CORMOHIPPARION SOFULARENSIS N. SP., A NEW HIPPARION SPECIES OF FROM THE LATE MIOCENE OF SOFULAR (TÜRKİYE, EARLY TUROLIAN)
DOI:
https://doi.org/10.54103/2039-4942/27705Keywords:
Cormohipparion; Sofular; Late Miocene; Turolian; TürkiyeAbstract
In this contribution, we study the hipparion remains from the Early Turolian locality of Sofular, Türkiye. We identify the presence of a new species, Cormohipparion sofularensis, with three other taxa which we refer to Hippotherium brachypus, Hipparion dietrichi and Cremohipparion moldavicum. We compare these taxa with other Late Miocene hipparions from Iran, Türkiye, Greece, Moldova, Austria, Germany, France, Spain, Algeria and Libya, by morphologic and morphometric comparisons of crania, mandibles, maxillary and mandibular dentitions, third metacarpals and third metatarsals to further evaluate the morphological and systematic comparisons. Recently, the Sofular locality is dated radioisotopically to 8.422±0.34 Ma, correlative with early MN11 (Early Turolian) faunas in our comparisons. The Sofular hipparions record the early evolutionary radiation of hipparion lineages in the Subparatethyan Province. Our analyses further reveal that Sofular Cormohipparion sofularensis is the youngest member of the genus Cormohipparion reported from Eurasia and Africa and is found to be derived in skull morphology compared to the Late Vallesian Cormohipparion cappadocium from Yeniyaylacık, Türkiye.
Downloads
References
Armour-Chelu M. & Bernor R.L. (2011) Equidae. In: Harrison T. (Ed.) - Geology and paleontology of Laetoli: 295-326. Springer-Verlag, New York.
Aydar E., Schmitt A.K., Cubukcu H E., Akin L., Erzoy O., Sen E., Duncan R.A. & Atici G. (2012) - Correlation of ignimbrites in the central Anatolian volcanic province using zircon and plagioclase ages and zircon compositions. Journal of Volcanology and Geothermal Research 213-214: 83-97.
Başoğlu O. (2017) - Vertebrate Fossil Beds in the Cappadocia Region and a New Hominid Site: Sofular. (Şükrü Devrez. Bilgin Kültür Sanat Şti. Ltd.).
Bernor R.L. (1983) - Geochronology and Zoogeographic Relationships of Miocene Hominoidea. Ciochon R.L. & Corruccini R. (Eds.) - New Interpretations of Ape and Human Ancestry: 21-64. Plenum Press, New York.
Bernor R.L.(1984) - A Zoogeographic Theater and Biochronologic Play: The Time/Biofacies Phenomena of Eurasian and African Miocene Mammal Provinces. Paleobiology Continentale. 14: 121-142.
Bernor R.L. (1985) - Systematics and Evolutionary Relationships of the Hipparionine horses from Maragheh. Iran. Paleovertebrata 15: 173-269.
Bernor R.L. & Hussain S.T. (1985) - An Assessment of the Systematic, Phylogenetic and Biogeographic Relationships of Siwalik Hipparionine Horses. Journal of Vertebrate Paleontology 5: 32-87.
Bernor R.L. & Armour-Chelu M. (1999) - Toward an Evolutionary History of African Hipparionine HorsesIn: Brommage T. & Schrenk F. (Eds.) - African Biogeography, Climate Change and Early Hominid Evolution: 189-215. Oxford Press, Oxford.
Bernor R.L. & Harris J. (2003) - Systematics and Evolutionary Biology of the late Miocene and early Pliocene Hipparionine horses from Lothagam, Kenya. — In: Leakey M. & Harris J. (Eds.) - Lothagam: The Dawn of Humanity in Eastern Africa: 387-438. Columbia University Press, New York.
Bernor R.L. & Scott R.S. (2003) - New Interpretations of the Systematics, Biogeography and Paleoecology of the Sahabi Hipparions (latest Miocene), Libya. Geodiversitas, 25: 297-319.
Bernor R.L. & Haile Selassie Y. (2009) - Equidae. In: Haile-Selassie Y. & Woldegabriel G. (Eds.) - Ardipithecus kadabba: Late Miocene Evidence from the Middle Awash, Ethiopia: 397-428. University of California Press, Berkeley.
Bernor R.L. & White T.D. (2009) - Systematics and Biogeography of “Cormohipparion” africanum, Early Vallesian (MN 9, ca. 10.5 Ma) of Bou Hanifia, Algeria. In: Albright III, B. (Ed.) - Papers on Geology, Vertebrate Paleontology, and Biostratigraphy in Honor of Michael O. Woodburne. Bulletin of the Museum of Northern Arizona, 65: 635-658.
Bernor R.L. & Sun B. (2015) - Morphology through Ontogeny of Chinese Proboscidipparion and Plesiohipparion and Observations on their Eurasian and African Relatives. Vertebrata Pal Asiatica, 53:73-92.
Bernor R.L., Qiu Z. & Tobien H. (1987) - Phylogenetic and Biogeographic bases for an Old World Hipparionine Horse Geochronology. Proceedings of the VIIIth International Congress of the Regional Committee on Mediterranean Neogene Stratigraphy. Annals of the Hungarian Geological Institute, 70: 43-53.
Bernor R.L., Qiu Z. & Hayek L.A.C. (1990a) - Systematic revision of Chinese Hipparion species described by Sefve, 1927. American Museum Novitates, 2984: 1-60.
Bernor R.L., Tobien H. & Woodburne M.O. (1990b) - Patterns of Old World Hipparionine Evolutionary Diversification. In: Lindsay E., Fahlbusch V. & Mein P. (Eds.) - European Neogene Mammal Chronology: 263-319. Plenum Press, New York.
Bernor R.L., Fahlbusch V., Andrews P., De Bruijn H., Fortelius M., Rögl F., Steininger F.F. & Werdelin L. (1996) - The evolution of Western Eurasian Neogene mammal faunas: a chronologic, systematic, biogeographic, and paleoenvironmental synthesis. In Bernor R.L., Fahlbusch V. & Mittman H.W. (Eds) - The Evolution of Western Eurasian Neogene Mammal Faunas, Columbia University Press, New York: 449-469.
Bernor R.L., Tobien H., Hayek L.-A., & Mittmann H.-W. (1997) - The Höwenegg hipparionine horses: systematics, stratigraphy, taphonomy and paleoenvironmental context. Andrias, 10: 1-230.
Bernor R.L., Scott R.S., Fortelius M., Kappelman J. & Sen S. (2003) - Systematics and Evolution of the Late Miocene Hipparions from Sinap, Türkiye. In: Fortelius M. Kappelman J., Sen S. & Bernor R.L. (Eds.) - The Geology and Paleontology of the Miocene Sinap Formation, Türkiye: 220-281. Columbia University Press, New York.
Bernor R.L., Scott R.S. & Haile-Selassie Y. (2005) - A contribution to the evolutionary history of Ethiopian hipparionine horses: morphometric evidence from the postcranial skeleton. Geodiversitas, 27: 133-158.
Bernor R.L., Armour-Chelu M., Gilbert H., Kaiser T. & Schulz E. (2010) - Equidae. In: Werdelin L. & Sanders B. (Eds.) - Cenozoic Mammals of Africa: 685-721. University of California Press, Berkeley.
Bernor R.L., Kaiser T.M., Nelson S.V. & Rook L. (2011) - Systematics and Paleobiology of Hippotherium malpassi n. sp. from the latest Miocene of Baccinello V3 (Tuscany, Italy). Bolletino della Società Paleontologica Italiana, 50: 175-208.
Bernor R.L., Gilbert H., Semprebon G., Simpson S. & Semaw S. (2013) - Eurygnathohippus woldegabrieli sp. nov. (Perissodactyla: Mammalia) from the Middle Pliocene of Aramis, Ethiopia (4.4 Ma.). Journal of Vertebrate Paleontology, 33: 1472-1485.
Bernor R.L., Meshida K. & Sun B. (2015) - Phylogenetic Signatures in the Juvenile Craniums and Cheek Teeth of Pleistocene Proboscidipparion sinense, China. Rivista Italiana di Paleontologia e Stratigrafia, 121: 255-264.
Bernor R.L., Mirzaie Ataabadi M., Meshida K. & Wolf D. (2016) - The Maragheh Hipparions; late Miocene of Azarbaijan, Iran. Palaeobiodiversity and Palaeoenvironments, 96: 453-488.
Bernor R.L., Goehlic U., Harzhauser M. & Semprebon G. (2017) - Stratigraphy, Chronology, Systematics and Paleoecology of the Vienna Basin Pannonian C Hipparions (basal MN9, late Miocene). Palaeogeography, Palaeoclimatology, Palaeoecology, 476: 28-41.
Bernor R.L., Boaz N.T., Cirilli O., El-Shawaihdi M. & Rook L. (2020) - Sahabi Eurygnathohippus feibeli: its systematic, stratigraphic, chronologic and biogeographic contexts. Rivista Italiana di Paleontologia e Stratigrafia, 126: 561-581.
Bernor R.L, Kaya F., Kakkinen A., Sarinnen J. & Fortelius M. (2021) - Old World Hipparionine Evolution, Biogeography, Climatology and Ecology. Earth-Science Reviews, 221: 103784.
Bernor R.L., Cirilli O. & Mittmann H.W. (2022) - Höwenegg Hippotherium primigenium: geological context, cranial and postcranial morphology, paleoecological and biogeographic importance. Historical Biology, 34: 1376-1390.
Bernor R.L., Ataabadi M., Başoglu O., Cirilli O., Kaya F., Pehlevan C., Niknahad M., Vaziri M.R. & Arab A.L. (2024) - Cormohipparion cappadocium, a new species from the late Miocene of Yeniyaylacık, Turkiye, and the emergence of western Eurasian hipparion bioprovinciality. Annales Zoologici Fennici, 61: 303-333. https://doi.org/10.1007/978-3-031-17491-9_3.
Bernor R.L., Morgan M., Nelson S. & Semprebon G. (2025) - 15. Siwalik Equidae. In: Badgley C. , Pilbeam D. & Morgan M. (Eds.) - At the Foot of the Himalayas: Paleontology and Ecosystem Dynamics of the Siwalik Record of Pakistan: 260-271. Johns Hopkins University Press, Baltimore.
Çiner A., Doğan U., Yıldırım C., Akçar N., Ivy-Ochs S., Alfimov V., Kubik P.W. & Schlüchter C. (2015) - Quaternary uplift rates of the Central Anatolian Plateau, Türkiye: insights from cosmogenic isochron-burial nuclide dating of the Kızılırmak River terraces. Quaternary Science Reviews 107: 81-97.
Cirilli O., Zouhri S., Boughabi S., Benvenuti M., Bernor R.L., Papini M. & Rook L. (2020) - The hipparionine horses (Perissodactyla: Mammalia) from the Late Miocene of Tizi N’Tadderht (southern Ouarzazate basin; Central High Atlas; Morocco). Rivista Italiana di Paleontologia e Stratigrafia 126:1-12.
Cirilli O., Bernor R.L. & Rook L. (2021) - The Early Pleistocene Equids from Roca-Neyra (France, Central Europe). Implications for the Hipparion Last Occurrence and the Equus First Appearance Datum in Europe. Journal of Paleontology 95: 406-425.
Cirilli O., Pandolfi L., Alba D.M., Madurell-Malapeira J., Bukhsianidze M., Kordos L., Lordkipanidze D., Rook L. & Bernor R.L. (2023) - The last Plio-Pleistocene hipparions of Western Eurasia. A review with remarks on their taxonomy, paleobiogepgraphy and evolution. Quaternary Science Reviews, 306: 107976.
Dogan U. & Şenkul Ç. (2020) - When did the drainage system of the Kızılırmak River form in Cappadocia (Anatolia, Türkiye)? A revised geological and geomorphological stratigraphy. Turkish Journal of Earth Science 29: 1100-1113.
Eisenmann V. (1995) - What metapodial morphometry has to say about some Miocene hipparions. In Vrba E.S., Denton G.H., Partridge T.C. & Burckle L.H. (Eds.) - Paleoclimate and evolution, with special emphasis on Human origins: 48-162. Yale University Press, New Haven.
Eisenmann V., Alberdi M.T., De Giuli C. & Staesche U. (1988) - Methodology. In: Woodburne M. & Sondaar P.Y. (Eds.) - Studying Fossil Horses: 1-71. E.J. Brill Press, Leiden.
Eronen J.T., Mirzaie Ataabadi M., Karme A. Micheels A., Bernor R.L. & Fortelius M. (2009) - Distribution History and Climatic Controls of the Late Miocene Pikermian Chronofauna. Proceedings of the National Academy of Sciences, 106: 11867-11871.
Fortelius M., Eronen J., Jernvall J., Liu L., Pushkina D., Rinne J., Tesakov A., Vislobokova I., Zhang Z. & Zhou L. (2002) - Fossil Mammals Resolve Regional Patterns of Eurasian Climate Change Over 20 Million Years. Evolutionary Ecology Research, 4(7): 1005-1016.
Gabunja L.K. (1959) - Histoire du genre Hipparion. USSR academy of Science, Moscou. [traduction en francais B.R.G.M.], 69 Fig., 151 Tab., 23 PI.; Paris.
Gilbert H. & Bernor R.L. (2008) - Equidae. In: Gilbert H. & Asfaw B. (Eds.) - Homo erectus - Pleistocene Evidence from the Middle Awash, Ethiopia: 133-166. University of California Press, Berkeley.
Gromova V.I. (1952) - Le genre Hipparion. Institut de Paléontologie, Académie des Sciences d’URSS (en Russe), traduit en français par Saint Aubin. Bureau de Recherches Géologiques et Minières, ACEDP, 12, 1-473. .
Innocenti F., Mazzuoli R., Pasquarè G., Di Brozolo F.R. & Villari L. (1975) - The Neogene calcalkaline volcanism of Central Anatolia: geochronological data on Kayseri-Nigde area. Geological Magazine, 112(4), 349-360.
Kaya F., Bibi F., Zliobaite I., Eronen J.T., Hui T. & Fortelius M. (2018) - The rise and fall of the Old World savanna fauna and the origins of the African savannah biome. Nature: Ecology and Evolution 2: 241-246, https://doi. org/10.1038/s41559-017-0414-1.
Koufos G.D. (2016) - Hipparion macedonicum revisited: New data on evolution of hipparionine horses from the late Miocene of Greece. Acta Palaeontologica Polonica, 61: 519-536.
Koufos G.D. & Vlachou T. (2005) - Equidae (Mammalia, Perissodactyla) from the late Miocene of Akkaşdağı, Türkiye. In: Sen S. (Ed.) - Geology, Mammals and Environments at Akkaşdağı, Late Miocene of Central Anatolia. Geodiversitas, 27: 633-705.
Koufos G.D., Kostopoulos D.S. & Vlachou T.D. (2016) - Synthesis. In: Koufos G.D. & Kostopoulos D.S. (Eds.) - Palaeontology of the Upper Miocene Vertebrate Localities of Nikiti (Chalkidiki Peninsula, Macedonia, Greece). Geobios, 49: 147-154.
Le Pennec J.L., Bourdier J.L., Froger J.L., Temel A.B.İ.D.İ.N., Camus G. & Gourgaud A. (1994) - Neogene ignimbrites of the Nevsehir plateau (Central Türkiye): stratigraphy, distribution and source constraints. Journal of Volcanology and Geothermal Research, 63: 59-87.
Lepetit P., Viereck L., Piper J.D., Sudo M., Gürel A., Çopuroğlu I. & Gürsoy H. (2014) - 40Ar/39Ar dating of ignimbrites and plinian air-fall layers from Cappadocia, Central Türkiye: Implications to chronostratigraphic and Eastern Mediterranean palaeoenvironmental record. Geochemistry, 74: 471-488.
Liu L., Puolamäki K., Eronen J.T., Ataabadi M.M., Hernesniemi E. & Fortelius M. (2012) - Dental functional traits of mammals resolve productivity in terrestrial ecosystems past and present. Proceedings of the Royal Society of London B: Biological Sciences, 279: 2793-2799. https://doi.org/10.1098/rspb.2012.0211.
MacFadden B.J. (1980) - The Miocene horse Hipparion from North America and from the type locality in southern France. Palaeontology, 23: 617-635.
MacFadden B.J. (1984) - Systematics and phylogeny of Hipparion, Neohipparion, Nannippus, and Cormohipparion (Mammalia, Equidae), from the Miocene and Pliocene of the New World. Bulletin of the American Museum of Natural History, 179: 1-196.
MacFadden B.J. & Skinner M.F. (1981) - Earliest Holarctic hipparion, Cormohipparion goorisi n. sp. (Mammalia, Equidae) from the Barstovian (medial Miocene) Texas Gulf Coastal Plain. Journal of Paleontology, 55: 619-627.
MacFadden B.J. & Woodburne M.O. (1982) - Systematics of the Neogene Siwalik Hipparions (Mammalia, Equidae) based on cranial and dental morphology. Journal of Vertebrate Paleontology, 2: 185-218.
Mouralis D. & Aydar E. (2019) - Quaternary Volcanic Landscapes and Prehistoric Sites in Southern Cappadocia: Göllüdağ, Acıgöl and Hasandağ. Landscapes and landforms of Turkey: 551-563.
Oksanen O., Žliobaitė I., Saarinen J., Lawing A.M. & Fortelius M. (2019) - A Humboldtian approach to life and climate of the geological past: Estimating palaeotemperature from dental traits of mammalian communities. Journal of Biogeography, 46: 1760-1776, https://doi.org/10.1111/jbi.13586.
Pasquarè G. (1968) - Geology of the Cenozoic Volcanic Area of Central Anatolia. Atti Accademia Nazionale dei Lincei, 9: 55-204.
Piper J. D.A., Koçbulut F., Gürsoy H., Tatar O., Viereck L., Lepetit P., Roberts A.P. & Akpınar Z. (2013) - Palaeomagnetism of the Cappadocian Volcanic Succession, Central Türkiye: Major ignimbrite emplacement during two short (Miocene) episodes and Neogene tectonics of the Anatolian collage. Journal of Volcanology and Geothermal Research, 262: 47-67.
Qiu Z., Huang W. & Guo Z. (1987) - The Chinese hipparionine fossils. Palaeontologia Sinica, N.S.C. 17: 1-250.
R Core Team (2023) - R: a Language and Environment for Statistical Computing. R Foundation for Statistical Computing, Vienna, Austria.
Wolf D., Bernor R.L. & Hussain S.T. (2013) - A systematic, biostratigraphic, and paleobiogeographic reevaluation of the Siwalik hipparionine horse assemblage from the Potwar Plateau, Northern Pakistan. Palaeontographica, 300: 1-115.
Tholt A., Başoğlu O., Bektaş Y., Bernor R., Carlson J.P., Dağ Ö., Doğan U., Erkman A.C., Kaya F., Kaymakçı N., Gözlük Kırmızıoğlu P., Meijers M.J.M., Kahya Parıldar Ö., Pehlevan C., Şimşek E., White, T. & Renne P., (2025) - Building better biochronology: New fossils and 40Ar/39Ar radioisotopic dates from Central Anatolia, Proceedings of the National Academy of Sciences USA, 122 (12) e2424428122, https://doi.org/10.1073/pnas.2424428122.
Woodburne M.O. (1989) - Hipparion horses: A pattern of endemic evolution and intercontinental dispersal. In: Prothero D.R. & Schoch R.M. (Eds.) - The Evolution of Perissodactyls: 197-23. Oxford University Press, New York.
Woodburne M.O. (2007) - Phyletic diversification of the Cormohipparion occidentale Complex (Mammalia; Perissodactyla, Equidae), late Miocene, North America, and the origin of the Old World Hippotherium Datum. Bulletin of the American Museum of Natural History, 306: 1-138.
Woodburne M.O. (2009) - The early Vallesian vertebrates of Atzelsdorf (late Miocene, Austria) 9. Hippotherium (Mammalia, Equidae). Annalen Naturhistorischen Museums Wien, 111A: 585-604.
Woodburne M.O. & Bernor R.L. (1980) - On Superspecific Groups of Some Old World Hipparionine Horses. Journal of Paleontology, 54: 1319-1348.
Woodburne M.O., MacFadden B.J. & Skinner M. (1981) - The North American “Hipparion Datum”, and implications for the Neogene of the Old World. Geobios, 14: 493-524.
Downloads
Additional Files
Published
How to Cite
Issue
Section
License
Copyright (c) 2025 ÖZGE KAHYA PARILDAR, OKŞAN BAŞOĞLU, OMAR CIRILLI, ÖMER DAĞ, FERHAT KAYA, PINAR GÖZLÜK KIRMIZIOĞLU, CESUR PEHLEVAN, EMRAH ŞİMŞEK, RAYMOND L. BERNOR
This work is licensed under a Creative Commons Attribution-ShareAlike 4.0 International License.
The journal allow the author(s) to hold the copyright without restrictions.
Accepted 2025-03-28
Published 2025-06-03
