Capoeta svanetica Roman, Afanasyev, Golub et Lietytska, 2022

Roman, A., Afanasyev, S., Golub, O. & Lietytska, O., 2022, Capoeta Svanetica (Teleostei, Cyprinidae), A New Species From The Luchunis River (Rioni River Drainage) In Georgia, Zoodiversity 56 (2), pp. 117-134 : 119-128

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 drainage 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).

Material. 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) .

Diagnosis. C. svanetica sp. n. is distinguished from the other genus Capoeta species 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 first gill arch (14 in holotype); serrae number in the last unbranched fin ray is 7–9; lateral head length (HL) is 22.1–27.7 % of SL (22.1 % in holotype); anal fin base length is 7.4–9.4 % of SL (7.4 % in holotype); eye diameter 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 figure 2 ; morphometric and meristic data are present in tables 2–5. The body is elongated and cylindrical with a slightly convex upper profile and less convex ventral one. The head is relatively short; the upper profile is slightly convex in the interorbital and is slightly concave at the level of the nostrils. The mouth is inferior, narrow, and slightly arched ( fig. 3 ). Lips are slightly fleshy. The 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 fin with 3 or 4 simple (unbranched) and 7 branched rays (4 and 7 in holotype, respectively), the outer margin is slightly concave, origin slightly in front of vertical through the pelvic-fin origin, last simple ray only slightly ossified, proximal two thirds rigid, and without serrae on posterior margin in small fish (less than near 100 mm standard length) or with 7–9 small serrae in adult ( fig. 4 ).

Pectoral fins do not extend to the pelvic-fin base; their outer margins are usually slightly convex with 16–22 branched rays (table 1) in total (18 in holotype). Pelvic fins (table 1)

* total used samples are different, see tables 2–4.

N o t e. Data on C. banarescui , C. baliki and C. tinca noted after 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.

with 1 simple and 8–9 branched rays (9 in holotype), it is not extending to anal-fin base, their outer margin straight or slightly convex; pelvic axillary scales present. Anal fin with 3 simple and 51 / 2 branched rays, the outer margin is convex. The caudal fin is long and deeply forked, its upper lobe often longer than the lower one.

Scales small, total lateral-line scales 70–74 (table 2), 10–12 between dorsal-fin origin and lateral line, and 7–8 between anal-fin 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 first-gill arch (table. 4). Pharyngeal teeth arranged in 3 rows as 4.3.2–2.3.4.

Coloration. In formaldehyde-fixed samples, the total body color is greyish on the back and upper part of the flank and light grey on the lower flank. There are no spots on the body and head. Dorsal and caudal fins are grey; pectoral, anal, and pelvic fins are yellowish ( fig. 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 finished. Thus, we can speak of sexual dimorphism as independent of the spawning period. But breeding tubercles present in anal-fin rays in males are more typical for the spawning period. Thus, this question needs to be investigated in detail. We can

13.3 14.7 30.1 19.3 8.3 20.4 21.3 24.5 20.6 only bercles conclude in adult about males the in the presence second of summer epithelium half and tu-

their absence in mature females at the same period.

Small epithelial tubercles ( fig. 5 ) are present on the head, anal fin rays, and flank scales (one per apical edge of scales) in males during the second half sum-

2 mer season at least.

In females, epithelium tubercles are absent during

4

the second half summer season. Any differences based

5 on meristic (tables 1–4) or morphometric ( table. 5)

data were not found as like as no clear differences in the lower lip shape ( fig. 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. The species occurs in medium-fast flowing rivers

1 3 1 with usually gravel substrates and clear waters. Sampling sites of the Luchunis River should be divided into

23

4 2 two types ( fig. 6 ), but both are characterized by a sub-

22

1 2 5 5 strate consisting of coarse gravel and boulders, and fast-

21

9 5 6 4 flowing The and first translucent riverbed type waters is (Afanasiev classified as et al a. braided 2022).

20

8 2 6 8 channel ( fig. 6, a and 6 , c). Two islands divide the main

19

6 4 3 6 are flow 80 into m length three and branches in average. The 9 first m width island (23 parameters m max.);

the second island had 25 m length and in average 5 m

4 6 2width (12 m max.). The riverbed cross-sections were

16 6 2 species 17 5 The made shape average; for banks each velocity were branch covered is separately 1.03 by m trees /s (, fig with, bushes. 6, c the). Valley, maximum and grass is U-. Capoeta 15 1 11 at ing 1.67 waves m/, s. and Flow unbroken types include standing chaotic waves, broken. The standright in 14 1 4 branch average width is 9 m (it is varied between 2 variation 12 1 2 13 3 2 the and tween maximum 7.5 3.8 m and); the at 9 left m 0.45). branch The m. Both average average river depth banks width is were 0.24 is 6 m mainly m with (bearch made by pebble and the riverbed was covered by cobgill ble (52 %), pebble (24 %), and 9 % of boulders. first a single The channel second riverbed type. The type average (fig. velocity 6, b) is classified is 0.99 m / as s, of 9 7 with the maximum at 1.39 m /s. Flow types include side 8 3 turbulent, broken, and unbroken standing waves and outer n 6 25 23 35 15 24 28 2 25 6 ripples, 5 m and. The 10 average m). Bed width elements is 8 m include (it is varied bars between, rapids, on riffles, rocks, and step/pools. The average depth is rakers

.

n. 0 evenly, 49 m covered with the by maximum cobble riverbed at 1.7 m .. Valley Among is substrate U-shape

Gill b

. e 4 l a sp svanetica . banarescui . sieboldii .. baliki oguzelii . ekmekciae . capoeta . kaput. tinca . gia from types where E the also t y m name this prevails o l o species g Svaneti y. cobble The occurred, name the and historical of pebble. the species. region is in derived Geor- T C C C C C C C C C

Note. Data on C. tinca , C. baliki and C. banaresqui after Turan et al., 2006 b.

Comparison with closely related species

Using morphological data analysis (tables 1–4) were compared C. svanetica sp. n. samples 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 flows. From the Sea of Marmara basin: C. tinca as previously known species distributed in the Rioni River drainage; and also from Caspian 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 fixed in formaldehyde during the year and they practically lost their coloration. We can state only complete any spots absence on the fish body, head, and fins in C. svanetica sp. n. ( fig. 2 ) the same as closely related C. banarescui ( fig. 7 , a) and C. baliki ( fig. 7 , d, Turan, et al., 2006 b) from Black Sea rivers basins and previously known for the same areal C. tinca ( fig. 8 , a, Turan et al., 2006 b) This live and formaldehyde preserved specimens are characterized by dark brown on back and flanks, 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. banaerscui are also characterized by two pairs of barbels, but the first of them ( C. baliki ) has 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 barbels length. But the last one clearly distinguishes from C. svanetica sp. n. by more highest serrae number in the last unbranched fin ray ( fig. 4 ; 12–20 in C. banaerscui compared to C. svanetica sp. n. without serrae in the last unbranched fin ray in small fish (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 fin rays (7–8 (mean 7.1) vs. 8–9 in other species); from C. sieboldii and C. oguzelii C. svanetica sp. n. also differs by the fewer number of pectoral fin rays (15–16 and 16 (common) — 17 (mean 16.6) respectively vs. 16–18 (mean 17.3)); and from C. ekmekciae the last one differs by the fewer number of pelvic fin 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.

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. The 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 first gill arch rakers number we should conclude C. svanetica sp. n. (12–15 with mean 13.3) is more similar to C. banarescui (12–16 with mean 14.7). C. oguzelii differenced from C. svanetica sp. n. by fewer (7–10, mean 8.3) gill rakers number. C. baliki , C. ekmekciae and C. sieboldii differenced 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 fin number than C. kaput . The 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 ossified last unbranched fin ray with high serrae number and also by highest gill rakers number on the first 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 features: 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. svanetica 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 respectively, 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 posterior 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 identified. These primarily are meristic (accountant) features ( fig. 1–4 ). Identification key is proposed primarily based on such ones.

Identification key for Capoeta Genus from Eastern and South-Eastern Black Sea basin rivers

Below we provide identification key and taxonomic accounts of Capoeta species distributed in Georgia and some Turkish rivers. Also to identification 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 fin ray well ossified, with 12 or more well-developed serrae............................................5

(4b) Last unbranched fin ray not ossified, 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 fin ray C. tinca (Marmara Sea basin)

(6b) Less than 24 serrae on the last unbranched fin ray .................... C. baliki (Sakarya and Kızılırmak rivers)

Figures

Fig. 2. The general body appearance of C. svanetica sp. n. (male — top, SL = 136 mm and female — bottom, SL = 118 mm).

Fig. 3. Mouth shape male (left) SL = 136 mm and female (right) SL = 118 mm.

Fig. 4. Last simple (unbranched) dorsal fin rays: a — C. tinca, 139 mm SL, female (after Turan et al., 2006 b); b — C.banarescui, 144 mm SL, female (after Turan et al., 2006 b); c — C. baliki, 148 mm SL, female (after Turan et al., 2006 b) and d —C. svanetica sp. n. 118 mm SL, female — only 7 serrae are present.

Fig. 5. Epithelial tubercles on each scale and anal fin rays of C. svanetica sp. n. males during the second half summer season.

Fig. 6. Type habitats of juvenile Capoeta svanetica sp. n. (a) and adult (b) with detailed river bed structure (c).

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), after Turan et al, 2006 b; d–f: C. baliki, holotype, 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), after 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), after Turan et al, 2006 b; d–f: C. oguzelii, holotype, FCME 2017-05a, 109 mm SL; Ezine Stream (d); lower lip (e); dorsal fin with soft last unbranched fin ray (f), after Elp et al., 2018.

Fig. 1. Genus Capoeta distribution in river basins of Georgia and some neighbors.