{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,11,10]],"date-time":"2026-11-10T15:06:10Z","timestamp":1794323170658,"version":"4.9.0"},"reference-count":67,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2018,5,31]],"date-time":"2018-05-31T00:00:00Z","timestamp":1527724800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2018,5,31]],"date-time":"2018-05-31T00:00:00Z","timestamp":1527724800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Sci Rep"],"abstract":"<jats:title>Abstract<\/jats:title><jats:p>The sequence of cheek teeth mineralization, eruption, and replacement of an extinct horse species is here documented with radiological techniques for the first time thanks to the exceptional preservation of <jats:italic>Hipparion<\/jats:italic> sp. mandibles from Cerro de los Batallones (Madrid Basin, Spain). The sequence of dental ontogeny in mammals provides valuable insights about life history traits, such as the pace of growth, and about the mode of formation of fossiliferous assemblages. We have determined that the order of permanent cheek teeth mineralization and eruption of hipparionine horses is m1, m2, (p2, p3), p4, m3. Cheek teeth mineralization timing of hipparionine horses coincides with the one observed in modern equids. In turn, there are differences in the eruption timing of the p4 and m3 between horses belonging to the Anchitheriinae and Hipparionini compared to equids of the <jats:italic>Equus<\/jats:italic> genus that might be related to the shorter durability of the deciduous tooth dp4 in anchitheriine and hipparionine horses and, more broadly, to an increased durability of equid teeth through their evolutionary history. Based on the dental eruption sequence, hipparionine horses are slow-growing, long-living mammals. The <jats:italic>Hipparion<\/jats:italic> sp. assemblage from Batallones-10 conforms to an attritional model, as individuals more vulnerable to natural mortality predominate.<\/jats:p>","DOI":"10.1038\/s41598-018-26817-3","type":"journal-article","created":{"date-parts":[[2018,5,25]],"date-time":"2018-05-25T14:08:34Z","timestamp":1527257314000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":23,"title":["First Radiological Study of a Complete Dental Ontogeny Sequence of an Extinct Equid: Implications for Equidae Life History and Taphonomy"],"prefix":"10.1038","volume":"8","author":[{"given":"M. Soledad","family":"Domingo","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Enrique","family":"Cantero","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Isabel","family":"Garc\u00eda-Real","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Manuel J.","family":"Chamorro Sancho","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"David M.","family":"Mart\u00edn Perea","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"M. Teresa","family":"Alberdi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jorge","family":"Morales","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2018,5,31]]},"reference":[{"key":"26817_CR1","doi-asserted-by":"crossref","unstructured":"Matthew, W. D. The evolution of the horse: a record and its interpretation. Q. Rev. Biol. 1, 139\u2013185, http:\/\/www.jstor.org\/stable\/2808222 (1926).","DOI":"10.1086\/394242"},{"key":"26817_CR2","unstructured":"Simpson, G. G. Horses. (Oxford University Press, 1951)."},{"key":"26817_CR3","unstructured":"MacFadden, B. J. Fossil Horses: Systematics, Paleobiology, and Evolution of the Family Equidae. (Cambridge University Press, 1992)."},{"key":"26817_CR4","doi-asserted-by":"publisher","first-page":"1728","DOI":"10.1126\/science.1105458","volume":"307","author":"BJ MacFadden","year":"2005","unstructured":"MacFadden, B. J. Fossil horses \u2013 Evidence for evolution. Science 307, 1728\u20131730, https:\/\/doi.org\/10.1126\/science.1105458 (2005).","journal-title":"Science"},{"key":"26817_CR5","doi-asserted-by":"publisher","first-page":"236","DOI":"10.1666\/0094-8373(2006)32[236:EOHIET]2.0.CO;2","volume":"32","author":"CAE Str\u00f6mberg","year":"2006","unstructured":"Str\u00f6mberg, C. A. E. Evolution of hypsodonty in equids: testing a hypothesis of adaptation. Paleobiology 32, 236\u2013258, https:\/\/doi.org\/10.1666\/0094-8373(2006)32[236:EOHIET]2.0.CO;2 (2006).","journal-title":"Paleobiology"},{"key":"26817_CR6","doi-asserted-by":"publisher","first-page":"627","DOI":"10.1126\/science.aag1772","volume":"355","author":"JL Cantalapiedra","year":"2017","unstructured":"Cantalapiedra, J. L., Prado, J. L., Hern\u00e1ndez Fern\u00e1ndez, M. & Alberdi, M. T. Decoupled ecomorphological evolution and diversification in Neogene-Quaternary horses. Science 355, 627\u2013630, https:\/\/doi.org\/10.1126\/science.aag1772 (2017).","journal-title":"Science"},{"key":"26817_CR7","doi-asserted-by":"publisher","first-page":"824","DOI":"10.1126\/science.283.5403.824","volume":"283","author":"BJ MacFadden","year":"1999","unstructured":"MacFadden, B. J., Solounias, N. & Cerling, T. E. Ancient diets, ecology, and extinction of 5-million-year-old horses from Florida. Science 283, 824\u2013827, https:\/\/doi.org\/10.1126\/science.283.5403.824 (1999).","journal-title":"Science"},{"key":"26817_CR8","doi-asserted-by":"publisher","first-page":"67","DOI":"10.1002\/jmor.10125","volume":"258","author":"TM Kaiser","year":"2003","unstructured":"Kaiser, T. M. & Fortelius, M. Differential mesowear in occluding upper and lower molars: opening mesowear analysis for lower molars and premolars in hypsodont horses. J. Morphol. 258, 67\u201383, https:\/\/doi.org\/10.1002\/jmor.10125 (2003).","journal-title":"J. Morphol."},{"key":"26817_CR9","doi-asserted-by":"publisher","first-page":"503","DOI":"10.1007\/s00114-008-0500-y","volume":"96","author":"L Domingo","year":"2009","unstructured":"Domingo, L., Grimes, S. T., Domingo, M. S. & Alberdi, M. T. Paleoenvironmental conditions in the Spanish Miocene-Pliocene boundary: isotopic analyses of Hipparion dental enamel. Naturwissenchaften 96, 503\u2013511, https:\/\/doi.org\/10.1007\/s00114-008-0500-y (2009).","journal-title":"Naturwissenchaften"},{"key":"26817_CR10","doi-asserted-by":"publisher","first-page":"1178","DOI":"10.1126\/science.1196166","volume":"331","author":"MC Mihlbachler","year":"2011","unstructured":"Mihlbachler, M. C., Rivals, F., Solounias, N. & Semprebon, G. M. Dietary change and evolution of horses in North America. Science 331, 1178\u20131181, https:\/\/doi.org\/10.1126\/science.1196166 (2011).","journal-title":"Science"},{"key":"26817_CR11","doi-asserted-by":"publisher","first-page":"10","DOI":"10.1016\/j.palaeo.2015.11.004","volume":"442","author":"GM Semprebon","year":"2016","unstructured":"Semprebon, G. M., Rivals, F., Solounias, N. & Hulbert, R. C. Jr. Paleodietary reconstruction of fossil horses from the Eocene through Pleistocene of North America. Palaeogeogr. Palaeocl. Palaeoecol. 442, 10\u2013127, https:\/\/doi.org\/10.1016\/j.palaeo.2015.11.004 (2016).","journal-title":"Palaeogeogr. Palaeocl. Palaeoecol."},{"key":"26817_CR12","doi-asserted-by":"publisher","first-page":"361","DOI":"10.1016\/j.palaeo.2007.08.004","volume":"255","author":"MS Domingo","year":"2007","unstructured":"Domingo, M. S., Alberdi, M. T. & Azanza, B. A new quantitative biochronological ordination for the Upper Neogene mammalian localities of Spain. Palaeogeogr. Palaeocl. Palaeoecol. 255, 361\u2013376, https:\/\/doi.org\/10.1016\/j.palaeo.2007.08.004 (2007).","journal-title":"Palaeogeogr. Palaeocl. Palaeoecol."},{"key":"26817_CR13","doi-asserted-by":"publisher","first-page":"28","DOI":"10.1016\/j.palaeo.2017.03.026","volume":"476","author":"RL Bernor","year":"2017","unstructured":"Bernor, R. L., G\u00f6hlich, U. B., Harzhauser, M. & Semprebon, G. M. The Pannonian C hipparions from the Vienna Basin. Palaeogeogr. Palaeocl. Palaeoecol. 476, 28\u201341, https:\/\/doi.org\/10.1016\/j.palaeo.2017.03.026 (2017).","journal-title":"Palaeogeogr. Palaeocl. Palaeoecol."},{"key":"26817_CR14","unstructured":"Schultz, A. H. Age Changes in Primates and their Modification in Man in Human Growth (ed. Tanner, J. M.) 1\u201320 (Pergamon Press, 1960)."},{"key":"26817_CR15","doi-asserted-by":"crossref","unstructured":"Smith, B. H. \u2018Schultz\u2019s Rule\u2019 and the Evolution of Tooth Emergence and Replacement Patterns in Primates and Ungulates in Development, Function and Evolution of Teeth (eds. Teaford, M. F., Smith, M. M. & Ferguson, M. W. J.) 212\u2013227 (Cambridge University Press, 2000).","DOI":"10.1017\/CBO9780511542626.015"},{"key":"26817_CR16","doi-asserted-by":"publisher","first-page":"e70743","DOI":"10.1371\/journal.pone.0070743","volume":"8","author":"X Jordana","year":"2013","unstructured":"Jordana, X. et al. First fossil evidence for the advance of replacement teeth coupled with life history evolution along an anagenetic mammalian lineage. PLOS ONE 8, e70743, https:\/\/doi.org\/10.1371\/journal.pone.0070743 (2013).","journal-title":"PLOS ONE"},{"key":"26817_CR17","doi-asserted-by":"publisher","first-page":"251","DOI":"10.1007\/s10914-015-9315-8","volume":"23","author":"K Veitschegger","year":"2016","unstructured":"Veitschegger, K. & S\u00e1nchez-Villagra, M. R. Tooth eruption sequences in cervids and the effect of morphology, life history, and phylogeny. J. Mammal Evol. 23, 251\u2013263, https:\/\/doi.org\/10.1007\/s10914-015-9315-8 (2016).","journal-title":"J. Mammal Evol."},{"key":"26817_CR18","doi-asserted-by":"publisher","first-page":"1069","DOI":"10.1073\/pnas.1614029114","volume":"114","author":"H Gomes Rodrigues","year":"2017","unstructured":"Gomes Rodrigues, H., Herrel, A. & Billet, G. Ontogenetic and life history trait changes associated with convergent ecological specializations in extinct ungulate mammals. Proc. Natl. Acad. Sci. 114, 1069\u20131074, https:\/\/doi.org\/10.1073\/pnas.1614029114 (2017).","journal-title":"Proc. Natl. Acad. Sci."},{"issue":"special paper 1","key":"26817_CR19","doi-asserted-by":"publisher","first-page":"1","DOI":"10.2113\/gsrocky.8.special_paper_1.1","volume":"8","author":"MR Voorhies","year":"1969","unstructured":"Voorhies, M. R. Taphonomy and population dynamics of an Early Pliocene vertebrate fauna, Knox County, Nebraska. Rocky Mountain Geology 8(special paper 1), 1\u201369, https:\/\/doi.org\/10.2113\/gsrocky.8.special_paper_1.1 (1969).","journal-title":"Rocky Mountain Geology"},{"key":"26817_CR20","doi-asserted-by":"publisher","first-page":"131","DOI":"10.1127\/0077-7749\/2012\/0251","volume":"265","author":"MS Domingo","year":"2012","unstructured":"Domingo, M. S., Alberdi, M. T., Azanza, B. & Morales, J. Mortality patterns and skeletal physical condition of the carnivorans from the Miocene assemblage of Batallones-1 (Madrid Basin, Spain). N. Jb. Geol. Pal\u00e4ont. Abh. 265, 131\u2013145, https:\/\/doi.org\/10.1127\/0077-7749\/2012\/0251 (2012).","journal-title":"N. Jb. Geol. Pal\u00e4ont. Abh."},{"key":"26817_CR21","doi-asserted-by":"publisher","first-page":"265","DOI":"10.1007\/s10914-015-9313-x","volume":"23","author":"C B\u00f6hmer","year":"2016","unstructured":"B\u00f6hmer, C., Heissig, K. & R\u00f6ssner, G. E. Dental eruption series and replacement pattern in Miocene Prosantorhinus (Rhinocerotidae) as revealed by macroscopy and X-ray: implications for ontogeny and mortality profile. J. Mammal Evol. 23, 265\u2013279, https:\/\/doi.org\/10.1007\/s10914-015-9313-x (2016).","journal-title":"J. Mammal Evol."},{"key":"26817_CR22","doi-asserted-by":"publisher","first-page":"93","DOI":"10.1111\/j.1463-6395.1964.tb00712.x","volume":"45","author":"L Van Valen","year":"1964","unstructured":"Van Valen, L. Age in two fossil horse populations. Acta Zool. 45, 93\u2013106, https:\/\/doi.org\/10.1111\/j.1463-6395.1964.tb00712.x (1964).","journal-title":"Acta Zool."},{"key":"26817_CR23","doi-asserted-by":"publisher","first-page":"159","DOI":"10.1017\/S0094837300004504","volume":"8","author":"RC Hulbert Jr.","year":"1982","unstructured":"Hulbert, R. C. Jr. Population dynamics of the three-toed horse Neohipparion from the Late Miocene of Florida. Paleobiology 8, 159\u2013167 (1982. http:\/\/www.jstor.org\/stable\/2400452","journal-title":"Paleobiology"},{"key":"26817_CR24","doi-asserted-by":"publisher","first-page":"547","DOI":"10.1080\/02724634.1984.10012030","volume":"4","author":"RC Hulbert Jr.","year":"1984","unstructured":"Hulbert, R. C. Jr. Paleoecology and population dynamics of the Early Miocene (Hemingordian) horse Parahippus leonensis from the Thomas Farm Site, Florida. J. Vertebr. Paleontol. 4, 547\u2013558 (1984).","journal-title":"J. Vertebr. Paleontol."},{"key":"26817_CR25","doi-asserted-by":"publisher","first-page":"412","DOI":"10.1666\/0094-8373(2003)029<0412:DOLMRT>2.0.CO;2","volume":"29","author":"MC Mihlbachler","year":"2003","unstructured":"Mihlbachler, M. C. Demography of late Miocene rhinoceroses (Teleoceras proterum and Aphelops malacorhinus) from Florida: linking mortality and sociality in fossil assemblages. Paleobiology 29, 412\u2013428, https:\/\/doi.org\/10.1666\/0094-8373(2003)0290412:DOLMRT2.0.CO;2 (2003).","journal-title":"Paleobiology"},{"key":"26817_CR26","first-page":"449","volume":"45","author":"JA O\u2019Sullivan","year":"2005","unstructured":"O\u2019Sullivan, J. A. Population dynamics of Archaeohippus blackbergi (Mammalia; Equidae) from the Miocene Thomas Farm fossil site of Florida. Bull. Flo. Mus. Nat. Hist. 45, 449\u2013463 (2005).","journal-title":"Bull. Flo. Mus. Nat. Hist."},{"key":"26817_CR27","doi-asserted-by":"publisher","first-page":"481","DOI":"10.1007\/s12542-013-0207-z","volume":"88","author":"K Bastl","year":"2014","unstructured":"Bastl, K. & Nagel, D. First evidence of the tooth eruption sequence of the upper jaw in Hyaenodon (Hyaenodontidae, Mammalia) and new information on the ontogenetic development of its dentition. Pal\u00e4ontol. Z. 88, 481\u2013494, https:\/\/doi.org\/10.1007\/s12542-013-0207-z (2014).","journal-title":"Pal\u00e4ontol. Z."},{"key":"26817_CR28","doi-asserted-by":"publisher","first-page":"127","DOI":"10.1127\/0077-7749\/2013\/0300","volume":"267","author":"M Hellmund","year":"2013","unstructured":"Hellmund, M. Odontological and osteological investigations on propalaeotheriids (Mammalia, Equidae) from the Eocene Geiseltal Fosillagerst\u00e4tte (Central Germany) \u2013 a full range of extraordinary phenomena. N. Jb. Geol. Pal\u00e4ont. Abh. 267, 127\u2013154, https:\/\/doi.org\/10.1127\/0077-7749\/2013\/0300 (2013).","journal-title":"N. Jb. Geol. Pal\u00e4ont. Abh."},{"key":"26817_CR29","doi-asserted-by":"publisher","first-page":"e0175460","DOI":"10.1371\/journal.pone.0175460","volume":"12","author":"Y Li","year":"2017","unstructured":"Li, Y., Deng, T., Hua, H., Li, Y. & Zhang, Y. Assessment of dental ontogeny in late Miocene hipparionines from the Lamagou fauna of Fugu, Shaanxi Province, China. PLOS ONE 12, e0175460, https:\/\/doi.org\/10.1371\/journal.pone.0175460 (2017).","journal-title":"PLOS ONE"},{"key":"26817_CR30","first-page":"41","volume":"8","author":"J Morales","year":"2008","unstructured":"Morales, J. et al. El sistema de yacimientos de mam\u00edferos miocenos del Cerro de los Batallones, Cuenca de Madrid: estado actual y perspectivas. Paleontologica Nova 8, 41\u2013117 (2008).","journal-title":"Paleontologica Nova"},{"key":"26817_CR31","doi-asserted-by":"publisher","first-page":"e63046","DOI":"10.1371\/journal.pone.0063046","volume":"8","author":"MS Domingo","year":"2013","unstructured":"Domingo, M. S., Alberdi, M. T., Azanza, B., Silva, P. G. & Morales, J. Origin of an assemblage massively dominated by carnivorans from the Miocene of Spain. PLOS ONE 8, e63046, https:\/\/doi.org\/10.1371\/journal.pone.0063046 (2013).","journal-title":"PLOS ONE"},{"key":"26817_CR32","doi-asserted-by":"publisher","first-page":"725","DOI":"10.1111\/j.1096-3642.2010.00618.x","volume":"160","author":"R L\u00f3pez-Anto\u00f1anzas","year":"2010","unstructured":"L\u00f3pez-Anto\u00f1anzas, R., Pel\u00e1ez-Campomanes, P., \u00c1lvarez-Sierra, M. A. & Garc\u00eda-Paredes, I. New species of Hispanomys (Rodentina, Cricetodontinae) from the Upper Miocene of Batallones (Madrid, Spain). Zool. J. Linn. Soc. 160, 725\u2013747, https:\/\/doi.org\/10.1111\/j.1096-3642.2010.00618.x (2010).","journal-title":"Zool. J. Linn. Soc."},{"key":"26817_CR33","doi-asserted-by":"publisher","first-page":"193","DOI":"10.1016\/j.palaeo.2011.05.014","volume":"307","author":"AR G\u00f3mez Cano","year":"2011","unstructured":"G\u00f3mez Cano, A. R., Hern\u00e1ndez Fern\u00e1ndez, M. & \u00c1lvarez-Sierra, M. A. 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. Palaeogeogr. Palaeocl. Palaeoecol. 307, 193\u2013204, https:\/\/doi.org\/10.1016\/j.palaeo.2011.05.014 (2011).","journal-title":"Palaeogeogr. Palaeocl. Palaeoecol."},{"key":"26817_CR34","doi-asserted-by":"publisher","first-page":"315","DOI":"10.1017\/S0094837300007077","volume":"8","author":"MO Woodburne","year":"1982","unstructured":"Woodburne, M. O. & MacFadden, B. J. A reappraisal of the systematics, biogeography, and evolution of fossil horses. Paleobiology 8, 315\u2013327 (1982).","journal-title":"Paleobiology"},{"key":"26817_CR35","doi-asserted-by":"publisher","first-page":"e103708","DOI":"10.1371\/journal.pone.0103708","volume":"9","author":"C Mart\u00ednez Maza","year":"2014","unstructured":"Mart\u00ednez Maza, C., Alberdi, M. T., Nieto-D\u00edaz, M. & Prado, J. L. Life-history traits of the Miocene Hipparion concudense (Spain) inferred from bone histological structure. PLOS ONE 9, e103708, https:\/\/doi.org\/10.1371\/journal.pone.0103708 (2014).","journal-title":"PLOS ONE"},{"key":"26817_CR36","first-page":"103","volume":"4","author":"GL Smuts","year":"1974","unstructured":"Smuts, G. L. Age determination in Burchell\u2019s zebra (Equus burchelli antiquorum) from the Kruger National Park. J. Sth. Afr. Wildl. Mgmt. Ass. 4, 103\u2013115 (1974).","journal-title":"J. Sth. Afr. Wildl. Mgmt. Ass."},{"key":"26817_CR37","unstructured":"Levine, M. A. The Use of Crown Height Measurements and Eruption-Wear Sequences to Age Horse Teeth in Ageing and Sexing Animal Bones from Archaeological Sites (eds Wilson, B., Grigson, C. & Payne, S.) 223\u2013250 (BAR British Series, 1982)."},{"key":"26817_CR38","doi-asserted-by":"crossref","unstructured":"Dixon, P. M. & Copeland, A. N. The radiological appearance of mandibular cheek teeth in ponies of different ages. Equine Vet. Educ. 5, 317\u2013323 1111\/j.2042-3292.1993.tb01061.x (1993).","DOI":"10.1111\/j.2042-3292.1993.tb01061.x"},{"key":"26817_CR39","first-page":"27","volume":"19","author":"NA Misk","year":"1997","unstructured":"Misk, N. A. & Seilem, S. M. Radiographic studies on the development of cheek teeth in donkeys. Equine Pract. 19, 27\u201338 (1997).","journal-title":"Equine Pract."},{"key":"26817_CR40","doi-asserted-by":"publisher","first-page":"355","DOI":"10.1016\/j.palaeo.2004.01.012","volume":"206","author":"KA Hoppe","year":"2004","unstructured":"Hoppe, K. A., Stover, S. M., Pascoe, J. R. & Amundson, R. Tooth enamel biomineralization in extant horses: implications for isotopic microsampling. Palaeogeogr. Palaeocl. Palaeoecol. 206, 355\u2013365, https:\/\/doi.org\/10.1016\/j.palaeo.2004.01.012 (2004).","journal-title":"Palaeogeogr. Palaeocl. Palaeoecol."},{"key":"26817_CR41","doi-asserted-by":"publisher","first-page":"77","DOI":"10.1159\/000083615","volume":"76","author":"LR Godfrey","year":"2005","unstructured":"Godfrey, L. R., Samonds, K. E., Wright, P. C. & King, S. J. Schultz\u2019s unruly rule: dental developmental sequences and schedules in small-bodied, folivorous lemurs. Folia Primatol. 76, 77\u201399, https:\/\/doi.org\/10.1159\/000083615 (2005).","journal-title":"Folia Primatol."},{"key":"26817_CR42","doi-asserted-by":"publisher","first-page":"355","DOI":"10.1017\/S0094837300003109","volume":"12","author":"BJ MacFadden","year":"1986","unstructured":"MacFadden, B. J. Fossil horses from \u201cEohippus\u201d (Hyracotherium) to Equus: scaling, Cope\u2019s Law, and the evolution of body size. Paleobiology 12, 355\u2013369 (1986. http:\/\/www.jstor.org\/stable\/2400511","journal-title":"Paleobiology"},{"key":"26817_CR43","doi-asserted-by":"publisher","first-page":"20122138","DOI":"10.1098\/rspb.2012.2138","volume":"280","author":"MS Domingo","year":"2013","unstructured":"Domingo, M. S., Domingo, L., Badgley, C., Sanisidro, O. & Morales, J. Resource partitioning among top predators in a Miocene food web. Proc. Royal Soc. B 280, 20122138, https:\/\/doi.org\/10.1098\/rspb.2012.2138 (2013).","journal-title":"Proc. Royal Soc. B"},{"key":"26817_CR44","doi-asserted-by":"publisher","first-page":"268","DOI":"10.1098\/rsbl.2006.0610","volume":"3","author":"V Veiberg","year":"2007","unstructured":"Veiberg, V. et al. Bigger teeth for longer life? Longevity and molar height in two roe deer populations. Biol. Lett. 3, 268\u2013270, https:\/\/doi.org\/10.1098\/rsbl.2006.0610 (2007).","journal-title":"Biol. Lett."},{"key":"26817_CR45","doi-asserted-by":"publisher","first-page":"3339","DOI":"10.1098\/rspb.2012.0689","volume":"279","author":"X Jordana","year":"2012","unstructured":"Jordana, X., Mar\u00edn-Moratalla, N., DeMiguel, D., Kaiser, T. M. & K\u00f6hler, M. Evidence of correlated evolution of hypsodonty and exceptional longevity in endemic insular mammals. Proc. Royal Soc. B 279, 3339\u20133346, https:\/\/doi.org\/10.1098\/rspb.2012.0689 (2012).","journal-title":"Proc. Royal Soc. B"},{"key":"26817_CR46","doi-asserted-by":"crossref","unstructured":"Bernor, R. L., Armour-Chelu, M. J., Gilbert, H., Kaiser, T. M. & Schulz, E. Equidae in Cenozoic mammals of Africa (eds. Werdelin, L. & Sanders, W. J.) 685\u2013721 (University of California Press, 2010).","DOI":"10.1525\/california\/9780520257214.003.0035"},{"key":"26817_CR47","first-page":"73","volume":"53","author":"RL Bernor","year":"2015","unstructured":"Bernor, R. L. & Sun, B.-Y. Morphology through ontogeny of Chinese Proboscidipparion and Plesiohipparion and observations of Eurasian and African relatives. Vertebrata PalAsiatica 53, 73\u201392 (2015).","journal-title":"Vertebrata PalAsiatica"},{"key":"26817_CR48","unstructured":"Janis, C. The Horse Series in Icons in Evolution (ed. Regal, B.) 251\u2013280 (Greenwood Press, 2007)."},{"key":"26817_CR49","doi-asserted-by":"publisher","first-page":"397","DOI":"10.2307\/3515249","volume":"11","author":"JD Bryant","year":"1996","unstructured":"Bryant, J. D., Froelich, P. N., Showers, W. J. & Genna, B. J. A tale of two quarries: biologic and taphonomic signatures in the oxygen isotope composition of tooth enamel phosphate from modern and Miocene equids. PALAIOS 11, 397\u2013408 (1996).","journal-title":"PALAIOS"},{"key":"26817_CR50","doi-asserted-by":"publisher","first-page":"337","DOI":"10.1016\/j.palaeo.2004.01.011","volume":"206","author":"P Higgins","year":"2004","unstructured":"Higgins, P. & MacFadden, B. J. \u201cAmount Effect\u201d recorded in oxygen isotopes of Late Glacial horse (Equus) and bison (Bison) teeth from the Sonoran and Chihuahuan deserts, southwestern United States. Palaeogeogr. Palaeocl. Palaeoecol. 206, 337\u2013353 (2004).","journal-title":"Palaeogeogr. Palaeocl. Palaeoecol."},{"key":"26817_CR51","doi-asserted-by":"crossref","unstructured":"McDonald, H. G. Population Structure of the Late Pliocene (Blancan) Zebra Equus simplicidens (Perissodactyla: Equidae) from the Hagerman Horse Quarry, Idaho in Palaeoecology and Palaeoenvironments of Late Cenozoic Mammals (eds. Stewart, K. M. & Seymour, K. L.) 134\u2013155 (University of Toronto Press, 1996).","DOI":"10.3138\/9781487574154-010"},{"key":"26817_CR52","first-page":"33","volume":"1930","author":"JW Gidley","year":"1930","unstructured":"Gidley, J. W. Continuation of the fossil horse round-up on the Old Oregon Trail. Smithsonian Explorations and Fieldwork 1930, 33\u201340 (1930).","journal-title":"Smithsonian Explorations and Fieldwork"},{"key":"26817_CR53","doi-asserted-by":"publisher","first-page":"281","DOI":"10.5479\/si.00963801.83-2985.281","volume":"83","author":"CL Gazin","year":"1936","unstructured":"Gazin, C. L. A study of the fossil horse remains from the Upper Pliocene of Idaho. Proceedings of the United States National Museum 83, 281\u2013320 (1936).","journal-title":"Proceedings of the United States National Museum"},{"key":"26817_CR54","unstructured":"Waring, G. H. Horse Behavior. The Behavioral Traits and Adaptations of Domestic and Wild Horses, Including Ponies. (Noyes Publications, 1983)."},{"key":"26817_CR55","doi-asserted-by":"publisher","first-page":"1681","DOI":"10.1111\/sed.12048","volume":"60","author":"JP Calvo","year":"2013","unstructured":"Calvo, J. P., Pozo, M., Silva, P. G. & Morales, J. Pattern of sedimentary infilling of fossil mammal traps formed in pseudokarst at Cerro de los Batallones, Madrid Basin, central Spain. Sedimentology 60, 1681\u20131708, https:\/\/doi.org\/10.1111\/sed.12048 (2013).","journal-title":"Sedimentology"},{"key":"26817_CR56","first-page":"49","volume":"1","author":"E Joubert","year":"1974","unstructured":"Joubert, E. Composition and limiting factors of a Khomas Hochland population of Hartmann zebra Equus zebra hartmannae. Madoqua 1, 49\u201353 (1974).","journal-title":"Madoqua"},{"key":"26817_CR57","doi-asserted-by":"publisher","first-page":"1403","DOI":"10.1126\/science.220.4604.1403","volume":"220","author":"J Berger","year":"1983","unstructured":"Berger, J. Ecology and catastrophic mortality in wild horses: implications for interpreting fossil assemblages. Science 220, 1403\u20131404, https:\/\/doi.org\/10.1126\/science.220.4604.1403 (1983).","journal-title":"Science"},{"key":"26817_CR58","first-page":"1","volume":"399","author":"JW Osborn","year":"1973","unstructured":"Osborn, J. W. & Crompton, A. W. The evolution of mammalian from reptilian dentitions. Breviora 399, 1\u201318 (1973).","journal-title":"Breviora"},{"key":"26817_CR59","first-page":"1","volume":"76","author":"B Kurt\u00e9n","year":"1953","unstructured":"Kurt\u00e9n, B. On the variation and population dynamics of fossil and recent mammal populations. Acta Zoologica Fennica 76, 1\u2013122 (1953).","journal-title":"Acta Zoologica Fennica"},{"key":"26817_CR60","doi-asserted-by":"publisher","first-page":"906","DOI":"10.2307\/1935638","volume":"47","author":"G Caughley","year":"1966","unstructured":"Caughley, G. Mortality patterns in mammals. Ecology 47, 906\u2013918 (1966).","journal-title":"Ecology"},{"key":"26817_CR61","doi-asserted-by":"publisher","first-page":"305","DOI":"10.1016\/0278-4165(90)90010-B","volume":"9","author":"MC Stiner","year":"1990","unstructured":"Stiner, M. C. The use of mortality patterns in archaeological studies of hominid predatory adaptations. J. Anthropol. Archaeol. 9, 305\u2013351, https:\/\/doi.org\/10.1016\/0278-4165(90)90010-B (1990).","journal-title":"J. Anthropol. Archaeol."},{"key":"26817_CR62","doi-asserted-by":"crossref","unstructured":"Lyman, R. L. Vertebrate Taphonomy (Cambridge University Press, 1994).","DOI":"10.1017\/CBO9781139878302"},{"key":"26817_CR63","doi-asserted-by":"publisher","first-page":"85","DOI":"10.12651\/JSR.2013.2.1.085","volume":"2","author":"D Lkhagvasuren","year":"2013","unstructured":"Lkhagvasuren, D. et al. Age determination of the Mongolian wild ass (Equus hemionus, 1775) by the dentition patterns and annual lines in tooth cementum. Journal of Species Research 2, 85\u201390 (2013).","journal-title":"Journal of Species Research"},{"key":"26817_CR64","doi-asserted-by":"publisher","first-page":"1","DOI":"10.2307\/3503962","volume":"157","author":"P Grubb","year":"1981","unstructured":"Grubb, P. Equus burchelli. Mammalian Species 157, 1\u20139 (1981).","journal-title":"Mammalian Species"},{"key":"26817_CR65","doi-asserted-by":"publisher","first-page":"89","DOI":"10.12651\/JSR.2013.2.1.085","volume":"25","author":"BL Penzhorn","year":"1982","unstructured":"Penzhorn, B. L. Age determination in Cape Mountain zebras Equus zebra zebra in the Mountain Zebra National Park. Koedoe 25, 89\u2013102, https:\/\/doi.org\/10.12651\/JSR.2013.2.1.085 (1982).","journal-title":"Koedoe"},{"key":"26817_CR66","first-page":"5","volume":"1","author":"E Joubert","year":"1972","unstructured":"Joubert, E. Tooth development and age determination in the Hartmann zebra Equus zebra hartmannae. Madoqua 1, 5\u201316 (1972).","journal-title":"Madoqua"},{"key":"26817_CR67","first-page":"1","volume":"179","author":"BJ MacFadden","year":"1984","unstructured":"MacFadden, B. J. Systematics and phylogeny of Hipparion, Neohipparion, Nannippus, and Cormohipparion (Mammalia, Equidae) from the Miocene and Pliocene of the New World. B. Am. Mus. Nat. Hist. 179, 1\u2013196 (1984).","journal-title":"B. Am. Mus. Nat. Hist."}],"container-title":["Scientific Reports"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41598-018-26817-3.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-018-26817-3","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-018-26817-3.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2022,12,21]],"date-time":"2022-12-21T06:10:08Z","timestamp":1671603008000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41598-018-26817-3"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2018,5,31]]},"references-count":67,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2018,12]]}},"alternative-id":["26817"],"URL":"https:\/\/doi.org\/10.1038\/s41598-018-26817-3","relation":{},"ISSN":["2045-2322"],"issn-type":[{"value":"2045-2322","type":"electronic"}],"subject":[],"published":{"date-parts":[[2018,5,31]]},"assertion":[{"value":"6 December 2017","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"16 May 2018","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"31 May 2018","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing interests.","order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing Interests"}}],"article-number":"8507"}}