University of Oulu

Cirilli, O., Machado, H., Arroyo-Cabrales, J., Barrón-Ortiz, C. I., Davis, E., Jass, C. N., Jukar, A. M., Landry, Z., Marín-Leyva, A. H., Pandolfi, L., Pushkina, D., Rook, L., Saarinen, J., Scott, E., Semprebon, G., Strani, F., Villavicencio, N. A., Kaya, F., & Bernor, R. L. (2022). Evolution of the family equidae, subfamily equinae, in north, central and south america, eurasia and africa during the plio-pleistocene. Biology, 11(9), 1258.

Evolution of the family Equidae, subfamily Equinae, in North, Central and South America, Eurasia and Africa during the plio-pleistocene

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Author: Cirilli, Omar1,2; Machado, Helena3; Arroyo-Cabrales, Joaquin4;
Organizations: 1Laboratory of Evolutionary Biology, Department of Anatomy, College of Medicine, Howard University, Washington, DC 20059, USA
2Earth Science Department, Paleo[Fab]Lab, University of Florence, Via La Pira 4, 50121 Firenze, Italy
3Earth Sciences Department, University of Oregon, 100 Cascade Hall, Eugene, OR 97403, USA
4Instituto Nacional de Antropología e Historia Laboratorio de Arqueozoología “M. en C. Ticul Álvarez Solórzano”, Subdirección de Laboratorios y Apoyo Académico, Ciudad de Mexico 00810, Mexico
5Quaternary Palaeontology Program, Royal Alberta Museum, 9810 103a Ave NW, Edmonton, AB T5J 0G2, Canada
6Clark Honors College, University of Oregon, Eugene, OR 97401, USA
7Department of Geosciences, University of Arizona, 1040 E 4th St., Tucson, AZ 85721, USA
8Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, 10th St and Constitution Ave NW, Washington, DC 20013, USA
9Department of Earth Sciences, University of Ottawa, 25 Templeton Street, Ottawa, ON K1N 6N5, Canada
10Laboratorio de Paleontología, Facultad de Biología, Universidad Michoacana de San Nicolás de Hidalgo, Edif. R 2. Piso. Ciudad Universitaria, Morelia 58030, Mexico
11Department of Science, University of Basilicata, Via dell’Ateneo Lucano 10, 85100 Potenza, Italy
12Department of Geosciences and Geography, University of Helsinki, FI-00014 Helsinki, Finland
13Cogstone Resource Management, Inc., 1518 W., Taft Avenue, Orange, CA 92865, USA
14Department of Biology, California State University, 5500 University Parkway, San Bernardino, CA 92407, USA
15Department of Biology, Bay Path University, 588 Longmeadow Street, Longmeadow, MA 01106, USA
16PaleoFactory, Dipartimento di Scienze della Terra, Sapienza Università di Roma, Piazzale Aldo Moro 5, 00185 Rome, Italy
17Departamento de Ciencias de La Tierra, Instituto Universitario de Investigación en Ciencias Ambientales de Aragón (IUCA), Universidad de Zaragoza, 50009 Zaragoza, Spain
18Corporación Laguna de Taguatagua, Av. Libertador Bernardo O’Higgins 351, Santiago 1030000, Chile
19Instituto de Ciencias Sociales, Universidad de O’Higgins, Av., Libertador Bernardo O’Higgins 611, Rancagua 2852046, Chile
20Biomolecular Laboratory, Institut Català de Paleoecologia Humana i Evolucio Social, 43007 Tarragona, Spain
21Department of Archaeology, University of Oulu, FI-90014 Oulu, Finland
22Human Origins Program, Department of Anthropology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20013, USA
Format: article
Version: published version
Access: open
Online Access: PDF Full Text (PDF, 3.3 MB)
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Language: English
Published: Multidisciplinary Digital Publishing Institute, 2022
Publish Date: 2023-02-03


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 Equus evolution, their biochronologic and biogeographic background and paleoecological and paleoclimatic contexts.

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Series: Biology
ISSN: 2079-7737
ISSN-E: 2079-7737
ISSN-L: 2079-7737
Volume: 11
Issue: 9
Article number: 1258
DOI: 10.3390/biology11091258
Type of Publication: A1 Journal article – refereed
Field of Science: 1181 Ecology, evolutionary biology
Funding: This research was funded by the National Science Foundation (DBI:ABI 1759882 and 1759821) to R.L. Bernor and E. Davis, respectively, under the aegis of the FuTRES Equid working group for which they, O. Cirilli and H. Machado have been funded. F.K. has been funded by Finnish Cultural Foundation (project nr. 00220063). Z.L. acknowledge the Admission Scholarship from University of Ottawa, and a Canadian Graduate Scholarship-Doctoral Program from Natural Sciences and Engineering Research Council of Canada. A.M.J. thanks the Yale Institute for Biospheric Studies for funding. L.P. thanks the European Commission’s Research Infrastructure Action, EU-SYNTHESYS projects AT-TAF-2550, DE-TAF-3049, GB-TAF-2825, HU-TAF-3593, HU-TAF-5477, ES-TAF-2997, the SYNTHESYS Project (access on 5 August 2022) which is financed by European Community Research Infrastructure Action under the FP7 “Capacities” Program, and the research project “Ecomorphology of fossil and extant Hippopotamids and Rhinocerotids” granted to L.P. by the University of Florence (“Progetto Giovani Ricercatori Protagonisti” initiative). J.S. wishes to acknowledge the Academy of Finland (AoF. project nr. 340775/346292, “NEPA-Non-analogue ecosystems in the past”). F.S. is supported by Sapienza “5 per mille” funds (ref. SPC: 2021-0070-1350-175998).
Dataset Reference: All data generated by this study are available in this manuscript and the accompanying Supplementary Materials (Supplementary Materials Tables S1–S6).
Copyright information: © 2022 by the authors. This article is an open access article distributed under the terms and conditions of the Creative Commons Attribution (CC BY) license (