Garra mondica, Sayyadzadeh, Golnaz, Esmaeili, Hamid Reza & Freyhof, Jörg, 2015

Sayyadzadeh, Golnaz, Esmaeili, Hamid Reza & Freyhof, Jörg, 2015, Garra mondica, a new species from the Mond River drainage with remarks on the genus Garra from the Persian Gulf basin in Iran (Teleostei: Cyprinidae), Zootaxa 4048 (1), pp. 75-89 : 78-88

Garra mondica , new species

( Figs. 2–6 )

Holotype. ZM-CBSU H1032, 66 mm SL; Iran: Fars prov.: Konar Siyah spring at Firuzabd, 28°43'40''N 52°25'20''E; G. Sayyadzadeh, M. Razbanian, A. Khajeh Panah and H. Zareiyan, 10 March 2015.

Paratypes. All from Iran. ZM-CBSU H1033, 8, 39–64 mm SL; same data as holotype.—ZM-CBSU K1080, 7, 41–58 mm SL; Iran: Fars prov.: Konar Siyah spring at Firuzabd, 28°43'40''N 52°25'20 ''E; S. Babaee and R. Khaefi, 15 Dec 2012.—ZM-CBSU B242, 1, 48 mm SL; ZM-CBSU B250, 1, 53 mm SL; ZM-CBSU B255, 1, 49 mm SL; Fars prov.: Tang-e-Mohr spring at Lamerd, 27°31'36''N, 52°51'47''E; 6 June 2008.

Additional material. All from Iran. ZM-CBSU B236, 2, 33–47 mm SL; ZM-CBSU B240, 1, 48 mm SL; ZM- CBSU B244, 1, 38 mm SL; ZM-CBSU B246, 4, 39–50 mm SL; ZM-CBSU B251, 4, 37–48 mm SL; Fars prov.: Tang-e-Mohr spring at Lamerd. 27°31'36''N, 52°51'47''E.—ZM-CBSU H1030, 2, 59–63 mm SL; mm SL;—ZM- CBSU H1041, 12, 32–71 mm SL;— FSJF 3488, 3, 53–61 mm SL; Fars prov.: Konar Siyah spring at Firuzabd, 28°43'22''N 52°25'21''E.

Material for molecular genetic analysis: ZM-CBSU M699; Fars prov.: Tang-e-Mohr spring at Lamerd, 27°31'36''N, 52°51'47''E, GenBank accession number: KM214741 .—ZM-CBSU M793–M794; Fars prov.: Konar Siyah spring at Firuzabd, 28°43'40''N 52°25'20''E, GenBank accession numbers: KM214762 , KM214736 .

Diagnosis. Garra mondica is distinguished from all other species of Garra in the rivers flowing to the Persian Gulf south of the Tigris by having a naked breast and anterior belly (vs. fully covered by scales) and a naked middorsal area in front of the dorsal-fin origin or dorsal midline with only few, usually embedded scales ( Fig. 2 ). The naked mid-dorsal area seems to be an unique character state of this species within its comparison group including the species of the Euphrates and Tigris drainages. A naked breast and anterior belly is also found in some populations of G. r u f a. Garra mondica is further distinguished from G. r u f a by having usually 7½ branched dorsalfin rays (vs. usually 8½). Garra mondica is distinguished from all the congeners included in this study by 2 fixed, diagnostic nucleotide substitutions ( Table 1 ), and a K2P nearest-neighbor distance of 3.9% to G. rufa ( Table 2 ).

Besides the naked breast, anterior belly and predorsal back, G. mondica is also distinguished from G. persica by having usually 9+8 caudal-fin rays (vs. usually 8+8). Garra mondica is distinguished from G. rossica by having a fully-developed mental disc (vs. reduced) and 18–23 total gill rakers on the first branchial arch (vs. 11–15), from G. variabilis by having a fully developed mental disc (vs. reduced) and two pairs of barbels (vs. one) and from G. typhlops and G. widdowsoni by having well-developed eyes and a brown or grey, usually mottled colour pattern (vs. absence of colour pattern).

Nucleotide position relative to Oryzias latipes complete mitochondrial genome ( AP004421 ).

Nucleotide position relative to Oryzias latipes complete mitochondrial genome ( AP004421 ).

Description. See Figures 2–6 for general appearance; morphometric data are given in Table 3 . Body elongated, moderately compressed laterally, more compressed in region of caudal peduncle. Dorsal head profile rising gently, slightly convex, more or less continuous with dorsal body profile to nape or about middle between nape and dorsal-fin origin. Ventral profile more or less straight to anal-fin origin. Head moderately large and depressed, with slightly convex or flat interorbital space; height at nape less than head length; width at nape greater or about equal to depth at nape. Snout roundish; transverse lobe with 11–21 tubercles, demarcated posteriorly by a shallow transverse groove in some individuals, no transverse groove in others. Proboscis covered with small or medium sized tubercles, largest on anterior margin of proboscis. Proboscis not or only slightly elevated from depressed rostral surface. Lateral surface of snout covered by small to medium sized tubercles reaching to anterior eye margin in some individuals, or to posterior nostril in others. Depressed rostral surface always without tubercles, anterior arm of depressed rostral surface not reaching to base of rostral barbel, clearly separating transverse lobe from lateral surface. No groove between transverse lobe and lateral surface in some individuals. No head tubercles in juveniles and in a few adults. Eye placed dorso-laterally in posterior half of head. Barbels in two pairs: rostral barbel anterolaterally located, shorter or about equal to eye diameter; maxillary barbel at corner of mouth, shorter than rostral barbel. Rostral cap well-developed, fimbriate, papillate on ventral surface. Upper lip present, as a thin band of papillae arranged in two ridges. Upper jaw almost or completely covered by rostral cap. Disc elliptical, shorter than wide and narrower than head width through base of maxillary barbel; papillae on anterior fold of same size, regularly arranged; groove between antero-median fold and central callous-pad narrow and deep, papillae on inner half of whole length of latero-posterior flap coarsely arranged; anterior marginal surface of central callous pad without or with sparsely arranged small papillae; posterior margin of latero-posterior flap extending vertical to middle of eye.

Dorsal fin with 3 simple and 7½ (28)–8½ (12) branched rays, last simple ray slightly shorter than head length; distal margin concave; origin closer to snout tip than to caudal-fin base; inserted anterior to vertical of pelvic-fin origin; first branched ray longest, tip of last branched ray reaching vertical to, or slightly in front of anus. Pectoral fin with 1 simple and 12–14 branched rays, reaching to a point 3–4 scales anterior to pelvic-fin origin, length shorter than head length. Pelvic fin with 1 simple and 7–8 branched rays, reaching to anus or anal-fin base, origin closer to anal-fin origin than to pectoral-fin origin, inserted below third or fourth branched dorsal-fin ray. Anal fin short, with 3 simple and 5½ branched rays; first branched ray longest; distal margin straight or slightly concave; origin closer to pelvic-fin origin than to caudal-fin base. Anus 2–3 scales in front of anal-fin origin. Caudal fin forked with 9+8 branched rays; tip of lobes pointed. Total gill rakers on first branchial arch 18–23 [18(3), 19(8), 20(6), 21(2), 23(1)]. Lateral line complete, with 28–32 [28(1), 29(2), 30(6), 31(9), 32(8)] scales on body and 2 scales on caudal-fin base. Transverse scale rows above lateral line 3–4; between lateral line and pelvic-fin origin 3 and between lateral line and anal-fin origin 3. Circumpeduncular scale rows 13–14. Usually, no scales on predorsal midline between dorsal-fin origin and nape (20 specimens). Seven specimens with 2–4 scales in front of dorsal-fin origin; nine individuals with a few, deeply embedded scales on predorsal midline. Scales on flank regularly arranged. Chest and belly naked. In some individuals belly covered with embedded scales between posterior tip of pectoral fin when folded back and pelvic-fin base. One long axillary scale at base of pelvic fin, 4–6 (usually 5) scales between posteriormost pelvic-fin base and anus, embedded in skin in some individuals.

Coloration. In ethanol: Head, dorsum and flank dark or pale brown. Single, or groups of dark-brown scales on flank. A very faint irregularly shaped midlateral stripe restricted to posterior flank, absent in some individuals, usually interrupted. Mouth, chest and abdomen yellowish white. A wide black or dark-brown bar at posteriormost caudal peduncle, faded in some individuals, bold, 2–3 scales wide in others. Bar reaching dorsal midline in most individuals, not reaching ventral midline. A small black blotch at anterior most lateral line. Lateral line pores cream whitish. Fins hyaline with irregularly set black spots on rays, or rays partly dusty grey or black. Base of last 4 – 6 branched dorsal-fin rays with a black spot or black in individuals larger than 40 mm SL, hyaline in others. In life: Fins hyaline with black spots. Head grey. Flank scales dark grey with individual or groups of pale grey scales forming a mottled pattern; whitish or pale grey on ventral flank and belly. Iris silvery orange. A pale blue dot at anterior-most lateral line reaching down to upper pectoral-fin base in some individuals. A faint irregularly shaped midlateral stripe in few individuals.

Distribution. Garra mondica was found in two small springs in the Mond River drainage in Iran, the Konar Siyah spring ( Fig. 7), which is situated 20 km south-west of Firouzabad and the Tang-e-Mohr spring, which is situated about 10 km south of Mohr.

Etymology. The species is named for the Mond River. An adjective.

Remarks. Garra mondica and G. rufa both occur in the Mond River drainage. While G. r u f a is widespread, it has not been found in sympatry with G. mondica , which has been recorded only from two small springs. Within the Mond River drainage, both species are distinguished by the naked breast, anterior belly and predorsal midline in G. mondica (vs. covered by scales in G. r u f a) and by usually 7½ branched dorsal-fin rays (vs. 8½).

Discognathus crenulatus was described by Heckel (1849) from the area of Shiraz, "Confluenten des Araxes, als aus den Quellen des Saadi und dem Kara-Agatsch [Confluence of Araxes Rivers as well as from the springs of Saadi and from Kara-Agatsch]". The “Confluence of Araxes Rivers” belongs to the Kor drainage, the “springs of Saadi belong to the Lake Mahalo basin and the “Kara-Agatsch” belongs to the Mond River drainage. Coad (1991) treated D. crenulatus as a synonym of G. rufa while Bianco & Banarescu (1982) considered it as a subspecies of G. rufa . JF examined syntypes of D. crenulatus at NRM (NMW 53236–37). All fishes have the breast and belly as well as the pre-dorsal midline fully covered by scales. No difference between the syntypes of D. crenulatus and G. rufa could be found and we follow Coad (1991) and treat D. crenulatus as a synonym of G. rufa .

Berg (1949) described G. rufa gymnothorax from the Iranian Karun drainage, a tributary of the lowermost Tigris, distinguishing G. r. gymnothorax from G. r. r u f a by its naked breast (vs. scaled). Alexander Naseka (Dolsko) examined syntypes (ZIN 13214–15) of this nominal species for us, and these fishes have indeed a naked breast as described by Berg (1949). They are distinguished from G. mondica by having the predorsal midline fully covered by scales (vs. naked in G. mondica ) and usually 8½ branched dorsal-fin rays (vs. usually 7½). Coad (1991) treated G. r. gymnothorax as a synonym of G. r u f a. Our molecular data suggest that fishes from the Karun River drainage might represent a distinct species, as they form a separate molecular group, well differentiated from G. r u f a ( Fig. 1 : G. cf. gymnothorax ). No Garra with a naked breast have been found by us in the Karun River drainage until now. All fishes included in the molecular analysis or examined by us at other occasions have their breast fully covered by scales. While G. gymnothorax has been distinguished as a distinct subspecies from G. r u f a by Berg (1949) due to its naked breast, this character seems to be quite variable and does not allow distinguishing our molecular lineage from G. r u f a. The squamation pattern of the breast and belly is quite variable in different populations of G. r u f a and does not allow distinguishing G. mondica from all populations of G. r u f a. Beside the syntypes of G. r. gymnothorax , the breast and the midline of the anterior belly are naked in the fishes examined from a tributary to Atatürk reservoir (Euphrates drainage) in Turkey (FSJF 2549). The abdomen is fully covered by embedded, overlapping scales and there are few isolated, deeply embedded scales on the breast in fishes collected from a second tributary to Atatürk reservoir (FSJF 2588). The breast and the anterior abdomen are fully covered by embedded scales except the distalmost pectoral-fin base and the anterior tip of the breast in fishes from the Karun (FSJF 2198), the Karkheh (ZM-CBSU J2501, ZM-CBSU J2404, ZM-CBSU J2084), the Sirvan (ZM-CBSU J2337, ZM-CBSU J2006) and the Zohreh River (ZM-CBSU J1051) and the breast and the anterior abdomen are fully covered by unembedded scales except the anterior tip of the breast, which is naked in fishes from a spring in the Iranian Lake Maharlo basin (FSJF 2238). Additional studies are needed to test whether there are morphological characters distinguishing the molecular lineage called here G. cf. gymnothorax from G. r u f a. It is beyond the aims of this study to clarify this issue.

In our molecular dataset, two additional populations of Iranian Garra are very closely related to G. mondica . These are fishes from the springs Golabi and Qalatoyeh, both situated in the Kol River drainage in the wider area of the Iranian city of Darab. Fishes from both these populations do not share the diagnostic characters of G. mondica , as both have the breast, anterior belly and predorsal midline covered by scales (vs. naked in G. mondica ) and a greater postorbital distance (40–48% HL vs. in posterior, 36–43% HL). These two populations might also represent G. mondica , which might have a variable squamation pattern as is the case in G. r u f a and the molecular lineage, which might be conspecific with G. gymnothorax . But fishes from the Golabi and Qalatoyeh springs are also distinguished from G. mondica by having frequently 8+8 branched caudal-fin rays (vs. almost always 9+8) and frequently 8½ branched dorsal-fin rays (vs. almost always 7½). Therefore, we do not identify these fishes as G. mondica .

As we cannot identify the fishes from the Golabi and Qalatoyeh springs as G. mondica , they might be identified as populations of G. persica or G. r u f a that have been introgressed by G. mondica due to postulated past hybridization events. Introgressed fishes can usually be identified as one species, but share the mtDNA with another species. Introgressive hybridization has been documented several times in cyprinid fishes ( Freyhof et al.

2005), but was found to be an uncommon phenomenon in Mediterranean freshwater fishes by Geiger et al. (2014). The springs Golabi and Qalatoyeh are situated in the upper Kol River drainage. Garra persica is widespread in the lower Kol River drainage. The fishes from the springs Golabi and Qalatoyeh are distinguished from G. p er s i c a by having 7½ (18) or 8½ (12) branched dorsal-fin rays (vs. 6½ (7)–7½ (10)) and by having frequently 8+8 (5) or 9+8 (22) branched caudal-fin rays (vs. almost always 8+8). If a hybridization of G. mondica with G. persica is suspected, this could lead to an increased frequency of fishes with 9+8 branched caudal-fin rays in the hybrid population, as G. mondica has almost always 9+8 and G. persica has almost always 8+8 branched caudal-fin rays. This scenario does not explain easily the increased frequency of fishes with 8½ branched dorsal-fin rays. Garra rufa is a species widely distributed in all rivers north of the Kol drainage diagnosed by having almost always 8½ branched dorsal-fin rays. While it might be speculated that the fishes from the Golabi and Qalatoyeh springs might have originated from successive hybridization events between G. mondica , G. persica and G. r u f a, or might represent a separate, still undescribed species, it is far too speculative and beyond the possibilities of this study to resolve this case. To clarify the evolutionary history of these two populations, in-depth molecular studies are required.

Comparative material. Garra r. gymnothorax : ZIN 13214, 7 syntypes, 39–57 mm SL; Iran: Kulikhan in Karun River drainage, H. Zarudny, 6 June 1904.

Garra cf. gymnothorax : FSJF 2198, 19, 40–94 mm SL; Iran: Kohkeloyeh va Boyrahmad prov.: Beshar River 20 km northeast of Yasooj, 30°44'9.12"N 51°29'31.32"E.— FSJF 2209, 9, 66–129 mm SL; Iran: Chaharmahal Bakhtiari prov.: Stream Sangan at Sangan, 31°15'41.52"N 51°17'9.00"E.

Garra persica : FSJF 2218, 20, 50–83 mm SL; Iran: Hormuzgan prov.: River Rodan about 6 km northeast of Rodan, 27°28'53.70"N 57°15'53.70"E.— FSJF 397, 20, 35–50 mm SL; Iran: Sistan & Baluchestan prov.: Karvander River at bridge north of Karvander, 27°51'18''N 60°46'03''E.—ZM-CBSU 10200, 30, 34–56 mm SL; Iran: Fars prov.: Rasool River at God e Gaz village, 27°17'28.8"N 54°29'20.7"E.

Garra rufa: Qweik drainage: NMW 53240, lectotype (101 mm SL) and 7 paralectotypes of D. rufus , 59–101 mm SL; Syria: Aleppo; T. Kotschy, 1842.— NMW 53238, syntypes of D. obtusus , 2, 43– 134 mm SL; Syria: Aleppo; T. Kotschy, 1842. Euphrates and Tigris drainage: NMW 53257, 6 syntypes of D. obtusus , 31–104 mm SL; Iraq: Mossul; T. Kotschy, 1842.— NMW 53236–37, syntypes of D. crenulatus , 4, 56– 74 mm SL; Iran: Qarah Aqaj River.— FSJF 2238, 22, 55–101 mm SL; Iran: Fars prov.: Spring Pirbanoo about 10 km south of Shiraz, 29°31'8.10"N 52°27'55.98"E.— FSJF 2549, 8, 43–82 mm SL; Turkey: Adıyaman prov.: stream Egri south of Adıyaman, a tributary to Atatürk reservoir, 37°44'30.00"N 38°20'6.48"E.— FSJF 2588, 5, 58–65 mm SL; Turkey: Adıyaman prov.: steam Çakal, 13 km west of Adıyaman, tributary to Atatürk reservoir, 37°43'20.53"N 38° 9'55.27"E.— FSJF 2621, 2, 60–67 mm SL; Turkey: Diyabakır prov.: Tigris south of Diyabakır, 37°53'13.80"N 40°13'47.28"E.— FSJF 2849, 7, 71–105 mm SL; Turkey: Diyarbakır prov.: stream Ambar at road to Silvan, 25 km east of Diyarbakır, 37°59'24.72"N 40°22'56.64"E.— FSJF 2864, 3, 98–129 mm SL; Turkey: Batman prov.: Tigris 5 km west of Hasankeyf, 37°43'25.68"N 41°21'37.80"E.— FSJF 2909, 1, 91 mm SL; Turkey: Sivas prov.: stream Kangal under railway bridge at Çetinkaya, 39°15'5.76"N 37°37'8.04"E.— FSJF 2955, 1, 62 mm SL; Turkey: Diyarbakır prov.: stream Savur between Bayındır and Ahmetli east of Tepe, 37°45'49.32"N 40°53'2.04"E.— FSJF 2999, 5, 60–94 mm SL; Turkey: Gaziantep prov.: stream Merziman south of Yavuzeli, 37°17'32.64"N 37°43'23.16"E.— FSJF 3371, 2, 92–100 mm SL; Iraq: stream Suraw near Suraw village, 35°45'45.60"N 45°59'5.40"E.— FSJF 3368, 18, 52–103 mm SL; Iraq: Nalparez River, 35°34'14.40"N 45°51'46.80"E.—ZM-CBSU J 2083, 25, 61–135 mm SL; Iran: Illam prov.: Seimareh River, 33°39'40.3 "N 47°03'29.2 "E.—ZM-CBSU J 2006, 10, 44–78 mm SL; Iran: Kermanshah prov.: Sirvan River, 34°49'37.9 "N 46°21'30 "E.—ZM-CBSU J 1051, 12, 60– 92 mm SL; Iran: Khuzestan prov.: Zohreh River, 30°13'58.56 "N 50°26'50.52 "E. —ZM-CBSU J 2237, 6, 50–89 mm SL; Iran: Kermanshah prov.: Sirvan River, 35°06'43.01 "N 46°15'23.7"E.—ZM-CBSU J 2501, 50, 32–92 mm SL; Iran: Illam prov.: Seimareh River, 33°41'38.1"N 46°42'57.4"E.—ZM-CBSU J2404, 38, 29-92 mm SL; Iran: Illam prov.: Karkheh River, 33°54'26.7"N 46°05'22.8"E.

Garra sp.: FSJF 2255, 22, 52–75 mm SL; ZM-CBSU E812, 87 mm SL; ZM-CBSU E809, 3, 92–125 mm SL; ZM-CBSU E813, 6, 57–84 mm SL; Iran: Fars prov.: Golabi spring about 35 km west of Darab, 28°47'18''N 54°22'15''E.—ZM-CBSU H1056, 20, 62–92 mm SL; Iran: Fars prov.: Rudbal River at Estahban, 28°58'23''N 54°23'25''E; 31 July 2011.—ZM-CBSU H1076, 12, 37–57 mm SL; ZM-CBSU B110, 21, 37–71 mm SL; Iran: Hormuzgan prov.: Haji Abad River about 12 km south-west of Haji Abad, 28°13’33''N 55°49'25''E.—ZM-CBSU H1088, 5, 37–59 mm SL; Iran: Fars prov.: Fadami stream at north of Fadami, 28°13'40''N 55°08'18''E.

Material used in the molecular genetic analysis. Garra barreimiae : FSJF DNA-2488; UAE: Wadi Shawkah, in Emirate of Ras Al-Khaimah, 25°05'54"N 56°06'33"E (GenBank accession numbers: KM214685 , KM214738 , KM214758 , KM214779 ).— FSJF DNA-2489; UAE: Wadi Wurayah in Emirate of Al-Fujaira, 25°23'54"N 56°16'10"E (GenBank accession numbers: KM214734 , KM214767 , KM214783 ).

Garra cf. gymnothorax : ZM-CBSU M845, M846, M848; Iran: Bashar River, 31°02'25.7"N 51°13'03.6"E (GenBank accession numbers: KM214803 , KM214735 , KM214763 ).—ZM-CBSU M837; Iran: Bashar River, 30°40'50.94"N 51°32'20.72"E (GenBank accession number: KM373234 ).

Garra cf. longipinnis : FSJF DNA-2493; Oman: Al-Saiq Platue at Jabal al Akhdar, 23°02'00"N 57°28'00"E (GenBank accession numbers: KM214752 , KM214756 ).

Garra ghorensis : FSJF DNA-1193; Jordan: Spring at Ghor al Hadithah, 31°17'47"N 35°32'33"E (GenBank accession numbers: KJ553478 , KJ553508 , KJ553520 ).— FSJF DNA-1225; Jordan: stream below Afra hot spring, 30°57'56"N 35°40'56"E (GenBank accession number: KJ553401 ).— FSJF DNA-2495; Jordan: Wadi Burbaita at the upper part of Wadi Al-Hassa, 30°59'01"N 35°40'11"E (GenBank accession numbers: KM214688 , KM214728 , KM214788 ).— FSJF DNA-2496; Jordan: Wadi Fifa in Southern Ghor, 30°55'52"N 35°28'52"E (GenBank accession number: KM214722 ).— FSJF DNA-2497; Jordan: Wadi Ibn-Hammad in Al-Karak Provience, 31°18'05"N 35°37'47"E (GenBank accession number: KM214798 ).

Garra jordanica : FSJF DNA-1186; Jordan: Spring in Suweyma, 31°46'14"N 35°36'10"E (GenBank accession numbers: KM214710 , KM214750 ).— FSJF DNA-1206; Syria: River Yarmuk at in Wadi Jallayn, 32°44'21"N 35°58'56"E (GenBank accession number: KJ553525 ).— FSJF DNA-2498; Jordan: lower most Mujib River, 31°27'22"N 35°35'08"E (GenBank accession numbers: KM214697 , KM214703 , KM214753 , KM214806 ).

Garra persica : ZM-CBSU M1135, M995; Iran: Rudan River, 27°45'49.4"N 57°14'32.8"E (GenBank accession number: KT808682 , KM373235 ).–ZMCBSU M700; Iran: Shur River, 27°19'40.58"N 56°33'10.79"E (GenBank accession number: KM214706 ). –ZM-CBSU M701; Iran: Kol River, 27°17'28.8"N 54°29'20.7"E (GenBank accession number: KM214807 ).

Garra rossica : ZM-CBSU M318; Iran: Sistan basin, 34°44'10.1"N 60°34'29.7"E (GenBank accession number: KM214771 ).—ZM-CBSU M792, M298; Iran: Karvandar River, 27°51'17.97"N 60°46'2.99"E (GenBank accession numbers: KM214796 , KM214770 ).—ZM-CBSU M367; Iran: Kavir basin, 32°52'46.74"N 59°3'17.39"E (GenBank accession number: KM214759 ).

Garra rufa : FSJF DNA-2414; Turkey: Balıklı kaplıca 10 km northeast of Kangal, 39°18'49.07"N 37°28'10.06"E (GenBank accession numbers: KM214792 - KM214794 ).— FSJF DNA-923; Turkey: stream Çakal, 13 km west of Adıyaman, tributary to Atatürk reservoir, 37°43'20.36"N 38°9'55.34"E (GenBank accession numbers: KM214692 , KM214695 , KM214800 ).— FSJF DNA-2234; Iraq: Tabin river south of Zarbi, 35°48'19.20"N 45°1'12.00"E.— FSJF DNA-2480; Iraq: Shatt al-Arab at Basrah, 30°32'22"N 47°49"E (GenBank accession number: KM214777 ).— FSJF DNA-918; Turkey: River Tigris south of Diyabakır, 37°53'13.80"N 40°13'47.40"E (GenBank accession number: KM214684 ). –ZM-CBSU M849, M850; Iran: Dalaki River, 29°28'15.40"N 51°18'14.60"E (GenBank accession: KT808680 , KT808681 ).—ZM-CBSU M704, M836; Iran: Koohmareh Sorkhi River at Helleh, 29°23'39.80"N 52°9'40.20"E (GenBank accession numbers: KM214760 , KM214810 ).—ZM-CBSU M847; Iran: Bashar River at Yasouj, 31° 2'25.68"N 51°13'3.60"E (GenBank accession number: KM214711 ).—ZM-CBSU M745; Iran: Doirej River at Ilam, 32°39'13.13"N 47°32'36.97"E (GenBank accession number: KM214733 ).—ZM-CBSU M713; Iran: Kavar River at Mond, 29°10'54.60"N 52°41'28.20"E (GenBank accession number: KM373232 ). –ZM-CBSU M842; Iran: stream at Saadi Tomb, 29°37'20.77"N 52°34'55.59"E (GenBank accession number: KM214714 ).—ZM-CBSU M711; Iran: Fahlian River, 30°11'4.40"N 51°31'25.57"E (GenBank accession number: KM214709 ).

Garra sp.: ZM-CBSU M299, M301, M302; Fars prov.: Golabi spring about 35 km west of Darab, 28°47'18''N 54°22'15''E (GenBank accession numbers: KM214691 , KM214782 , KM214791 ).

Garra typhlops : FSJF DNA-2169; Iran: Well at Kaaje-Ru, 33°04'39"N 48°35'33"E (GenBank accession numbers: KM214717 , KM214731 ).

Garra variabilis : FSJF DNA-1159; Syria: Nahr al Barid at Nahr al Barid, 35°18'08"N 36°20'43"E (GenBank accession numbers: KJ553391 , KJ553567 )— FSJF DNA-1168; Syria: Orontes at Shayzar, 35°16'18"N 36°33'46"E (GenBank accession numbers: KJ553422 , KJ553524 ).

Garra widdowsoni : FSJF DNA-2301; Iraq: cavern 25 feet below ground, 6 miles north of Haditha, 34° 4'0.00"N 42°23'60.00"E (GenBank accession numbers: KM214769 , KM214795 ).

Figures

FIGURE 1. Maximum Likelihood (ML) estimation of the phylogenetic relationships of Garra species having a mental disc form the Middle East based on the mitochondrial COI barcode region. Nucleotide positions with less than 95 % site coverage were eliminated before analysis. Numbers of major nodes indicate bootstrap values from the Maximum Likelihood-method from 1000 pseudo-replicates, followed by Bayesian posterior probabilities.

FIGURE 2. Garra mondica, ZM-CBSU H 1032, holotype, 66 mm SL; Iran: spring Konar Siyah.

FIGURE 3. Garra mondica, ZM-CBSU K 1080, paratypes, a, 58 mm SL, b, 51 mm SL, c, 46 mm SL; Iran: spring Konar Siyah.

FIGURE 4. Garra mondica, ZM-CBSU K 1080, paratypes, a, 58 mm SL, b, 51 mm SL, c, 46 mm SL; Iran: spring Konar Siyah.

Tables

TABLE 1. Diagnostic nucleotide substitutions found in the COI barcode region of Garra species studied.

Nucleotide position	N	5 9 2 8	5 9 3 4	5 9 4 3	5 9 4 6	5 9 7 3	5 9 7 9	5 9 9 1	6 0 0 0	6 0 0 3	6 0 0 6	6 0 0 9	6 0 2 7	6 0 3 3	6 0 3 4	6 0 3 6	6 0 4 0	6 0 5 1	6 0 5 4	6 0 5 8	6 0 6 0	6 0 7 5	6 0 7 8	6 1 1 7	6 1 2 6	6 1 3 2	6 1 4 1	6 1 4 4	6 1 4 7	6 1 5 4
G. barreimiae	7	T	A	T	T	C	C	T	A	C	C	G	C	A	C	A	C	G	A	C	A	A	G	C	A	A	T	T	T	C
G. ghorensis	9	T	G	T	T	C	C	T	A	A	C	G	C	A	C	A	C	G	G	C	A	A	G	C	A	A	T	T	T	C
G. jordanica	7	T	G	T	T	C	C	T	A	C	C	G	C	A	C	A	C	G	A	C	A	A	G	C	A	A	T	T	T	C
G. cf. longipinnis	2	T	G	T	T	C	C	T	A	C	C	G	C	A	T	A	T	G	A	C	G	A	G	C	A	A	T	T	T	C
G. mondica	3	T	G	C	T	C	C	C	A	C	C	G	C	A	C	A	C	A	A	C	A	A	G	C	A	A	T	T	T	C
G. persica	4	T	G	T	T	C	C	T	A	C	C	A	C	A	C	A	C	A	A	C	A	A	G	C	A	A	T	T	T	C
G. rossica	4	T	G	T	T	C	C	T	T	T	C	A	T	G	C	A	C	T	C	T	A	A	A	C	A	T	T	C	C	T
G. r u f a	18	T	G	T	T	C	C	T	A	C	C	G	C	A	C	A	C	A	A	C	A	A	G	C	A	A	T	T	T	C
G. typhlops	2	T	G	T	T	C	C	T	A	C	C	G	C	A	C	A	C	G	T	C	A	A	G	C	A	A	T	T	T	C
G. variabilis	4	T	G	T	C	T	T	T	C	T	T	T	C	T	C	A	C	A	C	C	A	A	G	C	G	C	T	T	T	C
G. widdowsoni	2	C	G	T	T	C	C	T	A	C	C	G	C	A	C	A	C	A	A	C	A	G	G	C	A	A	C	T	T	C
G. cf. gymnothorax	4	T	G	T	T	C	C	T	A	C	C	G	C	A	C	G	C	G	A	C	A	A	G	T	A	A	T	T	T	C
continued.

TABLE 2. Estimates of evolutionary divergence (%) over sequence pairs between species found in the COI barcode region of Garra species studied.

G. r u f a	N 18	rufa . G	mondica . G	Kol. sp. G	persica . G	gymnothorax . cf. G	typhlops . G	widdowsoni . G	jordanica . G	barreimiae . G	longipinnis ghorensis . cf.. G G	a c i s s ro. G
G. mondica	3	3.88
G. sp. Kol	3	3.51	0.73
G. persica	4	2.43	4.13	3.8
G. cf. gymnothorax	4	5.67	5.29	5.8	5.48
G. typhlops	2	5.12	6.31	6.08	4.77	4.49
G. widdowsoni	2	3.17	4.05	3.69	3.8	7.16	6.99
G. jordanica	7	4.15	6.16	5.71	5.26	6.32	6.15	6.39
G. barreimiae	7	6.67	7.46	7.68	6.22	7.39	7.45	7.69	7.3
G. cf. longipinnis	2	7.09	6.97	6.74	6.19	6.68	5.4	8.12	7.73	7.66
G. ghorensis	9	5.57	7.06	6.75	5.98	7.6	6.52	7.45	4.39	8.37	7.42
G. rossica	4	13.6	13.7	13.4	13	14.1	11.5	15	14.7	15.3	13.9 13.69
G. variabilis	2	13.8	16.3	15.7	12.7	14.6	11.9	15.5	13.9	17	13.6 13.52	11.43

TABLE 3. Morphometric data of Garra mondica holotype ZM-CBSU H 1032, paratypes, ZM-CBSU K 1079 - K 1086, ZM-CBSU H 1033 - H 1040, ZM-CBSU B 242, ZM-CBSU B 250, ZM-CBSU B 255, n = 20.

	Holotype	Range	Mean	SD
Standard length (mm)	65.9	38.5–65.9
In percent of standard length
Head length	25.7	24.0–27.4	25.9	1.0
Body depth at dorsal-fin origin	18.5	16.2–23.4	18.5	2.0
Predorsal length	47.5	46.1–50.3	48.2	1.1
Postdorsal length	40.2	37.4–41.9	39.6	1.5
Preanal length	72.0	71.1–74.6	73.1	1.0
Prepelvic length	51.5	49.6–55.3	52.8	1.5
Distance between pectoral and pelvic-fin origins	30.0	28.2–31.6	30.0	1.1
Distance between pelvic and anal-fin origins	19.5	17.4–21.6	19.6	1.1
Depth of caudal peduncle	11.1	10.5–12.4	11.6	0.6
Length of caudal peduncle	20.6	18.8–22.2	20.4	0.8
Dorsal-fin base length	13.6	12.0–15.1	13.2	0.9
Anal-fin base length	8.4	6.9–8.4	7.5	0.4
Pectoral-fin length	24.3	23.0–27.7	24.8	1.4
Pelvic-fin length	20.1	18.1–21.3	19.8	0.9
In percent of head length
Head depth at eye	59	55–66	59.8	4.1
Snout length	47	38–47	43.0	2.6
Postorbital distance	40	36–43	39.5	1.9
Interorbital width	40	37–46	41.8	2.3
Eye diameter	15	13–22	17.8	2.5
Maximum head width	68	66–74	68.6	2.3
Rostral barbel length	18	16–20	17.9	1.5
Maxillary barbel length	13	12–17	14.5	1.1