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New insights on the Early Pleistocene equids from Roca-Neyra (France, central Europe): implications for the Hipparion LAD and the Equus FAD in Europe

Published online by Cambridge University Press:  17 December 2020

Omar Cirilli Open the ORCID record for Omar Cirilli [Opens in a new window] ,
Raymond L. Bernor Open the ORCID record for Raymond L. Bernor [Opens in a new window]  and
Lorenzo Rook Open the ORCID record for Lorenzo Rook [Opens in a new window]
Omar Cirilli*
Affiliation:
Dottorato di Ricerca in Scienze della Terra, Università di Pisa, Via S. Maria 53, 56126Pisa, Italy Dipartimento di Scienze della Terra, Paleo[Fab]Lab, Università degli Studi di Firenze, Via G. La Pira 4, 50121Firenze, Italy
Raymond L. Bernor
Affiliation:
College of Medicine, Department of Anatomy, Laboratory of Evolutionary Biology, 520 W St. N.W., 20059, WashingtonD.C., USA Human Origins Program, Department of Anthropology, Smithsonian Institution, 20560, WashingtonD.C., USA
Lorenzo Rook
Affiliation:
Dipartimento di Scienze della Terra, Paleo[Fab]Lab, Università degli Studi di Firenze, Via G. La Pira 4, 50121Firenze, Italy
*
*Corresponding Author
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Abstract

We undertake a redescription of the equid sample from the Early Pleistocene of Roca-Neyra, France. This locality has been recently calibrated at the Pliocene/Pleistocene boundary (2.6 ± 0.2 Ma) and therefore it is of interest for the first appearance of the genus Equus and last appearance of hipparionine horses. The Roca-Neyra equid sample, re-analyzed herein using morphological, morphometrical, and statistical analyses, has revealed the co-occurrence of Plesiohipparion cf. ?P. rocinantis and Equus cf. E. livenzovensis. The analysis undertaken on several European, African, and Asian “Hipparion” sensu lato species from late Miocene to Early Pleistocene has revealed different remnant Hipparion lineages in the Plio-Pleistocene of Europe: Plesiohipparion, Proboscidippaion, and likely Cremohipparion. The discovery of the first European monodactyl horse, Equus cf. E. livenzovensis correlates Roca-Neyra with other 2.6 Ma European localities in Italy, Spain, and in the Khapry area (Azov Sea region). The morphological description of the Equus cf. E. livenzovensis lower cheek teeth has highlighted intermediate features between the North American Pliocene species Equus simplicidens and Early Pleistocene European Equus stenonis. Our study supports the hypothesis that E. livenzovensis is a plausible evolutionary predecessor for the Equus stenonis group. These observations underscore the importance of Roca-Neyra as an important locality for the last European hipparions and the first Equus in the Early Pleistocene of Europe.

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Journal of Paleontology , Volume 95 , Issue 2 , March 2021 , pp. 406 - 425
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Copyright © The Author(s), 2020. Published by Cambridge University Press on behalf of The Paleontological Society

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References

Abel, O., 1926, Die Geschichte der Equiden auf dem Boden Noramerikas: Verhandlungen Zoologisch-Botanischen Gesellschaft, v. 74, p. 159–164.Google Scholar
Agusti, J., Vekua, A., Oms, O., Lordkipanidze, D., Bukhsianidze, M., Kiladze, G., and Rook, L., 2009, The Pliocene–Pleistocene succession of Kvabebi (Georgia) and the background to the early human occupation of Southern Caucasus: Quaternary Science Review, v. 28, p. 3275–3280.CrossRefGoogle Scholar
Alberdi, M.T., 1974, El género Hipparion en España, revision e historia evolutive: Trabajos sobre Neógeno-Cuaternario, v. 1, p. 1–139.Google Scholar
Alberdi, M.T., 1986, The Pliocene species of Hipparion and their biostratigraphical meanings: Geobios, v. 19, p. 517–522.CrossRefGoogle Scholar
Alberdi, M.T., 1989, A review of Old World hipparionine horses, in Prothero, D.R., and Schoch, R.M., eds., The Evolution of Perissodactils: New York and London, Oxford University Press, p. 234–261.Google Scholar
Alberdi, M.T., and Alcalà, L., 1999, A study of the new samples of the Pliocene Hipparion (Equidae, Mammalia) from Spain and Bulgaria: Transactions of the Royal Society of Edinburgh: Earth Science, v. 89, p. 167–186.CrossRefGoogle Scholar
Alberdi, M.T., Cerdeño, E., López-Martínez, , Morales, J., and Soria, M.D., 1997, La fauna Villafranquiense de El Rincón-1 (Albacete, Castilla-La Mancha): Estudios Geologicos, v. 53, p. 69–93.CrossRefGoogle Scholar
Alberdi, M.T., Ortiz Jaureguizar, E., and Prado, J.L., 1998, A quantitative review of European stenonoid horses: Journal of Paleontology, v. 72, p. 371–387.CrossRefGoogle Scholar
Alexeeva [Alekseeva], L.I., and Garutt, V.E., 1965, Novye dannye ob evolyutsii slonov roda Archidiskodon: Byulleten komissii po lzucheniyu Chetvertichnogo Perioda, AN SSSR, v. 30, p. 161–166. [in Russian]Google Scholar
Arambourg, C., 1959, Vertébrés continentaux du Miocène supérieur de l'Afrique du Nord: Mémoire du Service de la Carte Géologique d'Algérie, Paléontologie, v. 4, p. 1–161.Google Scholar
Athanassiou, A., 2018, A Villafranchian Hipparion-bearing mammal fauna from Sésklo (E. Thessaly, Greece): implications for the question of Hipparion-Equus sympatry in Europe: Quaternary, v. 1, 12. p. 1–24. doi:10.3390/quat1020012.CrossRefGoogle Scholar
Aymard, A., 1854, Des Terrains Fossiliferes du Bassin Superior de la Lorire: Comptes Rendus de l'Academia des Science de France, Series D, v. 38, p. 673–677.Google Scholar
Azanza, B., Morales, M., Andrés, M., Cerdeño, E., and Alberdi, M.T., 2016, Los macromamíferos y la datación del yacimiento clásico de Villarroya, in Alberdi, M.T., Azanza, B., and Cervante, E., eds, Villarroya, Yacimiento Clave de la Paleontologìa Riojana: Instituto de Estudios Riojanos, v. 56, p. 267–280.Google Scholar
Azzaroli, A., 1964, The two Villafranchian horses of the Upper Valdarno: Palaeontographia Italica, v. 59, p. 1–12.Google Scholar
Azzaroli, A., 1982, On Villafranchian Palaearctic equids and their allies: Palaeontographia Italica, v. 72, p. 74–97.Google Scholar
Azzaroli, A., 1983, Quaternary mammals and the “end-Villafranchian” dispersal event—a turning point in the history of Eurasia: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 44, p. 117–139. doi:10.1016/0031-0182(83)90008-1.CrossRefGoogle Scholar
Azzaroli, A., 1992, Ascent and decline of monodactyl equids: a case for prehistoric overkill: Annales Zoologici Fennici, v. 28, p. 151–163.Google Scholar
Azzaroli, A., 2000, On Equus livenzovensis Bajgusheva, 1978 and the “stenonid” lineage of equids: Palaeontographia Italica, v. 87, p. 1–17.Google Scholar
Azzaroli, A., 2002, Phylogeny of the genus Equus L.: Palaeontographia Italica, v. 84, p. 11–16.Google Scholar
Azzaroli, A., and Voorhies, M.R., 1993, The Genus Equus in North America. The Blancan species: Palaeontographia Italica, v. 80, p. 175–198.Google Scholar
Bajgusheva [Baigusheva], V.S., 1978, Krupnaya loshad Khaprovskogo kompleksa iz allyuviya Severo-vostochnogo Priazoviya: Izvestiya Severo-Kavkazkogo Nauchnogo Tsentra Vyschei Shkoly, Estestvennye Nauki, v. 1, p. 98–102. [in Russian]Google Scholar
Bajgusheva, V.S., Titov, V.V., and Tesakov, A.S., 2001, The sequence of Plio-Pleistocene mammal faunas from south Russian Plain (the Azov Region): Bollettino della Società Paleontologica Italiana, v. 40, p. 133–138.Google Scholar
Barrón-Ortiz, C.I., Avilla, L.S., Jass, C.N., Bravo-Cuevas, V.M., Machado, H., and Mothé, D., 2019, What is Equus? Reconciling taxonomy and phylogenetic analyses: Frontiers in Ecology and Evolution, v. 7, n. 343. doi:10.3389/fevo.2019.00343.CrossRefGoogle Scholar
Bartolini Lucenti, S., 2017, Nyctereutes megamastoides (Canidae, Mammalia) from the early and middle Villafranchian (late Pliocene and Early Pleistocene) of the Lower Valdarno (Firenze and Pisa, Tuscany, Italy): Rivista Italiana di Paleontologia e Stratigrafia, v. 123, p. 211–218.Google Scholar
Berggren, W.A., and Van Couvering, J.A., 1974, The late Neogene: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 16, p. 1–215.Google Scholar
Bernor, R.L., and Armour-Chelu, M., 1999, Towards an evolutionary history of African hipparionine horses, in Bromage, T., and Schrenk, F., eds., African Biogeography, Climate Change and Early Hominid Evolution: New York, Oxford University Press, p. 189–215.Google Scholar
Bernor, R.L., and Harris, J.M., 2003, Systematics and evolutionary biology of the late Miocene and early Pliocene hipparionine horses from Lothagam, Kenya, in Harris, J.M., and Leakey, M.G., eds., Lothagam: The Dawn of Humanity in Eastern Africa: New York, Columbia University Press, p. 387–438.Google Scholar
Bernor, R.L., and Hussain, S.T., 1985, An assessment of the systematic, phylogenetic and biogeographic relationships of Siwalik hipparionine horses: Journal of Vertebrate Paleontology, v. 5, p. 32–87.CrossRefGoogle Scholar
Bernor, R.L., and Lipscomb, D., 1991, The systematic position of “Plesiohipparion” aff. huangeense (Equidae, Hipparionini) from Gülyazi, Turkey: Mitteilungen der Bayerischen Staatssammlung für Paläontologie und Historiesche Geologie, v. 31, p. 107–123.Google Scholar
Bernor, R.L., and Sen, S., 2017, The early Pliocene Plesiohipparion and Probiscidipparion (Equidae, Hipparionini) from Çalta, Turkey (Ruscinian Age, c. 4.0 Ma): Geodiversitas, v. 39, p. 28–41.CrossRefGoogle Scholar
Bernor, R.L., and Sun, B.Y., 2015, Morphology through ontogeny of Chinese Proboscidipparion and Plesiohipparion and observations on their Eurasian and African relatives: Vertebrata Palasiatica, v. 53, p. 73–92.Google Scholar
Bernor, R.L., and White, T.D., 2009, Systematics and biogeography of “Cormohipparion” africanum, Early Vallesian (MN 9, ca. 10.5 Ma) of Bou Hanifia, Algeria, in Albright, L.B., III, ed., Papers on Geology, Vertebrate Paleontology, and Biostratigraphy in Honour of Michael O. Woodburne: Museum of North Arizona Bulletin, v. 65, p. 635–658.Google Scholar
Bernor, R.L., Qiu, Z., and Hayek, L.A.C., 1989, Systematic revision of Chinese Hipparion species described by Sfeve, 1927: American Museum Novitates. v. 2984, p. 1–60.Google Scholar
Bernor, R.L., Tobien, H., and Woodburne, M.O., 1990, Patterns of Old Word hipparionine evolutionary diversification and biogeographic dxtension, in Lindsay, E.H., Fahlbusch, V., and Mein, P., eds, European Neogene Mammal Chronology: New York, Plenum Press, p. 263–319.CrossRefGoogle Scholar
Bernor, R.L., Koufos, G.D., Woodburne, M.O., and Fortelius, M., 1996, The evolutionary history and biochronology of European and southwestern Asian late Miocene and Pliocene hipparionine horses, in Bernor, R.L., Fahlbusch, V., and Mittman, H.W., eds., The Evolution of Western Eurasian Neogene Mammal Faunas: New York, Columbia University Press, p. 307–338.Google Scholar
Bernor, R.L., Tobien, H., Hayek, L.A.C., and Mittmann, H.W., 1997, Hippotherium primigenium (Equidae, Mammalia) from the late Miocene of Höwenegg (Hegau, Germany): Andrias, v. 10, p. 1–230.Google Scholar
Bernor, R.L., Scott, R. S., Fortelius, M., Kappelman, J., and Sen, S., 2003, Systematics and evolution of the late Miocene hipparions from Sinap, Turkey, in Fortelius, M., Kappelman, J., Sen, S., and Bernor, R.L., eds., The Geology and Paleontology of the Miocene Sinap Formation, Turkey: New York, Columbia University Press, p. 220–281.Google Scholar
Bernor, R.L., Scott, R.S., and Haile-Selassie, Y., 2005, A contribution to the evolutionary history of Ethiopian hipparionine horse: morphometric evidence from postcranial skeleton: Geodiveristas, v. 27, p. 133–158.Google Scholar
Bernor, R.L., Armour-Chelu, M., Gilbert, H., Kaiser, T.M., and Schulz, E., 2010, Equidae, in Werdelin, L., and Sanders, W.J., eds., Cenozoic Mammals of Africa: Berkeley, CA, University of California Press, p. 685–721.CrossRefGoogle Scholar
Bernor, R.L., Kaiser, T.M., Nelson, S.V., and Rook, L., 2011, Systematics and paleobiology of Hippotherium malpassi n. sp. from the latest Miocene of Baccinello V3 (Tuscany, Italy): Bollettino della Societa Paleontologica Italiana, v. 50, p. 175–208.Google Scholar
Bernor, R.L., Boaz, N.T., and Rook, L., 2012, Eurygnathohippus feibeli (Perissodacyla: Mammalia) from the late Miocene of As Sahabi (Libya) and its evolutionary and biogeographic significance: Bollettino della Società Paleontologica Italiana, v. 51, p. 39–48.Google Scholar
Bernor, R.L., Gilbert, H., Semprebon, G.M., Simpson, S., and Sileshi Semaw, S., 2013, Eurygnathohippus woldegabrieli, sp. nov. (Perissodactyla, Mammalia), from the middle Pliocene of Aramis, Ethiopia: Journal of Vertebrate Paleontology, v. 33, p. 1472–1485. doi:10.1080/02724634.2013.829741.CrossRefGoogle Scholar
Bernor, R.L., Sun, B., and Chen, Y., 2015, Plesiohipparion shanxiense n. sp. from the Early Pleistocene (Nihowanian) of E Shanxi, China: Bollettino della Società Paleontologica Italiana, v. 54, p. 197–210.Google Scholar
Bernor, R.L., Ataabadi, M.M., Meshida, K., and Wolf, D., 2016, The Maragheh hipparions, late Miocene of Azarbaijan, Iran: Paleobiodiversity and Palaeoenviroments, v. 96, p. 453–488. doi:10.1007/s12549-016-0235-2.CrossRefGoogle Scholar
Bernor, R.L., Göhlich, U.B., Harzhauser, M., and Semprebon, G.M., 2017, The Pannonian C hipparions from the Vienna basin: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 476, p. 28–41.CrossRefGoogle Scholar
Bernor, R.L., Wang, S., Liu, Y., Chen, Y., and Sun, B., 2018a, Shanxihippus dermatorhinus comb. nov. comparisons to Old World hipparions with specialized nasal apparati: Rivista Italiana di Paleontologia e Stratigrafia, v. 124, p. 361–386.Google Scholar
Bernor, R.L., Cirilli, O., Wang, S.Q., and Rook, L., 2018b, Equus cf. livenzovenzis from Montopoli, Italy (Early Pleistocene; MN16b; ca. 2.6 Ma): Bollettino della Socità Paleontologica Italiana, v. 57, p. 203–216. doi:10.4435/BSPI.2018.13.Google Scholar
Bernor, R.L., Cirilli, O., Jukar, A.M., Potts, R., Bukhsianidze, M., and Rook, L., 2019, Evolution of early Equus in Italy, Georgia, the Indian Subcontinent, East Africa, and the origins of African zebras: Frontiers in Ecology and Evolution, v. 7, n. 166. doi:10.3389/fevo.2019.00166.CrossRefGoogle Scholar
Bernor, R.L., Boaz, N., Cirilli, O., El-Shawaihdi, M.H., and Rook, L., 2020, Sahabi Eurygnathohippus feibeli: its systematic, stratigraphic, chronologic and biogeographic contexts: Rivista Italiana di Paleontologia e Stratigrafia, v. 126, p. 561–581.Google Scholar
Bout, P., 1960, Le Villafranchien du Valey et du Bassin Hydrographique Moyen et Supérieur de l'Allier: Correlations Francaises et Européennes [Ph.D. thesis]: Paris, Université de Paris, Imprimerie Jeanne d'Arc, le Puy, 344 p.Google Scholar
Bravo-Cuevas, V.M., and Jiménez-Hidalgo, E., 2019, Evolutionary significance of Equinae From the Mexican Neogene: Frontiers in Ecology and Evolution, v. 7, n. 287. doi:10.3389/fevo.2019.00287.CrossRefGoogle Scholar
Carranza-Castañeda, O., 2019, Dinohippus mexicanus (early-late, late, and latest Hemphillian) and the transition to genus Equus, in Central Mexico faunas: Frontiers in Earth Science, v. 7, n. 89. doi:10.3389/feart.2019.00089.CrossRefGoogle Scholar
Cherin, M., Iurino, D.A., and Sardella, R., 2013, Earliest occurrence of Puma pardoides (Owen, 1846) (Carnivora, Felidae) at the Plio/Pleistocene transition in western Europe: new evidence from the middle Villafranchian assemblage of Montopoli, Italy: Compes Rendus Palevol, v. 12, p. 165–171.CrossRefGoogle Scholar
Cirilli, O., Zouhri, S., El Boughabi, S., Benvenuti, M.G., Papini, M., Bernor, R.L., and 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, v. 126, p. 1–12.Google Scholar
Cirilli, O., Pandolfi, L., and Bernor, R.L., in press, The Villafranchian perissodactyls of Italy: knowledge of the fossil record and future research perspectives: Geobios. doi:10.1016/j.geobios.2020.09.001.CrossRefGoogle Scholar
Clauzon, G., Le Strat, P., Duvail, C., Do Couto, D., Suc, J.P., Molliex, S., Bache, F., Besson, D., Lindsay, E.H., Opdyke, N.D., Rubino, J.L., Popescu, S.M., Haq, B.U., and Gorini, C., 2015, The Roussilon basin (S. France): a case-study to distinguish local and regional events between 6 and 3 Ma: Marine and Petroleum Geology, v. 66, p. 18–40.CrossRefGoogle Scholar
Cocchi, I., 1867, L'uomo fossile dell'Italia Centrale: Memorie della Società Italiana di Scienze Naturali, v. 2, p. 1–180.Google Scholar
Cope, E.D., 1892, A contribution to the vertebrate paleontology of Texas: Proceedings of the American Philosophical Society, v. 30, p. 123–131.Google Scholar
Croizet, J.B., and Jobert, A., 1828, Recherches sur les ossemens fossils du Département du Puy-de-Dôme: Paris, Thibaut-Landriot, 224 p.Google Scholar
Crusafont, P.M., and Sondaar, P., 1971, Una nouvelle espece d’Hipparion du Pliocéne terminal d'Espagne: Palaeovertebrata, v. 4, p. 59–66.CrossRefGoogle Scholar
Dawkins, W., 1868, On a new species of deer from the Norwich Crag: Quarterly Journal of the Geological Society of London, v. 24, p. 516–518.CrossRefGoogle Scholar
Del Campana, D., 1914, La Lycyaena lunensis n. sp. dell'ossario di Olivola (Val di Magra): Palaeontographia Italica, v. 20, p. 87–104.Google Scholar
Delson, E., Faure, M., Guérin, C., Aprile, L., Argant, J., Blackwell, B.A.B., Debard, E., Harcourt-Smith, W., Martin-Suarez, E., Monguillon, A., Parenti, F., Pastre, J.F., Sen, S., Skinner, A.R., Swisher, C.C. III, and Valli, A.M.F., 2006, Franco-American renewed research at the late Villafranchian locality of Senèze (Haute-Loire, France): Courier Forschungsinstitut Senckenberg, v. 256, p. 275–290.Google Scholar
Deng, T., and Xue, X., 1999, Equus qingyangensis sp. nov. (Perissodactyla, Equidae) from the Early Pleistocene of Qingyang, Gansu, China: Vertebrata Palasiatica, v. 37, p. 62–74.Google Scholar
Depéret, C., 1884, Nouvelles etudes sur les Ruminants Pliocenes et Quaternaires d'Auvergne: Bulletin de la Société Géologique de France, v. 12, p. 247–284.Google Scholar
Depéret, C., 1890, Les animaux pliocènes du Roussilon: Mémoires de la Société Géologique de France, v. 3, p. 1–198.Google Scholar
Devèze de Chabriol, J.S., and Bouillet, J.B., 1827, Essai géologique et minéralogique sur les environs d'Issoire, Département du Puy-de-Dôme, et principalement sur la Montagne de Boulade: avec la description et les figures lithographiées des ossemens fossiles qui y ont eté recueillis: Clermont-Ferrand, France, Imprimerie de Thibaud-Landriot, 104 p.Google Scholar
Domingo, M.S., Alberdi, M.T., and Azanza, B., 2007, A new quantitative biochronological ordination for the upper Neogene mammalian localities of Spain: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 255, p. 361–376.CrossRefGoogle Scholar
Domingo, L., Koch, P.L., Fernández, M.H., Fox, D.L., Domingo, M.S., and Alberdi, M.T., 2013, Late Neogene and early Quaternary paleoenvironmental and paleoclimatic conditions in southwestern Europe: isotopic analyses on mammalian taxa: PLoS One, v. 8, n. 63739. doi.org/10.1371/journal.pone.0063739.CrossRefGoogle ScholarPubMed
Eisenmann, V., 1976, Nouveaux crânes d'hipparions (Mammalia, Perissodactyla) Plio-Pleistocènes d'Afrique orientale (Ethiopie et Kenya): Hipparion sp., Hipparion cf. ethiopicum et Hipparion afarense nov. sp: Géobios, v. 5, p. 577–605.CrossRefGoogle Scholar
Eisenmann, V., 1983, Family Equidae, in Harris, J.M., ed., Koobi Fora Research Project Volume 2, The Fossil Ungulates: Proboscidea, Perissodactyla and Suidae: Oxford, UK, Clarendon Press, p. 156–214.Google Scholar
Eisenmann, V., 1995, What metapodial morphometry has to say about some Miocene hipparions, in Vrba, E.S., Denton, G.H., Partridge, T.C., and Burckle, L.H., eds., Paleoclimate and Evolution: With Emphasis on Human Origins: New Haven, CT, Yale University Press, p. 148–163.Google Scholar
Eisenmann, V., 2019, Website, https://vera-eisenmann.comGoogle Scholar
Eisenmann, V., and Brunet, J., 1973, Présence simultanée de cheval et d’Hipparion dans le Villafranchian moyen de France, à Roccaneyra (Puy-de-Dôme); Etude critique de cas semblables (Europe et Proche-Orient): International Colloqium on the Problem: “The Boundary between Neogene and Quaternary,” Moscow, Collection of papers IV, p. 104–122.Google Scholar
Eisenmann, V., and Sondaar, P., 1998, Pliocene vertebrate locality of Calta, Ankara, Turkey. Hipparion: Geodiversitas, v. 20, p. 409–439.Google Scholar
Eisenmann, V., and Baylac, M., 2000, Extant and fossil Equus (Mammalia, Perissodactyla) skulls: a morphometric definition of the genus Equus: Zoologica Scripta, v. 29, p. 89–100.CrossRefGoogle Scholar
Eisenmann, V., and Deng, T., 2005, Equus qingyangensis (Equidae, Perissodactyla) of the upper Pliocne of Bajiazui, China: evidence for North American origin of an Old World lineage distinct from E. stenonis: Quaternaire, v. 2, p. 113–122.Google Scholar
Eisenmann, V., and Geraads, D., 2007, Hipparion pomeli sp. nov from the late Pliocene of Ahl al Oughlam, Morocco, and a revision of the relationships of Pliocene and Pleistocene African hipparions: Palaeontologia Africana, v. 42, p. 51–98.Google Scholar
Eisenmann, V., Alberdi, M.T., De Giuli, C., and Staesche, U., 1988, Methodology, in Woodburne, M., and Sondaar, P.Y., eds., Studying Fossil Horses: Leiden, The Netherlands, EJ Brill Press, p. 1–71.Google Scholar
Falconer, H., 1868, Palaeontological Memoirs of the Late Hugh Falconer, A.M., M.D., with a Biographical Sketch of the Author. Vol. II, Mastodon, Elephat, Rinoceros, Ossiferous Caves, Primeval Man and His Contemporaries: London, Robert Hardwicke, 860 p.Google Scholar
Forsten, A., 1968, Revision of the Palearctic Hipparion: Acta Zoologica Fennica, v. 119, p. 1–134.Google Scholar
Forsten, A., 1998, The fossil horses (Equidae, Mammalia) from the Plio-Pleistocene of Liventsovka near Rostov-Don, Russia: Geobios, v. 31, p. 645–657.CrossRefGoogle Scholar
Forsten, A., 2001, The hipparions (Mammalia, Equidae) of Suffolk, England: Transactions of the Royal Society of Edinburgh: Earth Science, v. 92, p. 115–120.Google Scholar
Forsten, A., 2002, Latest Hipparion Cristol, 1832 in Europe. A review of the Pliocene Hipparion crassum Gervais Group and other finds (Mammalia, Equidae): Geodiversitas, v. 24, p. 465–486.Google Scholar
Gervais, P., 1859, Zoologie et Paléontologie Française (Animaux Vertébrés): deuxième edition: Paris, Arthus Bertrand, 544 p.Google Scholar
Gray, H., 1821, On the natural arrangement of vertebrose animals: London Medical Repository Review, v. 15, p. 296–310.Google Scholar
Gromova, V., 1952, The Genus Hipparion: Trudy Paleontologie Institut, v. 36, 478 p. [in Russian]Google Scholar
Guérin, C., 1972, Une nouvelle espèce de rhinocéros, Mammalia, Perissodactyla) à Vialette, Haute-Loire, France) et dans d'autres gisements du Villafranchien Inférieur européen: Dicerorhinus jeanvireti n. sp: Documents des Laboratoires de Géologie de la Faculté des Sciences de Lyon, v. 49, p. 53–161.Google Scholar
Guérin, C., 1990, Biozones or Mammal Units? Methods and limits in biochronology, in Lindsay, E.H., Fahlbusch, V., and Mein, P., eds, European Neogene mammal chronology: New York, Plenum Press, p. 119–130.CrossRefGoogle Scholar
Hays, I., 1834, Descriptions of the specimens of inferior maxillary bones of Mastodons in the cabinet of the American Philosophical Society, with remarks on the genus Tetracaulodon (Godman): Transactions of the American Philosophical Society, v. 4, p. 317–339.CrossRefGoogle Scholar
Heintz, E., 1970, Les Cervides Villafranchiens de France et d'Espagne: Memoires du Museum national d'histoire naturelle, Series C, v. 22, p. 1–302.Google Scholar
Hernández-Pacheco, E., 1921, La llanura manchega y sus mamíferos fósiles: yacimiento de La Puebla de Almoradier: Comisión de Investigaciones Pleontológicas y Prehistóricas, Memoria, v. 28, p. 1–41.Google Scholar
Hooijer, D.A., and Maglio, V.J., 1974, Hipparions from the late Miocene and Pliocene of Northwestern Kenya: Zoologische Verhandelingen, v. 134, p. 1–34Google Scholar
Jukar, A., Sun, B., and Bernor, R.L., 2018, The first occurrence of Plesiohipparion huangheense (Qiu, Huang & Guo, 1987) (Equidae, Hipparionini) from the late Pliocene of India: Bollettino della Società Paleontologica Italiana, v. 57, p. 125–132.Google Scholar
Jukar, A.M., Sun, B., Nanda, A.K., and Bernor, R.L., 2019, The first occurrence of Eurygnathohippus (Mammalia, Perissodatyla, Equidae) outside Africa and its biogeographic significance: Bollettino della Società Paleontologica Italiana, v. 58, p. 171–179.Google Scholar
Koufos, G.D., 1986, Study of the Vallesian hipparions of the lower Axios Valley (Macedonia, Greece): Geobios, v. 19, p. 61–79.CrossRefGoogle Scholar
Koufos, G.D., and Vlachou, T., 2005, Equidae (Mammalia, Perissodactyla) from the late Miocene of Akkaşdağı, Turkey: Geodiversitas, v. 27, p. 633–705.Google Scholar
Kurten, B., 1963, Villafranchian faunal evolution: Commentationes Biologicae, Societatis Scientiarium Fennica, v. 26, p. 1-18.Google Scholar
Lindsay, E.H., Opdyke, N.D., and Johnson, N.M., 1980, Pliocene dispersal of the horse Equus and late Cenozoic mammal dispersal events: Nature, v. 287, p. 135–138. doi:10.1038/287135a0.CrossRefGoogle Scholar
Linnaeus, C., 1758, Systema naturae per regna tria naturae, secundum Classes, Ordines, Genera, Species cum characteribus, differentiis, synonimis, locis: Editio decima, reformata. Stockholm, Laurentii Salvii, 824 p.Google Scholar
López-Martínez, N., 1989, Revisión sistemática y biostratigráfica de los Lagomorpha (Mammalia) del Terciario y Cuaternario de España: Memorias del Museo Paleontologico de la Universidad de Zaragoza, v. 3, p. 5–343.Google Scholar
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, v. 179, p. 1–195.Google Scholar
Mein, P., 1999, European Miocene mammal biochronology, in Rössner, G.E., and Heissig, K., eds, The Miocene Land Mammals of Europe: München, Verlag Dr. Friedrich Pfeil, p. 25–38.Google Scholar
Nomade, S., Pastre, J.F., Guillou, H., Faure, M., Guérin, C., Delson, E., Debard, E., Voinchet, P., and Messager, E., 2014, 40Ar/39Ar constraints on some French landmark late Pliocene to Early Pleistocene large mammalian paleofaunas: paleoenvironmental and paleoecological implications: Quaternary Geochronology, v. 21, p. 2–15. doi:10.1016/j.quageo.2012.12.006.CrossRefGoogle Scholar
Oustalet, E., 1882, Une nouvelle espèce de zèbre, le zèbre de Grévy (Equus grevyi): La Nature, v. 10, p. 12–14.Google Scholar
Owen, R., 1846, A History of British Fossil Mammals, and Birds: London, John Van Voorst, 560 p.CrossRefGoogle Scholar
Owen, R., 1848, Description of the teeth and portions of jaws of two extinct anthracotheroid quadrupeds (Hyopotamus vectianus and Hyop. 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 Geological Society of London, v. 4, p. 103–141.CrossRefGoogle Scholar
Palombo, M.R., and Valli, A.M.F., 2004, Remarks on the biochronology of mammalian faunal complexes from the Pliocene to the Middle Pleistocene in France: Geologica Romana, v. 37, p. 145–163.Google Scholar
Palombo, M.R., and Alberdi, M.T., 2017, Light and shadows in the evolution of South European stenonoid horses: Fossil Imprint, v. 73, p. 115–140.CrossRefGoogle Scholar
Palombo, M.R., Alberdi, M.T., Bellucci, L., and Sardella, R., 2017, An intriguing middle-sized horse from Coste San Giacomo (Anagni Basin, central Italy): Quaternary Research, v. 87, p. 347–362.CrossRefGoogle Scholar
Pandolfi, L., 2013, New and revised occurrences of Dihoplus megarhinus (Mammalia Rhinocerotidae) in the Pliocene of Italy: Swiss Journal of Palaeontology, v. 132, p. 239–255.CrossRefGoogle Scholar
Pandolfi, L., Cerdeño, E., Codrea, V., and Kotsakis, T., 2017, Biogeography and chronology of the Eurasian extinct rhinoceros Stephanorhinus etruscus, Mammalia, Rhinocerotidae: Comptes Rendus Palevol, v. 16, p. 762–773.CrossRefGoogle Scholar
Pandolfi, L., Popuscu, A., and Codrea, V., 2019, Stephanorhinus jeanvireti (Mammalia, Rhinocerotidae) from the Early Pleistocene of Colțești, southwestern Romania: Comptes Rendus Palevol, v. 18, p. 1041–1056.CrossRefGoogle Scholar
Pastre, J.F., 2004, The Perrier Plateau: a Plio-Pleistocene long fluvial record in the river Allier basin, Massif Central, France: Quaternaire, v. 15, p. 87–102.CrossRefGoogle Scholar
Pilgrim, G.E., 1910, Notices of new mammalian genera and species from the Tertiaries of India: Records of the Geological Survey of India, v. 40, p. 63–71.Google Scholar
Pirlot, P.R., 1956, Les formes européennes du genre Hipparion: Memorias y comunicaciones del Instituto Geológico, Diputación Provincial de Barcelona, v. 14, p. 1–122.Google Scholar
Pomel, M., 1842, Nouvelle espèce de chien fossile découverte dans les alluvions volcaniques de l'Auvergne: Bulletin de la Societé géologique de France, v. 14, p. 38–41.Google Scholar
Pomel, A., 1853, Catalogue méthodique et descriptif des vertébrés fossiles découverts dans le bassin hydrographique supérieur de la Loire, et surtout dans la vallée de son affluent principal l'Allier: Paris, J.B. Ballière, 193 p.Google Scholar
Prat, F., 1964, Contribution a la classification des Equidés Villafranchiens: Procès Verbaux de la Socíété Linnéenne de Bordeaux, v. 101, p. 14–32.Google Scholar
Pueyo, E.L., Muñoz, A., Laplana, C., and Parés, J.M., 2016, The last appearance datum of Hipparion in Western Europe: magnetostratigraphy along the Pliocene-Pleistocene boundary in the Villarroya basin (northern Spain): International Journal of Earth Science, v. 105, p. 2203–2220.CrossRefGoogle Scholar
Qiu, Z., Huang, W., and Guo, Z., 1987, The Chinese hipparionine fossils: Palaeontologia Sinica, New series C, v. 17, p. 1–250.Google Scholar
Qiu, Z.X., Deng, T., and Wang, B.Y., 2004, Early Pleistocene mammalian fauna from Longdan, Dongxiang, Gansu, China: Palaeontologia Sinica, New Series C, v. 27, p. 162–164.Google Scholar
Quinn, J.H., 1955, Miocene Equidae of the Texas Gulf Coastal Plain: Bureau of Economic Geology, The University of Texas Report of Investigations, v. 5516, p. 1–102.Google Scholar
R Core Team, 2013, R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria, ISBN 3-900051-07-0. http://www.R-project.org/Google Scholar
Radulescu, C., and Samson, P., 1967, Sur la signifaction de certains Equidés du Plèistocéne inférieur et moyen de Roumanie: Neues Jahrbuch für Geologie und Paläontologie Abhandlungen, v. 127, p. 157–178.Google Scholar
Rook, L., and Martínez-Navarro, B., 2010, Villafranchian: the long story of a Plio-Pleistocene European large mammal biochronologic unit: Quaternary International, v. 219, p. 134–144.Google Scholar
Rook, L., Cirilli, O., and Bernor, R.L., 2017, A late occurring “Hipparion” from the middle Villafranchian of Montopoli, Italy (Early Pleistocene; MN16b; ca. 2.5 Ma): Bollettino della Società Paleontologica Italiana, v. 56, p. 333–339.Google Scholar
Rook, L., Bernor, R.L., Avilla, L.D., Cirilli, O., Flynn, L., Jukar, A.M., Sanders, W., Scott, E., and Wang, X., 2019, Mammal biochronology (Land Mammal Ages) around the world from the late Miocene to Middle Pleistocene and major events in horse evolutionary history: Frontiers in Ecology and Evolution, v. 7, n. 278. doi:10.3389/fevo.2019.00278.CrossRefGoogle Scholar
Roth, J., and Wagner, A., 1855, Die fissilen Knochenüberreste von Pikermi: Bayerischen Akademie Wisseenschaften, Abhandlungen, v. 7, p. 361–464.Google Scholar
Schaub, S., 1923, Neue und wenig bekannte Cavicornier von Seneze: Eclogae Geologicae Helvetiae, v. 18, p. 281–295.Google Scholar
Schaub, S., 1944, Die oberpliocäne Säugetierfauna von Senèze (Haute Loire) und ihre verbreitungsgeschichtliche Stellung: Eclogae Geologicae Helveticae, v. 36, p. 270–289.Google Scholar
Sefve, I., 1927, Die Hipparionen Nord-Chinas: Palaeontologia Sinica, series C, v. 4, p. 1–54.Google Scholar
Sesé, C., 2006, Los roedores y lagomorfos del Neógeno de España: Estudios Geológicos, v. 62, p. 429–480.CrossRefGoogle Scholar
Stehlin, H.G., 1904, Une faune à Hipparion à Perrier: Bulletin de la Société Géologique de France, v. 28, p. 432–444.Google Scholar
Sun, B., and Deng, T., 2019, The Equus datum and the early radiation of Equus in China: Frontiers in Ecology and Evolution, v. 7, n. 429. doi:10.3389/fevo.2019.00429.CrossRefGoogle Scholar
Sun, B., Zhang, X., Liu, Y., and Bernor, R.L., 2018, Sivalhippus ptychodus and Sivalhippus platyodus (Perissodactyla, Mammalia) from the late Miocene of China: Rivista Italiana di Paleontologia e Stratigrafia, v. 124, p. 1–22.Google Scholar
Team QD, 2016, QGIS geographic information system: Open source geospatial foundation project. https://qgis.org.Google Scholar
Teilhard de Chardin, P., and Piveteau, J., 1930, Les mammifères fossiles de fossiles de Nihowan (Chine): Annales de Paleontologie, v. 19, p. 87–119.Google Scholar
Teilhard de Chardin, P., and Young, C., 1931, Fossil mammals from late Cenozoic of northern China: Paleontologica Sinica, Serie C, v. 9, p. 1–67.Google Scholar
Van Hoepen, E.C.N., 1930, Fossiele Pferde van Cornelia: Paleontologiese Navorsing van die Nasionale Museum, Bloemfontein, v. 2, p. 13–24.Google Scholar
Vekua, A., 1972, Kvabebi fauna of Akchagylian Vertebrates: Moscow, Nauka, p. 96–112. [in Russian]Google Scholar
Villalta, J.F., 1948, Una nueva especie de Hipparion en el Villafranquiense: Museo de la Ciudad de Sabadell, v. 4, p. 5–14.Google Scholar
Villalta, J.F., 1952, Contribución al conocimiento de la fauna de mamíferos fósiles del Plioceno de Villarroya (Logroño): Boletín del Instituto Geológico y Minero de España, v. 64, p. 1–201.Google Scholar
von Meyer, H., 1829, Taschenbuch für die gesamte Mineralogie: Zeitschrift für Mineralogie, Neue Folge, v. 23, p. 150–152.Google Scholar
Wolf, D., Bernor, R.L., and Hussain, S.T., 2013, A systematic, biostratigraphic, and paleobiogeographic reevaluation of the Siwalik hipparionine horse assemblage from the Potwar Plateau, Northern Pakistan: Palaeontographica Abteilung A, v. 300, p. 1–115.CrossRefGoogle Scholar
Woodburne, M., and Bernor, R.L., 1980, On superspecific groups of some Old World hipparionine horses: Journal of Paleontology, v. 54, p. 1319–1348.Google Scholar
Wüst, E., 1900, Untersuchungen über das Pliozän und das Älteste Pleistozän Thüringens, nördlich vom Thüringer Walde und westlich von der Saale: Abhandlungen der Naturforschenden Gesellschaft zu Halle, v. 23, p. 1–352.Google Scholar
Zhegallo, V.I., 1978, The hipparions of Central Asia: Trudy Soviet Mongolian Palaeontological Expedition, v. 7, p. 1–152.Google Scholar
Zouhri, S., and Bensalmia, A., 2005, Révision systématique des Hipparion sensu lato (Peryssodactyla, Equidae) de l'Ancient Monde: Estudios Geologicos, v. 61, p. 61–99.CrossRefGoogle Scholar