THE KAYSERI HIPPARION HORSES: NEW INSIGHTS INTO EQUID EVOLUTION IN TÜRKIYE DURING THE EARLY AND MIDDLE TUROLIAN (MN11-12, LATE MIOCENE)
DOI:
https://doi.org/10.54103/2039-4942/30263Keywords:
Hipparion horses; Kayseri; Late Miocene; Paleobiogeography; Biochronology.Abstract
The Yamula Reservoir shores have yielded a series of vertebrate localities rich in diverse ungulates and carnivores which we refer to the Kayseri faunas, Türkiye. We report herein the geological context and hipparion horses excavated from fluvial and lake deposits Çevril, Taşhan, and Hırka sites. Published 40Ar/39Ar geochronologies place the deposits within the late Tortonian, respectively the late European Mammal Neogene (MN) Units late MN11– early MN12 (early to middle Turolian). These localities can therefore be correlated with others of the Greek–Türkish–Iranian region, as Pikermi and Samos (Greece) and Maragheh (Iran). We undertake both morphologic and morphometric analyses in identifying taxa and comparing them to a broad suite of taxa from Western Eurasian localities. Hipparion horses identified in this study include Hippotherium brachypus, Hipparion dietrichi, Cremohipparion moldavicum, Cremohipparion aff. proboscideum and Plesiohipparion longipes. Our paleoclimate reconstruction of Upper Miocene Western Eurasian faunas based on the mean ordinated hypsodonty analysis of fossil mammal assemblages from MN10–12 shows a substantial climate shift, expressed in the expansion and diversification of the classic Pikermian ungulate and carnivore “savanna-like” chronofauna.
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References
Abel O. (1926) - Die Geschichte der Equiden auf dem boden Nordamerikas. Verhandlungen der Zoologisch-Botanischen Gesellschaft in Wien, 74-75: 159-164
Abel O. (1927) - Lebensbilder aus der Tierwelt der Vorzeit. Verlag von Gustav Fischer, Jena, 676 pp.
Ameghino F. (1904) - Recherches de morphologic phylogenetique sur les molaires superiores des ongules: Anales del Museo Nacional de Historia Natural de Buenos Aires, 3rd. Ser., 3: 1-541.
Arambourg C. (1959) - Vertebres continentaux du Miocene superieur de l’Afrique du Nord. Service Carte Geologie Algerie Paleontologie Memoire, Nouveaux Serie 4: 1-159.
Armour-Chelu M. & Bernor R.L. (2011) - Equidae. In: Harrison T. (Eds) - Geology and paleontology of Laetoli: 295-326. Springer-Verlag, New York.
Atabaadi M.M, Bernor R.L., Kostopoulos D.S., Wolf D., Orak Z., Zare G., Nakaya H., Watabe M. & Fortelius M. (2013) - Recent Advances in Paleobiological Research of the Late Miocene Maragheh Fauna, Northwest Iran. In: Wang X., Flynn L.J. & Fortelius M. (Eds) - Fossil Mammal of Asia: Neogene Biostratigraphy and Chronology: 539-546. Columbia University Press, 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.
Bernor R.L. (1983) - Geochronology and Zoogeographic Relationships of Miocene Hominoidea. In: 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. (1986) - Mammalian biostratigraphy, geochronology, and zoogeographic relationships of the Late Miocene Maragheh fauna, Iran. Journal of Vertebrate Paleontology, 6: 76- 95.
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. & Tobien H. (1989) - Two small species of Cremohipparion (Equidae, Mammalia) from Samos, Greece. Mitteilungen der Bayerischen Staatssammlung für Paläontologie und Historische Geologie, 29: 207-226
Bernor R.L. & Armour-Chelu M. (1999) - Toward an Evolutionary History of African Hipparionine Horses. In: 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. Museum of Northern Arizona Bulletin, 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 PalAsiatica, 53: 73-92.
Bernor R.L., Woodburne M.O. & Van Couvering J.A. (1980) - A contribution to the chronology of some Old World faunas based on hipparionine horses. Geobios, 13: 25-59.
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. (1996a) - The CORRELATION OF THREE CLASSICAL “Pikennian” mammal faunas - Maragheh, Samos, and Pikermi - wilh the European MN Unit System. In: Bernor R.L., Fahlbusch V. & Mittman H.W. (Eds) - The Evolution of Western Eurasian Neogene Mammal Faunas: 137-156. Columbia University Press, New York.
Bernor R.L., Koufos G.D., Woodburne M.O. & Fortelius M. (1996b). The evolutionary history ancl biochronology of European and Southwest Asian Late Miocene and Pliocene hipparionine horses. In: Bernor R.L., Fahlbusch V. & Mittman H.W. (Eds) - The evolution of Western Eurasian Neogene mammal faunas: 307-338. Columbia University Press, New York.
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, Turkey. In: Fortelius M., Kappelman J., Sen S. & Bernor R.L. (Eds) - The Geology and paleontology of the Miocene Sinap Formation, Turkey: 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.M., 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. & Sen S. (2017) - The Early Pliocene Plesiohipparion and Proboscidipparion (Equidae, Hipparionini) from Çalta, Turkey (Ruscinian Age, c. 4.0 Ma). Geodiversitas, 39: 285-314.
Bernor R.L., Göhlic U., Harzhauser M. & Semprebon G. (2017) - The Pannonian C hipparions from the Vienna Basin. Palaegeography, Palaeoclimatology, Palaeoecology, 476: 28-41.
Bernor R.L., Wang S., Liu Y., Chen Y. & Sun B. (2018) - Shanxihippus dermatorhinus comb. nov. with comparisons to Old World hipparions with specialized nasal apparati. Rivista Italiana di Paleontologia e Stratigrafia, 124: 361-386.
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 hipparion evolution, biogeography, climatology and ecology. Earth-Science Reviews, 221: 103784.
Bernor R.L., Cirilli O. & Mittmann H.W. (2022a) - Höwenegg Hippotherium primigenium: geological context, cranial and postcranial morphology, paleoecological and biogeographic importance. Historical Biology, 34: 1376-1390.
Bernor R.L., Beech M.J. & Bibi F. (2022b) - Equidae from the Baynunah Formation. In: Bibi F., Kraatz B., Beech M.J. & Hill A. (Eds) - Sands of Time. Ancient life in the Late Miocene of Abu Dhabi, United Arab Emirates: 261-282. Springer, Vertebrate Paleobiology and Paleoanthropology, Cham, Switzerland.
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 Yeniyaylacik, Turkiye, and the emergence of western Eurasian hipparion bioprovinciality. Annales Zoologici Fennici, 61: 303-333.
Bernor R.L., Morgan M., Nelson S. & Semprebon G. (2025) - 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.
Böhme M., Spassov N., Ebner M., Geraads D., Hristova L., Kirscher U., Kötter S., Linnemann U., Prieto J., Roussiakis S., Theodorou G., Uhlig G. & Winklhofer M. (2017) - Messinian age and savannah environment of the possible hominin Graecopithecus from Europe. PLoS ONE 12(5): e0177347.
Campbell B.G., Amini M.H., Bernor R.L., Dickenson W., Drake W., Morris R., Van Couvering J.A. & Van Couvering J.A.H. (1980) - Maragheh: a classical late Miocene vertebrate locality in northwestern Iran. Nature, 287: 837-841.
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.
Christol de J. (1832). Description d’Hipparion. Annales des Sciences et de l’Industrie du Midi de France, 1: 180-181
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. & Sondaar P. (1998) - Hipparion. Geodiversitas, 20: 409-439.
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., Micheels A., Karme 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.
Ezquerro L., Luzón A., Simón J.L. & Liesa L.C. (2022) - A review of the European Neogene Mammal zones from integration of litho-, bio- and magnetostratigraphy in the Teruel Basin. Earth-Science Reviews, 234 (2022): 104223.
Falconer H. & Cautley P.T. (1846) - Fauna Antiqua Sivalensis, being the fossil zoology of the Sewalik Hills, in the north of India. Part 9. Equidae, Camelidae, and Siuatherium. Smith, Elder and Co., London, 136 pp.
Forsten A.M. (1980) - How many Hipparion species at Samos? Neues Jahrbuch für Geologie und Paleontology, 7: 391-396.
Forsten A.M. (1978) - A review of the Bulgarian Hipparion (Mammalia, Perissodactyla). Geobios, 11: 31-41.
Forsten A.M. & Garevski R. (1989) - Hipparions (Mammalia, Perissodactyla) from Macedonia/Yugoslavia. Geologia Macedonica, Series T, 3: 159-206.
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: 1005-1016.
Fortelius M., Werdelin L., Andrews P., Bernor R.L., Gentry A., Humphrey L., Mittmann H.W. & Viranta S. (1996) - Provinciality, diversity, turnover, and paleoecology in land mammal faunas of the later Miocene of western Eurasia. In: Bernor R.L., Fahlbusch V., Mittmann W. (Eds) - The evolution of Western Eurasian Neogene mammal faunas: 414-448. Columbia Univ. Press.
Friedrichs B., Schmitt A.K., Lovera O.M. & Atıcı G. (2021) - Zircon as a recorder of contrasting magma recharge and eruptive recurrence patterns. Earth and Planetary Science Letters, 571: 117104
Gabunja L.K. (1959) - Histoire du genre Hipparion. USSR academy of Science, Moscow.
Gervais P. (1959) - Zoologie et paleontologie francaises. Nouvelles recherches animaux vertebres. Arthur Bertrand, Paris. 544 pp.
Gidley J. (1903) - A new three-toed horse. Bulletin of the American Museum of Natural History, 19: 465-476.
Gray H. (1821) - On the natural arrangement of vertebrose animals. London Medical Repository, 15: 296-310.
Gromova V. (1952) - Le genre Hipparion. Bureau de Recherches géologiques et Minières, 12: 1-288.
Cano A.R.G., Fernández M.H. & Álvarez-Sierra M.Á. (2011) - Biogeographic provincialism in rodent faunas from the Iberoccitanian Region (southwestern Europe) generates severe diachrony within the Mammalian Neogene (MN) biochronologic scale during the Late Miocene. Palaeogeography, Palaeoclimatology, Palaeoecology, 307(1-4): 193-204.
Hensel R. (1892) - Uber die Reste einiger Saugethierarten von Pikermi in der Munchener Sammlung. Monatsberichte der Königlich Preußischen Akademie der Wissenschaften zu Berlin, 27 : 560-569
Higgins M., Schoenbohm L.M., Brocard G., Kaymakcı N., Gosse J.C. & Cosca M.A. (2015) - New kinematic and geochronologic evidence for the Quaternary evolution of the Central Anatolian fault zone (CAFZ). Tectonics, 34: 2118-2141.
Hilgen F.J., Lourens L.J., Van Dam J.A., Beu A.G., Boyes A.F., Cooper R.A., Krijgsman W., Ogg J.G., Piller W.E. & Wilson D.S. (2012) - The Neogene Period. In: Gradstein F.M., Ogg J.G., Schmitz M.D. & Ogg G.M. (Eds) - The Geologic Time Scale: 923-978. Elsevier, Boston.
Kahya-Parıldar Ö., Başoğlu O., Cirilli O., Dağ Ö., Kaya F., Gözlük Kırmızıoğlu P., Pehlevan C., Şimşek E. & Bernor R.L. (2025) - Cormohipparion sofularensis n. sp., a new hipparion species from the Late Miocene of Sofular (Türkiye, Early Turolian). Rivista Italiana di Paleontologia e Stratigrafia, 131: 305-329.
Kaiser T.M., Bernor R.L., Scott R.S., Franzen J.L., Solounias N. (2003) - New interpretations of the systematics and palaeoecology of the Dorn-Duerkheim 1 Hipparions Late Miocene, Turolian Age [MN11], Rheinhessen, Germany. Senckenbergiana Lethea, 83(1/2): 103-133.
Kaup J.J. (1835) - Die zvei Urweltlichen pferdeartige Thiere whelche in tertiären Sande bei Eppelsheim gefunden werden. Nova Acta Physico-Medica Academiae Caesareae Leopoldino-Carolinae, 17: 173-182.
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.
Kostopoulos D.S., Sen S. & Koufos G. (2003) - Magnetostratigraphy and revised chronology of the late Miocene mammal localities of Samos, Greece. International Journal of Earth Sciences, 92: 779-794.
Kostopoulos D.S. (2009) - The Pikermian Event: temporal and spatial resolution of the Turolian large mammal fauna in SE Europe. Palaeogeography, Palaeoclimatology, Palaeoecology, 274(1-2): 82-95.
Koufos G.D. (1984) - A new hipparion (Mammalia, Perissodactyla) from the Vallesian (late Miocene) of Greece. Paläontologische Zeitschrift, 58: 307-317
Koufos G.D. (1987a) - Study of the Pikermi hipparions. In: Part I: Generalities and taxonomy. Bulletin du Muséum national d’Histoire naturelle Paris 4e series 9, section C, 2: 197-252.
Koufos G.D. (1987b) - Study of the Pikermi hipparions. In: Part II: Comparisons and odontograms. Bulletin du Muséum national d’Histoire naturelle Paris 4e series 9, section C, 3: 327-363.
Koufos G.D. (1987c) - Study of the Turolian hipparions of the lower Axios Valley (Macedonia, Greece). 1. Locality “Ravin des Zouaves-5” (RZO). Geobios, 20: 293-312.
Koufos G.D. (1987d) - Study of the Turolian hipparions of the lower Axios Valley (Macedonia, Greece). 2. Locality “Prochoma-1” (PXM). Palaontologische Zeitschrift, 61: 339-358.
Koufos G.D. (1988) - Study of the Turolian hipparions of the lower Axios valley (Macedonia, Greece). 4. Localities of Dytiko. Palaeovertebrata, 18: 187-239.
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. (2024) - Updating the fauna and age of the Neogene-Quaternary large mammal sites of Greece. Geobios, 85: 35-57.
Koufos G.D. & Kostopoulos D.S. (1994) - The late Miocene mammal localities of Kemiklitepe (Turkey). 3. Equidae. Bulletin du Muséum National d’Histoire Naturelle Paris, 4eme sér., 16: 41-80.
Koufos G.D. & Vlachou T. (2005) - Equidae (Mammalia, Perissodactyla) from the late Miocene of Akkaşdağı, Türkiye. Geodiversitas, 27: 633-705.
Koufos G.D. & Vlachou T.D. (2016) - Equidae. In: Koufos G.D. & Kostopoulos D.S. (Eds) - Palaeontology of the upper Miocene vertebrate localities of Nikiti (Chalkidiki peninsula, Macedonia, Greece). Geobios 49: 85-118
Koufos G.D. & Vlachou T.D. (2019) - New material and revision of hipparions from the upper Miocene locality Ravin des Zouaves-5, RZO of Axios Valley (Macedonia, Greece). Geobios, 57: 33-52.
Koufos G.D., Kostopoulos D.S. & Vlachou T.D. (2009) - The Late Miocene Mammal Faunas of the Mytilinii Basin, Samos Island, Greece: New Collection 16. Biochronology. Beiträge zur Paläontologie, 31: 397-408.
Kormos T. (1914) - Évben végzett âsatâsaim eredményi. Külônlenyomat A Magy. Kiraly Fôldtani Intezet 1913, Evi Jelentesebol: 506-523.
Kretzoi M. (1983) - Gerinces Indexfajok Felsö-neozói rétegtanunkban Hipparion. Magyr Állami Földtani Intézet Évi Jelentése AZ.: 513-521.
Leidy J. (1856) - Notices of some remains of extinct Mammalia, recently discovered by Dr. F. V. Hayden, in the badlands of Nebraska. Proceedings of the Academy of Natural Sciences of Philadelphia, 8: 59.
Linnaeus C. (1758) - Systema Naturae per Regna Tria Naturae, Secundum Classes, Ordines, Genera, Species Cum Characteribus, Differentiis, Synonimis, Locis: Editio Decima, Reformata. Laurentii Salvii, Stockholm, 824 pp.
Liu T., Li C. & Zhai R. (1978) - Pliocene vertebrates of Lantian, Shensi. Professional Papers of Stratigraphy and Palaeontology, 7: 149-200.
Lyddeker R. (1877) - Notices of new and other vertebrata from Indian Tertiary and Secondary Rocks. Record of the Geological Survey of India, 10: 30-43.
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.
Matthew W.D. (1926) -The evolution of the horse; a record and its interpretation. The Quarterly Review of Biology, 1, 139-185.
Meijers M.J.M., Brocard G.Y., Cosca M.A., Lüdecke T., Teyssier C., Whitney D.L. & Mulch A. (2018) - Rapid late Miocene surface uplift of the Central Anatolian Plateau margin. Earth and Planetary Science Letters, 497: 29-41.
Meijers M.J.M., Mikes T., Rojay F.B., Aydar E., Çubukçu H.E., Wagner T., Lüdecke T., Krsnik E., Fiebig J. & Mulch A. (2025) - Miocene decoupling of surface uplift of the Central Anatolian Plateau interior (Türkiye) and its modern mountainous margins. Earth and Planetary Science Letters, 663: 119414.
The NOW Community (2026) - New and Old Worlds Database of Fossil Mammals (NOW). Licensed under CC BY 4.0. Retrieved 2025) from https://nowdatabase.org/now/database/
Owen R. (1848) - Description of the teeth and portions of jaws of two extinct anthracotheroid quadrupeds (Hyopotamus vectianus and Hyopotamus bovinus) discovered by the Marchioness of Hastings in the Eocene deposits of the N.W. coast of the Isle of Wight: with an attempt to develop Cuvier’s idea of the classification of Pachyderms by the number of their toes. Quarterly Journal of the Geological Society of London, 4: 103-141.
Pirlot P.L. (1956) - Les formes européennes du genre Hipparion. Memorias y Comunicaciones del Instituto Geológico, Diputación Barcelona, 14: 1-121.
Qiu Z., Huang W. & Guo Z. (1987) - The Chinese hipparionine fossils. Palaeontologia Sinica New Series C, 17: 1-250.
Quinn J.H. (1955) - Miocene Equidae of the Texas Gulf Costal Plain. Bureau of Economic Geology, The University of Texas, Austin, 97 pp.
R Core Team (2025) - R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. URL https://www.R-project.org/
Roth J. & Wagner A. (1855) - Die fossilen Knochenüberreste von Pikermi in Griechenland. Abhandlungen der Mathematisch-Physikalischen Klasse der Königlich Bayerischen Akademie der Wissenschaften, 7: 371-464.
Saarinen J. & Liu L. (2024) - Quantitative paleoenvironmental reconstructions based on large mammal communities in Björn Kurtén’s work and since then—revising the case of later Late Miocene Old World “Hipparion faunas”. In: Annales Zoologici Fennici, 61(1): 179-232. Finnish Zoological and Botanical Publishing Board.
Sefve I. (1927) - Die Hipparionen Nord-Chinas. Palaeontologia Sinica, series C, 2: 1-54.
Sen S. (1997) - Magnetostratigraphic calibration of the European Neogene mammal chronology. Palaeogeography, Palaeoclimatology, Palaeoecology, 133: 181-204.
Solounias N. (1981) - The Turolian fauna from the island of Samos, Greece. Contributions to Vertebrate Evolution, 6: 1-67.
Sondaar P. (1961) - Les Hipparion de l’Aragón meridional. Estudios Geologicos, 17 : 209-305.
Sondaar P.Y. (1971) - The Samos Hipparion. Proceedings of the Koninklijke Nederlandse Akademie van Wetenschappen, Series B, 74: 417-441.
Steininger F.F., Berggren W.A., Kent D.V., Bernor R.L., Sen S. & Agusti J. (1996) - Circum-Mediterranean Neogene (Miocene and Pliocene) marine-continental chronologic correlations of European Mammal Units. In: Bernor R.L., Fahlbusch V. & Mittmann H.-W. (Eds) - The Evolution of Western Eurasian Neogene Mammal Faunas: 7-46. Columbia University Press, New York.
Studer T. (1911) - Eine neue Equidenform aus dem Obermiocän von Samos. Verhandlungen der Zoologisch-Botanischen Gesellschaft, 1911: 192-200.
Skinner M.F. & MacFadden B.J. (1977) - Cormohipparion n. gen. (Mammalia, Equidae) from the North American Miocene (Barstovian-Clarendonian). Journal of Paleontology, 51: 912-926.
Swisher III C.C. (1996) - New 40Ar/39Ar dates and their contribution toward a revised chronology for the Late Miocene of Europe and West Asia. In: Bernor R.L., Fahlbusch V. & Mittmann H.-W. (Eds) - The Evolution of Western Eurasian Neogene Mammal Faunas: 64-77. Columbia University Press, New York.
Şengör A.M.C. & Yilmaz Y. (1981) - Tethyan evolution of Turkey: A plate tectonic approach. Tectonophysics, 75: 181-241.
Studer T. (1911) - Eine neue Equidenform aus dem Obermiocän von Samos. Verhandlungen der Deutschen Zoologischen Gessllschaft, 20-21: 192-200.
Tholt A., Başoğlu O., Bektaş Y., Bernor R.L., Carlson J.P., Dağ ö., Doğan U., Erkman A.C., Kaya F., Kaymakçı N., Kırmızıoğlu P.G., Meijers M.J.M., Kahya 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, 122: e2424428122.
Van Hopen E.C.N. (1930) - Fossiele Pferde van Conrelia, O.V.S. Paleontologiese Navorsing van die Nasionale Museum Bloemfontein, 2: 13-24.
Van Hopen E.C.N. (1932) - Die Stamlyn van die Sebras. Paleontologiese Navorsing van die Nasionale Museum Bloemfontein, 2: 25-37.
Venables W.N. & Ripley B.D. (2008) - Modern Applied Statistics with R. Springer-Verlag, New York, 498 pp.
Villalta J.F. & Crusafont M. (1957) - Dos nuevas especies de Hipparion del Pikermiese Español. Cursillos y Conferencias del Instituto Lucas Mallada, 4: 65-69.
Vlachou T.D. & Koufos G.D. (2002) - The hipparions (Mammalia, Perissodactyla) from the Turolian locality of Nikiti-2, Chalkidiki, Macedonia, Greece. Annales de Paleontologie, 88: 167-186.
Vlachou T.D. & Koufos G.D. (2004) - Preliminary tests on the study of the Turolian hipparions of Greece. Proceedings of the 5th International Symposium on Eastern Mediterranean, 1-5.
Vlachou T.D. & Koufos G.D. (2006) - The late Miocene vertebrate locality of Perivolaki, Thessaly, Greece. 6. Equidae. Palaeontographica Abteilung A, 276: 81-119.
Vlachou T.D. & Koufos G.D. (2009) - The Late Miocene mammal faunas of the Mytilinii Basin, Samos Island, Greece: new collection. Equidae. Beiträge zur Paläontologie, 31: 207-281.
von Meyer H. (1829) - In: von Leonhard K.C.R. (Eds) - Taschenbuch fur die gesammte Mineralogie mit Hinsicht auf die neuesten Entdeckungen herausgegeben: 150-152. Akademische Buchhandlung von J. C. B. Mohr.
Weidmann M., Solounias N., Drake R.E & Curtis G.H. (1984) - Neogene stratigraphy of the eastern basin, Samos Island, Greece. Geobios, 17: 477-490.
Wickham H. (2016) - ggplot2: Elegant Graphics for Data Analysis. Springer-Verlag, New York, 213 pp.
Wehrli H. (1941) - Beitrag zur Kenntnis der Hipparionen von Samos. Paläeontologie Zeitschrift, 22: 321-386.
Whitney D.L., Delph J.R., Thomson S.N., Beck S.L., Brocard G.Y., Cosca M.A., Darin M.H., Kaymakci N., Meijers M.J.M., Okay A.I., Rojay B., Teyssier C. & Umhoefer P. J. (2023) - Breaking plates: creation of the East Anatolian Fault, the Anatolian Plate, and a tectonic escape system. Geology, 51: 673-677.
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.
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). Annals of the Natural History Museum of 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.
Zhegallo V.I. (1971) - Hipparions from the Neogene deposits of Western Mongolia and Tuva. Sovm. Trudy Sovmestnoy Sovetsko-Mongol’skoy Paleontologicheskoy Ekspeditsii, 3: 98-119.
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Copyright (c) 2026 ÖMER DAĞ, OKŞAN BAŞOĞLU, OMAR CIRILLI, FERHAT KAYA, PINAR GÖZLÜK-KIRMIZIOĞLU, ÖZGE KAHYA-PARILDAR, CESUR PEHLEVAN, MAUD J. M. MEIJERS, EMRAH ŞIMŞEK, RAYMOND L. BERNOR
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Accepted 2026-03-25
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