. J. Ichthyol. (September 2017), 4(3): 231-269 Received: July 27, 2017 © 2017 Iranian Society of Ichthyology Accepted: September 10, 2017 P-ISSN: 2383-1561; E-ISSN: 2383-0964 doi: 10.22034/iji.v4i3.23902.015 http://www.ijichthyol.org

Research Article

Mitochondrial phylogeny and taxonomic status of the damascina species group (: ) in Iran with description of a new species

Halimeh ZAREIAN, Hamid Reza ESMAEILI*

Ichthyology and Molecular Systematics Research Laboratory, Zoology Section, Department of Biology, College of Sciences, Shiraz University, Shiraz, Iran. * Email: [email protected]

Abstract: The members of the genus Capoeta show an abstruse taxonomic status, a complex evolutionary history with the highest diversification in the Middle East and are closely related to the genus Luciobarbus. Our molecular and morphological results confirm the existence of eight species within the Capoeta damascina group in Iran. In this paper all eight recognized species are reviewed, and diagnoses are presented for all species. Capoeta birunii is considered as new species from Zayandehrud drainage basins. This species is distinguished from other small scaled Capoeta in the C. damascina group by a combination of morphological and molecular characters. Keywords: Capoeta, Molecular analyses, Systematics, Distribution. Citation: Zareian, H. & Esmaeili, H.R. 2017. Mitochondrial phylogeny and taxonomic status the Capoeta damascina species group (Actinopterygii: Cyprinidae) in Iran with description of a new species. Iranian Journal of Ichthyology 4(3): 231-269.

Introduction includes about 34 species widely distributed in many Iran has one of the most diverse and species-rich river drainages and basins in southwest Asia except freshwater ichthyofaunas in the Middle East the Arabian Peninsula (Jouladeh-Roudbar et al. 2015; (Esmaeili et al. 2017). However, unresolved Alwan et al. 2016a; Ghanavi et al. 2016; Keivany et taxonomic problems could result in higher fish al. 2016; Esmaeili et al. 2016, 2017; Zareian et al. diversity including cyprinid fishes of the genus 2016a, b; Eschmeyer & Fricke 2017). They have Capoeta Valenciennes, 1842 (Kücük et al. 2009; been classified into three groups, namely the Esmaeili et al. 2017). Members of the genus Capoeta C. capoeta, C. damascina and C. trutta species groups exist in the Middle East and southwest Asia. This (see Jouladeh-Roudbar et al. 2015; Alwan et al. region is an important crossroads of biotic exchange 2016a; Ghanavi et al. 2016; Zareian et al. 2016a, b). with complex geological events such as novel The Capoeta damascina species group, orogeny (Durand et al. 2002; Krupp et al. 2009) morphologically characterized by having small which is the consequence of the ongoing northward scales, was originally defined by Schöter et al. convergence of the African, Arabian and Indian (2009), and is an important issue in taxonomic study plates towards Eurasia (Zhao & Xie 2016) resulting of cyprinid fishes in recent years. Alwan et al. (2011, in isolated river basins promoting speciation events. 2016a, b) studied the Capoeta damascina species As presently recognized, the genus Capoeta group using COI and LSU genes, using a few Iranian 231

Iran. J. Ichthyol. (September 2017), 4(3): 231-269

Fig.1. Map of Iran showing different basins (M= Maharlu). After Esmaeili et al. (2017). specimens and later Ghanavi et al. (2016) studied populations which were not identified as any diversity of Capoeta species in Iran using the cytb described species (Ghanavi et al. 2016). Three of gene. The latter study proposed the presence of five these suggested new taxa were small scaled Capoeta new species of Capoeta in Iranian basins, three of distributed in the Tigris River tributaries of the them belong to the C. damascina species group and Persian Gulf basin, traditionally identified as then Jouladeh-Roudbar et al. (2017) described those C. damascina or C. saadii (Jouladeh-Roudbar et al. three new small scaled Capoeta. 2015b; Esmaeili et al. 2010; 2017). The Capoeta damascina species group occurs in Hence, the main aim of this study is to clarify the the entire Levant, Mesopotamia, Orontes, Iran and taxonomic status of the Capoeta damascina species the southern and eastern parts of . In Iran, it is group in Iran, to provide a taxonomic review and reported from the Tigris (part of the Tigris-Euphrates mitochondrial phylogeny on this group and describe River system), Namak Lake, Esfahan, Kor, Maharlu, one new species. Persis, Kerman-Nain and Hormuz basins (Coad 1995; 2008; Esmaeili et al. 2010, 2015, 2017). The Materials and Methods taxonomic status of this group is largely unsettled Taxon sampling: Fish samples were collected from (Krupp & Schneider 1989; Esmaeili et al. 2010; endorheic (Caspian Sea, Urmia Lake, Namak Lake, 2017; Alwan et al. 2016a, b; Zareian et al. 2016a, b). Kor, Kavir, Zayandehrud: Esfahan) and exorheic A detailed study of the populations of the Capoeta (Persis: Mond, Tigris, flowing into the Persian Gulf) damascina species group in Iran found some basins (Fig. 1). After anesthesia (using Clove oil), 232 Zareian and Esmaeili - Phylogeny and of Capoeta damascina species group in Iran

Table 1. Primer used in this study.

Name Sequence Gene Reference

FishF1 5'TCAACCAACCACAAAGACATTGGCAC3' FishR1 5'TAGACTTCTGGGTGGCCAAAGAATCA3' COI Ward et al. 2005 L14724 5'GTGACTTGAAAAACCACCGTTG3' H15915 5'CAACGATCTCCGGTTTAGAAGAC3' cytb Xiao et al. 2001 GluF 5'AACCACCGTTGTATTCAACTACAA3' H15560 5`TAGGCRAATAGG AAR TATCA3` cytb Machordom & Doadrio 2001 specimens were fixed in 10% formaldehyde and later oxidase subunit 1: 650 bp) and cytochrome b (cytb: stored in 70% ethanol for the morphological study. 900 bp) region were amplified using a primer pairs The right pectoral fin or tissue from below the dorsal listed in Table 1. fin on the right side of each specimen was removed Purification and sequencing of the PCR products using sterile techniques and utensils and preserved in were conducted at Macrogen Korea Laboratories 96% ethanol and numbered separately at the using aforementioned primer pairs. Data processing sampling sites for the molecular study implied by and sequence assembly was done in BioEdit 7.2.5 sterile techniques. (Hall 1999) and MEGA6 (Tamura et al. 2013) was Morphological analyses: Twenty-two morphometric used to create a DNA sequence alignment using measurements were conducted with an electronic Clustal W program for COI and cytb. No indications digital caliper, to the nearest 0.01mm. For measuring of unexpected stop-codons occurred in any sequence. fish greater than 200mm total length, a ruler or a All generated DNA barcodes and cytb are deposited measuring tape was used. All measurements were in the NCBI GenBank, and given with their made point to point, and never by projections. respective accession numbers (Table 2). The most Measurements and counts mainly followed Hubbs & appropriate sequence evolution model for the given Lagler (1958), Krupp (1983) and Kottelat & Freyhof data was determined with Modeltest (Posada & (2007). Fin ray counts separate unbranched and Crandall 1998) as implemented in the MEGA6 branched rays. The last two branched rays software. The model with the lowest BIC (Bayesian articulating on a single pterygiophore in the dorsal Information Criterion) scores is considered to best and anal fins are counted as "1½". Principle describe the substitution pattern. To explore species Components and Discriminant Factor analysis were phylogenetic affinities, we generated Maximum performed using SPSS software (Ver. 22). Likelihood phylogenetic trees with 10,000 bootstrap Abbreviations: SL, standard length; HL, lateral head replicates in RaxML software 7.2.5 (Stamatakis length; ALL, scale rows between lateral line and 2006) under the GTR+G model of nucleotide dorsal fin origin, BLL, scale rows between lateral substitution, with fast bootstrap for both genes and line and anal fin origin, NMW, Naturhistorisches also Bayesian analysis (BA) of nucleotide sequences Museum Wien, Vienna; ZM-CBSU, Zoological was performed, using the Markov Chain Monte Carlo Museum of Shiraz University, Collection of Biology algorithm (MCMC), with 10,000,000 generations Department, Shiraz. under the most generalizing model (GTR+G+I) using Molecular analyses: DNA extraction from either Mr. Bayes 3.1.1 (Huelsenbeck & Ronquist 2001). muscle tissues or fin clips were done using the Salt Screening for diagnostic nucleotide substitutions was method (Bruford et al. 1992). The standard vertebrate performed manually from the sequence alignment. DNA barcode region of the COI (cytochrome c Cyprinus carpio was used as outgroup.

233 Iran. J. Ichthyol. (September 2017), 4(3): 231-269

Table 2. List of species used in this study with museum number. * indicate accession numbers of COI genes and others are cytb gene accession numbers.

Species Basin Museum N. Accession N. Species Basin Museum N. Accession N. C. buhsei Namak ZM-CBSU 1289 KU564292* C. anamisensis Minab ZM-CBSU 1416 KU312342* MF621312 KU312379 C. buhsei Namak ZM-CBSU 1290 KU564293* C. anamisensis Minab ZM-CBSU 1417 KU312343* MF621311 KU312380 C. buhsei Namak ZM-CBSU 1292 MF621275* C. anamisensis Hasan Langi ZM-CBSU 1475 KU312341* MF621310 KU312381 C. buhsei Namak ZM-CBSU 1299 KU312349* C. pyragyi Tigris ZM-CBSU 1474 MF621294* KU312369 MF621321 C. buhsei Namak ZM-CBSU 1300 KU312350* C. pyragyi Tigris ZM-CBSU 1470 MF621293* KU312370 MF621322 C. coadi Tigris ZM-CBSU 1447 KU564297* C. shajariani Tigris ZM-CBSU 1835 MF621299* KU564303 MF621328 C. coadi Tigris ZM-CBSU 1448 KU564298* C. shajariani Tigris ZM-CBSU 1836 MF621300* KU564304 MF621329 C. coadi Tigris ZM-CBSU 1458 KU564294* C. shajariani Tigris ZM-CBSU 1837 MF621301* KU564305 - C. coadi Tigris ZM-CBSU 1459 KU564295* C. shajariani Tigris ZM-CBSU 1476 MF621298* KU564306 - C. saadii Persis ZM-CBSU 822 KU312362* C. ferdowsii Zohreh ZM-CBSU 1832 MF621295* KU312374 MF621325 C. saadii Persis ZM-CBSU 824 KU312357* C. ferdowsii Zohreh ZM-CBSU 1833 MF621296* MF621302 MF621326 C. saadii Persis ZM-CBSU 825 KU312361* C. ferdowsii Zohreh ZM-CBSU 1834 MF621297* KU312373 MF621327 C. saadii Persis ZM-CBSU 1421 MF621266* C. birunii Zayandehrud ZM-CBSU 1413 MF621292* KU564309 MF621323 C. saadii Persis ZM-CBSU 1422 MF621267* C. birunii Zayandehrud ZM-CBSU 1155 - KU564310 MF621303 C. saadii Persis ZM-CBSU 1423 MF621268* C. birunii Zayandehrud ZM-CBSU 1159 - MF621318 MF621304 C. saadii Kor ZM-CBSU 1426 KU564299* C. buhsei Iran, Namak JF798283 KU564312 C. coadi Iran, Tigris KU564304 C. saadii Kor ZM-CBSU 1427 KU564300* C. coadi Iran, Tigris KU564305 KU564313 C. coadi Iran, Tigris KU564306 C. umbla Tigris ZM-CBSU 1469 MF621281* C. saadii Iran, Kor JF798326 - C. saadii Iran, Minab JF798327 C. umbla Tigris ZM-CBSU 1472 MF621283* C. saadii Iran, KU564311 - C. saadii Iran,Maharlou Persis KU312374 C. umbla Tigris ZM-CBSU 1477 MF621284* C. saadii Iran, Kor JF798326 - C. saadii Iran, Minab JF798327 C. umbla Tigris ZM-CBSU 1480 MF621285* Cyprinus carpio KU050703 -

Results morphological variation (Table 3). Discriminant Morphological analysis: According to the results Function Analysis (DFA) on relative warps classified from the PCA, seven functions were derived. The specimens into the correct C. damascina species first function explained 27.643% and the second one groups at the rate of 95.8% in original data (Table 4), explained 23.452% of the observed variation in the indicating a high differentiation among the groups. data. Two principal component functions explained DFA based on the morphometric and meristic 51.095% of the total variance in the observed data of characters, distinguished 8 species of C. damascina

234 Zareian and Esmaeili - Phylogeny and taxonomy of Capoeta damascina species group in Iran

Table 3. The Principle Component Analysis (PCA) of Capoeta damascina species group.

Extraction Sums of Squared Loadings Component Total % of Variance Cumulative % PCI 5.529 27.643 27.643 PCII 4.690 23.452 51.095 PCIII 3.534 17.668 68.763 PCIV 1.855 9.276 78.040 PCV 1.649 8.244 86.283 PCVI 1.337 6.686 92.969 PCVII 1.126 5.631 98.601

Table 4. Classification matrix: number and percentage of individuals that were correctly classified in the Capoeta damascina species group.

Predicted Group Membership Species C. C. C. C. C. C. C. C. Total buhsei coadi saadii umbla shajariani pyragyi ferdowsii birunii C. buhsei 27 0 0 0 0 0 0 0 27 C. coadi 0 17 0 0 1 1 0 0 19 C. saadii 0 0 14 0 0 0 0 1 15 Count C. umbla 0 0 0 13 1 0 0 0 14 C. shajariani 0 0 0 0 9 0 0 0 9 C. pyragyi 0 1 0 0 0 15 0 0 16 C. ferdowsii 0 0 0 0 0 0 9 0 9 Original C. birunii 0 0 0 0 0 0 0 11 11 C. buhsei 100.0 .0 0 0 0 0 0 0 100 C. coadi 0 89.5 0 0 5.3 5.3 0 0 100 C. saadii 0 0 93.3 0 0 0 0 6.7 100 C. umbla 0 0 0 92.9 7.1 0 0 0 100 % C. shajariani 0 0 0 0 100 0 0 0 100 C. pyragyi .0 6.3 0 0 0 93.8 0 0 100 C. ferdowsii 0 0 0 0 0 0 100 0 100 C. birunii 0 0 0 0 0 0 0 100 100 group along with 2 canonical axes (Fig. 2). range of genetic distance for COI among small scaled Molecular Analysis: Capoeta species varied from 0.33 (between COI: Tables 5 and 6 list the diagnostic nucleotide C. damascina and C. pyragyi) to 4.53 (between substitutions and pairwise genetic distances C. caelestis and C. birunii). Of those, C. buhsei, (respectively) in the Capoeta damascina species C. coadi, C. saadii, C. shajariani, C. pyragyi, group for COI nucleotide sequences. The two C. ferdowsii and C. birunii are endemic to the Iranian different phylogenetic analyses including Bayesian basins. Capoeta umbla is the other species, which and Maximum Likelihood (Fig. 3) produced similar exists in the Tigris tributaries of Iran, , Turkey topologies. In both trees, members of the and . In these trees, samples of the C. coadi and C. damascina group form a monophyletic group C. birunii form a well-supported monophyletic including C. buhsei, C. caelestis, C. coadi, group, sister to C. buhsei. We have only a single COI C. damascina, C. ferdowsi, C. pyragyi, C. saadii, sequence of C. birunii because of experimental C. shajarianii, C. umbla and C. birunii sp. n. The problems, but the validity of this species by more

235 Iran. J. Ichthyol. (September 2017), 4(3): 231-269

Fig.2. Discriminant function scores for the morphological characters of Capoeta damascina species group (Wilk’lambda:0.001 for first function and 0.013 for second function; P<0.001).

Table 5. Diagnostic nucleotide substitutions in COI barcode region of the Capoeta damascina species group.

5636 5711 5717 5756 5774 5804 5810 5840 5849 5852 5861 5891 5909 5936 5966 5978 5996 6053 6080 6086

C. buhsei C A A T G G G T C G G A G A A C C G G A C. caelestis T G C T G A A G T G A A A C G T T G G G C. coadi C A A T A G G T C G G A G A A C C G A A C. damascina T A G T G A A G T A A A G C A T T A G G C. saadii T A G/A C G G G/A T C A A A G A A C C G T A C. umbla C/T A G T G A A G T A A G G C A T T G A G

C. shajariani T A A T G A A T T G A A G A A T T G G A shajarianiC. pyragyi T A G T G A A G T A A G G C A T T G G G

C. ferdowsii T A A T G G G T C G A A G A A C C G A A C. birunii C/T A A C G/A A A T C/T G A/G A G A A C/T C/T G A A

236 Zareian and Esmaeili - Phylogeny and taxonomy of Capoeta damascina species group in Iran

Table 6. Genetic distances between the Capoeta damascina species group based on COI gene.

C. C. buhsei C. caelestis C. coadi C. damascina C. saadii C. shajariani C. pyragyi C. ferdowsii C. buhsei C. caelestis 2.58 C. coadi 0.39 2.98 C. damascina 2.37 1.24 2.76 C. saadii 1.39 3.11 1.57 2.44 C. shajariani 1.34 1.58 1.72 1.37 1.90 C. pyragyi 2.32 1.19 2.71 0.33 2.39 1.32 C. ferdowsii 0.58 2.37 0.56 2.16 0.99 1.13 2.11 C. birunii 3.15 4.37 2.94 4.15 3.52 2.85 4.10 2.89 cytb sequences and also by morphologic diagnostic Using Haploview, 35 haplotypes were recovered characters. These three species were sister group to based on COI sequences of 111 individuals of the C. saadii and these four, sister to C. ferdowsii. Other C. damascina group (Fig. 4). Each species has its species (C. caelestis, C. damascina, C. umbla, own haplotypes. The haplotype network also C. shajariani, C. pyragyi and C. birunii) form a indicated a close relationship between C. damascina monophyletic clade. All of these species have close and C. umbla and also for C. buhsei and C. coadi. relationship (we do not have COI sequence from Cytb: Tables 7 and 8 list the diagnostic nucleotide C. kosswigi). Despite the intraspecific genetic substitutions and pairwise genetic distances diversity in C. umbla populations; based on these (respectively) in members of the C. damascina group trees, C. umbla specimens formed a mono-phyletic for cytb nucleotide sequences. The range of genetic clade and thus was treated as a distinct species. distance for cytb among members of the Results of Alwan et al. (2016b) also showed the C. damascina group varied from 0.56 (between intraspecific genetic diversity in C. damascina and C. kosswigi and C. damascina) to 3.87 (between C. umbla populations based on phylogenetic trees C. shajariani and C. birunii). The two different and haplotype networks. phylogenetic analyses include Bayesian and We examined large number of C. saadii specimens Maximum Likelihood (Fig. 5) and also produced from different drainage basins. Although individuals similar topologies based on cytb data set. In both of C. saadii were nested in several sub clades, but we trees, members of the C. damascina group form a did not find enough morphological diagnostic well-supported monophyletic group including characters or high genetic distance to consider them C. buhsei, C. caelestis, C. coadi, C. damascina, as different species. There are unresolved C. kosswigi, C. saadii, C. umbla, C. shajariani, phylogenetic relationships between C. tinca C. pyragyi, C. ferdowsii and C. birunii. In these trees, populations based on sequences which were obtained samples of the C. coadi and C. birunii form from Genebank. Sequences of this species were monophyletic group, sister to C. buhsei. As with the nested in the two different clades, probably because COI trees, these three species were sister group to these sequences belong to different species, due to C. saadii and all these sister to C. ferdowsii. Other misidentification. species (C. caelestis, C. damascina, C. kosswigi,

237 Iran. J. Ichthyol. (September 2017), 4(3): 231-269

Fig.3. Bayesian tree inferred from COI sequences. Numbers left of the slash indicate posterior probabilities; numbers right of the slash are the bootstrap support values for 10,000 replicates in Maximum Likelihood, using RaxML. Asterisks (*) indicate less than 50% Maximum Likelihood support for the node.

238 Zareian and Esmaeili - Phylogeny and taxonomy of Capoeta damascina species group in Iran

Table 7. Diagnostic nucleotide substitutions in COI barcode region of the Capoeta damascina species group.

14485 14506 14521 14542 14548 14554 14563 14596 14612 14623 14635 14641 14644 14665 14668 14669 14686 14701 14713 14752

C. buhsei A A C G T A A A G C T G A C T T C T A C C. caelestis A A T A C G A G G C T G A C T T T C A C C. coadi A A T G C A G A G C T A A C T T C C A C C. damascina A A T A C A A G A C T A A C T T C T A C C. kosswigi A A T A C G A G A C T A A C T T C T A C A C. saadii A A T G C A A / G C T A A T T T C T G C G C. umbla A A T A C G A/G G A C T A A C T T C T A C C. shajariani A A T A C A A A G C T G A C C A C T A T C. pyragyi A A T A C A A G A C T G A C T T C T A C C. ferdowsii A A T G C A A A G T C G C C T T C T A C C. birunii G G T G C G G A G C T A A C T T C T A C

14755 14759 14769 14773 14779 14785 14788 14833 14836 14842 14857 14884 14896 14905 14908 14938 14951 14952 14963 14986 15013

A/ C. buhsei A G A T G A A A C A T A T C A G C A C G G C. caelestis A A A C G A A A C A T A T A G A G C A C A C. coadi G G A T G A A A C A T A T G G A G C A C G C. damascina A A A T G G A A C A T G T A G A G C A C G C. kosswigi A A A T G A A A T A T G T A G A G C A C G C. saadii A/G A A T G A A/ A C A C A T G A A A C A C A/G G C. umbla A A A T G A A A T A T G T A G A G C A C G C. shajariani A A A T G G G A C A T A T A G A G T A C G C. pyragyi A A G T G A A G C A T G T A G A G C G C G C. ferdowsii A G A T A A A A C G T A T G G A G C A C G C. birunii G G G T G A A A C A T A T G G G G C G T G

15016 15021 15022 15028 15058 15061 15070 15076 15102 15118 15124 15130 15131 15136 15145 15154 15161 15188 15190 15193

C. buhsei G A A G C C G G C C A A T G G C A A A A C. caelestis G A A G C C G G T T A G T A A T A A A A C. coadi G A A G C C G G C C A A T G A C A A A A C. damascina A A A G C C G G C T A G T A A T A A A A C. kosswigi A A A G C C G G C T A G T A A T A C A A A/ A/ G A A G C T A A C T A A T A C A A A C. saadii G C C. umbla A A A G C C G G C T A G T A A T A C A A C. shajariani G A A G C C G G C C A A T G A C A A A A C. pyragyi G G G G A C G A C T A A C A A T G A A A C. ferdowsii G A A G C C G G C T G A T A A T A A G G A/ G A A A C C G G C T A A T A A C A A A C. birunii G

239 Iran. J. Ichthyol. (September 2017), 4(3): 231-269

Fig.4. Haplotype networks of Capoeta damascina species group based on of the COI. Each species is marked with different color and each circle represents one haplotype. Circle area is proportional to haplotype frequency.

Table 8. Genetic distance between the Capoeta damascina species group based on cytb gene.

C. C. coadi

C. C. saadii

C. C. buhsei

C. C. pyragyi

C. C. caelestis C. kosswigi C. ferdowsi

C. C. shajariani

C. C. damascina

C. buhsei C. caealestis 2.38 C. coadi 1.89 2.40 C. damascina 2.64 2.02 2.71 C. kosswigi 2.64 2.02 2.71 0.56 C. saadii 2.35 2.56 2.12 2.30 2.30 C. shajariani 2.44 2.38 3.07 2.07 2.68 2.76 C. pyragyi 2.69 2.02 3.33 1.72 1.72 2.87 2.68 C. ferdowsii 1.44 3.23 2.73 3.54 3.54 3.24 3.28 3.54 C. birunii 2.58 3.82 1.51 2.87 2.87 2.86 3.87 3.62 3.48

C. umbla and C. pyragyi) form a monophyletic individuals (Fig. 6). Each species has its own clade. Capoeta shajariani is sister group to all haplotypes. Cytb Haplotype network revealed a close C. damascina species group. Based on these trees, all relationship among C. damascina, C. kosswigi and studied species formed a monophyletic clade. Using C. umbla. These results revealed wide variation in Haploview, 50 haplotypes were recovered based on C. saadii populations. Unresolved phylogenetic cytb sequences of 119 small scaled Capoeta relationship was found between C. baliki andC. tinca 240 Zareian and Esmaeili - Phylogeny and taxonomy of Capoeta damascina species group in Iran

Fig.5. Bayesian tree inferred from cytb sequences. Numbers left of the slash indicate posterior probabilities; numbers right of the slash are bootstrap support values for 10,000 replicates in Maximum Likelihood, using RaxML. Asterisks (*) indicate less than 50% Maximum Likelihood support for the node.

241 Iran. J. Ichthyol. (September 2017), 4(3): 231-269

Fig.6. Haplotype networks of Capoeta damascina species group based on of the cytb gene. Each species is marked with different color and each circle represents one haplotype. Circle area is proportional to haplotype frequency. which requires further study based on more arch……………………………………………C. pyragyi molecular and morphological characters. - Less than 13 gill rakers on the lower limb of first gill Combined data set: The combined data set based tree arch……………………………………………………...4 was only performed for the Iranian C. damascina 4. Barbel length more than 19% of the head group, for which both cytb and COI sequences exist length…………………………………………...C. birunii - Barbel length less than 19% of the head length...C. coadi for each specimen. The Capoeta damascina group 5. Horizontal eye diameter more than 70% preorbital also form the well supported clade in this tree (Fig. length………………………………………..C. ferdowsii 7). Based on the tree there are also four full species - Horizontal eye diameter less than 70% preorbital within the former C. damascina species in the Iranian length...... 6 basins. 6. Barbel does not reach vertical of middle Key to Iranian C. damascina species group eye…………………………………………..C. shajariani 1. More than 18 scales above the lateral line…….C. umbla - Barbel pass vertical of middle eye…………………….7 - Less than 18 scales above the lateral line………………2 7. Less than 29 circumpeduncular scales (modally 25- 2. Deep depression anterior to nasal pore on the snout…..3 26)……………………………………………….C. saadii - No depression or shallow grove anterior to nasal pore on - More than 28 circumpeduncular scales……….C. buhsei the snout………………………………………………...5 Point distribution map of the Iranian Capoeta 3. More than 13 gill rakers on the lower limb of first gill damascina species group is shown in the Figure 8. 242 Zareian and Esmaeili - Phylogeny and taxonomy of Capoeta damascina species group in Iran

Fig.7. Bayesian tree inferred from combined data set. Numbers: posterior probabilities of the Bayesian Blue: Small scaled Capoeta; Green: C. trutta group and Red: C. capoeta group. 1982: 88, near Tehran, Djodje. Capoeta buhsei Kessler, 1877 Capoeta buhsei. – Coad 1995: 14, Namak Lake (Fig. 9) basin. Capoeta buhsei Kessler, 1877: 85, Persia (“iz Persii”, Capoeta buhsei. – Coad, 2008: 75-77, Namak apparently near Tehran). Lake basin (in the Damavand River, Jajrud, Karaj Varicorhinus nikolskii Derzhavin, 1929: 74-75, River and Dam, Sharra River, Kar River, Qareh Su, fig. 2, Karaj River near Tehran (“Keredsh flumen”). Abhar River and Qom River) and Kavir basin (in the Varicorhinus buhsei. – Berg 1949: 805, fig. 16, Hableh Rud and the Shurab near Semnan). vicinity of Tehran. Material examined: — ZM-CBSU Z218-229, 12, Capoeta buhsei. – Karaman, 1969: 36-37, figs 2, 104-149mm SL; Iran: Semnan Prov.: Hableh Rud at 3, plate 1 (fig. 2), plate 6 (fig. 5), vicinity of Tehran. Garmsar, Kavir basin, 35°18'06"N 52°24'57"E. — Varicorhinus damascinus (partim). – Saadati, ZM-CBSU Z260-274, 15, 88-130mm SL; Iran: 1977: 74-75, Namak Lake basin. Albourz Prov.: Kordan River at Karaj, Namak Lake Capoeta buhsei (partim). – Bianco & Banarescu, basin, 35°57'12"N 56°50'18"E.

243 Iran. J. Ichthyol. (September 2017), 4(3): 231-269

Fig.8. Point distribution map of small scaled Capoeta species in Iran.

Fig.9. Capoeta buhsei, Qom, Emamzadeh River, Namak Lake basin. Diagnosis: This species is distinguished from all profile convex; snout rounded; mouth inferior, large other species of the C. damascina group by the and a shallow horseshoe-shape, horny lower jaw following combination of characters: low number of layer weakly developed, lower lip corners in gill rakers on lower limb of first gill arch (8-10), particular fleshy and well-developed; rostral cap well number of total gill rakers on first gill arch (12–14) developed, in most specimens partly overlapping and depth of caudal peduncle in percent of standard upper lip; one posterior pair of barbels. Dorsal fin length (8.6–10.8). inserted anterior to pelvic-fin origin, its outer margin Description: See Figure 9 for general appearance. usually straight but sometimes concave or slightly Body usually elongate and cylindrical; greatest body concave with 3-4 unbranched and 8 branched rays, depth at level of dorsal fin origin; predorsal body last unbranched dorsal fin ray weakly ossified with profile smoothly convex to dorsal fin origin; nuchal thin and flexible distal part; pectoral fin not hump present in well-fed specimens; dorsal head extending to pelvic fin base, its tip rounded or 244 Zareian and Esmaeili - Phylogeny and taxonomy of Capoeta damascina species group in Iran

Fig.10. Qom, Emamzadeh River, Namak Lake basin, habitat of Capoeta buhsei. pointed, pectoral fin branched rays 15-18; pelvic fin Coloration: Overall colour is brownish in not extending to anal-fin base, its outer margins preservative without spots or any distinctive straight or slightly rounded, pelvic fin branched rays markings. The back is dark. The peritoneum is dark 7-10, pelvic axillary scale present; anal fin short not brown to black in preserved fish. reaching base of caudal fin, its outer margin convex Distribution: Namak Lake and Kavir basins (Figs. 8, or slightly convex, anal fin with 3 unbranched and 5 10). branched rays; caudal fin forked with 17 branched fin rays; scales small, lateral line scales 72- 89; total gill Capoeta coadi Alwan, Zareian & Esmaeili, 2016 rakers on first gill arch 12-14 (7-10 gill rakers on (Fig. 11). lower limb of the first gill arch). Material examined: Holotype. — ZM-CBSU Z190, According to Coad (2017) meristics are as 157mm SL; Iran: Kohgiluyeh and Boyer Ahmad follows: dorsal fin branched rays 8(35) or 9(3); anal Prov., Beshar River at Tale Gah village, Karun River fin branched rays 5(38); pectoral fin branched rays drainage, 30°47'27"N 51°25'13"E. — ZM-CBSU 14(1), 17(20), 18(11) or 19(6); pelvic fin branched Z191, 6, 91-157mm SL; all from Iran: Kohgiluyeh rays 7(1), 8(5), 9(31); lateral line scales 72(2), 73(1), and Boyer Ahmad Prov.: Beshar River at Tale Gah 75(3), 76(4), 77(2), 78(1), 79(7), 80(4), 81(5), 82(2), village, Karun River drainage, 30°47'27"N 83(2), 84(2), 86(2) or 91(1); total gill rakers 9(2), 51°25'13"E. ZM-CBSU J520, 1, 107mm SL; — ZM- 10(2), 11(4), 12(12), 13(11), 14(4), 15(2) or 17(1); CBSU Z275, 12, 105-152mm SL; Iran: Kohgiluyeh pharyngeal teeth 2,3,4,-4,3,2(7), 2,3,4-5,3,2(6), and Boyer Ahmad Prov.: Beshar River at Tale Gah 2,3,5-4,3,2(5), 2,3,5-5,3,2(1), 1,3,4-4,3,2 (1) or 2,3,4- village, Karun River drainage, 30°47'27"N 4,3,1(1); and total vertebrae 43(1) and 44(6) (USNM 51°25'13"E. — ZM-CBSU J526, 1, 98mm SL; — 20593 and the syntypes). ZM-CBSU J533, 1, 114mm SL; — ZM-CBSU J535,

245 Iran. J. Ichthyol. (September 2017), 4(3): 231-269

Fig.11. Capoeta coadi, Beshar River, Tigris River basin.

Fig.12. Beshar River at Taleh Gah village, Tigris River tributary, type locality of Capoeta coadi. 1, 97mm SL. — ZM-CBSU J540, 1, 67mm SL; all River, a tributary of Beshar River, north of Sepidan from Iran: Kohgiluyeh and Boyer Ahmad Prov.: city, Karun River drainage, 30°21.283'N Beshar River at Tange sorkh, Karun River drainage, 51°45.754'E. 30°26'14"N 51°45'48"E. — ZM-CBSU J444, 2, 73- Diagnosis: Capoeta coadi is distinguished from all 90mm SL; — ZM-CBSU J447, 2, 76-111mm SL; — other species of Capoeta by the following ZM-CBSU J450, 1, 86mm SL; ZM-CBSU J452, 1, combination of characters: last unbranched dorsal-fin 107mm SL; — ZM-CBSU J459, 2, 104-120mm SL; ray weakly to moderately ossified and serrated in 1/3- ZM-CBSU J464, 1, 110mm SL; all from Iran: Kohgi- 2/3 of its length; scales small, 70-84 total lateral line luyeh and Boyer Ahmad Prov.; Beshar River at scales, 12-17 scales between dorsal-fin origin and Mokhtar village, Karun River drainage, 30°40'31"N lateral line, 9-11 scales between anal-fin origin and 51°31'26"E. — ZM-CBSU 7880–7881, 2, 96.69– lateral line, 26-32 encircling least circumference of 158.12mm SL; Iran: Fars Prov.: Sepidan city, Gorgu caudal peduncle; total gill rakers 14-18, 10-13 gill

246 Zareian and Esmaeili - Phylogeny and taxonomy of Capoeta damascina species group in Iran

Fig.13. Capoeta ferdowsii from Doshmanzeyari, Doshmanzeyari River, Tigris River drainage. rakers on lower limb of first gill arch; 45-47 total 29˚59′16.30″N 51˚55′14.84″E; H. Zareian, vertebrae; one posterior pair of barbels. S. Ghasemian, R. Sadeghi & H. Darvishneia, 25 Dec. Description: See Figure 11 for general appearance. 2016. Body elongate and cylindrical; predorsal body profile Diagnosis: Capoeta ferdowsii is distinguished from smoothly convex with no marked discontinuity all other Capoeta species by the following between head and body except when a nuchal hump combination of characters: elongate and usually is present in few specimens; greatest body depth at cylindrical body; 7-9 branched dorsal-fin rays; last level of dorsal-fin origin. Dorsal-fin origin anterior to unbranched dorsal-fin ray weakly ossified; 7-8 total pelvic-fin origin, its outer margin usually straight to pelvic-fin rays; scales small, 71-80 scales in lateral- concave; pectoral fins not extending to pelvic fin line series; 13-17 scales between dorsal-fin origin base; their outer margins usually slightly convex; and lateral line, 8-11 scales between anal-fin origin pelvic fins not extending to anal fin base, their outer and lateral line (modally 9), 26-30 encircling least margin straight or slightly convex and blunt; pelvic circumference of caudal peduncle; 10-12 gill rakers axillary scale present; outer margin of anal fin on lower limb and 13-15 total gill rakers on first gill straight or slightly convex. arch; one pair of posterior barbels. Habitat and distribution: Capoeta coadi occurs in Description: Morphometric data are given in Table 9. medium-fast flowing rivers with usually gravel See Figures 13 and 14 for general appearance. Body substrates and clear waters. It is known only from the usually elongate and cylindrical; greatest body depth Karun River drainage, a system that constitutes the at level of dorsal fin origin; dorsal head profile southeastern part of the Tigris-Euphrates River convex; predorsal body profile smoothly convex to system (Figs. 8, 12). dorsal-fin origin; snout slightly rounded; mouth ventral; lower lip covered with a sharp-edged horny Capoeta ferdowsii Jouladeh-Roudbar, Eagderi, sheath. Murillo-Ramos, Ghanavi & Doadrio, 2017 Dorsal fin inserted anterior to pelvic-fin origin, its (Figs. 13-14) outer margin straight or concave, dorsal fin with 4-5 Capoeta damascina. - Esmaeili et al.: 2010: Tigris unbranched and 7-9 branched rays, last unbranched River drainage. dorsal-fin ray weakly ossified, flexible and soft at the Capoeta damascina. - Jouladeh-Roudbar et al.: tip, pectoral fins not extending to pelvic-fin base, its 2015: Tigris River drainage. outer margins usually slightly convex with 15-18 Capoeta saadii. - Esmaeili et al.: 2017: Tigris rays in total; pelvic fins not extending to anal fin River drainage. base, its outer margin slightly rounded to straight Material Examined: — ZM-CBSU Z700-708, 49- with 7-8 total rays, pelvic axillary scale present; anal 58mm SL; Iran: Fars Prov.: Doshmanzeyari River fin with 3 unbranched and 5 branched rays; caudal fin near Nourabad, Zohreh River drainage, forked, its tips pointed and its lobes not always of 247 Iran. J. Ichthyol. (September 2017), 4(3): 231-269

Fig.14. Capoeta ferdowsii, a) ZM-CBSU Z700; 58mm SL; b) ZM-CBSU Z701; 57mm SL; c) ZM-CBSU Z702; 50mm SL.

Fig.15. Doshmanzeyari River, habitat of Capoeta ferdowsii, Zohreh River basin. equal size with 16-18 branched rays; scales small, 71- between anal-fin origin and lateral line, 26-30 scales 83 scales in the lateral-line series, 13-17 scales encircling least circumference of caudal peduncle, between dorsal-fin origin and lateral line, 8-10 scales ventral midline and pectoral region covered with 248 Zareian and Esmaeili - Phylogeny and taxonomy of Capoeta damascina species group in Iran

Table 9. Morphometric characters of Capoeta ferdowsii in Zohreh basin; Doshmanzeyari River.

Character N Minimum Maximum Mean Std. Deviation In percent of standard length Head length 9 24.88 28.35 25.8740 1.09581 Maximum Head depth 9 17.33 20.10 18.2524 .86416 Head width 9 14.28 15.87 15.0067 .55809 Postorbital length 9 6.31 9.19 8.0892 .80345 Preorbital length 9 10.41 12.12 11.3668 .52500 Dorsal-fin base length 9 12.45 14.64 13.8593 .75297 Longest dorsal fin ray length 9 16.28 21.06 18.1936 1.49721 Predorsal length 9 49.08 53.56 51.5197 1.35301 Postdorsal length 9 54.72 59.05 56.3499 1.28205 Pectoral fin length 9 15.71 19.72 17.3320 1.12113 Pelvic fin length 9 13.73 18.21 14.7496 1.36906 Pelvic fin base length 9 4.75 6.10 5.3072 .39134 Prepelvic length 9 51.71 55.96 53.3261 1.36686 Anal fin base length 9 7.45 9.72 8.2512 .68233 Preanal length 9 72.29 76.54 73.6381 1.44113 Anal fin length 9 14.77 17.89 15.8016 .94345 Length of caudal peduncle 9 17.54 21.84 19.1192 1.59924 Depth of caudal peduncle 9 9.53 11.54 10.4986 .64771 Body depth at dorsal-fin origin 9 9.90 21.55 19.2029 3.59632 In percent of head length Maximum Head depth 9 67.44 73.05 70.5485 1.74439 Head width 9 55.48 61.75 58.0432 2.14035 Postorbital length 9 39.92 46.31 43.9695 2.12386 Preorbital length 9 25.22 34.40 31.2767 2.99530 Interorbital distance 9 34.28 46.36 39.1437 3.77473 Eye diameter 9 23.16 27.49 25.2023 1.67871 Intranasal distance 9 23.52 27.93 26.1616 1.35809 Mouth width 9 28.27 34.76 32.1692 2.01823 Barbel length 9 13.29 28.27 17.7897 4.26271 deeply embedded scales of reduced size; 10-12 gill C. umbla by number of scales (ALL: 13-17 vs. 18- rakers on lower limb of first gill arch and 13-16 gill 25) and number of gill rakers on the firs gill arch (13- rakers in total. 16 vs. 17-20). Capoeta ferdowsii is distinguished Capoeta ferdowsii is distinguished from C. buhsei from C. shajariani by head depth at eye in percent of by number of gill rakers on the lower limb of the first postorbital length (90-102 vs. 105-126). Capoeta gill arch (10-12 vs. 7-10) and postrorbital in percent ferdowsii is distinguished from C. pyragyi by number of head length (39.9-46.3 vs. 47.6-60.6). Capoeta of gill rakers on lower limb of first gill arch (10-12 ferdowsii is distinguished from C. coadi by vs. 13-15). Capoeta ferdowsii is distinguished from horizontal eye diameter in percent of preorbital C. birunii by postorbital in percent of head length length (70.21-84.91 vs. 39.55-64.63). Capoeta (39.9-46.3 vs. 46.9-51.8), horizontal eye diameter in ferdowsii is distinguished from C. saadii by percent of periorbital length (70.2-84.9 vs. 43.1-61.0) postrorbital in percent of head length (39.92-46.31 and head depth at eye in percent of postorbital (106- vs. 46.58-54.84) and horizontal eye diameter in 132 vs. 90-102). percent of preorbital length (70.21-84.91 vs. 33.37- Coloration: Dorsum, head and sides above the lateral 68.79). Capoeta ferdowsii is distinguished from line in the C. ferdowsii from the Doshmanzeyari

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Fig.16. Capoeta pyragyi from the Tireh River, Doroud, Tigris River drainage. River are brown, below the lateral line white (Fig. of characters: elongate and usually cylindrical body; 13); all fins mostly colourless; base of dorsal, pelvic 4-5 unbranched (modally 5) and 8-9 (modally 9) and pectoral fins orange. Some specimens have a branched dorsal-fin rays; last unbranched dorsal-fin diffuse lateral band along the sides and small diffuse ray weakly ossified; 8-9 total pelvic-fin rays; small black spots above the lateral line. scales, 69-79 scales in the lateral-line series, 12-16 Habitat and distribution: Capoeta ferdowsii is found scales between dorsal-fin origin and lateral line, 7-10 in medium to fast flowing rivers with muddy bottoms scales between anal-fin origin and lateral line and cold water. This species inhabits the Zohreh (modally 8 and 9), 27-29 encircling least River basin in Iran (Figs. 8, 15). circumference of caudal peduncle; 13-15 gill rakers on lower limb and 16-19 total gill rakers on first gill Capoeta pyragyi Jouladeh-Roudbar, Eagderi, arch; one pair of barbels. Murillo-Ramos, Ghanavi & Doadrio, 2017 Description: Morphometric data are given in Table (Figs. 16-17) 10. See Figures 16 and 17 for general appearance. Capoeta damascina. - Esmaeili et al.: 2010, Tigris Body usually elongate and cylindrical; greatest body River basin. depth at level of dorsal fin origin; dorsal head profile Capoeta damascina. - Jouladeh-Roudbar et al.: straight or convex; predorsal body profile straight or 2015: Tigris River drainage. smoothly convex to dorsal-fin origin, snout usually Capoeta saadii. - Esmaeili et al.: 2017: Tigris pointed; deep depression anterior the nasal pore on River drainage the snout; mouth ventral; lower lip covered with a Material Examined: — ZM-CBSU Z750-757,48- sharp-edged horny sheath; one pair of barbels. Dorsal 72mm SL; Iran: Lorestan Prov.: Sezar River near fin inserted anterior to pelvic-fin origin, its outer Doroud, Tigris River basin, 29˚59′16.30″N margin usually straight or concave with 4-5 51˚55′14.84″E; A. Gholamhosseini, R. Khaefi, unbranched and 8-9 branched rays, last unbranched H. Darvishnia, R. Sadeghi, H. Mehraban & dorsal-fin ray weakly ossified, flexible and soft at the M. Razbanian, 20 Aug. 2015. — ZM-CBSU Z760- tip; pectoral fin short not extending to pelvic fin base 767, 95-155mm SL; Iran: Lorestan Prov.: Tireh River outer margins usually slightly convex with 16-19 near Doroud, Tigris River basin, 33˚27′59.4″N rays in total; pelvic fin not extending to anal-fin base, 49˚04′06.2″E; A. Gholamhosseini, R. Khaefi, its outer margin slightly rounded to straight with 8-9 H. Darvishnia, R. Sadeghi, H. Mehraban & rays in total; pelvic axillary scale present; anal fin M. Razbanian, 20 Aug. 2015. with 3 unbranched and 5 branched rays; caudal fin Diagnosis: Capoeta pyragyi is distinguished from all forked, its tips pointed with 16-19 branched rays; other Capoeta species by the following combination scales small, 63-79 scales in the lateral-line series, 250 Zareian and Esmaeili - Phylogeny and taxonomy of Capoeta damascina species group in Iran

Fig.17. Capoeta pyragyi, a) ZM-CBSU Z750; 72mm SL; b) ZM-CBSU Z751; 70mm SL; c) ZM-CBSU Z752; 64mm SL.

Fig.18. Tireh River, habitat of Capoeta pyragyi, the Tigris River drainage.

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Table 10. Morphometric characters of Capoeta pyragyi in the Tigris River drainage; Tireh and Sezar Rivers.

Character N Minimum Maximum Mean Std. Deviation In percent of standard length Head length 16 24.45 26.31 25.3419 .55892 Maximum Head depth 16 16.63 25.70 18.2414 2.15746 Head width 16 14.71 18.01 15.6623 .88727 Postorbital length 16 5.05 10.30 9.0423 1.17736 Preorbital length 16 10.74 15.91 12.6664 1.25430 Dorsal fin base length 16 13.04 16.19 14.4473 .88534 Longest dorsal fin ray length 16 18.32 21.64 19.8421 .83667 Predorsal length 16 48.01 52.51 50.7118 1.29211 Postdorsal length 16 53.96 61.46 57.3108 2.41510 Pectoral fin length 16 15.94 20.26 18.3297 1.12949 Pelvic fin length 16 13.34 17.14 15.4127 1.10176 Pelvic fin base length 16 4.22 6.19 5.0012 .47807 Prepelvic length 16 49.23 57.31 53.7692 1.76983 Anal fin base length 16 7.14 8.23 7.6129 .36704 Preanal length 16 71.55 76.25 73.3946 1.09215 Anal fin length 16 14.60 18.19 16.3146 .90151 Length of caudal peduncle 16 16.78 21.20 19.7929 1.17388 Depth of caudal peduncle 16 10.57 12.25 11.2703 .47408 Body depth at dorsal-fin origin 16 21.09 25.34 23.4426 1.29295 In percent of head length Maximum Head depth 16 64.62 100.04 71.9934 8.37580 Head width 16 56.66 70.10 61.8163 3.43026 Postorbital length 16 41.02 61.91 49.9792 4.74372 Preorbital length 16 19.28 39.66 35.7170 4.74588 Interorbital distance 16 25.79 43.62 38.7544 4.10139 Eye diameter 16 13.96 59.11 22.0521 10.38563 Intranasal distance 16 18.77 27.97 24.2398 2.62545 Mouth width 16 30.73 38.01 35.0518 1.97708 Barbel length 16 12.44 20.21 16.3547 1.93766

12-16 scales between dorsal-fin origin and lateral depression anterior to nasal pore on snout in line, 7-10 scales between anal-fin origin and lateral C. pyragyi vs. absence or shallow grove in the line, 27-29 scales encircling least circumference of C. shajariani and C. saadii). Capoeta pyragyi is caudal peduncle, ventral midline and pectoral region distinguished from C. umbla by number of scales covered with deeply embedded scales of reduced (LL: 69-81 vs. 82-100; ALL: 12-16 vs. 18-25). size; gill rakers slightly hooked; 13-15 gill rakers on Capoeta pyragyi is distinguished from C. ferdowsii lower limb of first gill arch. and C. birunii by number of gill rakers on lower limb Capoeta pyragyi is distinguished from C. buhsei of first gill arch (13-15 vs. 10-12 and 10-13, by higher number of total gill rakers (16-19 vs. 12- respectively). 14) and higher number of gill rakers on lower limb of Coloration: Dorsum dark olive, sides golden-yellow the first gill arch (13-15 vs. 7-10). Capoeta pyragyi is or golden-orange along the lateral line and white distinguished from C. coadi by number of gill rakers cream belly (Fig. 16). Dorsal view of head olive and on lower limb of first gill arch (13-15 vs. 10-13). operculums light orange. Dorsal, anal and caudal fins Capoeta pyragyi is distinguished from C. saadii and dark yellow with dark rays; pectoral and pelvic fins C. shajariani by shape of snout (presence of a deep golden-yellow. Few dark blotches are present on the

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body of some specimens. Capoeta saadi. – Bianco & Banarescu 1982: 88, Habitat and distribution: Capoeta pyragyi is found in Pulwar near Persepolis, Mand River near Akbar and cold and slow flowing rivers with rock and stone near Dasht-e-Arzhan, springs of Kul River basin near bottoms and gravel substrate and shade of trees. Darab (Persepolis). Capoeta pyragyi inhabits the Tireh and Sezar Rivers Capoeta buhsei (partim). – Bianco & Banarescu in the Tigris River drainage, Iran (Figs. 8, 18). 1982: 88, springs of Kul River basin near Darab, vicinity of Kerman. Capoeta saadii (Heckel, 1849) Capoeta damascina (partim). – Coad 1995: 14, (Fig. 19) Kor River, Lake Maharlu, Sirjan, Kerman-Na’in and Scaphiodon saadii Heckel, 1849: 260, sources of Hormuz basins (listed, distribution). Saadi at Schiraz (Shiraz), near Persepolis (“Quellen Capoeta saadii Esmaeili et al. 2017:40. Kor, des Saadi”, possibly Kor River). Esfahan, Persis, Tigris, Maharlu, Sirjan, Kerman- Scaphiodon amir Heckel, 1849: 258, Araxes. Na’in and Hormuz. Scaphiodon niger Heckel, 1849: 258, Araxes or Material examined: — ZM-CBSU Z136-146, 11, 78- Benth-Amir. 121mm SL; ZM-CBSU 2504, 1, 82mm SL; — ZM- Scaphiodon chebisiensis Keyserling, 1861: 5, CBSU 2508, 1, 69 mm SL; ZM-CBSU 2520-2521, 2, plate 2, canal in Chebis (“Wasserleitung in Chebis”). 51-62 mm SL; — ZM-CBSU 2524-2528, 5, 113- Scaphiodon rostratus Keyserling, 1861: 7, plate 3, 231mm SL; Iran: Fars Prov.: Ghadamgah spring, canal in the vicinity of Jezd, Meibut Doroodzan, 30°15'11"N 54°25'32"E. (“Wasserleitungen in der Umgegend von Jezd”). Diagnosis: Capoeta saadii is distinguished from all Capoeta chebisiensis. – De Filippi 1865: 359 other Capoeta species by the following combination Capoeta saadi. – De Filippi 1865: 359, Shiraz. of characters: elongate and usually cylindrical body; Capoeta amir. – Günther 1868: 79, Araxes River. 8-9 branched dorsal fin rays; last unbranched dorsal- Scaphiodon niger. – Günther 1868: 79, Araxes. fin ray weakly ossified and serrated in 1/3-3/4 of its Capoëta chebisiensis. – Tortonese 1934: 6, Persia. length; 8-10 total pelvic fin rays (modally 9); scales Capoëta saadii. – Tortonese 1934: 6, Shiraz. small, 63-79 scales in the lateral-line series,11-16 Varicorhinus damascinus (partim). – Berg 1949: scales between dorsal-fin origin and lateral line 800-803, Shiraz, Kerman. (modally 14 and 15), 7-10 scales between anal-fin Varicorhinus damascinus (partim). – Ladiges origin and lateral line (modally 8 and 9), 23-28 1960: 130, Persepolis. encircling least circumference of caudal peduncle Capoeta amir. – Kähsbauer 1963: 328, Araxes in (modally 24 and 26); 9-15 gill rakers on lower limb Persepolis, Chun-i-Kaka (Sargado, possibly Sarhad of first gill arch (modally 10-11); 42-46 total region in eastern Kerman). vertebrae (modally 43-44); light golden-yellowish or Capoeta chebisiensis. – Kähsbauer 1963: 328, silvery-greyish body coloration in life; presence of a canal in Chebis. few black blotches on the body of some specimens. Capoeta niger. – Kähsbauer 1963: 329, Araxes in Description: See Figure 19 for general appearance. Persepolis. Body usually elongate and cylindrical; greatest body Capoeta rostratus. – Kähsbauer 1963: 329, canal depth at level of dorsal fin origin; dorsal head profile in Meibut near Jezd (Jazd). straight (very pronounced in specimens from the Kor Capoeta saadii. – Kähsbauer 1963: 329, Saadi River basin) or convex; predorsal body profile River in Shiraz, Araxes River in Persepolis. smoothly convex to dorsal-fin origin, slightly convex Varicorhinus damascinus (partim). – Saadati in some specimens, nuchal hump present in well-fed 1977: 74, 192, 193, Jazd, Maharlu, Sirjan. specimens; snout usually pointed; mouth ventral; lips 253 Iran. J. Ichthyol. (September 2017), 4(3): 231-269

Fig.19. Capoeta saadii, a) Kor River, b) Malousjan Spring; Kor River basin. a little fleshy, being more pronounced at mouth branched rays; caudal fin forked, its tips pointed and corners; lower lip covered with a sharp-edged horny its lobes not always of equal size with 16-18 sheath (sometimes not very well developed), its branched rays. Scales small, 63-79 scales in lateral- anterior margin straight in adult specimens and line series; 11-16 scales between dorsal-fin origin rounded to almost crescent-shaped in juveniles, with and lateral line; 7-10 scales between anal-fin origin a considerable degree of individual variation; rostral and lateral line; 23-28 scales encircling least cap well developed, partly overlapping upper lip in circumference of caudal peduncle; ventral midline most specimens; one posterior pair of barbels. Dorsal and pectoral region covered with deeply embedded fin inserted anterior to pelvic-fin origin, its outer scales of reduced size; gill rakers slightly hooked; 9- margin usually straight but occasionally concave 15 gill rakers on the lower limb of first gill arch. with 3-6 unbranched and 8-9 branched rays, last Coloration: Live specimens: dorsum, head and sides unbranched dorsal-fin ray weakly ossified, flexible of C. saadii from the Kor River basin golden-yellow, and soft at the tip, serrated in 1/3-2/3 of its length in darker dorsally and lighter below the lateral line (Fig. adults and subadults and 1/2-3/4 in juveniles; 19); dorsal, anal and caudal fins light golden; pectoral pectoral fin not extending to pelvic-fin base, its outer and pelvic fins golden-yellow. Upper half of body of margin usually slightly convex with 17-20 rays in C. saadii from other river drainages light golden or total; pelvic fin not extending to anal-fin base, its silvery-grey and lower half white with or without outer margins slightly rounded to straight with 8-10 grey tinge; fins white-yellowish or dusty grey. Few total rays; pelvic axillary scale present; anal fin not dark blotches are present on the body of some adult, reaching caudal-fin base with 3 unbranched and 5 subadult and juvenile specimens. In addition,

254 Zareian and Esmaeili - Phylogeny and taxonomy of Capoeta damascina species group in Iran

Fig.20. Kor River, habitat of Capoeta saadii, Kor River basin. juvenile specimens have a diffuse lateral stripe along Material examined: — ZM-CBSU Z800-808, 73- the sides and small diffuse black spots above the 109mm SL; Iran: Kermanshah Prov.: Gamasiyab lateral line. Preserved specimens: dorsum, head and River at Darakeh, Tigris River basin, 34˚23′31.0″N sides grey dorsally and beige or yellow ventrally; fins 47˚42′57.5″E; A. Gholamhosseini, R. Khaefi, beige or dusty grey; dark lateral band (in juveniles), H. Darvishnia, R. Sadeghi, H. Mehraban & spots (in juveniles) and blotches well discernible. M. Razbanian, 30 Aug. 2015. Habitat and Distribution: Capoeta saadii is found in Diagnosis: Capoeta shajariani is distinguished from slow to medium flowing rivers with muddy bottoms all other Capoeta species by the following and gravel substrates. Capoeta saadii inhabits the combination of characters: elongate and usually Kor, Persis, Hormuz, Kerman and Maharlu Lake cylindrical body; 8-9, modally 9, branched dorsal-fin basins in Iran (Figs. 8, 20). rays; 8-9 total pelvic-fin rays (modally 8); small scales, 13-14 above the lateral line (modally 14), 8-9 Capoeta shajariani Jouladeh-Roudbar, Eagderi, below the lateral line, 25-31 encircling least Murillo-Ramos, Ghanavi & Doadrio, 2017 circumference of caudal peduncle, 71-80 scales in the (Figs. 21-22) lateral-line series; 11-14 gill rakers on lower limb of Capoeta damascina. - Coad: 2017. first gill arch (modally 12) and 15-18 total gill rakers Capoeta damascina. - Samaee & Patzner: 2011, on the first gill arch; one posterior pair of barbels. Gamasiyab. Description: Morphometric data are given in Table Capoeta damascina. - Esmaeili et al.: 2010, Tigris 11. See Figures 21 and 22 for general appearance. River basin. Body usually elongate and cylindrical; greatest body Capoeta damascina. - Jouladeh-Roudbar et al.: depth at level of dorsal fin origin; dorsal head profile 2015: Tigris River basin. straight or convex; predorsal body profile smoothly Capoeta saadii. – Esmaeili et al.: 2017: Tigris convex to dorsal-fin origin; snout usually pointed; River. mouth ventral; lower lip covered with a sharp-edged 255 Iran. J. Ichthyol. (September 2017), 4(3): 231-269

Fig.21. Capoeta shajariani; a) ZM-CBSU Z800; 102mm SL; b) ZM-CBSU Z801; 95mm SL; c) ZM-CBSU Z802; 73mm SL.

Fig.22. Capoeta shajariani from Gamasyiab River, Darakeh, Tigris River drainage. horny sheath, rostral cap well developed, partly (modally 9) branched rays, last unbranched dorsal-fin overlapping upper lip in most specimens; one pairs of ray weakly ossified, flexible and soft at the tip; posterior barbels. Dorsal fin inserted anterior to pectoral fin short and not extending to pelvic-fin pelvic-fin origin, its outer margin usually straight but base, its outer margin usually slightly convex with occasionally concave with 4 unbranched and 8-9 16-18 pectoral-fin rays in total; pelvic fins not

256 Zareian and Esmaeili - Phylogeny and taxonomy of Capoeta damascina species group in Iran

Table 11. Morphometric characters of Capoeta shajariani in the Tigris River drainage; Gamasyiab River.

Character N Minimum Maximum Mean Std. Deviation In percent of standard length Head length 9 23.82 26.54 25.5221 0.90088 Maximum Head depth 9 16.58 18.63 17.3966 0.63626 Head width 9 14.16 15.75 14.8414 0.58811 Postorbital length 9 8.49 10.40 9.1804 0.56927 Preorbital length 9 11.11 12.21 11.6238 0.35816 Dorsal-fin base length 9 13.69 16.02 14.8018 0.72076 Longest dorsal fin ray length 9 18.58 21.94 20.3918 0.94828 Predorsal length 9 49.01 51.81 50.3010 .88831 Postdorsal length 9 56.71 62.43 59.0954 1.58613 Pectoral fin length 9 17.34 19.34 18.4967 0.68756 Pelvic fin length 9 15.01 17.17 16.0594 0.63766 Pelvic fin base length 9 4.37 5.73 4.8416 0.41739 Prepelvic length 9 52.38 55.79 53.8050 1.24606 Anal fin base length 9 7.11 10.53 8.1038 1.04827 Preanal length 9 70.53 75.63 73.6299 1.53081 Anal fin length 9 15.65 18.62 17.3255 0.93747 Length of caudal peduncle 9 17.47 20.34 18.7302 1.08973 Depth of caudal peduncle 9 9.71 11.01 10.6448 0.39633 Body depth at dorsal-fin origin 9 21.92 26.10 23.2567 1.21763 In percent of head length Maximum Head depth 9 64.79 72.02 68.2015 2.49229 Head width 9 53.57 60.97 58.2086 2.87991 Postorbital length 9 43.70 46.83 45.5650 1.21498 Preorbital length 9 32.24 39.98 35.9880 2.13829 Interorbital distance 9 37.87 44.05 39.7216 1.73838 Eye diameter 9 18.84 23.99 21.1676 1.54110 Intranasal distance 9 23.49 25.98 24.9368 .74755 Mouth width 9 29.29 38.10 32.9250 2.68655 Barbel length 9 13.72 17.60 15.7116 1.11948 extending to anal-fin base, its outer margin slightly distinguished from C. coadi by number of scales rounded to straight with 8-9 rays in total; anal fin with between anal fin origin and lateral line (8-9 vs. 9-12). 3 unbranched and 5 branched rays; caudal fin forked, Capoeta shajariani is distinguished from C. saadii by its tips pointed with 17 branched rays; scales small, size of barbels (reach to anterior margin of eye vs. 71-80 scales in lateral-line series, 13-14 scales pass half of horizontal eye diameter). Capoeta between dorsal-fin origin and lateral line; 8-9 scales shajariani is distinguished from C. umbla by number between anal-fin origin and lateral line, 25-31 of scales (LL: 71-80 vs. 82-100; ALL: 13-14 vs. 18- encircling least circumference of caudal peduncle; 25; BLL: 8-9 vs. 10-14) and number of gill rakers on 11-14 gill rakers on lower limb and 15-18 total gill lower limb of first gill arch (11-14 vs. 15-16). rakers on first gill arch. Capoeta shajariani is Capoeta shajariani is distinguished from C. pyragyi distinguished from C. buhsei by number of gill rakers by shape of snout (vs. absence or shallow grove in (14-18 vs. 12-14) and gill rakers on lower limb of C. shajariani vs. presence of a deep depression first gill arch (11-14 vs. 7-10). Capoeta shajariani is anterior to the nasal pore on snout in C. pyragyi). 257 Iran. J. Ichthyol. (September 2017), 4(3): 231-269

Fig.23. Gamasyiab River, habitat of Capoeta shajariani in Darakeh, Tigris River drainage. Capoeta shajariani is distinguished from C. ferdowsi Capoeta umbla. – Günther 1868: 79, Tigris River. by head depth at eye in percent of postorbital length Capoeta umbla. – Sauvage 1884: 5, 16, Tigris, (105-126 vs. 90-102). Capoeta shajariani is Biredjik (Euphrates). distinguished from C. birunii by barbel length in Varicorhinus umbla. – Kosswig & Battalgil 1943: percent of mouth width (39-55 vs. 57-79). 24, Mossul, Şitak, Batman suyu, Hazer (Hazar) Gölü. Coloration: Dorsum, head and sides of C. shajariani Varicorhinus umbla. – Berg 1949: 379, fig. 17, dark olive, darker dorsally and lighter below the Tigris basin from Mosul to the lower reaches. lateral line (Fig. 22); silvery sides and white belly; Varicorhinus umbla. – Ladiges 1960: 132 (listed). dorsal fin dark olive in the base and lighter in the Varicorhinus damascinus umbla. – Khalaf 1961: margin; pectoral and pelvic fins dark yellow; few 34, Iraqi waters. dark blotches are present on the body of some small Varicorhinus umbla. – Beckman 1962: 149, Tigris specimens. River. Habitat and distribution: Capoeta shajariani can be Capoeta capoeta umbla. – Karaman 1969: 32, found in slow to medium flowing rivers with gravel upper reaches of the Tigris-Euphrates system. substrates and riparian vegetation (reed). Capoeta Capoeta capoeta umbla. – Kuru 1971: 139, shajariani inhabits the Gamasiyab River (Figs. 8, 23) Euphrates, Murat suyu (Murat Nehri), Karasu. in the Tigris River drainage. ? Varicorhinus damascinus (partim). – Saadati 1977: 74, 192-193, Tigris basin in Iran. Capoeta umbla (Heckel, 1843) Varicorhinus umbla. – Saadati 1977: 76, 194, (Figs. 24-25) Tigris River system. Scaphiodon umbla Heckel, 1843: 1060, plate 5, fig. Capoeta capoeta umbla. – Coad 1980: 91, upper 3, Tigris River in Mossul (“Tigris bei Mossul”). Tigris-Euphrates system.

258 Zareian and Esmaeili - Phylogeny and taxonomy of Capoeta damascina species group in Iran

Fig.24. Capoeta umbla, syntype, NMW 55933, 177.4mm SL.

Fig.25. Live specimens of Capoeta umbla, Iran: Kurdistan, Garran River, Tigris River drainage. Varicorhinus umbla. – Banister 1980: 103, Tigris unbranched dorsal-fin ray weakly to moderately and Euphrates. ossified and serrated in 1/2-3/4 of its length; very Capoeta capoeta umbla. – Kuru 1980: 494, Tigris small scales, 18-25 scales between dorsal-fin origin and Euphrates. and lateral line, 10-15 scales between anal-fin origin Capoeta capoeta umbla. – Bianco & Banarescu and the lateral line, 31-39 encircling least 1982: 87, Tigris-Euphrates drainage circumference of the caudal peduncle, 82-104 total Capoeta damascina (partim). – Coad 1991: 15, lateral line scales; 12-16 gill rakers on lower limb of Tigris-Euphrates basin (listed). first gill arch; 17-20 total gill rakers on first gill arch; Capoeta umbla. – Banarescu 1999: 386, Tigris, 46-50 total vertebrae; one posterior pair of barbels. Euphrates. Description: General morphology of C. umbla is Capoeta capoeta umbla. – Kiliç demirok & Ünlü shown in Figures 24-25. Body elongate and slightly 2001: 389-393, Tigris River (biology). compressed; greatest body depth at level of dorsal-fin Capoeta capoeta umbla. – Türkmen et al. 2002: origin; pre-dorsal body profile smoothly convex to 317-328, Aşkale region of the Karasu River dorsal- fin origin; nuchal hump present in well-fed (biology). specimens; snout pointed; mouth ventral; lower lip Capoeta umbla (partim). – schöter et al. 2009: 234, covered with a sharp-edged horny sheath and its Tigris-Euphrates system. anterior margin straight or rounded to almost Diagnosis: Capoeta umbla is distinguished from all crescent-shaped. Dorsal fin inserted anterior to other Capoeta species by the combination of the pelvic-fin origin and its outer margin usually following characters: very elongate and slightly concave; 4-6 unbranched and 8-10 branched dorsal- compressed body; 8-10 branched dorsal-fin rays; last fin rays; last unbranched dorsal-fin ray weakly to 259 Iran. J. Ichthyol. (September 2017), 4(3): 231-269

Fig.26. Garan River; habitat of Capoeta umbla in Marivan, Tigris River drainage. moderately ossified, soft and flexible at the tip, (Persian Gulf basin), in Iran, Iraq, Syria and Turkey serrated in 1/2-3/4 of its length; pectoral fins not (Fig. 8). It inhabits a wide range of water bodies from extending to pelvic-fin base, outer margins usually small headwater streams (Fig. 26) down to large slightly convex; pelvic fins not extending to anal fin lowland rivers, marshes, lakes and reservoirs (see base, outer margins straight or slightly rounded; Freyhof 2014). pelvic axillary scale present; anal fin with 3 Material examined: unbranched and 5 branched rays, outer margin Syntypes: — NMW 55932, 1, 183.55mm SL, Iraq: straight or slightly convex; caudal fin forked and its Tigris at Mosul. — NMW 55933, 1, 177.45mm SL, tips pointed; 17-18 branched caudal-fin rays. Scales Iraq: Tigris at Mosul. — NMW 79373, 1, ca. 263mm very small, 86-104 scales in the lateral-line series, SL, Iraq: Mosul. — NMW 79374 (Dry). 18-24 between dorsal-fin origin and lateral line, 11.5- Other material examined: — NMW 90541, (1) 1, 15.5 between lateral line and anal-fin origin, 32-39 175.81mm SL, Turkey: Palu/Murat (source of encircling least circumference of caudal peduncle, Euphrates), ca. 38°43'N 39°56'E, V. Pietschmann. — ventral midline and pectoral region covered with FSJF 1494 (in 96% alcohol), (2) 1, 42.43mm SL; deeply embedded scales of reduced size; gill rakers Turkey: outflow of Hazar Gölü at Plajköy, slightly hooked, 15-16 gill rakers on the lower limb 38°30.187'N 39°30.423' E, 24. IX. 2008, M. Özuluğ of the first gill arch; 46-50 total vertebrae; one & J. Freyhof. — NMW 90543, (3) 1, bent specimen, posterior pair of barbels; pharyngeal teeth arranged Turkey: Hazar Gölü, ca. 38°30'N 39°30'E, in three rows in the following manner: 2.3.5-5.3.2 22/23.VII.1914, V. Pietschmann. — NMW 91066, (f1) and very similar in shape to those of (4) 2, 212.22-220.14mm SL, Turkey: Kürk at Hazar C. damascina (see Alwan 2011). Gölü, draining to Tigris, ca. 38°30'N 39°30'E, V. Habitat and Distribution: Capoeta umbla is Pietschmann, 1914. — NMW 91067, (5) 6, 156.73- distributed in the Tigris and Euphrates River system 216.18mm SL; Turkey: Kürk at Hazar Gölü, draining 260 Zareian and Esmaeili - Phylogeny and taxonomy of Capoeta damascina species group in Iran

to Tigris, ca. 38°30'N 39°30'E, V. Pietschmann. — Esfahan Prov.: Daran River near Daran, NMW 91069, (6) 3, 153.47-228.75mm SL; Turkey: Zayandehrud basin, 32˚49′25.8″N 50˚25′47.4″E; Kürk at Hazar Gölü, draining to Tigris, ca. 38°30'N H.R. Esmaeili, R. Zammaniannejad, G. Sayyadzadeh 39°30'E, V. Pietschmann. — NMW 90543, (7) 3, & S. Ghasemian, 16 Aug 201. bent specimen with damaged snout-ca. 227.28mm Paratypes: ZM-CBSU Z651-660, 10, 90-165mm SL; SL; Turkey: Wadi Mahmedian Çay, (source of Same data as holotype. Tigris), ca. 38°20'N 40°45'E, 1914, V. Pietschmann. Diagnosis: Morphometric data are given in Table 12. — NMW 91062, (8) 2, bent specimen with damaged Capoeta birunii is distinguished from all other snout-224.04mm SL; Turkey: Wadi Mahmedian Capoeta species by the following combination of Çay, (source of Tigris), ca. 38°20'N 40°45'E, 1914, characters: elongate and usually cylindrical body; 7- V. Pietschmann. — FSJF 1425 (in 96 % alcohol), (9) 8 branched dorsal-fin rays; 8-10 total pelvic-fin rays; Fin clip, Turkey: Tigris River, 5km east of Bismil, small scales, 68-82 total lateral line scales, 13-15 37°50.314'N 40°41.620'E, 25.IX.2008, M. Özuluğ & scales between dorsal-fin origin and lateral line, 7-11 J. Freyhof, specimen identified by J. Freyhof. — scale between anal-fin origin and lateral line, 25-29 NMW 90536, (10) 2, 123.26-141.95mm SL; Turkey: scales encircling least circumference of caudal Batman (a tributary to Tigris), ca. 37°50'N 41°00'E, peduncle; total gill rakers on first gill arch 13-17 and V. Pietschmann. — NMW 91078, (11) 2, 132.67- 10-13 gill rakers on lower limb. 152.50mm SL; Syria: Jerablus at Euphrates, ca. Description: See Figures 27-28 for general 36°49'N 38°02'E, V. Pietschmann. — BMNH appearance and Table 12 for morphometric data. 1974.2.22.1390. (12) 1, bent specimen (ca. 100mm Body usually elongate and cylindrical; predorsal SL), Iraq: Nahr Rawanduz (Great Zab), N. Sarsam. body profile smoothly convex; greatest body depth at — BMNH 1974.2.22.1392-1393. (13) 2, bent level of dorsal-fin origin; dorsal head profile straight; specimens (ca. 178-ca. 201mm SL), Iraq: Nahr mouth inferior; lips slightly fleshy, especially at the Rawanduz and Serokani stream (Rawanduz), N. mouth corners; lower lip covered with a sharp-edged Sarsam. — BMNH 1974.2.22.1394- 1395. (14) 2, horny sheath, its anterior margin straight in adult 173.15-197.74mm SL; Iraq: Great Zab near Aski specimens and rounded to almost crescent-shaped in Kalak (Eski Kelek) and near Bekhme Dam, N. juveniles, with a considerable degree of individual Sarsam. — BMNH 1974.2.22.1391, (15) 1, bent variation. Dorsal-fin origin inserted anterior to specimen (ca. 197mm SL), Iraq: Siwel River, Little pelvic-fin origin, its outer margin usually straight to Zab, N. Sarsam. — BMNH 1974.2.22.1396, – (16) 1, concave with 4-5 unbranched and 7-8 branched rays, bent specimen, Iraq: Nahr Qiziljah, Little Zab, last unbranched dorsal-fin ray weakly to moderately N. Sarsam. — ZM-CBSU uncatalogued (# 11) (in ossified, flexible and soft at the tip; pectoral fin not 96% alcohol), (17) Fin clip, Iran: Rud-e Garan extending to pelvic-fin base; its outer margins usually (Garan River), Marivan, Kurdistan, 26.VI.2009, H.R. slightly convex with 18-21 rays in total; pelvic fin Esmaeili. — ZM-CBSU Z290-304, (17) 15, 71.3- short, not extending to anal fin base, its outer margin 148.3mm SL; Iran: Garan River, Tigris River basin, straight or slightly convex and blunt with 8-10 rays, at Marivan, Kurdistan Province. pelvic axillary scale present; anal fin with 3 unbranched and 5 branched rays, its outer margin Capoeta birunii sp. n. straight or slightly convex; caudal fin forked with 17- (Figs. 27-28) 18 branched rays. Scales small, total lateral-line http://zoobank.org:LSIDurn:lsid:zoobank.org:act:3E scales 68-82, 13-15 scales between dorsal- fin origin FC9C1A-10F4-4D0D-8CCA-7B233B4FBE27 and lateral line, 7-11 between anal-fin origin and Holotype: ZM-CBSU Z650, 141mm SL; Iran: lateral line; 20-26 circum-peduncle scales; total gill 261 Iran. J. Ichthyol. (September 2017), 4(3): 231-269

Fig.27. Capoeta birunii, a) holotype: ZM-CBSU Z650; 141mm SL; b) ZM-CBSU Z651; 105mm SL; c) ZM-CBSU Z652; 104mm SL.

Fig.28. Capoeta birunii from Zayandehrud tributary, Daran, Zayandehrud basin. rakers on first gill arch 13-17 (10-13 on lower limb). shallow grove in the C. saadii). Capoeta birunii is Capoeta birunii is distinguished from C. buhsei in distinguished from C. umbla by number of scales having more gill rakers on lower limb of first gill arch (LL: 68-82 vs. 82-95, ALL: 13-15 vs. 18-25) and (10-13 vs. 7-10) and number of lateral line scales (68- number of gill rakers on first gill arch (10-13 vs. 17- 82 vs. 82-95). Capoeta birunii is distinguished from 20). Capoeta birunii is distinguished from C. saadii by shape of snout (presence of a depression C. shajariani by barbel length in percent of mouth anterior to nasal pore on snout (vs. absence or width (57-79% vs. 39-55%). Capoeta birunii is

262 Zareian and Esmaeili - Phylogeny and taxonomy of Capoeta damascina species group in Iran

Table 12. Morphometric characters of Capoeta birunii in Zayandehrud basin; Zayandeh River.

Character N Minimum Maximum Mean Std. Deviation In percent of standard length Head length 11 15.14 18.56 16.9155 .85830 Maximum Head depth 11 13.53 15.52 14.2875 .54013 Head width 11 8.40 9.59 8.8696 .38965 Postorbital length 11 11.89 12.89 12.5457 .31520 Preorbital length 11 24.29 25.93 25.3524 .44284 Dorsal-fin base length 11 12.23 14.28 13.1362 .51240 Longest dorsal fin ray length 11 17.90 21.15 19.0871 1.12174 Predorsal length 11 50.76 54.72 52.6474 1.49585 Postdorsal length 11 53.49 58.40 56.7173 1.36313 Pectoral fin length 11 16.69 18.78 17.9346 .60671 Pelvic fin length 11 14.42 16.19 15.1953 .56419 Pelvic fin base length 11 4.00 4.85 4.3979 .30854 Prepelvic length 11 52.47 74.29 56.0030 6.16107 Anal fin base length 11 6.15 7.75 6.9802 .49606 Preanal length 11 72.86 75.93 74.8069 .88483 Anal fin length 11 13.41 20.54 16.8508 1.95401 Length of caudal peduncle 11 17.98 20.40 19.3589 .74230 Depth of caudal peduncle 11 10.00 11.55 10.7729 .47153 Body depth at dorsal-fin origin 11 20.71 24.61 22.9544 1.25759 In percent of head length Maximum Head depth 11 62.32 72.37 66.7089 2.91057 Head width 11 53.89 59.85 56.3549 1.83983 Post orbital length 11 46.48 51.83 49.4966 1.42034 Preorbital length 11 33.35 38.18 34.9941 1.64399 Interorbital distance 11 33.19 38.38 36.1064 1.42872 Eye diameter 11 15.83 20.82 18.7753 1.80347 Intranasal distance 11 19.78 24.90 22.0553 1.45878 Mouth width 11 29.20 37.11 32.7226 2.48099 Barbel length 11 19.34 26.36 22.4304 2.02363 distinguished from C. pyragyi by number of gill Habitat and distribution: Capoeta birunii can be rakers on lower limb of first gill arch (13-15 vs. 10- found in slow to medium flowing rivers with muddy 13). Capoeta birunii is distinguished from bottoms and riparian vegetation. This species C. ferdowsi by horizontal eye diameter in percent of inhabits the Zayandeh River basin in Iran (Figs. 8, preorbital length (43.13-61.04 vs. 70.21-84.91). 29). Coloration: Dorsum, head and sides of C. birunii Etymology: The new species is named after Abū golden brown, darker dorsally and lighter below the Rayḥān Muḥammad ibn Aḥmad Al-Bīrūnī Abū Rayḥān ابوریحان بیرونی :lateral line (Fig. 28); white belly; overall body silvery (Chorasmian/Persian or golden in different specimens; pectoral and pelvic Bērōnī; New Persian: Abū Rayḥān Bīrūnī (4 fins golden-yellow. White operculum; outer margin September 973 – 9 December 1048), known as Al- of eyes golden; base of caudal fin golden. in English. He was an Iranian (البیرونی :Biruni (Arabic

263 Iran. J. Ichthyol. (September 2017), 4(3): 231-269

Fig.29. Zayandeh River, Daran, habitat of Capoeta birunii, Zayandehrud basin.

Table 13. Frequency distribution of the number of scales between lateral line and origin of anal fin.

Species 6 7 8 9 10 11 12 13 14 C. buhsei 3 13 7 3 1 C. coadi 1 11 12 8 1 C. saadii 2 4 3 9 2 C. umbla 2 2 4 1 C. shajariani 5 4 C. pyragyi 1 9 4 2 C. ferdowsii 2 5 1 C. birunii 1 2 5 2

Table 14. Frequency distribution of the number of scales between lateral line and origin of dorsal fin.

Species 11 12 13 14 15 16 17 18 20 21 22 25 C. buhsei 3 6 4 12 2 C. coadi 1 8 12 11 1 C. saadii 2 5 7 3 3 C. umbla 1 3 3 1 1 C. shajariani 3 6 C. pyragyi 1 8 4 3 C. ferdowsii 1 4 C. birunii 3 4 4

264 Zareian and Esmaeili - Phylogeny and taxonomy of Capoeta damascina species group in Iran

Table 15. Frequency distribution of the number of gill rakers on the first arch.

Species 12 13 14 15 16 17 18 19 20 C. buhsei 10 13 4 C. coadi 1 1 7 13 6 5 C. saadii 1 9 6 1 2 1 C. umbla 1 4 3 1 C. shajariani 2 3 2 2 C. pyragyi 3 4 4 4 C. ferdowsii 1 3 4 1 C. birunii 1 3 2 2 3

Table 16. Frequency distribution of the number of scales of the lateral line.

scholar and polymath. and also the existence of a new species of Capoeta in Some meristic characters of Iranian small scaled the Zayandeh River basin. The comparative Capoeta are present in Tables 13-16. morphological analyses clearly support the relationships between the studied species as Discussion and conclusions suggested by the molecular analysis. So now small For a long time, all the small scaled Capoeta which scaled Capoeta populations belong to six species, were distributed in the Iranian basins (e.g., Tigris, which had been previously designated as Esfahan, Kor, Maharlu Lake) were considered as C. damascina or C. saadii, of which, four species C. damascina or recently as C. saadii and C. buhsei including C. coadi, C. saadii, C. shajariani and (see Coad 1995; Esmaeili et al. 2010; 2014; 2017; C. pyragyi occur in the Iranian Tigris River basin. Jouladeh-Roudbal et al. 2015). However, recent This high diversity might be due to habitat studies revealed the presence of a complex group of heterogeneity, discontinuity of the river drainages, species (Alwan 2016b). Alwan et al. (2016a) and reducing gene flow in this large river basin. considered populations from the Karun River system as new species, Capoeta coadi. Ghanavi et al. (2016) Acknowledgments also considered three additional species of small This study was carried out in strict accordance with scaled Capoeta are present in the Iranian Tigris basin applicable national and international guidelines. We based on only the cytb gene without describing these thank E. Mikschi, and A. Palandacic (all NMW) for species. providing Capoeta umbla syntype photos. We also Our molecular (based on the COI and cytb genes) would like to thank A. Gholamhosseini, R. Sadeghi, and morphological results confirm differences R. Khaefi, M. Masoudi, H. Mehraban, G. Sayyad- between populations of this genus, presence of zadeh, H. Darvishneia, M. Ghasemian and undescribed species in the Iranian drainage basins M. Razbani for their help during field work. We 265 Iran. J. Ichthyol. (September 2017), 4(3): 231-269

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268 Iran. J. Ichthyol. (September 2017), 4(3): 231–269 Received: July 27, 2017 © 2016 Iranian Society of Ichthyology Accepted: September 10, 2017 P-ISSN: 2383-1561; E-ISSN: 2383-0964 doi: 10.22034/iji.v4i3.23902.02 http://www.ijichthyol.org

مقاله پژوهشی تبارشناسی ميتوکندريايی و وضعيت آرايهشناختی گروه گونه Capoeta damascina )پرتوبالگان: کپورماهيان( و توصيف يک گونه جديد از ايران

حليمه زارعيان، حميد رضا اسماعيلی*

آزمايشگاه تحقيقاتي ماهيشناسي و سيستماتيک مولکولي، بخش جانورشناسي، گروه زيستشناسي، دانشکده علوم، دانشگاه شيراز، شيراز، ايران.

چکيده: گونههاي جنس سياه ماهي )Capoeta( که از تنوع بااليي در ناحيه خاورميانه برخوردارند و داراي تاريخچه تکاملي و وضعيت آرايهشناختي پيچيدهاي باشند از وابستگان نزديک جنس Luciobarbus هستند. نتايج مولکولي و ريختشناسي حاضر نشان دهنده وجود هشت گونه از گروه گونهاي Capoeta damascina در ايران ميباشد. در اين مطالعه همه هشت گونه بررسي شده و صفات تشخيصي براي گونه Capoeta birunii بهعنوان گونه جديداً توصيف شده از حوضههاي آبريز زايندهرود آورده شده است. اين گونه از ديگر گونههاي گروه گونهاي C. damascina بهوسيله مجموعهاي از صفات ريختي و مولکولي قابل تشخيص هستند. کلماتکليدی: سياهماهي، آناليز مولکولي، سيستماتيک، پراکنش.

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