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

Pablo Librado, Naveed Khan, Antoine Fages, Mariya A. Kusliy, Tomasz Suchan, Laure Tonasso-Calvière, Stéphanie Schiavinato, Duha Alioglu, Aurore Fromentier, Aude Perdereau, Jean-Marc Aury, Charleen Gaunitz, Lorelei Chauvey, Andaine Seguin-Orlando, Clio Der Sarkissian, John Southon, Beth Shapiro, Alexey A. Tishkin, Alexey A. Kovalev, Saleh Alquraishi, Ahmed H. Alfarhan, Khaled A. S. Al-Rasheid, Timo Seregély, Lutz Klassen, Rune Iversen, Olivier Bignon-Lau, Pierre Bodu, Monique Olive, Jean-Christophe Castel, Myriam Boudadi-Maligne, Nadir Alvarez, Mietje Germonpré, Magdalena Moskal-del Hoyo, Jarosław Wilczyński, Sylwia Pospuła, Anna Lasota-Kuś, Krzysztof Tunia, Marek Nowak, Eve Rannamäe, Urmas Saarma, Gennady Boeskorov, Lembi Lōugas, René Kyselý, Lubomír Peške, Adrian Bălășescu, Valentin Dumitrașcu, Roxana Dobrescu, Daniel Gerber, Viktória Kiss, Anna Szécsényi-Nagy, Balázs G. Mende, Zsolt Gallina, Krisztina Somogyi, Gabriella Kulcsár, Erika Gál, Robin Bendrey, Morten E. Allentoft, Ghenadie Sirbu, Valentin Dergachev, Henry Shephard, Noémie Tomadini, Sandrine Grouard, Aleksei Kasparov, Alexander E. Basilyan, Mikhail A. Anisimov, Pavel A. Nikolskiy, Elena Y. Pavlova, Vladimir Pitulko, Gottfried Brem, Barbara Wallner, Christoph Schwall, Marcel Keller, Keiko Kitagawa, Alexander N. Bessudnov, Alexander Bessudnov, William Taylor, Jérome Magail, Jamiyan-Ombo Gantulga, Jamsranjav Bayarsaikhan, Diimaajav Erdenebaatar, Kubatbeek Tabaldiev, Enkhbayar Mijiddorj, Bazartseren Boldgiv, Turbat Tsagaan, Mélanie Pruvost, Sandra Olsen, Cheryl A. Makarewicz, Silvia Valenzuela Lamas, Silvia Albizuri Canadell, Ariadna Nieto Espinet, Ma Pilar Iborra, Jaime Lira Garrido, Esther Rodríguez González, Sebastián Celestino, Carmen Olària, Juan Luis Arsuaga, Nadiia Kotova, Alexander Pryor, Pam Crabtree, Rinat Zhumatayev, Abdesh Toleubaev, Nina L. Morgunova, Tatiana Kuznetsova, David Lordkipanize, Matilde Marzullo, Ornella Prato, Giovanna Bagnasco Gianni, Umberto Tecchiati, Benoit Clavel, Sébastien Lepetz, Hossein Davoudi, Marjan Mashkour, Natalia Ya. Berezina, Philipp W. Stockhammer, Johannes Krause, Wolfgang Haak, Arturo Morales-Muñiz, Norbert Benecke, Michael Hofreiter, Arne Ludwig, Alexander S. Graphodatsky, Joris Peters, Kirill Yu. Kiryushin, Tumur-Ochir Iderkhangai, Nikolay A. Bokovenko, Sergey K. Vasiliev, Nikolai N. Seregin, Konstantin V. Chugunov, Natalya A. Plasteeva, Gennady F. Baryshnikov, Ekaterina Petrova, Mikhail Sablin, Elina Ananyevskaya, Andrey Logvin, Irina Shevnina, Victor Logvin, Saule Kalieva, Valeriy Loman, Igor Kukushkin, Ilya Merz, Victor Merz, Sergazy Sakenov, Victor Varfolomeyev, Emma Usmanova, Viktor Zaibert, Benjamin Arbuckle, Andrey B. Belinskiy, Alexej Kalmykov, Sabine Reinhold, Svend Hansen, Aleksandr I. Yudin, Alekandr A. Vybornov, Andrey Epimakhov, Natalia S. Berezina, Natalia Roslyakova, Pavel A. Kosintsev, Pavel F. Kuznetsov, David Anthony, Guus J. Kroonen, Kristian Kristiansen, Patrick Wincker, Alan Outram and Ludovic Orlando ()
Additional contact information
Pablo Librado: Université Paul Sabatier
Naveed Khan: Université Paul Sabatier
Antoine Fages: Université Paul Sabatier
Mariya A. Kusliy: Université Paul Sabatier
Tomasz Suchan: Université Paul Sabatier
Laure Tonasso-Calvière: Université Paul Sabatier
Stéphanie Schiavinato: Université Paul Sabatier
Duha Alioglu: Université Paul Sabatier
Aurore Fromentier: Université Paul Sabatier
Aude Perdereau: Université Paris-Saclay
Jean-Marc Aury: Université d’Evry, Université Paris-Saclay
Charleen Gaunitz: Université Paul Sabatier
Lorelei Chauvey: Université Paul Sabatier
Andaine Seguin-Orlando: Université Paul Sabatier
Clio Der Sarkissian: Université Paul Sabatier
John Southon: University of California, Irvine
Beth Shapiro: University of California, Santa Cruz
Alexey A. Tishkin: Altai State University
Alexey A. Kovalev: Institute of Archaeology of the Russian Academy of Sciences
Saleh Alquraishi: King Saud University
Ahmed H. Alfarhan: King Saud University
Khaled A. S. Al-Rasheid: King Saud University
Timo Seregély: University of Bamberg
Lutz Klassen: Museum Østjylland
Rune Iversen: University of Copenhagen
Olivier Bignon-Lau: ArScAn-UMR 7041, Equipe Ethnologie préhistorique, CNRS, MSH-Mondes
Pierre Bodu: ArScAn-UMR 7041, Equipe Ethnologie préhistorique, CNRS, MSH-Mondes
Monique Olive: ArScAn-UMR 7041, Equipe Ethnologie préhistorique, CNRS, MSH-Mondes
Jean-Christophe Castel: Muséum d’histoire naturelle, Secteur des Vertébrés
Myriam Boudadi-Maligne: Université de Bordeaux
Nadir Alvarez: Geneva Natural History Museum
Mietje Germonpré: Royal Belgian Institute of Natural Sciences
Magdalena Moskal-del Hoyo: Polish Academy of Sciences
Jarosław Wilczyński: Polish Academy of Sciences
Sylwia Pospuła: Polish Academy of Sciences
Anna Lasota-Kuś: Institute of Archaeology and Ethnology Polish Academy of Sciences
Krzysztof Tunia: Institute of Archaeology and Ethnology Polish Academy of Sciences
Marek Nowak: Jagiellonian University
Eve Rannamäe: Institute of History and Archaeology
Urmas Saarma: University of Tartu
Gennady Boeskorov: Diamond and Precious Metals Geology Institute, SB RAS
Lembi Lōugas: Tallinn University
René Kyselý: Institute of Archaeology of the Czech Academy of Sciences
Adrian Bălășescu: Romanian Academy
Valentin Dumitrașcu: Romanian Academy
Roxana Dobrescu: Romanian Academy
Daniel Gerber: Eötvös Loránd Research Network
Viktória Kiss: Eötvös Loránd Research Network
Anna Szécsényi-Nagy: Eötvös Loránd Research Network
Balázs G. Mende: Eötvös Loránd Research Network
Zsolt Gallina: Ásatárs Ltd.
Krisztina Somogyi: Rippl-Rónai Municipal Museum with Country Scope
Gabriella Kulcsár: Eötvös Loránd Research Network
Erika Gál: Eötvös Loránd Research Network
Robin Bendrey: University of Edinburgh, Old Medical School
Morten E. Allentoft: Curtin University
Ghenadie Sirbu: Academy of Science of Moldova
Valentin Dergachev: Academy of Science of Moldova
Henry Shephard: Archaeological Institute of America
Noémie Tomadini: Centre National de Recherche Scientifique, Muséum national d’Histoire naturelle, Archéozoologie, Archéobotanique (AASPE), CP 56
Sandrine Grouard: Centre National de Recherche Scientifique, Muséum national d’Histoire naturelle, Archéozoologie, Archéobotanique (AASPE), CP 56
Aleksei Kasparov: Institute for the History of Material Culture, Russian Academy of Sciences (IHMC RAS)
Alexander E. Basilyan: Geological Institute, Russian Academy of Sciences
Mikhail A. Anisimov: Arctic and Antarctic Research Institute
Pavel A. Nikolskiy: Geological Institute, Russian Academy of Sciences
Elena Y. Pavlova: Arctic and Antarctic Research Institute
Vladimir Pitulko: Institute for the History of Material Culture, Russian Academy of Sciences (IHMC RAS)
Gottfried Brem: University of Veterinary Medicine Vienna
Barbara Wallner: University of Veterinary Medicine Vienna
Christoph Schwall: Austrian Archaeological Institute, Austrian Academy of Sciences
Marcel Keller: University of Tartu
Keiko Kitagawa: University of Tübingen
Alexander N. Bessudnov: Semenov-Tyan-Shanskii Lipetsk State Pedagogical University
Alexander Bessudnov: Institute for the History of Material Culture, Russian Academy of Sciences (IHMC RAS)
William Taylor: University of Colorado-Boulder
Jérome Magail: Musée d’Anthropologie préhistorique de Monaco
Jamiyan-Ombo Gantulga: Institute of Archaeology, Mongolian Academy of Sciences
Jamsranjav Bayarsaikhan: Max Planck Institute for the Science of Human History
Diimaajav Erdenebaatar: Ulaanbaatar State University
Kubatbeek Tabaldiev: Kyrgyz-Turkish Manas University
Enkhbayar Mijiddorj: Ulaanbaatar State University
Bazartseren Boldgiv: National University of Mongolia
Turbat Tsagaan: Institute of Archaeology, Mongolian Academy of Sciences
Mélanie Pruvost: Université de Bordeaux
Sandra Olsen: University of Kansas
Cheryl A. Makarewicz: Kiel University
Silvia Valenzuela Lamas: Archaeology of Social Dynamics, Institució Milà i Fontanals d’Humanitats, Consejo Superior de Investigaciones Científicas (IMF-CSIC)
Silvia Albizuri Canadell: Universitat de Barcelona
Ariadna Nieto Espinet: Universitat de Lleida
Jaime Lira Garrido: Universidad de Extremadura
Esther Rodríguez González: Instituto de Arqueología (CSIC–Junta de Extremadura)
Sebastián Celestino: Instituto de Arqueología (CSIC–Junta de Extremadura)
Carmen Olària: Universitat Jaume I
Juan Luis Arsuaga: Centro Mixto UCM-ISCIII de Evolución y Comportamiento Humanos
Nadiia Kotova: Institute of Archaeology National Academy of Sciences of Ukraine
Alexander Pryor: University of Exeter
Pam Crabtree: New York University
Rinat Zhumatayev: Al Farabi Kazakh National University
Abdesh Toleubaev: Al Farabi Kazakh National University
Nina L. Morgunova: Orenburg State Pedagogical University
Tatiana Kuznetsova: Faculty of Geology, Moscow State University
David Lordkipanize: Georgian National Museum
Matilde Marzullo: Università degli Studi di Milano, Dipartimento di Beni Culturali e Ambientali
Ornella Prato: Università degli Studi di Milano, Dipartimento di Beni Culturali e Ambientali
Giovanna Bagnasco Gianni: Università degli Studi di Milano, Dipartimento di Beni Culturali e Ambientali
Umberto Tecchiati: Università degli Studi di Milano, Dipartimento di Beni Culturali e Ambientali
Benoit Clavel: Centre National de Recherche Scientifique, Muséum national d’Histoire naturelle, Archéozoologie, Archéobotanique (AASPE), CP 56
Sébastien Lepetz: Centre National de Recherche Scientifique, Muséum national d’Histoire naturelle, Archéozoologie, Archéobotanique (AASPE), CP 56
Hossein Davoudi: University of Tehran, Central Laboratory, Bioarchaeology Laboratory, Archaeozoology Section
Marjan Mashkour: Centre National de Recherche Scientifique, Muséum national d’Histoire naturelle, Archéozoologie, Archéobotanique (AASPE), CP 56
Natalia Ya. Berezina: Lomonosov Moscow State University
Philipp W. Stockhammer: Max Planck Institute for Evolutionary Anthropology
Johannes Krause: Max Planck Institute for the Science of Human History
Wolfgang Haak: Max Planck Institute for the Science of Human History
Arturo Morales-Muñiz: Universidad Autónoma de Madrid
Norbert Benecke: Eurasia Department of the German Archaeological Institute
Michael Hofreiter: University of Potsdam
Arne Ludwig: Leibniz-Institute for Zoo and Wildlife Research
Alexander S. Graphodatsky: Institute of Molecular and Cellular Biology SB RAS
Joris Peters: LMU Munich
Kirill Yu. Kiryushin: Altai State University
Tumur-Ochir Iderkhangai: Ulaanbaatar State University
Nikolay A. Bokovenko: Institute for the History of Material Culture, Russian Academy of Sciences (IHMC RAS)
Sergey K. Vasiliev: ArchaeoZOOlogy in Siberia and Central Asia—ZooSCAn International Research Laboratory, Institute of Archeology and Ethnography of the Siberian Branch of the RAS
Nikolai N. Seregin: Altai State University
Konstantin V. Chugunov: State Hermitage Museum
Natalya A. Plasteeva: Paleoecology Laboratory, Institute of Plant and Animal Ecology, Ural Branch of the Russian Academy of Sciences
Gennady F. Baryshnikov: Zoological Institute, Russian Academy of Sciences
Ekaterina Petrova: Vilnius University
Mikhail Sablin: Zoological Institute, Russian Academy of Sciences
Elina Ananyevskaya: Vilnius University
Andrey Logvin: Kostanay State University
Irina Shevnina: Kostanay State University
Victor Logvin: Surgut Governmental University
Saule Kalieva: Surgut Governmental University
Valeriy Loman: Buketov Karaganda University
Igor Kukushkin: Buketov Karaganda University
Ilya Merz: Toraighyrov University, Joint Research Center for Archeological Studies
Victor Merz: Toraighyrov University, Joint Research Center for Archeological Studies
Sergazy Sakenov: Faculty of History, L. N. Gumilev Eurasian National University
Victor Varfolomeyev: Buketov Karaganda University
Emma Usmanova: Buketov Karaganda University
Viktor Zaibert: Al-Farabi Kazakh National University
Benjamin Arbuckle: University of North Carolina at Chapel Hill
Andrey B. Belinskiy: Nasledie Cultural Heritage Unit
Alexej Kalmykov: Nasledie Cultural Heritage Unit
Sabine Reinhold: Eurasia Department of the German Archaeological Institute
Svend Hansen: Eurasia Department of the German Archaeological Institute
Aleksandr I. Yudin: Research Center for the Preservation of Cultural Heritage
Alekandr A. Vybornov: Samara State University of Social Sciences and Education
Andrey Epimakhov: South Ural State University
Natalia S. Berezina: Archaeological School, Chuvash State Institute of Humanities
Natalia Roslyakova: Samara State University of Social Sciences and Education
Pavel A. Kosintsev: Paleoecology Laboratory, Institute of Plant and Animal Ecology, Ural Branch of the Russian Academy of Sciences
Pavel F. Kuznetsov: Samara State University of Social Sciences and Education
David Anthony: Harvard University
Guus J. Kroonen: University of Copenhagen
Kristian Kristiansen: University of Gothenburg
Patrick Wincker: Université d’Evry, Université Paris-Saclay
Alan Outram: University of Exeter
Ludovic Orlando: Université Paul Sabatier

Nature, 2021, vol. 598, issue 7882, 634-640

Abstract: Abstract 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 corralling2–4 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 bc8,9 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 culture11,12.

Date: 2021
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DOI: 10.1038/s41586-021-04018-9

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