FREE SHIPPING over $40
OVER 10000 FIVE-STAR REVIEWS
5% OFF ALL SUBSCRIPTIONS
100% HAPPINESS GUARANTEE
Analyze Diet
0
Biology2022; 11(9); 1258; doi: 10.3390/biology11091258

Evolution of the Family Equidae, Subfamily Equinae, in North, Central and South America, Eurasia and Africa during the Plio-Pleistocene.

Abstract: Studies of horse evolution arose during the middle of the 19th century, and several hypotheses have been proposed for their taxonomy, paleobiogeography, paleoecology and evolution. The present contribution represents a collaboration of 19 multinational experts with the goal of providing an updated summary of Pliocene and Pleistocene North, Central and South American, Eurasian and African horses. At the present time, we recognize 114 valid species across these continents, plus 4 North African species in need of further investigation. Our biochronology and biogeography sections integrate Equinae taxonomic records with their chronologic and geographic ranges recognizing regional biochronologic frameworks. The paleoecology section provides insights into paleobotany and diet utilizing both the mesowear and light microscopic methods, along with calculation of body masses. We provide a temporal sequence of maps that render paleoclimatic conditions across these continents integrated with Equinae occurrences. These records reveal a succession of extinctions of primitive lineages and the rise and diversification of more modern taxa. Two recent morphological-based cladistic analyses are presented here as competing hypotheses, with reference to molecular-based phylogenies. Our contribution represents a state-of-the art understanding of Plio-Pleistocene evolution, their biochronologic and biogeographic background and paleoecological and paleoclimatic contexts.
Get Full Text
Publication Date: 2022-08-24 PubMed ID: 36138737PubMed Central: PMC9495906DOI: 10.3390/biology11091258Google Scholar: Lookup
The Equine Research Bank provides access to a large database of publicly available scientific literature. Inclusion in the Research Bank does not imply endorsement of study methods or findings by Mad Barn.
LinkedInCopy
  • Journal Article

Summary

This research summary has been generated with artificial intelligence and may contain errors and omissions. Refer to the original study to confirm details provided. Submit correction.

The research article is an intensive study on the evolution of horses, specifically the Equinae subfamily, during the Pliocene-Pleistocene era across various continents like North, Central and South America, Eurasia and Africa. A collective team of 19 international experts aimed to present an up-to-date summary of horse species, their chronological and geographical information, paleobotanical diet and body mass data, and the correlation with paleoclimatic conditions, as well as comparing different cladistic analyses.

Scope of the Research

  • The research involves a comprehensive investigation into horse evolution, specifically from the middle of the 19th century. The focus is on the taxonomy, paleobiogeography, paleoecology, and evolution aspects.
  • This research is a collaborative effort of 19 international experts aiming to provide a thorough and updated summary of the Pliocene and Pleistocene ages’ horse species in various continents like North, Central and South America, Eurasia and Africa.

Key Findings

  • Through this research, 114 valid species across these continents are recognized, along with four species in North Africa that need a closer look.
  • The team has also categorized the horses based on chronological and geographic ranges, creating regional biochronology frameworks.
  • In terms of paleoecology, the research offers insights into their past diets and body masses, derived from mesowear and light microscopic methods. Furthermore, the research provides a guide to temporal sequences that depict how the paleoclimatic conditions affected these horses over time.
  • The findings revealed a sequential pattern of extinctions of primitive lineages and the emergence of modern taxa.

Comparative Analysis and Hypotheses

  • The article presents comparative studies between two morphological-based cladistic analyses. These are presented as competing hypotheses regarding the horse evolution process.
  • The contribution represents an in-depth understanding of the Plio-Pleistocene equine species, their chronological and geographical background, along with their paleoecological and paleoclimatic contexts.

Cite This Article

APA
(2022). Evolution of the Family Equidae, Subfamily Equinae, in North, Central and South America, Eurasia and Africa during the Plio-Pleistocene. Biology (Basel), 11(9), 1258. https://doi.org/10.3390/biology11091258

Publication

Biology
ISSN: 2079-7737
NlmUniqueID: 101587988
Country: Switzerland
Language: English
Volume: 11
Issue: 9
PII: 1258

Researcher Affiliations

Conflict of Interest Statement

The authors declare no conflict of interest.

References

This article includes 443 references
  1. Marsh C.O.. Polydactyle Horses, Recent and Extinct.. Am. J. Sci. 1879;17:499–505.
    doi: 10.2475/ajs.s3-17.102.499google scholar: lookup
  2. Gidley J.W.. Revision of the Miocene and Pliocene Equidae in North America.. Bull. Am. Mus. Nat. Hist. 1907;23:865–934.
  3. Osborn H.F.. Equidae of the Oligocene, Miocene, and Pliocene of North America. Iconographic type revision.. Mem. Am. Mus. Nat. Hist. New Ser. 1918;2:1–326.
  4. Matthew W.D.. A new link in the ancestry of the horse.. Am. Mus. Novit. 1924;131:130–131.
  5. Stirton R.A.. Phylogeny of North American Equidae.. Univ. Calif. Publ. Geol. Sci. 1940;25:165–198.
  6. Simpson G.G.. Horses: The Story of the Horse Family in the Modern World and through Sixty Million Years of History.. 1951. pp. 1–247.
  7. Macfadden BJ. Evolution. Fossil horses--evidence for evolution.. Science 2005 Mar 18;307(5716):1728-30.
    doi: 10.1126/science.1105458pubmed: 15774746google scholar: lookup
  8. Bernor R.L., Kaya F., Kaakinen A., Saarinen J., Fortelius M.. Old World hipparion evolution, biogeography, climatology and ecology.. Earth Sci. Rev. 2021;221:103748.
    doi: 10.1016/j.earscirev.2021.103784google scholar: lookup
  9. Barrón-Ortiz C.I., Avilla L.S., Jass C.N., Bravo-Cuevas V.M., Machado H., Mothé D.. What is Equus? Reconciling taxonomy and phylogenetic analyses.. Front. Ecol. Evol. 2019;7:343.
    doi: 10.3389/fevo.2019.00343google scholar: lookup
  10. Cirilli O, Pandolfi L, Rook L, Bernor RL. Evolution of Old World Equus and origin of the zebra-ass clade.. Sci Rep 2021 May 12;11(1):10156.
    doi: 10.1038/s41598-021-89440-9pmc: PMC8114910pubmed: 33980921google scholar: lookup
  11. Eisenmann V., Alberdi M.T., De Giuli C., Staesche U.. Methodology.. 1988. pp. 1–71.
  12. Bernor R.L., Tobien H., Hayek L.A.C., Mittmann H.W.. Hippotherium primigenium (Equidae, Mammalia) from the late Miocene of Höwenegg (Hegau, Germany). Andrias 1997;10:1–230.
  13. Cirilli O., Saarinen J., Pandolfi L., Rook L., Bernor R.L.. An updated review on Equus stenonis (Mammalia, Equidae): New implications for the European Early Pleistocene Equus taxonomy and paleoecology, and remarks on the Old World Equus evolution.. Quat. Sci. Rev. 2021;269:107155.
    doi: 10.1016/j.quascirev.2021.107155google scholar: lookup
  14. Bernor RL, Cirilli O, Bukhsianidze M, Lordkipanidze D, Rook L. The Dmanisi Equus: Systematics, biogeography, and paleoecology.. J Hum Evol 2021 Sep;158:103051.
    doi: 10.1016/j.jhevol.2021.103051pubmed: 34365132google scholar: lookup
  15. Fortelius M., Solounias N.. Functional characterization of ungulate molars using the abrasion-attrition wear gradient: A new method for reconstructing paleodiets.. Am. Mus. Novit. 2000;3301:1–36.
    doi: 10.1206/0003-0082(2000)301<0001:FCOUMU>2.0.CO;2google scholar: lookup
  16. Solounias N., Semprebon G.. Advances in the reconstruction of ungulate ecomorphology with application to early fossil equids.. Am. Mus. Novit. 2002;3366:1–49.
    doi: 10.1206/0003-0082(2002)366<0001:AITROU>2.0.CO;2google scholar: lookup
  17. Oksanen O., Žliobaitė I., Saarinen J., Lawing A.M., Fortelius M.. A Humboldtian approach to life and climate of the geological past: Estimating palaeotemperature from dental traits of mammalian communities.. J. Biogeogr. 2019;46:1760–1776.
    doi: 10.1111/jbi.13586google scholar: lookup
  18. Scott K.M.. Postcranial dimensions of ungulates as predictors of body mass.. 1990. pp. 301–355.
  19. Saarinen J., Cirilli O., Strani F., Meshida K., Bernor R.L.. Testing equid body mass estimate equations on modern zebras—with implications to understanding the relationship of body size, diet and habitats of Equus in the Pleistocene of Europe.. Front. Ecol. Evol. 2021;9:622412.
    doi: 10.3389/fevo.2021.622412google scholar: lookup
  20. Saarinen J., Eronen J., Fortelius M., Seppä H., Lister A.M.. Patterns of diet and body mass of large ungulates from the Pleistocene of Western Europe, and their relation to vegetation.. Palaeontol. Electron. 2016;19:1–58.
    doi: 10.26879/443google scholar: lookup
  21. Mihlbachler MC, Rivals F, Solounias N, Semprebon GM. Dietary change and evolution of horses in North America.. Science 2011 Mar 4;331(6021):1178-81.
    doi: 10.1126/science.1196166pubmed: 21385712google scholar: lookup
  22. The NOW Community. New and Old World Database of Fossil Mammals (NOW). Licensed under CC BY 4.0. 2022.. .
  23. Fortelius M., Eronen J.T., Jernvall J., Liu L., Pushkina D., Rinne J., Tesakov A., Vislobokova I.A., Zhang Z., Zhou L.. Fossil mammals resolve regional patterns ofEurasian climate change during 20 million years.. Evol. Ecol. Res. 2002;4:1005–1016.
  24. Eronen JT, Evans AR, Fortelius M, Jernvall J. The impact of regional climate on the evolution of mammals: a case study using fossil horses.. Evolution 2010 Feb 1;64(2):398-408.
    doi: 10.1111/j.1558-5646.2009.00830.xpubmed: 19735525google scholar: lookup
  25. Eronen JT, Polly PD, Fred M, Damuth J, Frank DC, Mosbrugger V, Scheidegger C, Stenseth NC, Fortelius M. Ecometrics: the traits that bind the past and present together.. Integr Zool 2010 Jun;5(2):88-101.
    doi: 10.1111/j.1749-4877.2010.00192.xpubmed: 21392327google scholar: lookup
  26. Liu L, Puolamäki K, Eronen JT, Ataabadi MM, Hernesniemi E, Fortelius M. Dental functional traits of mammals resolve productivity in terrestrial ecosystems past and present.. Proc Biol Sci 2012 Jul 22;279(1739):2793-9.
    pmc: PMC3367784pubmed: 22456884doi: 10.1098/rspb.2012.0211google scholar: lookup
  27. Orlando L, Ginolhac A, Zhang G, Froese D, Albrechtsen A, Stiller M, Schubert M, Cappellini E, Petersen B, Moltke I, Johnson PL, Fumagalli M, Vilstrup JT, Raghavan M, Korneliussen T, Malaspinas AS, Vogt J, Szklarczyk D, Kelstrup CD, Vinther J, Dolocan A, Stenderup J, Velazquez AM, Cahill J, Rasmussen M, Wang X, Min J, Zazula GD, Seguin-Orlando A, Mortensen C, Magnussen K, Thompson JF, Weinstock J, Gregersen K, Røed KH, Eisenmann V, Rubin CJ, Miller DC, Antczak DF, Bertelsen MF, Brunak S, Al-Rasheid KA, Ryder O, Andersson L, Mundy J, Krogh A, Gilbert MT, Kjær K, Sicheritz-Ponten T, Jensen LJ, Olsen JV, Hofreiter M, Nielsen R, Shapiro B, Wang J, Willerslev E. Recalibrating Equus evolution using the genome sequence of an early Middle Pleistocene horse.. Nature 2013 Jul 4;499(7456):74-8.
    doi: 10.1038/nature12323pubmed: 23803765google scholar: lookup
  28. Jónsson H, Schubert M, Seguin-Orlando A, Ginolhac A, Petersen L, Fumagalli M, Albrechtsen A, Petersen B, Korneliussen TS, Vilstrup JT, Lear T, Myka JL, Lundquist J, Miller DC, Alfarhan AH, Alquraishi SA, Al-Rasheid KA, Stagegaard J, Strauss G, Bertelsen MF, Sicheritz-Ponten T, Antczak DF, Bailey E, Nielsen R, Willerslev E, Orlando L. Speciation with gene flow in equids despite extensive chromosomal plasticity.. Proc Natl Acad Sci U S A 2014 Dec 30;111(52):18655-60.
    doi: 10.1073/pnas.1412627111pmc: PMC4284605pubmed: 25453089google scholar: lookup
  29. Heintzman PD, Zazula GD, MacPhee R, Scott E, Cahill JA, McHorse BK, Kapp JD, Stiller M, Wooller MJ, Orlando L, Southon J, Froese DG, Shapiro B. A new genus of horse from Pleistocene North America.. Elife 2017 Nov 28;6.
    doi: 10.7554/eLife.29944pmc: PMC5705217pubmed: 29182148google scholar: lookup
  30. Hulbert R.C.. Calippus and Protohippus (Mammalia, Perissodactyla, Equidae) from the Miocene (Barstovian-early Hemphillian) of the Gulf Coastal Plain.. Bull. Fla. State Mus. Biol. Sci. 1988;32:221–340.
  31. Olson E.C., McGrew P.O.. Mammalian fauna from the Pliocene of Honduras.. Bullettin Geol. Soc. Am. 1941;52:1219–1244.
    doi: 10.1130/GSAB-52-1219google scholar: lookup
  32. Drescher A.B.. Later Tertiary Equidae from the Tejon Hills, California.. Carneg. Inst. Wash. Pub. 1941;530:1–23.
  33. Cope E.D.. A new Hippidium.. Am. Nat. 1880;14:223.
  34. Matthew W.D., Stirton R.A.. Equidae from the Pliocene of Texas.. Bullettin. 1930;19:349–396.
  35. Lance J.F.. Palaeontología y estratigrafía del Pliocene de Yepómera, estado de Chihuahua 1a Parte: Equidos, excepto Neohipparion.. Inst. Geol. Univ. Nac. Autónoma México. 1950;54:83–94.
  36. Prado J.E., Alberdi M.T.. A cladistic analysis of the horses of the tribe Equini.. J. Paleontol. 1996;39:663–680.
  37. Kelly T.S.. New Middle Miocene equid crania from California and their implications for the phylogeny of the Equini.. Nat. Hist. Mus. Los Angel. Cty. Contrib. Sci. 1998;473:1–43.
    doi: 10.5962/p.226798google scholar: lookup
  38. Cope E.D.. A Preliminary Report on the Vertebrate Paleontology of the Llano Estacado.. 1893. pp. 1–136.
  39. Leidy J.. Notices of some remains of extinct Mammalia, recently discovered by Dr. F.V. Hayden, in the badlands of Nebraska.. Proc. Acad. Nat. Sci. Phila. 1856;8:59.
  40. Mooser O.. Fossil Equidae from the middle Pliocene of the Central Plateau of Mexico.. Southwest. Nat. 1968;13:1–12.
    doi: 10.2307/3668809google scholar: lookup
  41. Cope E.D.. Pliocene horses of southwestern Texas.. Am. Nat. 1885;19:1208–1209.
  42. Sondaar P.Y.. The osteology ofthe manus of fossil and Recent Equidae.. Ned. Akad. Wettenschappen. 1968;25:1–76.
  43. MacFadden B.J.. Systematics and phylogeny of Hipparion, Neohipparion, Nannippus, and Cormohipparion (Mammalia, Equidae), from the Miocene and Pliocene of the New World.. Bull. Am. Mus. Nat. Hist. 1984;179:196.
  44. MacFadden B.J.. Fossil Horses: Systematics, Paleobiology, and Evolution of the Family Equidae.. 1992.
  45. Merriam J.C.. New Species of the Hipparion Group from the Pacific Coast and Great Basin Provinces of North America.. 1915. pp. 1–8.
  46. Hulbert R.C., Harrington R.. An early Pliocene hipparionine horse from the Canadian Arctic.. Paleontology 1999;42:1017–1025.
    doi: 10.1111/1475-4983.00108google scholar: lookup
  47. MacFadden B.J., Skinner M.. Diversification and biogeography of the one-toed horses Onohippidium and Hippidion.. Yale Peabody Mus. Nat. Hist. Postilla. 1979;175:1–9.
  48. Hulbert R.C.. Late Neogene Neohipparion (Mammalia, Equidae) from the Gulf Coastal Plain of Florida and Texas.. J. Paleontol. 1987;61:809–830.
    doi: 10.1017/S0022336000029152google scholar: lookup
  49. Webb S.D., Hulbert R.C.. Systematics and evolution of Pseudhipparion (Mammalia, Equidae) from the late Neogene of the Gulf Coastal Plain and the Great Plains.. 1986. pp. 237–272.
  50. Merriam JC. CORRELATION BETWEEN THE TERTIARY OF THE GREAT BASIN AND THAT OF THE MARGINAL MARINE PROVINCE IN CALIFORNIA.. Science 1914 Oct 30;40(1035):643-5.
    doi: 10.1126/science.40.1035.643pubmed: 17789176google scholar: lookup
  51. Dalquest W.W., Donovan T.J.. A new three-toed horse (Nannippus) from the late Pliocene of Scurry County, Texas.. J. Paleont. 1973;47:34–45.
  52. Cope E.D.. A contribution to the vertebrate paleontology of Texas.. Proc. Am. Philos. Soc. 1892;30:123–131.
  53. Dalquest W.W., Schultz G.E.. Ice Age Mammals of Northwestern Texas.. 1992.
  54. Orlando M.F.. Order Perissodactyla.. 1972. pp. 117–125.
  55. Gidley J.W.. A new Pliocene horse from Idaho.. J. Mammal. 1930;16:52–60.
    doi: 10.2307/1374150google scholar: lookup
  56. Winans M.C.. Ph.D. Thesis.. 1985. Revision of North American fossil species of the genus Equus (Mammalia: Perissodactyla: Equidae) p. 264.
  57. Forsten A., Eisenmann V.. Equus (Plesippus) simplicidens (Cope), not Dolichohippus.. Mammalia. 1995;59:85–89.
    doi: 10.1515/mamm.1995.59.1.85google scholar: lookup
  58. Gazin C.L.. A study of the fossil horses remains from the Upper Pliocene of Idaho.. Proc. U.S. Natl. Mus. 1936;83:281–320.
    doi: 10.5479/si.00963801.83-2985.281google scholar: lookup
  59. Merriam J.C.. New Mammalia from the Idaho Formation.. Bull. Dep. Geol. 1918;10:523–530.
  60. Scott E.. Equus idahoensis from the Pliocene of Arizona, and Its Role in Plesippine Evolution in the American Southwest.. 2005.
  61. Downs T., Miller G.J.. Late Cenozoic equids from the Anza-Borrego Desert of California.. Nat. Hist. Mus. Los Angel. Co. Contrib. Sci. 1994;440:1–90.
    doi: 10.5962/p.208078google scholar: lookup
  62. Forsten A.. Mitochondrial-DNA time-table and ethe evolution of Equus: Comparison of molecular and paleontological evidence.. Ann. Zool. Fenn. 1992;28:301–309.
  63. Azzaroli A.. The genus Equus in Europe.. 1990. pp. 339–355.
  64. Troxell E.L.. The Vertebrate Fossils of Rock Creek, Texas.. Am. J. Sci. 1915;39:613–638.
    doi: 10.2475/ajs.s4-39.234.613google scholar: lookup
  65. Hibbard C.W.. Pleistocene vertebrates from the Upper Becerra (Becerra Superior) formation, Valley of Tequixquiac, Mexico, with notes on other Pleistocene forms.. Contrib. Mus. Paleontol. Univ. Mich. 1955;12:47–96.
  66. Eisenmann V., Howe J., Pichardo M.. Old World Hemiones and New World slender species (Mammalia, Equidae). Palaeovertebrata 2008;36:159–233.
    doi: 10.18563/pv.36.1-4.159-233google scholar: lookup
  67. Hulbert R.C.. Equus from Leisey Shell Pit 1A and other Irvingtonian localities from Florida.. Bull. Fla. Mus. Nat. Hist. 1995;37:553–602.
  68. Gidley J.W.. A new species of Pleistocene horse from the Staked Plains of Texas.. Bull. Am. Mus. Nat. Hist. 1900;13:111–116.
  69. Azzaroli A., Voorhies M.. The genus Equus in North America: The Blancan species.. Palaeontogr. Ital. 1993;80:175–198.
  70. Owen R.. On Fossil Remains of Equines from Centrai and South America.. Philos. Trans. Lond. 1869;159:559–573.
  71. Mooser O., Dalquest W.W.. Pleistocene mammals from Aguascalientes, Central Mexico.. J. Mammal. 1975;56:781–820.
    doi: 10.2307/1379653google scholar: lookup
  72. Cope E.D.. Extinct Mammalia of the Valley of Mexico.. Proc. Am. Philos. Soc. 1884;22:1–15.
  73. Gidley J.W.. Tooth characters and revision of the North American species of the genus.. Bull. Am. Mus. Nat. Hist. 1901;14:91–142.
  74. Azzaroli A.. The genus Equus in North America—The Pleistocene species.. Paleontogr. Ital. 1998;85:1–60.
  75. Hay O.P.. Description of a new species of Pleistocene horse, Equus lambei, from the Pleistocene of Yukon Territory.. Proc. U. S. Natl. Mus. 1917;53:435–443.
    doi: 10.5479/si.00963801.53-2212.435google scholar: lookup
  76. Hay O.P.. Contributions to the knowledge of the mammals of the Pleistocene of North America.. Proc. U. S. Natl. Mus. 1915;48:515–575.
    doi: 10.5479/si.00963801.48-2086.515google scholar: lookup
  77. Weinstock J, Willerslev E, Sher A, Tong W, Ho SY, Rubenstein D, Storer J, Burns J, Martin L, Bravi C, Prieto A, Froese D, Scott E, Xulong L, Cooper A. Evolution, systematics, and phylogeography of pleistocene horses in the new world: a molecular perspective.. PLoS Biol 2005 Aug;3(8):e241.
    doi: 10.1371/journal.pbio.0030241pmc: PMC1159165pubmed: 15974804google scholar: lookup
  78. Vershinina AO, Heintzman PD, Froese DG, Zazula G, Cassatt-Johnstone M, Dalén L, Der Sarkissian C, Dunn SG, Ermini L, Gamba C, Groves P, Kapp JD, Mann DH, Seguin-Orlando A, Southon J, Stiller M, Wooller MJ, Baryshnikov G, Gimranov D, Scott E, Hall E, Hewitson S, Kirillova I, Kosintsev P, Shidlovsky F, Tong HW, Tiunov MP, Vartanyan S, Orlando L, Corbett-Detig R, MacPhee RD, Shapiro B. Ancient horse genomes reveal the timing and extent of dispersals across the Bering Land Bridge.. Mol Ecol 2021 Dec;30(23):6144-6161.
    doi: 10.1111/mec.15977pubmed: 33971056google scholar: lookup
  79. Leidy J.. Description of Vertebrate Fossils.. 1860. pp. 99–122.
  80. Azzaroli A.. A Synopsis of the Quaternary species of Equus in North America.. Boll. Soc. Paleont. Ital. 1995;34:205–221.
  81. Quinn J.H.. Pleistocene Equidae of Texas.. Econ. Geol. Rep. Investig. 1957;33:1–51.
  82. von Reichenau W.. Beitrage zur naheren Kenntnis fossiler Pferde aus deutschem Pleistozan, insbesondere über die Entwicklung und die Abkaustadiem del Gebisses von Hochterrassenpferde (Equus mosbachensis v. R.). Abh. Gross. Hess. Geolog. Darmstadt. 1915;7:1–55.
  83. Alberdi M.T., Arroyo-Cabrales J., Marin-Leyva A.H., Polaco O.J.. Study of Cedral Horses and their place in the Mexican Quaternary.. Rev. Mex. Cienc. Geológicas. 2014;31:221–237.
  84. Kurten B., Anderson E.. Pleistocene Mammals of North America.. 1980. pp. 1–442.
  85. Winans M.C.. A quantitative study of the North American fossil species of the genus Equus.. Evol. Perissodactyls. 1989;72:262–297.
  86. Sher A.V.. Mlekopitaiushchie i Stratigrafia Pleistotsena Krainego Severo-Vostoka SSSR i Severnoi Ameriki.. 1971.
  87. Eisenmann V. Sussemionus, a new subgenus of Equus (Perissodactyla, Mammalia).. C R Biol 2010 Mar;333(3):235-40.
    doi: 10.1016/j.crvi.2009.12.013pubmed: 20338542google scholar: lookup
  88. Eisenmann V.. The equids from Liventsovka and other localities of the Khaprovskii Faunal Complex, Russia: A revision.. Geobios. 2022;70:17–33.
    doi: 10.1016/j.geobios.2021.11.001google scholar: lookup
  89. Leidy J.. Bones and teeth of horses from California and Oregon.. Proc. Acad. Nat. Sci. Phila. 1865;17:94.
  90. Merriam J.C.. Preliminary report on the horses of Rancho La Brea.. Bull. Dep. Geol. Sci. 1913;7:397–418.
  91. Savage D.E.. Late Cenozoic vertebrates of the San Francisco Bay Region.. Bull. Dep. Geol. Sci. 1951;28:215–314.
  92. Miller W.E.. Pleistocene vertebrates of the Los Angeles Basin and vicinity (exclusive of Rancho La Brea). Bull. Los Angel. Cty. Mus. 1971;10:1–124.
  93. Brown K.E., Akersten W.A., Scott E.. Equus occidentalis Leidy from “Asphalto”, Kern County, California.. 2015. pp. 81–89.
  94. Barrón-Ortiz CI, Rodrigues AT, Theodor JM, Kooyman BP, Yang DY, Speller CF. Cheek tooth morphology and ancient mitochondrial DNA of late Pleistocene horses from the western interior of North America: Implications for the taxonomy of North American Late Pleistocene Equus.. PLoS One 2017;12(8):e0183045.
    doi: 10.1371/journal.pone.0183045pmc: PMC5560644pubmed: 28817644google scholar: lookup
  95. Marín-Leyva A.H., Arroyo-Cabrales J., García-Zepeda M.L., Ponce-Saavedra J., Schaaf P., Pérez-Crespo V.P., Pedro Morales-Puente P., Cienfuegos-Alvarado E., Alberdi M.T.. Feeding ecology and habitat of Late Pleistocene Equus horses fromwest-central Mexico using carbon and oxygen isotopes variation.. Rev. Mex. Cienc. Geol. 2016;33:157–169.
  96. Jimenez-Hidalgo E., Diaz-Sibaja R.. Was Equus cedralensis a non-stilt legged horse? Taxonomical implication for the Mexican Pleistocene horses.. Ameghiniana. 2020;57:284–288.
    doi: 10.5710/AMGH.06.01.2020.3262google scholar: lookup
  97. Barrón-Ortiz C.. The Late Pleistocene Extinction in North America: An Investigation of Horse and Bison Fossil Material and Its Implication for Nutritional Extinction Models.. 2016.
    doi: 10.11575/PRISM/28229google scholar: lookup
  98. Prado J.L., Alberdi M.T.. Fossil Horses of South America.. 2017. pp. 1–150.
  99. Machado H., Grillo O., Scott E., Avilla L.. Following the footsteps of the South American Equus: Are autopodia taxonomically informative?. J. Mammal. Evol. 2018;25:397–405.
    doi: 10.1007/s10914-017-9389-6google scholar: lookup
  100. Machado H., Avilla L.. The Diversity of South American Equus. Did Size Really Matter?. Front. Ecol. Evol. 2019;7:235.
    doi: 10.3389/fevo.2019.00235google scholar: lookup
  101. Lund P.W.. Nouvelles Recherches sur la Faune fossile du Brésil.. Ann. Sci. Nat. 1840;13:310–319.
  102. MacFadden B.J., Siles O., Zeitler P., Johnson N.M., Campbell K.E.. Magnetic polarity stratigraphy of the middle Pleistocene (Ensenadan) Tarija Formation of southern Bolivia.. Quat. Res. 1983;19:172–187.
    doi: 10.1016/0033-5894(83)90003-0google scholar: lookup
  103. MacFadden B.J., Zeitler P.K., Anaya F., Cottle J.M.. Confirmation of the middle Pleistocene age of the sedimentary sequence from Tarija, Bolivia.. Quat. Res. 2013;79:268–273.
    doi: 10.1016/j.yqres.2012.12.009google scholar: lookup
  104. Prado J.L., Martinez-Maza C., Alberdi M.T.. Megafauna extinction in South America: A new chronology for theArgentine Pampas.. Palaeogeogr. Palaeoclim. Palaeoecol. 2015;425:41–49.
    doi: 10.1016/j.palaeo.2015.02.026google scholar: lookup
  105. Roth S.. El mamífero misterioso de la Patagonia Grypotherium domesticum. II. Descripción de los restos encontrados en la caverna de Última Esperanza.. Rev. Mus. Plata. 1899;9:421–453.
  106. Labarca R., Caro F.J., Villavicencio N.A., Capriles J.M., Briones E., Latorre C., Santoro C.M.. A Partially Complete Skeleton of Hippidion saldiasi Roth, 1899 (Mammalia: Perissodactyla) from the Late Pleistocene of the High Andes in Northern Chile.. J. Vertebr. Paleontol. 2021;40:e1862132.
    doi: 10.1080/02724634.2020.1862132google scholar: lookup
  107. Paunero R.S., Rosales G., Prado J.L., Alberdi M.T.. Cerro bombero: Registro de hippidion saldiasi roth, 1899 (equidae, perissodactyla) en el holoceno temprano de Patagonia (Santa Cruz, Argentina). Estud. Geol. 2008;64:89–98.
    doi: 10.3989/egeol.08641430google scholar: lookup
  108. Massone M., Prieto A.. Evaluacion de la modalidad cultural Fell 1 en Magallanes.. Chungara. 2004;1:303–315.
    doi: 10.4067/S0717-73562004000300033google scholar: lookup
  109. Lund P.W.. Meddelelse af det ubbytte de i 1844 undersögte Knoglehuler have avgivet til Kundskaben om Brasiliens Dyreverden för sidste Jordomvaeltning.. Ebenda. 1846;12:1–94.
  110. Montané J. Paleo-Indian remains from laguna de tagua tagua, central chile.. Science 1968 Sep 13;161(3846):1137-8.
    doi: 10.1126/science.161.3846.1137pubmed: 17812286google scholar: lookup
  111. Gervais P.. In: Recherches sur les Mammiferes Fossiles de l’Amerique Meridionale.. 1855. pp. 1–63.
  112. Shockey B.J., Salas-Gismondi R., Baby P., Guyot J.-L., Baltazar M.C., Huamán L., Clack A., Stucchi M., Pujos F., Emerson J.M.. New Pleistocene Cave Faunas of the Andes of Central Perú: Radiocarbon Ages and the Survival of Low Latitude, Pleistocene DNA.. Palaeontol. Electron. 2009;12:15.
  113. Villavicencio N.A., Werdelin L.. The casa del diablo cave (puno, peru) and the late pleistocene demise of megafauna in the andean altiplano.. Quat. Sci. Rev. 2018;195:21–31.
    doi: 10.1016/j.quascirev.2018.07.013google scholar: lookup
  114. Sun B., Deng T.. The Equus Datum and the Early Radiation of Equus in China.. Front. Ecol. Evol. 2019;7:429.
    doi: 10.3389/fevo.2019.00429google scholar: lookup
  115. Sun B., Liu W., Liu J., Liu L., Jin C.. Equus qiyngyangensis in Jinyan Cave and its paleozoographic significance.. Quat. Int. 2020;591:35–46.
    doi: 10.1016/j.quaint.2020.10.076google scholar: lookup
  116. Teilhard de Chardin P., Young C.C.. Fossil mammals from the late Cenozoic of northern China.. Palaeontol. Sin. 1931;9:1–67.
  117. Flynn L.J., Bernor R.L.. Late Tertiary mammals from the Mongolian People’s Republic.. Am. Mus. Novit. 1987;2872:1–16.
  118. Wang X., Li Q., Xie G., Saylor J.E., Tseng Z.J., Takeuchi G.T., Deng T., Wang Y., Hou S., Liu J.. Mio-Pleistocene Zanda Basin biostratigraphy and geochronology, pre-Ice Age fauna, and mammalian evolution in western Himalaya.. Palaeogeogr. Palaeoclimatol. Palaeoecol. 2013;374:81–95.
    doi: 10.1016/j.palaeo.2013.01.007google scholar: lookup
  119. Jukar A., Sun B., Bernor R.L.. The first occurrence of Plesiohipparion huangheense (Qiu, Huang & Guo, 1987) (Equidae, Hipparionini) from the late Pliocene of India.. Boll. Soc. Paleo. Ital. 2018;57:125–132.
  120. Qiu Z.X., Qiu Z.D., Deng T., Li C.K., Zhang Z.Q., Wang B.Y., Wang X.. Neogene land mammal stages/ages of China.. 2013. pp. 29–90.
  121. Matsumoto H.. On Hipparion richthofeni Koken.. Sci. Rep. Tohoku Univ. 1927;10:59–75.
  122. Qiu Z., Huang W., Guo Z.. The Chinese hipparionine fossils.. Palaeontol. Sin. New Ser. 1987;175:1–250.
  123. Bernor R.L., Lipscomb D.. The systematic position of “Plesiohipparion” aff. huangheense (Equidae, Hipparionini) from Gülyazi, Turkey.. Mitt. Bayer. Staatssamml. Paläontol. Hist. Geol. 1991;31:107–123.
  124. Bernor R.L., Sun B., Chen Y.. Plesiohipparion shanxiense n. sp. from the Early Pleistocene (Nihowanian) of E Shanxi, China.. Boll. Soc. Paleontol. Ital. 2015;54:197–210.
  125. Sefve I.. Die Hipparionen Nord-Chinas.. Palaeo. Sin. Ser. 1927;4:1–54.
  126. Liu P., Lovlie R.. Magnetostratigraphic age of Pleistocene loess/paleosol sections at Kehe, Shanxi.. J. Stratigr. 2007;31:240–246.
  127. Qiu Z., Deng T., Banyue W.. Early Pleistocene mammalian fauna from Longdan, Dongxiang, Gansu, China.. Palaeontol. Sin. New Ser. 2004;191:1–245.
  128. Teilhard de Chardin P., Piveteau J.. Les mammifères fossiles de Nihowan (Chine). Ann. Paléont. 1930;19:l–134.
  129. Kuzmina I.E.. In: Horses of North Eurasia from the Pliocene till the Present Time.. 1997. pp. 1–224.
  130. Deng T., Xue X.. Chinese Fossil Horses of Equus and Their Environment.. 1999.
  131. Chow M., Liu H.. Fossil equine teeth from Shansi.. Vert. PalAsiat. 1959;1:133–136.
  132. Ao H., An Z., Dekkers M.J., Li Y., Xiao G., Zhao H., Qiang X.. Pleistocene magnetochronology of the fauna and Paleolithic sites in the Nihewan Basin: Significance for environmental and hominin evolution in North China.. Quat. Geochronol. 2013;18:78–92.
    doi: 10.1016/j.quageo.2013.06.004google scholar: lookup
  133. Colbert E.. Pleistocene mammals from the Ma Kai Valley of northern Yunnan, China.. Am. Mus. Novit. 1940;1099:1–10.
  134. Eisenmann V.. Nouvelles interpretations des restes d’équidés (Mammalia, Perissodactyla) de Nihowan (Pléistocène inférieur de la Chine du Nord): Equus teilhardi nov. sp.. Geobios. 1975;8:125–134.
    doi: 10.1016/S0016-6995(75)80009-Xgoogle scholar: lookup
  135. Sun B., Deng T., Liu Y.. Early Pleistocene Equus (Equidae, Perissodactyla) from Andersson Loc. 32 in Qixian, Shanxi, China.. Hist. Biol. 2017;31:211–222.
    doi: 10.1080/08912963.2017.1357718google scholar: lookup
  136. Eisenmann V., Deng T.. Equus qingyangensis (Equidae, Perissodactyla) of the Upper Pliocene of Bajiazui, China: Evidence for the North American origin of an Old World lineage distinct from E. stenonis.. Quaternaire. 2005;2:113–122.
  137. Sharapov S.. Kuruksaiskii Kompleks Pozdnepliotsenovykh Mlekopitajushchikh Afgano-Adzhikistanskoi Depressii.. 1986. p. 270.
  138. Forsten A., Sharapov S.. Fossil equids (Mammalia, Equidae) from the Neogene and Pleistocene of Tadzhikistan.. Geodiversitas. 2000;22:293–314.
  139. Vangengeim E.A., Sotnikova M.V., Alekseeva L.I., Vislobokova I.A., Zhegallo V.I., Zazhigin V.S., Shevyreva N.S.. Biostratigrafiya Pozdnego Pliotsena—Rannego Pleistotsena Adzhikistana.. 1988. p. 125.
  140. Kuznetsova T.V., Zhegallo V.I.. Taxonomic Diversity of Equids from the Nalaikha Locality (Mongolia); Proceedings of the International Conference on the Conditions of the Theriofauna in Russia and Adjacent Countries; Moscow, Russia.. 1995;1–3 February. pp. 174–178.
  141. Kuznetsova T.V., Zhegallo V.I.. Equus (Hemionus) nalaikhaensis from the Pleistocene of Mongolia, the Earliest Kulan of Central Asia.. Paleontol. J. 2009;43:574–583.
    doi: 10.1134/S0031030109050141google scholar: lookup
  142. Lazarev P.A.. Anthropogenic Horses of Yakutia.. 1980. p. 190.
  143. Rusanov B.S.. Biostratigraphy of Cenoizoic Deposits of Southern Yakutia.. 1968. p. 459.
  144. Lazarev P.A.. Large mammals of Anthropogene in Yakutia.. 2008. p. 160.
  145. Spasskaya N., Pavlinov I., Sharko F., Boulygina E., Tsygankova S., Nedoluzhko A., Mashchenko E.. Morphometric and genetic analyses of diversity of the Lena horse (Equus lenensis Russanov, 1968; Mammalia: Equidae). Russ. J. Theriol. 2021;20:82–95.
    doi: 10.15298/rusjtheriol.20.1.09google scholar: lookup
  146. Liu H.Y.. A new species of Equus from Locality 21 of Zhoukoudian.. Vert PalAsiat. 1963;7:318–322.
  147. Forsten A.. Chinese fossil horses of the genus Equus.. Acta Zool. Fenn. 1986;181:1–40.
  148. Gromova V.I.. O novoj iskopaiemoj loshadi iz srednej Azii.. Dokl. Akad. Nauk SSSR. 1946;54:349–351.
  149. Eisenmann V., Helmer D., Segui M.S.. The big Equus from Geometric Kebaran of Umm el Tlel, Syria: Equus valeriani, Equus capensis or Equus caballus?. Proc. Fifth Int. Symp. Archaeozoology Southwest. Asia Adjac. Areas. 2002;1:62–73.
  150. Zhou X., Sun Y., Xu Q., Li Y.. Note on a new Late Pleistocene Equus from Dalian.. Vertebr. Palasiat. 1985;23:69–76.
  151. Yuan JX, Hou XD, Barlow A, Preick M, Taron UH, Alberti F, Basler N, Deng T, Lai XL, Hofreiter M, Sheng GL. Molecular identification of late and terminal Pleistocene Equus ovodovi from northeastern China.. PLoS One 2019;14(5):e0216883.
    doi: 10.1371/journal.pone.0216883pmc: PMC6522033pubmed: 31095634google scholar: lookup
  152. Eisenmann V., Vasiliev S.. Unexpected finding of a new Equus species (Mammalia, Perissodactyla) belonging to a supposedly extinct subgenus in Late Pleistocene deposits of Khakassia (southwestern Siberia). Geodiversitas. 2011;33:519–530.
    doi: 10.5252/g2011n3a5google scholar: lookup
  153. Orlando L, Metcalf JL, Alberdi MT, Telles-Antunes M, Bonjean D, Otte M, Martin F, Eisenmann V, Mashkour M, Morello F, Prado JL, Salas-Gismondi R, Shockey BJ, Wrinn PJ, Vasil'ev SK, Ovodov ND, Cherry MI, Hopwood B, Male D, Austin JJ, Hänni C, Cooper A. Revising the recent evolutionary history of equids using ancient DNA.. Proc Natl Acad Sci U S A 2009 Dec 22;106(51):21754-9.
    doi: 10.1073/pnas.0903672106pmc: PMC2799835pubmed: 20007379google scholar: lookup
  154. Yuan J., Sheng G., Preick M., Sun B., Hou X., Chen S., Taron U.H., Barlow A., Wang L., Hu J.. Mitochondrial genomes of Late Pleistocene caballines horses from China belong to a separate clade.. Quat. Sci. Rev. 2020;250:106691.
    doi: 10.1016/j.quascirev.2020.106691google scholar: lookup
  155. Plasteeva N., Vasiliev S., Kosintsev P.. Equus (Sussemionus) ovodovi Eisenmann et Vasiliev, 2011 from the Laye Pleistocene of Western Siberia.. Russ. J. Theriol. 2015;14:187–200.
    doi: 10.15298/rusjtheriol.14.2.07google scholar: lookup
  156. Pallas P.S.. Equus hemionus.. Novi. Comment. Acad. Sci. Imp. Petropol. 1774;19:394–417.
  157. Biswas S.K.. Regional Tectonic Framework, Structure and Evolution of the Western Marginal Basins of India.. Tectonophysics. 1987;135:302–327.
  158. Costa A.G.. A new Late Pleistocene fauna from arid coastal India: Implications for inundated coastal refugia and human dispersals.. Quat. Int. 2017;436:253–269.
    doi: 10.1016/j.quaint.2015.07.002google scholar: lookup
  159. Dassarma D.C., Biswas S.. Newly discovered Late Quaternary vertebrates from the terraced alluvial fills of the Son Valley.. Indian J. Earth Sci. 1977;4:122–136.
  160. Turvey S.T., Sathe V., Crees J.J., Jukar A.M., Chakraborty P., Lister A.M.. Late Quaternary megafaunal extinctions in India: How much do we know?. Quat. Sci. Rev. 2021;252:106740.
    doi: 10.1016/j.quascirev.2020.106740google scholar: lookup
  161. Stewart M.J., Louys G.J., Price N.A., Drake H.S., Petraglia M.D.. Middle and Late Pleistocene mammal fossils of Arabia and surrounding regions: Implications for biogeography and hominin dispersals.. Quat. Int. 2019;515:12–29.
    doi: 10.1016/j.quaint.2017.11.052google scholar: lookup
  162. Moorcroft W.. Travels in the Provinces of Hindustan and the Panjab.. 1841. p. 312.
  163. Shah N.. Status and action plan for the kiang (Equus kiang). 2002. pp. 72–81.
  164. Wang Y.X.. A Complete Check-List of Mammal Species and Subspecies in China: A Taxonomic and Geographic Reference.. 2002.
  165. Schaller G.B.. Wildlife of the Tibetan Steppe.. 1998.
  166. Poliakov I.S.. Izvestija Russkogo Geograficheskogo Obshchestva.. 1881. pp. 1–19.
  167. Groves C.. Morphology, habitat, and taxonomy.. 1994. pp. 39–59.
  168. Groves C., Grubb P.. Ungulate Taxonomy.. 2011.
  169. Heptner V., Nasimovich A.A., Bannikov A.G.. Artiodactyla and Perissodactyla.. 1961.
  170. Deng T.. Ages of some Late Pleistocene faunas based on the presence of Equus przewalskii (Perissodactyla, Equidae). Vertebr. Palasiat. 1999;23:51–56.
  171. Gaunitz C, Fages A, Hanghøj K, Albrechtsen A, Khan N, Schubert M, Seguin-Orlando A, Owens IJ, Felkel S, Bignon-Lau O, de Barros Damgaard P, Mittnik A, Mohaseb AF, Davoudi H, Alquraishi S, Alfarhan AH, Al-Rasheid KAS, Crubézy E, Benecke N, Olsen S, Brown D, Anthony D, Massy K, Pitulko V, Kasparov A, Brem G, Hofreiter M, Mukhtarova G, Baimukhanov N, Lõugas L, Onar V, Stockhammer PW, Krause J, Boldgiv B, Undrakhbold S, Erdenebaatar D, Lepetz S, Mashkour M, Ludwig A, Wallner B, Merz V, Merz I, Zaibert V, Willerslev E, Librado P, Outram AK, Orlando L. Ancient genomes revisit the ancestry of domestic and Przewalski's horses.. Science 2018 Apr 6;360(6384):111-114.
    doi: 10.1126/science.aao3297pubmed: 29472442google scholar: lookup
  172. Librado P, Orlando L. Genomics and the Evolutionary History of Equids.. Annu Rev Anim Biosci 2021 Feb 16;9:81-101.
    doi: 10.1146/annurev-animal-061220-023118pubmed: 33197207google scholar: lookup
  173. Taylor WTT, Barrón-Ortiz CI. Rethinking the evidence for early horse domestication at Botai.. Sci Rep 2021 Apr 2;11(1):7440.
    doi: 10.1038/s41598-021-86832-9pmc: PMC8018961pubmed: 33811228google scholar: lookup
  174. Wolf D., Bernor R.L., Hussain S.T.. A systematic, biostratigraphic, and paleobiogeographic reevaluation of the Siwalik hipparionine horse assemblage from the Potwar Plateau, Northern Pakistan.. Palaeontogr. Abt. 2013;300:1–115.
    doi: 10.1127/pala/300/2013/1google scholar: lookup
  175. Jukar A.M., Sun B., Nanda A.C., Bernor R.L.. The first occurrence of Eurygnathohippus (Mammalia, Perissodatyla, Equidae) outside Africa and its biogeographic significance.. Boll. Soc. Paleo. Ital. 2019;58:171–179.
  176. Rook L., Bernor R.L., Avilla L.D., Cirilli O., Flynn L., Jukar A.M., Sanders W., Scott E., Wang X.. Mammal Biochronology (Land Mammal Ages) around the World from the Late Miocene to Middle Pleistocene and Major Events in Horse Evolutionary History.. Front. Ecol. Evol. 2019;7:278.
    doi: 10.3389/fevo.2019.00278google scholar: lookup
  177. Jukar A.M., Lyons S.K., Wagner P.J., Uhen M.D.. Late Quaternary extinctions in the Indian Subcontinent.. Paleogeol. Paleoclim. Paleoecol. 2021;562:110137.
    doi: 10.1016/j.palaeo.2020.110137google scholar: lookup
  178. Thomas P.K., Joglekar P.. Holocene Faunal Studies.. Man Environ. 1994;19:179–203.
    doi: 10.2307/2840291google scholar: lookup
  179. Falconer H., Cautley P.T.. Fauna Antiqua Sivalensis, Being the Fossil Zoology of the Sewalik Hills in the North of India.. 1849.
  180. Hussain S.T.. Revision of Hipparion (Equidae, Mammalia) from the Siwalik Hills of Pakistan and India.. Geol. Mijnb. 1971;147:80–81.
  181. MacFadden B.J., Woodburne M.O.. Systematics of the Neogene Siwalik Hipparions (Mammalia, Equidae) based on cranial and dental morphology.. J. Vert. Paleo. 1982;2:185–218.
    doi: 10.1080/02724634.1982.10011930google scholar: lookup
  182. Bernor R.L., Hussain S.T.. An Assessment of the Systematic, Phylogenetic and Biogeographic Relationships of Siwalik Hipparionine horses.. J. Vert. Paleo. 1985;5:32–87.
    doi: 10.1080/02724634.1985.10011845google scholar: lookup
  183. Lydekker R.. Catalogue of the Fossil Mammalia in the British Museum (Natural History) Part III.. 1886.
  184. Bernor R.L., Cirilli O., Jukar A.M., Potts R., Buskianidze M., Rook L.. Evolution of Early Equus in Italy, Georgia, the Indian Subcontinent, East Africa, and the Origins of African Zebras.. Front. Ecol. Evol. 2019;7:166.
    doi: 10.3389/fevo.2019.00166google scholar: lookup
  185. Patel R., Nainwal H.C., Sharma T., Rana R.S.. Equus cf. sivalensis from the Tatrot Formation (Upper Pliocene) of Jhil-Bankabara Area Sirmaur District, Himachal Pradesh, India.. 2017. pp. 51–53.
  186. Moigne A.M., Dambricourt Malassé A., Singh M., Kaur A., Gaillard C., Karir B., Pal S., Bhardwaj V., Abdessadok S., Chapon Sao C.. The faunal assemblage of the paleonto-archeological localities of the Late Pliocene Quranwala Zone, Mosol Formation, Siwalik Range, NW India.. Palevol. 2016;15:359–378.
    doi: 10.1016/j.crpv.2015.09.016google scholar: lookup
  187. Gaur R., Chopra S.R.K.. Taphonomy, fauna, environment and ecology of Upper Sivaliks (Plio-Pleistocene) near Chandigarh, India.. Nature. 1984;308:353–355.
    doi: 10.1038/308353a0google scholar: lookup
  188. Hussain S.T., van den Bergh G., Steensma K.J., de Visser J.A., de Vos J., Mohammad A., van Dam J., Sondaar P., Malik S.B.. Biostratigraphy of the Plio-Pleistocene continental sediments (Upper Siwaliks) of the Mangla-Samwal anticline, Azad Kashmir.. Proc. K. Ned. Akad. Wet. 1992;95:65–80.
  189. Dennell R., Coard R., Turner A.. The biostratigraphy and magnetic polarity zonation of the Pabbi Hills, northern Pakistan: An Upper Siwalik (Pinjor Stage) Upper Pliocene–Lower Pleistocene fluvial sequence.. Palaeogeogr. Palaeoclimatol. Palaeoecol. 2006;234:168–185.
    doi: 10.1016/j.palaeo.2005.10.008google scholar: lookup
  190. Sonakia A., Biswas S.. Fossil mammals including early man from the Quaternary deposits of the Narmada and Son basins of Madhya Pradesh, India.. Palaeontol. Indica. 2011;53:1–84.
  191. Lydekker R.. Siwalik and Narbada Equidae.. Mem. Geol. Soc. India Palaeontol. Indica Ser. 1882;2:67–98.
  192. Cirilli O., Bernor R.L., Rook L.. New insights on the Early Pleistocene equids from Roca-Neyra (France, central Europe): Implications for the Hipparion LAD and the Equus FAD in Europe.. J. Paleontol. 2021;95:406–425.
    doi: 10.1017/jpa.2020.99google scholar: lookup
  193. Gromova V.. Le Genre Hipparion.. 1952. pp. 1–288.
  194. Bernor R.L., Sen S.. The Early Pliocene Plesiohipparion and Proboscidipparion (Equidae, Hipparionini) from Çalta, Turkey (Ruscinian Age, c. 4.0 Ma). Geodiversitas. 2017;39:285–314.
    doi: 10.5252/g2017n2a7google scholar: lookup
  195. Bernor R.L., Beech M., Bibi F.. Equidae from the Baynunah Formation. 2022. pp. 259–280.
  196. Crusafont P.M., Sondaar P.. Una nouvelle espece d’Hipparion du Pliocéne terminal d’Espagne.. Palaeovertebrata. 1971;4:59–66.
    doi: 10.18563/pv.4.2.59-66google scholar: lookup
  197. Alberdi M.T., Alcalà L.. A study of the new samples of the Pliocene Hipparion (Equidae, Mammalia) from Spain and Bulgaria.. Trans. R. Soc. Edinb. Earth Sci. 1999;89:167–186.
    doi: 10.1017/S0263593300007112google scholar: lookup
  198. Piñero P., Agustí J., Oms O., Fierro I., Montoya P., Mansino S., Ruiz-Sánchez F., Alba D.M., Alberdi M.T., Blain H.A.. Early Pliocene continental vertebrate Fauna at Puerto de la Cadena (SE Spain) and its bearing on the marine-continental correlation of the Late Neogene of Eastern Betics.. Paleogeogr. Palaeoclim. Palaeoecol. 2017;479:102–114.
    doi: 10.1016/j.palaeo.2017.04.020google scholar: lookup
  199. Gervais P.. Zoologie et Paléontologie Française (Animaux Vertébrés): Deuxième Edition.. 1859. pp. 1–544.
  200. Depéret C.. Les animaux pliocènes du Roussilon.. Mem. Soc. Geol. Fr. 1890;3:1–198.
  201. Forsten A.. The hipparions (Mammalia, Equidae) of Suffolk, England.. Trans. R. Soc. Edinb. Earth Sci. 2001;92:115–120.
    doi: 10.1017/S0263593300000080google scholar: lookup
  202. Forsten A.. Latest Hipparion Cristol, 1832 in Europe. A review of the Pliocene Hipparion crassum Gervais Group and other finds (Mammalia, Equidae). Geodiversitas. 2002;24:465–486.
  203. Eisenmann V., Sondaar P.. Pliocene vertebrate locality of Calta, Ankara, Turkey. Hipparion.. Geodiversitas. 1998;20:409–439.
  204. Hernández-Pacheco E.. La llanura manchega y sus mamíferos fósiles: Yacimiento de La Puebla de Almoradier.. Com. Investig. Pleontol. Mem. 1921;28:1–41.
  205. Alberdi M.T.. El género Hipparion en España, revision e historia evolutive.. Trab. Sobre Neógeno Cuatern. 1974;1:1–139.
  206. Bernor R.L., Koufos G.D., Woodburne M.O., Fortelius M.. The evolutionary history and biochronology of European and southwestern Asian late Miocene and Pliocene hipparionine horses.. 1996. pp. 307–338.
  207. Bernor R.L., Sun B.Y.. Morphology through ontogeny of Chinese Proboscidipparion and Plesiohipparion and observations on their Eurasian and African relatives.. Vertebr. Palasiat. 2015;53:73–92.
  208. Agustì J., Oms O.. On the age of the last hipparionine faunas in western Europe.. Comptes Rendus Acad. Sci. Paris Sci. 2001;332:291–297.
    doi: 10.1016/S1251-8050(01)01523-3google scholar: lookup
  209. Azanza B., Morales M., Andrés M., Cerdeño E., Alberdi M.T.. Los macromamíferos y la datación del yacimiento clásico de Villarroya.. 2016. pp. 267–280.
  210. Athanassiou A.. A Villafranchian Hipparion-bearing mammal fauna from Sésklo (E. Thessaly, Greece): Implications for the question of Hipparion-Equus sympatry in Europe.. Quaternary. 2018;12:12.
    doi: 10.3390/quat1020012google scholar: lookup
  211. Rook L., Cirilli O., Bernor R.L.. A late occurring “Hipparion” from the middle Villafranchian of Montopoli, Italy (Early Pleistocene; MN16b; ca. 2.5 Ma). Boll. Soc. Paleontol. Ital. 2017;56:333–339.
  212. Bajgusheava V.S.. Krupnaja Loshad Khaprovskogo Kompleksa is alluvija Severo Vostochnoto Priazovija.. Invesija Servo Kavk. Nauchnogo Zentra Vychei Shkoly. 1978;1:98–102.
  213. Gromova V.I.. Istorija loshadej (roda Equus) v Starom Svete. Chast’ 1. Obzor i opisanie form.. Tr. Paleontol. Inst. 1949;17:1–373.
  214. Forsten A.. The fossil horses (Equidae, Mammalia) from the Plio–Pleistocene of Liventsovka near Rostov–Don, Russia.. Geobios. 1997;31:645–657.
    doi: 10.1016/S0016-6995(98)80052-1google scholar: lookup
  215. Alberdi M.T., Cerdeño E., López–Martínez N., Morales J., Soria M.D.. La Fauna Villafranquiense de El Rincon–1 (Albacete, Castilla—La Mancha). Estud. Geol. 1997;53:69–93.
    doi: 10.3989/egeol.97531-2248google scholar: lookup
  216. Alberdi M.T., Ortiz–Jaureguizar E., Prado J.L.. A quantitative review of European stenonoid horses.. J. Paleontol. 1998;72:371–387.
    doi: 10.1017/S0022336000036350google scholar: lookup
  217. Azzaroli A.. On Equus livenzovensis Baigusheva 1978 and the “stenonid” lineage of Equids.. Paleontogr. Ital. 2000;87:1–17.
  218. Bernor R.L., Cirilli O., Wang S.Q., Rook L.. Equus cf. livenzovenzis from Montopoli, Italy (early Pleistocene; MN16b; ca. 2.6 Ma). Boll. Soc. Paleontol. Ital. 2018;57:203–216.
  219. Cherin M., Cirilli O., Azzarà B., Bernor R.L.. Equus stenonis (Equidae, Mammalia) from the Early Pleistocene of Pantalla (Italy) and the dispersion of stenonines horses in Europe.. Boll. Soc. Paleontol. Ital. 2021;60:1–18.
  220. Terhune C., Curran S., Croitor R., Drăguşin V., Gaudin T., Petrulescu A., Robinson C., Robu M., Werdelin L.. Early Pleistocene fauna of the Olteţ Valley of Romania: Biochronological and biogeographical implications.. Quat. Int. 2020;553:14–33.
    doi: 10.1016/j.quaint.2020.06.020google scholar: lookup
  221. Delafond F., Depéret C.. Les Terrains Tertiaires de la Bresse et Leurs Gites de Lignites et de Minerais de Fer.. 1893. pp. 1–332.
  222. Pomel A.. In: Catalogue Méthodique et Descriptif des Vertebres Fossiles Découverts dans le Bassin Hydrographique Supérieur de la Loire et Surtout Dans la Vallée de Son Affluent Principal.. 1853. pp. 1–123.
  223. Boule M.. Bulletin de la Société Géologique de France.. 1891;18:801.
  224. Viret J.. Le loess à bancs durcis de Saint Vallier (Drome) et sa faune de mammifères villagranchiens.. Nouv. Arch. Mus. D’histoire Nat. Lyon. 1954;4:1–200.
  225. Samson P.. Les équidés fossiles de Roumanie (Pliocène moyen—Pléistocène supérieur). Geol. Romana. 1975;14:165–354.
  226. Cocchi I.. L’uomo fossile dell’Italia Centrale.. Mem. Della Soc. Ital. Sci. Nat. 1867;2:1–180.
  227. Prat F.. Les equidés villafranchiens en France, genre Equus.. Cah. Quat. 1980;2:1–290.
  228. Palombo M.R., Alberdi M.T., Bellucci L., Sardella R.. An intriguing middle–sized horse from Coste San Giacomo (Anagni Basin, central Italy). Quat. Res. 2017;87:347–362.
    doi: 10.1017/qua.2017.6google scholar: lookup
  229. Cirilli O.. Equus stehlini Azzaroli, 1964 (Perissodactyla, Equidae). A revision of the most enigmatic horse from the Early Pleistocene of Europe, with new insights on the evolutionary history of European medium- and small-sized horses.. Riv. Paleont. Strat. 2022;128:241–265.
    doi: 10.54103/2039-4942/15744google scholar: lookup
  230. Azzaroli A.. The two Villafranchian Horses of the Upper Valdarno.. Palaeontogr. Ital. 1964;59:1–12.
  231. Alberdi M.T., Palombo M.R.. The late Early to early Middle Pleistocene stenonoid horses from Italy.. Quat. Int. 2013;288:25–44.
    doi: 10.1016/j.quaint.2011.12.005google scholar: lookup
  232. Palombo M.R., Alberdi M.T.. Light and shadows in the evolution of South European stenonoid horses.. Foss. Impr. 2017;73:115–140.
    doi: 10.2478/if-2017-0006google scholar: lookup
  233. Rook L., Martínez–Navarro B.. Villafranchian: The long story of a Plio–Pleistocene European large mammal biochronologic unit.. Quat. Int. 2010;219:134–144.
    doi: 10.1016/j.quaint.2010.01.007google scholar: lookup
  234. Bartolini-Lucenti S, Cirilli O, Pandolfi L, Bernor RL, Bukhsianidze M, Carotenuto F, Lordkipanidze D, Tsikaridze N, Rook L. Zoogeographic significance of Dmanisi large mammal assemblage.. J Hum Evol 2022 Feb;163:103125.
    doi: 10.1016/j.jhevol.2021.103125pubmed: 34954399google scholar: lookup
  235. Vekua A.K.. The Lower Pleistocene Mammalian Fauna of Akhalkalaki.. 1962. pp. 1–191.
  236. Koufos G.D.. Early Pleistocene equids from Mygdonia basin (Macedonia, Greece). Palaeontogr. Ital. 1992;79:167–199.
  237. Eisenmann V.. Equus granatensis of Venta Micena and evidence for primitive non–stenonid horses in the Lower Pleistocene.. 1995. pp. 175–189.
  238. Koufos G.D., Vlachou T., Gkeme A.. The Fossil Record of Equids (Mammalia: Perissodactyla: Equidae) in Greece.. 2021. pp. 351–401.
  239. Guerrero-Alba S., Palmqvist P.. Estudio morfometrico del caballo de Venta Micena (Orce, Granada) y su comparacion con los equidos modernos y del Plio- Pleistoceno en el viejo mundo.. Paleontol. Evol. 1997;30:93–148.
  240. Belmaker M.. Early Pleistocene faunal connections between Africa and Eurasia: An ecological perspective.. 2010. pp. 183–205.
  241. Wüst E.. Untersuchungen über das Pliozan und das Alteste Pleistozan Thüringens, nordlich vom Thüringer Walde und westlich von der Saale.. Abh. Nat. Ges. Halle. 1900;23:1–352.
  242. Vekua A.K.. The Lower Pleistocene mammalian fauna of Akhalkalaki (Southern Georgia, USSR). Palaeontogr. Ital. 1986;74:63–96.
  243. Musil R.. Die Pferdefunde der lokalität Stránská Skála.. Anthropos. 1972;20:185–192.
  244. Boulbes N., Van Asperen E.N.. Biostratigraphy and palaeoecology of European Equus.. Front. Ecol. Evol. 2019;7:301.
    doi: 10.3389/fevo.2019.00301google scholar: lookup
  245. Alberdi M.T.. Estudio de los caballos de los yacimientos de Fuente Nueva–3 y Barranco León–5 (Granada). 2010. pp. 291–306.
  246. Koufos G.D., Kostopoulos D.S., Sylvestrou I.A.. Equus apolloniensis n. sp. (Mammalia, Equidae) from the latest Villafranchian locality of Apollonia, macedonia, Greece.. Paleontol. Evol. 1997;30:49–76.
  247. Gkeme A.G., Koufos G.D., Kostopoulos D.S.. Reconsidering the Equids from the Early Pleistocene Fauna of Apollonia 1 (Mygdonia Basin, Greece). Quaternary. 2021;4:12.
    doi: 10.3390/quat4020012google scholar: lookup
  248. Eisenmann V., Boulbes N.. New results on equids from the Early Pleistocene site of Untermassfeld.. 2020. pp. 1295–1321.
  249. Musil R.. Die Equiden–Reste aus dem Unterpleistozän von Untermassfeld.. 2001. pp. 557–587.
  250. Forsten A.. A review of Equus stenonis Cocchi (Perissodactyla, Equidae) and related forms.. Quat. Sci. Rev. 2000;18:1373–1408.
    doi: 10.1016/S0277-3791(98)00073-0google scholar: lookup
  251. Lister A.M., Parfitt S.A., Owen F.J., Collinge S.E., Breda M.. Metric analysis of ungulate mammals in the early Middle Pleistocene of Britain, in relation to taxonomy and biostratigraphy: II: Cervidae, Equidae and Suidae.. Quat. Int. 2010;228:157–179.
    doi: 10.1016/j.quaint.2010.05.014google scholar: lookup
  252. Tsoukala E.. Contribution to the Study of the Pleistocene Fauna of Large Mammals (Carnivora, Perissodactyla, Artiodactyla) from Petralona Cave Chalkidiki (N. Greece). 1989.
  253. Azzaroli A.. On a Late Pleistocene ass from Tuscany with notes on thehistory of ass.. Palaeontol. Ital. 1979;71:27–47.
  254. Azzaroli A.. Pleistocene and living Horses of the old World.. Palaeontogr. Ital. 1966;61:1–15.
  255. Caloi L.. Il genere Equus nell’Italia centrale.. Studi Geol. Camerti. 1995:469–486.
  256. Eisenmann V.. Pliocene and Pleistocene Equids: Palaeontology versus molecular biology.. Cour. Forsch. Senckenberg. 2006;256:71–89.
  257. Regalia E.. Sull’Equus (Asinus) hydruntinus Regalia della grotta di Romanelli (Castro, Lecce). Arch. Antropol. Etnol. 1907;37:375–390.
  258. Vishnyatsky L.B.. The Palaeolithic of Central Asia.. J. World Prehistory. 1999;13:69–122.
    doi: 10.1023/A:1022538427684google scholar: lookup
  259. Musil R.. Die Equiden–Reste aus dem Pleistozän von Süssenborn bei Weimar.. Paläontologische Abh. Paläozool. 1969;3:617–666.
  260. Azzaroli A.. Ascent and decline of monodactyl equids: A case for prehistoric overkill.. Ann. Zool. Fenn. 1992;28:151–163.
  261. Bennett EA, Champlot S, Peters J, Arbuckle BS, Guimaraes S, Pruvost M, Bar-David S, Davis SJM, Gautier M, Kaczensky P, Kuehn R, Mashkour M, Morales-Muñiz A, Pucher E, Tournepiche JF, Uerpmann HP, Bălăşescu A, Germonpré M, Gündem CY, Hemami MR, Moullé PE, Ötzan A, Uerpmann M, Walzer C, Grange T, Geigl EM. Taming the late Quaternary phylogeography of the Eurasiatic wild ass through ancient and modern DNA.. PLoS One 2017;12(4):e0174216.
    pmc: PMC5396879pubmed: 28422966doi: 10.1371/journal.pone.0174216google scholar: lookup
  262. Aurell-Garrido J., Madurell-Malapeira J., Alba D.M., Moyà-Solà S.. The stenonian and caballoid equids from the Pleistocene section of Vallparadís (Terrassa, Catalonia, Spain). Cidaris. 2010;30:61–66.
  263. Madurell-Malapeira J., Minwer-Barakat R., Alba D.M., Garcés M., Gómez M., Aurell-Garrido J., Ros-Montoya S., Moya-Sola S., Berastegui X.. The Vallparadís section (Terrassa, Iberian peninsula) and the latest Villafranchian faunas of Europe.. Quat. Sci. Rev. 2010;29:2972–2982.
    doi: 10.1016/j.quascirev.2010.09.020google scholar: lookup
  264. Boddaert P.. Elenchus Animalium, Volumen 1: Sistens Quadrupedia Huc Usque Nota, Erorumque Varietates.. 1785. p. 174.
  265. Eisenmann V., Kuznetsova T.V.. Early Pleistocene equids (Mammalia, Perissodactyla) of Nalaikha (Mongolia) and the emergence of modern Equus Linnaeus, 1758.. Geodiversitas. 2004;26:535–561.
  266. van Asperen E.N.. Implications of age variation and sexual dimorphism in modern equids forMiddle Pleistocene equid taxonomy.. Int. J. Osteoarchaeol. 2013;23:1–12.
    doi: 10.1002/oa.1231google scholar: lookup
  267. van Asperen E.N.. Ecomorphological adaptations to climate and substrate in late middle Pleistocene caballoid horses.. Palaeogeogr. Palaeoclimatol. Palaeoecol. 2010;297:584–596.
    doi: 10.1016/j.palaeo.2010.09.007google scholar: lookup
  268. Churcher C.S., Richardson M.L.. Equidae.. 1978. pp. 379–422.
  269. Bernor R.L., Armour-Chelu M., Gilbert H., Kaiser T., Schulz E.. Equidae.. 2010. pp. 685–721.
  270. Duval M., Sahnouni M., Parés J.P., van der Made J., Abdessadok S., Harichane Z., Cheheb R.C., Boulaghraif K., Pérez-Gonzalez A.. The Plio-Pleistocene sequesnce of Oued Boucherti (Algeria): A unique chronologically-constrained archeological and paleontological record in North Africa.. Quat. Sci. Rev. 2021;271:107116.
    doi: 10.1016/j.quascirev.2021.107116google scholar: lookup
  271. Sahnouni M, Parés JM, Duval M, Cáceres I, Harichane Z, van der Made J, Pérez-González A, Abdessadok S, Kandi N, Derradji A, Medig M, Boulaghraif K, Semaw S. 1.9-million- and 2.4-million-year-old artifacts and stone tool-cutmarked bones from Ain Boucherit, Algeria.. Science 2018 Dec 14;362(6420):1297-1301.
    doi: 10.1126/science.aabpubmed: 30498166google scholar: lookup
  272. Villalta J.F., Crusafont M.. Les gisements de mammifères du Néogène espagnol.. Compt. Rendus Soc. Geol. Fr. 1957;1:9–10.
  273. Cirilli O., Zouhri S., Boughabi S., Benvenuti M., Bernor R.L., Papini M., Rook L.. The hipparionine horses (Perissodactyla: Mammalia) from the Late Miocene of Tizi N’Tadderht (southern Ouarzazate basin; Central High Atlas; Morocco). Riv. Paleontol. Strat. 2020;126:1–12.
  274. Bernor R.L., Boaz N.T., Cirilli O., El-Shawaihdi M., Rook L.. Sahabi Eurygnathohippus feibeli; its systematic, stratigraphic, chronologic and biogeographic contexts.. Riv. Paleont. Strat. 2020;126:561–581.
  275. Bernor R.L., Harris J.. Systematics and Evolutionary Biology of the Late Miocene and Early Pliocene Hipparionine Horses from Lothagam, Kenya.. 2003. pp. 387–438.
  276. Bernor R.L., Scott R.S.. New Interpretations of the Systematics, Biogeography and Paleoecology of the Sahabi Hipparions (latest Miocene), Libya.. Geodiversitas. 2003;25:297–319.
  277. Pomel A.. Les equidés: Carte géologie de l’Algérie.. Monogr. Paléontol. 1897;12:1–44.
  278. Hooijer D.A., Maglio V.J.. The earliest Hipparion south of the Sahara, in the late Miocene of Kenya.. Proc. K. Ned. Akad. Wet. 1973;76:311–315.
  279. Sun B., Zhang X., Liu Y., Bernor R.L.. Sivalhippus ptychodus and Sivalhippus platyodus (Perissodactyla, Mammalia) from the late Miocene of China.. Riv. Ital. Paleo. Strat. 2018;124:1–22.
  280. Bernor R.L., Kaiser T.. Systematics and Paleoecology of the Earliest Pliocene Equid, Eurygnathohippus hooijeri n. sp. from Langebaanweg, South Africa.. Mitt. Aus Hamburischen Zool. Mus. Inst. 2006;103:147–183.
  281. Hooijer D.A.. The late Pliocene Equidae of Langebaanweg, Cape Province, South Africa.. Zool. Verh. 1976;148:3–39.
  282. Bernor R.L., Gilbert H., Semprebon G., Simpson S., Semaw S.. Eurygnathohippus woldegabrieli sp. nov. (Perissodactyla: Mammalia) from the Middle Pliocene of Aramis, Ethiopia (4.4 Ma.). J. Vert. Paleo. 2013;33:1472–1485.
    doi: 10.1080/02724634.2013.829741google scholar: lookup
  283. Eisenmann V.. Le protostylide: Valeur systematique et signification phylétique chez les espèces actuelles et fossiles du genre Equus (Perissodactyla, Mammalia). Z. Saugetierkd. 1976;41:349–365.
  284. Eisenmann V.. Family Equidae. The Fossil Ungulates: Proboscidea, Perissodactyla, and Suidae.. 1983. pp. 156–214.
  285. Bernor R.L., Scott R.S., Haile-Selassie Y.. A Contribution to the Evolutionary History of Ethiopian Hipparionine Horses: Morphometric Evidence from the Postcranial Skeleton.. Geodiversitas. 2005;27:133–158.
  286. Eisenmann V., Geraads D.. 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.. Palaeo Afr. 2006;42:51–98.
  287. van Hoepen E.C.N.. Fossiele Pferde van Cornelia.. 1930;2:13–24.
  288. Leakey L.S.B.. Olduvai Gorge 1951–1961.. 1965.
  289. Armour-Chelu M., Bernor R.L., Mittmann H.-W.. Hooijer’s hypodigm for “Hipparion” cf. ethiopicum (Equidae, Hipparioninae) from Olduvai, Tanzania and comparative material from the East African Plio-Pleistocene.. Abh. Bayer. Akad. Wiss. 2006;30:15–24.
  290. Armour-Chelu M., Bernor R.L.. Equidae.. 2011. pp. 295–326.
  291. van der Made J., Boulaghraif K., Chelli Cheheb R., Caceres I., Harichane Z., Sahnouni M.. The last North African hipparions —Hipparion decline and extinction follows a common pattern.. N. Jb. Geol. Palaont. Abh. 2022;303:39–97.
    doi: 10.1127/njgpa/2022/1037google scholar: lookup
  292. Bernor R.L., Coillot T., Wolf D.. Phylogenetic Signatures in the Juvenile Skull and Dentition of Olduvai Gorge Eurygnathohippus cornelianus (Mammalia: Equidae). Riv. Ital. Paleontol. Stratigr. 2014;120:243–252.
  293. Hopwood A.T.. Die fossilen Pferde von Oldoway.. Wiss. Ergeb. Oldoway Exped. 1937;4:112–136.
  294. Thomas P.. Recherches stratigraphiques et paleontologiques sur quelques formations d’eau douce de l’Algerie.. Mem. Soc. Geol. Fr. 1884;3:1–50.
  295. Eisenmann V.. Les Chevaux (Equus Sensu Lato) Fossiles et Actuels: Crânes et Dents Jugales Supérieures.. 1980.
  296. Arambourg C.. Vertébrés continentaux du Miocène supérieur de l’Afrique du Nord.. Publ. Serv. Cart. Geol. L’algerie. 1959;4:1–161.
  297. Arambourg C.. Les Vertébrés du Pléistocène de l’Afrique du Nord.. Arch. Mus. Natl. D’histoiry Nat. 1970;10:1–127.
  298. Azzaroli A.. On Villafranchian Palaearctic Equids and their allies.. Palaeontogr. Ital. 1982;72:74–97.
  299. Geraads D, Raynal JP, Eisenmann V. The earliest human occupation of North Africa: a reply to Sahnouni et al. (2002).. J Hum Evol 2004 Jun;46(6):751-61; discussion 763-75.
    doi: 10.1016/j.jhevol.2004.01.008pubmed: 15183674google scholar: lookup
  300. Tchernov E.. The Lower Pleistocene Mammals of ‘Ubeidiya (Jordan Valley) Association. 1986.
  301. Yeshurun R, Zaidner Y, Eisenmann V, Martínez-Navarro B, Bar-Oz G. Lower Paleolithic hominin ecology at the fringe of the desert: Faunal remains from Bizat Ruhama and Nahal Hesi, Northern Negev, Israel.. J Hum Evol 2011 Apr;60(4):492-507.
    doi: 10.1016/j.jhevol.2010.01.008pubmed: 20303144google scholar: lookup
  302. Tchernov E.. The Faunal Sequence of the Southwest Asian Middle Paleolithic in Relation to Hominid Dispersal Events.. 2002. pp. 77–94.
  303. Broom R.. On evidence of a large horse recently extinct in South Africa.. Ann. S. Afr. Mus. 1909;7:281–282.
  304. Eisenmann V.. Equus capensis (Mammalia, Perissodactyla) from Elandsfontein.. Palaeontol. Afr. 2000;36:91–96.
  305. Churcher C.S.. The fossil equidae from the Krugersdorp Caves.. Ann. Transvaal Mus. 1970;26:145–168.
  306. Marom N, Lazagabaster IA, Shafir R, Natalio F, Eisenmann V, Horwitz LK. The Late Middle Pleistocene mammalian fauna of Oumm Qatafa Cave, Judean Desert: taxonomy, taphonomy and palaeoenvironment.. J Quat Sci 2022 May;37(4):612-638.
    doi: 10.1002/jqs.3414pmc: PMC9314136pubmed: 35915614google scholar: lookup
  307. Bagtache B., Hadjouis D., Eisenmann V.. Presence d’un Equus caballine (E. algericus n. sp.) et d’une autre espece nouvelle d’Equus (E. melkiensis n. sp.) dans l’Atérien des Allbroges, Algerie.. Comptes Rendus Séances L’académie Sci. Paris. 1984;298:609–612.
  308. Aouraghe H., Debenath A.. Les Equides du Pleistocene Superior de la grotte Zourah a el Harhoura, Maroc.. Quaternarie. 1999;10:283–292.
    doi: 10.3406/quate.1999.1649google scholar: lookup
  309. Oustalet E.. Une nouvelle espèce de zèbre, le zèbre de Grévy (Equus grevyi). Nature. 1882;10:12–14.
  310. O’Brien K., Tryon C.A., Blegen N., Kimeu B., Rowan J., Faith J.T.. First appearance of Grévy’s zebra (Equus grevyi), from the Middle Pleistocene of the Kapthurin Formation, Kenya, sheds lights on the evolution and paleoecology of large zebras.. Quat. Sci. Rev. 2021;256:106835.
    doi: 10.1016/j.quascirev.2021.106835google scholar: lookup
  311. Leonard JA, Rohland N, Glaberman S, Fleischer RC, Caccone A, Hofreiter M. A rapid loss of stripes: the evolutionary history of the extinct quagga.. Biol Lett 2005 Sep 22;1(3):291-5.
    doi: 10.1098/rsbl.2005.0323pmc: PMC1617154pubmed: 17148190google scholar: lookup
  312. Pedersen CT, Albrechtsen A, Etter PD, Johnson EA, Orlando L, Chikhi L, Siegismund HR, Heller R. A southern African origin and cryptic structure in the highly mobile plains zebra.. Nat Ecol Evol 2018 Mar;2(3):491-498.
    doi: 10.1038/s41559-017-0453-7pubmed: 29358610google scholar: lookup
  313. Linnaeus C.. Systema Naturae per Regna Tria Naturae, Secundum Classes, Ordines, Genera, Species cum Characteribus, Differentiis, Synonimis, Locis: Editio Decima, Reformata.. 1758. pp. 1–824.
  314. von Heuglin T.. Systematische Übersicht der Vögel Nord-Ost-Afrika’s: Mit Einschluss der Arabischen Küste des Rothen Meeres und der Nil-Quellen-Länder Südwärts bis zum IV. Grade Nördl. Breite.. 1866.
  315. Churcher C.S.. Oldest ass recovered from Olduvai Gorge, Tanzania, and the origin of asses.. J. Paleontol. 1982;56:1124–1132.
  316. Noack N.. Neues aus der Tierhandlung von Karl Hagenbeck, sowie aus dem Zoologischen Garten in Hamburg.. Zool. Gart. 1884;25:100–115.
  317. Groves C.P.. The taxonomy, distribution, and adaptations of recent equids.. 1986. pp. 11–65.
  318. Denzau G., Denzau H.. Wildesel.. 1999. pp. 1–221.
  319. Groves C.P., Smeenk C.. The nomenclature of the African wild ass.. Zool. Meded. 2007;81:121–135.
  320. Sam Y.. African origins of modern asses as seen from paleontology and DNA: What about the Atlas wild ass?. Geobios. 2020;58:73–84.
    doi: 10.1016/j.geobios.2019.12.001google scholar: lookup
  321. Berggren W.A., Van Couvering J.A.. The late Neogene: Biostratigraphy, geochronology and paleoclimatology of the last 15 million years in marine and continental sequences.. Palaeogeogr. Palaeoclimatol. Palaeoecol. 1974;16:1–216.
  322. Bell C.J., Lundelius E.L., Jr., Barnosky A.D., Graham R.W., Lindsay E.H., Ruez D.R., Jr., Semken H.A., Webb S.D., Zakrzwski R.J.. The Blancan, Irvingtonian, and Rancholabrean Mammal Ages.. 2004. pp. 232–314.
  323. Woodburne MO. The Great American Biotic Interchange: Dispersals, Tectonics, Climate, Sea Level and Holding Pens.. J Mamm Evol 2010 Dec;17(4):245-264.
    doi: 10.1007/s10914-010-9144-8pmc: PMC2987556pubmed: 21125025google scholar: lookup
  324. Avilla L.S., Bernardes C., Mothé D.. A new genus for Onohippidium galushai Macfadden and Skinner, 1979 (Mammalia, Equidae), from the late Hemphillian of North America.. J. Vertebr. Paleontol. 2015;35:e925909.
    doi: 10.1080/02724634.2014.925909google scholar: lookup
  325. Coltorti M, Abbazzi L, Ferretti MP, Iacumin P, Rios FP, Pellegrini M, Pieruccini P, Rustioni M, Tito G, Rook L. Last Glacial mammals in South America: a new scenario from the Tarija Basin (Bolivia).. Naturwissenschaften 2007 Apr;94(4):288-99.
    doi: 10.1007/s00114-006-0196-9pubmed: 17180614google scholar: lookup
  326. Bernor R.L., Golich U.B., Harzahauser M., Semprebon G.M.. The Pannonian C hipparions from the Vienna Basin.. Palaeogeogr. Palaeoclimatol. Palaeoecol. 2017;476:28–41.
    doi: 10.1016/j.palaeo.2017.03.026google scholar: lookup
  327. Barry J.C., Behrensmeyer A.K., Badgley C.E., Flynn L.J., Peltonen H., Cheema I.U., Pilbeam D., Lindsay E.H., Raza S.M., Rajpar A.R.. The Neogene Siwaliks of the Potwar Plateau, Pakistan.. 2013. pp. 373–399.
  328. Opdyke N.D., Lindsay E., Johnson G.D., Johnson N., Tahirkheli R.A.K., Mirza M.A.. Magnetic polarity stratigraphy and vertebrate paleontology of the upper siwalik subgroup of northern Pakistan.. Palaeogeogr. Palaeoclimatol. Palaeoecol. 1979;27:1–34.
    doi: 10.1016/0031-0182(79)90091-9google scholar: lookup
  329. Patnaik R.. Indian Neogene Siwalik mammalian biostratigraphy: An overview.. 2013. pp. 423–444.
  330. Gaur R., Chopra S.R.K.. On a new subspecies of Equus from Pinjor Formation of Upper Sivaliks—With remarks on Sivalik Equus.. J. Palaeontol. Soc. India. 1984;29:19–25.
  331. Zhu RX, Potts R, Pan YX, Yao HT, Lü LQ, Zhao X, Gao X, Chen LW, Gao F, Deng CL. Early evidence of the genus Homo in East Asia.. J Hum Evol 2008 Dec;55(6):1075-85.
    doi: 10.1016/j.jhevol.2008.08.005pubmed: 18842287google scholar: lookup
  332. Chauhan P.R.. Large mammal fossil occurrences and associated archaeological evidence in Pleistocene contexts of peninsular India and Sri Lanka.. Quarter. Int. 2008;192:20–42.
    doi: 10.1016/j.quaint.2007.06.034google scholar: lookup
  333. Van Couvering J., Delson E.. African Land Mammal Ages.. J. Vert. Pal. 2020;40:5.
    doi: 10.1080/02724634.2020.1803340google scholar: lookup
  334. Werdelin L.. Chronology of Neogene Mammal localities.. 2010. pp. 27–43.
  335. Pickford M.. Preliminary Miocene mammalian biostratigraphy for Western Kenya.. J. Hum. Evol. 1981;10:73–97.
    doi: 10.1016/S0047-2484(81)80026-7google scholar: lookup
  336. Gilbert H., Bernor R.L.. Equidae.. 2008. pp. 133–166.
  337. Geist V.. The relation of social evolution and dispersal in ungulates during the Pleistocene, with emphasis on Old World deer and the genus Bison.. Quat. Res. 1971;1:285–315.
    doi: 10.1016/0033-5894(71)90067-6google scholar: lookup
  338. Guthrie D.R.. Frozen Fauna of the Mammoth Steppe—The Story of Blue Babe.. 1990.
  339. McNab BK. Geographic and temporal correlations of mammalian size reconsidered: a resource rule.. Oecologia 2010 Sep;164(1):13-23.
    doi: 10.1007/s00442-010-1621-5pubmed: 20364270google scholar: lookup
  340. Saarinen J.. Ecometrics of Large Herbivorous Land Mammals in Relation to Climatic and Environmental Changes during the Pleistocene.. 2014.
  341. Damuth J, Janis CM. On the relationship between hypsodonty and feeding ecology in ungulate mammals, and its utility in palaeoecology.. Biol Rev Camb Philos Soc 2011 Aug;86(3):733-58.
    doi: 10.1111/j.1469-185X.2011.00176.xpubmed: 21418504google scholar: lookup
  342. Strömberg C.. Using phytolith assemblages to reconstruct the origin and spread of grass-dominated habitats in the great plains of North America during the late Eocene to early Miocene.. Palaeogeo. Palaeoclim. Palaeoecol. 2004;207:239–275.
    doi: 10.1016/j.palaeo.2003.09.028google scholar: lookup
  343. Strömberg CA. Decoupled taxonomic radiation and ecological expansion of open-habitat grasses in the Cenozoic of North America.. Proc Natl Acad Sci U S A 2005 Aug 23;102(34):11980-4.
    doi: 10.1073/pnas.0505700102pmc: PMC1189350pubmed: 16099827google scholar: lookup
  344. Strömberg C.. Evolution of Grasses and Grassland Ecosystems.. Annu. Rev. Earth Planet. Sci. 2011;39:517–544.
    doi: 10.1146/annurev-earth-040809-152402google scholar: lookup
  345. Janis C., Sahney S., Benton M.. Grit not grass: Concordant patterns of early origin of hypsodonty in Great Plains ungulates and Glires.. Palaeogeo. Palaeoclim. Palaeoecol. 2012;366:1–10.
  346. Semprebon G.M., Rivals F., Janis C.M.. The role of grass vs. exogenous abrasives in the paleodietary patterns of North American ungulates.. Front. Ecol. Evol. 2019;7:65.
    doi: 10.3389/fevo.2019.00065google scholar: lookup
  347. Strömberg C., McInerney F.. The Neogene transition from C3 to C4 grasslands in North America: Assemblage analysis of fossil phytoliths.. Paleobiology. 2011;37:50–71.
    doi: 10.1666/09067.1google scholar: lookup
  348. Pérez-Crespo V.A., Bravo-Cuevas V.M., Arroyo-Cabrales J.. Feeding habits of Equus conversidens and Haringtonhippus francisci from Valsequillo, Puebla, México.. Hist. Biol. 2021;34:1252–1259.
    doi: 10.1080/08912963.2021.1973452google scholar: lookup
  349. Hulbert R.C.. Taxonomic evolution in North American Neogene horses (Subfamily Equinae): The rise and fall of an adaptive radiation.. Paleobiology. 1993;19:216–234.
    doi: 10.1017/S0094837300015888google scholar: lookup
  350. Hulbert R.C., MacFadden B.J.. Morphological transformation and cladogenesis at the base of the adaptive radiation of Miocene hypsodont horses.. Am. Mus. Novit. 1991;3000:1–61.
  351. Radinsky L. Allometry and reorganization in horse skull proportions.. Science 1983 Sep 16;221(4616):1189-91.
    doi: 10.1126/science.221.4616.1189pubmed: 17811522google scholar: lookup
  352. Radinsky L. ONTOGENY AND PHYLOGENY IN HORSE SKULL EVOLUTION.. Evolution 1984 Jan;38(1):1-15.
    doi: 10.1111/j.1558-5646.1984.tb00254.xpubmed: 28556070google scholar: lookup
  353. Hulbert R.C.. Phylogenetic interrelationships and evolution of North American late Neogene Equidae.. 1989. pp. 176–196.
  354. Osborn H.F.. The Age of Mammals.. 1910.
  355. Scott W.B.. A History of Land Mammals in the Western Hemisphere.. 1937.
  356. Simpson G.G.. Tempo and Mode in Evolution.. 1944.
  357. Stirton R.A.. Observation of evolutionary rates in hypsodonty.. Evolution. 1947;1:32–41.
    doi: 10.1111/j.1558-5646.1947.tb02711.xgoogle scholar: lookup
  358. Webb S.D.. A history of savanna vertebrates in the New World. Part I: North America.. Annu. Rev. Ecol. Syst. 1977;8:355–380.
    doi: 10.1146/annurev.es.08.110177.002035google scholar: lookup
  359. Webb S.D.. The rise and fall of the late Miocene ungulate fauna in North America.. 1983. pp. 267–306.
  360. Janis C.M.. The significance of fossil ungulate communities as indicators of vegetation structure and climate.. 1984. pp. 85–104.
  361. Janis C.M.. Tertiary mammal evolution in the context of changing climates, vegetation, and tectonic events.. Annu. Rev. Ecol. Syst. 1993;24:467–500.
    doi: 10.1146/annurev.es.24.110193.002343google scholar: lookup
  362. Webb S.D., Opdyke N.D.. Effects of Past Global Change on Life.. 1995.
  363. Cohen J., DeSantis L.R.G., Lindsay E.L., Meachen J.A., O’Keefe F.R., Southon J.R., Binder W.J.. Dietary stability inferred from dental mesowear analysis in large ungulates from Racho La Brea and opportunistic feeding during the late Pleistocene.. Palaeogeo. Palaeoclim. Palaeoecol. 2021;570:110360.
    doi: 10.1016/j.palaeo.2021.110360google scholar: lookup
  364. Jones D.B., DeSantis L.R.G.. Dietary ecology od ungulates from La Brea tar pits in southern California: A multi-proxy approach.. Palaeogeo. Palaeoclim. Palaeoecol. 2017;466:110–117.
    doi: 10.1016/j.palaeo.2016.11.019google scholar: lookup
  365. Kaiser T.M., Solounias N., Fortelius M., Bernor R.L., Schrenk F.. Tooth mesowear analysis on Hippotherium primigenium from the Vallesian Dinotheriensande (Germany). A blind test study.. Carolinea. 2000;58:103–114.
  366. Kaiser T.M., Bernor R.L., Scott R.S., Franzen J.L., Solounias N.. New Interpretations of the Systematics and Palaeoecology of the Dorn Dürkheim 1 Hipparions (Late Miocene, Turolian Age [MN11]) Rheinhessen, German.. Senckenberg Lethea. 2003;83:103.
    doi: 10.1007/BF03043308google scholar: lookup
  367. Rivals F., Lister A.M.. Dietary flexibility and niche partitioning of large herbivores through the Pleistocene of Britain.. Quat. Sci. Rev. 2016;146:116–133.
    doi: 10.1016/j.quascirev.2016.06.007google scholar: lookup
  368. Head M.J.. Pollen and dinoflagellates from the Red Crag at Walton-on-the-Naze, Essex: Evidence for a mild climatic phase during the early Late Pliocene of eastern England.. Geol. Mag. 1998;135:803–817.
    doi: 10.1017/S0016756898001745google scholar: lookup
  369. Zalasiewicz J.A., Mathers S.J., Hughes M.J., Gibbard P.L., Peglar S.M., Harland R., Nicholson R.A., Boulton G.S., Cambridge P., Wealthall G.P.. Stratigraphy and Palaeoenvironments of the Red Crag and Norwich Crag Formations between Aldeburgh and Sizewell, Suffolk, England.. Philos. Trans. R. Soc. Lond. 1988;322:221–272.
  370. Rivals F., Athanassiou A.. Dietary adaptations in an ungulate community from the late pliocene of greece.. Palaeogeogr. Palaeoclimatol. Palaeoecol. 2008;265:134–139.
    doi: 10.1016/j.palaeo.2008.05.001google scholar: lookup
  371. Strani F., DeMiguel D., Bellucci L., Sardella R.. Dietary response of early pleistocene ungulate communities to the climate oscillations of the Gelasian/Calabrian transition in central Italy.. Palaeogeogr. Palaeoclimatol. Palaeoecol. 2018;499:102–111.
    doi: 10.1016/j.palaeo.2018.03.021google scholar: lookup
  372. Strani F., DeMiguel D., Sardella R., Bellucci L.. Paleoenvironments and climatic changes in the italian peninsula during the early pleistocene: Evidence from dental wear patterns of the ungulate community of Coste San Giacomo.. Quat. Sci. Rev. 2015;121:28–35.
    doi: 10.1016/j.quascirev.2015.05.008google scholar: lookup
  373. Strani F., Pushkina D., Bocherens H., Bellucci L., Sardella R., DeMiguel D.. Dietary adaptations of early and middle pleistocene equids from the anagni basin (Frosinone, Central Italy). Front. Ecol. Evol. 2019;7:176.
    doi: 10.3389/fevo.2019.00176google scholar: lookup
  374. Saarinen J., Oksanen O., Zliobaite I., Fortelius M., DeMiguel D., Azanza B., Bocherens H., Luzón C., Solano-Garcia J., Yravedra J.. Pliocene to Middle Pleistocene climate history in the Guadix-Baza Basin, and the environmental conditions of early Homo in Europe.. Quat. Sci. Rev. 2021;268:107132.
    doi: 10.1016/j.quascirev.2021.107132google scholar: lookup
  375. Strani F., DeMiguel D., Alba D., Moyà-Solà S., Bellucci L., Sardella R., Madurell-Malapeira J.. The effects of the “0.9 Ma event” on the Mediterranean ecosystems during the Early-Middle Pleistocene transition as revealed by dental wear patterns of fossil ungulates.. Quat. Sci. Rev. 2019;210:80–89.
    doi: 10.1016/j.quascirev.2019.02.027google scholar: lookup
  376. Blain H.-A., Lozano-Fernández I., Agustí J., Bailon S., Menéndez L., Patrocinio M., Ros-Montoya S., Manuel J., Toro-Moyano I., Martínez-Navarro B.. Refining upon the climatic background of the early Pleistocene hominid settlement in western Europe: Barranco León and Fuente Nueva-3 (Guadix- Baza Basin, SE Spain). Quat. Sci. Rev. 2016;144:132–144.
    doi: 10.1016/j.quascirev.2016.05.020google scholar: lookup
  377. Sánchez-Bandera C., Oms O., Blain H.-A., Lozano-Fernández I., Bispal-Chinesta J.F., Agustí J., Saarinen J., Fortelius M., Titton S., Serrano-Ramos A.. New stratigraphically constrained palaeoenvironmental reconstructions for the first human settlement in Western Europe: The Early Pleistocene herpetofaunal assemblages from Barranco León and Fuente Nueva 3 (Granada, SE Spain). Quat. Sci. Rev. 2020;243:106466.
    doi: 10.1016/j.quascirev.2020.106466google scholar: lookup
  378. Kahlke R.D., García N., Kostopoulos D.S., Lacombat F., Lister A.M., Mazza P.P., Spassov N., Titov V.V.. Western Palaearctic palaeoenvironmental conditions during the Early and early Middle Pleistocene inferred from large mammal communities, and implications for hominin dispersal in Europe.. Quat. Sci. Rev. 2011;30:1368–1395.
    doi: 10.1016/j.quascirev.2010.07.020google scholar: lookup
  379. Rivals F, Julien MA, Kuitems M, Van Kolfschoten T, Serangeli J, Drucker DG, Bocherens H, Conard NJ. Investigation of equid paleodiet from Schöningen 13 II-4 through dental wear and isotopic analyses: Archaeological implications.. J Hum Evol 2015 Dec;89:129-37.
    doi: 10.1016/j.jhevol.2014.04.002pubmed: 25242064google scholar: lookup
  380. Roberts M.B.. Excavation of the Lower Paleolithic site at Amey’s Eartham Pit, Boxgrove, West Sussex: A preliminary report.. Proc. Prehist. Soc. 1986;52:215–245.
    doi: 10.1017/S0079497X00006666google scholar: lookup
  381. Urban B, Bigga G. Environmental reconstruction and biostratigraphy of late Middle Pleistocene lakeshore deposits at Schöningen.. J Hum Evol 2015 Dec;89:57-70.
    doi: 10.1016/j.jhevol.2015.10.002pubmed: 26638875google scholar: lookup
  382. Strani F., DeMiguel D., Bona F., Sardella R., Biddittu I., Bruni L., De Castro A., Guadagnoli F., Bellucci L.. Ungulate dietary adaptations and palaeoecology of the Middle Pleistocene site of Fontana Ranuccio (anagni, central italy). Palaeogeogr. Palaeoclimatol. Palaeoecol. 2018;496:238–247.
    doi: 10.1016/j.palaeo.2018.01.041google scholar: lookup
  383. Leroi-Gourhan A.. Pollen analysis of sediment samples from Gough’s Cave, Cheddar.. Proc. Univ. Bristol. Spelaeol. Soc. 1986;17:141–144.
  384. Bates M.R., Bates C.R., Gibbard P.L., Macphail R.I., Owen F.J., Parfitt S.A., Preece R.C., Roberts M.B., Robinson J.E., Whittaker J.E.. Late Middle Pleistocene deposits at Norton Farm on the West Sussex coastal plain, southern England.. J. Quat. Sci. 2000;15:61–89.
    doi: 10.1002/(SICI)1099-1417(200001)15:1<61::AID-JQS463>3.0.CO;2-Kgoogle scholar: lookup
  385. West R.G., Lambert C.A., Sparks B.W., Dickson J.H.. Interglacial deposits at Ilford, Essex.. Philos. Trans. R. Soc. 1964;247:185–212.
  386. Gibbard P.L.. Pleistocene History of the Lower Thames Valley.. 1994.
  387. Sedov S.N., Khokhlova O.S., Sinitsyn A.A., Korkka A.V., Ortega B., Solleiro E., Rozanova M.S., Kuznetsova A.M.. Kazdym Late Pleistocene paleosol sequences as an instrument for the local paleographic reconstruction of the Kostenki 14 key section (Voronezh oblast) as an example.. Eurasian Soil Sc. 2010;43:876–892.
    doi: 10.1134/S1064229310080053google scholar: lookup
  388. Ukraintseva V.V.. Mammoths and the Environment.. 2013.
  389. van Asperen E.N., Stefaniak K., Proskurnyak I., Ridush B.. Equids from Emine-Bair-Khosar Cave (Crimea, Ukraine): Co-occurrence of the stenonid Equus hydruntinus and the caballoid E. ferus latipes based on skull and postcranial remains.. Palaeontol. Electron. 2012;15:28.
    doi: 10.26879/280google scholar: lookup
  390. Bernor R.L., Kaiser T.M., Nelson S.V.. The Oldest Ethiopian Hipparion (Equinae, Perissodactyla) from Chorora: Systematics, Paleodiet and Paleoclimate.. Cour. Forsch. Inst. Senckenberg. 2004;246:213–226.
  391. Bernor R.L.. The Latest Miocene Hipparionine (Equidae) from Lemundong’o, Kenya.. Kirtlandia. 2007;56:148–151.
  392. Bernor R.L., Kaiser T.M., Wolf D.. Revisiting Sahabi Equid Species Diversity, Biogeographic Patterns and Diet Preferences.. Gharyounis Bull. 2008;5:159–167.
  393. Bernor R.L., Haile Selassie Y.. Equidae.. 2009. pp. 397–428.
  394. Bernor R.L., Boaz N.T., Rook L.. Eurygnathohippus feibeli (Perissodactyla: Mammalia) from the late Miocene of Sahabi (Libya) and its evolutionary and Biogeographic Significance.. Bolletino Soc. Paleontol. Ital. 2012;51:39–48.
  395. Schulz E., Kaiser T.M.. Historical distribution, habitat requirements and feeding ecology of the genus Equus (Perissodactyla). Mammal. Rev. 2013;43:111–123.
    doi: 10.1111/j.1365-2907.2012.00210.xgoogle scholar: lookup
  396. Goloboff P., Farris J., Nixon K.. TNT, a free program for phylogenetic analysis.. Cladistics. 2008;24:774–786.
    doi: 10.1111/j.1096-0031.2008.00217.xgoogle scholar: lookup
  397. Bennett D.K.. Stripes do not a zebra make, Part I: A cladistic analysis of Equus.. Syst. Zool. 1980;29:272–287.
    doi: 10.2307/2412662google scholar: lookup
  398. Antoine P.O.. Phylogénie et évolution des Elasmotheriina (Mammalia, Rhinocerotidae). Mémoir. Mus. Natl. Hist. 2002;188:1–359.
  399. Antoine P.O., Downing K.F., Crochet J.Y., Duranthon F., Flynn L.J., Marivaux L., Métais G., Rajpar A.R., Roohi G.. A revision of Aceratherium blanfordi Lydekker, 1884 (Mammalia: Rhinocerotidae) from the Early Miocene of Pakistan: Postcranials as a key.. Zool. J. Linn. Soc. Lond. 2010;160:139–194.
    doi: 10.1111/j.1096-3642.2009.00597.xgoogle scholar: lookup
  400. Pandolfi L.. Persiatherium rodleri, gen. et sp. nov. (Mammalia, Rhinocerotidae), from the Late Miocene of Maragheh (Northwestern Iran). J. Vert Paleontol. 2015;36:e1040118.
    doi: 10.1080/02724634.2015.1040118google scholar: lookup
  401. de Queiroz K., Gauthier J.. Phylogenetic taxonomy.. Annu. Rev. Ecol. Syst. 1992;23:449–480.
    doi: 10.1146/annurev.es.23.110192.002313google scholar: lookup
  402. Ereshefsky M.. The Poverty of the Linnaean hierarchy. A Philosophical Study of Biological Taxonomy.. 2001.
  403. Papavero N., Llorente-Bousquets J., Minoro Abel J.. Proposal of a new system of nomenclature for phylogenetic systematics.. Arq. Zool. 2001;36:1–145.
    doi: 10.11606/issn.2176-7793.v36i1p1-145google scholar: lookup
  404. Béthoux O.. Propositions for a character-state-based biological taxonomy.. Zool. Scr. 2007;36:409–416.
    doi: 10.1111/j.1463-6409.2007.00287.xgoogle scholar: lookup
  405. Cantino P.D., de Queiroz K.. PhyloCode: International Code of Phylogenetic Nomenclature (Version 4c) 2010.. 2010.
  406. Zachos F.E.. Linnaean ranks, temporal banding, and time-clipping: Why not slaughter the sacred cow?. Biol. J. Linn. Soc. 2011;103:732–734.
    doi: 10.1111/j.1095-8312.2011.01711.xgoogle scholar: lookup
  407. Vences M, Guayasamin JM, Miralles A, De la Riva I. To name or not to name: Criteria to promote economy of change in Linnaean classification schemes.. Zootaxa 2013;3636:201-44.
    doi: 10.11646/zootaxa.3636.2.1pubmed: 26042291google scholar: lookup
  408. Hennig W.. Phylogenetic Systematics.. 1966.
  409. Mayr E.. Principles of Systematic Zoology.. 1969.
  410. Groves C.P.. Primate Taxonomy.. 2001.
  411. Groves C.P.. The what, why, and how of primate taxonomy.. Int. J. Primatol. 2004;25:1105–1126.
    doi: 10.1023/B:IJOP.0000043354.36778.55google scholar: lookup
  412. Simpson G.G.. The Major Features of Evolution.. 1953.
  413. MAYR E. Taxonomic categories in fossil hominids.. Cold Spring Harb Symp Quant Biol 1950;15:109-18.
    doi: 10.1101/SQB.1950.015.01.013pubmed: 14942702google scholar: lookup
  414. de Queiroz K.. Replacement of an essentialistic perspective on taxonomic definitions as exemplified by the definition of “Mammalia”.. Syst. Biol. 1994;43:497–510.
    doi: 10.1093/sysbio/43.4.497google scholar: lookup
  415. Janis C.M., Bernor R.L.. The Evolution of Equid Monodactyly: A Review Including a New Hypothesis.. Front. Ecol. Evol. 2019;7:119.
    doi: 10.3389/fevo.2019.00119google scholar: lookup
  416. Schultz J.R.. Plesippus francescana (Frick) from the late Pliocene, Coso Mountains, California, with a review of the genus Plesippus;. Publications of the Carnegie Institution of Washington. 1936;1–13.
  417. Eisenmann V., Baylac M.. Extant and fossil Equus (Mammalia, Perissodactyla) skulls: A morphometric definition of the subgenus Equus.. Zool. Scr. 2000;29:89–100.
    doi: 10.1046/j.1463-6409.2000.00034.xgoogle scholar: lookup
  418. Brochu A.B., Sumrall C.D.. Phylogenetic Nomenclature and Paleontology.. J. Paleontol. 2001;75:754–757.
    doi: 10.1666/0022-3360(2001)075<0754:PNAP>2.0.CO;2google scholar: lookup
  419. Bryant H.N.. Explicitness, Stability, and Universality in the Phylogenetic Definition and Usage of Taxon Names: A Case Study of the Phylogenetic Taxonomy of the Carnivora (Mammalia). Syst. Biol. 1996;45:174–189.
    doi: 10.1093/sysbio/45.2.174google scholar: lookup
  420. Cantino P.D., Olmstead R.G., Wagstaff S.J.. A Comparison of Phylogenetic Nomenclature with the Current System: A Botanical Case Study.. Syst. Biol. 1997;46:313–331.
    doi: 10.1093/sysbio/46.2.313google scholar: lookup
  421. Schulte JA, Macey JR, Larson A, Papenfuss TJ. Molecular tests of phylogenetic taxonomies: a general procedure and example using four subfamilies of the lizard family Iguanidae.. Mol Phylogenet Evol 1998 Dec;10(3):367-76.
    doi: 10.1006/mpev.1998.0541pubmed: 10051389google scholar: lookup
  422. de Queiroz K., Gauthier J.. Phylogeny as a Central Principle in Taxonomy: Phylogenetic Definitions of Taxon Names.. Syst. Biol. 1990;39:307–322.
    doi: 10.2307/2992353google scholar: lookup
  423. de Queiroz K, Gauthier J. Toward a phylogenetic system of biological nomenclature.. Trends Ecol Evol 1994 Jan;9(1):27-31.
    doi: 10.1016/0169-5347(94)90231-3pubmed: 21236760google scholar: lookup
  424. de Queiroz K., Cantino P.D.. Taxon Names, Not Taxa, Are Defined.. Taxon. 2001;50:821–826.
    doi: 10.2307/1223709google scholar: lookup
  425. Vilstrup JT, Seguin-Orlando A, Stiller M, Ginolhac A, Raghavan M, Nielsen SC, Weinstock J, Froese D, Vasiliev SK, Ovodov ND, Clary J, Helgen KM, Fleischer RC, Cooper A, Shapiro B, Orlando L. Mitochondrial phylogenomics of modern and ancient equids.. PLoS One 2013;8(2):e55950.
    pmc: PMC3577844pubmed: 23437078doi: 10.1371/journal.pone.0055950google scholar: lookup
  426. MacFadden BJ, Solounias N, Cerling TE. Ancient diets, ecology, and extinction of 5-million-year-Old horses from florida.. Science 1999 Feb 5;283(5403):824-7.
    doi: 10.1126/science.283.5403.824pubmed: 9933161google scholar: lookup
  427. Semprebon G.M., Rivals F., Solounias N., Hulbert R.C.. Paleodietary reconstruction of fossil horses from the Eocene through Pleistocene of North America.. Palaeogeogr. Palaeoclim. Palaeoecol. 2016;442:110–127.
    doi: 10.1016/j.palaeo.2015.11.004google scholar: lookup
  428. Miller W.E.. The Late Pliocene Las Tunas Local Fauna from Southernmost Baja California, Mexico.. J. Paleontol. 1980;54:762–805.
  429. MacFadden B.J., Carranza-Castañeda O.. Cranium of Dinohippus mexicanus (Mammalia: Equidae) from the Early Pliocene (latest Hemphillian) of Central Mexico, and the origin of Equus.. Bull. Fla. Mus. Nat. Hist. 2002;43:163–185.
  430. Carranza-Castañeda O.. Dinohippus mexicanus (Early—Late, Late, and Latest Hemphillian) and the transition to genus Equus, in Central Mexico faunas.. Front. Earth Sci. 2019;7:89.
    doi: 10.3389/feart.2019.00089google scholar: lookup
  431. Lowenstein JM, Ryder OA. Immunological systematics of the extinct quagga (Equidae).. Experientia 1985 Sep 15;41(9):1192-3.
    doi: 10.1007/BF01951724pubmed: 4043335google scholar: lookup
  432. George M Jr, Ryder OA. Mitochondrial DNA evolution in the genus Equus.. Mol Biol Evol 1986 Nov;3(6):535-46.
    pubmed: 2832696doi: 10.1093/oxfordjournals.molbev.a040414google scholar: lookup
  433. Oakenfull EA, Clegg JB. Phylogenetic relationships within the genus Equus and the evolution of alpha and theta globin genes.. J Mol Evol 1998 Dec;47(6):772-83.
    doi: 10.1007/PL00006436pubmed: 9847419google scholar: lookup
  434. Krüger K, Gaillard C, Stranzinger G, Rieder S. Phylogenetic analysis and species allocation of individual equids using microsatellite data.. J Anim Breed Genet 2005 Apr;122 Suppl 1:78-86.
    doi: 10.1111/j.1439-0388.2005.00505.xpubmed: 16130461google scholar: lookup
  435. Steiner C.C., Ryder O.A.. Molecular phylogeny and evolution of the Perissodactyla.. Zool. J. Linn. Soc. 2011;163:1289–1303.
    doi: 10.1111/j.1096-3642.2011.00752.xgoogle scholar: lookup
  436. Orlando L, Mashkour M, Burke A, Douady CJ, Eisenmann V, Hänni C. Geographic distribution of an extinct equid (Equus hydruntinus: Mammalia, Equidae) revealed by morphological and genetical analyses of fossils.. Mol Ecol 2006 Jul;15(8):2083-93.
    doi: 10.1111/j.1365-294X.2006.02922.xpubmed: 16780426google scholar: lookup
  437. Orlando L, Male D, Alberdi MT, Prado JL, Prieto A, Cooper A, Hänni C. Ancient DNA clarifies the evolutionary history of American Late Pleistocene equids.. J Mol Evol 2008 May;66(5):533-8.
    doi: 10.1007/s00239-008-9100-xpubmed: 18398561google scholar: lookup
  438. Flint J., Ryder O.A., Clegg J.B.. Comparison of theα-globin gene cluster structure in Perissodactyla.. J. Mol. Evol. 1990;30:36–42.
    doi: 10.1007/BF02102451google scholar: lookup
  439. Wallner B, Brem G, Müller M, Achmann R. Fixed nucleotide differences on the Y chromosome indicate clear divergence between Equus przewalskii and Equus caballus.. Anim Genet 2003 Dec;34(6):453-6.
    doi: 10.1046/j.0268-9146.2003.01044.xpubmed: 14687077google scholar: lookup
  440. Kaminski M. The biochemical evolution of the horse.. Comp Biochem Physiol B 1979;63(2):175-8.
    doi: 10.1016/0305-0491(79)90025-7pubmed: 400961google scholar: lookup
  441. Ishida N, Oyunsuren T, Mashima S, Mukoyama H, Saitou N. Mitochondrial DNA sequences of various species of the genus Equus with special reference to the phylogenetic relationship between Przewalskii's wild horse and domestic horse.. J Mol Evol 1995 Aug;41(2):180-8.
    doi: 10.1007/BF00170671pubmed: 7666447google scholar: lookup
  442. Oakenfull E.A., Lim H.N., Ryder O.A.. A survey of equid mitochondrial DNA: Implications for the evolution, genetic diversity and conservation of Equus.. Conserv. Genet. 2000;1:341–355.
    doi: 10.1023/A:1011559200897google scholar: lookup
  443. Cai D, Zhu S, Gong M, Zhang N, Wen J, Liang Q, Sun W, Shao X, Guo Y, Cai Y, Zheng Z, Zhang W, Hu S, Wang X, Tian H, Li Y, Liu W, Yang M, Yang J, Wu D, Orlando L, Jiang Y. Radiocarbon and genomic evidence for the survival of Equus Sussemionus until the late Holocene.. Elife 2022 May 11;11.
    doi: 10.7554/eLife.73346pmc: PMC9142152pubmed: 35543411google scholar: lookup

Citations

This article has been cited 5 times.
  1. Schweitzer MH. Paleontology in the 21st Century. Biology (Basel) 2023 Mar 22;12(3).
    doi: 10.3390/biology12030487pubmed: 36979178google scholar: lookup
  2. Jafari H, Abebe BK, Cong L, Ahmed Z, Zhaofei W, Sun M, Muhatai G, Chuzhao L, Dang R. Review: Genomic insights into the adaptive traits and stress resistance in modern horses. Stress Biol 2026 Jan 12;6(1):5.
    doi: 10.1007/s44154-025-00274-1pubmed: 41521281google scholar: lookup
  3. Matoshko AV, Gibbard P. The river Dniester valley: a long record of late-Cenozoic fluvial evolution within the Eastern Carpathian foreland and East European Platform margin. R Soc Open Sci 2025 Oct;12(10):250523.
    doi: 10.1098/rsos.250523pubmed: 41103769google scholar: lookup
  4. Du W, Sun Q, Hu S, Yu P, Kan S, Zhang W. Equus mitochondrial pangenome reveals independent domestication imprints in donkeys and horses. Sci Rep 2025 Feb 25;15(1):6803.
    doi: 10.1038/s41598-025-91564-1pubmed: 40000832google scholar: lookup
  5. Steklis NG, Peñaherrera-Aguirre M, Steklis HD, Herrera I. Why Were Zebras Not Domesticated? A Review of Domesticability Traits and Tests of Their Role in Ungulate Domestications with Macroevolutionary Models. Animals (Basel) 2024 Aug 14;14(16).
    doi: 10.3390/ani14162355pubmed: 39199888google scholar: lookup
Subscribe to our newsletter
Join 99,727 horse owners like you who receive our email updates on horse health and nutrition.