Analyze Diet
Nature2021; 598(7882); 634-640; doi: 10.1038/s41586-021-04018-9

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

Abstract: Domestication of horses fundamentally transformed long-range mobility and warfare. However, modern domesticated breeds do not descend from the earliest domestic horse lineage associated with archaeological evidence of bridling, milking and corralling at Botai, Central Asia around 3500 BC. Other longstanding candidate regions for horse domestication, such as Iberia and Anatolia, 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 association between horseback riding and the massive expansion of Yamnaya steppe pastoralists into Europe around 3000 BC driving the spread of Indo-European languages. This contrasts with the scenario in Asia where Indo-Iranian languages, chariots and horses spread together, following the early second millennium BC Sintashta culture.
Publication Date: 2021-10-20 PubMed ID: 34671162PubMed Central: PMC8550961DOI: 10.1038/s41586-021-04018-9Google 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.
  • Journal Article
  • Research Support
  • Non-U.S. Gov't

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.

This research article explores the origins and spread of domestic horses, tracing their homeland to the Western Eurasian steppes, especially the lower Volga-Don region. The study disputes previously held beliefs, detailing that the domestication of horses actually coincided with the rapid spread of these animals across Eurasia around 2000 BC, alongside the rise of equestrian culture.

Background and Purpose of the Study

  • The study sets out to unravel mysteries around the domestication of horses, an event which greatly altered long-range mobility and warfare. The origins of modern domestic horses have remained unclear, with evidence contradicting early theories about regions like Botai, Central Asia.
  • The researchers aim to determine the genetic, geographic, and temporal origins of these horses, as well as mapping the population changes that came with domestication.

Findings of the Study

  • The study pinpoints the Western Eurasian steppes, particularly the lower Volga-Don region, as the likely homeland of modern domestic horses.
  • It analyses 273 ancient horse genomes to illustrate population changes following domestication.
  • The findings suggest that domestic horses replaced virtually all other local populations as they swiftly expanded across Eurasia from about 2000 BC, synchronously with the rise of equestrian material culture.
  • Strong selection for crucial locomotor and behavioural adaptations at the GSDMC and ZFPM1 genes were noted, indicating a strong focus on equestrianism as part of this widespread cultural transition.

Implications and Conclusions

  • The research contradicts the often-accepted relationship between horseback riding and the large-scale expansion of Yamnaya steppe pastoralists into Europe around 3000 BC, which was previously thought to have carried the spread of Indo-European languages.
  • This contrasts with the scenario in Asia, where Indo-Iranian languages, chariots, and horses spread together following the early second millennium BC Sintashta culture.
  • The researchers’ findings offer significant insight into the historical interaction between human cultures and domesticated animals, in this case highlighting horses’ central role in the development and spread of significant aspects of human culture and travel.

Cite This Article

APA
Librado P, Khan N, Fages A, Kusliy MA, Suchan T, Tonasso-Calvière L, Schiavinato S, Alioglu D, Fromentier A, Perdereau A, Aury JM, Gaunitz C, Chauvey L, Seguin-Orlando A, Der Sarkissian C, Southon J, Shapiro B, Tishkin AA, Kovalev AA, Alquraishi S, Alfarhan AH, Al-Rasheid KAS, Seregély T, Klassen L, Iversen R, Bignon-Lau O, Bodu P, Olive M, Castel JC, 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 BG, Gallina Z, Somogyi K, Kulcsár G, Gál E, Bendrey R, Allentoft ME, Sirbu G, Dergachev V, Shephard H, Tomadini N, Grouard S, Kasparov A, Basilyan AE, Anisimov MA, Nikolskiy PA, Pavlova EY, Pitulko V, Brem G, Wallner B, Schwall C, Keller M, Kitagawa K, Bessudnov AN, Bessudnov A, Taylor W, Magail J, Gantulga JO, Bayarsaikhan J, Erdenebaatar D, Tabaldiev K, Mijiddorj E, Boldgiv B, Tsagaan T, Pruvost M, Olsen S, Makarewicz CA, Valenzuela Lamas S, Albizuri Canadell S, Nieto Espinet A, Iborra MP, Lira Garrido J, Rodríguez González E, Celestino S, Olària C, Arsuaga JL, Kotova N, Pryor A, Crabtree P, Zhumatayev R, Toleubaev A, Morgunova NL, Kuznetsova T, Lordkipanize D, Marzullo M, Prato O, Bagnasco Gianni G, Tecchiati U, Clavel B, Lepetz S, Davoudi H, Mashkour M, Berezina NY, Stockhammer PW, Krause J, Haak W, Morales-Muñiz A, Benecke N, Hofreiter M, Ludwig A, Graphodatsky AS, Peters J, Kiryushin KY, Iderkhangai TO, Bokovenko NA, Vasiliev SK, Seregin NN, Chugunov KV, Plasteeva NA, Baryshnikov GF, 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 AB, Kalmykov A, Reinhold S, Hansen S, Yudin AI, Vybornov AA, Epimakhov A, Berezina NS, Roslyakova N, Kosintsev PA, Kuznetsov PF, Anthony D, Kroonen GJ, Kristiansen K, Wincker P, Outram A, Orlando L. (2021). The origins and spread of domestic horses from the Western Eurasian steppes. Nature, 598(7882), 634-640. https://doi.org/10.1038/s41586-021-04018-9

Publication

ISSN: 1476-4687
NlmUniqueID: 0410462
Country: England
Language: English
Volume: 598
Issue: 7882
Pages: 634-640

Researcher Affiliations

Librado, Pablo
  • Centre d'Anthropobiologie et de Génomique de Toulouse, Université Paul Sabatier, Toulouse, France.
Khan, Naveed
  • Centre d'Anthropobiologie et de Génomique de Toulouse, Université Paul Sabatier, Toulouse, France.
  • Department of Biotechnology, Abdul Wali Khan University, Mardan, Pakistan.
Fages, Antoine
  • Centre d'Anthropobiologie et de Génomique de Toulouse, Université Paul Sabatier, Toulouse, France.
Kusliy, Mariya A
  • 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, Russia.
Suchan, Tomasz
  • Centre d'Anthropobiologie et de Génomique de Toulouse, Université Paul Sabatier, Toulouse, France.
  • W. Szafer Institute of Botany, Polish Academy of Sciences, Kraków, Poland.
Tonasso-Calvière, Laure
  • Centre d'Anthropobiologie et de Génomique de Toulouse, Université Paul Sabatier, Toulouse, France.
Schiavinato, Stéphanie
  • Centre d'Anthropobiologie et de Génomique de Toulouse, Université Paul Sabatier, Toulouse, France.
Alioglu, Duha
  • Centre d'Anthropobiologie et de Génomique de Toulouse, Université Paul Sabatier, Toulouse, France.
Fromentier, Aurore
  • Centre d'Anthropobiologie et de Génomique de Toulouse, Université Paul Sabatier, Toulouse, France.
Perdereau, Aude
  • Genoscope, Institut de biologie François-Jacob, Commissariat à l'Energie Atomique (CEA), Université Paris-Saclay, Evry, France.
Aury, Jean-Marc
  • Génomique Métabolique, Genoscope, Institut de biologie François Jacob, CEA, CNRS, Université d'Evry, Université Paris-Saclay, Evry, France.
Gaunitz, Charleen
  • Centre d'Anthropobiologie et de Génomique de Toulouse, Université Paul Sabatier, Toulouse, France.
Chauvey, Lorelei
  • Centre d'Anthropobiologie et de Génomique de Toulouse, Université Paul Sabatier, Toulouse, France.
Seguin-Orlando, Andaine
  • Centre d'Anthropobiologie et de Génomique de Toulouse, Université Paul Sabatier, Toulouse, France.
Der Sarkissian, Clio
  • Centre d'Anthropobiologie et de Génomique de Toulouse, Université Paul Sabatier, Toulouse, France.
Southon, John
  • Earth System Science Department, University of California, Irvine, Irvine, CA, USA.
Shapiro, Beth
  • Department of Ecology and Evolutionary Biology, University of California, Santa Cruz, Santa Cruz, CA, USA.
  • Howard Hughes Medical Institute, University of California, Santa Cruz, Santa Cruz, CA, USA.
Tishkin, Alexey A
  • Department of Archaeology, Ethnography and Museology, Altai State University, Barnaul, Russia.
Kovalev, Alexey A
  • Department of Archaeological Heritage Preservation, Institute of Archaeology of the Russian Academy of Sciences, Moscow, Russia.
Alquraishi, Saleh
  • Zoology Department, College of Science, King Saud University, Riyadh, Saudi Arabia.
Alfarhan, Ahmed H
  • Zoology Department, College of Science, King Saud University, Riyadh, Saudi Arabia.
Al-Rasheid, Khaled A S
  • Zoology Department, College of Science, King Saud University, Riyadh, Saudi Arabia.
Seregély, Timo
  • Institute for Archaeology, Heritage Conservation Studies and Art History, University of Bamberg, Bamberg, Germany.
Klassen, Lutz
  • Museum Østjylland, Randers, Denmark.
Iversen, Rune
  • Saxo Institute, section of Archaeology, University of Copenhagen, Copenhagen, Denmark.
Bignon-Lau, Olivier
  • ArScAn-UMR 7041, Equipe Ethnologie préhistorique, CNRS, MSH-Mondes, Nanterre Cedex, France.
Bodu, Pierre
  • ArScAn-UMR 7041, Equipe Ethnologie préhistorique, CNRS, MSH-Mondes, Nanterre Cedex, France.
Olive, Monique
  • ArScAn-UMR 7041, Equipe Ethnologie préhistorique, CNRS, MSH-Mondes, Nanterre Cedex, France.
Castel, Jean-Christophe
  • Muséum d'histoire naturelle, Secteur des Vertébrés, Geneva, Switzerland.
Boudadi-Maligne, Myriam
  • UMR 5199 De la Préhistoire à l'Actuel : Culture, Environnement et Anthropologie (PACEA), CNRS, Université de Bordeaux, Pessac Cedex, France.
Alvarez, Nadir
  • Geneva Natural History Museum, Geneva, Switzerland.
  • Department of Genetics and Evolution, University of Geneva, Geneva, Switzerland.
Germonpré, Mietje
  • OD Earth & History of Life, Royal Belgian Institute of Natural Sciences, Brussels, Belgium.
Moskal-Del Hoyo, Magdalena
  • W. Szafer Institute of Botany, Polish Academy of Sciences, Kraków, Poland.
Wilczyński, Jarosław
  • Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Kraków, Poland.
Pospuła, Sylwia
  • Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Kraków, Poland.
Lasota-Kuś, Anna
  • Institute of Archaeology and Ethnology Polish Academy of Sciences, Kraków, Poland.
Tunia, Krzysztof
  • Institute of Archaeology and Ethnology Polish Academy of Sciences, Kraków, Poland.
Nowak, Marek
  • Institute of Archaeology, Jagiellonian University, Kraków, Poland.
Rannamäe, Eve
  • Department of Archaeology, Institute of History and Archaeology, Tartu, Estonia.
Saarma, Urmas
  • Department of Zoology, Institute of Ecology and Earth Sciences, University of Tartu, Tartu, Estonia.
Boeskorov, Gennady
  • Diamond and Precious Metals Geology Institute, SB RAS, Yakutsk, Russia.
Lōugas, Lembi
  • Archaeological Research Collection, Tallinn University, Tallinn, Estonia.
Kyselý, René
  • Department of Natural Sciences and Archaeometry, Institute of Archaeology of the Czech Academy of Sciences, Prague, Czechia.
Peške, Lubomír
  • , Prague, Czechia.
Bălășescu, Adrian
  • Vasile Pârvan Institute of Archaeology, Department of Bioarchaeology, Romanian Academy, Bucharest, Romania.
Dumitrașcu, Valentin
  • Vasile Pârvan Institute of Archaeology, Department of Bioarchaeology, Romanian Academy, Bucharest, Romania.
Dobrescu, Roxana
  • Vasile Pârvan Institute of Archaeology, Department of Bioarchaeology, Romanian Academy, Bucharest, Romania.
Gerber, Daniel
  • Institute of Archaeogenomics, Research Centre for the Humanities, Eötvös Loránd Research Network, Budapest, Hungary.
  • Department of Genetics, Eötvös Loránd University, Budapest, Hungary.
Kiss, Viktória
  • Institute of Archaeology, Research Centre for the Humanities, Eötvös Loránd Research Network, Budapest, Hungary.
Szécsényi-Nagy, Anna
  • Institute of Archaeogenomics, Research Centre for the Humanities, Eötvös Loránd Research Network, Budapest, Hungary.
Mende, Balázs G
  • Institute of Archaeogenomics, Research Centre for the Humanities, Eötvös Loránd Research Network, Budapest, Hungary.
Gallina, Zsolt
  • Ásatárs Ltd., Kecskemét, Hungary.
Somogyi, Krisztina
  • Rippl-Rónai Municipal Museum with Country Scope, Kaposvár, Hungary.
Kulcsár, Gabriella
  • Institute of Archaeology, Research Centre for the Humanities, Eötvös Loránd Research Network, Budapest, Hungary.
Gál, Erika
  • Institute of Archaeology, Research Centre for the Humanities, Eötvös Loránd Research Network, Budapest, Hungary.
Bendrey, Robin
  • School of History, Classics and Archaeology, University of Edinburgh, Old Medical School, Edinburgh, UK.
Allentoft, Morten E
  • Trace and Environmental DNA (TrEnD) Lab, School of Molecular and Life Sciences, Curtin University, Perth, Western Australia, Australia.
  • Lundbeck Foundation GeoGenetics Centre, GLOBE Institute, University of Copenhagen, Copenhagen, Denmark.
Sirbu, Ghenadie
  • Department of Academic Management, Academy of Science of Moldova, Chișinău, Republic of Moldova.
Dergachev, Valentin
  • Center of Archaeology, Institute of Cultural Heritage, Academy of Science of Moldova, Chișinău, Republic of Moldova.
Shephard, Henry
  • Archaeological Institute of America, Boston, MA, USA.
Tomadini, Noémie
  • Centre National de Recherche Scientifique, Muséum national d'Histoire naturelle, Archéozoologie, Archéobotanique (AASPE), CP 56, Paris, France.
Grouard, Sandrine
  • Centre National de Recherche Scientifique, Muséum national d'Histoire naturelle, Archéozoologie, Archéobotanique (AASPE), CP 56, Paris, France.
Kasparov, Aleksei
  • Institute for the History of Material Culture, Russian Academy of Sciences (IHMC RAS), St Petersburg, Russia.
Basilyan, Alexander E
  • Geological Institute, Russian Academy of Sciences, Moscow, Russia.
Anisimov, Mikhail A
  • Arctic and Antarctic Research Institute, St Petersburg, Russia.
Nikolskiy, Pavel A
  • Geological Institute, Russian Academy of Sciences, Moscow, Russia.
Pavlova, Elena Y
  • Arctic and Antarctic Research Institute, St Petersburg, Russia.
Pitulko, Vladimir
  • Institute for the History of Material Culture, Russian Academy of Sciences (IHMC RAS), St Petersburg, Russia.
Brem, Gottfried
  • Institute of Animal Breeding and Genetics, University of Veterinary Medicine Vienna, Vienna, Austria.
Wallner, Barbara
  • Institute of Animal Breeding and Genetics, University of Veterinary Medicine Vienna, Vienna, Austria.
Schwall, Christoph
  • Department of Prehistory and Western Asian/Northeast African Archaeology, Austrian Archaeological Institute, Austrian Academy of Sciences, Vienna, Austria.
Keller, Marcel
  • Estonian Biocentre, Institute of Genomics, University of Tartu, Tartu, Estonia.
  • Department of Archaeogenetics, Max Planck Institute for the Science of Human History, Jena, Germany.
Kitagawa, Keiko
  • 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.
Bessudnov, Alexander N
  • Semenov-Tyan-Shanskii Lipetsk State Pedagogical University, Lipetsk, Russia.
Bessudnov, Alexander
  • Institute for the History of Material Culture, Russian Academy of Sciences (IHMC RAS), St Petersburg, Russia.
Taylor, William
  • Museum of Natural History, University of Colorado-Boulder, Boulder, CO, USA.
Magail, Jérome
  • Musée d'Anthropologie préhistorique de Monaco, Monaco, Monaco.
Gantulga, Jamiyan-Ombo
  • Institute of Archaeology, Mongolian Academy of Sciences, Ulaanbaatar, Mongolia.
Bayarsaikhan, Jamsranjav
  • Department of Archaeology, Max Planck Institute for the Science of Human History, Jena, Germany.
  • Chinggis Khaan Museum, Ulaanbaatar, Mongolia.
Erdenebaatar, Diimaajav
  • Department of Archaeology, Ulaanbaatar State University, Ulaanbaatar, Mongolia.
Tabaldiev, Kubatbeek
  • Department of History, Kyrgyz-Turkish Manas University, Bishkek, Kyrgyzstan.
Mijiddorj, Enkhbayar
  • Department of Archaeology, Ulaanbaatar State University, Ulaanbaatar, Mongolia.
Boldgiv, Bazartseren
  • Department of Biology, National University of Mongolia, Ulaanbaatar, Mongolia.
Tsagaan, Turbat
  • Institute of Archaeology, Mongolian Academy of Sciences, Ulaanbaatar, Mongolia.
Pruvost, Mélanie
  • UMR 5199 De la Préhistoire à l'Actuel : Culture, Environnement et Anthropologie (PACEA), CNRS, Université de Bordeaux, Pessac Cedex, France.
Olsen, Sandra
  • Division of Archaeology, Biodiversity Institute, University of Kansas, Lawrence, KS, USA.
Makarewicz, Cheryl A
  • Institute for Prehistoric and Protohistoric Archaeology, Kiel University, Kiel, Germany.
  • ROOTS Excellence Cluster, Kiel University, Kiel, Germany.
Valenzuela Lamas, Silvia
  • Archaeology of Social Dynamics, Institució Milà i Fontanals d'Humanitats, Consejo Superior de Investigaciones Científicas (IMF-CSIC), Barcelona, Spain.
Albizuri Canadell, Silvia
  • Departament d'Història i Arqueologia-SERP, Universitat de Barcelona, Barcelona, Spain.
Nieto Espinet, Ariadna
  • Grup d'Investigació Prehistòrica, Universitat de Lleida, PID2019-110022GB-I00, Lleida, Spain.
Iborra, Ma Pilar
  • , Valencia, Spain.
Lira Garrido, Jaime
  • Departamento de Medicina Animal, Facultad de Veterinaria, Universidad de Extremadura, Cáceres, Spain.
  • Centro Mixto UCM-ISCIII de Evolución y Comportamiento Humanos, Madrid, Spain.
Rodríguez González, Esther
  • Instituto de Arqueología (CSIC-Junta de Extremadura), Mérida, Spain.
Celestino, Sebastián
  • Instituto de Arqueología (CSIC-Junta de Extremadura), Mérida, Spain.
Olària, Carmen
  • Laboratori d'Arqueologia Prehistòrica, Universitat Jaume I, Castelló de la Plana, Spain.
Arsuaga, Juan Luis
  • Centro Mixto UCM-ISCIII de Evolución y Comportamiento Humanos, Madrid, Spain.
  • Departamento de Geodinámica, Estratigrafía y Paleontología, Facultad de Ciencias Geológicas, Universidad Complutense de Madrid, Madrid, Spain.
Kotova, Nadiia
  • Department of Eneolithic and Bronze Age, Institute of Archaeology National Academy of Sciences of Ukraine, Kyiv, Ukraine.
Pryor, Alexander
  • Department of Archaeology, University of Exeter, Exeter, UK.
Crabtree, Pam
  • Center for the Study of Human Origins, Anthropology Department, New York University, New York, NY, USA.
Zhumatayev, Rinat
  • Department of Archaeology, Ethnology and Museology, Al Farabi Kazakh National University, Almaty, Kazakhstan.
Toleubaev, Abdesh
  • Department of Archaeology, Ethnology and Museology, Al Farabi Kazakh National University, Almaty, Kazakhstan.
Morgunova, Nina L
  • Scientific Research Department, Orenburg State Pedagogical University, Orenburg, Russia.
Kuznetsova, Tatiana
  • Department of paleontology, Faculty of Geology, Moscow State University, Moscow, Russia.
  • Institute of Geology and Petroleum Technologies, Kazan Federal University, Kazan, Russia.
Lordkipanize, David
  • Georgian National Museum, Tbilisi, Georgia.
  • Tbilisi State University, Tbilisi, Georgia.
Marzullo, Matilde
  • Università degli Studi di Milano, Dipartimento di Beni Culturali e Ambientali, Milan, Italy.
Prato, Ornella
  • Università degli Studi di Milano, Dipartimento di Beni Culturali e Ambientali, Milan, Italy.
Bagnasco Gianni, Giovanna
  • Università degli Studi di Milano, Dipartimento di Beni Culturali e Ambientali, Milan, Italy.
Tecchiati, Umberto
  • Università degli Studi di Milano, Dipartimento di Beni Culturali e Ambientali, Milan, Italy.
Clavel, Benoit
  • Centre National de Recherche Scientifique, Muséum national d'Histoire naturelle, Archéozoologie, Archéobotanique (AASPE), CP 56, Paris, France.
Lepetz, Sébastien
  • Centre National de Recherche Scientifique, Muséum national d'Histoire naturelle, Archéozoologie, Archéobotanique (AASPE), CP 56, Paris, France.
Davoudi, Hossein
  • University of Tehran, Central Laboratory, Bioarchaeology Laboratory, Archaeozoology Section, Tehran, Iran.
Mashkour, Marjan
  • Centre National de Recherche Scientifique, Muséum national d'Histoire naturelle, Archéozoologie, Archéobotanique (AASPE), CP 56, Paris, France.
  • University of Tehran, Central Laboratory, Bioarchaeology Laboratory, Archaeozoology Section, Tehran, Iran.
Berezina, Natalia Ya
  • Research Institute and Museum of Anthropology, Lomonosov Moscow State University, Moscow, Russia.
Stockhammer, Philipp W
  • 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.
Krause, Johannes
  • Department of Archaeogenetics, Max Planck Institute for the Science of Human History, Jena, Germany.
  • Department of Archaeogenetics, Max Planck Institute for Evolutionary Anthropology, Leipzig, Germany.
Haak, Wolfgang
  • Department of Archaeogenetics, Max Planck Institute for the Science of Human History, Jena, Germany.
  • Department of Archaeogenetics, Max Planck Institute for Evolutionary Anthropology, Leipzig, Germany.
  • School of Biological Sciences, The University of Adelaide, Adelaide, South Australia, Australia.
Morales-Muñiz, Arturo
  • Department of Biology, Universidad Autónoma de Madrid, Madrid, Spain.
Benecke, Norbert
  • Eurasia Department of the German Archaeological Institute, Berlin, Germany.
Hofreiter, Michael
  • Evolutionary Adaptive Genomics, Institute of Biochemistry and Biology, Faculty of Mathematics and Science, University of Potsdam, Potsdam, Germany.
Ludwig, Arne
  • 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.
Graphodatsky, Alexander S
  • Department of the Diversity and Evolution of Genomes, Institute of Molecular and Cellular Biology SB RAS, Novosibirsk, Russia.
Peters, Joris
  • 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.
Kiryushin, Kirill Yu
  • Department of Archaeology, Ethnography and Museology, Altai State University, Barnaul, Russia.
Iderkhangai, Tumur-Ochir
  • Department of Archaeology, Ulaanbaatar State University, Ulaanbaatar, Mongolia.
Bokovenko, Nikolay A
  • Institute for the History of Material Culture, Russian Academy of Sciences (IHMC RAS), St Petersburg, Russia.
Vasiliev, Sergey K
  • ArchaeoZOOlogy in Siberia and Central Asia-ZooSCAn International Research Laboratory, Institute of Archeology and Ethnography of the Siberian Branch of the RAS, Novosibirsk, Russia.
Seregin, Nikolai N
  • Department of Archaeology, Ethnography and Museology, Altai State University, Barnaul, Russia.
Chugunov, Konstantin V
  • Department of Eastern European and Siberian Archaeology, State Hermitage Museum, St Petersburg, Russia.
Plasteeva, Natalya A
  • Paleoecology Laboratory, Institute of Plant and Animal Ecology, Ural Branch of the Russian Academy of Sciences, Ekaterinburg, Russia.
Baryshnikov, Gennady F
  • Zoological Institute, Russian Academy of Sciences, St Petersburg, Russia.
Petrova, Ekaterina
  • Department of Archaeology, History Faculty, Vilnius University, Vilnius, Lithuania.
Sablin, Mikhail
  • Zoological Institute, Russian Academy of Sciences, St Petersburg, Russia.
Ananyevskaya, Elina
  • Department of Archaeology, History Faculty, Vilnius University, Vilnius, Lithuania.
Logvin, Andrey
  • Laboratory for Archaeological Research, Faculty of History and Law, Kostanay State University, Kostanay, Kazakhstan.
Shevnina, Irina
  • Laboratory for Archaeological Research, Faculty of History and Law, Kostanay State University, Kostanay, Kazakhstan.
Logvin, Victor
  • Department of History and Archaeology, Surgut Governmental University, Surgut, Russia.
Kalieva, Saule
  • Department of History and Archaeology, Surgut Governmental University, Surgut, Russia.
Loman, Valeriy
  • Saryarka Archaeological Institute, Buketov Karaganda University, Karaganda, Kazakhstan.
Kukushkin, Igor
  • Saryarka Archaeological Institute, Buketov Karaganda University, Karaganda, Kazakhstan.
Merz, Ilya
  • Toraighyrov University, Joint Research Center for Archeological Studies, Pavlodar, Kazakhstan.
Merz, Victor
  • Toraighyrov University, Joint Research Center for Archeological Studies, Pavlodar, Kazakhstan.
Sakenov, Sergazy
  • Faculty of History, L. N. Gumilev Eurasian National University, Nur-Sultan, Kazakhstan.
Varfolomeyev, Victor
  • Saryarka Archaeological Institute, Buketov Karaganda University, Karaganda, Kazakhstan.
Usmanova, Emma
  • Saryarka Archaeological Institute, Buketov Karaganda University, Karaganda, Kazakhstan.
Zaibert, Viktor
  • Institute of Archaeology and Steppe Civilizations, Al-Farabi Kazakh National University, Almaty, Kazakhstan.
Arbuckle, Benjamin
  • Department of Anthropology, Alumni Building, University of North Carolina at Chapel Hill, Chapel Hill, NC, USA.
Belinskiy, Andrey B
  • Nasledie Cultural Heritage Unit, Stavropol, Russia.
Kalmykov, Alexej
  • Nasledie Cultural Heritage Unit, Stavropol, Russia.
Reinhold, Sabine
  • Eurasia Department of the German Archaeological Institute, Berlin, Germany.
Hansen, Svend
  • Eurasia Department of the German Archaeological Institute, Berlin, Germany.
Yudin, Aleksandr I
  • Research Center for the Preservation of Cultural Heritage, Saratov, Russia.
Vybornov, Alekandr A
  • Department of Russian History and Archaeology, Samara State University of Social Sciences and Education, Samara, Russia.
Epimakhov, Andrey
  • Russian and Foreign History Department, South Ural State University, Chelyabinsk, Russia.
  • South Ural Department, Institute of History and Archaeology, Ural Branch of the Russian Academy of Sciences, Ekaterinburg, Russia.
Berezina, Natalia S
  • Archaeological School, Chuvash State Institute of Humanities, Cheboksary, Russia.
Roslyakova, Natalia
  • Department of Russian History and Archaeology, Samara State University of Social Sciences and Education, Samara, Russia.
Kosintsev, Pavel A
  • Paleoecology Laboratory, Institute of Plant and Animal Ecology, Ural Branch of the Russian Academy of Sciences, Ekaterinburg, Russia.
  • Department of History of the Institute of Humanities, Ural Federal University, Ekaterinburg, Russia.
Kuznetsov, Pavel F
  • Department of Russian History and Archaeology, Samara State University of Social Sciences and Education, Samara, Russia.
Anthony, David
  • Department of Human Evolutionary Biology, Harvard University, Cambridge, MA, USA.
  • Anthropology Faculty, Hartwick College, Oneonta, NY, USA.
Kroonen, Guus J
  • Department of Nordic Studies and Linguistics, University of Copenhagen, Copenhagen, Denmark.
  • Leiden University Center for Linguistics, Leiden University, Leiden, The Netherlands.
Kristiansen, Kristian
  • Department of Historical Studies, University of Gothenburg, Gothenburg, Sweden.
  • Lundbeck Foundation GeoGenetics Centre, Copenhagen, Denmark.
Wincker, Patrick
  • Génomique Métabolique, Genoscope, Institut de biologie François Jacob, CEA, CNRS, Université d'Evry, Université Paris-Saclay, Evry, France.
Outram, Alan
  • Department of Archaeology, University of Exeter, Exeter, UK.
Orlando, Ludovic
  • Centre d'Anthropobiologie et de Génomique de Toulouse, Université Paul Sabatier, Toulouse, France. ludovic.orlando@univ-tlse3.fr.

MeSH Terms

  • Animals
  • Archaeology
  • Asia
  • DNA, Ancient
  • Domestication
  • Europe
  • Genetics, Population
  • Genome
  • Grassland
  • Horses / genetics
  • Phylogeny

Conflict of Interest Statement

The authors declare no competing interests.

References

This article includes 91 references
  1. Kelekna P. The Horse in Human History. Cambridge Univ. Press, 2009.
  2. Outram AK, Stear NA, Bendrey R, Olsen S, Kasparov A, Zaibert V, Thorpe N, Evershed RP. The earliest horse harnessing and milking.. Science 2009 Mar 6;323(5919):1332-5.
    doi: 10.1126/science.1168594pubmed: 19265018google scholar: lookup
  3. 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
  4. Olsen, S. L. in Horses and Humans: The Evolution of Human Equine Relationships (eds Olsen S. L.et al.) 81–113 (Archaeopress, 2006).
  5. Fages A, Hanghøj K, Khan N, Gaunitz C, Seguin-Orlando A, Leonardi M, McCrory Constantz C, Gamba C, Al-Rasheid KAS, Albizuri S, Alfarhan AH, Allentoft M, Alquraishi S, Anthony D, Baimukhanov N, Barrett JH, Bayarsaikhan J, Benecke N, Bernáldez-Sánchez E, Berrocal-Rangel L, Biglari F, Boessenkool S, Boldgiv B, Brem G, Brown D, Burger J, Crubézy E, Daugnora L, Davoudi H, de Barros Damgaard P, de Los Ángeles de Chorro Y de Villa-Ceballos M, Deschler-Erb S, Detry C, Dill N, do Mar Oom M, Dohr A, Ellingvåg S, Erdenebaatar D, Fathi H, Felkel S, Fernández-Rodríguez C, García-Viñas E, Germonpré M, Granado JD, Hallsson JH, Hemmer H, Hofreiter M, Kasparov A, Khasanov M, Khazaeli R, Kosintsev P, Kristiansen K, Kubatbek T, Kuderna L, Kuznetsov P, Laleh H, Leonard JA, Lhuillier J, Liesau von Lettow-Vorbeck C, Logvin A, Lõugas L, Ludwig A, Luis C, Arruda AM, Marques-Bonet T, Matoso Silva R, Merz V, Mijiddorj E, Miller BK, Monchalov O, Mohaseb FA, Morales A, Nieto-Espinet A, Nistelberger H, Onar V, Pálsdóttir AH, Pitulko V, Pitskhelauri K, Pruvost M, Rajic Sikanjic P, Rapan Papeša A, Roslyakova N, Sardari A, Sauer E, Schafberg R, Scheu A, Schibler J, Schlumbaum A, Serrand N, Serres-Armero A, Shapiro B, Sheikhi Seno S, Shevnina I, Shidrang S, Southon J, Star B, Sykes N, Taheri K, Taylor W, Teegen WR, Trbojević Vukičević T, Trixl S, Tumen D, Undrakhbold S, Usmanova E, Vahdati A, Valenzuela-Lamas S, Viegas C, Wallner B, Weinstock J, Zaibert V, Clavel B, Lepetz S, Mashkour M, Helgason A, Stefánsson K, Barrey E, Willerslev E, Outram AK, Librado P, Orlando L. Tracking Five Millennia of Horse Management with Extensive Ancient Genome Time Series.. Cell 2019 May 30;177(6):1419-1435.e31.
    doi: 10.1016/j.cell.2019.03.049pmc: PMC6547883pubmed: 31056281google scholar: lookup
  6. Guimaraes S, Arbuckle BS, Peters J, Adcock SE, Buitenhuis H, Chazin H, Manaseryan N, Uerpmann HP, Grange T, Geigl EM. Ancient DNA shows domestic horses were introduced in the southern Caucasus and Anatolia during the Bronze Age.. Sci Adv 2020 Sep;6(38).
    pmc: PMC7494339pubmed: 32938680doi: 10.1126/sciadv.abb0030google scholar: lookup
  7. Anthony D W. The Horse, the Wheel and Language. Princeton Univ. Press, 2007.
  8. Haak W, Lazaridis I, Patterson N, Rohland N, Mallick S, Llamas B, Brandt G, Nordenfelt S, Harney E, Stewardson K, Fu Q, Mittnik A, Bánffy E, Economou C, Francken M, Friederich S, Pena RG, Hallgren F, Khartanovich V, Khokhlov A, Kunst M, Kuznetsov P, Meller H, Mochalov O, Moiseyev V, Nicklisch N, Pichler SL, Risch R, Rojo Guerra MA, Roth C, Szécsényi-Nagy A, Wahl J, Meyer M, Krause J, Brown D, Anthony D, Cooper A, Alt KW, Reich D. Massive migration from the steppe was a source for Indo-European languages in Europe.. Nature 2015 Jun 11;522(7555):207-11.
    doi: 10.1038/nature14317pmc: PMC5048219pubmed: 25731166google scholar: lookup
  9. Allentoft ME, Sikora M, Sjögren KG, Rasmussen S, Rasmussen M, Stenderup J, Damgaard PB, Schroeder H, Ahlström T, Vinner L, Malaspinas AS, Margaryan A, Higham T, Chivall D, Lynnerup N, Harvig L, Baron J, Della Casa P, Dąbrowski P, Duffy PR, Ebel AV, Epimakhov A, Frei K, Furmanek M, Gralak T, Gromov A, Gronkiewicz S, Grupe G, Hajdu T, Jarysz R, Khartanovich V, Khokhlov A, Kiss V, Kolář J, Kriiska A, Lasak I, Longhi C, McGlynn G, Merkevicius A, Merkyte I, Metspalu M, Mkrtchyan R, Moiseyev V, Paja L, Pálfi G, Pokutta D, Pospieszny Ł, Price TD, Saag L, Sablin M, Shishlina N, Smrčka V, Soenov VI, Szeverényi V, Tóth G, Trifanova SV, Varul L, Vicze M, Yepiskoposyan L, Zhitenev V, Orlando L, Sicheritz-Pontén T, Brunak S, Nielsen R, Kristiansen K, Willerslev E. Population genomics of Bronze Age Eurasia.. Nature 2015 Jun 11;522(7555):167-72.
    doi: 10.1038/nature14507pubmed: 26062507google scholar: lookup
  10. Demoule J P. Mais où sont passés les Indo-Européens ? Le mythe d'origine de l'Occident. Le Seuil, 2014.
  11. Damgaard PB, Marchi N, Rasmussen S, Peyrot M, Renaud G, Korneliussen T, Moreno-Mayar JV, Pedersen MW, Goldberg A, Usmanova E, Baimukhanov N, Loman V, Hedeager L, Pedersen AG, Nielsen K, Afanasiev G, Akmatov K, Aldashev A, Alpaslan A, Baimbetov G, Bazaliiskii VI, Beisenov A, Boldbaatar B, Boldgiv B, Dorzhu C, Ellingvag S, Erdenebaatar D, Dajani R, Dmitriev E, Evdokimov V, Frei KM, Gromov A, Goryachev A, Hakonarson H, Hegay T, Khachatryan Z, Khaskhanov R, Kitov E, Kolbina A, Kubatbek T, Kukushkin A, Kukushkin I, Lau N, Margaryan A, Merkyte I, Mertz IV, Mertz VK, Mijiddorj E, Moiyesev V, Mukhtarova G, Nurmukhanbetov B, Orozbekova Z, Panyushkina I, Pieta K, Smrčka V, Shevnina I, Logvin A, Sjögren KG, Štolcová T, Taravella AM, Tashbaeva K, Tkachev A, Tulegenov T, Voyakin D, Yepiskoposyan L, Undrakhbold S, Varfolomeev V, Weber A, Wilson Sayres MA, Kradin N, Allentoft ME, Orlando L, Nielsen R, Sikora M, Heyer E, Kristiansen K, Willerslev E. 137 ancient human genomes from across the Eurasian steppes.. Nature 2018 May;557(7705):369-374.
    doi: 10.1038/s41586-018-0094-2pubmed: 29743675google scholar: lookup
  12. Narasimhan VM, Patterson N, Moorjani P, Rohland N, Bernardos R, Mallick S, Lazaridis I, Nakatsuka N, Olalde I, Lipson M, Kim AM, Olivieri LM, Coppa A, Vidale M, Mallory J, Moiseyev V, Kitov E, Monge J, Adamski N, Alex N, Broomandkhoshbacht N, Candilio F, Callan K, Cheronet O, Culleton BJ, Ferry M, Fernandes D, Freilich S, Gamarra B, Gaudio D, Hajdinjak M, Harney É, Harper TK, Keating D, Lawson AM, Mah M, Mandl K, Michel M, Novak M, Oppenheimer J, Rai N, Sirak K, Slon V, Stewardson K, Zalzala F, Zhang Z, Akhatov G, Bagashev AN, Bagnera A, Baitanayev B, Bendezu-Sarmiento J, Bissembaev AA, Bonora GL, Chargynov TT, Chikisheva T, Dashkovskiy PK, Derevianko A, Dobeš M, Douka K, Dubova N, Duisengali MN, Enshin D, Epimakhov A, Fribus AV, Fuller D, Goryachev A, Gromov A, Grushin SP, Hanks B, Judd M, Kazizov E, Khokhlov A, Krygin AP, Kupriyanova E, Kuznetsov P, Luiselli D, Maksudov F, Mamedov AM, Mamirov TB, Meiklejohn C, Merrett DC, Micheli R, Mochalov O, Mustafokulov S, Nayak A, Pettener D, Potts R, Razhev D, Rykun M, Sarno S, Savenkova TM, Sikhymbaeva K, Slepchenko SM, Soltobaev OA, Stepanova N, Svyatko S, Tabaldiev K, Teschler-Nicola M, Tishkin AA, Tkachev VV, Vasilyev S, Velemínský P, Voyakin D, Yermolayeva A, Zahir M, Zubkov VS, Zubova A, Shinde VS, Lalueza-Fox C, Meyer M, Anthony D, Boivin N, Thangaraj K, Kennett DJ, Frachetti M, Pinhasi R, Reich D. The formation of human populations in South and Central Asia.. Science 2019 Sep 6;365(6457).
    doi: 10.1126/science.aat7487pmc: PMC6822619pubmed: 31488661google scholar: lookup
  13. Rohland N, Harney E, Mallick S, Nordenfelt S, Reich D. Partial uracil-DNA-glycosylase treatment for screening of ancient DNA.. Philos Trans R Soc Lond B Biol Sci 2015 Jan 19;370(1660):20130624.
    doi: 10.1098/rstb.2013.0624pmc: PMC4275898pubmed: 25487342google scholar: lookup
  14. Schubert M, Jónsson H, Chang D, Der Sarkissian C, Ermini L, Ginolhac A, Albrechtsen A, Dupanloup I, Foucal A, Petersen B, Fumagalli M, Raghavan M, Seguin-Orlando A, Korneliussen TS, Velazquez AM, Stenderup J, Hoover CA, Rubin CJ, Alfarhan AH, Alquraishi SA, Al-Rasheid KA, MacHugh DE, Kalbfleisch T, MacLeod JN, Rubin EM, Sicheritz-Ponten T, Andersson L, Hofreiter M, Marques-Bonet T, Gilbert MT, Nielsen R, Excoffier L, Willerslev E, Shapiro B, Orlando L. Prehistoric genomes reveal the genetic foundation and cost of horse domestication.. Proc Natl Acad Sci U S A 2014 Dec 30;111(52):E5661-9.
    doi: 10.1073/pnas.1416991111pmc: PMC4284583pubmed: 25512547google scholar: lookup
  15. Librado P, Der Sarkissian C, Ermini L, Schubert M, Jónsson H, Albrechtsen A, Fumagalli M, Yang MA, Gamba C, Seguin-Orlando A, Mortensen CD, Petersen B, Hoover CA, Lorente-Galdos B, Nedoluzhko A, Boulygina E, Tsygankova S, Neuditschko M, Jagannathan V, Thèves C, Alfarhan AH, Alquraishi SA, Al-Rasheid KA, Sicheritz-Ponten T, Popov R, Grigoriev S, Alekseev AN, Rubin EM, McCue M, Rieder S, Leeb T, Tikhonov A, Crubézy E, Slatkin M, Marques-Bonet T, Nielsen R, Willerslev E, Kantanen J, Prokhortchouk E, Orlando L. Tracking the origins of Yakutian horses and the genetic basis for their fast adaptation to subarctic environments.. Proc Natl Acad Sci U S A 2015 Dec 15;112(50):E6889-97.
    doi: 10.1073/pnas.1513696112pmc: PMC4687531pubmed: 26598656google scholar: lookup
  16. Petkova D, Novembre J, Stephens M. Visualizing spatial population structure with estimated effective migration surfaces.. Nat Genet 2016 Jan;48(1):94-100.
    doi: 10.1038/ng.3464pmc: PMC4696895pubmed: 26642242google scholar: lookup
  17. Patterson N, Moorjani P, Luo Y, Mallick S, Rohland N, Zhan Y, Genschoreck T, Webster T, Reich D. Ancient admixture in human history.. Genetics 2012 Nov;192(3):1065-93.
    doi: 10.1534/genetics.112.145037pmc: PMC3522152pubmed: 22960212google scholar: lookup
  18. Harney É, Patterson N, Reich D, Wakeley J. Assessing the performance of qpAdm: a statistical tool for studying population admixture.. Genetics 2021 Apr 15;217(4).
    doi: 10.1093/genetics/iyaa045pmc: PMC8049561pubmed: 33772284google scholar: lookup
  19. Molloy EK, Durvasula A, Sankararaman S. Advancing admixture graph estimation via maximum likelihood network orientation.. Bioinformatics 2021 Jul 12;37(Suppl_1):i142-i150.
  20. Battey CJ, Ralph PL, Kern AD. Predicting geographic location from genetic variation with deep neural networks.. Elife 2020 Jun 8;9.
    doi: 10.7554/eLife.54507pmc: PMC7324158pubmed: 32511092google scholar: lookup
  21. de Barros Damgaard P, Martiniano R, Kamm J, Moreno-Mayar JV, Kroonen G, Peyrot M, Barjamovic G, Rasmussen S, Zacho C, Baimukhanov N, Zaibert V, Merz V, Biddanda A, Merz I, Loman V, Evdokimov V, Usmanova E, Hemphill B, Seguin-Orlando A, Yediay FE, Ullah I, Sjögren KG, Iversen KH, Choin J, de la Fuente C, Ilardo M, Schroeder H, Moiseyev V, Gromov A, Polyakov A, Omura S, Senyurt SY, Ahmad H, McKenzie C, Margaryan A, Hameed A, Samad A, Gul N, Khokhar MH, Goriunova OI, Bazaliiskii VI, Novembre J, Weber AW, Orlando L, Allentoft ME, Nielsen R, Kristiansen K, Sikora M, Outram AK, Durbin R, Willerslev E. The first horse herders and the impact of early Bronze Age steppe expansions into Asia.. Science 2018 Jun 29;360(6396).
    doi: 10.1126/science.aar7711pmc: PMC6748862pubmed: 29743352google scholar: lookup
  22. Reinhold, S. et al. in Appropriating Innovations: Entangled Knowledge in Eurasia, 5000–1500 bce (eds Stockhammer, P. W. & Maran, J.) 78–97 (Oxbow Books, 2017).
  23. Kristiansen, K. in Trade and Civilization. Economic Networks and Cultural Ties, from Prehistory to the Early Modern Period (eds Kristiansen, K. et al.) (Cambridge Univ. Press, 2018).
  24. Chechushkov I. V., & Epimakhov, A. V. in The Puzzle of Indo-European Origins and Dispersals: Archeology, Linguistics and Genetics (eds Kristiansen, K. et al.) (Cambridge Univ. Press, in the press).
  25. Littauer M A, Crouwel J H. The origin of the true chariot. Antiquity 1996;70:934–939.
    doi: 10.1017/S0003598X00084192google scholar: lookup
  26. Lindner S. Chariots in the Eurasian Steppe: a Bayesian approach to the emergence of horse-drawn transport in the early second millennium BC. Antiquity 2020;94:361–380.
    doi: 10.15184/aqy.2020.37google scholar: lookup
  27. Moorey P R S. Pictorial evidence for the history of horse-riding in Iraq before the Kassite period. Iraq 1970;32:36–50.
    doi: 10.2307/4199890google scholar: lookup
  28. Kanne K. Riding, ruling, and resistance equestrianism and political authority in the Hungarian Bronze Age. Curr. Anthropol. in the press.
  29. Suri P, Palmer MR, Tsepilov YA, Freidin MB, Boer CG, Yau MS, Evans DS, Gelemanovic A, Bartz TM, Nethander M, Arbeeva L, Karssen L, Neogi T, Campbell A, Mellstrom D, Ohlsson C, Marshall LM, Orwoll E, Uitterlinden A, Rotter JI, Lauc G, Psaty BM, Karlsson MK, Lane NE, Jarvik GP, Polasek O, Hochberg M, Jordan JM, Van Meurs JBJ, Jackson R, Nielson CM, Mitchell BD, Smith BH, Hayward C, Smith NL, Aulchenko YS, Williams FMK. Genome-wide meta-analysis of 158,000 individuals of European ancestry identifies three loci associated with chronic back pain.. PLoS Genet 2018 Sep;14(9):e1007601.
  30. Jiang H, Moro A, Liu Y, Wang J, Meng D, Zhan X, Wei Q. Two GWAS-identified variants are associated with lumbar spinal stenosis and Gasdermin-C expression in Chinese population.. Sci Rep 2020 Dec 3;10(1):21069.
    doi: 10.1038/s41598-020-78249-7pmc: PMC7713291pubmed: 33273635google scholar: lookup
  31. Tikker L, Casarotto P, Singh P, Biojone C, Piepponen TP, Estartús N, Seelbach A, Sridharan R, Laukkanen L, Castrén E, Partanen J. Inactivation of the GATA Cofactor ZFPM1 Results in Abnormal Development of Dorsal Raphe Serotonergic Neuron Subtypes and Increased Anxiety-Like Behavior.. J Neurosci 2020 Nov 4;40(45):8669-8682.
  32. Takahashi A, Miczek KA. Neurogenetics of aggressive behavior: studies in rodents.. Curr Top Behav Neurosci 2014;17:3-44.
    doi: 10.1007/7854_2013_263pmc: PMC4092042pubmed: 24318936google scholar: lookup
  33. Schmitt T, Varga Z. Extra-Mediterranean refugia: The rule and not the exception?. Front Zool 2012 Sep 6;9(1):22.
    doi: 10.1186/1742-9994-9-22pmc: PMC3462695pubmed: 22953783google scholar: lookup
  34. Spasskaya, N. N., & Pavlinov, I. in Zoological Research (Arch. Zoological Museum, Moscow State Univ., 2016).
  35. Colledge S, Conolly J, Crema E, Shennan S. Neolithic population crash in northwest Europe associated with agricultural crisis. Quat. Res. 2019;92:686–707.
    doi: 10.1017/qua.2019.42google scholar: lookup
  36. Outram, A. K. & Bogaard, A. Subsistence and Society in Prehistory: New Directions in Economic Archaeology (Cambridge Univ. Press, 2019).
  37. Anthony, D. W. in Social Complexity in Prehistoric Eurasia: Monuments, Metals and Mobility (eds Hanks, B. K. & Lindruff, K. M.) Ch. 4 (2009).
  38. Maran J, Bajenaru R, Ailincai S-C, Popescu A-D, Hansen S I. Objects, ideas and travelers. Contacts between the Balkans, the Aegean and Western Anatolia during the Bronze and Early Iron Age. Proc. of the Conference in Tulcea 10-13 November, 2017 (Rudolf Habelt, 2020).
  39. Glob P V. Denmark: An Archaeological History from the Stone Age to the Vikings. Cornell Univ. Press, 1971.
  40. Gimbutas M. The first wave of Eurasian Steppe pastoralists into Copper Age Europe. J. Indo. Eur. Stud. 1977;5:277–338.
  41. Anthony D W. The “Kurgan Culture,” Indo-European origins, and the domestication of the horse: a reconsideration. Curr. Anthropol. 1986;27:291–313.
    doi: 10.1086/203441google scholar: lookup
  42. Renfrew C. They ride horses, don’t they?: Mallory on the Indo-Europeans. Antiquity 1989;63:843–847.
    doi: 10.1017/S0003598X00077012google scholar: lookup
  43. Vandkilde H. Culture and Change in Central European Prehistory. Aarhus Univ. Press, 2007.
  44. Häusler, A. in Indogermanen und das Pferd (eds Hänsel, B. & Zimmer, S.) 217–257 (Archaeolingua Alapitvany, 1994).
  45. Kroonen G, Barjamovic G, Peyrot M. Linguistic supplement to de Barros Damgaard et al. 2018: Early Indo-European languages, Anatolian, Tocharian and Indo-Iranian. https://zenodo.org/record/1240524#.YFtLgGjTVMQ (2018).
  46. South A. rworldmap: a new R package for mapping global data. R J. 2011;3:35–43.
    doi: 10.32614/RJ-2011-006google scholar: lookup
  47. Brownrigg, R. maps: draw geographical maps. R package version 3.3.0 https://CRAN.R-project.org/package=maps (2018).
  48. Reimer P. The IntCal20 Northern Hemisphere radiocarbon age calibration curve (0–55 cal kBP). Radiocarbon 2020;62:725.
    doi: 10.1017/RDC.2020.41google scholar: lookup
  49. Ramsey C B. Bayesian analysis of radiocarbon dates. Radiocarbon 2009;51:337–360.
    doi: 10.1017/S0033822200033865google scholar: lookup
  50. Seguin-Orlando A, Donat R, Der Sarkissian C, Southon J, Thèves C, Manen C, Tchérémissinoff Y, Crubézy E, Shapiro B, Deleuze JF, Dalén L, Guilaine J, Orlando L. Heterogeneous Hunter-Gatherer and Steppe-Related Ancestries in Late Neolithic and Bell Beaker Genomes from Present-Day France.. Curr Biol 2021 Mar 8;31(5):1072-1083.e10.
    doi: 10.1016/j.cub.2020.12.015pubmed: 33434506google scholar: lookup
  51. Gamba C, Hanghøj K, Gaunitz C, Alfarhan AH, Alquraishi SA, Al-Rasheid KA, Bradley DG, Orlando L. Comparing the performance of three ancient DNA extraction methods for high-throughput sequencing.. Mol Ecol Resour 2016 Mar;16(2):459-69.
    doi: 10.1111/1755-0998.12470pubmed: 26401836google scholar: lookup
  52. Schubert M, Lindgreen S, Orlando L. AdapterRemoval v2: rapid adapter trimming, identification, and read merging.. BMC Res Notes 2016 Feb 12;9:88.
    doi: 10.1186/s13104-016-1900-2pmc: PMC4751634pubmed: 26868221google scholar: lookup
  53. Kalbfleisch TS, Rice ES, DePriest MS Jr, Walenz BP, Hestand MS, Vermeesch JR, O Connell BL, Fiddes IT, Vershinina AO, Saremi NF, Petersen JL, Finno CJ, Bellone RR, McCue ME, Brooks SA, Bailey E, Orlando L, Green RE, Miller DC, Antczak DF, MacLeod JN. Improved reference genome for the domestic horse increases assembly contiguity and composition.. Commun Biol 2018;1:197.
    doi: 10.1038/s42003-018-0199-zpmc: PMC6240028pubmed: 30456315google scholar: lookup
  54. Xu X, Arnason U. The complete mitochondrial DNA sequence of the horse, Equus caballus: extensive heteroplasmy of the control region.. Gene 1994 Oct 21;148(2):357-62.
    doi: 10.1016/0378-1119(94)90713-7pubmed: 7958969google scholar: lookup
  55. Felkel S, Vogl C, Rigler D, Dobretsberger V, Chowdhary BP, Distl O, Fries R, Jagannathan V, Janečka JE, Leeb T, Lindgren G, McCue M, Metzger J, Neuditschko M, Rattei T, Raudsepp T, Rieder S, Rubin CJ, Schaefer R, Schlötterer C, Thaller G, Tetens J, Velie B, Brem G, Wallner B. The horse Y chromosome as an informative marker for tracing sire lines.. Sci Rep 2019 Apr 15;9(1):6095.
    doi: 10.1038/s41598-019-42640-wpmc: PMC6465346pubmed: 30988347google scholar: lookup
  56. Poullet M, Orlando L. Assessing DNA sequence alignment methods for characterizing ancient genomes and methylomes. Front. Ecol. Evol. 2020;8:105.
    doi: 10.3389/fevo.2020.00105google scholar: lookup
  57. Jónsson H, Ginolhac A, Schubert M, Johnson PL, Orlando L. mapDamage2.0: fast approximate Bayesian estimates of ancient DNA damage parameters.. Bioinformatics 2013 Jul 1;29(13):1682-4.
  58. 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
  59. 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
  60. Korneliussen TS, Albrechtsen A, Nielsen R. ANGSD: Analysis of Next Generation Sequencing Data.. BMC Bioinformatics 2014 Nov 25;15(1):356.
    doi: 10.1186/s12859-014-0356-4pmc: PMC4248462pubmed: 25420514google scholar: lookup
  61. Skoglund P, Northoff BH, Shunkov MV, Derevianko AP, Pääbo S, Krause J, Jakobsson M. Separating endogenous ancient DNA from modern day contamination in a Siberian Neandertal.. Proc Natl Acad Sci U S A 2014 Feb 11;111(6):2229-34.
    doi: 10.1073/pnas.1318934111pmc: PMC3926038pubmed: 24469802google scholar: lookup
  62. Librado P, Gamba C, Gaunitz C, Der Sarkissian C, Pruvost M, Albrechtsen A, Fages A, Khan N, Schubert M, Jagannathan V, Serres-Armero A, Kuderna LFK, Povolotskaya IS, Seguin-Orlando A, Lepetz S, Neuditschko M, Thèves C, Alquraishi S, Alfarhan AH, Al-Rasheid K, Rieder S, Samashev Z, Francfort HP, Benecke N, Hofreiter M, Ludwig A, Keyser C, Marques-Bonet T, Ludes B, Crubézy E, Leeb T, Willerslev E, Orlando L. Ancient genomic changes associated with domestication of the horse.. Science 2017 Apr 28;356(6336):442-445.
    doi: 10.1126/science.aam5298pubmed: 28450643google scholar: lookup
  63. Der Sarkissian C, Ermini L, Schubert M, Yang MA, Librado P, Fumagalli M, Jónsson H, Bar-Gal GK, Albrechtsen A, Vieira FG, Petersen B, Ginolhac A, Seguin-Orlando A, Magnussen K, Fages A, Gamba C, Lorente-Galdos B, Polani S, Steiner C, Neuditschko M, Jagannathan V, Feh C, Greenblatt CL, Ludwig A, Abramson NI, Zimmermann W, Schafberg R, Tikhonov A, Sicheritz-Ponten T, Willerslev E, Marques-Bonet T, Ryder OA, McCue M, Rieder S, Leeb T, Slatkin M, Orlando L. Evolutionary Genomics and Conservation of the Endangered Przewalski's Horse.. Curr Biol 2015 Oct 5;25(19):2577-83.
    doi: 10.1016/j.cub.2015.08.032pmc: PMC5104162pubmed: 26412128google scholar: lookup
  64. Katoh K, Standley DM. MAFFT multiple sequence alignment software version 7: improvements in performance and usability.. Mol Biol Evol 2013 Apr;30(4):772-80.
    doi: 10.1093/molbev/mst010pmc: PMC3603318pubmed: 23329690google scholar: lookup
  65. Stamatakis A. RAxML version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies.. Bioinformatics 2014 May 1;30(9):1312-3.
  66. Bouckaert R, Vaughan TG, Barido-Sottani J, Duchêne S, Fourment M, Gavryushkina A, Heled J, Jones G, Kühnert D, De Maio N, Matschiner M, Mendes FK, Müller NF, Ogilvie HA, du Plessis L, Popinga A, Rambaut A, Rasmussen D, Siveroni I, Suchard MA, Wu CH, Xie D, Zhang C, Stadler T, Drummond AJ. BEAST 2.5: An advanced software platform for Bayesian evolutionary analysis.. PLoS Comput Biol 2019 Apr;15(4):e1006650.
  67. Heller R, Chikhi L, Siegismund HR. The confounding effect of population structure on Bayesian skyline plot inferences of demographic history.. PLoS One 2013;8(5):e62992.
  68. Keane TM, Creevey CJ, Pentony MM, Naughton TJ, Mclnerney JO. Assessment of methods for amino acid matrix selection and their use on empirical data shows that ad hoc assumptions for choice of matrix are not justified.. BMC Evol Biol 2006 Mar 24;6:29.
    doi: 10.1186/1471-2148-6-29pmc: PMC1435933pubmed: 16563161google scholar: lookup
  69. Drummond AJ, Rambaut A, Shapiro B, Pybus OG. Bayesian coalescent inference of past population dynamics from molecular sequences.. Mol Biol Evol 2005 May;22(5):1185-92.
    doi: 10.1093/molbev/msi103pubmed: 15703244google scholar: lookup
  70. Rambaut A, Drummond AJ, Xie D, Baele G, Suchard MA. Posterior Summarization in Bayesian Phylogenetics Using Tracer 1.7.. Syst Biol 2018 Sep 1;67(5):901-904.
    doi: 10.1093/sysbio/syy032pmc: PMC6101584pubmed: 29718447google scholar: lookup
  71. Nguyen LT, Schmidt HA, von Haeseler A, Minh BQ. IQ-TREE: a fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies.. Mol Biol Evol 2015 Jan;32(1):268-74.
    doi: 10.1093/molbev/msu300pmc: PMC4271533pubmed: 25371430google scholar: lookup
  72. Hoang DT, Chernomor O, von Haeseler A, Minh BQ, Vinh LS. UFBoot2: Improving the Ultrafast Bootstrap Approximation.. Mol Biol Evol 2018 Feb 1;35(2):518-522.
    doi: 10.1093/molbev/msx281pmc: PMC5850222pubmed: 29077904google scholar: lookup
  73. Lefort V, Desper R, Gascuel O. FastME 2.0: A Comprehensive, Accurate, and Fast Distance-Based Phylogeny Inference Program.. Mol Biol Evol 2015 Oct;32(10):2798-800.
    doi: 10.1093/molbev/msv150pmc: PMC4576710pubmed: 26130081google scholar: lookup
  74. Paradis E, Schliep K. ape 5.0: an environment for modern phylogenetics and evolutionary analyses in R.. Bioinformatics 2019 Feb 1;35(3):526-528.
    doi: 10.1093/bioinformatics/bty633pubmed: 30016406google scholar: lookup
  75. Schraiber JG. Assessing the Relationship of Ancient and Modern Populations.. Genetics 2018 Jan;208(1):383-398.
    doi: 10.1534/genetics.117.300448pmc: PMC5753871pubmed: 29167200google scholar: lookup
  76. Purcell S, Neale B, Todd-Brown K, Thomas L, Ferreira MA, Bender D, Maller J, Sklar P, de Bakker PI, Daly MJ, Sham PC. PLINK: a tool set for whole-genome association and population-based linkage analyses.. Am J Hum Genet 2007 Sep;81(3):559-75.
    doi: 10.1086/519795pmc: PMC1950838pubmed: 17701901google scholar: lookup
  77. Alexander DH, Novembre J, Lange K. Fast model-based estimation of ancestry in unrelated individuals.. Genome Res 2009 Sep;19(9):1655-64.
    doi: 10.1101/gr.094052.109pmc: PMC2752134pubmed: 19648217google scholar: lookup
  78. Cheng JY, Mailund T, Nielsen R. Fast admixture analysis and population tree estimation for SNP and NGS data.. Bioinformatics 2017 Jul 15;33(14):2148-2155.
  79. Lawson DJ, van Dorp L, Falush D. A tutorial on how not to over-interpret STRUCTURE and ADMIXTURE bar plots.. Nat Commun 2018 Aug 14;9(1):3258.
    doi: 10.1038/s41467-018-05257-7pmc: PMC6092366pubmed: 30108219google scholar: lookup
  80. Excoffier L, Dupanloup I, Huerta-Sánchez E, Sousa VC, Foll M. Robust demographic inference from genomic and SNP data.. PLoS Genet 2013 Oct;9(10):e1003905.
  81. Gerritsen, H. mapplots: data visualisation on maps. R package version 1.5.1 https://CRAN.R-project.org/package=mapplots (2018).
  82. Bjornstad, O. N. & Cai, J. ncf: spatial covariance functions. R package version 1.2-9 http://ento.psu.edu/directory/onb1 (2020).
  83. Loog L, Mirazón Lahr M, Kovacevic M, Manica A, Eriksson A, Thomas MG. Estimating mobility using sparse data: Application to human genetic variation.. Proc Natl Acad Sci U S A 2017 Nov 14;114(46):12213-12218.
    doi: 10.1073/pnas.1703642114pmc: PMC5699029pubmed: 29087301google scholar: lookup
  84. Hijmans, R. J., Williams, E. & Vennes, C. E.. geosphere: spherical trigonometry. R package version 1.5.1 (2019).
  85. Boyle, J. GeoRange: calculating geographic range from occurrence data. R package version 0.1.0. (2017).
  86. Hahne F, Ivanek R. Visualizing Genomic Data Using Gviz and Bioconductor.. Methods Mol Biol 2016;1418:335-51.
    doi: 10.1007/978-1-4939-3578-9_16pubmed: 27008022google scholar: lookup
  87. Renaud G, Petersen B, Seguin-Orlando A, Bertelsen MF, Waller A, Newton R, Paillot R, Bryant N, Vaudin M, Librado P, Orlando L. Improved de novo genomic assembly for the domestic donkey.. Sci Adv 2018 Apr;4(4):eaaq0392.
    doi: 10.1126/sciadv.aaq0392pmc: PMC5938232pubmed: 29740610google scholar: lookup
  88. Jagannathan V, Gerber V, Rieder S, Tetens J, Thaller G, Drögemüller C, Leeb T. Comprehensive characterization of horse genome variation by whole-genome sequencing of 88 horses.. Anim Genet 2019 Feb;50(1):74-77.
    doi: 10.1111/age.12753pubmed: 30525216google scholar: lookup
  89. Andersson LS, Larhammar M, Memic F, Wootz H, Schwochow D, Rubin CJ, Patra K, Arnason T, Wellbring L, Hjälm G, Imsland F, Petersen JL, McCue ME, Mickelson JR, Cothran G, Ahituv N, Roepstorff L, Mikko S, Vallstedt A, Lindgren G, Andersson L, Kullander K. Mutations in DMRT3 affect locomotion in horses and spinal circuit function in mice.. Nature 2012 Aug 30;488(7413):642-6.
    doi: 10.1038/nature11399pmc: PMC3523687pubmed: 22932389google scholar: lookup
  90. Teufer M. Ein Scheibenknebel aus Dzarkutan (Süduzbekistan). Archäologische Mitteilungen aus Iran und Turan Band. 1999;31:69–142.
  91. Chechushkov I V. Wheel Complex of the Late Bronze Age Era of Steppe and Forest-Steppe Eurasia (from Dnieper to Irtysh). PhD thesis. Department of Archeology and Ethnography of the Federal State Budgetary Institution of Science, Institute of History and Archeology of the Ural Branch of the Russian Academy of Sciences (2013).

Citations

This article has been cited 95 times.