Main Article Content
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.
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How to Cite
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. https://doi.org/10.22034/iji.v4i3.239
References
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Alwan, N. 2011. Systematics, taxonomy, phylogeny and zoogeography of the Capoeta damascina species complex (Pisces: Teleostei: Cyprinidae) inferred from comparative morphology and molecular markers. PhD thesis, Frankfurt a.M., Germany: Goethe Universität.
Alwan, N.; Zareian, H. & Esmaeili H.R. 2016a. Capoeta coadi, a new species of cyprinid fish from the Karun River drainage, Iran based on morphological and molecular evidences (Teleostei, Cyprinidae). Zookeys 573: 155-180.
Alwan, N.; Esmaeili, H.R. & Krupp, F. 2016b. Molecular Phylogeny and Zoogeography of the Capoeta damascina Species Complex (Pisces: Teleostei: Cyprinidae). PLOS ONE 11(6): e0156434.
Beckman, W.C. 1962. The freshwater fishes of Syria. FAO Fisheries Biology Technical Paper 8: 1-297.
Berg, L.S. 1949. Freshwater Fishes of the USSR and Adjacent Countries. Academy of Sciences of the USSR.
Bianco, P.G. & Bănărescu, P. 1982. A contribution to the knowledge of the Cyprinidae of Iran (Pisces, Cypriniformes). Cybium 6: 75-96.
Bruford, M.W.; Hanotte, O.; Brokfield, J.F.Y. & Burke, T. 1992. Single-locus and multilocus DNA fingerprinting. In: (A.R. Hoelzel, ed.) Molecular genetic analysis of populations: a practical approach. Oxford University Press, New York. pp. 225-269.
Coad, B.W. 1995. Freshwater Fishes of Iran. Acta Scientiarum Naturalium Academiae Scientiarum Bohemicae 29(1):1-64.
Coad, B.W. 2008. Fishes of Tehran Province and adjacent areas. Shabpareh Publications, Tehran.
Coad, B.W. 2017. Freshwater fishes of Iran. Retrieved from http://www.briancoad.com. Accessed: February 2017.
De Filippi, F. 1865. Note de un viaggio in Persia nei 1862. Pesci, pp. 3, 63, 141, 357-360.
Derzhavin, A.N. 1929. Kurinskoe rybnoe khozyastvo i melioratsiya mugani [The fisheries of the Kura and the melioration of the Mugan]. Izvestiya Bakinskoi Ikhtiologicheskoi Laboratorii 2(2): 1-68.
Durand, J.D; Tsigenopoulos, C.S.; Unlü, E. & Berrebi, P. 2002. Phylogeny and biogeography of the family Cyprinidae in the Middle East inferred from cytochrome b DNA- evolutionary significance of this region. Molecular Phylogenetics and Evolution 22(1): 91-100.
Eschmeyer, W.N. & Fricke, R. 2017. Catalog of Fishes: Genera, Species, References.10/ 10/2016.
Esmaeili, H.R.; Coad, B.W.; Gholamifard, A.; Nazari, N. & Teimory, A. 2010. Annotated checklist of the freshwater fishes of Iran. Zoosystematica Rossica 19: 361-386.
Esmaeili H.R.; Coad, B.W.; Mehraban, H.R.; Masoudi, M.; Khaefi, R.; Abbasi, K.; Mostavavi, H. & Vatandoust, S. 2014. An updated checklist of fishes of the Caspian Sea basin of Iran with a note on their zoogeography. Iranian Journal of Ichthyology 1(3): 152-184.
Esmaeili, H.R.; Babai, S.; Gholamifard, A.; Pazira, A.; Gholamhosseini, A. & Coad, B.W. 2015. Fishes of the Persis region of Iran: an updated checklist and ichthyogeography. Iranian Journal of Ichthyology 2: 201-223.
Esmaeili, H.R.; Zareian, H.; Eagderi, S. & Alwan, N. 2016. Review on the taxonomy of Tigris scraper, Capoeta umbla (Heckel, 1843) and its confirmation record from the Iranian part of Tigris River, Persian Gulf basin (Teleostei: Cyprinidae). FishTaxa 1: 35-44.
Esmaeili, H.R.; Mehraban, H.; Abbasi, K.; Keivany, Y. & Coad, B.W. 2017. Review and updated checklist of freshwater fishes of Iran: Taxonomy, distribution and conservation status. Iranian Journal of Ichthyology 4(Suppl. 1): 1-114.
Freyhof, J. 2014. Capoeta umbla. The IUCN Red List of Threatened Species.
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.
Günther, A. 1868. Catalogue of the fishes in the British Museum. Vol. 7. British Museum, London. 512 p.
Hall, T.A. 1999. BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symposium Series 41: 95–98. [London] Heckel, J.J. 1849. Die Fische Persiens gesammelt von Theodor Kotschy. In J. Russegger (Ed.), Reisen in Europa, Asien und Afrika (pp. 255-335). Volume 2. Stuttgart: Schweitzerbart’sche Verlagsbuchhandlung.
Heckel, J.J. 1843. Ichthyologie [von Syrien]. In: J. von Russegger. Reisen in Europa, Asien und Africa, mit besonderer Rücksicht auf die naturwissenschaftlichen Verhältnisse der betreffenden Länder unternommen in den Jahren 1835 bis 1841, etC. E. Schweizerbart'sche Verlagshandlung 1(2): 990-1099.
Hubbs, C.L. & Lagler, K.F. 1958. Fishes of the Great Lakes Region. Ann Arbor, University of Michigan Press.
Huelsenbeck, J.P. & Ronquist, F. 2001. MrBayes: Bayesian inference of phylogeny. Bioinformatics 17: 754-755.
Jouladeh-Roudbar, A.; Vatandoust, S.; Eagderi, S.; Jafari-Kenari, S. & Mousavi-Sabet, H. 2015. Freshwater fishes of Iran; an updated checklist. AACL Bioflux 8(6): 855-909.
Jouladeh-Roudbar, A.; Eagderi, S.; Murillo-Ramos, L.; Ghanavi, H. R. & Doadrio I. 2017. Three new species of algae-scraping cyprinid from Tigris River drainage in Iran (Teleostei: Cyprinidae). FishTaxa 2(3): 134-155.
Kähsbauer, P. 1963. Zur Kenntnis der Ichthyofauna von Iran (II. Teil). Annalen des naturhistorischen Museums in Wien 67: 453-475.
Karaman, M.S. 1969. Süßwasserfische der Türkei. 7. Teil. Revision der kleinasiatischen und vorderasiatischen Arten des Genus Capoeta (Varicorhinus, partim). Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut 66: 17-54.
Keivany, Y.; Nasri, M.; Abbasi, K. & Abdoli, A. 2016. Atlas of inland water fishes of Iran. Iran Department of Environment Press. 218 p.
Keyserling, E. 1861. Neue Cypriniden aus Persien. Zeitschrift für die Gesammten Naturwissenschaften, Halle 17(1):1-24, Taf. I-IX.
Kottelat, M. & Freyhof, J. 2007. Handbook of European Freshwater Fishes. Published by the authors, Cornol.
Kosswig, C. & Battalgil, F. 1943. Türkiye tatlı su balıklarının zoogeografik ehemmiyeti. Türk Fiziki Tabii İlimler Sosyetesi Yıllık Bildiriler Arşivi II, Sayı 8: 18-31.
Krupp, F. & Schneider, W. 1989. The fishes of the Jordan River drainage basin and Azraq Oasis. Fauna of Saudi Arabia 10: 347-416.
Krupp, F. 1983. Freshwater fishes of Saudi Arabia and adjacent regions of the Arabian Peninsula. Fauna of Saudi Arabia 5: 568-636.
Krupp, F.; Al-Jumaily, M.; Bariche, M.; Khalaf, M.; Malek, M. & Streit, B. 2009. The Middle Eastern Biodiversity Network: Generating and sharing knowledge for ecosystem management and conservation. ZooKeys 31: 3-15.
Kuru, M. 1971. The freshwater fish fauna of eastern Anatolia. Istanbul Üniversitesi Fen Fakültesi Mecmuasi, Serisi B 36: 137-147.
Kuru, M. 1980. Van gölünde saptanan yeni bir balık türü (Cyprinidae: Alburnus timarensis sp. nov.). Tubitak VII. Bilim Kongresi-Aydın, Biyoloji Tebliğleri 7: 493-504.
Küçük, F.; Turan, D.; Şahin, C. & Gülle, İ. 2009. Capoeta mauricii n. sp., a new species of cyprinid fish from Lake Beyşehir, Turkey. Zoology in the Middle East 47(1): 71-82.
Ladiges, W. 1960. Süßwasserfische der Türkei, 1. Teil Cyprinidae. Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut 58: 105-150.
Machordom, A. & Doadrio, I. 2001. Evidence of a Cenozoic Betic–Kabilian Connection Based on Freshwater Fish Phylogeography (Luciobarbus, Cyprinidae). Molecular Phylogenetics and Evolution 18: 252-263.
Posada, D. & Crandall, K.A. 1998. MODELTEST: testing the model of DNA substitution. Bioinformatics, 14: 817-818.
Saadati, M. 1977. Taxonomy and distribution of the freshwater fishes of Iran. M. SC. Thesis. Colorado State University, Fort Collins, Colorado.
Schöter, C.; Özuluğ, M. & Freyhof, J. 2009. Capoeta caelestis, a new species from Göksu River, Turkey (Teleostei: Cyprinidae). Ichthyological Exploration of Freshwaters 20(3): 229-236.
Stamatakis, A. 2006. RAxML-VI-HPC: maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics 22: 2688-2690.
Tamura, K.; Stecher, G.; Peterson, D.; Filipski, A. & Kumar, S. 2013. MEGA6: Molecular Evolutionary Genetics Analysis Version 6.0. Molecular Biology and Evolution 30: 2725-2729.
Tortonese, E. 1934. Pesci della Persia raccolti dal Marchese Giacomo Doria (1862). Bollettino dei Museo di Zoologia e di Anatomia Comparata della R. Università di Torino 44, III(49): 153-171.
Ward, R.D.; Zemlak T.S.; Innes, H.B.; Last, R.P. & Hebert, P.D.N. 2005. DNA barcoding Australia’s fish species. Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences 360 (1462): 1847-1857.
Xiao, W.; Zhang, Y. & Liu, H. 2001. Molecular systematics of Xenocyprinae (Teleostei: Cyprinidae): taxonomy, biogeography, and coevolution of a special group restricted in East Asia. Molecular Phylogenetics and Evolution 18: 163-173.
Zareian, H.; Esmaeili, H.R.; Heidari, A.; Khoshkholgh, M.R. & Mousavi-Sabet, H. 2016a. Contribution to the molecular systematics of the genus Capoeta from the south Caspian Sea basin using mitochondrial cytochrome b sequences (Teleostei: Cyprinidae). Molecular Biology Research Communications 5: 65-75.
Zareian, H.; Esmaeili, H.R. & Freyhof, J. 2016b. Capoeta anamisensis, a new species from the Minab and Hasan Langhi River drainages in Iran (Teleostei: Cyprinidae). Zootaxa 4083(1): 126-142.
Zhao, L.F. & Xie, X.B. 2016. Strong Lg-wave attenuation in the Middle East continental collision orogenic belt. Tectonophysics 674: 135-146.
References
Alwan, N. 2011. Systematics, taxonomy, phylogeny and zoogeography of the Capoeta damascina species complex (Pisces: Teleostei: Cyprinidae) inferred from comparative morphology and molecular markers. PhD thesis, Frankfurt a.M., Germany: Goethe Universität.
Alwan, N.; Zareian, H. & Esmaeili H.R. 2016a. Capoeta coadi, a new species of cyprinid fish from the Karun River drainage, Iran based on morphological and molecular evidences (Teleostei, Cyprinidae). Zookeys 573: 155-180.
Alwan, N.; Esmaeili, H.R. & Krupp, F. 2016b. Molecular Phylogeny and Zoogeography of the Capoeta damascina Species Complex (Pisces: Teleostei: Cyprinidae). PLOS ONE 11(6): e0156434.
Beckman, W.C. 1962. The freshwater fishes of Syria. FAO Fisheries Biology Technical Paper 8: 1-297.
Berg, L.S. 1949. Freshwater Fishes of the USSR and Adjacent Countries. Academy of Sciences of the USSR.
Bianco, P.G. & Bănărescu, P. 1982. A contribution to the knowledge of the Cyprinidae of Iran (Pisces, Cypriniformes). Cybium 6: 75-96.
Bruford, M.W.; Hanotte, O.; Brokfield, J.F.Y. & Burke, T. 1992. Single-locus and multilocus DNA fingerprinting. In: (A.R. Hoelzel, ed.) Molecular genetic analysis of populations: a practical approach. Oxford University Press, New York. pp. 225-269.
Coad, B.W. 1995. Freshwater Fishes of Iran. Acta Scientiarum Naturalium Academiae Scientiarum Bohemicae 29(1):1-64.
Coad, B.W. 2008. Fishes of Tehran Province and adjacent areas. Shabpareh Publications, Tehran.
Coad, B.W. 2017. Freshwater fishes of Iran. Retrieved from http://www.briancoad.com. Accessed: February 2017.
De Filippi, F. 1865. Note de un viaggio in Persia nei 1862. Pesci, pp. 3, 63, 141, 357-360.
Derzhavin, A.N. 1929. Kurinskoe rybnoe khozyastvo i melioratsiya mugani [The fisheries of the Kura and the melioration of the Mugan]. Izvestiya Bakinskoi Ikhtiologicheskoi Laboratorii 2(2): 1-68.
Durand, J.D; Tsigenopoulos, C.S.; Unlü, E. & Berrebi, P. 2002. Phylogeny and biogeography of the family Cyprinidae in the Middle East inferred from cytochrome b DNA- evolutionary significance of this region. Molecular Phylogenetics and Evolution 22(1): 91-100.
Eschmeyer, W.N. & Fricke, R. 2017. Catalog of Fishes: Genera, Species, References.10/ 10/2016.
Esmaeili, H.R.; Coad, B.W.; Gholamifard, A.; Nazari, N. & Teimory, A. 2010. Annotated checklist of the freshwater fishes of Iran. Zoosystematica Rossica 19: 361-386.
Esmaeili H.R.; Coad, B.W.; Mehraban, H.R.; Masoudi, M.; Khaefi, R.; Abbasi, K.; Mostavavi, H. & Vatandoust, S. 2014. An updated checklist of fishes of the Caspian Sea basin of Iran with a note on their zoogeography. Iranian Journal of Ichthyology 1(3): 152-184.
Esmaeili, H.R.; Babai, S.; Gholamifard, A.; Pazira, A.; Gholamhosseini, A. & Coad, B.W. 2015. Fishes of the Persis region of Iran: an updated checklist and ichthyogeography. Iranian Journal of Ichthyology 2: 201-223.
Esmaeili, H.R.; Zareian, H.; Eagderi, S. & Alwan, N. 2016. Review on the taxonomy of Tigris scraper, Capoeta umbla (Heckel, 1843) and its confirmation record from the Iranian part of Tigris River, Persian Gulf basin (Teleostei: Cyprinidae). FishTaxa 1: 35-44.
Esmaeili, H.R.; Mehraban, H.; Abbasi, K.; Keivany, Y. & Coad, B.W. 2017. Review and updated checklist of freshwater fishes of Iran: Taxonomy, distribution and conservation status. Iranian Journal of Ichthyology 4(Suppl. 1): 1-114.
Freyhof, J. 2014. Capoeta umbla. The IUCN Red List of Threatened Species.
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.
Günther, A. 1868. Catalogue of the fishes in the British Museum. Vol. 7. British Museum, London. 512 p.
Hall, T.A. 1999. BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symposium Series 41: 95–98. [London] Heckel, J.J. 1849. Die Fische Persiens gesammelt von Theodor Kotschy. In J. Russegger (Ed.), Reisen in Europa, Asien und Afrika (pp. 255-335). Volume 2. Stuttgart: Schweitzerbart’sche Verlagsbuchhandlung.
Heckel, J.J. 1843. Ichthyologie [von Syrien]. In: J. von Russegger. Reisen in Europa, Asien und Africa, mit besonderer Rücksicht auf die naturwissenschaftlichen Verhältnisse der betreffenden Länder unternommen in den Jahren 1835 bis 1841, etC. E. Schweizerbart'sche Verlagshandlung 1(2): 990-1099.
Hubbs, C.L. & Lagler, K.F. 1958. Fishes of the Great Lakes Region. Ann Arbor, University of Michigan Press.
Huelsenbeck, J.P. & Ronquist, F. 2001. MrBayes: Bayesian inference of phylogeny. Bioinformatics 17: 754-755.
Jouladeh-Roudbar, A.; Vatandoust, S.; Eagderi, S.; Jafari-Kenari, S. & Mousavi-Sabet, H. 2015. Freshwater fishes of Iran; an updated checklist. AACL Bioflux 8(6): 855-909.
Jouladeh-Roudbar, A.; Eagderi, S.; Murillo-Ramos, L.; Ghanavi, H. R. & Doadrio I. 2017. Three new species of algae-scraping cyprinid from Tigris River drainage in Iran (Teleostei: Cyprinidae). FishTaxa 2(3): 134-155.
Kähsbauer, P. 1963. Zur Kenntnis der Ichthyofauna von Iran (II. Teil). Annalen des naturhistorischen Museums in Wien 67: 453-475.
Karaman, M.S. 1969. Süßwasserfische der Türkei. 7. Teil. Revision der kleinasiatischen und vorderasiatischen Arten des Genus Capoeta (Varicorhinus, partim). Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut 66: 17-54.
Keivany, Y.; Nasri, M.; Abbasi, K. & Abdoli, A. 2016. Atlas of inland water fishes of Iran. Iran Department of Environment Press. 218 p.
Keyserling, E. 1861. Neue Cypriniden aus Persien. Zeitschrift für die Gesammten Naturwissenschaften, Halle 17(1):1-24, Taf. I-IX.
Kottelat, M. & Freyhof, J. 2007. Handbook of European Freshwater Fishes. Published by the authors, Cornol.
Kosswig, C. & Battalgil, F. 1943. Türkiye tatlı su balıklarının zoogeografik ehemmiyeti. Türk Fiziki Tabii İlimler Sosyetesi Yıllık Bildiriler Arşivi II, Sayı 8: 18-31.
Krupp, F. & Schneider, W. 1989. The fishes of the Jordan River drainage basin and Azraq Oasis. Fauna of Saudi Arabia 10: 347-416.
Krupp, F. 1983. Freshwater fishes of Saudi Arabia and adjacent regions of the Arabian Peninsula. Fauna of Saudi Arabia 5: 568-636.
Krupp, F.; Al-Jumaily, M.; Bariche, M.; Khalaf, M.; Malek, M. & Streit, B. 2009. The Middle Eastern Biodiversity Network: Generating and sharing knowledge for ecosystem management and conservation. ZooKeys 31: 3-15.
Kuru, M. 1971. The freshwater fish fauna of eastern Anatolia. Istanbul Üniversitesi Fen Fakültesi Mecmuasi, Serisi B 36: 137-147.
Kuru, M. 1980. Van gölünde saptanan yeni bir balık türü (Cyprinidae: Alburnus timarensis sp. nov.). Tubitak VII. Bilim Kongresi-Aydın, Biyoloji Tebliğleri 7: 493-504.
Küçük, F.; Turan, D.; Şahin, C. & Gülle, İ. 2009. Capoeta mauricii n. sp., a new species of cyprinid fish from Lake Beyşehir, Turkey. Zoology in the Middle East 47(1): 71-82.
Ladiges, W. 1960. Süßwasserfische der Türkei, 1. Teil Cyprinidae. Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut 58: 105-150.
Machordom, A. & Doadrio, I. 2001. Evidence of a Cenozoic Betic–Kabilian Connection Based on Freshwater Fish Phylogeography (Luciobarbus, Cyprinidae). Molecular Phylogenetics and Evolution 18: 252-263.
Posada, D. & Crandall, K.A. 1998. MODELTEST: testing the model of DNA substitution. Bioinformatics, 14: 817-818.
Saadati, M. 1977. Taxonomy and distribution of the freshwater fishes of Iran. M. SC. Thesis. Colorado State University, Fort Collins, Colorado.
Schöter, C.; Özuluğ, M. & Freyhof, J. 2009. Capoeta caelestis, a new species from Göksu River, Turkey (Teleostei: Cyprinidae). Ichthyological Exploration of Freshwaters 20(3): 229-236.
Stamatakis, A. 2006. RAxML-VI-HPC: maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics 22: 2688-2690.
Tamura, K.; Stecher, G.; Peterson, D.; Filipski, A. & Kumar, S. 2013. MEGA6: Molecular Evolutionary Genetics Analysis Version 6.0. Molecular Biology and Evolution 30: 2725-2729.
Tortonese, E. 1934. Pesci della Persia raccolti dal Marchese Giacomo Doria (1862). Bollettino dei Museo di Zoologia e di Anatomia Comparata della R. Università di Torino 44, III(49): 153-171.
Ward, R.D.; Zemlak T.S.; Innes, H.B.; Last, R.P. & Hebert, P.D.N. 2005. DNA barcoding Australia’s fish species. Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences 360 (1462): 1847-1857.
Xiao, W.; Zhang, Y. & Liu, H. 2001. Molecular systematics of Xenocyprinae (Teleostei: Cyprinidae): taxonomy, biogeography, and coevolution of a special group restricted in East Asia. Molecular Phylogenetics and Evolution 18: 163-173.
Zareian, H.; Esmaeili, H.R.; Heidari, A.; Khoshkholgh, M.R. & Mousavi-Sabet, H. 2016a. Contribution to the molecular systematics of the genus Capoeta from the south Caspian Sea basin using mitochondrial cytochrome b sequences (Teleostei: Cyprinidae). Molecular Biology Research Communications 5: 65-75.
Zareian, H.; Esmaeili, H.R. & Freyhof, J. 2016b. Capoeta anamisensis, a new species from the Minab and Hasan Langhi River drainages in Iran (Teleostei: Cyprinidae). Zootaxa 4083(1): 126-142.
Zhao, L.F. & Xie, X.B. 2016. Strong Lg-wave attenuation in the Middle East continental collision orogenic belt. Tectonophysics 674: 135-146.
Alwan, N.; Zareian, H. & Esmaeili H.R. 2016a. Capoeta coadi, a new species of cyprinid fish from the Karun River drainage, Iran based on morphological and molecular evidences (Teleostei, Cyprinidae). Zookeys 573: 155-180.
Alwan, N.; Esmaeili, H.R. & Krupp, F. 2016b. Molecular Phylogeny and Zoogeography of the Capoeta damascina Species Complex (Pisces: Teleostei: Cyprinidae). PLOS ONE 11(6): e0156434.
Beckman, W.C. 1962. The freshwater fishes of Syria. FAO Fisheries Biology Technical Paper 8: 1-297.
Berg, L.S. 1949. Freshwater Fishes of the USSR and Adjacent Countries. Academy of Sciences of the USSR.
Bianco, P.G. & Bănărescu, P. 1982. A contribution to the knowledge of the Cyprinidae of Iran (Pisces, Cypriniformes). Cybium 6: 75-96.
Bruford, M.W.; Hanotte, O.; Brokfield, J.F.Y. & Burke, T. 1992. Single-locus and multilocus DNA fingerprinting. In: (A.R. Hoelzel, ed.) Molecular genetic analysis of populations: a practical approach. Oxford University Press, New York. pp. 225-269.
Coad, B.W. 1995. Freshwater Fishes of Iran. Acta Scientiarum Naturalium Academiae Scientiarum Bohemicae 29(1):1-64.
Coad, B.W. 2008. Fishes of Tehran Province and adjacent areas. Shabpareh Publications, Tehran.
Coad, B.W. 2017. Freshwater fishes of Iran. Retrieved from http://www.briancoad.com. Accessed: February 2017.
De Filippi, F. 1865. Note de un viaggio in Persia nei 1862. Pesci, pp. 3, 63, 141, 357-360.
Derzhavin, A.N. 1929. Kurinskoe rybnoe khozyastvo i melioratsiya mugani [The fisheries of the Kura and the melioration of the Mugan]. Izvestiya Bakinskoi Ikhtiologicheskoi Laboratorii 2(2): 1-68.
Durand, J.D; Tsigenopoulos, C.S.; Unlü, E. & Berrebi, P. 2002. Phylogeny and biogeography of the family Cyprinidae in the Middle East inferred from cytochrome b DNA- evolutionary significance of this region. Molecular Phylogenetics and Evolution 22(1): 91-100.
Eschmeyer, W.N. & Fricke, R. 2017. Catalog of Fishes: Genera, Species, References.10/ 10/2016.
Esmaeili, H.R.; Coad, B.W.; Gholamifard, A.; Nazari, N. & Teimory, A. 2010. Annotated checklist of the freshwater fishes of Iran. Zoosystematica Rossica 19: 361-386.
Esmaeili H.R.; Coad, B.W.; Mehraban, H.R.; Masoudi, M.; Khaefi, R.; Abbasi, K.; Mostavavi, H. & Vatandoust, S. 2014. An updated checklist of fishes of the Caspian Sea basin of Iran with a note on their zoogeography. Iranian Journal of Ichthyology 1(3): 152-184.
Esmaeili, H.R.; Babai, S.; Gholamifard, A.; Pazira, A.; Gholamhosseini, A. & Coad, B.W. 2015. Fishes of the Persis region of Iran: an updated checklist and ichthyogeography. Iranian Journal of Ichthyology 2: 201-223.
Esmaeili, H.R.; Zareian, H.; Eagderi, S. & Alwan, N. 2016. Review on the taxonomy of Tigris scraper, Capoeta umbla (Heckel, 1843) and its confirmation record from the Iranian part of Tigris River, Persian Gulf basin (Teleostei: Cyprinidae). FishTaxa 1: 35-44.
Esmaeili, H.R.; Mehraban, H.; Abbasi, K.; Keivany, Y. & Coad, B.W. 2017. Review and updated checklist of freshwater fishes of Iran: Taxonomy, distribution and conservation status. Iranian Journal of Ichthyology 4(Suppl. 1): 1-114.
Freyhof, J. 2014. Capoeta umbla. The IUCN Red List of Threatened Species.
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.
Günther, A. 1868. Catalogue of the fishes in the British Museum. Vol. 7. British Museum, London. 512 p.
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