al. (2002) on clove oil having no significant difference with MS222, another anaesthetic used in fish farms; Dorafshan et al. (2003) on induction of spawning using pituitary extract and GnRH analogue in combination with domperidone; Sharifpour et al. (2003) on the anaesthetic effects of clove oil under various pH and temperature regimes; Vahabzadeh Roodsari et al. (2003) comparing malachite green and hydrogen peroxide to control fungal infections (the latter is effective and less dangerous); Akhlaghi et al. (2004) on phagocytosis in relation to immunostimulants; Ebrahimi (2004) on the deleterious effects of copper, a pollutant, on sperm anatomy; Moini and Basimy (2004) on production of fish cake according to various recipes and its shelf life; Sharifpour (2004) on the histology of the response and the circumstances of wound healing; Oryan et al. (2005) on  the effects of baclofen on the pituitary system; Sharifpour et al. (2005) on the highly toxic effects of the pesticide endosulfan (with an LC50 of less than 0.1 mg/l); Yousefian (2005) on generating gynogenetic carp through irradiation; Baradaran Noveiri et al. (2006) on cryopreservation of spermatozoa using different extenders; Ghiasi and Mirzagar (2006) on lysozyme content in sub-lethal concentrations of cadmium; Kazemipour and Keivany (2006) on the use of garlic, mallow and motherwort in healing superficial wounds - garlic reduced recovery by one week; Kazemipour et al. (2006) on carp kept in aquaria at low concentrations of garlic (0.1 g/L) which healed superficial wounds more quickly than controls or aquaria having mallow or motherwort; Kohodabandeh and Abtahi (2006) on the use of sodium chloride, iodine and formalin to control Saprolegnia sp. on eggs (sodium chloride was recommended); Naji et al. (2007) on the 96h LC50 value of cobalt chloride at 327-328 mg/l; Naji et al. (2007) on the toxic effect of zinc sulphate on gill tissues which suffer hypertrophy and hyperplasia; Darafsh et al. (2008) on the use of scales as an indicator of heavy metal pollution; Safari et al. (2008) on changes in muscle chemistry during maturation; Rostami Mina and Soltani (2009) on the histopathological effects of the aquaculture pollutant copper sulphate; Sheykhzadeh et al. (2009) on the effects of Eucalyptus essential oil on immunological variables; Ghiasi et al. (2010) on the effects of low concentrations of cadmium on the immune response in winter; Soltani et al. (2010) on the immune responses to Zataria multiflora (Persian thyme) essential oil used as an anti-fungal for carp eggs; etc.

This species is actively angled for along the Caspian shore of Iran and in its rivers (e.g. see Noorbakhsh (1993a)), appears regularly in fish markets of Ahvaz, Khuzestan and is caught by anglers there using bread or potato as bait.

It is characterised as a fatty fish according to a lipid content 9-14% by wet weight of muscle in autumn (Hantoush et al., 1999). Al-Aswad et al. (1980) detailed the chemical composition of this species in Dukan Lake, Iraq including seasonal levels of moisture, fat, protein and ash, and the various types of fatty acids and amino acids. Hindi et al. (1996a) give the chemical composition of flesh of this species as 78.87% moisture, 2.46% fat, 17.06% protein and 1.35% ash, indicating a valuable food fish characterised as lean to medium fatty. Hindi et al. (1996b) give chemical indices for assessing fish freshness according to the month of capture and marketing (pH 6.28, total volatile nitrogen bases 11.07 mg/N/100g fish, thiobarbituric acid 0.47 mg, and free fatty acids 0.62%).

The roe or eggs of this species have been implicated in poisoning (Halstead, 1967-1970; Coad, 1979b) and should be avoided (see under the genus Schizothorax for more information on egg poisoning). Fish should be carefully cleaned in the spawning season to remove the eggs and ensure against contamination of flesh. Severe cases of egg poisoning in other species have resulted in death.

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 textbooks, for sport, as an experimental species and because it has been introduced outside its natural range. Brightly coloured varieties of carp are known as "koi" and are kept as ornamental fish. Colours include red, orange, white, black, blue and yellow in various combinations.

Conservation

Vladykov (1964) recommended that fishing for this species in Mazandaran and Gorgan be prohibited for 5 years because of reduced stocks. 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. In 1999-2000, 20 million juveniles were released into the Caspian Sea (Iranian Fisheries Research Organization Newsletter, 23:4, 2000). From October to March 2000, 3 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). Poaching was a problem in the Caspian basin of Iran (Razivi et al., 1972) and no doubt continues. The Atrak River stocks are an important fishery for both Iran and Turkmenistan but are susceptible to loss through the absence of flooding of the spawning grounds. Fish passes are needed to ensure access to the spawning grounds, timely release of water from a reservoir to flood the spawning areas in years of low water flow, enforcement of catch limits, and continued stocking (Petr, 1987). Yousefian et al. (2009) examined the normative reproduction values and genetic characters for native carp in the Caspian Sea. The average weight of sampled fish was 1441.6 g and standard length was 50.5 cm. Fertilisation rate was 62-79%, absolute fecundity was 114,347 and relative fecundity 82,007. There was no significant differences in haplotype distribution between areas in the southern Caspian Sea.

Lelek (1987) considered that the wild form, as opposed to domestic stocks, was vulnerable to endangered in Europe because of habitat modifications. 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), present in other water bodies in Iran, and present outside the Caspian Sea basin. Karami et al. (2005) found river fish had greater exposure to organophosphorus compounds than Caspian Sea and well-fed fish.

Further work

Where this species is introduced, its interactions with native species should be carefully studied and monitored in an effort to protect the native fauna.

Sources

A recent summary on this species is Bănărescu, Barus and Peňáz in Bănărescu and Paepke (2002).

Iranian material: CMNFI 1970-0510, 1, 47.8 mm standard length, Gilan, Golshan River (37º26'N, 49º40'E); CMNFI 1970-0522, 1, 108.5 mm standard length, Gilan, Safid River at Astaneh Bridge (37º16'30"N, 49º56'E); CMNFI 1970-0563, 3, 28.6-58.6 mm standard length, Gilan, Caspian Sea at Kazian Beach (ca. 37º29'N, ca. 49º29'E); CMNFI 1970-0568, 1, 125.8 mm standard length, Gilan, Caspian Sea at Kazian Beach (ca. 37º29'N, ca. 49º29'E); CMNFI 1970-0582, 3, 63.3- 134.0 mm standard length, Mazandaran, Aliabad Reservoir (36º56'N, 54º50'E); CMNFI 1970-0587, 2, 47.5-76.7 mm standard length, Mazandaran, Babol River at Babol Sar (36º43'N, 52º39'E); CMNFI 1979-0455, 6, 46.9-67.1 mm standard length, Markazi, Manjil Dam (36º45'N, 49º17'E); CMNFI 1979-0476, 3, 44.1-85.3 mm standard length, Mazandaran, Qareh Su near Kord Kuy (36º51'N, 54º05'E); CMNFI 1979-0479, 19, 28.0-143.8 mm standard length, Mazandaran, dam on Gorgan River (37º09'30"N, 54º41'30"E); CMNFI 1979-0685, 2, 31.5-45.7 mm standard length, Gilan, Safid River (ca. 37º22'N, ca. 49º57'E); CMNFI 1979-0788, 2, 104.2-123.5 mm standard length, Mazandaran, Gorgan River at Khadje Nafas (37º00'N, 54º07'E); CMNFI 1980-0128, 3, 60.2-75.2 mm standard length, Mazandaran, Qareh Su (36º49'30"N, 54º03'30"E); CMNFI 1980-0132, 4, 56.0-64.6 mm standard length, Gilan, Safid River at Kisom (37º12'N, 49º54'E); CMNFI 1980-0157, 1, 126.3 mm standard length, Mazandaran, Gorgan River estuary (36º59'N, 53º59'30"E); CMNFI 1980-0905, 2, 78.1-92.9 mm standard length, Mazandaran, Gorgan River at Khadje Nafas (37º00'N, 54º07'E); CMNFI 1980-0908, 1, 54.4 mm standard length, Gilan, Safid River estuary (ca. 37º28'N, ca. 49º54'E).

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.

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 Caspian Sea basin.

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 = 0.013L3.04. 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).

This species is also recorded from the Karakum Canal and Kopetdag Reservoir in Turkmenistan (Shakirova and Sukhanova, 1994; Sal'nikov, 1995) and may eventually reach Iranian waters in the Tedzhen (= Hari) River basin.

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 Mesoptamichthys
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.

© Brian W. Coad (www.briancoad.com)