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Horse domestication revolutionized warfare and accelerated travel, trade, and the geographic expansion of languages. Here, we present the largest DNA time series for a non-human organism to date, including genome-scale data from 149 ancient animals and 129 ancient genomes (≥1-fold coverage), 87 of which are new. This extensive dataset allows us to assess the modern legacy of past equestrian civilizations. We find that two extinct horse lineages existed during early domestication, one at the far western (Iberia) and the other at the far eastern range (Siberia) of Eurasia. None of these contributed significantly to modern diversity. We show that the influence of Persian-related horse lineages increased following the Islamic conquests in Europe and Asia. Multiple alleles associated with elite-racing, including at the MSTN "speed gene," only rose in popularity within the last millennium. Finally, the development of modern breeding impacted genetic diversity more dramatically than the previous millennia of human management. Genome-wide data from 278 ancient equids provide insights into how ancient equestrian civilizations managed, exchanged, and bred horses and indicate vast loss of genetic diversity as well as the existence of two extinct lineages of horses that failed to contribute to modern domestic animals. ; Taylor Family-Asia Foundation Endowed Chair in Ecology and Conservation Biology. M.L. was supported by a Marie-Curie Individual Fellowship (MSCA-IF-67852). L.L. was supported by the Estonian Research Council (PRG29). C.L. was supported by FCT (SFRH/BPD/100511/2014). P.K., N.R., and O.M. were supported by the Ministry of Educations and Science of Russian Federation (33.1907, 2017/P4) and the Russian Scientific Foundation (18-18-00137). T.M.-B. was supported by the BFU2017-86471-P (MINECO/FEDER, UE), the U01 MH106874 grant, Howard Hughes International Early Career, Obra Social ''La Caixa,'' and Secretaria d'Universitats i Recerca del Departament d'Economia i Coneixement de la Generalitat de Catalunya. V.P. was supported by Russian Science Foundation (16-18-10265) e Danish National Research Foundation (DNRF94), the Initiative d'Excellence Chaires d'attractivite´ , Universite´ de Toulouse (OURASI), the International Highly Cited Research Group Program (HCRC#15-101), Deanship of Scientific Research, King Saud University, the Villum Fonden miGENEPI research project, the Swiss National Science Foundation (CR13I1_140638), the Research Council of Norway (project 230821/F20); the investigation grant HAR2016-77600-P, Ministerio de Economía y Competitividad, Spain, and the National Science Foundation ANS1417036). This project has received funding from the European Research Council (ERC) under the European Union's Horizon 2020 research and innovation programme (grant agreement 681605)

Horse domestication revolutionized warfare and accelerated travel, trade, and the geographic expansion of languages. Here, we present the largest DNA time series for a non-human organism to date, including genome-scale data from 149 ancient animals and 129 ancient genomes (≥1-fold coverage), 87 of which are new. This extensive dataset allows us to assess the modern legacy of past equestrian civilizations. We find that two extinct horse lineages existed during early domestication, one at the far western (Iberia) and the other at the far eastern range (Siberia) of Eurasia. None of these contributed significantly to modern diversity. We show that the influence of Persian-related horse lineages increased following the Islamic conquests in Europe and Asia. Multiple alleles associated with elite-racing, including at the MSTN "speed gene," only rose in popularity within the last millennium. Finally, the development of modern breeding impacted genetic diversity more dramatically than the previous millennia of human management. ; B.B. was supported by the Taylor Family-Asia Foundation Endowed Chair in Ecology and Conservation Biology. M.L. was supported by a Marie-Curie Individual Fellowship (MSCA-IF-67852). L.L. was supported by the Estonian Research Council (PRG29). C.L. was supported by FCT (SFRH/BPD/100511/2014). P.K., N.R., and O.M. were supported by the Ministry of Educations and Science of Russian Federation (33.1907, 2017/Π4) and the Russian Scientific Foundation (18-18-00137). T.M.-B. was supported by the BFU2017-86471-P (MINECO/FEDER, UE), the U01 MH106874 grant, Howard Hughes International Early Career, Obra Social "La Caixa," and Secretaria d'Universitats i Recerca del Departament d'Economia i Coneixement de la Generalitat de Catalunya. V.P. was supported by Russian Science Foundation (16-18-10265). This research received support from the SYNTHESYS Project (http://www.synthesys.info/), which is financed by European Community Research Infrastructure Action under the Seventh Framework "Capacities" Programme. This work was supported by the Danish National Research Foundation (DNRF94), the Initiative d'Excellence Chaires d'attractivité, Université de Toulouse (OURASI), the International Highly Cited Research Group Program (HCRC#15-101), Deanship of Scientific Research, King Saud University, the Villum Fonden miGENEPI research project, the Swiss National Science Foundation (CR13I1_140638), the Research Council of Norway (project 230821/F20); the investigation grant HAR2016-77600-P, Ministerio de Economía y Competitividad, Spain, and the National Science Foundation (ANS-1417036). This project has received funding from the European Research Council (ERC) under the European Union's Horizon 2020 research and innovation programme (grant agreement 681605). ; Peer reviewed

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 bc3. 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. ; The work by G. Boeskorov is done on state assignment of DPMGI SB RAS. This project was supported by the University Paul Sabatier IDEX Chaire d'Excellence (OURASI); Villum Funden miGENEPI research programme; the CNRS 'Programme de Recherche Conjoint' (PRC); the CNRS International Research Project (IRP AMADEUS); the France Génomique Appel à Grand Projet (ANR-10-INBS-09-08, BUCEPHALE project); IB10131 and IB18060, both funded by Junta de Extremadura (Spain) and European Regional Development Fund; Czech Academy of Sciences (RVO:67985912); the Zoological Institute ZIN RAS (АААА-А19-119032590102-7); and King Saud University Researchers Supporting Project (NSRSP–2020/2). The research was carried out with the financial support of the Russian Foundation for Basic Research (19-59-15001 and 20-04-00213), the Russian Science Foundation (16-18-10265, 20-78-10151, and 21-18-00457), the Government of the Russian Federation (FENU-2020-0021), the Estonian Research Council (PRG29), the Estonian Ministry of Education and Research (PRG1209), the Hungarian Scientific Research Fund (Project NF 104792), the Hungarian Academy of Sciences (Momentum Mobility Research Project of the Institute of Archaeology, Research Centre for the Humanities); and the Polish National Science Centre (2013/11/B/HS3/03822). This project has received funding from the European Union's Horizon 2020 research and innovation programme under the Marie Skłodowska-Curie (grant agreement 797449). This project has received funding from the European Research Council (ERC) under the European Union's Horizon 2020 research and innovation programme (grant agreements 681605, 716732 and 834616). ; Peer reviewed