Capoeta Capoeta are medium-sized fishes found throughout West Asia. All 27 species of the genus are present in the region, with the distribution range of C. heratensis extending to the Amu Darya drainage in the Aral basin. Capoeta are unmistakable, with their 1–2 pairs of short barbels, straight or rounded lower jaw, horny cutting sheath, and strong, serrated last unbranched dorsal ray. Some Capoeta have two pairs of barbels, while most have one. There is some variation in this character, with occasional individuals having one, two, or even three barbels. The molecular diversity of Capoeta was the focus of several detailed studies, resulting in the description of 12 new species since the year 2000. This makes Capoeta the second-largest genus of Cyprinids in West Asia (after Garra ). These studies also resulted in the description of several “molecular” species that are hardly distinguishable by external characters. Several of the populations that were proposed as species could not be confirmed by subsequent studies and are treated as synonyms. Capoeta frequently forms hybrids with sympatric Luciobarbus . Many species of Capoeta show remarkable variation in the shape of the mouth, especially the lower jaw, which is often wide, straight in some individuals, narrow, and arched in others. The hypothesis that these differences in mouth shape are attributed to gender was not supported when larger series were studied. The genus Paracapoeta was proposed for C. anamisensis , C. barroisi , C. erhani , C. mandica , and C. trutta . We reject this genus as valid. This species group forms a monophyletic group, very closely related to Capoeta , and morphological characters poorly distinguish it. Capoeta anamisensis does not show any of the proposed diagnostic character states for Paracapoeta . Capoeta forms an important component of many fish communities in West Asia, which are often dominated by these cyprinids by biomass. They are very abundant in many places and found in almost all kinds of water bodies, from cold mountain streams to large lowland lakes and small, brackish desert streams, as long as the fish have access to small patches of stony substrate and flowing waters. Spawning also takes place at the outflow of springs or wave-washed shores of lakes. Some Capoeta species are likely very powerful invasive species, but there have been few documented cases of translocations so far.

Capoeta Capoeta are medium-sized fishes found throughout West Asia. All 27 species of the genus are present in the region, with the distribution range of C. heratensis extending to the Amu Darya drainage in the Aral basin. Capoeta are unmistakable, with their 1–2 pairs of short barbels, straight or rounded lower jaw, horny cutting sheath, and strong, serrated last unbranched dorsal ray. Some Capoeta have two pairs of barbels, while most have one. There is some variation in this character, with occasional individuals having one, two, or even three barbels. The molecular diversity of Capoeta was the focus of several detailed studies, resulting in the description of 12 new species since the year 2000. This makes Capoeta the second-largest genus of Cyprinids in West Asia (after Garra). These studies also resulted in the description of several “molecular” species that are hardly distinguishable by external characters. Several of the populations that were proposed as species could not be confirmed by subsequent studies and are treated as synonyms. Capoeta frequently forms hybrids with sympatric Luciobarbus. Many species of Capoeta show remarkable variation in the shape of the mouth, especially the lower jaw, which is often wide, straight in some individuals, narrow, and arched in others. The hypothesis that these differences in mouth shape are attributed to gender was not supported when larger series were studied. The genus Paracapoeta was proposed for C. anamisensis, C. barroisi, C. erhani, C. mandica, and C. trutta. We reject this genus as valid. This species group forms a monophyletic group, very closely related to Capoeta, and morphological characters poorly distinguish it. Capoeta anamisensis does not show any of the proposed diagnostic character states for Paracapoeta. Capoeta forms an important component of many fish communities in West Asia, which are often dominated by these cyprinids by biomass. They are very abundant in many places and found in almost all kinds of water bodies, from cold mountain streams to large lowland lakes and small, brackish desert streams, as long as the fish have access to small patches of stony substrate and flowing waters. Spawning also takes place at the outflow of springs or wave-washed shores of lakes. Some Capoeta species are likely very powerful invasive species, but there have been few documented cases of translocations so far.

Further reading. Levin et al. 2012 (phylogeny); Ghanavi et al. 2016 (phylogeny, Iran); Geiger et al. 2014 (phylogeny, Mediterranean); Bektaş et al. 2017 (phylogeny); Turan et al. 2022a ( Paracapoeta ).

Keys to species of Capoeta

Marmara and Mediterranean basins, including endorheic basins in Central Anatolia, the Damascus, and the Dead Sea basins

1a - Two pairs of barbels. ………………2

1b - One pair of barbels. ………………3

2a - Posterior edge of last unbranched dorsal ray smooth, without serrae; 49–57 total lateral-line scales. ……………… C. antalyensis

2b - Posterior edge of last unbranched dorsal ray with serrae; 69–87 total lateral-line scales. ……………… C. tinca

3a - Edge of lower jaw not cornified; lips fleshy; snout long and pointed. ……………… C. pestai

3b - Edge of lower jaw cornified; lips narrow, not fleshy; snout short and blunt. ………………4

4a - Flank silvery or brown with many small black spots; last unbranched dorsal ray very strongly ossified, with strong serrae reaching almost to tip of ray. ………………5

4b - Flank beige or brown without small black spots; juveniles and some adults with large black blotches; last unbranched dorsal ray moderately ossified, without serrae or with small serrae, not or poorly ossified and without serrae on tip of ray. ………………6

5a - 76–83 total lateral-line scales; pelvic–hypural distance when carried forward, always falling in front of tip of snout; predorsal keel not or very slightly elevated; black spots on dorsal head smaller than on predorsal flank. ……………… C. barroisi

5b - 63–79 total lateral-line scales; pelvic–hypural distance when carried forward, falling in between midpoint of nostril and tip of snout and center of eye; predorsal keel elevated; black spots on dorsal head about equally sized than on predorsal flank. ……………… C. erhani

6a - Last unbranched dorsal ray weak, without serrae. ……………… C. caelestis

6b - Last unbranched dorsal ray ossified, with serrae. ………………7

7a - Usually 68–82 total lateral-line scales; usually 9½ branched dorsal rays. ……………… C. damascina

7b - Usually 58–71 total lateral-line scales; usually 8½ branched dorsal rays. ………………8

8a - Last unbranched dorsal ray with 21–27 small serrae; posterior margin of dorsal concave. ……………… C. bergamae

8b - Last unbranched dorsal ray with 7–20 small serrae; posterior margin of dorsal straight. ……………… C. aydinensis

Black Sea basin

1a - Two pairs of barbels. ………………2 1b - One pair of barbels. ………………4

2a - Last dorsal ray without serrae in individuals smaller than 100 mm SL, with 7–9 serrae in larger individuals. ……………… C. svanetica 2b - Last dorsal ray with 12–28 serrae. ………………3

3a - 12–14 scale rows between lateral line and dorsal origin; 12–18 gill rakers; eye diameter smaller than cheek length; shape of lower jaw monomorphic, wide and straight in all individuals. ……………… C. banarescui 3b - 14–17 scale rows between lateral line and dorsal origin; 16–22 gill rakers; eye diameter equal to or greater than cheek length; shape of lower jaw polymorphic, wide and straight or arched and narrow in some individuals. ……………… C. tinca

4a - 7–10 gill rakers; 7½ branched dorsal rays; 68–82 total lateral-line scales; 0–8 serrae along posterior margin of last simple dorsal ray. ……………… C. oguzelii 4b - 18–35 gill rakers; 8–9½ branched dorsal rays; 46–60 total lateral-line scales; many serrae along posterior margin of last simple dorsal ray. ………………5

5a - Lower and upper lips pleated, well developed and fleshy; edge of lower jaw not cornified; 28–35 gill rakers. ……………… C. sieboldii 5b - Lower and upper lips smooth and narrow; edge of lower jaw cornified; 17–26 gill rakers. ……………… C. capoeta

Caspian basin, Lake Urmia and Namak basins, Hari drainage, rivers flowing to the Persian Gulf south of Zohreh and Iranian endorheic basins

1a - Flank plain silvery or silvery with many small black spots, rarely brown with many black spots; last unbranched dorsal ray moderate to strongly ossified, with strong serrae. ………………2

1b - Flank beige, golden or brown without small black spots, juveniles and some adults with large black blotches; last unbranched dorsal ray soft or moderately ossified, with small serrae. ………………3

2a - Flank plain silvery or pale brown, often with grey stripe. ……………… C. anamisensis

2b - Flank silvery or brown with many small black spots. ……………… C. mandica

3a - Two pairs of barbels. ……………… C. heratensis

3b - One pair of barbels. ………………4

4a - 63–95 total lateral-line scales. ………………5

4b - 32–63 total lateral-line scales. ………………7

5a - A deep groove in front of nares. ……………… C. coadi

5b - No groove in front of nares. ………………6

6a - 23–28 circumpeduncular scales; 7–10 scales between lateral line and anal origin. ……………… C. saadii

6b - 29–31 circumpeduncular scales; 10–11 scales between lateral line and anal origin. ……………… C. buhsei

7a - Usually 7½ branched dorsal rays; 12–15 gill rakers. ……………… C. fusca

7b - Usually 8–9½ branched dorsal rays; 15–25 gill rakers. ……………… 8

8a - Usually 9½ branched dorsal rays. ……………… C. kaput

8b - Usually 8½ branched dorsal rays. ……………… 9

9a - 36–44 total lateral-line scales. ……………… C. aculeata

9b - 46–59 total lateral-line scales. ……………… 10

10a - 44–47 total vertebrae. ……………… C. capoeta

10b - Less than 44 total vertebrae. ……………… C. razii

Euphrates and Tigris drainages, including Lake Van basin, excluding Karkheh, Karun, and Zohreh drainages in Iran

1a - Flank silvery with many small black spots, rarely brown with many black spots; last unbranched dorsal ray very strongly ossified, as long or longer than head, rarely shorter, with strong serrae. ……………… C. trutta

1b - Flank beige, golden or brown without small black spots, juveniles and some adults with large black blotches; last unbranched dorsal ray soft or moderately ossified, shorter than head, with small serrae. ……………… C. damascina

Karkheh, Karun, and Zohreh drainages in Iran

1a - Last unbranched dorsal ray as long as or longer than head, rarely shorter; adults larger than 150 mm SL, usually silvery with many small dark-brown or black blotches. ……………… C. trutta

1b - Last unbranched dorsal ray shorter than head; adults larger 150 mm SL brown or silvery, without or with minute dark-brown or black spots. ………………2

2a - 32–44 total lateral-line scales. ……………… C. aculeata

2b - 56–104 total lateral-line scales. ………………3

3a - A deep groove in front of nares. ………………4

3b - No groove in front of nares. ………………5 4a - 12–18 gill rakers on lower limb. ……………… C. damascina

4b - 10–13 gill rakers on lower limb. ……………… C. coadi

5a - Head depth at eye 90–102 % of postorbital length. ……………… C. ferdowsii

5b - Head depth at eye 105–126 % of postorbital length. ……………… C. shajariani