Main Article Content
Abstract
The Hormuz River System (HRS) is located in the southern coastal plain of the Zagros Mountains. Two main river drainages i.e., Mehran and Kol are known from this region, which are directly discharged to the Persian Gulf. The HRS has surprising diversity for tooth-carp fishes (family Aphaniidae). The present study aims to accentuate this notable diversity and discuss the possible cause of diversification of the tooth-carps in the HRS. To date, 15 Aphaniid species have been discovered from the Iranian drainage basins. Among them, 10 species belong to the genus Aphanius, four species belong to the genus Aphaniops, and a single species belonging to the genus Paraphanius. The HRS by having four species (22% of the Iranian tooth-carps) demonstrated the highest diversity among the other Iranian basins. Of these, one belongs to the genus Aphanius (A. darabensis) and three species belong to the genus Aphaniops (A. furcatus, A. ginaonis, and A. hormuzensis). By considering the degree of endemicity of the tooth-carps fauna relative to the territorial boundaries of drainages, the HRS has the highest endemicity, in which 31% of the Iranian endemic tooth-carp species present only in the HRS. The last connection of the Makran Basin with the HRS dates back to the late Pliocene and lower Pleistocene about 1.8 million years ago. Also, the other vicariance-based events like those events related to the last glacial maximum (21000-18000 BP), population migration during the Early Holocene sea-level rise, and the Holocene to present-day have promoted the intra- and interspecific differentiation of tooth-carps in the HRS. Also, the geological situation of the HRS provided various extreme environmental conditions, which caused a particular increase in the diversification of the tooth-carp species.
Keywords
Aphanius
Aphaniops
Paraphanius
Extreme ecological condition
Geological history.
Article Details
License
The authors transfer the copyrights of their papers to the Iranian Society of Ichthyology. However, the information could be used in accordance with the Creative Commons licence (Attribution-NonCommercial 4.0 International (CC BY-NC 4.0)
How to Cite
TEIMORI, A., & ESMAEILI, H. R. (2020). Hormuz River System as a micro hotspot of diversification for the aphaniid fishes in Iran. Iranian Journal of Ichthyology, 7(2), 181–196. https://doi.org/10.22034/iji.v7i2.505
References
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Ariyaee, M.; Hamidian, A.H.; Eagderi, S.; Ashrafi, S. & Poorbagher, H. 2018. Cadmium and arsenic bioaccumulation and bio-concentration in the endemic toothed carp Aphanius arakensis in salt water. International Journal of Aquatic Biology 6(1): 25-30.
Barton, N.H. & Charlesworth, B. 1984. Genetic revolutions, founder effects, and speciation. Annual Review of Ecology and Systematics 15: 133-164.
Buj, I.; Miočić-Stošić, J.; Marčić, Z.; Mustafić, P.; Zanella, D.; Mrakovčić, M. &. Ćaleta, M. 2015. Population genetic structure and demographic history of Aphanius fasciatus (Cyprinodontidae: Cyprinodontiformes) from hypersaline habitats in the eastern Adriatic. Scientia Marina 79: 399-408.
Cavraro, F.; Malavasi, S.; Torricelli, P.; Gkenas, C.; Liousia, V.; Leonardos, I. &…Triantafyllidis, A. 2017 Genetic structure of the South European toothcarp Aphanius fasciatus (Actinopterygii: Cyprinodontidae) populations in the Mediterranean basin with a focus on the Venice lagoon. Search Results Web results. European Zoological Journal 84: 153-166.
Coad, B.W. 1980. A re-description of Aphanius ginaonis (Holly, 1929) from southern Iran (Osteichthyes; Cyprinodontiformes). Journal of Natural Histology 14: 33-40.
Coad, B.W. 1996. Fishes from the qanats of Iran. Publicaciones Especiales Instituto Español de Oceanografía 21: 63-79.
Coad, B.W. 1998. Threatened fishes of the world: Lebias ginaonis (Holly, 1929) (Cyprinodontidae). Environmental Biology of Fishes 51(3): 284.
Coad, B.W. 2000. Distribution of Aphanius species in Iran. Journal of the American Killifish Association 33: 183-191.
Coad, B. 2009. A new species of tooth-carp, Aphanius mesopotamicus, from Iran and Iraq (Actinopterygii, Cyprinodontidae). Zookeys 31:149.
Day, F. 1872. Notes on fish, collected by Dr. Stoliczka in Kachh. Journal of the Asiatic Society of Bengal 41: 1-4.
Echelle, A.A. & Echelle, A.F. 1992. Mode and pattern of speciation in the evolution of Inland pupfishes in the Cyprinodon variegatus complex (Teleostei: Cyprinodontidae): An ancestor-descendent hypothesis, in Systematics, Historical Ecology, and North American Freshwater Fishes, Mayden, R.L., Eds., Stanford, Stanford University Press California, pp. 691-709.
Esmaeili, H.R. & Gholami, Z. 2007. Investigations on the surface ultrastructure of scale of Geno tooth-carp, Aphanius ginaonis (Holly, 1929) (Actinopterygii: cyprinodontidae) using scanning electron microscope. Iranian Journal of Biology 20: 307-314.
Esmaeili, H.R.; Coad, B.W.; Gholamifard, A.; Nazari, N. & Teimori, A. 2010. Annotated сhecklist of the freshwater fishes of Iran. Zoosystematica Rossica 19: 361-386.
Esmaeili, H.R.; Teimori, A.; Sayyadzadeh, G.; Masoudi, M. & Reichenbacher, B. 2014a. Phylogenetic relationships of the tooth-carp Aphanius (Teleostei: Cyprinodontidae) in the river systems of southern and south-western Iran based on mtDNA sequences. Zoology in the Middle East 60: 29-38.
Esmaeili, H.R.; Teimori, A.; Gholami, Z. & Reichenbacher, B. 2014b. Two new species of the tooth-carp Aphanius (Teleostei: Cyprinodontidae) and the evolutionary history of the Iranian inland and inland-related Aphanius species. Zootaxa 3786: 246-268.
Esmaeili, H.R.; Sayyadzadeh, G. & Seehausen, O. 2016. Iranocichla persa, a new cichlid species from southern Iran (Teleostei, Cichlidae). ZooKeys 636: 141.
Esmaeili, H.R.; Sayyadzadeh, G.; Eagderi, S. & Abbasi, K. 2018. Checklist of freshwater fishes of Iran. FishTaxa 3: 1-95.
Esmaeili, H.R.; Teimori, A.; Zarei, F. & Sayyadzadeh, G. 2020. DNA barcoding and species delimitation of the Old-World tooth-carps, family Aphaniidae Hoedeman, 1949 (Teleostei: Cyprinodontiformes). PLoS ONE 15(4): e0231717.
Ferrito, V.; Mannino, M.C.; Pappalardo, A.M. & Tigano, C. 2007. Morphological variation between Italian populations of Aphanius fasciatus Nardo, 1827 (Teleostei, Cyprinodontidae). Journal of Fish Biology 70: 1-20.
Freyhof, J.; Weissenbacher, A. & Geiger, M. 2017. Aphanius kruppi, a new killifish from Oman with comments on the A. dispar species group (Cyprinodontiformes: Aphaniidae). Zootaxa 4338: 557-573.
Gaudant, J. 1993. Un exemple de “régression évolutive” chez des poissons Cyprinodontidae du Miocène supérieur d'Espagne: Aphanius illunensis nov. sp. Geobios 26: 449-454.
Grimm, H. 1980. Investigations on the problem of scale reduction and sulphate tolerance of West Anatolian cyprinodonts (Pisces). Internationale Revue der Gesamten Hydrobiologie und Hydrographie 65: 517-533.
Hatzfeld, D.; Authemayou, C.; Van Der Beek, P.; Bellier, O.; Lavé, J.; Oveisi, B.; Tatar, M.; Tavakoli, F.; Walpersdorf, A. & Yamini-Fard, F. 2010. The kinematics of the Zagros mountains (Iran). Geological Society, London, Special Publications 330: 19-42.
Heckel, J.J. 1849. Über eine neue fossile Fishgattung, Chirocentrites, und die ersten Ueberreste eines Siluroiden aus der Vorwelt. Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften, Mathematisch-Naturwissenschaftlichen Classe Wien, 2: 16-19.
Holly, L.F. Holly Pneumatic Systems Inc. 1929. Apparatus and method for loading loose-bulk cargoes. U.S. Patent 1,708,176.
Hrbek, T. & Meyer, A. 2003. Closing of the Tethys Sea and the phylogeny of Eurasian killifishes (Cyprinodontiformes: Cyprinodontidae). Journal of Evolutionary Biology 16: 17-36.
Hrbek, T.; Küçük, F.; Frickey, T.; Stölting, K.N.; Wildekamp, R.H. & Meyer, A. 2002. Molecular phylogeny and historical biogeography of the Aphanius (Pisces, Cyprinodontiformes) species complex of central Anatolia, Turkey. Molecular Phylogenetics and Evolution 25: 125-137.
Hrbek, T.; Keivany, Y. & Coad, B.W. 2006. New species of Aphanius (Teleostei, Cyprinodontidae) from Isfahan Province of Iran and a reanalysis of other Iranian species. Copeia 2006: 244-255.
Iranmanesh, A. & Motamedi, M. 2018. Histopathological changes of hepatorenal toxicity induced by gentamicin in killifish, Aphanius hormuzensis (Aphaniidae) and its kidney regeneration through nephron neogenesis. Journal of Ichthyology 58: 932-938.
Jenkins, J.T. 1910. XII. Notes on fish from India and Persia, with descriptions of new species. Records of the Indian Museum 10: 123-140.
Motamedi, M.; Shamsaldini, F.; Teimori, A. & Hesni, M.A. 2019a. Histomicroscopy and normal anatomy of the adult killifish Aphanius hormuzensis (Teleostei; Aphaniidae) from the Persian Gulf coastal environment. Microscopy Research and Technique 82: 466-480.
Motamedi, M.; Teimori, A. & Iranmanesh, A. 2019b. Ontogenetic pattern, morphological sexual and side dimorphism in the saccular otolith of a scaleless killifish Aphanius furcatus (Teleostei: Aphaniidae). Acta Zoologica. (In Press)
Nasrolah Pourmoghadam, M.; Poorbagher, H. & Eagderi, S. 2014. Histopathological effects of cypermethrin on liver of Aphanius sophiae (Heckel, 1849) using rank-based estimation for linear models. International Journal of Aquatic Biology 2(4): 184-187.
Parenti, L.R. 1981. A phylogenetic and biogeographic analysis of cyprinodontiform fishes (Teleostei, Atherinomorpha). Bulletin of the American Museum of Natural History. 168: 335-557.
Porkhial, S.; Nouraliee, J.; Rahmani, M. & Ebrahimi, D. 2013. Resource assessment of Vartun geothermal region, central Iran. Journal of Tethys 1: 29-40.
Regard, V.; Bellier, O.; Thomas, J.C.; Abbassi, M.R.; Mercier, J.; Shabanian, E.; Feghhi, K.H. & Soleymani, S. 2004. Accommodation of Arabia-Eurasia convergence in the Zagros-Makran transfer zone, SE Iran: A transition between collision and subduction through a young deforming system. Tectonics 23: TC4007
Reichenbacher, B.; Kamrani, E.; Esmaeili, H.R. & Teimori, A. 2009. The endangered cyprinodont Aphanius ginaonis (Holly, 1929) from southern Iran is a valid species: evidence from otolith morphology. Environmental Biology of Fishes 86(4): 507.
Rocco, L.; Ferrito, V.; Costagliola, D.; Marsilio, A.; Pappalardo, A.M.; Stingo, V. & Tigano, C. 2007. Genetic divergence among and within four Italian populations of Aphanius fasciatus (Teleostei, Cyprinodontiformes). Italian Journal of Zoology 74: 371-379.
Schwarzer, J.; Shabani, N.; Esmaeili, H.R.; Mwaiko, S. & Seehausen, O. 2017. Allopatric speciation in the desert: diversification of cichlids at their geographical and ecological range limit in Iran. Hydrobiologia 791: 193-207.
Sienknecht, U.1999. Kreuzungsgenetische Analyse des Uberganges von Paarflossen bis zu fehlendem Extremitatengurtel: entwickeltamBeispiel der normogenetischen Ventralflossenbildung von Aphanius iberus Cuv and Val 1846 im Vergleich zur Anormogenese seiner Hybriden mit A. apodus Gervais, 1853. Cyprinodontidae Der Andere Verlag, Bad Iburg, Teleostei.
Teimori, A.; Schulz-Mirbach, T.; Esmaeili, H.R. & Reichenbacher, B. 2012a. Geographical differentiation of Aphanius dispar (Teleostei: Cyprinodontidae) from southern Iran. Journal of Zoological Systematics and Evolutionary Research 50: 289-304.
Teimori, A.; Esmaeili, H.R.; Gholami, Z.; Zarei, N. & Reichenbacher, B. 2012b. Aphanius arakensis, a new species of tooth-carp (Actinopterygii, Cyprinodontidae) from the endorheic Namak Lake basin in Iran. ZooKeys 215: 55–76.
Teimori, A.; Jawad, L.A.J.; Al-Kharusi, L.H.; Al-Mamry, J.M. & Reichenbacher, B. 2012c. Late Pleistocene to Holocene diversification and historical zoogeography of the Arabian killifish (Aphanius dispar) inferred from otolith morphology. Scientia Marina 76: 637-645.
Teimori, A.; Esmaeili, H.R.; Erpenbeck, D. & Reichenbacher, B. 2014. A new and unique species of the genus Aphanius Nardo, 1827 (Teleostei: Cyprinodontidae) from Southern Iran: A case of regressive evolution. Zoologischer Anzeiger-A Journal of Comparative Zoology 253: 327-337.
Teimori, A.; Esmaeili, H.R.; Hamidan, N. & Reichenbacher, B. 2018. Systematics and historical biogeography of the Aphanius dispar species group (Teleostei: Aphaniidae) and description of a new species from Southern Iran. Journal of Zoological Systematics and Evolutionary Research 56: 579-598.
Teimori, A.; Ebrahimi, M.; Mostafavi, H.; Esmaeili, H.R. & Masoudi, M. 2020. Distribution and abundance of the endemic scaleless tooth-carp, Aphanius furcatus Teimori, Esmaeili, Erpenbeck & Reichenbacher, 2014. Turkish Journal of Fisheries and Aquatic Science 20: 61-67.
Tigano, C.; Canapa, A.; Ferrito, V., Barucca, M.; Arcidiacono, I.; Deidun, A.; Schembri, P.J. & Olmo, E. 2006. A study of osteological and molecular differences in populations of Aphanius fasciatus Nardo 1827, from the central Mediterranean (Teleostei, Cyprinodontidae). Marine Biology 149: 1539-1550.
Villwock, V.W. 1964. Genetische Untersuchungen an altweltlichen Zahnkarpfen der Tribus Aphaniini (Pisces, Cyprinodontidae) nach Gesichtspunkten der Neuen Systematik. Journal of Zoological Systematics and Evolutionary Research 2: 267-382.
Wildekamp, R.H. 1993. A world of killies: atlas of the oviparous cyprinodontiform fishes of the world. American Killifish Association.
Wildekamp, R.H.; Küçük, F.; Ünlüsayin, M. & Van Neer, W. 1999. Species and subspecies of the genus Aphanius Nardo 1897 (Pisces: Cyprinodontidae) in Turkey. Turkish Journal of Zoology 23: 23-44.
Zeinali, F. & Motamedi, M. 2017. The regeneration capacity of caudal fin in the common tooth-carp, Aphanius dispar (Rüppell, 1829) (Teleostei: Cyprinodontidae). International Journal of Aquatic Biology 5: 321-327.
References
Ariyaee, M.; Hamidian, A.H.; Eagderi, S.; Ashrafi, S. & Poorbagher, H. 2018. Cadmium and arsenic bioaccumulation and bio-concentration in the endemic toothed carp Aphanius arakensis in salt water. International Journal of Aquatic Biology 6(1): 25-30.
Barton, N.H. & Charlesworth, B. 1984. Genetic revolutions, founder effects, and speciation. Annual Review of Ecology and Systematics 15: 133-164.
Buj, I.; Miočić-Stošić, J.; Marčić, Z.; Mustafić, P.; Zanella, D.; Mrakovčić, M. &. Ćaleta, M. 2015. Population genetic structure and demographic history of Aphanius fasciatus (Cyprinodontidae: Cyprinodontiformes) from hypersaline habitats in the eastern Adriatic. Scientia Marina 79: 399-408.
Cavraro, F.; Malavasi, S.; Torricelli, P.; Gkenas, C.; Liousia, V.; Leonardos, I. &…Triantafyllidis, A. 2017 Genetic structure of the South European toothcarp Aphanius fasciatus (Actinopterygii: Cyprinodontidae) populations in the Mediterranean basin with a focus on the Venice lagoon. Search Results Web results. European Zoological Journal 84: 153-166.
Coad, B.W. 1980. A re-description of Aphanius ginaonis (Holly, 1929) from southern Iran (Osteichthyes; Cyprinodontiformes). Journal of Natural Histology 14: 33-40.
Coad, B.W. 1996. Fishes from the qanats of Iran. Publicaciones Especiales Instituto Español de Oceanografía 21: 63-79.
Coad, B.W. 1998. Threatened fishes of the world: Lebias ginaonis (Holly, 1929) (Cyprinodontidae). Environmental Biology of Fishes 51(3): 284.
Coad, B.W. 2000. Distribution of Aphanius species in Iran. Journal of the American Killifish Association 33: 183-191.
Coad, B. 2009. A new species of tooth-carp, Aphanius mesopotamicus, from Iran and Iraq (Actinopterygii, Cyprinodontidae). Zookeys 31:149.
Day, F. 1872. Notes on fish, collected by Dr. Stoliczka in Kachh. Journal of the Asiatic Society of Bengal 41: 1-4.
Echelle, A.A. & Echelle, A.F. 1992. Mode and pattern of speciation in the evolution of Inland pupfishes in the Cyprinodon variegatus complex (Teleostei: Cyprinodontidae): An ancestor-descendent hypothesis, in Systematics, Historical Ecology, and North American Freshwater Fishes, Mayden, R.L., Eds., Stanford, Stanford University Press California, pp. 691-709.
Esmaeili, H.R. & Gholami, Z. 2007. Investigations on the surface ultrastructure of scale of Geno tooth-carp, Aphanius ginaonis (Holly, 1929) (Actinopterygii: cyprinodontidae) using scanning electron microscope. Iranian Journal of Biology 20: 307-314.
Esmaeili, H.R.; Coad, B.W.; Gholamifard, A.; Nazari, N. & Teimori, A. 2010. Annotated сhecklist of the freshwater fishes of Iran. Zoosystematica Rossica 19: 361-386.
Esmaeili, H.R.; Teimori, A.; Sayyadzadeh, G.; Masoudi, M. & Reichenbacher, B. 2014a. Phylogenetic relationships of the tooth-carp Aphanius (Teleostei: Cyprinodontidae) in the river systems of southern and south-western Iran based on mtDNA sequences. Zoology in the Middle East 60: 29-38.
Esmaeili, H.R.; Teimori, A.; Gholami, Z. & Reichenbacher, B. 2014b. Two new species of the tooth-carp Aphanius (Teleostei: Cyprinodontidae) and the evolutionary history of the Iranian inland and inland-related Aphanius species. Zootaxa 3786: 246-268.
Esmaeili, H.R.; Sayyadzadeh, G. & Seehausen, O. 2016. Iranocichla persa, a new cichlid species from southern Iran (Teleostei, Cichlidae). ZooKeys 636: 141.
Esmaeili, H.R.; Sayyadzadeh, G.; Eagderi, S. & Abbasi, K. 2018. Checklist of freshwater fishes of Iran. FishTaxa 3: 1-95.
Esmaeili, H.R.; Teimori, A.; Zarei, F. & Sayyadzadeh, G. 2020. DNA barcoding and species delimitation of the Old-World tooth-carps, family Aphaniidae Hoedeman, 1949 (Teleostei: Cyprinodontiformes). PLoS ONE 15(4): e0231717.
Ferrito, V.; Mannino, M.C.; Pappalardo, A.M. & Tigano, C. 2007. Morphological variation between Italian populations of Aphanius fasciatus Nardo, 1827 (Teleostei, Cyprinodontidae). Journal of Fish Biology 70: 1-20.
Freyhof, J.; Weissenbacher, A. & Geiger, M. 2017. Aphanius kruppi, a new killifish from Oman with comments on the A. dispar species group (Cyprinodontiformes: Aphaniidae). Zootaxa 4338: 557-573.
Gaudant, J. 1993. Un exemple de “régression évolutive” chez des poissons Cyprinodontidae du Miocène supérieur d'Espagne: Aphanius illunensis nov. sp. Geobios 26: 449-454.
Grimm, H. 1980. Investigations on the problem of scale reduction and sulphate tolerance of West Anatolian cyprinodonts (Pisces). Internationale Revue der Gesamten Hydrobiologie und Hydrographie 65: 517-533.
Hatzfeld, D.; Authemayou, C.; Van Der Beek, P.; Bellier, O.; Lavé, J.; Oveisi, B.; Tatar, M.; Tavakoli, F.; Walpersdorf, A. & Yamini-Fard, F. 2010. The kinematics of the Zagros mountains (Iran). Geological Society, London, Special Publications 330: 19-42.
Heckel, J.J. 1849. Über eine neue fossile Fishgattung, Chirocentrites, und die ersten Ueberreste eines Siluroiden aus der Vorwelt. Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften, Mathematisch-Naturwissenschaftlichen Classe Wien, 2: 16-19.
Holly, L.F. Holly Pneumatic Systems Inc. 1929. Apparatus and method for loading loose-bulk cargoes. U.S. Patent 1,708,176.
Hrbek, T. & Meyer, A. 2003. Closing of the Tethys Sea and the phylogeny of Eurasian killifishes (Cyprinodontiformes: Cyprinodontidae). Journal of Evolutionary Biology 16: 17-36.
Hrbek, T.; Küçük, F.; Frickey, T.; Stölting, K.N.; Wildekamp, R.H. & Meyer, A. 2002. Molecular phylogeny and historical biogeography of the Aphanius (Pisces, Cyprinodontiformes) species complex of central Anatolia, Turkey. Molecular Phylogenetics and Evolution 25: 125-137.
Hrbek, T.; Keivany, Y. & Coad, B.W. 2006. New species of Aphanius (Teleostei, Cyprinodontidae) from Isfahan Province of Iran and a reanalysis of other Iranian species. Copeia 2006: 244-255.
Iranmanesh, A. & Motamedi, M. 2018. Histopathological changes of hepatorenal toxicity induced by gentamicin in killifish, Aphanius hormuzensis (Aphaniidae) and its kidney regeneration through nephron neogenesis. Journal of Ichthyology 58: 932-938.
Jenkins, J.T. 1910. XII. Notes on fish from India and Persia, with descriptions of new species. Records of the Indian Museum 10: 123-140.
Motamedi, M.; Shamsaldini, F.; Teimori, A. & Hesni, M.A. 2019a. Histomicroscopy and normal anatomy of the adult killifish Aphanius hormuzensis (Teleostei; Aphaniidae) from the Persian Gulf coastal environment. Microscopy Research and Technique 82: 466-480.
Motamedi, M.; Teimori, A. & Iranmanesh, A. 2019b. Ontogenetic pattern, morphological sexual and side dimorphism in the saccular otolith of a scaleless killifish Aphanius furcatus (Teleostei: Aphaniidae). Acta Zoologica. (In Press)
Nasrolah Pourmoghadam, M.; Poorbagher, H. & Eagderi, S. 2014. Histopathological effects of cypermethrin on liver of Aphanius sophiae (Heckel, 1849) using rank-based estimation for linear models. International Journal of Aquatic Biology 2(4): 184-187.
Parenti, L.R. 1981. A phylogenetic and biogeographic analysis of cyprinodontiform fishes (Teleostei, Atherinomorpha). Bulletin of the American Museum of Natural History. 168: 335-557.
Porkhial, S.; Nouraliee, J.; Rahmani, M. & Ebrahimi, D. 2013. Resource assessment of Vartun geothermal region, central Iran. Journal of Tethys 1: 29-40.
Regard, V.; Bellier, O.; Thomas, J.C.; Abbassi, M.R.; Mercier, J.; Shabanian, E.; Feghhi, K.H. & Soleymani, S. 2004. Accommodation of Arabia-Eurasia convergence in the Zagros-Makran transfer zone, SE Iran: A transition between collision and subduction through a young deforming system. Tectonics 23: TC4007
Reichenbacher, B.; Kamrani, E.; Esmaeili, H.R. & Teimori, A. 2009. The endangered cyprinodont Aphanius ginaonis (Holly, 1929) from southern Iran is a valid species: evidence from otolith morphology. Environmental Biology of Fishes 86(4): 507.
Rocco, L.; Ferrito, V.; Costagliola, D.; Marsilio, A.; Pappalardo, A.M.; Stingo, V. & Tigano, C. 2007. Genetic divergence among and within four Italian populations of Aphanius fasciatus (Teleostei, Cyprinodontiformes). Italian Journal of Zoology 74: 371-379.
Schwarzer, J.; Shabani, N.; Esmaeili, H.R.; Mwaiko, S. & Seehausen, O. 2017. Allopatric speciation in the desert: diversification of cichlids at their geographical and ecological range limit in Iran. Hydrobiologia 791: 193-207.
Sienknecht, U.1999. Kreuzungsgenetische Analyse des Uberganges von Paarflossen bis zu fehlendem Extremitatengurtel: entwickeltamBeispiel der normogenetischen Ventralflossenbildung von Aphanius iberus Cuv and Val 1846 im Vergleich zur Anormogenese seiner Hybriden mit A. apodus Gervais, 1853. Cyprinodontidae Der Andere Verlag, Bad Iburg, Teleostei.
Teimori, A.; Schulz-Mirbach, T.; Esmaeili, H.R. & Reichenbacher, B. 2012a. Geographical differentiation of Aphanius dispar (Teleostei: Cyprinodontidae) from southern Iran. Journal of Zoological Systematics and Evolutionary Research 50: 289-304.
Teimori, A.; Esmaeili, H.R.; Gholami, Z.; Zarei, N. & Reichenbacher, B. 2012b. Aphanius arakensis, a new species of tooth-carp (Actinopterygii, Cyprinodontidae) from the endorheic Namak Lake basin in Iran. ZooKeys 215: 55–76.
Teimori, A.; Jawad, L.A.J.; Al-Kharusi, L.H.; Al-Mamry, J.M. & Reichenbacher, B. 2012c. Late Pleistocene to Holocene diversification and historical zoogeography of the Arabian killifish (Aphanius dispar) inferred from otolith morphology. Scientia Marina 76: 637-645.
Teimori, A.; Esmaeili, H.R.; Erpenbeck, D. & Reichenbacher, B. 2014. A new and unique species of the genus Aphanius Nardo, 1827 (Teleostei: Cyprinodontidae) from Southern Iran: A case of regressive evolution. Zoologischer Anzeiger-A Journal of Comparative Zoology 253: 327-337.
Teimori, A.; Esmaeili, H.R.; Hamidan, N. & Reichenbacher, B. 2018. Systematics and historical biogeography of the Aphanius dispar species group (Teleostei: Aphaniidae) and description of a new species from Southern Iran. Journal of Zoological Systematics and Evolutionary Research 56: 579-598.
Teimori, A.; Ebrahimi, M.; Mostafavi, H.; Esmaeili, H.R. & Masoudi, M. 2020. Distribution and abundance of the endemic scaleless tooth-carp, Aphanius furcatus Teimori, Esmaeili, Erpenbeck & Reichenbacher, 2014. Turkish Journal of Fisheries and Aquatic Science 20: 61-67.
Tigano, C.; Canapa, A.; Ferrito, V., Barucca, M.; Arcidiacono, I.; Deidun, A.; Schembri, P.J. & Olmo, E. 2006. A study of osteological and molecular differences in populations of Aphanius fasciatus Nardo 1827, from the central Mediterranean (Teleostei, Cyprinodontidae). Marine Biology 149: 1539-1550.
Villwock, V.W. 1964. Genetische Untersuchungen an altweltlichen Zahnkarpfen der Tribus Aphaniini (Pisces, Cyprinodontidae) nach Gesichtspunkten der Neuen Systematik. Journal of Zoological Systematics and Evolutionary Research 2: 267-382.
Wildekamp, R.H. 1993. A world of killies: atlas of the oviparous cyprinodontiform fishes of the world. American Killifish Association.
Wildekamp, R.H.; Küçük, F.; Ünlüsayin, M. & Van Neer, W. 1999. Species and subspecies of the genus Aphanius Nardo 1897 (Pisces: Cyprinodontidae) in Turkey. Turkish Journal of Zoology 23: 23-44.
Zeinali, F. & Motamedi, M. 2017. The regeneration capacity of caudal fin in the common tooth-carp, Aphanius dispar (Rüppell, 1829) (Teleostei: Cyprinodontidae). International Journal of Aquatic Biology 5: 321-327.
Barton, N.H. & Charlesworth, B. 1984. Genetic revolutions, founder effects, and speciation. Annual Review of Ecology and Systematics 15: 133-164.
Buj, I.; Miočić-Stošić, J.; Marčić, Z.; Mustafić, P.; Zanella, D.; Mrakovčić, M. &. Ćaleta, M. 2015. Population genetic structure and demographic history of Aphanius fasciatus (Cyprinodontidae: Cyprinodontiformes) from hypersaline habitats in the eastern Adriatic. Scientia Marina 79: 399-408.
Cavraro, F.; Malavasi, S.; Torricelli, P.; Gkenas, C.; Liousia, V.; Leonardos, I. &…Triantafyllidis, A. 2017 Genetic structure of the South European toothcarp Aphanius fasciatus (Actinopterygii: Cyprinodontidae) populations in the Mediterranean basin with a focus on the Venice lagoon. Search Results Web results. European Zoological Journal 84: 153-166.
Coad, B.W. 1980. A re-description of Aphanius ginaonis (Holly, 1929) from southern Iran (Osteichthyes; Cyprinodontiformes). Journal of Natural Histology 14: 33-40.
Coad, B.W. 1996. Fishes from the qanats of Iran. Publicaciones Especiales Instituto Español de Oceanografía 21: 63-79.
Coad, B.W. 1998. Threatened fishes of the world: Lebias ginaonis (Holly, 1929) (Cyprinodontidae). Environmental Biology of Fishes 51(3): 284.
Coad, B.W. 2000. Distribution of Aphanius species in Iran. Journal of the American Killifish Association 33: 183-191.
Coad, B. 2009. A new species of tooth-carp, Aphanius mesopotamicus, from Iran and Iraq (Actinopterygii, Cyprinodontidae). Zookeys 31:149.
Day, F. 1872. Notes on fish, collected by Dr. Stoliczka in Kachh. Journal of the Asiatic Society of Bengal 41: 1-4.
Echelle, A.A. & Echelle, A.F. 1992. Mode and pattern of speciation in the evolution of Inland pupfishes in the Cyprinodon variegatus complex (Teleostei: Cyprinodontidae): An ancestor-descendent hypothesis, in Systematics, Historical Ecology, and North American Freshwater Fishes, Mayden, R.L., Eds., Stanford, Stanford University Press California, pp. 691-709.
Esmaeili, H.R. & Gholami, Z. 2007. Investigations on the surface ultrastructure of scale of Geno tooth-carp, Aphanius ginaonis (Holly, 1929) (Actinopterygii: cyprinodontidae) using scanning electron microscope. Iranian Journal of Biology 20: 307-314.
Esmaeili, H.R.; Coad, B.W.; Gholamifard, A.; Nazari, N. & Teimori, A. 2010. Annotated сhecklist of the freshwater fishes of Iran. Zoosystematica Rossica 19: 361-386.
Esmaeili, H.R.; Teimori, A.; Sayyadzadeh, G.; Masoudi, M. & Reichenbacher, B. 2014a. Phylogenetic relationships of the tooth-carp Aphanius (Teleostei: Cyprinodontidae) in the river systems of southern and south-western Iran based on mtDNA sequences. Zoology in the Middle East 60: 29-38.
Esmaeili, H.R.; Teimori, A.; Gholami, Z. & Reichenbacher, B. 2014b. Two new species of the tooth-carp Aphanius (Teleostei: Cyprinodontidae) and the evolutionary history of the Iranian inland and inland-related Aphanius species. Zootaxa 3786: 246-268.
Esmaeili, H.R.; Sayyadzadeh, G. & Seehausen, O. 2016. Iranocichla persa, a new cichlid species from southern Iran (Teleostei, Cichlidae). ZooKeys 636: 141.
Esmaeili, H.R.; Sayyadzadeh, G.; Eagderi, S. & Abbasi, K. 2018. Checklist of freshwater fishes of Iran. FishTaxa 3: 1-95.
Esmaeili, H.R.; Teimori, A.; Zarei, F. & Sayyadzadeh, G. 2020. DNA barcoding and species delimitation of the Old-World tooth-carps, family Aphaniidae Hoedeman, 1949 (Teleostei: Cyprinodontiformes). PLoS ONE 15(4): e0231717.
Ferrito, V.; Mannino, M.C.; Pappalardo, A.M. & Tigano, C. 2007. Morphological variation between Italian populations of Aphanius fasciatus Nardo, 1827 (Teleostei, Cyprinodontidae). Journal of Fish Biology 70: 1-20.
Freyhof, J.; Weissenbacher, A. & Geiger, M. 2017. Aphanius kruppi, a new killifish from Oman with comments on the A. dispar species group (Cyprinodontiformes: Aphaniidae). Zootaxa 4338: 557-573.
Gaudant, J. 1993. Un exemple de “régression évolutive” chez des poissons Cyprinodontidae du Miocène supérieur d'Espagne: Aphanius illunensis nov. sp. Geobios 26: 449-454.
Grimm, H. 1980. Investigations on the problem of scale reduction and sulphate tolerance of West Anatolian cyprinodonts (Pisces). Internationale Revue der Gesamten Hydrobiologie und Hydrographie 65: 517-533.
Hatzfeld, D.; Authemayou, C.; Van Der Beek, P.; Bellier, O.; Lavé, J.; Oveisi, B.; Tatar, M.; Tavakoli, F.; Walpersdorf, A. & Yamini-Fard, F. 2010. The kinematics of the Zagros mountains (Iran). Geological Society, London, Special Publications 330: 19-42.
Heckel, J.J. 1849. Über eine neue fossile Fishgattung, Chirocentrites, und die ersten Ueberreste eines Siluroiden aus der Vorwelt. Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften, Mathematisch-Naturwissenschaftlichen Classe Wien, 2: 16-19.
Holly, L.F. Holly Pneumatic Systems Inc. 1929. Apparatus and method for loading loose-bulk cargoes. U.S. Patent 1,708,176.
Hrbek, T. & Meyer, A. 2003. Closing of the Tethys Sea and the phylogeny of Eurasian killifishes (Cyprinodontiformes: Cyprinodontidae). Journal of Evolutionary Biology 16: 17-36.
Hrbek, T.; Küçük, F.; Frickey, T.; Stölting, K.N.; Wildekamp, R.H. & Meyer, A. 2002. Molecular phylogeny and historical biogeography of the Aphanius (Pisces, Cyprinodontiformes) species complex of central Anatolia, Turkey. Molecular Phylogenetics and Evolution 25: 125-137.
Hrbek, T.; Keivany, Y. & Coad, B.W. 2006. New species of Aphanius (Teleostei, Cyprinodontidae) from Isfahan Province of Iran and a reanalysis of other Iranian species. Copeia 2006: 244-255.
Iranmanesh, A. & Motamedi, M. 2018. Histopathological changes of hepatorenal toxicity induced by gentamicin in killifish, Aphanius hormuzensis (Aphaniidae) and its kidney regeneration through nephron neogenesis. Journal of Ichthyology 58: 932-938.
Jenkins, J.T. 1910. XII. Notes on fish from India and Persia, with descriptions of new species. Records of the Indian Museum 10: 123-140.
Motamedi, M.; Shamsaldini, F.; Teimori, A. & Hesni, M.A. 2019a. Histomicroscopy and normal anatomy of the adult killifish Aphanius hormuzensis (Teleostei; Aphaniidae) from the Persian Gulf coastal environment. Microscopy Research and Technique 82: 466-480.
Motamedi, M.; Teimori, A. & Iranmanesh, A. 2019b. Ontogenetic pattern, morphological sexual and side dimorphism in the saccular otolith of a scaleless killifish Aphanius furcatus (Teleostei: Aphaniidae). Acta Zoologica. (In Press)
Nasrolah Pourmoghadam, M.; Poorbagher, H. & Eagderi, S. 2014. Histopathological effects of cypermethrin on liver of Aphanius sophiae (Heckel, 1849) using rank-based estimation for linear models. International Journal of Aquatic Biology 2(4): 184-187.
Parenti, L.R. 1981. A phylogenetic and biogeographic analysis of cyprinodontiform fishes (Teleostei, Atherinomorpha). Bulletin of the American Museum of Natural History. 168: 335-557.
Porkhial, S.; Nouraliee, J.; Rahmani, M. & Ebrahimi, D. 2013. Resource assessment of Vartun geothermal region, central Iran. Journal of Tethys 1: 29-40.
Regard, V.; Bellier, O.; Thomas, J.C.; Abbassi, M.R.; Mercier, J.; Shabanian, E.; Feghhi, K.H. & Soleymani, S. 2004. Accommodation of Arabia-Eurasia convergence in the Zagros-Makran transfer zone, SE Iran: A transition between collision and subduction through a young deforming system. Tectonics 23: TC4007
Reichenbacher, B.; Kamrani, E.; Esmaeili, H.R. & Teimori, A. 2009. The endangered cyprinodont Aphanius ginaonis (Holly, 1929) from southern Iran is a valid species: evidence from otolith morphology. Environmental Biology of Fishes 86(4): 507.
Rocco, L.; Ferrito, V.; Costagliola, D.; Marsilio, A.; Pappalardo, A.M.; Stingo, V. & Tigano, C. 2007. Genetic divergence among and within four Italian populations of Aphanius fasciatus (Teleostei, Cyprinodontiformes). Italian Journal of Zoology 74: 371-379.
Schwarzer, J.; Shabani, N.; Esmaeili, H.R.; Mwaiko, S. & Seehausen, O. 2017. Allopatric speciation in the desert: diversification of cichlids at their geographical and ecological range limit in Iran. Hydrobiologia 791: 193-207.
Sienknecht, U.1999. Kreuzungsgenetische Analyse des Uberganges von Paarflossen bis zu fehlendem Extremitatengurtel: entwickeltamBeispiel der normogenetischen Ventralflossenbildung von Aphanius iberus Cuv and Val 1846 im Vergleich zur Anormogenese seiner Hybriden mit A. apodus Gervais, 1853. Cyprinodontidae Der Andere Verlag, Bad Iburg, Teleostei.
Teimori, A.; Schulz-Mirbach, T.; Esmaeili, H.R. & Reichenbacher, B. 2012a. Geographical differentiation of Aphanius dispar (Teleostei: Cyprinodontidae) from southern Iran. Journal of Zoological Systematics and Evolutionary Research 50: 289-304.
Teimori, A.; Esmaeili, H.R.; Gholami, Z.; Zarei, N. & Reichenbacher, B. 2012b. Aphanius arakensis, a new species of tooth-carp (Actinopterygii, Cyprinodontidae) from the endorheic Namak Lake basin in Iran. ZooKeys 215: 55–76.
Teimori, A.; Jawad, L.A.J.; Al-Kharusi, L.H.; Al-Mamry, J.M. & Reichenbacher, B. 2012c. Late Pleistocene to Holocene diversification and historical zoogeography of the Arabian killifish (Aphanius dispar) inferred from otolith morphology. Scientia Marina 76: 637-645.
Teimori, A.; Esmaeili, H.R.; Erpenbeck, D. & Reichenbacher, B. 2014. A new and unique species of the genus Aphanius Nardo, 1827 (Teleostei: Cyprinodontidae) from Southern Iran: A case of regressive evolution. Zoologischer Anzeiger-A Journal of Comparative Zoology 253: 327-337.
Teimori, A.; Esmaeili, H.R.; Hamidan, N. & Reichenbacher, B. 2018. Systematics and historical biogeography of the Aphanius dispar species group (Teleostei: Aphaniidae) and description of a new species from Southern Iran. Journal of Zoological Systematics and Evolutionary Research 56: 579-598.
Teimori, A.; Ebrahimi, M.; Mostafavi, H.; Esmaeili, H.R. & Masoudi, M. 2020. Distribution and abundance of the endemic scaleless tooth-carp, Aphanius furcatus Teimori, Esmaeili, Erpenbeck & Reichenbacher, 2014. Turkish Journal of Fisheries and Aquatic Science 20: 61-67.
Tigano, C.; Canapa, A.; Ferrito, V., Barucca, M.; Arcidiacono, I.; Deidun, A.; Schembri, P.J. & Olmo, E. 2006. A study of osteological and molecular differences in populations of Aphanius fasciatus Nardo 1827, from the central Mediterranean (Teleostei, Cyprinodontidae). Marine Biology 149: 1539-1550.
Villwock, V.W. 1964. Genetische Untersuchungen an altweltlichen Zahnkarpfen der Tribus Aphaniini (Pisces, Cyprinodontidae) nach Gesichtspunkten der Neuen Systematik. Journal of Zoological Systematics and Evolutionary Research 2: 267-382.
Wildekamp, R.H. 1993. A world of killies: atlas of the oviparous cyprinodontiform fishes of the world. American Killifish Association.
Wildekamp, R.H.; Küçük, F.; Ünlüsayin, M. & Van Neer, W. 1999. Species and subspecies of the genus Aphanius Nardo 1897 (Pisces: Cyprinodontidae) in Turkey. Turkish Journal of Zoology 23: 23-44.
Zeinali, F. & Motamedi, M. 2017. The regeneration capacity of caudal fin in the common tooth-carp, Aphanius dispar (Rüppell, 1829) (Teleostei: Cyprinodontidae). International Journal of Aquatic Biology 5: 321-327.