ZM-CBSU Z190, 157 mm SL; Iran, Kohgiluyeh and Boyer Ahmad prov., Beshar (Bashar) River at Tale Gah village, Karun River drainage, 30°47'27"N, 51°25'13"E.

ZM-CBSU Z191, 6, 91–157 mm SL; same data as holotype. ZM-CBSU J520, 1, 107 mm SL; ZM-CBSU Z275, 12, 105–152 mm SL; Iran, Kohgiluyeh and Boyer Ahmad prov., Beshar (Bashar) River at Tale Gah village, Karun River drainage, 30°47'27"N, 51°25'13"E. 15 December 2014, G. Sayyadzadeh, R. Khaefi, A. Khajehpanah. ZM-CBSU J526, 1, 98 mm SL; ZM-CBSU J533, 1, 114 mm SL; ZM-CBSU J535, 1, 97 mm SL; ZM-CBSU J540, 1, 67 mm SL; All from Iran, Kohgiluyeh and Boyer Ahmad prov., Beshar River at Tange sorkh, Karun River drainage, 30°26'14"N, 51°45'48"E. 24 July 2011, R. Zamaneian Nejad, S. Mirgheiasi, S. Ghasemian. ZM-CBSU J444, 2, 73–90 mm SL; ZM-CBSU J447, 2, 76–111 mm SL; ZM-CBSU J450, 1, 86 mm SL; ZM-CBSU J452, 1, 107 mm SL; ZM-CBSU J459, 2, 104–120 mm SL; ZM-CBSU J464, 1, 110 mm SL; all from Iran, Kohgiluyeh and Boyer Ahmad prov., Beshar River at Mokhtar village, Karun River drainage, 30°40'31"N, 51°31'26"E. 25 May 2011, R. Zamaneian Nejad.

ZM-CBSU 7880–7881, 2, 96.69–158.12 mm SL; Iran, Fars prov., Sepidan city, Gorgu River, a tributary of Beshar River, north of Sepidan city, Karun River drainage, 30°21.283'N, 51°45.754'E. 2006. H.R. Esmaeili, A. Teimori, M. Ebrahimi and A. Gholamhoseini. SMF 33337, 1, 48.86 mm SL; Iran, Lorestan prov., Hadi River between Zagheh and Polehoru, 33°31.138'N, 48°46.340'E. 04 March 2008. N. Alwan, K. Borkenhagen, M. Ghanbari Fardi and A. Kazemi. FSJF 2213, 11, 107.92–143.94 mm SL; Iran, Chaharmahal and Bakhtiari Prov., Sandgan River (Sandgan stream) at Sandgan, 31°15.692'N, 51°17.150'E. 19 April 2007, A. Abdoli and J. Freyhof. FSJF 2233, 2, 156.22–162.23 mm SL; Iran, Kohgiluyeh and Boyer Ahmad prov., Beshar River, 20 km northeast of Yasuj, 30°44.152'N, 51°29.522'E. 19 April 2007. A. Abdoli and J. Freyhof. SMF 30865, 1, 26.94 mm SL; Iran, Kohgiluyeh and Boyer Ahmad prov., Beshar River at Tang-e Sorkh, 30°27.680'N, 51°44.907'E. 28 November 2007, K. Borkenhagen, H. R. Esmaeili and F. Wicker (in 96% alcohol). SMF 30871, 1, 28.34 mm SL; Iran, Kohgiluyeh and Boyer Ahmad prov., Beshar River at Tang-e Sorkh, 30°27.680'N, 51°44.907'E. 28 November 2007. K. Borkenhagen, H. R. Esmaeili and F. Wicker (in 96% alcohol). SMF 33316, 7, 35.22–166.87 mm SL; Iran, Kohgiluyeh and Boyer Ahmad prov., Beshar River at Tang-e Sorkh, 30°27.680'N, 51°44.907'E. 28 November 2007, K. Borkenhagen, H. R. Esmaeili and F. Wicker. SMF 30872, 1, 29.70 mm SL; Iran, Fars prov., Sepidan, Tang-e Tizab, 30°23.470'N, 51°46.710'E, 28 November 2007, K. Borkenhagen, H. R. Esmaeili and F. Wicker (in 96% alcohol).

ZM-CBSU M1275,1, Iran, Kohgiluyeh and Boyer Ahmad prov., Beshar River at Dehno village, Karun River drainage, 30°38'55"N, 51°37'05"E. 16 January 2014, H.R. Esmaeili, G. Sayyadzadeh, H.R. Mehraban, M. Razbani. GenBank accession number: (COI: KU564296); ZM-CBSU M1447, 2, GenBank accession number: (COI: KU564297, KU564298; cytb: KU564303, KU564304) ZM-CBSU M1458, 2); Iran, Kohgiluyeh and Boyer Ahmad prov., Beshar River at Tale Gah village, Karun River drainage, 30°47'27"N, 51°25'13"E. 14 December 2013. G. Sayyadzadeh, A. Khajehpanah, R. Khaefi. GenBank accession number: (COI: KU564294, KU564295; cytb: KU564305, KU564306).

Capoeta coadi sp. n. is distinguished from all other species of Capoeta by the following combination of characters: last unbranched dorsal-fin ray weakly to moderately ossified and serrated in 1/3–2/3 of its length; scales small, 70–84 total lateral line scales (84 in holotype), 12–17 scales between dorsal-fin origin and lateral line (16 in holotype), 9–11 scales between anal-fin origin and lateral line (11 in holotype), 26–32 encircling least circumference of caudal peduncle (31 in holotype); total gill rakers 14–18 (17 in holotype), 10–13 gill rakers on lower limb of first gill arch (12 in holotype); 45–47 total vertebrae; one posterior pair of barbels; length of the longest dorsal-fin ray 14.92–21.58% SL (18.90 in holotype); head length 22.87–26.33% SL (23.76 in holotype); mouth width 7.48–9.77% SL (8.65 in holotype); bright golden-greenish or silvery body coloration in life.

General body shape and appearance are shown in Figs 1–3, morphometric data in Table 1 and meristic data are summarized in Tables 2–9. Body elongate and cylindrical; predorsal body profile smoothly convex with no marked discontinuity between head and body except when a nuchal hump is present in few specimens; greatest body depth at level of dorsal-fin origin; snout rounded (in 20 specimens) or pointed (in 14 specimens) and not size dependent; mouth inferior; lips slightly fleshy, especially at the mouth corners; lower lip covered with a sharp-edged horny sheath, its anterior margin straight in adult specimens and rounded to almost crescent-shaped in juveniles, with a considerable degree of individual variation.

Dorsal-fin origin anterior to pelvic-fin origin, its outer margin usually straight to concave with 3–5 unbranched and 8–9 branched rays (3 and 8 in holotype, respectively); last unbranched dorsal-fin ray weakly to moderately ossified, flexible and soft at the tip, serrated in 1/2–2/3 of its length (Fig. 4); pectoral fins not extending to pelvic-fin base; their outer margins usually slightly convex with 16–22 rays in total (19 in holotype) (Table 2); pelvic fins not extending to anal fin base, their outer margin straight or slightly convex and blunt with 7–11 rays in total (8 in holotype) (Table 2); pelvic axillary scale present; anal fin with 3 unbranched and 5 branched rays, outer margin straight or slightly convex; caudal fin forked with 16–19 branched rays (17 in holotype) (Table 3), its tip pointed and its upper lobe often longer than lower one.

Scales small, total lateral-line scales 70–84; 12–17 scales between dorsal-fin origin and lateral line (Table 4); 9–11 scales between anal-fin origin and lateral line (Table 4); 26–32 circum-peduncle scales (Table 5); ventral midline and pectoral region covered with deeply embedded scales of reduced size; gill rakers slightly hooked, total gill rakers 14–18 (10–13 gill rakers on lower limb) of first gill arch (Table 8–9); 45–47 total vertebrae; usually one posterior pair of barbels present (very rarely two, 1 out of 51 individual); pharyngeal teeth arranged in 3 rows in the following manner: 2.3.5–5.3.2 and very similar in shape to those of Capoeta damascina; teeth in the main row spatulate or spoon-shaped and crowns flat, narrow and curved.

Coloration. Live specimens. Dorsum and sides bright golden-green or silvery, darker dorsally and lighter below the lateral line; dorsal head bright golden-green or light pink-brown; dorsal, anal and caudal fins beige to light brown with light pink to red tinge; pectoral and pelvic-fins beige to light brown or golden with brown tinge on the first few rays (Fig. 3); few large black blotches present on the body of some specimens whereas small diffuse black spots are present only on the body of some juveniles (above the lateral line).

Dorsum, head and sides grey or brownish-grey dorsally and beige or yellow ventrally; dorsal and caudal fins dusky grey; pectoral, pelvic and anal fins white or beige with or without grey tinge; blotches and spots well discernible (Figs 1–2).

Breeding tubercles present in both sexes, being bigger and more pronounced in males. Tubercles present on the sides of the snout but may also cover the entire body surface, on and above the lateral line with one or two tubercles per scale but not on each scale, below the lateral line especially in the area above the anal fin and on the branched anal-fin rays; tip of anal fin reaching to or beyond the vertical of the caudal-fin base in females and to about 2/3 of the caudal peduncle in males.

Capoeta coadi sp. n. occurs in medium-fast flowing rivers with usually gravel substrates and clear waters (Fig. 5). At the Beshar River sampling site, the river is about 25 m wide, with substrate consisting of coarse gravel and boulders, and fast-flowing and semi-transparent waters. The physicochemical parameters at the spot were: dissolved oxygen, 9.89 mg/L; total dissolved solids, 190.2 mg/L; salinity, 0.19‰; conductivity, 395 µs/cm; pH: 8.5 and water temperature 23.4 °C. It is known only from the Karun River drainage, a system that constitutes the southeastern part of the Tigris-Euphrates River system.

The new species is named after Brian W. Coad, a well-known ichthyologist for his valuable contribution to the knowledge of freshwater fishes of Iran.

The presence of one pair of barbels in Capoeta coadi sets the species apart from Capoeta antalyensis, Capoeta baliki, Capoeta banarescui, Capoeta tinca, and Capoeta heratensis, all of which have two pairs of barbels based on data from Turan et al. (2006a) and this study. The new species is further distinguished from Capoeta antalyensis by the presence of serrae on the last unbranched dorsal-fin ray (vs. absence) (Fig. 4), and by number of scales between dorsal-fin origin and lateral line (12–17 vs. 10–12 in Capoeta antalyensis) (Table 4), between anal-fin origin and lateral line (9–11 vs. 7), and by total number of the lateral-line scales (70–84 vs. 51–57) (Table 7). Capoeta coadi is distinguished from Capoeta banarescui by number of scales between anal-fin origin and lateral line (9–11 vs. 8–9) (Table 4). Data for Capoeta antalyensis and Capoeta banarescui are from Turan et al. (2006a).

Capoeta coadi is distinguished from Capoeta mandica, Capoeta erhani, and Capoeta trutta by having 10–13 gill rakers on the lower limb of the first gill arch (vs. 17–24 in Capoeta mandica, 20–22 in Capoeta erhani and 18–25 in Capoeta trutta [data from Krupp 1985, Turan et al. 2008, Table 8]). The total number of gill rakers in Capoeta coadi specimens is 14–18 that is lower than in Capoeta mandica (23–27), Capoeta barroisi (28–30), Capoeta turani (25–30) and Capoeta trutta (21–31) [data from Turan et al. (2006b), Özuluğ and Freyhof (2008), and this study] Table 9. Capoeta coadi is further distinguished from Capoeta mandica by having fewer pectoral fin rays (16–22 vs. 13–16) (Table 2). Capoeta coadi is distinguished from Capoeta bergamae, Capoeta capoeta and Capoeta sieboldii by number of scales between dorsal-fin origin and lateral line (12–17 in Capoeta coadi vs. 8–10 in Capoeta capoeta and 9–11 in Capoeta sieboldii); number of scales between anal-fin origin and lateral line (9–11 in Capoeta coadi vs. 7–9 in Capoeta bergamae, 6–10 in Capoeta capoeta and 8–10 in Capoeta sieboldii); total lateral line scales (70–84 in Capoeta coadi vs. 48–66 in Capoeta capoeta and 52–60 in Capoeta sieboldii) [data from Banarescu 1999, Turan et al. 2006b, Tables 4, 7]. In addition to the presence of serrae on the unbranched dorsal-fin ray, Capoeta coadi is set apart from Capoeta caelestis by the number of scales between the dorsal-fin origin and lateral line (12–17 in Capoeta coadi vs. 10–13.5 in Capoeta caelestis); scales between anal-fin origin and lateral line (9–11 in Capoeta coadi vs. 7–8 in Capoeta caelestis); circum-peduncular scales (26–32 in Capoeta coadi vs. 23–24 in Capoeta caelestis) (Tables 4–5) and probably vertebral counts (45–47 in Capoeta coadi vs. 44 in Capoeta caelestis) [data from Schöter et al. 2009].

It is distinguished from Capoeta damascina by having 11–13, modally 13, gill rakers on the lower limb of the first gill arch (vs. 12–18, modally 14–15) (Alwan 2011, Table 8). Capoeta coadi is clearly distinguished from Capoeta ekmekciae by number of scales between dorsal-fin origin and lateral line (12–17 in Capoeta coadi vs. 9–10 in Capoeta ekmekciae); number of scales between anal-fin origin and lateral line (9–11 in Capoeta coadi vs. 6–7 in Capoeta ekmekciae) (Table 4); number of lateral line scales (70–84 in Capoeta coadi vs. 55–61 in Capoeta ekmekciae [data from Turan et al. 2006b; Alwan 2011].

Capoeta coadi is distinguished from Capoeta kosswigi by total number of gill rakers (Table 9): 14–18 in Capoeta coadi vs. 19–28 in Capoeta kosswigi (see Karaman 1969; Turan et al. 2006b; Turan 2008).

Capoeta coadi is distinguished from Capoeta mauricii and Capoeta pestai by having a weaker, thinner and less ossified last unbranched dorsal-fin ray in juveniles and adults and fewer scales between dorsal-fin origin and lateral line (12–17 in Capoeta coadi vs. 18–22 in Capoeta mauricii and 16–19 in Capoeta pestai [data from Özuluğ and Freyhof 2008, Küçük et al. 2009]). It is further distinguished from Capoeta pestai by the absence of spots on the body except in juveniles (vs. presence of many on the body [see Özuluğ and Freyhof 2008, Küçük et al. 2009]). Capoeta coadi is distinguished from Capoeta umbla by total number of lateral line scales (70–84 vs. 86–104), number of scales between dorsal-fin origin and lateral line (12–17 vs. 18–24), number of scales between anal-fin origin and lateral line (9–11 vs. 11.5–15.5), and circum-pendicular scales (26–32 vs. 32–39) (see Alwan 2011, Tables 4–7).

Compared to other Iranian species of Capoeta, Capoeta coadi has more scales and fewer gill rakers than Capoeta aculeata (number of scales between dorsal-fin origin and lateral line: 12–17 vs. 6–10; number of scales between anal-fin origin and lateral line: 9–11 vs. 5–8; circum-peduncular scales: 26–32 vs. 13–23; total number of lateral line scales: 70–84 vs. 36–52; caudal peduncle scales: 14–18 vs. 10–12; gill rakers on the lower limb of the first gill arch: 10–13 vs. 15–18 [data from Coad and Krupp 1994] and this study (Tables 4–9)). Capoeta coadi is distinguished from Capoeta fusca by more total vertebrae (45–47 vs. 44), and more total lateral-line scales (70–84 vs. 40–62) (see Coad 2008, Johari et al. 2009).

Capoeta coadi differs from its sister species (see Figs 6–7), Capoeta buhsei in having more gill rakers on lower limb of first gill arch (10–13 vs. 8–10), more gill rakers on the whole first gill arch (14–18 vs. 12–14, see Tables 8–9) and by depth of caudal peduncle in percent of standard length (10.03–11.61 vs. 8.58–10.84). Capoeta coadi is distinguished from another closely related species, Capoeta saadii by having more scales below the lateral line (9–11 vs. 6–10, modally 9) (Table 4) and more circum-pendicular scales (26–32 vs. 23–28, modally 25–26) [data from Alwan (2011)].