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Type: Article
Published: 2022-04-20
Page range: 151-194
Abstract views: 471
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A practical approach to revise the Oxynoemacheilus bergianus species group (Teleostei: Nemacheilidae)

Museum für Naturkunde, Leibniz Institute for Evolution and Biodiversity Science, 10115 Berlin, Germany.
Recep Tayyip Erdogan University, Faculty of Fisheries, 53100 Rize, Turkey.
Zoological Research Museum Alexander Koenig, Leibniz Institute for Animal Biodiversity, Adenauerallee 160, 53113 Bonn, Germany.
Pisces Freshwater fish Taxonomy Cytochrome oxidase I Middle East

Abstract

The Oxynoemacheilus bergianus species group is revised based on tree topology (ML, NJ, MP), distance (K2P and ASAP) and Poisson tree process analyses of DNA barcode data tested against morphometric and morphological characters including colour patterns. The O. bergianus species group is distinguished from other Oxynoemacheilus groups based on morphological characters: its constituent species have a slender caudal peduncle, a suborbital flap in the male, a mottled or blotched colour pattern, and lack bold, black spots on the caudal-fin base. It is also supported as a monophyletic unit in our molecular analysis.

                The O. bergianus group includes 10 molecular clades following congruently well-supported NJ, MP and ML based entities. Species described as O. bergianus, O. banarescui, O. erdali, O. fatsaensis, O. samanticus, and O. simavicus from Turkey, O. lenkoranensis from Azerbaijan, and O. longipinnis and O. parvinae from Iran belong to this species group. The group includes also four unnamed molecular clades. We were unable to detect external differences between any of the molecular clades in colour pattern or any morphometric or morphological characters examined. In the 10 molecular clades in the O. bergianus species group, the intraclade K2P distance ranges from 0.0–1.8% while the distances between molecular clades ranges from 0.6–5.9%. To resolve the species diversity of this group, we also analysed the intraspecific and interspecific variability in the K2P distance of DNA barcode data from 53 other Oxynoemacheilus species. Here, the intraspecific variability ranges from 0.0–2.4% while the interspecific K2P distance ranges from 1.2–20.8%. In the O. bergianus species group, only four groups are detected by the mPTP species delimitation approach distinguished by a K2P distance of 2.9% or more. We treat these four groups as valid species, corresponding to O. banarescui, O. bergianus, O. fatsaensis, and O. simavicus. Oxynoemacheilus samanticus from the Kızılırmak and Seyhan drainages, O. lenkoranensis from the Caspian basin, O. erdali from the Euphrates, and O. longipinnis and O. parvinae from the Tigris drainage are treated as synonyms of O. bergianus. Fishes from an unnamed molecular clade from the upper Tigris, and from a second unnamed clade from the upper Euphrates, are both identified as O. bergianus. Oxynoemacheilus bergianus might be a junior synonym of O. bergi from the Kura. The distribution range of O. simavicus, described from the Simav drainage in the Marmara basin, is expanded to the east and two molecularly differentiated population groups occur in the Sakarya drainage, the Büyük Melen River and potentially in other adjacent coastal streams. Oxynoemacheilus fatsaensis, described from the coastal stream Elekçi in northern Anatolia, is also widespread in the Yeşilırmak drainage. Morphological characters proposed to distinguish O. fatsaensis from the other species of the O. bergianus group could not be confirmed by our data on fishes from the Yeşilırmak. This study also discusses the theoretical background, our reasons for conducting this revision in the way we did, and what the alternatives would be.

 

References

  1. Abdurakhmanov, Y. A. (1962) Freshwater fishes of Azerbaidjan. Akademii nauk Azerbaidzhanskoi SSR, Baku, 407 pp. [in Russian]
    Ahrens, D., Ahyong, S.T., Ballerio, A., Barclay, M.V.L., Eberle, J., Espeland, M., Huber, B.A., Mengual, X., Pacheco, T.L., Peters, R.S., Rulik, B., Vaz-De-Mello, F., Wesener, T. & Krell, F.-T. (2021) Is it time to describe new species without diagnoses?—A comment on Sharkey et al. (2021). Zootaxa, 5027 (2), 151–159. https://doi.org/10.11646/zootaxa.5027.2.1
    Akkuş, M., Sarı, M., Ekmekçi, F.G. & Yoğurtçuoğlu, B. (2021) The discovery of a microbialite-associated freshwater fish in the world’s largest saline soda lake, Lake Van (Turkey). Zoosystematics and Evolution, 97 (1), 181–189. https://doi.org/10.3897/zse.97.62120
    Antal, L., László, B., Kotlík, P., Mozsár, A., Czeglédi, I., Oldal, M., Kemenesi, G., Jakab, F. & Nagy, S.A. (2016) Phylogenetic evidence for a new species of Barbus in the Danube River basin. Molecular Phylogenetics and Evolution, 96, 187–194. https://doi.org/10.1016/j.ympev.2015.11.023
    April, J., Mayden, R.L., Hanner, R.H. & Bernatchez, L. (2011) Genetic calibration of species diversity among North America’s freshwater fishes. Proceedings of the National Academy of Sciences of the United States of America, 108, 10602–10607. https://doi.org/10.1073/pnas.1016437108
    Bănărescu, P.M., Nalbant, T.T. & Balik, S. (1978) Süßwasserfische der Türkei. 11. Teil. Die Gattung Orthrias in der Türkei und in Südbulgarien (Pisces, Cobitidae, Noemacheilinae). Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut, 75, 255–266.
    Bayçelebi, E., Kaya, C., Turan, D., Ergüden, S.A. & Freyhof, J. (2018) Redescription of Garra turcica from southern Anatolia (Teleostei: Cyprinidae). Zootaxa, 4524 (2), 227–236. https://doi.org/10.11646/zootaxa.4524.2.6
    Bayçelebi, E., Kaya, C., Turan, D. & Freyhof, J. (2021) Garra orontesi, a new species from the Orontes River drainage (Teleostei: Cyprinidae). Zootaxa, 4952 (1), 169–180. https://doi.org/10.11646/zootaxa.4952.1.10
    Berg, L.S. (1899) Beiträge zur Ichthyofauna des Kaukasus. Mitteilungen des Kaukasischen Museums [Bulletin du Musée du caucase; Izvestija Kavkazskogo Muzeja], 1 (3), 1–80. [in Russian and German]
    Bianco, P.G. & Delmastro, G.B. (2011) Recenti novità tassonomiche riguardanti i pesci d‘acqua dolce autoctoni in Italiae e descrizione di una nuova specie di luccio. Researches on Wildlife Conservation, IGF Publishing, USA, 2 (Supplement), 1–13
    Biomatters (2013) Geneious Pro. Available from: http://www.geneious.com (accessed 29 March 2022)
    Britz, R., Hundsdörfer, A. & Fritz, U. (2020) Funding, training, permits—the three big challenges of taxonomy. Megataxa, 1 (1), 49–52. https://doi.org/10.11646/megataxa.1.1.10
    Buj, I., Vukić, J., Šanda, R., Perea, S., Ćaleta, M., Marčić, Z., Bogut, I., Povž, M. & Markovčić, M. (2010) Morphological comparison of bleaks (Alburnus, Cyprinidae) from the Adriatic basin with the description of a new species. Folia Zoologica: international journal of vertebrate zoology, 59, 129–141. https://doi.org/10.25225/fozo.v59.i2.a8.2010
    Butcher, B.A., Smith, M.A., Sharkey, M.J. & Quicke, D.L.I. (2012) A turbo-taxonomic study of Thai Aleiodes (Aleiodes) and Aleiodes (Arcaleiodes) (Hymenoptera: Braconidae: Rogadinae) based largely on COI barcoded specimens, with rapid descriptions of 179 new species. Zootaxa, 3457 (1), 1–232. https://doi.org/10.11646/zootaxa.3457.1.1
    Çiçek, E., Eagderi, S. & Sungur, S. (2018) Oxynoemacheilus veyseli, a new nemacheilid species from the upper Aras River drainage of Turkey (Teleostei: Nemacheilidae). Iranian Journal of Ichthyology, 5 (3), 232–242. https://doi.org/10.22034/iji.v5i3.302
    Coad, B.W. & Nalbant, T.T. (2005) A new genus and a new species of a remarkable nemacheilid fish from Iran (Pisces: Ostariophysi: Nemacheilidae). Travaux du Muséum National d’Histoire Naturelle “Grigore Antipa”, Bucharest, 48, 303–308.
    Collins, R.A. & Cruickshank, R.H. (2013) The seven deadly sins of DNA barcoding. Molecular Ecology Resources, 13, 969–975 https://doi.org/10.1111/1755-0998.1204
    Darriba, D., Taboada, G.L., Doallo, R. & Posada, D. (2012) jModelTest 2: more models, new heuristics and parallel computing. Nature Methods, 9, 772. https://doi.org/10.1038/nmeth.2109
    de Queiroz, K. (2007) Species concepts and species delimitation. Systematic Biology, 56, 879–886. https://doi.org/10.1080/10635150701701083
    Delmastro, G.B. (1982) Un nuovo cobite dai tributari del Mar Nero in Asia Minore (Osteichthyes, Cobitidae). Rivista Piemontese di Storia Naturale, 3, 53–59.
    Derjavin, A.N. (1934) Freshwater fish of the south coast of the Caspian Sea. Trudy Azerbaidzhanskogo otdela Zakavkazskogo filiala Akademii Nauk SSSR, Sektor Zoologii, 7, 91–126. [In Russian, English summary.]
    Doadrio, I. & Perdices, A. (1997) Taxonomic study of the Iberian Cobitis (Osteichthyes, Cobitidae), with description of a new species. Zoological Journal of the Linnean Society, 119, 51–67. https://doi.org/10.1111/j.1096-3642.1997.tb00135.x
    Dvŏrák, T., Šlechtová, V. & Bohlen, J. (2022) Using species groups to approach the large and taxonomically unresolved freshwater fish family Nemacheilidae (Teleostei: Cypriniformes). Biology, 11, 175. https://doi.org/10.3390/biology11020175
    Eberle, J., Ahrens, D., Mayer, C., Niehuis, O. & Misof, B. (2020) A plea for standardized nuclear markers in metazoan DNA taxonomy. Trends in Ecology & Evolution, 35 (4), 336–345. https://doi.org/10.1016/j.tree.2019.12.003
    Edgar, R.C. (2004) MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic acids research, 32, 1792–1797. https://doi.org/10.1093/nar/gkh340
    Epitashvili, G., Geiger, M., Astrin, J.J., Herder, F., Japoshvili, B. & Mumladze, L. (2020) Towards retrieving the Promethean treasure: a first molecular assessment of the freshwater fish diversity of Georgia. Biodiversity Data Journal, 8, e57862. https://doi.org/10.3897/BDJ.8.e57862
    Erk’akan, F., Nalbant, T.T. & Özeren, S.C. (2007) Seven new species of Barbatula, three new species of Schistura and a new species of Seminemacheilus (Ostariophysi: Balitoridae: Nemacheilinae) from Turkey. Journal of Fisheries International [electronic journal], 2 (1), 69–85.
    Esmaeili, H.R. & Sayyadzadeh, G. & Zarei, F. & Kafaei, S. & Coad, B.W. (2019) Phylogeographic pattern and population structure of the Persian stone loach, Oxynoemacheilus persa (Heckel 1847) (family: Nemacheilidae) in southern Iran with implications for conservation. Environmental Biology of Fishes, 103, 77–88. https://doi.org/10.1007/s10641-019-00934-y
    Esmaeili, H.R., Sayyadzadeh, G., Coad, B.W. & Eagderi, S. (2016) Review of the genus Garra Hamilton, 1822 in Iran with description of a new species: a morpho-molecular approach (Teleostei: Cyprinidae). Iranian Journal of Ichthyology, 3 (2), 82–121. https://doi.org/10.22034/iji.v3i2.173
    Freyhof, J. (2016) Oxynoemacheilus karunensis, a new species from the Persian Gulf basin Teleostei: Nemacheilidae). Zootaxa, 4175 (1), 94–100. https://doi.org/10.11646/zootaxa.4175.1.9 
    Freyhof, J., Erk’akan, F., Özeren, C. & Perdices, A.J. (2012) An overview of the western Palaearctic loach genus Oxynoemacheilus (Teleostei: Nemacheilidae). Ichthyological Exploration of Freshwaters, 22, 301–312.
    Freyhof, J., Esmaeili, H.R., Sayyadzadeh, G. & Geiger, M. (2014) Review of the crested loaches of the genus Paracobitis from Iran and Iraq with the description of four new species (Teleostei: Nemacheilidae). Ichthyological Exploration of Freshwaters, 25, 11–38.
    Freyhof, J. & Geiger, M.F. (2017) Oxynoemacheilus zarzianus, a new loach from the Lesser Zab River drainage in Iraqi Kurdistan (Teleostei: Nemacheilidae). Zootaxa, 4273 (2), 258–270. https://doi.org/10.11646/zootaxa.4273.2.6
    Freyhof, J. & Geiger, M.F. (2021) Oxynoemacheilus shehabi, a new nemacheilid loach from the upper Orontes in southern Syria (Teleostei: Nemacheilidae). Zootaxa, 4908 (4), 571–583. https://doi.org/10.11646/zootaxa.4908.4.9
    Freyhof, J., Kaya, C., Bayçelebi, E., Geiger, M. & Turan, D. (2018) Generic assignment of Leuciscus kurui Bogutskaya from the upper Tigris drainage, and a replacement name for Alburnus kurui Mangit & Yerli (Teleostei: Leuciscidae). Zootaxa, 4410 (1), 113–135. https://doi.org/10.11646/zootaxa.4410.1.6
    Freyhof, J., Kaya, C., Epitashvili, G. & Geiger, M.F. (2021) Oxynoemacheilus phasicus, a new nemacheilid loach from the eastern Black Sea basin with some remarks on other Caucasian Oxynoemacheilus (Teleostei: Nemacheilidae). Zootaxa, 4952 (1), 135–151. https://doi.org/10.11646/zootaxa.4952.1.8
    Freyhof, J., Kaya, C., Turan, D. & Geiger, M.F. (2019) Review of the Oxynoemacheilus tigris group with the description of two new species from the Euphrates drainage (Teleostei: Nemacheilidae). Zootaxa, 4612 (1), 29–57. https://doi.org/10.11646/zootaxa.4612.1.2
    Geiger, M.F., Herder, F., Monaghan, M.T., Almada, V., Barbieri, R., Bariche, M., Berrebi, P., Bohlen, J., Casal-Lopez, M., Delmastro, G.B., Denys, G.P.J., Dettai, A., Doadrio, I., Kalogianni, E., Kärst, H., Kottelat, M., Kovačić, M., Laporte, M., Lorenzoni, M., Marčić, Z., Özuluğ, M., Perdices, A., Perea, S., Persat, H., Porcelotti, S., Puzzi, C., Robalo, J., Šanda, R., Schneider, M., Šlechtová, V., Stoumboudi, M., Walter, S. & Freyhof, J. (2014) Spatial heterogeneity in the Mediterranean Biodiversity Hotspot affects barcoding accuracy of its freshwater fishes. Molecular Ecology Resources, 14, 1210–1221. https://doi.org/10.1111/1755-0998.12257
    Giangrande, A. (2003) Biodiversity, conservation, and the ‘Taxonomic impediment’. Aquatic Conservation: Marine and Freshwater Ecosystems, 13, 451–459. https://doi.org/10.1002/aqc.584
    Golzarianpour, K., Abdoli, A. & Freyhof, J. (2011) Oxynoemacheilus kiabii, a new loach from Karkheh River drainage, Iran (Teleostei: Nemacheilidae). Ichthyological Exploration of Freshwaters, 22, 201–208.
    Gratzianov, V.I. (1907) Opyt’ obzora ryby Rossiskoi Imperii v’ sistematicheskom’ i geograficescom’ otnoshenii. Trudy Otdela Ichtiologii Imperatorskago Russkago Obshchestva Akklimatizacii Zivotnych‘ i Rastenii, 4, i–xxx + 1–567.
    Hashemzadeh Segherloo, I., Freyhof, J., Berrebi, P. Ferchaud, A.-L., Geiger, M. Laroche, J., Levin, B.A., Normandeau, E. & Bernatchez, L. (2021) A genomic perspective on an old question: Salmo trouts or Salmo trutta (Teleostei: Salmonidae)? Molecular Phylogenetics and Evolution, 162, 107204. https://doi.org/10.1016/j.ympev.2021.107204
    Hebert, P.D., Cywinska, A., Ball, S.L. & DeWaard, J.R. (2003) Biological identifications through DNA barcodes. Proceedings of the Royal Society of London. Series B: Biological Sciences, 270 (1512), 313–321. https://doi.org/10.1098/rspb.2002.2218
    Hill, G.E. (2016) Mitonuclear coevolution as the genesis of speciation and the mitochondrial DNA barcode gap. Ecology and Evolution, 6 (16), 5831–5842. https://doi.org/10.1002/ece3.2338
    Hubert, N., Hanner, R., Holm, E., Mandrak, N.E., Taylor, E., Burridge, M., Watkinson, D., Dumont, P., Curry, A., Bentzen, P., Zhang, J., April, J. & Bernatchez, L. (2008) Identifying Canadian freshwater fishes through DNA barcodes. PLoS ONE, 3 (6), e2490. https://doi.org/10.1371/journal.pone.0002490
    Ivanova, N.V., Zemlak, T.S., Hanner, R.H. & Hebert, P.D.N. (2007) Universal primer cocktails for fish DNA barcoding. Molecular Ecology Notes, 7, 544–548. https://doi.org/10.1111/j.1471-8286.2007.01748.x
    Jouladeh-Roudbar, A., Eagderi, S. & Hosseinpour, T. (2016) Oxynoemacheilus freyhofi, a new nemacheilid species (Teleostei, Nemacheilidae) from the Tigris basin, Iran. FishTaxa, 1 (2), 94–107. https://doi.org/10.7508/fishtaxa.2016.02.005
    Kapli, P., Lutteropp, S., Zhang, J., Kobert, K., Pavlidis, P., Stamatakis A. & Flouri, T. (2017) Multi-rate Poisson Tree Processes for single-locus species delimitation under Maximum Likelihood and Markov Chain Monte Carlo. Bioinformatics, 33 (11) 1630–1638. https://doi.org/10.1093/bioinformatics/btx025
    Kaya, C., Turan, D., Bayçelebi, E., Kalayci, G. & Freyhof, J. (2020) Oxynoemacheilus cilicicus, a new nemacheilid loach from the Göksu River in southern Anatolia (Teleostei: Nemacheilidae). Zootaxa, 4808 (2), 284–300. https://doi.org/10.11646/zootaxa.4808.2.3
    Khaefi, R., Esmaeili, H.R., Geiger, M.F. & Eagderi, S. (2017) Taxonomic review of the cryptic Barbus lacerta species group with description of a new species (Teleostei: Cyprinidae). FishTaxa, 2 (2), 90–115.
    Korshunova, T., Martynov, A., Bakken, T. & Picton, B. (2017) External diversity is restrained by internal conservatism: new nudibranch mollusc contributes to the cryptic species problem. Zoologica Scripta, 46, 692–683. https://doi.org/10.1111/zsc.12253
    Kotlík, P., Tsigenopoulos, C.S., Ráb, P. & Berrebi, P. (2002) Two new Barbus species from the Danube River basin, with redescription of B. petenyi (Teleostei: Cyprinidae). Folia Zoologica: international journal of vertebrate zoology, 51 (3), 227–240.
    Kottelat, M. (1990) Indochinese nemacheilines. A revision of nemacheiline loaches (Pisces: Cypriniformes) of Thailand, Burma, Laos, Cambodia and southern Viet Nam. Verlag Dr. Friedrich Pfeil, München, 262 pp.
    Kottelat, M. (1997) European freshwater fishes. An heuristic checklist of the freshwater fishes of Europe (exclusive of former USSR), with an introduction for non-systematists and comments on nomenclature and conservation. Biologia, Bratislava, Section Zoology, 52 (Supplement 5), 1–271.
    Kottelat, M. (2012) Conspectus cobitidum: an inventory of the loaches of the world (Teleostei: Cypriniformes: Cobitoidei). The Raffles Bulletin of Zoology, 26 (Supplement), 1–199.
    Kottelat, M. & Freyhof, J. (2007) Handbook of European freshwater fishes. Kottelat, Cornol and Freyhof, Berlin, xiv + 646 pp.
    Kumar, S., Stecher, G., Li, M., Knyaz, C. & Tamura, K. (2018) MEGA X: Molecular Evolutionary Genetics Analysis across computing platforms. Molecular Biology and Evolution, 35, 1547–1549. https://doi.org/10.1093/molbev/msy096
    Kunz, W. (2007) Do species exist? Principles of taxonomic classification. Wiley-Blackwell, Weinheim, xxxiii + 245 pp.
    Kwong, S., Srivathsan, A., Vaidya, G. & Meier, R. (2012) Is the COI barcoding gene involved in speciation through intergenomic conflict? Molecular Phylogenetics and Evolution, 62 1009–1012. https://doi.org/10.1016/j.ympev.2011.11.034
    John, L., Philip, S., Dahanukar, N., Hamsa Anvar Ali, P., Tharian, J., Raghavan, R. & Antunes, A. (2013) Morphological and genetic evidence for multiple evolutionary distinct lineages in the endangered and commercially exploited red lined torpedo barbs endemic to the Western Ghats of India. PLoS ONE, 8 (8), 10.1371/annotation/bf1622ea-bb3e-49f7-9d16-f549c05fc584. https://doi.org/10.1371/journal.pone.0069741
    Lavoué, S., Miya, M., Arnegard, M.E., McIntyre, P.B., Mamonekene V. & Nishida, M. (2010) Remarkable morphological stasis in an extant vertebrate despite tens of millions of years of divergence. Proceedings of the Royal Society, B. 2781003–1008. https://doi.org/10.1098/rspb.2010.1639
    Lorenzoni, M., Carosi, A., Quadroni, S., De Santis, V., Vanetti, I., Delmastro, G.B. & Zaccara, S. (2021) Cryptic diversity within endemic Italian barbels: revalidation and description of new Barbus species (Teleostei: Cyprinidae). Journal of Fish Biology, 98 (5), 1433–1449. https://doi.org/10.1111/jfb.14688
    Lucentini, L., Puletti, M.E., Ricciolini, C., Gigliarelli, L., Fontaneto, D., Lanfaloni, L., Bilò, F., Natali, M. & Panara, F. (2011) Molecular and phenotypic evidence of a new species of genus Esox (Esocidae, Esociformes, Actinopterygii): the southern pike, Esox flaviae. PLoS ONE, 6 (12), 1–14. https://doi.org/10.1371/journal.pone.0025218
    Masters, B.C., Fan, V. & Ross, H.A. (2011) Species delimitation – a geneious plugin for the exploration of species boundaries. Molecular Ecology Resources, 11, 154–157. https://doi.org/ 10.1111/j.1755-0998.2010.02896.x
    Mateus, C.S., Alves, M.J., Quintella, B.R. & Almeida, P.R. (2013) Three new cryptic species of the lamprey genus Lampetra Bonnaterre, 1788 (Petromyzontiformes: Petromyzontidae) from the Iberian Peninsula. Contributions to Zoology, 82 (1), 37–53. https://doi.org/10.1371/journal.pone.0025218
    Mayden, R.L. (1997) A hierarchy of species concepts: the denouement in the saga of the species problem. In: Claridge, M.F., Dawah, H.A. & Wilson, M.R., Species: The Units of Biodiversity. Chapman & Hall, London, pp. 381–424
    Meier, R., Shiyang, K., Vaidya, G., & Ng, P.K.L. (2006) DNA Barcoding and Taxonomy in Diptera: A Tale of High Intraspecific Variability and Low Identification Success. Systematic Biology, 55 (5), 715–728. https://doi.org/10.1080/10635150600969864
    Palandačić, A., Naseka, A., Ramler D. & Ahnelt, H. (2017) Contrasting morphology with molecular data: an approach to revision of species complexes based on the example of European Phoxinus (Cyprinidae). BMC Evolutionary Biology 17 (184), 1–17. https://doi.org/10.1186/s12862-017-1032-x
    Pentinsaari, M., Vos, R. & Mutanen, M. (2017) Algorithmic single-locus species delimitation: effects of sampling effort, variation and nonmonophyly in four methods and 1870 species of beetles. Molecular Ecology Resources, 17 (3), 393–404.
    https://doi.org/10.1111/1755-0998.12557
    Pereira, L.H.G., Hanner, R., Foresti, F. & Oliveira, C. (2013) Can DNA barcoding accurately discriminate megadiverse Neotropical freshwater fish fauna? BMC Genetics, 14, 20. https://doi.org/10.1186/1471-2156-14-20
    Puillandre, N., Brouillet, S. & Achaz, G. (2021). ASAP: Assemble species by automatic partitioning. Molecular Ecology Resources, 21, 609–620. https://doi.org/10.1111/1755-0998.13281
    Puillandre, N., Lambert, A., Brouillet, S. & Achaz, G. (2012). ABGD, Automatic Barcode Gap Discovery for primary species delimitation. Molecular Ecology, 21, 1864–1877. https://doi.org/10.1111/j.1365-294X.2011.05239.x
    Riedel, A. & Narakusumo, R.P. (2019) One hundred and three new species of Trigonopterus weevils from Sulawesi. ZooKeys, 828, 1–153. https://doi.org/10.3897/zookeys.828.32200
    Riedel, A., Sagata, K., Suhardjono, Y.R., Tänzler, R. & Balke, M. (2013) Integrative taxonomy on the fast track – towards more sustainability in biodiversity research. Frontiers in Zoology, 10, 15. https://doi.org/10.1186/1742-9994-10-15
    Saitou, N. & Nei, M. (1987) The neighbor-joining method: a new method for reconstructing phylogenetic trees. Molecular Biology and Evolution, 4 (4), 406–425. https://doi.org/10.1093/oxfordjournals.molbev.a040454
    Saygun, S., Ağdamar, S. & Özuluğ, M. (2021) Oxynoemacheilus fatsaensis, a new nemacheilid loach from the Elekçi stream in northern Anatolia (Teleostei: Nemacheilidae). Zoologischer Anzeiger, 294, 39–49. https://doi.org/10.1016/j.jcz.2021.07.011
    Sayyadzadeh, G., Eagderi, S. & Esmaeili, H.R. (2016) A new loach of the genus Oxynoemacheilus from the Tigris River drainage and its phylogenetic relationships among the nemacheilid fishes (Teleostei: Nemacheilidae) in the Middle East based on mtDNA COI sequences. Iranian Journal of Ichthyology, 3, 236–250. https://doi.org/10.22034/iji.v3i4.205
    Sayyadzadeh, G. & Esmaeili, H.R. (2020) Oxynoemacheilus marunensis, a new loach species from the Persian Gulf basin with remarks on O. frenatus (Teleostei: Nemacheilidae). Zootaxa, 4885 (2), 189–206. https://doi.org/10.11646/zootaxa.4885.2.2
    Sayyadzadeh, G.; Esmaeili, H.R., Eagderi, S., Jouladeh-Roudbar, A.; Masoudi, M. & Vatandoust, S. (2017) Re-description of Oxynoemacheilus longipinnis from the Persian Gulf basin (Teleostei: Nemacheilidae). Zoology in the Middle East, 63 (3), 228–238. https://doi.org/10.1080/09397140.2017.1349243
    Schindel, D.E. & Miller, S.E. (2005) DNA barcoding a useful tool for taxonomists. Nature, 435, 17.
    Stankowski, S. & Ravinet, M. (2021) Quantifying the use of species concepts. Current Biology, 31, (9), R428–R429. https://doi.org/10.1016/j.cub.2021.03.060
    Sharkey, M.J., Janzen, D.H., Hallwachs, W., Chapman, E.G., Smith, M.A., Dapkey, T., Brown, A., Ratnasingham, S., Naik, S., Manjunath, R., Perez, K., Milton, M., Hebert, P., Sgaw, S.R., Kittel, R.N., Solis, M.A. Metz, M.A., Goldstein, P.Z., Brown, J.W., Quicke, D.L.J., van Achterberg, C., Brown, B.V. & Burns, J.M. (2021) Minimalist revision and description of 403 new species in 11 subfamilies of Costa Rican braconid parasitoid wasps, including host records for 219 species. ZooKeys, 1013, 1–666. https://doi.org/10.3897/zookeys.1013.55600
    Srivathsan, A., Hartop, E., Puniamorrthy, J., Lee, W.T., Kutty, S.N., Kurina, O. & Meier, R. (2019) Rapid, large-scale species discovery in hyperdiverse taxa using 1D MinION sequencing. BMC Biology, 17, 96. https://doi.org/10.1186/s12915-019-0706-9
    Struck, T.H & Cerca, J. (2019) Cryptic species and their evolutionary significance. In: eLS. John Wiley & Sons, Ltd, Chichester. [published online] https://doi.org/10.1002/9780470015902.a0028292
    Sudasinghe, H., Pethiyagoda, R., Meegaskumbura, M., Maduwage, K. & Britz, R. (2020) Channa kelaartii, a valid species of dwarf snakehead from Sri Lanka and southern peninsular India (Teleostei: Channidae). Vertebrate Zoology, 70 (2), 157–170. https://doi.org/10.26049/VZ70-2-2020-05
    Sungur, S., Jalili, P. & Eagderi, S. (2017) Oxynoemacheilus ciceki, new nemacheilid species (Teleostei, Nemacheilidae) from the Sultan Marsh, Kayseri Province, Turkey. Iranian Journal of Ichthyology 4 (4), 375–383. https://doi.org/10.22034/iji.v4i4.258
    Swofford, D.L. (2002) PAUP*. Phylogenetic analysis using parsimony (*and other methods). Version 4. Sinauer Associates, Sunderland, Massachusetts.
    Tautz, D., Arctander, P., Minelli, A., Thomas, R.H. & Vogler, A.P. (2003) A plea for DNA taxonomy. Trends in Ecology and Evolution, 18, 70–74. https://doi.org/10.1016/S0169-5347(02)00041-1
    Turan, D., Kaya, C., Kalaycı, G., Bayçelebi, E. & Aksu, İ. (2019) Oxynoemacheilus cemali, a new species of stone loach (Teleostei: Nemacheilidae) from the Çoruh River drainage. Journal of Fish Biology, 94 (3), 458–468. https://doi.org/10.1111/jfb.13909
    Yeates, D.K., Seago, A., Nelson, L., Cameron, S.L., Joseph, L. & Trueman, J.W.H. (2011) Integrative taxonomy, or iterative taxonomy? Systematic Entomology, 36, 209–217. https://doi.org/10.1111/j.1365-3113.2010.00558.x
    Yoğurtçuoğlu, B. & Freyhof, J. (2018) Aphanius irregularis, a new killifish from south-western Anatolia (Cyprinodontiformes: Aphaniidae). Zootaxa, 4410 (2), 319–330. https://doi.org/10.11646/zootaxa.4410.2.4
    Yoğurtçuoğlu, B., Kaya, C., Geiger, M.F. & Freyhof, J. (2020) Revision of the genus Seminemacheilus, with the description of three new species (Teleostei: Nemacheilidae). Zootaxa, 4802 (3), 477–501. https://doi.org/10.11646/zootaxa.4802.3.5 
    Yoğurtçuoğlu, B., Kaya, C. & Freyhof, J. (2021a) Oxynoemacheilus nasreddini, a new nemacheilid loach from the Central Anatolia (Teleostei: Nemacheilidae). Zootaxa, 4974 (1), 135–150. https://doi.org/10.11646/zootaxa.4974.1.5
    Yoğurtçuoğlu, B., Kaya, C., Özuluğ, M. & Freyhof, J. (2021b) Oxynoemacheilus isauricus, a new nemacheilid loach from Central Anatolia (Teleostei: Nemacheilidae). Zootaxa, 4975 (2), 369–378. https://doi.org/10.11646/zootaxa.4975.2.7
    Zhang, J., Kapli, P., Pavlidis, P. & Stamatakis, A. (2013) A general species delimitation method with applications to phylogenetic placements. Bioinformatics, 29 (22), 2869–2876. https://doi.org/10.1093/bioinformatics/btt499
    Zhou, Z., Guo, H., Han, L., Chai, J., Che, X. & Shi, F. (2019) Singleton molecular species delimitation based on COI-5P barcode sequences revealed high cryptic/undescribed diversity for Chinese katydids (Orthoptera: Tettigoniidae). BMC Evolutionary Biology, 19, 79. https://doi.org/10.1186/s1286 2-019-1404-5

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