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Bernor, R. L., Kaya, F., Kaakinen, A., Saarinen, J., & Fortelius, M. (2021). Old world hipparion evolution, biogeography, climatology and ecology. Earth-Science Reviews, 221, 103784.

Old world hipparion evolution, biogeography, climatology and ecology

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Author: Bernor, Raymond L.1,2; Kaya, Ferhat3,4; Kaakinen, Anu3;
Organizations: 1College of Medicine, Department of Anatomy, Laboratory of Evolutionary Biology, Howard University, 520 W St. N.W, Washington D.C 20059, USA
2Human Origins Program, Department of Anthropology, Smithsonian Institution, 10th St. & Constitution Ave. NE, Washington D.C 20002, USA
3Department of Geosciences and Geography, P.O. Box 64, 00014, University of Helsinki, Finland
4Department of Archaeology, University of Oulu, Oulu, Finland
5Finnish Museum of Natural History (LUOMUS), Helsinki, Finland
Format: article
Version: published version
Access: open
Online Access: PDF Full Text (PDF, 6.3 MB)
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Language: English
Published: Elsevier, 2021
Publish Date: 2021-09-28


Nearly five decades ago Berggren and Van Couvering proposed an Old World “Hipparion Datum” wherein a North American Hipparion extended its range across Eurasia and Africa as an “instantaneous prochoresis” populating the Old World. Four decades ago Woodburne and Bernor examined European and North African hipparion assemblages and proposed a number of distinct hipparion lineages, sharply departing from the mono-generic paradigm of previous work. Through the 1980s until now, hipparion systematic studies have delineated multiple superspecific groups of hipparions. Herein, we define 10 recognizable genus-rank Eurasian and African taxa delineating their chronologic occurrences, geographic extent and where data exists, their body mass and paleodietary preferences. Our study supports the current interpretation that a species of North American Cormohipparion extended its range into the Old World in the early late Miocene. Regional first occurrences of Cormohipparion are recognized in the Potwar Plateau, Pakistan and Sinap Tepe, Turkey 10.8 Ma. The slightly derived lineage Hippotherium is recorded earlier in the Pannonian C of the Vienna Basin, 11.4–11.0 Ma marking the chronologic “Hipparion” Datum at the lower boundary of Mammal Neogene (MN) Unit 9. Within MN 9, 11.2–9.9 Ma, Cormohipparion underwent a minor diversification whereas Hippotherium diversified in Central and Western Europe and China and Sivalhippus (S. nagriensis) originated in the Indian Subcontinent. Whereas Cormohipparion did not survive into the late Vallesian, MN10 (9.9–8.9 Ma), Hippotherium and Sivalhippus did and the Cremohipparion and Hipparion s.s. lineages originated. During the early and middle Turolian (MN11–12, 8.9–6.8 Ma) Hippotherium, Sivalhippus, Cremohipparion and Hipparion persisted and new lineages, Eurygnathohippus, Plesiohipparion, Baryhipparion and Shanxihippus originated. An initial extinction interval occurred at the end of the Miocene, MN13 (6.8–5.3 Ma) wherein all but one endemic species of Hippotherium, H. malpassi (Italy), Hipparion and several species of Cremohipparion became extinct. Lineage and species reduction continued across the Mio-Pliocene boundary so that by the beginning of the Pliocene (MN14, 5.3 Ma) only African species of Eurygnathohippus, Chinese Plesiohipparion houfenense and Proboscidipparion sinense remained. The later Pliocene (MN15–16, ca. 5.0–2.5 Ma) documents the persistence of endemic Chinese Baryhipparion insperatum, modest diversification of African Eurygnatohippus spp. and Chinese Plesiohipparion and Proboscidipparion spp. Eurygnatohippus made a limited geographic extension into the Indian subcontinent during MN16, whereas Pleisohipparion and Proboscidipparion extended their ranges into Eurasia during MN15 and MN16. The latest occurring hipparions are Proboscidipparion sinense at 1.0 Ma in China and Eurygnatohippus cornelianus in Africa <1.0 Ma. Old World hipparion lineages early on increased their body mass in MN9. During the Turolian interval (MN11–13) hipparion lineages diversified their body mass from very small (<100 kg) to heavy forms (>300 kg), with the smaller forms being predominately grass feeders and larger ones being mixed feeders. Decreased hipparion lineage and species diversity in the Pliocene was accompanied by increased average body size and hypsodonty probably in response to more seasonal Eurasian and African environments. There is no evidence that hipparions ever adapted to cold and dry Old World Pleistocene environments.

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Series: Earth-science reviews
ISSN: 0012-8252
ISSN-E: 1872-6828
ISSN-L: 0012-8252
Volume: 221
Article number: 103784
DOI: 10.1016/j.earscirev.2021.103784
Type of Publication: A1 Journal article – refereed
Field of Science: 1181 Ecology, evolutionary biology
1171 Geosciences
615 History and archaeology
Funding: Bernor wishes to acknowledge research funding by NSF EAR grants 8806645, 0125009, 1113175, 1558586, DBI grant 1759882 for the FuTRES database, and support from HERC, U.C. Berkley, the Smithsonian Human Origins Program. This work was financially supported by the project “Tracing the winds” (316799) funded by the Academy of Finland (AK). FK was financially supported by Finnish Cultural Foundation during this work. JS was working on an Academy of Finland -funded postdoctoral research project Behavioural and morphological adaptation to environmental change: the example of African Neogene to Quaternary Proboscidea (Academy of Finland n:o 315691) during this work.
Copyright information: © 2021 The Authors. Published by Elsevier B.V. This is an open access article under the CC BY license (