Roman.indd UDC 597.5(479.22) CAPOETA SVANETICA (TELEOSTEI, CYPRINIDAE), A NEW SPECIES FROM THE LUCHUNIS RIVER (RIONI RIVER DRAINAGE) IN GEORGIA A. Roman1,2, S. Afanasyev2, O. Golub2, O. Lietytska2 1Technical University “Metinvest Politecnica”, Sechenova st., 71 a, Mariupol, 87524 Ukraine E-mail: anatoliy.roman@mipolytech.education 2Institute of Hydrobiology of NAS of Ukraine, Prospect Geroiv Stalingradu, 12, Kyiv, 04210 Ukraine E-mail: aroman.fi sh@gmail.com E-mail: safanasyev@ukr.net E-mail: oleggolyb@gmail.com E-mail: lietitska@ukr.net S. Afanasyev (https://orcid.org/0000-0002-5247-3542) O. Lietytska (https://orcid.org/0000-0001-7026-4093) urn:lsid:zoobank.org:pub:18A322D8-BED6-44A0-AAF3-966EA98BB142 Capoeta svanetica (Teleostei, Cyprinidae), a New Species from the Luchunis River (Rioni River Drainage) in Georgia. Roman, A., Afanasyev, S., Golub, O., Lietytska, O. — Recent Georgia’s fi sh fauna includes four Capoeta species: C. capoeta (Kura River basin), C. kaput (Araxes River and its left tributar- ies, the Akhuryan and Mezamor rivers), C.  sieboldii (East and South-East Black Sea river basins), and C.  banarescuii (Chorokh River basin). Based on morphological data analysis new species of the genus Capoeta — C. svanetica sp. n. is described. It is more similar to the Capoeta damascina complex group (Anatolian-Iranian group) which also includes C.  banarescui, C.  baliki, and C.  sieboldii spread at East and South-East Black Sea coastal rivers. Th e Anatolian-Iranian group, also known as small-scales spe- cies group well distinguished from Aral-Caspian species (C. capoeta and C. kaput in Georgia) by highest scales number in lateral line. C. svanetica sp. n. is distinguished from other Capoeta species of Black Sea basin rivers in Georgia and adjacent waters by combination of characters: two pairs of barbels are present (C. sieboldii and C. oguzelii have one barbels pair); no spots on the body (C. oguzelii has small black spots); lower lip has keratinised edge without fringe (C. sieboldii and C. oguzelii are characterized by fringed lips); scales small, 70–74 total lateral line scales (in C. sieboldii only 52–60 scales); 10–12 scale rows above lat- eral line (C. banarescui, C. baliki and C. oguzelii have more than 12 scale rows) and 7–8 scale rows below lateral line (C. baliki and C. oguzelii have more than 10 scale rows); 12–15 gill rakers on the fi rst gill arch (C.  sieboldii and C.  baliki have more then 16); last unbranched fi n ray soft , serrae number on it is 7–9 in adult and 0 in juvenile samples (all other Capoeta species of Black Sea basin rivers have well ossifi ed last unbranched ray with high serrae number); anal fi n base length is only 7.4–9.4 % of SL (more shorter than C. banarescui, C. baliki and also C. tinca); as well as mouth width (25.4–29.4 % of HL); eye diameter (19.0–28.7 % of HL) is biggest than same in C. banarescui, C. baliki and also C. tinca. K e y w o r d s : new species, Capoeta, Svaneti, Capoeta damascina complex group, West Georgia. Zoodiversity, 56(2):117–134, 2022 DOI 10.15407/zoo2022.02.117 118 A. Roman, S. Afanasyev, O. Golub, O. Lietytska Introduction According to Ninua and Japoshvili (2008) in Georgia’s water fl ows were known four taxons of the ge- nus Capoeta: Capoeta capoeta (Güldenstädt, 1773) and it subspecies Capoeta capoeta sevangi de Filippi, 1865 (Caspian Sea basin rivers); Capoeta tinca (Heckel, 1843) and Capoeta sieboldii (Steindachner, 1864) — both from Black Sea basin rivers. Later (Baycelebi et al., 2015) for Chorokh [Çoruh] River fi sh fauna were listed also Capoeta ekmekciae Turan, Kottelat, Kirankaya & Engin, 2006 (Turan et al., 2006 a) and C. banarescui Turan, Kottelat, Ekmekci, Imamoglu, 2006 (Turan et al., 2006 b), instead of C. tinca which was noted only for the Sea of Marmara basin (Turan et al., 2006 b). Subspecies Capoeta capoeta sevangi based on DNA analysis data was con- sidered as a distinct species Capoeta sevangi de Filippi, 1865 (Zareian et al., 2016; 2018). Th us, based on main re- visions of fi sh fauna of Georgia should include fi ve species of the genus Capoeta: C. capoeta (Kura River basin), C. sevangi (Sevan Lake), C. sieboldii (all Georgian Black Sea basin Rivers from Rioni to Chorokh), C. banarescuii and С. ecmekcii (in Georgia Chorokh River basin only). C. banarescuii according to D. Turan’s data (Turan et al., 2006 b) was known from the Chorokh River drainage from the Kachkar Mountains in Turkey, but based on the Chorokh River lowermost course fl owing in Georgia which fl ows to the Black Sea at Batumi it was noted as fi sh fauna of Georgia potential component. As well as С. ecmekcii also known for the Chorokh River drainage but only for Turkey. Recently (Kuljanishvili et al., 2020) Caucasus freshwater fi shes were revised and it was noted four Capoeta species for freshwater fauna of Georgia. Th ey are C. capoeta (Kura River basin), Capoeta kaput Levin, Prokofi ev et Roubenyan, 2019 (Araxes River and its left tributaries, the Akhuryan and Mezamor rivers.), C. sieboldii (West Georgia’s river basins), and C. banarescuii (Choroh River basin). At one of the worst investigated mountain regions of Georgia–Kvemo–Svaneti on the Luchunis River (Rioni River tributary) at Uravi were found samples of Capoeta genus which could not be noted as any one of four known species. Aft er detailed morphological data analysis showed these samples as distinct ones. Material and methods Fish specimens were sampled at the high mountain region of Kvemo–Svaneti during the spring-fall pe- riod (fi g. 1). In total, 7 specimens were collected using hand-nets. All of them were caught in the Luchunis River (Rioni River drainage) at Uravi (Kvemo–Svaneti, Georgia). All used samples are deposited in the National Museum of Natural History of NAS of Ukraine (NMNH NASU). Sampled fi sh specimens were fi xed in 4 % formaldehyde and stored in 70 % ethanol. Measurements were recorded using a dial caliper (with 0.1  mm accuracy). All measurements were made point-to-point on fi xed fi sh samples. Th e methods used for counts and measurements followed those of Turan et al. (2006 b), for best comparison results, and previously it was taken from Kottelat & Freyhof (2007). Th e standard length (SL) of the fi sh was measured from the tip of the snout to the end of the hypural complex. Th e last two-branched rays ar- ticulating on a single pterygiophore in the dorsal and anal fi ns were noted as ‘1½’ (Kottelat, Freyhof, 2007). Also used next abbreviation: Du — number of unbranched (simple) rays in dorsal fi n; Db — number of branched rays in dorsal fi n; P — number of branched (soft ) rays in pectoral fi n; V — number of branched rays in ventral fi n; l.l. — scales number in lateral line; sc. ab. l.l. — scales number above lateral line (between l.l. and dorsal-fi n origin); sc. bl. l.l. — scales number below the lateral line (between l.l. and ventral fi n origin). Fig. 1. Genus Capoeta distribution in river basins of Georgia and some neighbors. 119Capoeta svanetica (Teleostei, Cyprinidae), a New Species from the Luchunis River… Morphological data of C. baliki, C. banarescui, C. capoeta, C. ekmekciae, C. sieboldii, C. sevangi, C. kaput, C.  tinca and Capoeta oguzelii Elp, Osmanoğlua, Kadak and Turan, 2018 were taken from Berg, 1912–1914; 1949; Turan et al., 2006 a; 2006 b; Levin et al., 2012; 2019; Elp et al., 2018. Also for studying morphological data were used additional materials. C. capoeta: Georgia. Borjomi Municipality, Samtskhe–Javakheti Region: Kura River at Akhaldaba [41.9104 N, 43.4848 E]; 25.09.1966; 1 individual (V. Pinchuk). C.  sevangi: Azerbaijan. Fuzuli District: Araxes River [39.3484 N, 47.2525 E]; 06.06.1970; 6 individuals (M. Golovushkin); Sharur District: Arpa River at Diza, Araxes River Basin [39.5987 N, 45.0709 E]; 27.08.1975; 599 individuals (M. Golubiev). C. banarescui: Georgia. Autonomous Republic of Imeretia, Kutaisi Region: Gubistskali River, Rioni River basin [42.3043 N, 42.5020 E]; 16.04.2016; 1 individual (A.  Roman); Autonomous Republic of Adjara, Khel- vachauri district: Chorokh River at Khelvachauri [41.5663 N, 41.6785 E]; 22.04.2016; 3 individuals (A. Roman). Results Capoeta svanetica Roman, Afanasyev, Golub et Lietytska sp. n. urn:lsid:zoobank.org:act:CCEB77F1-25D8-4770-AED2-BA375E2EE11D Capoeta tinca (non Heckel, 1843): Berg, 1912–1914: 0168 (Olty-tschai, Chorokh drain- age and Rion drainage). Varicorhinus tinca (non Heckel, 1843): Berg, 1912–1914: 554 (Olty-tschai, Chorokh and Rion drainages), 1948–1949: 684 (in part; Chorokh and Rion drainages, Georgia). M a t e r i a l . Type. Holotype: NMNH NASU 10420, {: 136  mm SL; Georgia: Kvemo–Svaneti Region: Luchunis River at Uravi, Rioni River Basin, [42.6123 N, 43.2623 E]; 12.08.2016 (S.  Afanasyev, O.  Golub). Paratypes. NMNH NASU 10421, 2 individuals, 44–118 mm SL; same data as holotype, 12.08.2016 (S. Afanasyev, O. Golub). Non-Type. Georgia: Kvemo–Svaneti Region: Luchunis River at Uravi, Rioni River basin [42.6123 N, 43.2623 E]; 14.08.2016; 5 individuals (S. Afanasyev, O. Golub). D i a g n o s i s . C.  svanetica  sp.  n. is distinguished from the other genus Capoeta spe- cies by combination of the following characters: two pairs of barbels are present; snout rounded without any spots; mouth is narrow and slightly arched; lower lip slightly arched and has keratinised edge without fringe; scales small, 70–74 total lateral line scales (73 in holotype); 10–12 scales rows above lateral line (12 in holotype) and 7–8 scales rows below lateral line (8 in holotype); 12–15 gill rakers on the fi rst gill arch (14 in holotype); serrae number in the last unbranched fi n ray is 7–9; lateral head length (HL) is 22.1–27.7 % of SL (22.1 % in holotype); anal fi n base length is 7.4–9.4 % of SL (7.4 % in holotype); eye diam- eter is 19.0–28.7 % of HL (19.0 % in holotype); snout depth at nostrils is 34.2–38.6 % of HL (37.7 % in holotype); length of anterior barbel is 13.9–20.1 % of HL (19.3 % in holotype); length of posterior barbel is 18.7–28.6 % of HL (22.7 % in holotype); mouth width is 25.4– 29.4 % of HL (29.0 % in holotype). D e s c r i p t i o n . General body appearance presents in fi gure 2; morphometric and mer- istic data are present in tables 2–5. Th e body is elongated and cylindrical with a slightly convex upper profi le and less convex ventral one. Th e head is relatively short; the upper profi le is slightly convex in the interorbital and is slightly concave at the level of the nostrils. Th e mouth is inferior, narrow, and slightly arched (fi g. 3). Lips are slightly fl eshy. Th e lower lip is slightly arched and covered with a sharp-edged horny sheath in both sexes. Anterior barbel reaches to the nostrils (it is 13.9–20.1 % of HL), and the posterior barbel is longer and reaches the center of the eye (it is 18.7–28.6 of HL). Dorsal fi n with 3 or 4 simple (unbranched) and 7 branched rays (4 and 7 in holo- type, respectively), the outer margin is slightly concave, origin slightly in front of verti- cal through the pelvic-fi n origin, last simple ray only slightly ossifi ed, proximal two thirds rigid, and without serrae on posterior margin in small fi sh (less than near 100 mm standard length) or with 7–9 small serrae in adult (fi g. 4). Pectoral fi ns do not extend to the pelvic-fi n base; their outer margins are usually slight- ly convex with 16–22 branched rays (table 1) in total (18 in holotype). Pelvic fi ns (table 1) 120 A. Roman, S. Afanasyev, O. Golub, O. Lietytska Fig. 2. Th e general body appearance of C. svanetica sp. n. (male — top, SL = 136 mm and female — bottom, SL = 118 mm). T a b l e 1 . Meristic (counted) features of Capoeta species from Black, Caspian and Marmara Seas basins С ou nt ed fe at ur es Black Sea basin Caspian Sea basin Marmara Sea basin C . s va ne tic a sp .n . С . b an ar es - cu i C . s ie bo ld ii C . b al ik i C . o gu ze lii C . e km ek ci ae C . c ap oe ta C . s ev an gi C . k ap ut C . t in ca n = 8 n = 25 n = 23 n = 35 n = 21 n = 24 n = 31 n = 6 n = 17 n = 25 barbels II II I II I I I I I II gill rakers 12–15 (13.3) 12–16 (14.7) 28–33 (30.1) 16–22 (19.3) 7–10 (8.3) 18–24 (60.4) 18–25 (21.3)* 21–24 (22.5) 24–25 (24.5) 19–23 (20.6) serrae 7–9 (7.4) 12-20 no data 17–23 (19.5) 0–12 no data 20–41 (26.0) 15–20 (17.0) 24–35 (30.7) 24–28 (26.6) Du IV III-IV III–IV III–IV IV III–IV IV III-IV IV III Db 7–8 (7.1) 7–9 (8.0) 8–9 (8.0) 8–9 (8.1) 7 (71/2) 8–9 (8.1) 7–9 (7.9) 7–8 (7.8) 8–9 (8.9) 8 (8.0) P 16–18 (17.3) 17–19 (17.8) 15–16 17–20 (18.4) 16–17 (16.6) 16–20 18 18–21 (19.0) no data 18–20 (18.8) V 8–9 (8.6) 9–10 (9.1) 8–9 9–10 (9.0) 8–9 (8.1) 12 10 10–11 (10.2) no data 8–9 (9.0) l.l. 70–74 (72.0) 64–77 (70.8) 52–60 (55.8) 72–86 (78.4) 72–82 (77.0) 55–61 (57.5) 47–59 (53.6)* 50–56 (53.0) 52–60 (55.9) 69–80 (74.9) sc. ab. l.l. 10–12 (11.1) 12–14 (12.8) 9–11 (9.6) 14–17 (14.9) 12–17 (13.8) 9–10 (9.4) 8–10 (9.0)* 8–9 (8.2) 9–12 (10.6) 14–17 (15.7) sc. bl. l.l. 7–8 (7.9) 8–9 (8.1) 8–10 (8.9) 10–11 (10.1) 11–13 (11.6) 6–7 (6.8) 6–8 (7.0)* 16–8 (6.5) 7–8 (7.4) 9–11 (9.5) * total used samples are diff erent, see tables 2–4. N o t e . Data on C. banarescui, C. baliki and C. tinca noted aft er Turan et al., 2006 b; data on C. ekmekciae were taken from Turan et al., 2006 a; data on C. oguzelii and C. sieboldii were taken from Elp et al. (2018); data on C. capoeta and C. kaput were taken from Levin et al., 2019; data on C. svanetica sp.n. and C. sevangi are ours. 121Capoeta svanetica (Teleostei, Cyprinidae), a New Species from the Luchunis River… with 1 simple and 8–9 branched rays (9 in holotype), it is not extending to anal-fi n base, their outer margin straight or slightly convex; pelvic axillary scales present. Anal fi n with 3 simple and 51/2 branched rays, the outer margin is convex. Th e caudal fi n is long and deeply forked, its upper lobe oft en longer than the lower one. Scales small, total lateral-line scales 70–74 (table 2), 10–12 between dorsal-fi n origin and lateral line, and 7–8 between anal-fi n origin and lateral line (table 1). Ventral mid-line and pectoral region covered with deeply embedded scales of reduced size. Gill rakers number 12 (1), 13 (3), 14 (1), and 15 (1) on the outer side of the fi rst-gill arch (table. 4). Pharyngeal teeth arranged in 3 rows as 4.3.2–2.3.4. C o l o r a t i o n . In formaldehyde-fi xed samples, the total body color is greyish on the back and upper part of the fl ank and light grey on the lower fl ank. Th ere are no spots on the body and head. Dorsal and caudal fi ns are grey; pectoral, anal, and pelvic fi ns are yellowish (fi g. 2). S e x u a l d i m o r p h i s m . All samples were collected at the second half summer period when spawning was fi nished. Th us, we can speak of sexual dimorphism as independent of the spawning period. But breeding tubercles present in anal-fi n rays in males are more typi- cal for the spawning period. Th us, this question needs to be investigated in detail. We can Fig. 3. Mouth shape male (left ) SL = 136 mm and female (right) SL = 118 mm. Fig. 4. Last simple (unbranched) dorsal fi n rays: a — C. tinca, 139 mm SL, female (aft er Turan et al., 2006 b); b — C. banarescui, 144 mm SL, female (aft er Turan et al., 2006 b); c — C. baliki, 148 mm SL, female (aft er Turan et al., 2006 b) and d —C. svanetica sp. n. 118 mm SL, female — only 7 serrae are present. 122 A. Roman, S. Afanasyev, O. Golub, O. Lietytska T ab le 2 . L at er al li ne sc al es n um be r va ri at io n in C ap oe ta sp ec ie s ( co lu m ns w it ho ut d at a ar e ex cl ud ed fr om ta bl e) Sp ec ie s N um be r o f l at er al li ne sc al es N 47 49 50 51 52 53 54 55 56 57 58 59 60 61 64 65 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 84 85 86 m ea n C . s va ne tic a sp . n . 7 2 1 1 2 1 72 .0 C . b an ar es cu i 25 1 2 1 2 2 4 2 4 3 1 1 70 .8 C . s ie bo ld ii 23 2 2 3 2 5 4 3 1 1 55 .8 C . b al ik i 35 1 2 4 2 5 7 4 3 1 1 3 1 1 78 .4 C . o gu ze lii 21 2 3 2 2 3 1 4 2 1 1 77 .0 C . e km ek ci ae 24 6 4 1 3 6 2 2 57 .5 C . c ap oe ta 61 1 3 2 2 11 9 10 12 4 5 1 1 53 .6 C . s ev an gi 6 1 2 1 1 1 53 .0 C . k ap ut 17 2 2 1 1 3 5 1 2 55 .9 C . t in ca 25 1 2 4 2 3 2 4 2 2 1 74 .9 T ab le 3 . B ra nc he d do rs al fi n ra ys a nd sc al es r ow s n um be r ab ov e an d be lo w la te ra l l in e va ri at io n in C ap oe ta sp ec ie s Sp ec ie s Br an ch ed D or sa l F in R ay s A bo ve L at er al L in e Be lo w L at er al L in e n 6 7 8 9 m ea n n 8 9 10 11 12 13 14 15 16 17 18 m ea n n 6 7 8 9 10 11 12 13 m ea n C . s va ne tic a sp . n . 7 6 1 7. 1 7 1 4 2 11 .1 7 1 6 7. 9 C . b an ar es cu i 25 25 8. 0 25 6 17 2 12 .8 25 23 2 8. 1 C . s ie bo ld ii 23 22 1 8. 0 23 11 10 2 9. 6 23 4 17 2 8. 9 C . b al ik i 35 33 2 8. 1 35 18 17 5 1 14 .9 35 23 2 10 .1 C . o gu ze lii 20 20 71 / 2 13 3 3 4 1 1 1 13 .8 13 6 6 1 11 .6 C . e km ek ci ae 23 21 2 8. 1 24 13 11 9. 4 24 5 19 6. 8 C . c ap oe ta 31 5 25 1 7. 9 30 6 18 * 6 9. 0 29 5* 20 * 4* 7. 0 C . s ev an gi 6 1 5 7. 8 6 5 1 8. 2 6 4 1 1 6. 5 C . k ap ut 10 1 9 8. 9 7 1 3 1 2 10 .6 9 5 4 7. 4 C . t in ca 25 25 8. 0 25 1 9 10 5 15 .7 25 16 6 3 9. 5 * L ev in e t a l., 2 01 9 ar e no te d al so 8 .5 a nd 9 .5 sc al es n um be rs a bo ve a nd 5 .5 ; 6 .5 a nd 7 .5 sc al es n um be rs g ra de s b el ow la te ra l l in e. W e ad de d th is n um be rs to n ex t f ul l n um be r fo r b es t c om pa ri so n. 123Capoeta svanetica (Teleostei, Cyprinidae), a New Species from the Luchunis River… T ab le 4 . G ill r ak er s o n ou te r si de o f fi r st g ill a rc h va ri at io n in C ap oe ta sp ec ie s Sp ec ie s N um be r o f g ill ra ke rs o n ou te r s id e of fi rs t g ill a rc h n 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 m ea n C . s va ne tic a sp . n . 6 1 3 1 1 13 .3 C . b an ar es cu i 25 2 2 4 11 6 14 .7 C . s ie bo ld ii 23 5 5 4 2 5 2 30 .1 C . b al ik i 35 2 5 4 6 8 9 1 19 .3 C . o gu ze lii 15 4 3 7 1 8. 3 C . e km ek ci ae 24 6 4 2 5 2 4 1 20 .4 C . c ap oe ta 28 2 3 6 6 5 3 3 21 .3 C . k ap ut 2 1 1 24 .5 C . t in ca 25 6 8 4 5 2 20 .6 only conclude about the presence of epithelium tu- bercles in adult males in the second summer half and their absence in mature females at the same period. Small epithelial tubercles (fi g.  5) are present on the head, anal fi n rays, and fl ank scales (one per apical edge of scales) in males during the second half sum- mer season at least. In females, epithelium tubercles are absent during the second half summer season. Any diff erences based on meristic (tables  1–4) or morphometric (table.  5) data were not found as like as no clear diff erences in the lower lip shape (fi g. 3) H a b i t a t a n d b i o l o g y . C. svanetica sp. n. is too rare species that was collected in the Luchunis River (Rioni River basin) at Kvemo–Svaneti Region at Uravi village (729 masl) in the middle and lower parts of the river. Th e species occurs in medium-fast fl owing rivers with usually gravel substrates and clear waters. Sam- pling sites of the Luchunis River should be divided into two types (fi g. 6), but both are characterized by a sub- strate consisting of coarse gravel and boulders, and fast- fl owing and translucent waters (Afanasiev et al. 2022). Th e fi rst riverbed type is classifi ed as a braided channel (fi g. 6, a and 6, c). Two islands divide the main fl ow into three branches. Th e fi rst island parameters are 80 m length and in average 9 m width (23 m max.); the second island had 25 m length and in average 5 m width (12  m max.). Th e riverbed cross-sections were made for each branch separately (fi g. 6, c). Valley is U- shape; banks were covered by trees, bushes, and grass. Th e average velocity is 1.03  m/s, with the maximum at 1.67 m/s. Flow types include chaotic, broken stand- ing waves, and unbroken standing waves. Th e right branch average width is 9  m (it is varied between 2 and 7.5 m); the left branch average width is 6 m (be- tween 3.8 and 9 m). Th e average depth is 0.24 m with the maximum at 0.45 m. Both river banks were mainly made by pebble and the riverbed was covered by cob- ble (52 %), pebble (24 %), and 9 % of boulders. Th e second riverbed type (fi g. 6, b) is classifi ed as a single channel type. Th e average velocity is 0.99 m/s, with the maximum at 1.39  m/s. Flow types include turbulent, broken, and unbroken standing waves and ripples. Th e average width is 8 m (it is varied between 6,5  m and 10  m). Bed elements include bars, rapids, riffl es, rocks, and step/pools. Th e average depth is 0,49 m with the maximum at 1.7 m. Valley is U-shape evenly covered by cobble riverbed. Among substrate types also prevails cobble and pebble. E t y m o l o g y . Th e name of the species is derived from the name Svaneti, the historical region in Geor- gia where this species occurred. 124 A. Roman, S. Afanasyev, O. Golub, O. Lietytska T a b l e 5 . Morphometric features of most similar to Capoeta svanetica species (Capoeta svanetica include holotype and paratypes Morphometric features C. svanetica sp.n.n = 7 C. banaresqui n = 26 C. tinca n = 24 C. baliki n = 25 Percentage of standard length Lateral head length 22.1–27.7 (24.6) 22.2–25.8 (24.4) 23.3–26.7 (24.9) 21.9–24.8 (23.5) Body depth at dorsal fi n origin 22.0–24.8 (23.2) 21.4–25.1 (23.0) 24.4–28.0 (26.0) 21.2–24.9 (23.2) Predorsal length 48.8–54.1 (51.4) 48.7–54.5 (50.6) 48.0–53.4 (51.0) 48.6–55.5 (51.3) Postdorsal length 37.5–42.1 (38.4) 34.9–40.1 (37.2) 34.8–39.3 (37.3) 34.4–38.8 (36.6) Prepelvic length 51.7–56.7 (54.1) 50.5–57.3 (54.7) 52.4–57.8 (54.5) 51.8–56.7 (54.1) Preanal length 72.5–79.5 (76.4) 73.3–79.9 (76.3) 74.9–79.8 (77.0) 74.7–79.3 (77.0) Pectoral-fi n origin to anal fi n 50.8–58.8 (54.2) 50.3–58.2 (54.7) 52.6–57.6 (55.1) 52.3–58.4 (55.3) Pectoral-fi n origin to pelvic fi n 28.6–34.6 (32.0) 29.1–35.3 (33.0) 30.1–34.9 (32.0) 28.9–34.0 (32.0) Pelvic-fi n origin to anal fi n 20.9–24.5 (22.9) 18.5–23.6 (21.5) 20.9–26.2 (23.5) 22.1–24.4 (23.3) Dorsal fi n length 12.6–18.5 (15.0) Dorsal fi n height 13.9–19.1 (16.3) 15.9–20.7 (18.6) 17.4–21.9 (19.2) 15.6–21.1 (18.3) Anal fi n base length 7.0–10.0 (8.7) 15.9–21.8 (18.7) 17.0–21.7 (18.8) 15.1–21.3 (17.0) Anal fi n height 17.4–20.2 (18.8) Pectoral fi n length 19.6–21.8 (20.4) 16.2–21.6 (19.4) 17.3–21.3 (18.9) 16.5–19.5 (18.3) Pelvic fi n length 15.0–19.0 (17.4) 14.0–17.0 (15.9) 15.2–17.7 (16.5) 14.4–16.8 (15.8) Upper caudal-fi n lobe 22.1–26.4 (23.7) 20.3–25.5 (22.9) 22.8–26.6 (24.7) 19.3–24.6 (22.3) Lower caudal-fi n lobe 22.5–26.4 (24.8) Length of middle caudal fi n rays 8.4–13.2 (10.1) 10.6–15.1 (13.3) 13.1–15.7 (14.2) 11.6–14.7 (13.4) Length of caudal peduncle 13.0–18.5 (16.5) 15.5–19.9 (17.4) 15.4–18.5 (16.8) 16.0–20.2 (17.6) Depth of caudal peduncle 9.8–16.2 (11.5) 9.8–11.7 (10.7) 10.8–13.4 (11.7) 9.7–12.3 (10.9) In percent of head length Snout length 29.5–38.9 (34.7) 35.4–41.2 (38.3) 33.1–40.4 (37.3) 33.7–40.6 (37.3) Eye diameter 19.0–28.7 (22.3) 13.2–18.9 (15.1) 14.2–18.6 (16.3) 13.1–18.8 (15.9) Postorbital distance 46.6–50.0 (48.8) Interorbital width 30.3–37.0 (33.5) 35.9–42.1 (38.3) 33.9–42.5 (38.7) 36.3–43.2 (39.5) Head width at anterior margin of eyes 35.2–45.0 (40.7) 40.9–46.8 (43.8) 39.8–46.5 (43.0) 42.4–49.6 (45.4) Head depth at interorbital region 50.6–54.9 (53.0) 45.7–53.3 (49.0) 45.4–54.4 (49.4) 49.1–59.2 (53.6) Snout width at nostrils 34.9–45.9 (37.3) 36.4–45.0 (40.4) 35.9–41.1 (38.3) 35.5–47.6 (40.3) Snout depth at nostrils 34.2–38.6 (36.3) 29.7–35.1 (32.7) 30.0–41.1 (34.2) 33.1–41.6 (37.3) Length of anterior barbel 13.9–20.1 (16.7) 12.4–20.8 (16.9) 8.1–14.1 (10.6) 9.8–14.3 (12.7) Length of posterior barbel 18.7–28.6 (22.2) 18.4–28.8 (21.9) 13.1–19.3 (15.4) 14.7–18.5 (16.5) Mouth width 25.4–29.4 (28.3) 29.5–37.9 (34.3) 27.4–34.2 (30.8) 29.5–38.5 (33.3) N o t e . Data on C. tinca, C. baliki and C. banaresqui aft er Turan et al., 2006 b. Fig.  5. Epithelial tubercles on each scale and anal fi n rays of C.  svanetica  sp.  n. males during the second half summer season. 125Capoeta svanetica (Teleostei, Cyprinidae), a New Species from the Luchunis River… Comparison with closely related species Using morphological data analysis (tables 1–4) were compared C. svanetica sp. n. sam- ples with such from Black Sea basin rivers: C. banarescui, C. sieboldii, and C. ekmekciae as the similar distributed species; C.  baliki (widest spread species at South Black Sea rivers basins), C.  oguzelii from adjacent water fl ows. From the Sea of Marmara basin: C.  tinca as previously known species distributed in the Rioni River drainage; and also from Cas- pian Sea basin: C. capoeta, C. sevangi, and C. kaput. For more detailed analysis also used body measurement analysis (table 5) most closely related to C.  svanetica  sp.  n. species: C. banarescui, C. baliki, and C. tinca. We did not use coloration in detail for comparison C.  svanetica  sp.  n. with other Capoeta species because all our samples were fi xed in formaldehyde during the year and they practically lost their coloration. We can state only complete any spots absence on the fi sh body, head, and fi ns in C. svanetica sp. n. (fi g. 2) the same as closely related C. banares- cui (fi g. 7, a) and C. baliki (fi g. 7, d, Turan, et al., 2006 b) from Black Sea rivers basins and previously known for the same areal C. tinca (fi g. 8, a, Turan et al., 2006 b) Th is live and formaldehyde preserved specimens are characterized by dark brown on back and fl anks, yellowish-white on the belly. As a result of C. svanetica sp. n. comparison with the Capoeta species from rivers of the Black Sea basin were found new species samples is distinguishing from C.  sieboldii, C. oguzelii and C. ekmekciae by two barbels pairs presence (table 1). C. baliki and C. ban- aerscui are also characterized by two pairs of barbels, but the fi rst of them (C. baliki) has Fig. 6. Type habitats of juvenile Capoeta svanetica sp. n. (a) and adult (b) with detailed river bed structure (c). 126 A. Roman, S. Afanasyev, O. Golub, O. Lietytska shorter anterior (mean 12.7; range 9.8–14.3) and posterior (mean 16.5; range 14.7–18.5) barbels than C. svanetica sp. n. (mean 16.7; range 13.9–16.7 and mean 22.2; range 18.7–28.6 respectively). Another species (C.  banaerscui) has approximately the same pairs of bar- bels length. But the last one clearly distinguishes from C. svanetica sp. n. by more highest serrae number in the last unbranched fi n ray (fi g. 4; 12–20 in C. banaerscui compared to C. svanetica sp. n. without serrae in the last unbranched fi n ray in small fi sh (less than near 100  mm standard length) or with 7–9 small serrae in adult). Also, C.  banaerscui has the highest scale rows number upper lateral line (12–14 (mean 12.8) compared to 10–12 (mean 11.1) in C. svanetica sp. n., table. 1 and table 3). All Black Sea basin rivers Capoeta species, excluding C. oguzelii, are also distinguished from C. svanetica sp. n. by the fewer number of branched dorsal fi n rays (7–8 (mean 7.1) vs. 8–9 in other species); from C. sieboldii and C. oguzelii C. svanetica sp. n. also diff ers by the fewer number of pectoral fi n rays (15–16 and 16 (common) — 17 (mean 16.6) respectively vs. 16–18 (mean 17.3)); and from C. ek- mekciae the last one diff ers by the fewer number of pelvic fi n rays — 12 in C.  ekmekciae and 8–9 (mean 8.6) in C. svanetica sp. n. C. oguzelii is also the species which characterized by the keratinized edge on the lower lip absence (Elp et al., 2018). And C. sieboldii is also characterized by fringed lips. Fig. 7. a–c: C. banarescui, holotype, ESFM-PISI/2004-072, 177 mm SL; Turkey, Chorokh drainage at Torum (a); female lower lip, 192 mm SL (b) and male lower lip, 178 mm SL (c), aft er Turan et al, 2006 b; d–f: C. baliki, ho- lotype, ESFM-PISI/2004-74, 202 mm SL; Turkey, Sakarya drainage at Kızılcahamam; female lower lip, 176 mm SL (e) and male lower lip, 164 mm SL (f), aft er Turan et al, 2006 b. Fig. 8. a–c: C. tinca, general body appearance, FFR 718, 129 mm SL; Turkey: Koca River (a); female lower lip, 129 mm SL (b) and male lower lip, 136 mm SL (c), aft er Turan et al, 2006 b; d–f: C. oguzelii, holotype, FCME 2017-05a, 109 mm SL; Ezine Stream (d); lower lip (e); dorsal fi n with soft last unbranched fi n ray (f), aft er Elp et al., 2018. 127Capoeta svanetica (Teleostei, Cyprinidae), a New Species from the Luchunis River… Both, C. sieboldii and C. ekmekciae are characterized by the fewer number of scales in lateral line (52–60 (mean 55.8) and 57–60 (mean ) respectively vs. 70–74 (mean 72.0)) than C. svanetica sp. n.; C. sieboldii and C. ekmekciae have also fewer scales rows number above lateral line (9–11 (mean 9.6) and 9–10 (mean 9.4) respectively) and C.  sieboldii have fewer number of scales below (8–10 (mean 8.9)) lateral line (table  2) than C. svanetica sp. n. Th e last one is characterized by the fewer number of scales rows above and below the lateral line (table 3) than C. oguzelii (12–17 (mean 13.8) above and 11–13 (mean 11.6) below l.l.) and C. baliki (14–17 (mean 14.9) above and 10–11 (10.1) below l.l.). Based on the outer side of the fi rst gill arch rakers number we should conclude C. sva- netica  sp.  n. (12–15 with mean 13.3) is more similar to C.  banarescui (12–16 with mean 14.7). C. oguzelii diff erenced from C. svanetica sp. n. by fewer (7–10, mean 8.3) gill rakers number. C.  baliki, C.  ekmekciae and C.  sieboldii diff erenced from C.  svanetica  sp.  n. by highest gill rakers number (16–22, mean 19.3; 18–24, mean 20.4 and 28–33, mean 30.1 respectively). Capoeta species from the Caspian Sea Basin (East Georgia) also clearly distinguished from C.  svanetica  sp.  n. by only one pair of barbels (table  1) and fewer scales number in lateral line (47–59 (mean 53.6) in C. capoeta, 50–56 (mean 53.0) in C. sevangi and 52–60 (mean 55.9) in C. kaput); above l.l. (8–10 (mean 9.0) in C. capoeta and 8–9 (mean 8.2) in C. sevangi) and below it (6–8 (mean 7.0) in C. capoeta and 6–8 (mean 6.5) in C. sevangi). C.  svanetica  sp.  n. as well as C.  capoeta and C.  sevangi characterized by fewer branched rays in dorsal fi n number than C. kaput. Th e last one has 8–9 (8.9) — only one sample had 8 rays, all others have 9. All studied Capoeta species from Caspian Sea Basin (C. capoeta, C. sevangi and C. kaput) are well distinguished from C. svanetica sp. n. by well ossifi ed last unbranched fi n ray with high serrae number and also by highest gill rakers number on the fi rst gill arch. For morphometric features comparing were used only the most similar species C.  banarescui and C.  baliki distributed in Black Sea basin rivers and also C.  tinca as previously known species for Rioni River drainage (table 5). Thus, C. svanetica sp. n. is distinguished from C. banarescui, C. baliki and C. tinca by next morphometric fea- tures: anal fin base length, eye diameter, snout depth at nostrils, length of anterior and posterior barbel and mouth width. C. svanetica sp. n. is characterized by shorter anal fin base length (7.0–10.0 (mean 8.7)) and less mouth width 25.4–29.4 (mean 28.3) than three other compared species (15.9–21.8 (mean 18.7) and 29.5–37.9 (mean 34.3) in C. banarescui, 17.0–21.7 (mean 18.8) and 27.4–34.2 (mean 30.8) in C. baliki, 15.1–21.3 (mean 17.0) and 29.5–38.5 (mean 33.3) in C. tinca respectively (table 5). Also C. sva- netica sp. n. has the biggest eye diameter (19.0–28.7 (mean 22.3)) and snout depth at nostrils (34.2–38.6 (mean 36.3)) than three other species (13.2–18.9 (mean 15.1) and 29.7–35.1 (mean 32.7) in C.  banarescui; 14.2–18.6 (mean 16.3) and 30.0–41.1 (mean 34.2) in C. baliki; 13.1–18.8 (mean 15.9) and 33.1–41.6 (mean 37.3) in C. tinca respec- tively, table.  5). C.  tinca is characterized by longer middle caudal fin rays 13.1–15.7 (mean 14.2) compared with C.  svanetica  sp.  n. 8.4–13.2 (mean 10.1) and also shorter anterior and posterior barbels like and C. baliki. C. svanetica sp. n. also characterized by 13.9–20.1 % (mean 16.7 %) and 18.7–28.6 % (mean 22.2 %) of anterior and pos- terior barbels length respectively compared to 8.1–14.1 % (mean 10.6 %) and 13.1– 19.3 % (mean 15.4 %) in C. tinca and 9.8–14.3 % (mean 12.7 %) and 14.7–18.5 % (mean 16.5 %) in C. baliki respectively. In total, for best taxonomic comparison were used only such features, which show the lowest variability level, do not overlap in related taxa and can be easily identifi ed. Th ese pri- marily are meristic (accountant) features (fi g. 1–4). Identifi cation key is proposed primarily based on such ones. 128 A. Roman, S. Afanasyev, O. Golub, O. Lietytska Identifi cation key for Capoeta Genus from Eastern and South-Eastern Black Sea basin rivers Below we provide identifi cation key and taxonomic accounts of Capoeta species distributed in Georgia and some Turkish rivers. Also to identifi cation is included previously known in such area C. tinca, recently renowned for the Sea of Marmara basin. (1a) Lateral line scales equal to or less than 61 (large scales Capoeta capoeta group, the Aral-Caspian group) (1b) 64 or more lateral line scales. If 61 or less — one pair of barbels ........ 2 (small scales Capoeta damascina complex group, the Anatolian-Iranian group) (2a) One pair of barbels, lips are fringed .........................C. sieboldi (East and South–East Black Sea drainage) (2b) Two pairs of barbels .............................................................................................................................................3 (3a) Usually 11 or more scales below lateral line; 7–10 total gill rakers; one pair of barbels .C. oguzelii (Ezine Stream (Black Sea basin) in Turkey) (3b) Usually less than 11 scales below lateral line; more than 12 total gill rakers ................................................4 (4a) Last unbranched fi n ray well ossifi ed, with 12 or more well-developed serrae ............................................5 (4b) Last unbranched fi n ray not ossifi ed, soft ; serrae are not well developed, their total number less than 7 or absent in samples with SL less than 100 mm ...C. svanetica sp. n. (Luchunis River (Rioni River basin) in Georgia) (5a) the number of scales rows below the lateral line is equal to 9 or less ........... С. banarescui (East Black Sea drainage from Rioni to Choroch) (5b) Number of scales rows below the lateral line is equal to 10 or more ............................................................6 (6a) More than 24 serrae on the last unbranched fi n ray C. tinca (Marmara Sea basin) (6b) Less than 24 serrae on the last unbranched fi n ray .................... C. baliki (Sakarya and Kızılırmak rivers) Capoeta sieboldi (Steindachner, 1864) Types. No types known. T y p e L o c a l i t y . Amasya. D i a g n o s i s . Meristic characters (tables 1–4): D: III–IV 8–9 (8.0), P: I 15–16, V: I 8–9, A: III 5½, lateral line: 52–60 (55.8), scales number above/below lateral line: 9–11 (9.6)/8–10 (8.9). C.  sieboldi is distinguished from other Capoeta species of East and South-East Black Sea rivers (C. svanetica sp. n., C. oguzelii, C. banarescui, C. baliki, C. ekmekciae, and also from C. tinca of Marmara Sea basin) by the combination of the following characters: one pair of barbels are present (only C. oguzelii have also one barbels pair); lower lip is fringed; scales large, only 52–60 total lateral line scales (all other compared Capoeta species of East and South-East Black Sea rivers have more than 64 scales); 9–11 scales rows above lateral line (fewer than for all other compared Capoeta species of East and South-East Black Sea rivers) as well as fewer gill rakers on the fi rst-gill arch number (28–33 (30.1)). D i s t r i b u t i o n . C. sieboldi is known from the western South Caucasus to the Sa- karya River (Turan et al., 2006 a). Later this species is noted in the Chorokh, Yesilırmak, and Sakarya rivers in Turkey (Elp et al., 2018) and total for Georgia (Kuljanishvili et al., 2020). Capoeta oguzelii Elp, Osmanoğlua, Kadak and Turan, 2018 Types. Holotype. FCME 2017-05a, 109 mm SL; Turkey: Kastamonu prov.: Ezine Stream at Devrekani, Black Sea drainage, 41°44'02" N, 33°52'58" E, M. Elp, A. Kadak, M. Osmanoglu, 29.06.2017. Paratypes. FCME 2017-05b, 9 individuals, 51–139 mm SL; same data as holotype. FCME 2017-34, 13 individuals, 54–78 mm SL; same location as holotype; 04.10.2017 (Elp et al., 2018). T y p e L o c a l i t y . Ezine Stream, a coastal stream in the southern Black Sea basin (Elp et al., 2018). D i a g n o s i s . Meristic characters (tables  1–4): D: IV 7 (71/2), P: I 16–17 (16.6), V: I 8–9 (8.1), A: III 5, lateral line: 72–82 (77.0), scales number above/below lateral line: 12–17 (13.8)/11–13 (11.6). 129Capoeta svanetica (Teleostei, Cyprinidae), a New Species from the Luchunis River… C. oguzelii is distinguished from other Capoeta species of East and South-East Black Sea rivers (C. svanetica sp. n., C. sieboldi, C. banarescui, C. baliki, C. ekmekciae, and also from C. tinca Sea of Marmara basin) by the combination of the following characters: one pair of barbels are present (only C. sieboldi have also one barbels pair); lower lip without keratinized edge; C.  oguzelii well distinguished from C.  sieboldi by small scales and its highest (72–82) total number; 11–13 scales rows below the lateral line (more than for all other compared Capoeta species of East and South-East Black Sea rivers); but only 7–10 (8.3) gill rakers (in comparison with previous); last unbranched dorsal-fi n ray not well ossifi ed (up to 20 %, Elp et al., 2018) with small serrae number, the similar to C. svanetica sp.nov. C. oguzelii is also characterized by small black spots presence. D i s t r i b u t i o n . Species is known only from Ezine Stream (Black Sea basin) in Turkey (Elp et al., 2018). Capoeta banarescui Turan, Kottelat, Ekmekçi and İmamoğlu, 2006 Types. Holotype. ESFM-PISI/2004-072, 177 mm SL; Turkey: Artvin: Tortum District: Çoruh drainage, stream Tortum, 100 km north of Erzurum; 40°34' N 41°36' E; D. Turan, F. Ekmekci, H. Imamoglu, O. Serdar, S.  Kırankaya, 19.07.2004. Paratypes. ESFM-PISI/2004-073, 4, 166-201 mm SL; FFR 712, 16, 85-232 mm SL; CMK 18474, 5, 135-193 mm SL; same data as holotype. — FFR 711, 9, 163 — 231 mm SL; CMK 18540, 9, 121 — 193 mm SL; Turkey: Artvin: Chorokh drainage, Bulanık stream, Savsat, 30 km east of Artvin, 41°34' N 42°14' E; D. Turan, F. Ekmekci, H. Imamoglu, O. Serdar, S. Kırankaya, 19.06.2004. — FFR 720, 3, 92 — 125 mm SL; CMK 18549, 1, 145 mm SL; Turkey: Cavuslu, Borcka, 41°21' N 41°42' E; D. Turan, 13.10.2004 (aft er Turan et al., 2006 b). T y p e L o c a l i t y . Chorokh River. D i a g n o s i s . Meristic characters (tables 1–4): D: III-IV 7–9 (8.0), P: I 17–19 (17.8), V: I 9–10 (9.1), A: III 5, lateral line: 64–77 (70.8), scales number above/below lateral line: 12–14 (12.8)/8–9 (8.1). C.  banarescui is distinguished from other Capoeta species of East and South–East Black Sea rivers (C. svanetica sp. n., C. sieboldi, C. oguzelii, C. baliki, C. ekmekciae, and also from C. tinca Sea of Marmara basin) by the combination of characters. Two pairs of barbels (C.  sieboldi and C.  oguzelii have only one pair); gill rakers number (12–16 (14.7)) higher than in C. oguzelii but fewer than for C. sieboldi, C. baliki, C. tinca, and C. ekmekciae; last unbranched dorsal-fi n ray well ossifi ed with the high number of serrae (unlike C. sieboldi and C. oguzelii); 8–9 scales rows below the lateral line (less than in C. baliki and C. oguzelii). C.  banarescui is also characterized by longer posterior barbels 18.4–28.8 (21.9) than the same parameter for C. baliki and C. tinca. D i s t r i b u t i o n . C. banarescui is known from Chorokh and Yesilırmak rivers (Turan et. al., 2006 a; Elp et al., 2018). Th is species is also noted for Georgian waters (Kuljanishvili et al., 2020). Some additional samples were studied from the Rioni River basin (Gubistskali River, fi g. 9) and Chorokh River (fi g. 10). Th ese individuals were recognized as C. banares- cui Th us, we should conclude C. banarescui is the widest distributed species in West Geor- gian rivers from Rioni to Chorokh. Th ere are no clear diff erences in morphological features (meristics and morphomet- rics, including mouth arching — fi g. 9, b and, 10 b) but some diff erences in general body appearance and coloration should be concluded. Specimen from Gubistskali River is slight- ly elongated with more concave dorsal and anal fi ns. Specimen from Chorokh River is slightly highest with straightly edged dorsal and anal fi ns. Th e coloration of the fi rst is gold- ish in total, darker on the back and lighter on the belly, with more dark (up to brown) fi ns. Chorokh’s specimen had more greyish coloration on the back and lighter (up to white) on the belly. All fi ns are gray. Th ese diff erences in coloration may be connected with condi- tions in the river. In the fi rst case, the specimen was sampled during fl oods, when river water was rich in sediments. Th e second case was diff erent — the specimen was collected in clear water. 130 A. Roman, S. Afanasyev, O. Golub, O. Lietytska Capoeta tinca (Heckel, 1843) Types. Lectotype: Naturhistorisches Museum Wien 55931:1, designated by Banarescu & Herzig-Straschil (aft er Banarescu, 1999). T y p e L o c a l i t y . “Brussa in Natolien” D i a g n o s i s . Meristic characters (tab.  1–4): D: III 8 (8.0), P: I 18–20 (18.8), V: I 8–9 (9.0), A: III 5, lateral line: 69–80 (74.9), scales number above/below lateral line: 14–17 (15.7)/9–11 (9.5). C.  tinca, previously known from the Rioni and Chorokh rivers was revised recently and noted only for the Marmara Sea basin in Turkey. Th is species is distinguished from the Capoeta species of East and South- East Black Sea rivers (C. svanetica sp. n., C. siebol- di, C. oguzelii, C. baliki, C. ekmekciae, and C. banarescui) by the combination of characters. Two pairs of barbels (C.  sieboldi and C.  oguzelii have only one pair); gill rakers number (19–23 (20.6)) higher than in C. svanetica sp. n., C. banarescui, C. baliki and C. oguzelii but fewer than for C. sieboldi; last unbranched dorsal-fi n ray well ossifi ed with the high number of serrae (unlike C.  sieboldi and C.  oguzelii); 14–17 scales above the lateral line are high- est than in C. svanetica sp. n., C. banarescui and C. sieboldi; 9–11 scales rows below lateral line are highest than in C.  svanetica  sp.  n. and C.  banarescui. C.  tinca also characterized by less length of anterior and posterior barbels 8.1–14.1 (mean 10.6) and 13.1–19.3 (mean 15.4) respectively than the same parameter for C. svanetica sp. n. (13.9–20.1 (mean 16.7) / 18.7–28.6 (mean 22.2)) and C. banarescui (12.4–20.8 (mean 16.9) / 18.4–28.8 (mean 21.9)). D i s t r i b u t i o n . C. tinca is known from the rivers draining to the southern shore of the Sea of Marmara (Turan et al., 2006 b). Capoeta baliki Turan, Kottelat, Ekmekçi and İmamoğlu, 2006 Types. Holotype. ESFM-PISI/2004-74, 202 mm SL; Turkey: Ankara: Sakarya River: Kızılcahamam Stream, Kızılcahamam, 60 km west of Ankara, 40°29' N 32°39' E; D. Turan, M. Turan, 15.04.2004. Paratypes. ESFM- PISI/2004-75, 4, 140–190 mm SL; FFR 713, 5, 121–219 mm SL; CMK 18541, 10, 128–188 mm SL; same data as Fig. 9. Live specimen of C. banarescui adult sample general body appearance (a), Gubistskali River and its lower lip structure (b). Fig.  10. Live specimen of C.  banarescui juvenile sample general body appearance (a), Chorokh River and its lower lip structure (b). 131Capoeta svanetica (Teleostei, Cyprinidae), a New Species from the Luchunis River… holotype. FFR 714, 5, 151-209 mm SL; Turkey: Ankara: Sakarya River, Ova Stream, Kazan, 50 km west of Anka- ra, 40°11' N 32°39' E; D. Turan, M. Turan, 15.04.2004. FFR 715, 5, 121-183 mm SL; same data, 16.06.2004. FFR 716, 10, 168-217 mm SL; Turkey: Sıvas: Kızılırmak River, Delice Stream; F. Ekmekci, S. Kırankaya, 22.11.2002 (aft er Turan et al., 2006 b). T y p e L o c a l i t y . Sakarya River. D i a g n o s i s . Meristic characters (tables 1–4): D: III–IV 8–9 (8.1), P: I 17–20 (18.4), V: I 9–10 (9.0), A: III 5, lateral line: 72–86 (78.4), scales number above/below lateral line: 14–17 (14.9)/10–11 (10.1). C.  baliki is distinguished from other Capoeta species of East and South-East Black Sea rivers (C. svanetica sp. n., C. sieboldi, C. oguzelii, C. banarescui, C. ekmekciae, and also from C. tinca of Marmara Sea basin) by the combination of characters. Two pairs of barbels (C.  sieboldi and C.  oguzelii have only one pair); gill rakers number (16–22 (19.3)) higher than in C. svanetica sp. n., C. banarescui and C. oguzelii but less than for C. sieboldi; last un- branched dorsal-fi n ray well ossifi ed with the high number of serrae (unlike C. sieboldi and C. oguzelii); 14–17 scales rows above the lateral line and 10–11 scales rows below the lateral line (more than in C. svanetica sp. n., C. sieboldi, and C. banarescui). C. baliki also charac- terized by less length of anterior and posterior barbels 9.8–14.3 (mean 12.7) and 14.7–18.5 (mean 16.5) respectively than the same parameter for C. svanetica sp. n. (13.9–20.1 (mean 16.7) / 18.7–28.6 (mean 22.2)) and C. banarescui (12.4–20.8 (mean 16.9) / 18.4–28.8 (mean 21.9)). D i s t r i b u t i o n . C.  baliki is presently known from the Sakarya and Kızılırmak river drainages (Turkey), including lakes and reservoirs (Turan et al., 2006 b; Elp et al., 2018). Capoeta ekmekciae Turan, Kottelat, Kirankaya and Engin 2006 Types. Holotype. ESFM-PISI/2004-076, 203, 203 mm SL. Paratypes. ESFM-PISI/2004-077, 4, 150–209 mm SL (aft er Turan et al., 2006 a). T y p e L o c a l i t y . Chorokh River. D i a g n o s i s . Meristic characters: D: III–IV 8–9 (8.1), P: I 16-20, V: I 12, A: III 5½, lat- eral line: 55–61 (57.5), scales number above/below lateral line: 9–10 (9.4)/6–7 (6.8). C. ekmekciae is well distinguished from other Capoeta species of East and South-East Black Sea rivers (C. svanetica sp. n., C. sieboldi, C. oguzelii, C. banarescui, C. baliki, C. ekme- kciae and also from C. tinca of the Sea of Marmara basin) by the combination of characters. Only one pair of barbels is present, 55–61 lateral line scales and the keratinized edge of the lower lip (C. sieboldi and C. oguzelii have only one pair, but fi rst also had fringed lower lip and the second had lower lip without keratinized edge and 72–82 scales in lateral line). Based on other meristic features C. ekmekciae is more similar to the Aral-Caspian group (tables 1–4). Th is group, known as large scales, is well distinguished by fewer scales number in lateral line (less than 61), fewer scales rows above the lateral line and below lateral it but higher gill rakes number. D i s t r i b u t i o n a n d s o m e t a x o n o m i c r e m a r k s . Based on the phylogenetic rela- tionship of Capoeta species using COI and cytb sequences was shown (Zareian et al., 2016; 2018) that C. ekmekciae belongs to the Capoeta capoeta complex (Aralo-Caspian group). Th is data is based on one sample analysis with an unknown locality. Capoeta capoeta com- plex includes species characterized by large scales, their fewer number in lateral lines (fewer than 64), absence of irregular black spots on the dorsal half of the body, and by only one pair of barbels. Our comparison (tables 1–4) shows it well similar to this group species. But, based on all known data (Baycelebi et al., 2015; Zareian et al., 2016; 2018; Elp et al., 2018 and others) including the fi rst description (Turan et al., 2006 a) C. ekmekciae is known only from the lower Chorokh River near Borcka and Cavuslu (Black Sea basin). Th us, this spe- cies recent distribution needs to be reviewed. 132 A. Roman, S. Afanasyev, O. Golub, O. Lietytska Discussion According to Berg (1912–1914 and 1949), only three species of the genus Capoeta were known for Caucasus water fl ows in total (and for Georgia in particular): Varicorhinus capoëta (Güldenstaedt, 1787), Varicorhinus sieboldii (Steindachner, 1864) and Varicorhinus tinca (Heckel, 1843). First species distributed within the Kura River basin and Araxes River basin particularly. Th e second species inhabited Northern Minor Asia coastal rivers and the Western Caucasus from the Sakarya River to Rioni. Th e third of them was distributed as in Minor Asian rivers (from Bursa to Trabzon towns) and the Chorokh River basin. Varico- rhinus capoëta as distinct species also included three subspecies: V. сapoëta sevangi (Sevan Lake, Araxes River basin and some Kura River tributaries), V. capoëta gracilis (Lenkoran Region rivers) and V. capoëta heratensis (Turkmenistan water fl ows). In 1969 (Karaman, 1969) all these taxons were replaced to the genus Capoeta and later (Zareian et al., 2016; 2018) three subspecies based on genetic data were redescribed as distinct species: Capoeta sevangi De Filippi, 1865, C. gracilis (Keyserling, 1861) and C. heratensis (Keyserling, 1861). Th ese three species belong to Capoeta capoeta complex (Aralo-Caspian group) which is characterized by larger scales and their fewer number in the lateral line and absence of ir- regular black spots on the dorsal half of the body (Zareian et al., 2016; 2018). Also, C. kaput (Levin et al., 2019) was described. C. svanetica sp. n. with two pairs of barbels, small scales and spotting absents is more similar to Capoeta damascina complex group (Anatolian– Iranian group (Levin et al., 2012; Zareian et al., 2018)) which also includes C. banarescui, C.  baliki and C.  sieboldii (Levin et al., 2012). Th is group of species also called the Young Evolutionary Group (Zareian et al., 2016): younger than Aral-Caspian one, widespread and diversifi ed group of species. Th e fi rst species to diverge within this group clade was C.  sieboldi (samples from the Kizilirmak River, Black Sea drainage (Levin et al., 2012)). Within this group, C. sieboldi a single species with a single pair of barbells, an arched mouth and fringed lips. Separation of the Black Sea clade (including also C. baliki and C. banares- cui) took place during Pliocene (Levin et al., 2012). Noted species are characterized by two pairs of barbels and large scales numbers in the lateral line and above it. Members of this group have 2–4 barbels and a horseshoe-shaped lower jaw (Karaman, 1969). C. tinca should also belong to the Anatolian–Iranian group because it shows high relatedness with C.  baliki (Zareian et al., 2018). Th e third Capoeta trutta complex group (spotted capoeta group or the Mesopotamian Capoeta group includes species spread at Tigris, Euphrates and Orontes drainages — Capoeta trutta (Heckel, 1843), at Seyhan drainage and southern Anatolia — Capoeta turani Özuluğ and Freyhof, 2008 and at Tigris and Euphrates drain- ages — Capoeta barroisi Lortet, 1894 (Zareian et al., 2016). Th ere is a clear correlation between molecular and morphological data in Capoeta species. Th us, Capoeta trutta complex or the Mesopotamian group is characterized by having numerous irregular black spots on the dorsal half of the body or fl ank and fi n. Capoeta capoeta complex or the Aralo-Caspian group characterized by large scales and plain body (absence of irregular black spots on the dorsal half of the body) distributed in Aralo-Caspian water bodies (e.g., Kura and Araxes River drainages, Lake Sevan drainages, and many rivers from Sefi drud to Atrak) in the Caspian Sea basin. Capoeta damasciana or the Anatolian–Iranian group characterized by small scales and plain body (absence of irregular black spots on the dorsal half of the body) encompasses many species occupying the majority of Capoeta’s range, including Anatolia, the Zagros Mountains, Mesopotamia, and the Iranian plateau (Zareian et al., 2016). It has also been noted that Capoeta species with four barbels, like all Luciobarbus species with four barbels, are more ancient than species with two barbles (Karaman, 1969; Levin et al., 2012). But this hypothesis was not supported due to phylogenetic relationships between three Capoeta species groups (Levin et al., 2012; Zareian et al., 2016; 2018). Species belonging to C. trutta group, the earliest lineage in the genus, have two barbels. Samples 133Capoeta svanetica (Teleostei, Cyprinidae), a New Species from the Luchunis River… belonging to the two other clades have 2–4 barbels. It should be noted (Levin et al., 2012) that the number of barbels may be retained in some taxons, whereas other species could rapidly lose them independently of their branch due to the specialization required to scrape algae from stones. It has been proposed that the number of barbels is an evolutionarily reversible character in Capoeta (Levin et al., 2012). It is worth mentioning that a correlation between molecular and morphological data in Capoeta species needs to be revised in detail. At a present time, there are no clear data on the phylogenetic structure between all taxons within the Capoeta species from all Caucasus drainage and neighbor areas. An example of C.  sieboldi phylogenetic history shows us morphological features analysis opportunities. Th e use of this set of data makes it possible to analyze the historical background for studying the formation and development of individual morphological characters. We had used only clear morphological diff erences for comparison of Capoeta species. Th e meristic features were preferred as less variable in comparison with metric features. But the last was used also in comparison to the most related morphologically species. We did not include large scales Capoeta capoeta (Aralo-Caspian) group into identifi cation keys because of their clear diff erences from small scales (Anatolian-Iranian) group, distributed in Eastern and South-Eastern rivers of the Black Sea basin. Conclusion Based on the results of comparing C. svanetica sp. n. with other Capoeta species from the rivers of the Black Sea basin, we can conclude that this is a separate species that stands out from the complex group Capoeta damascina, also known as the Anatolian-Iranian group (C. baliki, C. banarescui, C. sieboldii and C. oguzelii) by the combination of the char- acters. New species is also well distinguished from the Capoeta capoeta complex group (large scales Capoeta group, the Aral-Caspian group) which include C. capoeta, C. sevangi, C.  kaput and may also include C.  ekmekciae as more similar to this group based on ge- netic and morphological data analysis. Finally, C. svanetica sp. n. is well distinguished from C. tinca previously known for the Chorokh and Rioni River drainage. C. svanetica sp. n. known only from the Luchunis River (Rioni basin). C.  banarescui is recognized as well spread species in East and South-East Black Sea rivers. Th is species is also noted for so high level of coloration changeability. At present, Georgian fi sh fauna includes fi ve species of the genus Capoeta: C. capoeta, C. kaput, C. banarescui, C. sieboldii, and C. svanetica sp. n. References Banarescu, P. 1999. Th e freshwater fi shes of Europe. Aula, Wiesbaden, 426 (5. Cyprinidae 2. Part I. Rhodeus to Capoeta). Berg, L. S. 1912–1914. [Faune de Russie et des pays limitrophes. Poissons (Marsipobranchii et Pisces). Vol. 3. Ostariophysi]. Izdatelstvo Akademii Nauk, St-Petersburg, 1 (1912), 1–336, 2 pls, 2 (1914), 337–704, 4 pls. [In Russian]. Berg, L. S. 1949. Fishes of Fresh Waters of the USSR and Adjacent Countries. AN SSSR, Moscow, 469–925 (Vol. 2) [In Russian]. Baycelebi, E., Turan, D., Japoshvili, B. 2015. Fish Fauna of Coruh River and Two First Record for Turkey. Turk- ish Journal of Fisheries and Aquatic Sciences, 15, 783–794. Elp, M., Osmanoğlu, M., Kadak, A., Turan, D. 2018. 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