The origins and spread of domestic horses from the Western Eurasian steppes

Librado P. Khan N. Fages A. Kusliy M.A. Suchan T. Tonasso-Calvière L. Schiavinato S. Alioglu D. Fromentier A. Perdereau A. Aury J.-M. Gaunitz C. Chauvey L. Seguin-Orlando A. Der Sarkissian C. Southon J. Shapiro B. Tishkin A.A. Kovalev A.A. Alquraishi S. Alfarhan A.H. Al-Rasheid K.A.S. Seregély T. Klassen L. Iversen R. Bignon-Lau O. Bodu P. Olive M. Castel J.-C. Boudadi-Maligne M. Alvarez N. Germonpré M. Moskal-del Hoyo M. Wilczyński J. Pospuła S. Lasota-Kuś A. Tunia K. Nowak M. Rannamäe E. Saarma U. Boeskorov G. Lōugas L. Kyselý R. Peške L. Bălășescu A. Dumitrașcu V. Dobrescu R. Gerber D. Kiss V. Szécsényi-Nagy A. Mende B.G. Gallina Z. Somogyi K. Kulcsár G. Gál E. Bendrey R. Allentoft M.E. Sirbu G. Dergachev V. Shephard H. Tomadini N. Grouard S. Kasparov A. Basilyan A.E. Anisimov M.A. Nikolskiy P.A. Pavlova E.Y. Pitulko V. Brem G. Wallner B. Schwall C. Keller M. Kitagawa K. Bessudnov A.N. Bessudnov A. Taylor W. Magail J. Gantulga J.-O. Bayarsaikhan J. Erdenebaatar D. Tabaldiev K. Mijiddorj E. Boldgiv B. Tsagaan T. Pruvost M. Olsen S. Makarewicz C.A. Valenzuela Lamas S. Albizuri Canadell S. Nieto Espinet A. Iborra M.P. Lira Garrido J. Rodríguez González E. Celestino S. Olària C. Arsuaga J.L. Kotova N. Pryor A. Crabtree P. Zhumatayev R. Toleubaev A. Morgunova N.L. Kuznetsova T. Lordkipanize D. Marzullo M. Prato O. Bagnasco Gianni G. Tecchiati U. Clavel B. Lepetz S. Davoudi H. Mashkour M. Berezina N.Y. Stockhammer P.W. Krause J. Haak W. Morales-Muñiz A. Benecke N. Hofreiter M. Ludwig A. Graphodatsky A.S. Peters J. Kiryushin K.Y. Iderkhangai T.-O. Bokovenko N.A. Vasiliev S.K. Seregin N.N. Chugunov K.V. Plasteeva N.A. Baryshnikov G.F. Petrova E. Sablin M. Ananyevskaya E. Logvin A. Shevnina I. Logvin V. Kalieva S. Loman V. Kukushkin I. Merz I. Merz V. Sakenov S. Varfolomeyev V. Usmanova E. Zaibert V. Arbuckle B. Belinskiy A.B. Kalmykov A. Reinhold S. Hansen S. Yudin A.I. Vybornov A.A. Epimakhov A. Berezina N.S. Roslyakova N. Kosintsev P.A. Kuznetsov P.F. Anthony D. Kroonen G.J. Kristiansen K. Wincker P. Outram A. Orlando L.
28 October 2021 Nature Research

Nature
2021 #598 Issue 7882 634 - 640 pp.

Domestication of horses fundamentally transformed long-range mobility and warfare¹. However, modern domesticated breeds do not descend from the earliest domestic horse lineage associated with archaeological evidence of bridling, milking and corralling^2–4 at Botai, Central Asia around 3500 bc³. Other longstanding candidate regions for horse domestication, such as Iberia⁵ and Anatolia⁶, have also recently been challenged. Thus, the genetic, geographic and temporal origins of modern domestic horses have remained unknown. Here we pinpoint the Western Eurasian steppes, especially the lower Volga-Don region, as the homeland of modern domestic horses. Furthermore, we map the population changes accompanying domestication from 273 ancient horse genomes. This reveals that modern domestic horses ultimately replaced almost all other local populations as they expanded rapidly across Eurasia from about 2000 bc, synchronously with equestrian material culture, including Sintashta spoke-wheeled chariots. We find that equestrianism involved strong selection for critical locomotor and behavioural adaptations at the GSDMC and ZFPM1 genes. Our results reject the commonly held association⁷ between horseback riding and the massive expansion of Yamnaya steppe pastoralists into Europe around 3000 bc^8,9 driving the spread of Indo-European languages¹⁰. This contrasts with the scenario in Asia where Indo-Iranian languages, chariots and horses spread together, following the early second millennium bc Sintashta culture^11,12.

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Centre d’Anthropobiologie et de Génomique de Toulouse, Université Paul Sabatier, Toulouse, France
Department of the Diversity and Evolution of Genomes, Institute of Molecular and Cellular Biology SB RAS, Novosibirsk, Russian Federation
W. Szafer Institute of Botany, Polish Academy of Sciences, Kraków, Poland
Genoscope, Institut de biologie François-Jacob, Commissariat à l’Energie Atomique (CEA), Université Paris-Saclay, Evry, France
Génomique Métabolique, Genoscope, Institut de biologie François Jacob, CEA, CNRS, Université d’Evry, Université Paris-Saclay, Evry, France
Earth System Science Department, University of California, Irvine, Irvine, CA, United States
Department of Ecology and Evolutionary Biology, University of California, Santa Cruz, Santa Cruz, CA, United States
Howard Hughes Medical Institute, University of California, Santa Cruz, Santa Cruz, CA, United States
Department of Archaeology, Ethnography and Museology, Altai State University, Barnaul, Russian Federation
Department of Archaeological Heritage Preservation, Institute of Archaeology of the Russian Academy of Sciences, Moscow, Russian Federation
Zoology Department, College of Science, King Saud University, Riyadh, Saudi Arabia
Institute for Archaeology, Heritage Conservation Studies and Art History, University of Bamberg, Bamberg, Germany
Museum Østjylland, Randers, Denmark
Saxo Institute, section of Archaeology, University of Copenhagen, Copenhagen, Denmark
ArScAn-UMR 7041, Equipe Ethnologie préhistorique, CNRS, MSH-Mondes, Nanterre Cedex, France
Muséum d’histoire naturelle, Secteur des Vertébrés, Geneva, Switzerland
UMR 5199 De la Préhistoire à l’Actuel : Culture, Environnement et Anthropologie (PACEA), CNRS, Université de Bordeaux, Pessac Cedex, France
Geneva Natural History Museum, Geneva, Switzerland
Department of Genetics and Evolution, University of Geneva, Geneva, Switzerland
OD Earth & History of Life, Royal Belgian Institute of Natural Sciences, Brussels, Belgium
Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Kraków, Poland
Institute of Archaeology and Ethnology Polish Academy of Sciences, Kraków, Poland
Institute of Archaeology, Jagiellonian University, Kraków, Poland
Department of Archaeology, Institute of History and Archaeology, Tartu, Estonia
Department of Zoology, Institute of Ecology and Earth Sciences, University of Tartu, Tartu, Estonia
Diamond and Precious Metals Geology Institute, SB RAS, Yakutsk, Russian Federation
Archaeological Research Collection, Tallinn University, Tallinn, Estonia
Department of Natural Sciences and Archaeometry, Institute of Archaeology of the Czech Academy of Sciences, Prague, Czech Republic
Prague, Czech Republic
Vasile Pârvan Institute of Archaeology, Department of Bioarchaeology, Romanian Academy, Bucharest, Romania
Institute of Archaeogenomics, Research Centre for the Humanities, Eötvös Loránd Research Network, Budapest, Hungary
Department of Genetics, Eötvös Loránd University, Budapest, Hungary
Institute of Archaeology, Research Centre for the Humanities, Eötvös Loránd Research Network, Budapest, Hungary
Ásatárs Ltd., Kecskemét, Hungary
Rippl-Rónai Municipal Museum with Country Scope, Kaposvár, Hungary
School of History, Classics and Archaeology, University of Edinburgh, Old Medical School, Edinburgh, United Kingdom
Trace and Environmental DNA (TrEnD) Lab, School of Molecular and Life Sciences, Curtin University, Perth, WA, Australia
Lundbeck Foundation GeoGenetics Centre, GLOBE Institute, University of Copenhagen, Copenhagen, Denmark
Department of Academic Management, Academy of Science of Moldova, Chișinău, Moldova
Center of Archaeology, Institute of Cultural Heritage, Academy of Science of Moldova, Chișinău, Moldova
Archaeological Institute of America, Boston, MA, United States
Centre National de Recherche Scientifique, Muséum national d’Histoire naturelle, Archéozoologie, Archéobotanique (AASPE), CP 56, Paris, France
Institute for the History of Material Culture, Russian Academy of Sciences (IHMC RAS), St Petersburg, Russian Federation
Geological Institute, Russian Academy of Sciences, Moscow, Russian Federation
Arctic and Antarctic Research Institute, St Petersburg, Russian Federation
Institute of Animal Breeding and Genetics, University of Veterinary Medicine Vienna, Vienna, Austria
Department of Prehistory and Western Asian/Northeast African Archaeology, Austrian Archaeological Institute, Austrian Academy of Sciences, Vienna, Austria
Estonian Biocentre, Institute of Genomics, University of Tartu, Tartu, Estonia
Department of Archaeogenetics, Max Planck Institute for the Science of Human History, Jena, Germany
SFB 1070 Resource Cultures, University of Tübingen, Tübingen, Germany
Department of Early Prehistory and Quaternary Ecology, University of Tübingen, Tübingen, Germany
UMR 7194 Muséum National d’Histoire Naturelle, CNRS, UPVD, Paris, France
Semenov-Tyan-Shanskii Lipetsk State Pedagogical University, Lipetsk, Russian Federation
Museum of Natural History, University of Colorado-Boulder, Boulder, CO, United States
Musée d’Anthropologie préhistorique de Monaco, Monaco, Monaco
Institute of Archaeology, Mongolian Academy of Sciences, Ulaanbaatar, Mongolia
Department of Archaeology, Max Planck Institute for the Science of Human History, Jena, Germany
Chinggis Khaan Museum, Ulaanbaatar, Mongolia
Department of Archaeology, Ulaanbaatar State University, Ulaanbaatar, Mongolia
Department of History, Kyrgyz-Turkish Manas University, Bishkek, Kyrgyzstan
Department of Biology, National University of Mongolia, Ulaanbaatar, Mongolia
Division of Archaeology, Biodiversity Institute, University of Kansas, Lawrence, KS, United States
Institute for Prehistoric and Protohistoric Archaeology, Kiel University, Kiel, Germany
ROOTS Excellence Cluster, Kiel University, Kiel, Germany
Archaeology of Social Dynamics, Institució Milà i Fontanals d’Humanitats, Consejo Superior de Investigaciones Científicas (IMF-CSIC), Barcelona, Spain
Departament d’Història i Arqueologia–SERP, Universitat de Barcelona, Barcelona, Spain
Grup d’Investigació Prehistòrica, Universitat de Lleida, PID2019-110022GB-I00, Lleida, Spain
Valencia, Spain
Departamento de Medicina Animal, Facultad de Veterinaria, Universidad de Extremadura, Cáceres, Spain
Centro Mixto UCM-ISCIII de Evolución y Comportamiento Humanos, Madrid, Spain
Instituto de Arqueología (CSIC–Junta de Extremadura), Mérida, Spain
Laboratori d’Arqueologia Prehistòrica, Universitat Jaume I, Castelló de la Plana, Spain
Departamento de Geodinámica, Estratigrafía y Paleontología, Facultad de Ciencias Geológicas, Universidad Complutense de Madrid, Madrid, Spain
Department of Eneolithic and Bronze Age, Institute of Archaeology National Academy of Sciences of Ukraine, Kyiv, Ukraine
Department of Archaeology, University of Exeter, Exeter, United Kingdom
Center for the Study of Human Origins, Anthropology Department, New York University, New York, NY, United States
Department of Archaeology, Ethnology and Museology, Al Farabi Kazakh National University, Almaty, Kazakhstan
Scientific Research Department, Orenburg State Pedagogical University, Orenburg, Russian Federation
Department of paleontology, Faculty of Geology, Moscow State University, Moscow, Russian Federation
Institute of Geology and Petroleum Technologies, Kazan Federal University, Kazan, Russian Federation
Georgian National Museum, Tbilisi, Georgia
Tbilisi State University, Tbilisi, Georgia
Università degli Studi di Milano, Dipartimento di Beni Culturali e Ambientali, Milan, Italy
University of Tehran, Central Laboratory, Bioarchaeology Laboratory, Archaeozoology Section, Tehran, Iran
Research Institute and Museum of Anthropology, Lomonosov Moscow State University, Moscow, Russian Federation
Department of Archaeogenetics, Max Planck Institute for Evolutionary Anthropology, Leipzig, Germany
Institute for Pre- and Protohistoric Archaeology and Archaeology of the Roman Provinces, Ludwig Maximilian University, Munich, Munich, Germany
School of Biological Sciences, The University of Adelaide, Adelaide, SA, Australia
Department of Biology, Universidad Autónoma de Madrid, Madrid, Spain
Eurasia Department of the German Archaeological Institute, Berlin, Germany
Evolutionary Adaptive Genomics, Institute of Biochemistry and Biology, Faculty of Mathematics and Science, University of Potsdam, Potsdam, Germany
Department of Evolutionary Genetics, Leibniz-Institute for Zoo and Wildlife Research, Berlin, Germany
Albrecht Daniel Thaer-Institute, Faculty of Life Sciences, Humboldt University Berlin, Berlin, Germany
ArchaeoBioCenter and Institute of Palaeoanatomy, Domestication Research and the History of Veterinary Medicine, LMU Munich, Munich, Germany
SNSB, State Collection of Anthropology and Palaeoanatomy, Munich, Germany
ArchaeoZOOlogy in Siberia and Central Asia—ZooSCAn International Research Laboratory, Institute of Archeology and Ethnography of the Siberian Branch of the RAS, Novosibirsk, Russian Federation
Department of Eastern European and Siberian Archaeology, State Hermitage Museum, St Petersburg, Russian Federation
Paleoecology Laboratory, Institute of Plant and Animal Ecology, Ural Branch of the Russian Academy of Sciences, Ekaterinburg, Russian Federation
Zoological Institute, Russian Academy of Sciences, St Petersburg, Russian Federation
Department of Archaeology, History Faculty, Vilnius University, Vilnius, Lithuania
Laboratory for Archaeological Research, Faculty of History and Law, Kostanay State University, Kostanay, Kazakhstan
Department of History and Archaeology, Surgut Governmental University, Surgut, Russian Federation
Saryarka Archaeological Institute, Buketov Karaganda University, Karaganda, Kazakhstan
Toraighyrov University, Joint Research Center for Archeological Studies, Pavlodar, Kazakhstan
Faculty of History, L. N. Gumilev Eurasian National University, Nur-Sultan, Kazakhstan
Institute of Archaeology and Steppe Civilizations, Al-Farabi Kazakh National University, Almaty, Kazakhstan
Department of Anthropology, Alumni Building, University of North Carolina at Chapel Hill, Chapel Hill, NC, United States
Nasledie Cultural Heritage Unit, Stavropol, Russian Federation
Research Center for the Preservation of Cultural Heritage, Saratov, Russian Federation
Department of Russian History and Archaeology, Samara State University of Social Sciences and Education, Samara, Russian Federation
Russian and Foreign History Department, South Ural State University, Chelyabinsk, Russian Federation
South Ural Department, Institute of History and Archaeology, Ural Branch of the Russian Academy of Sciences, Ekaterinburg, Russian Federation
Archaeological School, Chuvash State Institute of Humanities, Cheboksary, Russian Federation
Department of History of the Institute of Humanities, Ural Federal University, Ekaterinburg, Russian Federation
Department of Human Evolutionary Biology, Harvard University, Cambridge, MA, United States
Anthropology Faculty, Hartwick College, NY, Oneonta, United States
Department of Nordic Studies and Linguistics, University of Copenhagen, Copenhagen, Denmark
Leiden University Center for Linguistics, Leiden University, Leiden, Netherlands
Department of Historical Studies, University of Gothenburg, Gothenburg, Sweden
Lundbeck Foundation GeoGenetics Centre, Copenhagen, Denmark
Department of Biotechnology, Abdul Wali Khan University, Mardan, Pakistan

Centre d’Anthropobiologie et de Génomique de Toulouse
Department of the Diversity and Evolution of Genomes
W. Szafer Institute of Botany
Genoscope
Génomique Métabolique
Earth System Science Department
Department of Ecology and Evolutionary Biology
Howard Hughes Medical Institute
Department of Archaeology
Department of Archaeological Heritage Preservation
Zoology Department
Institute for Archaeology
Museum Østjylland
Saxo Institute
ArScAn-UMR 7041
Muséum d’histoire naturelle
UMR 5199 De la Préhistoire à l’Actuel : Culture
Geneva Natural History Museum
Department of Genetics and Evolution
OD Earth & History of Life
Institute of Systematics and Evolution of Animals
Institute of Archaeology and Ethnology Polish Academy of Sciences
Institute of Archaeology
Department of Archaeology
Department of Zoology
Diamond and Precious Metals Geology Institute
Archaeological Research Collection
Department of Natural Sciences and Archaeometry
Prague
Vasile Pârvan Institute of Archaeology
Institute of Archaeogenomics
Department of Genetics
Institute of Archaeology
Ásatárs Ltd.
Rippl-Rónai Municipal Museum with Country Scope
School of History
Trace and Environmental DNA (TrEnD) Lab
Lundbeck Foundation GeoGenetics Centre
Department of Academic Management
Center of Archaeology
Archaeological Institute of America
Centre National de Recherche Scientifique
Institute for the History of Material Culture
Geological Institute
Arctic and Antarctic Research Institute
Institute of Animal Breeding and Genetics
Department of Prehistory and Western Asian/Northeast African Archaeology
Estonian Biocentre
Department of Archaeogenetics
SFB 1070 Resource Cultures
Department of Early Prehistory and Quaternary Ecology
UMR 7194 Muséum National d’Histoire Naturelle
Semenov-Tyan-Shanskii Lipetsk State Pedagogical University
Museum of Natural History
Musée d’Anthropologie préhistorique de Monaco
Institute of Archaeology
Department of Archaeology
Chinggis Khaan Museum
Department of Archaeology
Department of History
Department of Biology
Division of Archaeology
Institute for Prehistoric and Protohistoric Archaeology
ROOTS Excellence Cluster
Archaeology of Social Dynamics
Departament d’Història i Arqueologia–SERP
Grup d’Investigació Prehistòrica
Valencia
Departamento de Medicina Animal
Centro Mixto UCM-ISCIII de Evolución y Comportamiento Humanos
Instituto de Arqueología (CSIC–Junta de Extremadura)
Laboratori d’Arqueologia Prehistòrica
Departamento de Geodinámica
Department of Eneolithic and Bronze Age
Department of Archaeology
Center for the Study of Human Origins
Department of Archaeology
Scientific Research Department
Department of paleontology
Institute of Geology and Petroleum Technologies
Georgian National Museum
Tbilisi State University
Università degli Studi di Milano
University of Tehran
Research Institute and Museum of Anthropology
Department of Archaeogenetics
Institute for Pre- and Protohistoric Archaeology and Archaeology of the Roman Provinces
School of Biological Sciences
Department of Biology
Eurasia Department of the German Archaeological Institute
Evolutionary Adaptive Genomics
Department of Evolutionary Genetics
Albrecht Daniel Thaer-Institute
ArchaeoBioCenter and Institute of Palaeoanatomy
SNSB
ArchaeoZOOlogy in Siberia and Central Asia—ZooSCAn International Research Laboratory
Department of Eastern European and Siberian Archaeology
Paleoecology Laboratory
Zoological Institute
Department of Archaeology
Laboratory for Archaeological Research
Department of History and Archaeology
Saryarka Archaeological Institute
Toraighyrov University
Faculty of History
Institute of Archaeology and Steppe Civilizations
Department of Anthropology
Nasledie Cultural Heritage Unit
Research Center for the Preservation of Cultural Heritage
Department of Russian History and Archaeology
Russian and Foreign History Department
South Ural Department
Archaeological School
Department of History of the Institute of Humanities
Department of Human Evolutionary Biology
Anthropology Faculty
Department of Nordic Studies and Linguistics
Leiden University Center for Linguistics
Department of Historical Studies
Lundbeck Foundation GeoGenetics Centre
Department of Biotechnology

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