Genus Capoeta
Valenciennes in Cuvier and 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 in Cuvier and 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 in Cuvier and 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, Kavir, Yazd, Esfahan, Kor River basins (Rainboth, 1981; Bianco and Banarescu, 1982; Ghorbani Chafi, 2000). Abdoli (2000) maps this species from theYazd basin generally; the upper Kal Shur, Jajarm and Jovein rivers in the 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.

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, ?, Khorasan, Jajarm (36º57'N, 56º23'E); BM(NH) 1975.1.17:255-258, 4, 103.0-161.0 mm standard length, Esfahan, Esfahan (?); 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 in Barrois, 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). 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.

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.

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 Hormozgan basin (Bianco and Banarescu (1982) and the 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 in Cuvier and 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.

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 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 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).

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 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 (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 L_t = 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 L_t = 190(1-exp{-0.462 [t+1]} for males and L_t = 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.

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).

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.

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 in Cuvier and 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 in Cuvier and 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, 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, Yazd, Lut, Hormozgan 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).

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 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.

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), Kavir, Bejestan, 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

None.

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 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