Int. J. Aquat. Biol. (2021) 9(1): 33-40 ISSN: 2322-5270; P-ISSN: 2383-0956 Journal homepage: www.ij-aquaticbiology.com © 2021 Iranian Society of Ichthyology Original Article Capoeta baliki Turan, Kottelat, Ekmekçi & Imamoglu, 2006 a junior synonym of Capoeta tinca (Heckel, 1843) (Teleostei: Cyprinidae) Erdoğan Çiçek* 1, Soheil Eagderi2, Sevil Sungur3, Burak Secer1 1Department of Biology, Faculty of Art and Sciences, Nevşehir Hacı Bektaş Veli University, Nevşehir, Turkey. 2Department of Fisheries, Faculty of Natural Resources, University of Tehran, Karaj, Iran. 3Health Services Vocational School, Nevşehir Hacı Bektaş Veli University, 50300, Nevşehir, Turkey. s Article history: Received 6 October 2020 Accepted 27 December 2020 Available online 2 5 February 2021 Keywords: Sakarya basin Kızılırmak basin Susurluk basin Interspecific variation Abstract: Capoeta baliki was described from Sakarya basin, Turkey. It was distinguished from its nearest congener i.e. C. tinca based on a combination of characters, including fewer serrae along posterior margin of last simple dorsal-fin ray, modally fewer scale rows between lateral line and dorsal-fin origin, fewer vertebrae, deeper and shorter head and slenderer caudal peduncle. We examined the synonymy hypothesis of C. baliki and C. tinca by comparing their morphometric, meristic and molecular characters. Based on the results, their morphometric and meristic characters largely overlapped and no character was found to distinguish them. In addition, a low K2P mean genetic divergence of 0.37% C. baliki and C. tinca based on cytb gene and clustering in same clad showed that they are identical in molecular characters. As no character could be found to clearly distinguish these species, we treat C. baliki as a junior synonym of C. tinca. Introduction Capoeta tinca Heckel (1843), described from Nilüfer River, Bursa Province, Susurluk basin of Turkey, is found throughout the Black Sea watersheds, as well as Sakarya and Kizilirmak basins in the central Anatolia (Geldiay and Balık, 2007; Çiçek at al., 2020). Turan et al. (2006) described the C. tinca populations of the central Anatolian basins i.e. Sakarya and Kizilirmak as a distinct species of C. baliki based on some morphological characters. However, further works (Özdemir, 2013, 2015; Kaya, 2019) rejected distinguishing characters of C. baliki from C. tinca suggesting C. baliki as a junior synonym of C. tinca. Additionally, pronounced discriminative differences was not found in the osteological characteristics of C. baliki and C. tinca (Küçük et al., 2008). In recent years, molecular studies (Özdemir, 2013; Bektaş et al., 2017, 2019; Ghanavi et al., 2016; Levin, 2012; Zareian and Esmaeili, 2017; Zareian et al., 2018) based on both COI and cytb genes of mitochondrion indicated clustering of C. tinca and C. baliki in the same clade supporting C. baliki as *Correspondence: Erdoğan Çiçek DOI: https://doi.org/10.22034/ijab.v9i1.1118 E-mail: erdogancicek@nevsehir.edu.tr junior synonym of C. tinca. Hence in the present study, we decided to examine the synonymy hypothesis of C. baliki and C. tinca by comparing their morphometric, meristic and molecular (mtDNA cytb gene) characters. Materials and Methods Specimens of C. tinca were collected from its type locality (Susurluk basin) and C. baliki from Sakarya basins. All specimens caught by electrofishing, and after anaesthesia by MS222, they were fixed into 5% buffered formaldehyde and stored in 70% ethanol after two weeks. Methods for counts and measurements followed Armbruster (2012). Measurements were made with a dial calliper to the nearest 0.1 mm. Gill rakers were counted on the outer margin of the anterior gill arch. The posterior pair of the branched rays articulating on a single pterygiophore in the dorsal and anal fins were noted as “1½”. Twenty morphometric characters were measured (Table 1) and then standardized using an allometric method to remove size-dependent variation using Madj 34 Çiçek et al./ Capoeta baliki a junior synonym of Capoeta tinca = M (Ls / L0)b, where M is the original measurement, Madj the size adjusted measurement, L0 the standard length of the fish, Ls the overall mean of the standard length for all fish from all samples in each analysis, and b was estimated for each character from the observed data as the slope of the regression of log M on log L0 using all fish in any group (Elliot, 1999). The results derived from the allometric method were confirmed by testing significance of the correlation between transformed variables and standard length (Buj et al., 2008). The morphometric data of the two species were analyzed using multivariate analyzes of Table 1. Cytochrome b sequences downloaded from NCBI GenBank with information on basin, country of origin and reference. Species Drainage Country Published by Genbank Acc. No. Capoeta baliki Black Sea basin, Sakarya Turkey Bektas et al. 2017 GQ424019 Capoeta baliki Black Sea basin, Sakarya Turkey Bektas et al. 2017 GQ424016 Capoeta baliki Black Sea basin, Sakarya Turkey Bektas et al. 2017 GQ424015 Capoeta baliki Black Sea basin, Sakarya Turkey Bektas et al. 2017 GQ424014 Capoeta baliki Black Sea Basin, Sakarya Turkey Bektas et al. 2017 GQ424013 Capoeta baliki Black Sea basin, Sakarya Turkey Bektas et al. 2017 GQ424012 Capoeta baliki Black Sea basin, Sakarya Turkey Bektas et al. 2017 GQ424011 Capoeta baliki Lake Iznik basin, Çakirca Stream Turkey Levin et al. 2012 JF798275 Capoeta baliki Sakarya basin, Kurtbogazı Dam Lake Turkey Levin et al. 2012 JF798274 Capoeta baliki Sakarya basin, Kurtbogazı Dam Lake Turkey Levin et al. 2012 JF798273 Capoeta baliki Black Sea Basin, Kelkit River Turkey Levin et al. 2012 JF798272 Capoeta baliki Black Sea Basin, Kızılırmak River Turkey Levin et al. 2012 JF798271 Capoeta tinca Eber Lake Basin, Afyon Turkey Bektas et al. 2017 GQ424010 Capoeta tinca Eber Lake Basin, Afyon Turkey Bektas et al. 2017 GQ424009 Capoeta tinca Marmara Basin, Susurluk Turkey Bektas et al. 2017 GQ424008 Capoeta tinca Marmara Basin, Susurluk Turkey Bektas et al. 2017 GQ424007 Capoeta tinca Marmara Basin, Susurluk Turkey Bektas et al. 2017 GQ424006 Capoeta tinca Marmara Basin, Susurluk Turkey Bektas et al. 2017 GQ424005 Capoeta tinca Marmara Basin, Susurluk Turkey Bektas et al. 2017 GQ424004 Capoeta damascina Mediterranean Sea basin, Orontes River Turkey Levin et al. 2012 JF798306 Capoeta damascina Mediterranean Sea basin, Orontes River Turkey Levin et al. 2012 JF798305 Capoeta aydinensis Aegean Sea Basin, B. Menderes Turkey Bektas et al. 2017 KY065274 Capoeta aydinensis Aegean Sea Basin, B. Menderes Turkey Bektas et al. 2017 KY065275 Capoeta caelestis Mediterranean Sea basin, Kargi Stream Turkey Levin et al. 2012 JF798287 Capoeta caelestis Mediterranean Sea basin, Kargi Stream Turkey Levin et al. 2012 JF798288 Capoeta antalyensis Mediterranean Sea Basin, Aksu at Gokdere Turkey Bektas et al. 2017 GQ424021 Capoeta antalyensis Mediterranean Sea Basin, Aksu at Gokdere Turkey Bektas et al. 2017 GQ424023 Capoeta sieboldii Black Sea Basin, Kelkit River Turkey Levin et al. 2012 JF798330 Capoeta sieboldii Black Sea Basin, Kızılırmak River Turkey Levin et al. 2012 JF798329 Capoeta sieboldii Black Sea Basin, Kizilirmak Turkey Bektas et al. 2017 KY065259 Capoeta sieboldii Black Sea Basin, Kizilirmak Turkey Bektas et al. 2017 KY065258 Capoeta sieboldii Black Sea Basin, Yesilirmak Turkey Bektas et al. 2017 KY065256 Capoeta bergamae Marmara Basin, Bakacak stream Turkey Levin et al. 2012 JF798282 Capoeta bergamae Bakırçay River Turkey Levin et al. 2012 JF798280 Capoeta banarescui Black Sea Basin, Coruh Turkey Bektas et al. 2017 GQ423988 Capoeta banarescui Black Sea Basin, Coruh Turkey Bektas et al. 2017 GQ423984 Capoeta banarescui Black Sea Basin, Coruh Turkey Bektas et al. 2017 GQ423992 Capoeta banarescui Black Sea Basin, Coruh Turkey Bektas et al. 2017 GQ423991 Capoeta banarescui Black Sea Basin, Coruh Turkey Bektas et al. 2017 GQ423990 Capoeta banarescui Black Sea Basin, Coruh Turkey Bektas et al. 2017 GQ423989 Capoeta capoeta Caspian Sea basin, Aras River Iran Ghanavi et al. 2016 KU167938 Capoeta trutta Karoun River Drainage, Lordegan Iran Ghanavi et al. 2016 KM459673 Luciobarbus esocinus Tigris River Turkey Yang et al. 2015 KP712264 35 Int. J. Aquat. Biol. (2021) 9(1): 33-40 principal component analysis (PCA) and P-value obtained from permutation test of one-way NPMANOVA. All outliers were removed from further analysis. All analyses were performed using PAST software. Molecular data analysis: For this study, we retrieved 43 cytb sequences of the published Capoeta from GenBank using the (BLASTn) basic local alignment search tool (Altschul et al., 1990) (Table 1). For phylogenetic reconstruction, the datasets were analysed by Bayesian Inference (BI) using MrBayes 3.1.2 (Ronquist et al., 2011) and the maximum likelihood (ML) method in IQ-TREE 1.6.0 (Nguyen et al., 2015). We determined the best-fit model of molecular evolution for the genomic dataset using the Bayesian information criterion (BIC) in IQTREE 1.6.0 (Kalyaanamoorthy et al., 2017). MrBayes was run with 6 substitution types (nst=6) and considered the gamma-distributed rate variation across sites plus a proportion of invariable sites (GTR) for the COI datasets. For BI, Bayesian inference was calculated with MrBayes v.3.2.6 (Ronquist et al., 2011). Two simultaneous analyses were run with each 2,000,000 generations and four MCMC chains sampling every 10,000 generations. Convergence was checked on Tracer 1.6 (Rambaut and Drummond, 2013). After discarding the first 10% of generations as burn-in, we obtained the 50% majority rule consensus tree and the posterior probabilities. For ML analyses, we conducted heuristic searches (1,000 runs) under a TN+F+G4 model. Uncorrected pairwise genetic distances (p-distances) were investigated based on Kimura two-parameter (K2P) distances (Tamura et al., 2013). Capoeta capoeta (KU167938), Capoeta trutta (KM459673) and Luciobarbus esocinus (KP712264) were used as outgroups. Abbreviations used. HL, Head length; SL, standard length; K2P, Kimura 2-parameter. Collection codes: NUIC, Ichthyological Collection of the Nevsehir Haci Bektas Veli University. Results General appearances of C. tinca and C. baliki are presented in Figure 1 showing their body shapes and colour patterns similarity. Tables 2 and 3 represent their morphometric measurements and meristic counts, respectively. All morphometric and meristic features of C. baliki are largely overlapped with those of C. tinca. We failed to find any non-overlapping morphological differences between the C. tinca and Figure 1. Lateral view of (A) Capoeta baliki, NUIC-1816, 106.3 mm SL; Derecik Stream, Ankara province, and (B) C. tinca, NUIC-1717, 119.2 mm SL Değirmen Stream, Balıkesir province (all from Turkey). 36 Çiçek et al./ Capoeta baliki a junior synonym of Capoeta tinca Table 2. Morphometric data of Capoeta tinca (n=20) and C. baliki populations (n=20). Morphometric characters Capoeta tinca (NUIC-1717) Capoeta baliki (NUIC-1816) min-max mean±SD min-max mean±SD Standard length (mm) 90.6-140.7 117.0±15.2 90.8-227.3 127.4±37.2 In percent of standard length Head length 23.4-24.9 24.3±0.5 21.4-26.4 24.2±1.3 Body depth at dorsal fin origin 22.9-26.3 24.5±1.1 20.2-24.8 23.1±1.4 Predorsal length 47.0-53.2 49.6±2.1 46.7-53.8 50.9±1.9 Prepelvic length 51.2-54.4 52.5±1.1 49.8-54.9 52.8±1.5 Preanal length 72.9-76.0 74.7±1.1 73.5-78.1 75.0±1.6 Distance between pectoral-fin origin to anal fin 51.1-55.7 53.9±1.5 43.3-58.1 52.7±3.5 Distance between pectoral-fin origin to pelvic fin 29.1-32.9 31.1±1.2 29.7-34.2 31.1±1.3 Distance between pelvic-fin origin to anal fin 21.1-23.9 22.6±0.9 20.2-24.2 22.3±1.0 Dorsal-fin height 19.3-22.8 21.4±1.1 18.1-22.7 20.4±1.5 Anal-fin length 16.0-18.4 16.9±0.9 16.0-22.2 18.3±2.1 Pectoral-fin length 17.6-19.7 18.7±0.7 17.3-19.6 18.6±0.8 Pelvic-fin length 15.0-16.6 15.8±0.6 14.7-17.1 15.7±0.7 Upper caudal-fin lobe 16.9-25.3 22.7±2.2 18.1-23.6 21.6±1.4 Length of caudal peduncle 17.6-19.3 18.6±0.7 16.3-20.3 18.0±1.4 Depth of caudal peduncle 11.5-12.8 12.1±0.4 10.9-12.9 12.2±0.6 In percent of Head length Head depth at eye 45.2-51.0 48.4±1.7 48.0-55.2 50.9±2.2 Snout length 32.4-38.2 35.2±1.4 35.3-40.4 37.5±1.6 Eye horizontal diameter 17.1-22.5 19.8±1.5 16.6-22.1 19.0±1.8 Interorbital width 37.5-42.0 39.9±1.7 37.4-44.2 41.1±2.0 Postorbital distance 46.2-51.2 48.8±1.8 46.2-52.3 49.1±1.7 Maximum head width 60.2-68.3 62.9±2.5 60.5-69.2 63.7±2.8 Table 3. Meristic data of Capoeta tinca (NUIC-1717) and C. baliki (NUIC-1816) (n=20 in each populations). Gill raker Examined materials 19 20 21 22 23 24 Capoeta tinca 3 6 10 1 Capoeta baliki 2 5 3 8 1 1 Lateral Line Scales Examined materials 75 76 77 78 79 80 Capoeta tinca 8 4 3 Capoeta baliki 12 5 3 Scales above lateral line Scales below lateral line Examined materials 14 15 16 17 18 7 8 9 Capoeta tinca 1 15 4 1 14 5 Capoeta baliki 1 12 6 1 18 2 Branched dorsal-fin rays Examined materials 7½ 8½ 9½ mode Capoeta tinca 4 16 9 Capoeta baliki 2 18 9 Branched anal-fin rays Examined materials 5 6 7 mode Capoeta tinca 2 18 6 Capoeta baliki 4 16 6 Pelvic-fin rays Examined materials 7 8 9 mode Capoeta tinca 2 18 8 Capoeta baliki 20 8 Pectoral-fin rays Examined materials 17 18 19 20 mode Capoeta tinca 2 16 2 18 Capoeta baliki 18 2 18 37 Int. J. Aquat. Biol. (2021) 9(1): 33-40 C. baliki. In PCA, the first three PCs accounted a total of 84.49% of the variances (PC1=41.02, PC2=20.90 and PC3=12.857) (Jolliffe cut-off=1.119). By plotting the first two PCs, the distribution of the studied species based on their morphometrics are presented in Figure 2, showing overlapping the specimens of both C. tinca and C. baliki. The result of multivariate one-way NPMANOVA showed no different between two species (P=0.1003, F=1.913). Based on Figure 3, Capoeta species, including C. tinca, C. baliki, C. banarescui and C. antalyensis were clustered in the same clade with those C. baliki and C. tinca merged in the same clade. A low K2P mean genetic divergence of 0.37% was calculated between C. baliki and C. tinca (Özdemir, 2013; Bektaş et al., 2019) (Table 4). Discussions According to Turan et al. (2006), C. baliki is distinguished from C. tinca by having fewer serrae along posterior margin of last simple dorsal-fin ray (17-23 vs. 24-28), modally fewer scale rows between lateral line and dorsal-fin origin (14 vs. 16), fewer vertebrae (43-44 vs. 44-46), shorter head (21.8-24.5 vs. 23.3-26.7% SL), deeper head (55.6-63.5 vs. 49.3- 56.5% HL) and lower caudal peduncle (9.5-12.2 vs. 10.8-13.4% SL). Based on the examined materials, our data in line with pervious findings (Özdemir, 2013, 2015; Kaya, 2019) showed overlapping of all above-mentioned distinguishing characters as well as others (Tables 2 and 3). Therefore, there is no morphological diagnostic characters to distinguish C. baliki from C. tinca. Capoeta tinca and C. baliki were clustered in the Figure 2. The PCA graph of morphometric characters in Capoeta tinca (NUIC-1717, n=10) and C. baliki (NUIC-1816, n=12). Table 4. Estimates of average K2P genetic divergence over sequence pairs between the studied Capoeta species. Species No 1 2 3 4 5 6 7 8 C. baliki 1 C. tinca 2 0.37 C. damascina 3 3.63 4.19 C. aydinensis 3 4.29 4.84 3.19 C. caelestis 4 4.11 4.47 1.43 3.93 C. antalyensis 5 1.81 2.02 3.77 4.71 4.04 C. sieboldi 6 4.49 5.07 2.99 4.15 3.58 4.74 C. bergamae 7 4.74 5.31 3.74 2.61 4.39 5.18 4.83 C. banarescu 8 4.30 4.33 3.51 5.05 4.18 3.56 5.40 5.53 38 Çiçek et al./ Capoeta baliki a junior synonym of Capoeta tinca same clade and cannot be considered as distinct species based on phylogenetic species concept (PSC). In addition, mean K2P genetic divergence of 0.37% between C. tinca and C. baliki is low for species delimitation criteria suggested by Geiger et al. (2014) for freshwater fishes of the Mediterranean region and also this genetic distance stands within intraspecific range in the genus Capoeta based on the pervious Figure 3. Bayesian Inference of the phylogenetic relationships based on the mitochondrial cytb barcode region (values at nodes correspond to BI posterior probability/ ML bootstrap). 39 Int. J. Aquat. Biol. (2021) 9(1): 33-40 studies (Bektaş et al., 2017, 2019; Ghanavi et al., 2016; Levin, 2012; Zareian and Esmaeili, 2017; Zareian et al., 2018). Furthermore, Bektaş et al. (2017) reported that the haplotypes of C. baliki (from Sakarya river drainage and Lake Eber) and C. tinca (from Susurluk drainage) are closely related. The basins drain to Black Sea and the Sea of Marmara are not fully isolate from each other and exchanges routes are still available e.g. via river capture (Yıldırım and Emre, 2004). Therefore, C. tinca and C. baliki have perhaps recently isolated. Moreover, even existence of minor molecular and morphological differences between populations of widespread freshwater fish species, is a well-studied phenomenon (Marcil et al., 2006). Since no morphological diagnostic characters to distinguish C. baliki from C. tinca is available and they are identical in molecular characters i.e. cytb gene, therefore we treat C. baliki as a junior synonym of C. tinca. Material examined. All from Turkey. Capoeta baliki, NUIC-1816, 20, 90.8-227.3 mm SL; Ankara prov.: Derecik Stream, Sakarya basin, 40°30'44''N 32°19'30''E; 15 May 2018. ⸺ ESFM- PISI/2004-74 (Holotype), 202 mm SL; Ankara prov.: Sakarya River: Kizilcahamam Stream, 60 km west of Ankara, 40°29'N 32°39'E; 15 April 2004. ⸺ ESFM- PISI/2004-75, 4, 140-190 mm SL; same data as holotype. Capoeta tinca, NUIC-1717, 20, 90.6-140.7 mm SL; Balıkesir prov.: Değirmen Stream, Susurluk basin, 39°54'50''N 27°33'50''E. Zoobank Registration: urn:lsid:zoobank.org:pub:72 9C2166-8359-401D-B3CA-102A9CBACB4B Acknowledgments The authors would like to thank Nevsehir Hacı Bektas Veli and Tehran Universities for financial supports. We also thank D. Turan, C. Kaya and E. Bayçelebi for helping us to examine the types of C. baliki. References Altschul S.F., Gish W., Miller W., Myers E.W., Lipman D.J. (1990). Basic local alignment search tool. Journal of Molecular Biology, 215: 403-410. Armbruster J.W. (2012). Standardized measurements, landmarks, and meristic counts for cypriniform fishes. Zootaxa, 3586: 8-16. Bektaş Y., Aksu İ., Kaya C., Turan D. (2019). DNA Barcoding of the Genus Capoeta (Actinopterygii: Cyprinidae) from Anatolia. Turkish Journal of Fisheries and Aquatic Sciences, 19(9): 739-752. Bektaş Y., Turan D., Aksu İ., Kaya C., Ciftci Y., Eroglu O., Kalaycı G. Beldüz A.O. (2017). Molecular phylogeny of the genus Capoeta (Teleostei: Cyprinidae) in Anatolia, Turkey. Biochemical Systematics and Ecology, 70: 80-94. Buj I., Podnar M., Mrakovčić M., Ćaleta M., Mustafić P., Zanella D., Marčić Z. (2008). Morphological and genetic diversity of Sabanejewia balcanica in Croatia. Folia Zoologica, 57(1): 100-110. Çiçek E., Fricke R., Sungur S., Eagderi E. (2018). Endemic freshwater fishes of Turkey. FishTaxa, 3(4): 1-39. Çiçek E., Sungur S., Fricke R. (2020). Freshwater lampreys and fishes of Turkey; a revised and updated annotated checklist 2020. Zootaxa, 4809(2): 241-270. Elliott N.G., Haskard K., Koslow J.A. (1995). Morphometric analysis of orange roughy (Hoplostethus atlanticus) off the continental slope of southern Australia. Journal of Fish Biology, 46: 202-220. Geiger M.F., Herder F., Monaghan M.T., Almada V., Barbieri R., Bariche M., Berrebi P., Bohlen J., Casal- Lopez M., Delmastro G.B., Denys G.P., Dettai A., Doadrio I., Kalogianni E., Karst H., Kottelat M., Kovacic M., Laporte M., Lorenzoni M., Marcic Z., Ozulug M., Perdices A., Perea S., Persat H., Porcelotti S., Puzzi C., Robalo J., Sanda R., Schneider M., Slechtova V., Stoumboudi M., Walter S., Freyhof J. (2014). Spatial heterogeneity in the Mediterranean biodiversity hotspot affects barcoding accuracy of its freshwater fishes. Molecular Ecology Resources, 14(6): 1210-1221. Geldiay R., Balik S. (2007). Freshwater Fishes of Turkey. V. Edition, Ege University Press, Bornova, Izmir. 638 p. Ghanavi H.R., Gonzalez E.G., Doadrio I. (2016). Phylogenetic relationships of freshwater fishes of the genus Capoeta (Actinopterygii, Cyprinidae) in Iran. Ecology and Evolution, 6: 8205-8222. Kalyaanamoorthy S., Minh B.Q., Wong T.K.F., von Haeseler V., Jermiin L.S. (2017). ModelFinder: fast model selection for accurate phylogenetic estimates. 40 Çiçek et al./ Capoeta baliki a junior synonym of Capoeta tinca Nature Methods, 14: 587-589. Kaya C. (2019). Taxonomic revision of the species belonge to the genus Capoeta by using both the distributed in Turkey. Ph.D. thesis, Recep Tayyip Erdogan University, Rize. 126 p. Küçük F., Turan D., Turna I.I. (2008). Comparing the osteological characteristics of some Capoeta (Teleostei: Cyprinidae) species distribution in Anatolia. E.U. Journal of Fisheries and Aquatic Sciences, 25(4): 267- 273. Levin B.A., Freyhof J., Lajbner Z., Perea S., Abdoli A., Gaffaroğlu M., Özulug M., Rubenyan H.R., Salnikov V.B., Doadrio I. (2012). Phylogenetic relationships of the algae scraping cyprinid genus Capoeta (Teleostei: Cyprinidae). Molecular Phylogenetics and Evolution, 62: 542-549. Marcil J., Swain D.P., Hutchings J.A. (2006). Genetic and environmental components of phenotypic variation in body shape among populations of Atlantic cod (Gadus morhua L.). Biological Journal of the Linnean Society, 88(3): 351-365. Nguyen N.D., Mirarab S., Kumar K. (2015). Ultra-large alignments using phylogeny-aware profiles. Genome Biology 16: 124. Özdemir F. 2013. The revision of species and subspecies of the genus Capoeta (Teleostei: Cyprinidae) by using both the classical systematic and molecular systematic methods in Turkey. Ph.D. thesis, Hacettepe University, Ankara. 171 p. Özdemir F. (2015). principle components analysis of two pairs of Barbels Species of the Genus Capoeta (Teleostei: Cyprinidae) in Turkey. Pakistan Journal of Zoology, 47(3): 753-762. Rambaut A., Drummond A.J. (2013). Tree Annotator v1.8.0 MCMC Output Analysis. Ronquist F., Teslenko M., van der Mark P., Ayres D.L., Darling A., Höhna S., Larget B., Liu L., Suchard M.A., Huelsenbeck J.P. (2012). MrBayes 3.2: Efficient bayesian phylogenetic ınference and model choice across a large model space. Bioanformatics, 61: 539- 542. Tamura K., Stecher G., Peterson D., Filipski A., Kumar S. (2013). MEGA6: Molecular Evolutionary Genetics Analysis Version 6.0. Molecular Biology and Evolution, 2013 Dec; 30(12): 2725-2729. Turan D., Kottelat M., Ekmekçi F.G., Imamoğlu H.O. (2006). A review of Capoeta tinca, with descriptions of two new species from Turkey (Teleostei: Cyprinidae). Revue Suisse de Zoologie, 113(2): 421-436. Yıldırım C., Emre O. (2004). Drainage evolution along the north Anatolian fault zone, eastern Marmara-Turkey. In: Denver Annual Meeting, 19-24. Zareian H., Esmaeili H.R. (2017). Mitochondrial phylogeny and taxonomic status of the Capoeta damascina species group (Actinopterygii: Cyprinidae) in Iran with description of a new species. Iranian Journal of Ichthyology, 4(3): 231-269. Zareian H., Esmaeili H.R., Gholamhosseini A., Japoshvili B., Ozulug M., Mayden R.L. (2018). Diversity, mitochondrial phylogeny, and ichthyogeography of the Capoeta capoeta complex (Teleostei: Cyprinidae). Hydrobiologia, 806: 363-409.