Research Article |
Corresponding author: Cüneyt Kaya ( cuneyt.kaya@erdogan.edu.tr ) Academic editor: Nicolas Hubert
© 2024 Cüneyt Kaya, Irmak Kurtul, İsmai̇l Aksu, Münevver Oral, Jörg Freyhof.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Kaya C, Kurtul I, Aksu İ, Oral M, Freyhof J (2024) Oxynoemacheilus chaboras, a new loach species from the Euphrates drainage in Türkiye (Teleostei, Nemacheilidae). Zoosystematics and Evolution 100(2): 457-468. https://doi.org/10.3897/zse.100.118612
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Oxynoemacheilus chaboras, new species, from the stream Beyazsu in the Euphrates drainage, belongs to the O. persa species group, being closely related to O. shehabi from the Orontes, O. sarus from the Seyhan and Ceyhan, O. euphraticus from the Euphrates and Tigris, O. karunensis from the Karkheh, and O. persa from Central Iran. The new species is distinguished from others in the O. persa group by having 8–9 pores in the supraorbital canal, two distinct black blotches at the caudal-fin base, a rudimentary and shallow pelvic axillary lobe, 6–10 irregularly shaped bars on the flank, and a deep head, body, and caudal peduncle. Oxynoemacheilus chaboras sp. nov. is most closely related to O. euphraticus, from which it is differentiated by a mean uncorrected p-distance of 3.24% (min. 3.09%) in its COI barcode gene.
Cypriniformes, Cytochrome c oxidase subunit I, freshwater fish, taxonomy, Western Asia
The genus Oxynoemacheilus Bănărescu & Nalbant, 1966, with 63 recognised species, is the most speciose genus of freshwater fishes in the western Palaearctic (
There are many tributaries to the upper and middle Euphrates. One of these rivers is the Khabur that has few springs in Türkiye, but mostly flows in Syria. The stream Beyazsu, located in the Turkish Mardin province, is one of the headwater streams in the Khabour drainage. It flows into Syria after crossing the border at the city of Nusaybin, only 17.5 km below its source, the spring Beyazsu (
Until now, only
The care of experimental animals was consistent with the Republic of Türkiye’s animal welfare laws, guidelines, and policies. After anaesthesia, fishes were fixed in 5% formalin and stored in 70% ethanol, fin clips directly fixed in 99% ethanol. Measurements were made with a dial calliper, recorded to 0.1 mm, from a precise point-to-point approach, never by projections. Methods for counts and measurements followed
Morphological data for Oxynoemacheilus zagrosensis Kamangar, Prokofiev, Ghaderi & Nalbant, 2014 are taken from
SL, standard length;
HL, head length;
Collection codes:
FFR, Recep Tayyip Erdogan University Zoology Collection of the Faculty of Fisheries, Rize;
FSJF, Fischsammlung J. Freyhof, Berlin, Germany.
Genomic DNA extraction of Oxynoemacheilus specimens was performed according to the application protocol recommended by the manufacturer using the DNeasy Blood & Tissue Kit (Qiagen, Hilden, Germany). Amplification of the barcode region of the cytochrome c oxidase subunit 1 (COI) gene of vertebrate mitochondrial DNA was performed according to
Oxynoemacheilus species distributed in the Euphrates and all other species, except O. zagrosensis, of the O. persa species group, as well as all other species known from the Euphrates drainage, were included in our dataset (Fig.
Maximum Likelihood (ML) phylogenetic tree was reconstructed based on the COI-Barcode gene. ML and BI methods resulted in generally similar topologies with minor differences, and therefore only the ML tree is shown. The bootstrap values of ML and posterior probability values of BI are indicated on nodes (ML/BI). The bootstrap percentage values (BP) ≥ 50% from ML analysis and Bayesian posterior probabilities (PP) ≥ 0.50 are shown on the nodes.
Phylogenetic relationships among Oxynoemacheilus species were estimated using Maximum Likelihood (ML;
Molecular analysis was conducted with four newly-generated DNA barcodes (see Genetic material section) and in addition already published data from NCBI GenBank. The average nucleotide frequency of four sequences of O. chaboras were A = 22.0%, T = 30.2%, C = 28.2% and G = 19.7%, and the nucleotide composition was A-T (52.2%) rich. Although the phylogenetic tree topologies reconstructed by both ML and BI methods indicated some minor differences from each other, they were generally compatible. In both topologies, some of the internal branches corresponding to the phylogenetic relationships between species, were weakly supported (Fig.
The members of the O. persa species group (as defined by
The interspecies genetic distances calculated by the uncorrected p-distance model for the Oxynoemacheilus species of Euphrates-Tigris and other O. persa species group.
Species | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 | |
1 | O. chaboras | ||||||||||
2 | O. euphraticus | 0.0324 | |||||||||
3 | O. shehabi | 0.0382 | 0.0300 | ||||||||
4 | O. persa | 0.0382 | 0.0265 | 0.0423 | |||||||
5 | O. marunensis | 0.0455 | 0.0340 | 0.0431 | 0.0431 | ||||||
6 | O. hanae | 0.0455 | 0.0349 | 0.0374 | 0.0439 | 0.0285 | |||||
7 | O. kurdistanicus | 0.0463 | 0.0366 | 0.0455 | 0.0512 | 0.0268 | 0.0301 | ||||
8 | O. kentritensis | 0.0472 | 0.0356 | 0.0415 | 0.0480 | 0.0390 | 0.0366 | 0.0415 | |||
9 | O. zagrosensis | 0.0488 | 0.0323 | 0.0480 | 0.0480 | 0.0260 | 0.0268 | 0.0220 | 0.0374 | ||
10 | O. sarus | 0.0488 | 0.0358 | 0.0415 | 0.0431 | 0.0520 | 0.0480 | 0.0545 | 0.0504 | 0.0537 | |
11 | O. argyrogramma | 0.0507 | 0.0406 | 0.0509 | 0.0472 | 0.0388 | 0.0390 | 0.0388 | 0.0480 | 0.0363 | 0.0604 |
12 | O. zarzianus | 0.0512 | 0.0395 | 0.0488 | 0.0423 | 0.0325 | 0.0341 | 0.0390 | 0.0415 | 0.0350 | 0.0545 |
13 | O. chomanicus | 0.0512 | 0.0332 | 0.0455 | 0.0488 | 0.0268 | 0.0309 | 0.0260 | 0.0415 | 0.0187 | 0.0545 |
14 | O. karunensis | 0.0520 | 0.0414 | 0.0488 | 0.0374 | 0.0593 | 0.0537 | 0.0577 | 0.0577 | 0.0569 | 0.0472 |
15 | O. araxensis | 0.0772 | 0.0706 | 0.0764 | 0.0732 | 0.0854 | 0.0829 | 0.0837 | 0.0724 | 0.0854 | 0.0732 |
16 | O. kaynaki | 0.0789 | 0.0683 | 0.0715 | 0.0764 | 0.0772 | 0.0780 | 0.0780 | 0.0756 | 0.0732 | 0.0894 |
17 | O. arsaniasus | 0.0821 | 0.0716 | 0.0715 | 0.0780 | 0.0756 | 0.0764 | 0.0748 | 0.0740 | 0.0691 | 0.0854 |
18 | O. tigris | 0.0821 | 0.0688 | 0.0699 | 0.0748 | 0.0789 | 0.0780 | 0.0846 | 0.0724 | 0.0780 | 0.0813 |
19 | O. muefiti | 0.0894 | 0.0796 | 0.0821 | 0.0846 | 0.0862 | 0.0854 | 0.0878 | 0.0813 | 0.0756 | 0.0967 |
20 | O. paucilepis | 0.1024 | 0.0948 | 0.0878 | 0.0976 | 0.0951 | 0.0959 | 0.0935 | 0.0870 | 0.0927 | 0.0967 |
21 | O. bergianus | 0.1122 | 0.1156 | 0.1220 | 0.1122 | 0.1228 | 0.1171 | 0.1146 | 0.1098 | 0.1146 | 0.1179 |
11 | 12 | 13 | 14 | 15 | 16 | 17 | 18 | 19 | 20 | ||
1 | O. chaboras | ||||||||||
2 | O. euphraticus | ||||||||||
3 | O. shehabi | ||||||||||
4 | O. persa | ||||||||||
5 | O. marunensis | ||||||||||
6 | O. hanae | ||||||||||
7 | O. kurdistanicus | ||||||||||
8 | O. kentritensis | ||||||||||
9 | O. zagrosensis | ||||||||||
10 | O. sarus | ||||||||||
11 | O. argyrogramma | ||||||||||
12 | O. zarzianus | 0.0455 | |||||||||
13 | O. chomanicus | 0.0390 | 0.0358 | ||||||||
14 | O. karunensis | 0.0515 | 0.0650 | 0.0537 | |||||||
15 | O. araxensis | 0.0808 | 0.0748 | 0.0862 | 0.0862 | ||||||
16 | O. kaynaki | 0.0705 | 0.0748 | 0.0683 | 0.0902 | 0.0878 | |||||
17 | O. arsaniasus | 0.0721 | 0.0732 | 0.0683 | 0.0902 | 0.0846 | 0.0228 | ||||
18 | O. tigris | 0.0770 | 0.0732 | 0.0715 | 0.0894 | 0.0862 | 0.0423 | 0.0488 | |||
19 | O. muefiti | 0.0851 | 0.0813 | 0.0829 | 0.1024 | 0.0967 | 0.0220 | 0.0358 | 0.0512 | ||
20 | O. paucilepis | 0.0981 | 0.0992 | 0.0943 | 0.1008 | 0.0878 | 0.0992 | 0.0959 | 0.0943 | 0.1033 | |
21 | O. bergianus | 0.1187 | 0.1203 | 0.1106 | 0.1163 | 0.1057 | 0.1098 | 0.1114 | 0.1203 | 0.1154 | 0.1008 |
Holotype. FFR 15646, 53 mm SL; Türkiye: Mardin prov.: stream Beyazsu 14 km north of Nusaybin, 37.1989, 41.3076.
Paratypes. FFR 1428, 11, 46–60 mm SL: same data as holotype. — FFR 15633, 2, 40–51 mm SL; FSJF 4116, 4, 46–55; Türkiye: Mardin prov.: stream Beyazsu 12 km north of Nusaybin, 37.1730, 41.2690.
Oxynoemacheilus araxensis, O. argyrogramma, Oxynoemacheilus arsaniasus Freyhof, Kaya, Turan & Geiger, 2019, Oxynoemacheilus bergianus (Derjavin, 1934), O. euphraticus, Oxynoemacheilus kaynaki Erk’akan, Özeren & Nalbant, 2008, Oxynoemacheilus muefiti Freyhof, Kaya, Turan & Geiger, 2019, Oxynoemacheilus paucilepis (Erk’akan, Nalbant & Özeren, 2007), and Oxynoemacheilus tigris (Heckel, 1843) are other species of Oxynoemacheilus known from the Euphrates drainage (Fig.
Oxynoemacheilus chaboras belongs to a group of species (O. argyrogramma, O. chaboras, O. euphraticus) having two bold, black, round or comma-shaped blotches on the caudal-fin base (vs. absent in Oxynoemacheilus araxensis, O. arsaniasus, O. bergianus, O. kaynaki, O. muefiti, O. paucilepis, and O. tigris). Furthermore, male O. chaboras have a suborbital groove (as in O. araxensis and O. bergianus vs. absent in O. arsaniasus, O. kaynaki, O. muefiti, O. paucilepis, and O. tigris).
Oxynoemacheilus chaboras is further distinguished from O. araxensis by having a forked caudal fin (vs. slightly emarginate), and it is further distinguished from O. bergianus by having a forked caudal fin (shortest middle caudal-fin ray is 57–70% of longest ray of the upper caudal-fin lobe, vs. deeply emarginated, 70–84), and a deeper caudal peduncle (depth 1.4–1.7 times in its length vs. 1.7–3.5).
Oxynoemacheilus chaboras is distinguished from O. argyrogramma and O. euphraticus by possessing a mid-lateral series of blotches (vs. marbled or mottled pattern in O. argyrogramma), without a mottling pattern above or below the blotches in front of dorsal-fin base (vs. irregularly mottled or marbled in O. euphraticus), and having no, or a very short, incision in the upper lip (vs. a deep median incision in O. euphraticus). It is further distinguished from O. euphraticus by having a deeper caudal peduncle (caudal-peduncle depth 1.4–1.7 times in its length vs. 2.0–2.8).
See Figs
Morphometric data of Oxynoemacheilus chaboras (holotype FFR 15646 and paratypes FFR 1428, n = 11).
Holotype | min | max | mean | SD | |
---|---|---|---|---|---|
Standard length (mm) | 53 | 46 | 60 | 51.9 | 4.0 |
In percent of standard length | |||||
Head length | 24.2 | 23.0 | 25.0 | 24.1 | 0.6 |
Body depth at dorsal-fin origin | 18.4 | 17.4 | 21.8 | 18.8 | 1.2 |
Predorsal length | 50.9 | 47.4 | 51.1 | 49.7 | 1.2 |
Postdorsal length | 35.8 | 33.8 | 37.4 | 35.5 | 1.2 |
Prepelvic length | 51.2 | 49.6 | 55.5 | 52.1 | 1.2 |
Preanal length | 72.7 | 72.7 | 78.7 | 75.7 | 1.8 |
Distance between pectoral and pelvic-fin origins | 30.1 | 26.2 | 30.6 | 28.4 | 1.6 |
Distance between pelvic and anal-fin origins | 21.4 | 21.4 | 25.2 | 22.8 | 1.0 |
Distance between vent and anal-fin origin | 3.0 | 2.1 | 3.7 | 2.9 | 0.5 |
Dorsal-fin height | 22.7 | 19.9 | 24.3 | 22.4 | 1.2 |
Anal-fin height | 18.9 | 15.8 | 19.2 | 17.9 | 1.0 |
Pectoral-fin length | 24.1 | 21.8 | 26.7 | 24.2 | 1.7 |
Pelvic-fin length | 18.7 | 16.5 | 19.3 | 18.1 | 0.8 |
Length of caudal peduncle | 19.8 | 16.9 | 19.8 | 18.2 | 0.9 |
Depth of caudal peduncle | 11.9 | 10.2 | 12.2 | 11.6 | 0.5 |
In percent of head length | |||||
Head depth at eye | 51 | 51 | 60 | 53.9 | 3.3 |
Maximum head width | 59 | 59 | 66 | 63.2 | 2.6 |
Snout length | 40 | 39 | 44 | 41.4 | 1.8 |
Eye diameter | 19 | 19 | 23 | 20.6 | 1.4 |
Postorbital distance | 45 | 40 | 50 | 45.2 | 2.7 |
Interorbital width | 21 | 18 | 24 | 21.6 | 2.3 |
Length of inner rostral barbel | 30 | 20 | 30 | 24.7 | 2.5 |
Length of outer rostral barbel | 41 | 34 | 44 | 38.5 | 3.2 |
Length of maxillary barbel | 40 | 33 | 44 | 39.6 | 3.3 |
Oxynoemacheilus species in the Euphrates drainages: a. O. bergianus, stream Sason, 61 mm SL; b. O. argyrogramma, stream Sünnep, 50 mm SL; c. O. euphraticus, Great Zap River, 55 mm SL; d. O. muefiti, Murat River, 69 mm SL; e. O. tigris, stream Sünnep, 55 mm SL; f. O. kaynaki, Göksu River, 68 mm SL; g. O. paucilepis, stream Balıklıtohma, 70 mm SL; h. O. arsaniasus, stream Kaleli, 90 mm SL; i. O. araxensis, stream Arkaçayırlar, 71 mm SL.
Dorsal fin with 9½–10½ branched rays, outer margin straight or slightly concave. Anal fin with 5½ branched rays, outer margin straight. Pectoral fin with 9–11 branched rays, outer margin straight or slightly convex, tip pointed in male. Pelvic fin with 6 branched rays, outer margin straight or slightly convex. Caudal fin forked with (8+8)9+8 branched rays, lobes pointed. Flank and back covered by cycloid scales. Chest and belly without scales. Lateral line complete, terminating between origin of hypural complex and caudal fin base. Anterior nostril opening at end of a low, ovoid, flap-like tube. Posterior tip of anterior nostril overlapping posterior nostril when folded backwards. One central pore and one lateral pore on each side of supratemporal head canal, 3(4) + 9–10 pores in infraorbital canal, 8–9 pores in supraorbital canal, and 9–10 pores in mandibular canal. A suborbital groove in male. Mouth small, arched. Lips thick without furrows, lower lip thicker than upper lip. A median interruption in lower lip. Upper lip without median incision, rarely with a very small and short median incision. Processus dentiformis narrow and rounded. Lower jaw rounded, without median notch. Barbels long; inner rostral barbel reaching base of maxillary barbel, outer reaching to vertical of posterior of anterior eye margin. Maxillary barbel reaching or almost reaching to vertical of posterior eye-margin.
Body with yellowish or cream background and dark-brown pattern in live and preserved individuals. Preserved individuals with a dark-grey, narrow inner-axial stripe, absent in life. Dorsal head and upper part of cheek brown, with marbled pattern. Ventral surface of head yellowish without pattern. Flank with 6–10 dark-brown bars or blotches, as much as, or thicker than, interspaces. Bars and blotches irregularly shaped and set, generally vertically elongated, sometimes oval, or horizontally elongated, usually extending to mid-dorsal saddles and meeting contra laterals. Back with 1–3 predorsal saddles, one saddle at dorsal-fin origin and one at posterior dorsal-fin base, and 3 saddles behind dorsal fin, as much as or thicker than interspaces. One dark-brown or black blotch at lower caudal-fin base, a second, much smaller blotch at uppermost caudal-fin base, both distinct in both live and preserved individuals. Dorsal, caudal and pectoral fins with many, small brown blotches on rays. These blotches forming 2–3 narrow bands on dorsal, and 3–5 on caudal. Pectoral, anal and pelvic fins hyaline, sometimes with a few dark-brown blotches on rays.
The species is named Chaboras, an ancient Greek name of the Khabur (Χαβώρας), as it was first documented by Ptolemy and Pliny the Elder ichthyofauna. A noun in genitive, indeclinable.
Following our molecular analysis, Oxynoemacheilus chaboras belongs to the O. persa species group as defined by
Oxynoemacheilus chaboras is most closely related to O. shehabi from the upper Orontes, O. sarus from the Seyhan and Ceyhan, O. euphraticus from the Euphrates and Tigris, O. karunensis from the Karkheh, and O. persa from Central Iran. Oxynoemacheilus argyrogramma, O. hanae, O. kurdistanicus, O. marunensis are placed in a second cluster of species within the O. persa species group and are not closely related. While all these species are well-supported in our molecular analysis (Table
Oxynoemacheilus chaboras is distinguished from O. shehabi and O. sarus by possessing 8–9 pores in the supraorbital canal (vs. 5–7), a rudimentary and shallow pelvic axillary lobe fully attached to the body (vs. well-developed with a free tip), a deeper body (body depth at dorsal fin origin 17–22% SL vs. 14–17 in O. shehabi), deeper caudal peduncle (10–12% SL vs. 8–9 in O. shehabi), deeper head (head depth at eye 51–60% HL vs. 44–51 in O. sarus) and a longer anal fin (anal-fin height 18–22% SL vs. 16–19 in O. sarus). It is distinguished from O. hanae by lacking isolated patches of dark-brown spots or blotches on the lower part of the flank (vs. present) and possessing two distinct black blotches at the caudal-fin base (vs. usually absent or very small, overlaid by a chevron shaped bar).
The new species is distinguished from O. karunensis and O. persa by lacking the dense mottling in the interspaces between the blotches on the flank in almost all individuals (vs. very dense mottling in all individuals), possessing a deeper caudal peduncle (caudal peduncle depth 1.4–1.7 times in its length vs. 1.7–3.1 in O. karunensis), and two distinct black blotches at the caudal-fin base (vs. two very large blotches, usually fused to an irregularly shaped bar in O. persa).
We were not able to compare O. chaboras to O. marunensis as we had no materials available. We noted that the description of this species by
Oxynoemacheilus chaboras is distinguished from O. kurdistanicus by possessing no, or rarely, a very short incision in the upper lip (vs. usually a deep, rarely a shallow median incision), and a series of mid-lateral blotches disconnected from the saddles on the back below the dorsal-fin origin (vs. bars connected to saddles in most, but not all individuals). All O. kurdistanicus examined have a pattern of bars on flank, while most O. chaboras have a series of mid-lateral blotches usually narrowly connected to saddles. In O. chaboras, this pattern is usually formed by two wide and dark elements (blotch and saddle) connected by a narrower and paler field of pigments while in O. kurdistanicus (and O. euphraticus and O. marunensis), bars are usually (not always) regularly shaped and not wider along the lateral midline.
Oxynoemacheilus araxensis ZMH 4827, holotype, 61 mm SL; ZMH 4826, paratypes, 5, 36–50 mm SL; ZMH 5951, paratypes, 4, 44–64 mm SL; Türkiye: Erzurum prov.: Kandili Karasu, Euphrates drainage.—FFR 1354, 11, 66–90 mm SL Türkiye: Erzurum prov.: stream Sırlı at Ilıca 40.2130°N, 41.0699°E.—FFR 1451, 2, 68–75 mm SL; Türkiye: Erzurum prov.: stream Ağarcık at Ilıca, 40.2460°N, 41.0710°E.—FFR 1468, 12, 53–70 mm SL; Türkiye: Erzurum prov.: stream Baş about 1 km west of Çayköy, 39.9470°N, 40.8040°E.—FSJF 3440, 6, 42–71 mm SL; Türkiye: Erzurum prov.: stream Arkaçayırlar at Paşayurdu, 39.9833°N, 40.9920°E.
Oxynoemacheilus argyrogramma FFR 15516, 26, 37–49 mm SL; Türkiye: Kilis prov.: stream Sünnep at northeastern Küplüce, 36.7640°N, 37.2540°E.—FFR 1574, 14, 41–62 mm; FFR 1448, 11, 37–48 mm SL; Türkiye: Gaziantep prov.: stream Merziman about 3 km south of Yavuzeli, 37.2910°N, 37.5730°E.
Oxynoemacheilus arsaniasus FFR 15531, paratypes, 5, 36–54 mm SL; Türkiye: Muş prov.: stream Kaynarca at Tepe, 39.1070°N, 41.4920°E.—FFR 1449, 1, 49 mm SL; Türkiye: Muş prov.: stream Kaynarca about 3 km southeast Tepe, 39.0680N, 41.5290°E.—FSJF 4019, 12, 46–97 mm SL; Türkiye: Bitlis prov.: Reservoir of stream Karasu in Kaleli, 38.5537°N, 42.0257°E.
Oxynoemacheilus bergianus FFR 1577, 19, 54–62 mm SL; Türkiye: Samsun prov.: stream Soruk 20 km east of Vezirköprü, 41.1189°N, 35.2269°E.—FFR 15561, 9, 35–69 mm SL; Türkiye: Kayseri prov.: stream Sarnaz a drainage of stream Zamantı at Taşçı, 38.1953°N, 35.7805°E.—FSJF 2983, 15, 38–77 mm SL; Türkiye: Kayseri prov.: stream Zamantı at Pınarbaşı, 38.7366°N, 36.4131°E. —FFR 1457, 11, 64 –72 mm SL; Türkiye: Malatya prov.: stream Sultansuyu 8 km east of Akçadağ, 38.3388°N, 38.0620°E.—FFR 1467, 28, 54–64 mm SL; Türkiye: Erzurum prov.: stream Baş 10 km east of Aşkale, 39.9478°N, 40.8040°E.—FFR 15506, 25, 33–59 mm SL; Türkiye: Ağrı prov.: Murat River 17 km west of Taşlıçay, 39.6785°N, 43.1887°E.
Oxynoemacheilus chomanicus FSJF 3644, 5, 33–61 mm SL; Iraq: Choman River at Alut, 35.9563°N, 45.6155°E.
Oxynoemacheilus euphraticus FFR 1434, 1, 56 mm SL; Türkiye: Sivas prov.: Euphrates at İliç, 39.4850°N, 38.5850°E.—FFR 1471, 25, 27–63 mm SL; Türkiye: Sivas prov.: stream Kangal about 1 km west of Çetinkaya, 39.2560°N, 37.6250°E.—FFR15520, 14, 41–57 mm SL; Malatya prov.: stream Sultan Suyu about 7 km east of Akçadağ, 38.3390°N, 38.0620°E.—FFR 15508, 13, 53–70 mm SL; Türkiye: Adıyaman prov.: stream Göksu at Düzbağ, 37.7950°N, 37.4710°E.
Own data: 39.9124°N, 40.8540°E. 39.9094°N, 40.8028°E. 36.943333°N, 44.19533°E. 39.2516°N, 37.6189°E. 38.9500°N, 40.0166°E. 38.9166°N, 41.2666°E. 39.2515°N, 37.6189°E. 39.2515°N, 37.6189°E. 39.2515°N, 37.61894°E. 39.2515°N, 37.6189°E. 38.9167°N, 41.2667°E. 39.3013°N, 37.6743°E. 37.8370°N, 37.6848°E. 39.1439°N, 37.2571°E. 39.2516°N, 37.6189°E. 36.9433°N, 44.1953°E. 37.7950°N, 37.4705°E. 37.8444°N, 37.6702°E. 37.6500°N, 39.0500°E. 37.0666°N, 37.9500°E. 39.9833°N, 40.9920°E. 36.7484°N, 44.2997°E. 36.9433°N, 44.1953°E. 36.6164°N, 44.8781°E. 36.6106°N, 44.8381°E.
Oxynoemacheilus hanae
Oxynoemacheilus karunensis FSJF 3525, 8, 33–55 mm SL; Iran: Hamadan prov.: Gamasiab River at Do Ab, 47.9167°N, 34.3724°E.—FSJF 3523, 6, 34–51 mm SL; Iran: Hamadan prov.: Haram Abad River at Ashmizan, 34.1105°N, 48.8704°E.—FSJF 3524, 7, 37–53 mm SL; Hamadan prov.: Dehno stream about 2 km south-west of Nahavand, 48.3532°N, 34.1691°E.—FSJF 3526, 2, 30–40 mm SL; Iran: Hamadan prov.: Gamasiab River at Chesme Mahi, 34.3382°N, 48.0324°E.—SMF IR7, 3, 36–44 mm SL; Iran: Khozestan prov.: Marun River near Behbehan, 30.6567°N, 50.1883°E.
Oxynoemacheilus kentritensis FFR 1566, holotype, 67 mm SL; Türkiye: FFR 01403, paratypes, 3, 57–68 mm SL; Bitlis prov.: stream Kesan about 1 km south of Güntepe, 38.3566°N, 42.6275°E.—FSJF 3645, paratypes, 3, 65–79 mm SL; Türkiye: Bitlis prov.: stream Horozdere east of Hizan, 38.2447°N, 42.4791°E.—FSJF 3646, paratypes, 2, 68–70 mm SL; Türkiye: Bitlis prov.: stream Oraniz about 1 km east of Dönertaş, 38.3141°N, 42.5655°E.
Oxynoemacheilus kurdistanicus FSJF 3369, 28, 40–61 mm SL; Iraq: Nalparez River, 35.5707°N, 45.8630°E.—FSJF 3347, 25, 50–62 mm SL; Iraq: stream north-west of Saburawa, a tributary of Tabin River, 35.8336°N, 45.1044°E.—FSJF 3353, 9, 40–61 mm SL; Iraq: stream KunaMassi in Sevanja, 35.7892°N, 45.4030°E.—FSJF 3373, 54, 35–62 mm SL; Iraq: stream Suraw near Suraw village, 35.7626°N, 45.9848°E.—FSJF 3643, 15, 36–62 mm SL; Iraq: Choman River at Alut, 35.9564°N, 45.6155°E.
Oxynoemacheilus muefiti FFR 15507, paratypes, 2, 29–45 mm SL; Ağrı prov.: Türkiye: Murat River at Ballıbostan; 39.6780°N, 43.1890°E.—FFR 1432, 7, 42–63 mm SL, Ağrı prov.: Türkiye: Murat River at Taşlıçay; 39.6460°N, 43.3670°E.—FSJF 3444, 4, 33–46 mm SL; Türkiye: Ağrı prov.: Murat River at Ballıbostan, 12 km east of Ağrı, 39.6789°N, 43.1896°E.—FSJF 2556, 3, 45–47 mm SL; Türkiye: Adıyaman prov.: stream Eğri south of Adıyaman, tributary to Atatürk Reservoir, 37.7417°N, 38.3351°E.—IUSHM 2019-1410, 3, 37–68 mm SL; Türkiye: Ağrı prov.: stream near Sarıköy, 16 km west of Eleşkirt, 39.8016°N, 42.4816°E.—IUSHM 2019-1411, 3, 43–52 mm SL; Türkiye: Ağrı prov.: Murat River at Balıbostan, 12 km east of Ağrı, 39.6789°N, 43.1896°E.
Oxynoemacheilus paucilepis FFR15510, 10, 34–73 mm SL; Türkiye: Sivas prov.: stream Balıklıtohma about 3 km south of Kocakurt, 39.1440°N, 37.2570°E.—FFR15521, 15, 41–76 mm SL; Türkiye: Sivas prov.: stream Balıklıtohma at Kuruayşe, 39.2070°N, 37.2010°E.—FSJF 2852, 50, 24–39 mm SL; Türkiye: Sivas prov.: stream Tersakan about 15 km southwest of Kangal, 39.1439°N, 37.2571°E.
Oxynoemacheilus persa NMW 48567, holotype, 50 mm SL; Iran: spring at Persepolis.—FSJF 3214 (earlier IZA 7826), 25 paratypes of O. farsicus, 34–56 mm SL; Iran: Fars prov.: Shur River at Dasht-e-Arzhan, a tributary of Mond River.— FSJF 2245, 44, 31–65 mm SL; Iran: Fars prov.: Kor River about 73 km north of Shiraz, 30.1936°N, 52.4657°E.
Oxynoemacheilus sarus FFR 15585, holotype, 52.5 mm SL; FFR 15522, paratypes, 4, 39–54 mm SL;Türkiye: Adana prov.: lower stream Çakıt, south of Salbaş, 37.1031°N, 35.1094°E.—FSJF 2327, paratypes, 10, 32–49 mm SL; Türkiye: Adana prov.: lower stream Çakıt, south of Salbaş, 37.0961°N, 35.1170°E.—FSJF 2377, paratypes, 2, 48–49 mm SL; Türkiye: Adana prov.: stream Körkün at Karakuyu, 37.1529°N, 35.1606°E.—FFR 15586, 3, 47–51 mm SL; Türkiye: Kahramanmaraş Prov.; stream Aksu at 8 km northeast of Pazarcık, 37.5390°N, 37.3480°E.—FSJF 2567, 1, 48 mm SL; Türkiye: Adıyaman prov.: stream Çelik at road south of Gölbaşı, 37.6239°N, 37.5034°E.
Oxynoemacheilus shehabi
Oxynoemacheilus zarzianus FSJF 3352, 28, 39–69 mm SL; Iraq: stream Kunamasi in Sevanja, 35.7892°N, 45.4030°E.—FSJF 3348, 16, 46–68 mm SL; Iraq: stream in Merga village, 36.0515°N, 45.0945°E.—FSJF 3651, 18, 54–75 mm SL: Iraq: stream Kunamasi in Kunamasi, 35.7967°N, 45.4136°E.—FSJF 3372, 30, 43–71 mm SL; Iraq: stream Suraw near Suraw village, 35.7626°N, 45.9848°E.
Authors would like to thank Fadil Kaya (Bitlis) for his help during the fieldwork. We also thank Müfit Özuluğ (IUSHM), Anja Palandacic (NMW), Serkan Wesel (ZFMK-ICH), and Ralf Thiel (ZMH) for allowing JF to examine materials under their care. Because the second author contributed to this manuscript at Bournemouth University, we would like to thank Bournemouth University for providing their facilities, and TÜBİTAK BİDEB (2219 Program) which supported her with one-year scholarships during her post-doc research at United Kingdom.