The origins and spread of domestic horses from the Western Eurasian steppes
Authors: Librado Pablo , Khan Naveed , Fages Antoine , Kusliy Mariya A. , Suchan Tomasz , Tonasso-Calvière Laure , Schiavinato Stéphanie , Alioglu Duha , Fromentier Aurore , Perdereau Aude , Aury Jean-Marc , Gaunitz Charleen , Chauvey Lorelei , Seguin-Orlando Andaine , Der Sarkissian Clio , Southon John , Shapiro Beth , Tishkin Alexey A. , Kovalev Alexey A. , Alquraishi Saleh , Alfarhan Ahmed H. , Al-Rasheid Khaled A. S. , Seregély Timo , Klassen Lutz , Iversen Rune , Bignon-Lau Olivier , Bodu Pierre , Olive Monique , Castel Jean-Christophe , Boudadi-Maligne Myriam , Alvarez Nadir , Germonpré Mietje , Moskal-del Hoyo Magdalena , Wilczyński Jarosław , Pospuła Sylwia , Lasota-Kuś Anna , Tunia Krzysztof , Nowak Marek , Rannamäe Eve , Saarma Urmas , Boeskorov Gennady , Lōugas Lembi , Kyselý René , Peške Lubomír , Bălășescu Adrian , Dumitrașcu Valentin , Dobrescu Roxana , Gerber Daniel , Kiss Viktória , Szécsényi-Nagy Anna , Mende Balázs G. , Gallina Zsolt , Somogyi Krisztina , Kulcsár Gabriella , Gál Erika , Bendrey Robin , Allentoft Morten E. , Sirbu Ghenadie , Dergachev Valentin , Shephard Henry , Tomadini Noémie , Grouard Sandrine , Kasparov Aleksei , Basilyan Alexander E. , Anisimov Mikhail A. , Nikolskiy Pavel A. , Pavlova Elena Y. , Pitulko Vladimir , Brem Gottfried , Wallner Barbara , Schwall Christoph , Keller Marcel , Kitagawa Keiko , Bessudnov Alexander N. , Bessudnov Alexander , Taylor William , Magail Jérome , Gantulga Jamiyan-Ombo , Bayarsaikhan Jamsranjav , Erdenebaatar Diimaajav , Tabaldiev Kubatbeek , Mijiddorj Enkhbayar , Boldgiv Bazartseren , Tsagaan Turbat , Pruvost Mélanie , Olsen Sandra , Makarewicz Cheryl A. , Valenzuela Lamas Silvia , Albizuri Canadell Silvia , Nieto Espinet Ariadna , Iborra Ma Pilar , Lira Garrido Jaime , Rodríguez González Esther , Celestino Sebastián , Olària Carmen , Arsuaga Juan Luis , Kotova Nadiia , Pryor Alexander , Crabtree Pam , Zhumatayev Rinat , Toleubaev Abdesh , Morgunova Nina L. , Kuznetsova Tatiana , Lordkipanize David , Marzullo Matilde , Prato Ornella , Bagnasco Gianni Giovanna , Tecchiati Umberto , Clavel Benoit , Lepetz Sébastien , Davoudi Hossein , Mashkour Marjan , Berezina Natalia Ya. , Stockhammer Philipp W. , Krause Johannes , Haak Wolfgang , Morales-Muñiz Arturo , Benecke Norbert , Hofreiter Michael , Ludwig Arne , Graphodatsky Alexander S. , Peters Joris , Kiryushin Kirill Yu. , Iderkhangai Tumur-Ochir , Bokovenko Nikolay A. , Vasiliev Sergey K. , Seregin Nikolai N. , Chugunov Konstantin V. , Plasteeva Natalya A. , Baryshnikov Gennady F. , Petrova Ekaterina , Sablin Mikhail , Ananyevskaya Elina , Logvin Andrey , Shevnina Irina , Logvin Victor , Kalieva Saule , Loman Valeriy , Kukushkin Igor , Merz Ilya , Merz Victor , Sakenov Sergazy , Varfolomeyev Victor , Usmanova Emma , Zaibert Viktor , Arbuckle Benjamin , Belinskiy Andrey B. , Kalmykov Alexej , Reinhold Sabine , Hansen Svend , Yudin Aleksandr I. , Vybornov Alekandr A. , Epimakhov Andrey , Berezina Natalia S. , Roslyakova Natalia , Kosintsev Pavel A. , Kuznetsov Pavel F. , Anthony David , Kroonen Guus J. , Kristiansen Kristian , Wincker Patrick , Outram Alan , Orlando Ludovic 
Published: October 2021

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.