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

      research-article
      1 , 1 , 121 , 1 , 1 , 2 , 1 , 3 , 1 , 1 , 1 , 1 , 4 , 5 , 1 , 1 , 1 , 1 , 6 , 7 , 8 , 9 , 10 , 11 , 11 , 11 , 12 , 13 , 14 , 15 , 15 , 15 , 16 , 17 , 18 , 19 , 20 , 3 , 21 , 21 , 22 , 22 , 23 , 24 , 25 , 26 , 27 , 28 , 29 , 30 , 30 , 30 , 31 , 32 , 33 , 31 , 31 , 34 , 35 , 33 , 33 , 36 , 37 , 38 , 39 , 40 , 41 , 42 , 42 , 43 , 44 , 45 , 44 , 45 , 43 , 46 , 46 , 47 , 48 , 49 , 50 , 51 , 52 , 53 , 43 , 54 , 55 , 56 , 57 , 58 , 59 , 60 , 59 , 61 , 56 , 17 , 62 , 63 , 64 , 65 , 66 , 67 , 68 , 69 , 70 , 71 , 71 , 72 , 70 , 73 , 74 , 75 , 76 , 77 , 77 , 78 , 79 , 80 , 81 , 82 , 83 , 83 , 83 , 83 , 42 , 42 , 84 , 42 , 84 , 85 , 86 , 87 , 49 , 86 , 49 , 86 , 88 , 89 , 90 , 91 , 92 , 93 , 2 , 94 , 95 , 9 , 59 , 43 , 96 , 9 , 97 , 98 , 99 , 100 , 99 , 100 , 101 , 101 , 102 , 102 , 103 , 103 , 104 , 104 , 105 , 103 , 103 , 106 , 107 , 108 , 108 , 90 , 90 , 109 , 110 , 111 , 112 , 113 , 110 , 98 , 114 , 110 , 115 , 116 , 117 , 118 , 119 , 120 , 5 , 75 , 1 ,
      Nature
      Nature Publishing Group UK
      Evolutionary genetics, Population genetics, Population genetics

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          Abstract

          Domestication of horses fundamentally transformed long-range mobility and warfare 1 . However, modern domesticated breeds do not descend from the earliest domestic horse lineage associated with archaeological evidence of bridling, milking and corralling 24 at Botai, Central Asia around 3500 bc 3 . Other longstanding candidate regions for horse domestication, such as Iberia 5 and Anatolia 6 , 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 7 between horseback riding and the massive expansion of Yamnaya steppe pastoralists into Europe around 3000 bc 8, 9 driving the spread of Indo-European languages 10 . 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 .

          Abstract

          Analysis of 273 ancient horse genomes reveals that modern domestic horses originated in the Western Eurasian steppes, especially the lower Volga-Don region.

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          Most cited references73

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          MAFFT Multiple Sequence Alignment Software Version 7: Improvements in Performance and Usability

          We report a major update of the MAFFT multiple sequence alignment program. This version has several new features, including options for adding unaligned sequences into an existing alignment, adjustment of direction in nucleotide alignment, constrained alignment and parallel processing, which were implemented after the previous major update. This report shows actual examples to explain how these features work, alone and in combination. Some examples incorrectly aligned by MAFFT are also shown to clarify its limitations. We discuss how to avoid misalignments, and our ongoing efforts to overcome such limitations.
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            RAxML version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies

            Motivation: Phylogenies are increasingly used in all fields of medical and biological research. Moreover, because of the next-generation sequencing revolution, datasets used for conducting phylogenetic analyses grow at an unprecedented pace. RAxML (Randomized Axelerated Maximum Likelihood) is a popular program for phylogenetic analyses of large datasets under maximum likelihood. Since the last RAxML paper in 2006, it has been continuously maintained and extended to accommodate the increasingly growing input datasets and to serve the needs of the user community. Results: I present some of the most notable new features and extensions of RAxML, such as a substantial extension of substitution models and supported data types, the introduction of SSE3, AVX and AVX2 vector intrinsics, techniques for reducing the memory requirements of the code and a plethora of operations for conducting post-analyses on sets of trees. In addition, an up-to-date 50-page user manual covering all new RAxML options is available. Availability and implementation: The code is available under GNU GPL at https://github.com/stamatak/standard-RAxML. Contact: alexandros.stamatakis@h-its.org Supplementary information: Supplementary data are available at Bioinformatics online.
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              IQ-TREE: A Fast and Effective Stochastic Algorithm for Estimating Maximum-Likelihood Phylogenies

              Large phylogenomics data sets require fast tree inference methods, especially for maximum-likelihood (ML) phylogenies. Fast programs exist, but due to inherent heuristics to find optimal trees, it is not clear whether the best tree is found. Thus, there is need for additional approaches that employ different search strategies to find ML trees and that are at the same time as fast as currently available ML programs. We show that a combination of hill-climbing approaches and a stochastic perturbation method can be time-efficiently implemented. If we allow the same CPU time as RAxML and PhyML, then our software IQ-TREE found higher likelihoods between 62.2% and 87.1% of the studied alignments, thus efficiently exploring the tree-space. If we use the IQ-TREE stopping rule, RAxML and PhyML are faster in 75.7% and 47.1% of the DNA alignments and 42.2% and 100% of the protein alignments, respectively. However, the range of obtaining higher likelihoods with IQ-TREE improves to 73.3-97.1%.
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                Author and article information

                Contributors
                ludovic.orlando@univ-tlse3.fr
                Journal
                Nature
                Nature
                Nature
                Nature Publishing Group UK (London )
                0028-0836
                1476-4687
                20 October 2021
                20 October 2021
                2021
                : 598
                : 7882
                : 634-640
                Affiliations
                [1 ]GRID grid.15781.3a, ISNI 0000 0001 0723 035X, Centre d’Anthropobiologie et de Génomique de Toulouse, , Université Paul Sabatier, ; Toulouse, France
                [2 ]GRID grid.415877.8, ISNI 0000 0001 2254 1834, Department of the Diversity and Evolution of Genomes, , Institute of Molecular and Cellular Biology SB RAS, ; Novosibirsk, Russia
                [3 ]GRID grid.413454.3, ISNI 0000 0001 1958 0162, W. Szafer Institute of Botany, , Polish Academy of Sciences, ; Kraków, Poland
                [4 ]GRID grid.460789.4, ISNI 0000 0004 4910 6535, Genoscope, Institut de biologie François-Jacob, Commissariat à l’Energie Atomique (CEA), , Université Paris-Saclay, ; Evry, France
                [5 ]GRID grid.8390.2, ISNI 0000 0001 2180 5818, Génomique Métabolique, Genoscope, Institut de biologie François Jacob, CEA, CNRS, , Université d’Evry, Université Paris-Saclay, ; Evry, France
                [6 ]GRID grid.266093.8, ISNI 0000 0001 0668 7243, Earth System Science Department, , University of California, Irvine, ; Irvine, CA USA
                [7 ]GRID grid.205975.c, ISNI 0000 0001 0740 6917, Department of Ecology and Evolutionary Biology, , University of California, Santa Cruz, ; Santa Cruz, CA USA
                [8 ]GRID grid.205975.c, ISNI 0000 0001 0740 6917, Howard Hughes Medical Institute, , University of California, Santa Cruz, ; Santa Cruz, CA USA
                [9 ]GRID grid.77225.35, ISNI 0000000112611077, Department of Archaeology, Ethnography and Museology, , Altai State University, ; Barnaul, Russia
                [10 ]GRID grid.465449.e, ISNI 0000 0001 1214 1108, Department of Archaeological Heritage Preservation, , Institute of Archaeology of the Russian Academy of Sciences, ; Moscow, Russia
                [11 ]GRID grid.56302.32, ISNI 0000 0004 1773 5396, Zoology Department, College of Science, , King Saud University, ; Riyadh, Saudi Arabia
                [12 ]GRID grid.7359.8, ISNI 0000 0001 2325 4853, Institute for Archaeology, Heritage Conservation Studies and Art History, , University of Bamberg, ; Bamberg, Germany
                [13 ]Museum Østjylland, Randers, Denmark
                [14 ]GRID grid.5254.6, ISNI 0000 0001 0674 042X, Saxo Institute, section of Archaeology, , University of Copenhagen, ; Copenhagen, Denmark
                [15 ]GRID grid.4444.0, ISNI 0000 0001 2112 9282, ArScAn-UMR 7041, Equipe Ethnologie préhistorique, CNRS, MSH-Mondes, ; Nanterre Cedex, France
                [16 ]Muséum d’histoire naturelle, Secteur des Vertébrés, Geneva, Switzerland
                [17 ]GRID grid.412041.2, ISNI 0000 0001 2106 639X, UMR 5199 De la Préhistoire à l’Actuel : Culture, Environnement et Anthropologie (PACEA), CNRS, , Université de Bordeaux, ; Pessac Cedex, France
                [18 ]GRID grid.466902.f, ISNI 0000 0001 2248 6951, Geneva Natural History Museum, ; Geneva, Switzerland
                [19 ]GRID grid.8591.5, ISNI 0000 0001 2322 4988, Department of Genetics and Evolution, , University of Geneva, ; Geneva, Switzerland
                [20 ]GRID grid.20478.39, ISNI 0000 0001 2171 9581, OD Earth & History of Life, , Royal Belgian Institute of Natural Sciences, ; Brussels, Belgium
                [21 ]GRID grid.413454.3, ISNI 0000 0001 1958 0162, Institute of Systematics and Evolution of Animals, , Polish Academy of Sciences, ; Kraków, Poland
                [22 ]GRID grid.413454.3, ISNI 0000 0001 1958 0162, Institute of Archaeology and Ethnology Polish Academy of Sciences, ; Kraków, Poland
                [23 ]GRID grid.5522.0, ISNI 0000 0001 2162 9631, Institute of Archaeology, , Jagiellonian University, ; Kraków, Poland
                [24 ]Department of Archaeology, Institute of History and Archaeology, Tartu, Estonia
                [25 ]GRID grid.10939.32, ISNI 0000 0001 0943 7661, Department of Zoology, Institute of Ecology and Earth Sciences, , University of Tartu, ; Tartu, Estonia
                [26 ]Diamond and Precious Metals Geology Institute, SB RAS, Yakutsk, Russia
                [27 ]GRID grid.8207.d, ISNI 0000 0000 9774 6466, Archaeological Research Collection, , Tallinn University, ; Tallinn, Estonia
                [28 ]GRID grid.447879.1, ISNI 0000 0001 0792 540X, Department of Natural Sciences and Archaeometry, , Institute of Archaeology of the Czech Academy of Sciences, ; Prague, Czechia
                [29 ]Prague, Czechia
                [30 ]GRID grid.418333.e, ISNI 0000 0004 1937 1389, Vasile Pârvan Institute of Archaeology, Department of Bioarchaeology, , Romanian Academy, ; Bucharest, Romania
                [31 ]GRID grid.481823.4, Institute of Archaeogenomics, Research Centre for the Humanities, , Eötvös Loránd Research Network, ; Budapest, Hungary
                [32 ]GRID grid.5591.8, ISNI 0000 0001 2294 6276, Department of Genetics, , Eötvös Loránd University, ; Budapest, Hungary
                [33 ]GRID grid.481830.6, ISNI 0000 0001 2238 5843, Institute of Archaeology, Research Centre for the Humanities, , Eötvös Loránd Research Network, ; Budapest, Hungary
                [34 ]Ásatárs Ltd., Kecskemét, Hungary
                [35 ]Rippl-Rónai Municipal Museum with Country Scope, Kaposvár, Hungary
                [36 ]GRID grid.4305.2, ISNI 0000 0004 1936 7988, School of History, Classics and Archaeology, , University of Edinburgh, Old Medical School, ; Edinburgh, UK
                [37 ]GRID grid.1032.0, ISNI 0000 0004 0375 4078, Trace and Environmental DNA (TrEnD) Lab, School of Molecular and Life Sciences, , Curtin University, ; Perth, Western Australia Australia
                [38 ]GRID grid.5254.6, ISNI 0000 0001 0674 042X, Lundbeck Foundation GeoGenetics Centre, GLOBE Institute, , University of Copenhagen, ; Copenhagen, Denmark
                [39 ]GRID grid.435140.7, Department of Academic Management, , Academy of Science of Moldova, ; Chișinău, Republic of Moldova
                [40 ]GRID grid.435140.7, Center of Archaeology, Institute of Cultural Heritage, , Academy of Science of Moldova, ; Chișinău, Republic of Moldova
                [41 ]GRID grid.446391.d, ISNI 0000 0001 2190 3450, Archaeological Institute of America, ; Boston, MA USA
                [42 ]Centre National de Recherche Scientifique, Muséum national d’Histoire naturelle, Archéozoologie, Archéobotanique (AASPE), CP 56, Paris, France
                [43 ]GRID grid.473277.2, ISNI 0000 0001 2291 1890, Institute for the History of Material Culture, Russian Academy of Sciences (IHMC RAS), ; St Petersburg, Russia
                [44 ]GRID grid.465388.4, Geological Institute, Russian Academy of Sciences, ; Moscow, Russia
                [45 ]GRID grid.424187.c, ISNI 0000 0001 1942 9788, Arctic and Antarctic Research Institute, ; St Petersburg, Russia
                [46 ]GRID grid.6583.8, ISNI 0000 0000 9686 6466, Institute of Animal Breeding and Genetics, , University of Veterinary Medicine Vienna, ; Vienna, Austria
                [47 ]GRID grid.466489.1, ISNI 0000 0001 2151 4674, Department of Prehistory and Western Asian/Northeast African Archaeology, , Austrian Archaeological Institute, Austrian Academy of Sciences, ; Vienna, Austria
                [48 ]GRID grid.10939.32, ISNI 0000 0001 0943 7661, Estonian Biocentre, Institute of Genomics, , University of Tartu, ; Tartu, Estonia
                [49 ]GRID grid.469873.7, ISNI 0000 0004 4914 1197, Department of Archaeogenetics, , Max Planck Institute for the Science of Human History, ; Jena, Germany
                [50 ]GRID grid.10392.39, ISNI 0000 0001 2190 1447, SFB 1070 Resource Cultures, , University of Tübingen, ; Tübingen, Germany
                [51 ]GRID grid.10392.39, ISNI 0000 0001 2190 1447, Department of Early Prehistory and Quaternary Ecology, , University of Tübingen, ; Tübingen, Germany
                [52 ]GRID grid.4444.0, ISNI 0000 0001 2112 9282, UMR 7194 Muséum National d’Histoire Naturelle, CNRS, UPVD, ; Paris, France
                [53 ]GRID grid.459698.f, ISNI 0000 0000 8989 8101, Semenov-Tyan-Shanskii Lipetsk State Pedagogical University, ; Lipetsk, Russia
                [54 ]GRID grid.266190.a, ISNI 0000000096214564, Museum of Natural History, , University of Colorado-Boulder, ; Boulder, CO USA
                [55 ]Musée d’Anthropologie préhistorique de Monaco, Monaco, Monaco
                [56 ]GRID grid.425564.4, ISNI 0000 0004 0587 3863, Institute of Archaeology, Mongolian Academy of Sciences, ; Ulaanbaatar, Mongolia
                [57 ]GRID grid.469873.7, ISNI 0000 0004 4914 1197, Department of Archaeology, , Max Planck Institute for the Science of Human History, ; Jena, Germany
                [58 ]Chinggis Khaan Museum, Ulaanbaatar, Mongolia
                [59 ]Department of Archaeology, Ulaanbaatar State University, Ulaanbaatar, Mongolia
                [60 ]GRID grid.444269.9, ISNI 0000 0004 0387 4627, Department of History, , Kyrgyz-Turkish Manas University, ; Bishkek, Kyrgyzstan
                [61 ]GRID grid.260731.1, ISNI 0000 0001 2324 0259, Department of Biology, , National University of Mongolia, ; Ulaanbaatar, Mongolia
                [62 ]GRID grid.266515.3, ISNI 0000 0001 2106 0692, Division of Archaeology, Biodiversity Institute, , University of Kansas, ; Lawrence, KS USA
                [63 ]GRID grid.9764.c, ISNI 0000 0001 2153 9986, Institute for Prehistoric and Protohistoric Archaeology, , Kiel University, ; Kiel, Germany
                [64 ]GRID grid.9764.c, ISNI 0000 0001 2153 9986, ROOTS Excellence Cluster, , Kiel University, ; Kiel, Germany
                [65 ]GRID grid.4711.3, ISNI 0000 0001 2183 4846, Archaeology of Social Dynamics, Institució Milà i Fontanals d’Humanitats, Consejo Superior de Investigaciones Científicas (IMF-CSIC), ; Barcelona, Spain
                [66 ]GRID grid.5841.8, ISNI 0000 0004 1937 0247, Departament d’Història i Arqueologia–SERP, , Universitat de Barcelona, ; Barcelona, Spain
                [67 ]GRID grid.15043.33, ISNI 0000 0001 2163 1432, Grup d’Investigació Prehistòrica, , Universitat de Lleida, ; PID2019-110022GB-I00, Lleida, Spain
                [68 ]Valencia, Spain
                [69 ]GRID grid.8393.1, ISNI 0000000119412521, Departamento de Medicina Animal, Facultad de Veterinaria, , Universidad de Extremadura, ; Cáceres, Spain
                [70 ]Centro Mixto UCM-ISCIII de Evolución y Comportamiento Humanos, Madrid, Spain
                [71 ]GRID grid.454770.5, ISNI 0000 0001 1945 3489, Instituto de Arqueología (CSIC–Junta de Extremadura), ; Mérida, Spain
                [72 ]GRID grid.9612.c, ISNI 0000 0001 1957 9153, Laboratori d’Arqueologia Prehistòrica, , Universitat Jaume I, ; Castelló de la Plana, Spain
                [73 ]GRID grid.4795.f, ISNI 0000 0001 2157 7667, Departamento de Geodinámica, Estratigrafía y Paleontología, Facultad de Ciencias Geológicas, , Universidad Complutense de Madrid, ; Madrid, Spain
                [74 ]GRID grid.418751.e, ISNI 0000 0004 0385 8977, Department of Eneolithic and Bronze Age, , Institute of Archaeology National Academy of Sciences of Ukraine, ; Kyiv, Ukraine
                [75 ]GRID grid.8391.3, ISNI 0000 0004 1936 8024, Department of Archaeology, , University of Exeter, ; Exeter, UK
                [76 ]GRID grid.137628.9, ISNI 0000 0004 1936 8753, Center for the Study of Human Origins, Anthropology Department, , New York University, ; New York, NY USA
                [77 ]GRID grid.77184.3d, ISNI 0000 0000 8887 5266, Department of Archaeology, Ethnology and Museology, , Al Farabi Kazakh National University, ; Almaty, Kazakhstan
                [78 ]GRID grid.445474.2, ISNI 0000 0001 1092 7131, Scientific Research Department, , Orenburg State Pedagogical University, ; Orenburg, Russia
                [79 ]GRID grid.14476.30, ISNI 0000 0001 2342 9668, Department of paleontology, , Faculty of Geology, Moscow State University, ; Moscow, Russia
                [80 ]GRID grid.77268.3c, ISNI 0000 0004 0543 9688, Institute of Geology and Petroleum Technologies, , Kazan Federal University, ; Kazan, Russia
                [81 ]GRID grid.452450.2, ISNI 0000 0001 0739 408X, Georgian National Museum, ; Tbilisi, Georgia
                [82 ]GRID grid.26193.3f, ISNI 0000 0001 2034 6082, Tbilisi State University, ; Tbilisi, Georgia
                [83 ]GRID grid.4708.b, ISNI 0000 0004 1757 2822, Università degli Studi di Milano, Dipartimento di Beni Culturali e Ambientali, ; Milan, Italy
                [84 ]GRID grid.46072.37, ISNI 0000 0004 0612 7950, University of Tehran, Central Laboratory, Bioarchaeology Laboratory, Archaeozoology Section, ; Tehran, Iran
                [85 ]GRID grid.14476.30, ISNI 0000 0001 2342 9668, Research Institute and Museum of Anthropology, , Lomonosov Moscow State University, ; Moscow, Russia
                [86 ]GRID grid.419518.0, ISNI 0000 0001 2159 1813, Department of Archaeogenetics, , Max Planck Institute for Evolutionary Anthropology, ; Leipzig, Germany
                [87 ]GRID grid.5252.0, ISNI 0000 0004 1936 973X, Institute for Pre- and Protohistoric Archaeology and Archaeology of the Roman Provinces, , Ludwig Maximilian University, Munich, ; Munich, Germany
                [88 ]GRID grid.1010.0, ISNI 0000 0004 1936 7304, School of Biological Sciences, , The University of Adelaide, ; Adelaide, South Australia Australia
                [89 ]GRID grid.5515.4, ISNI 0000000119578126, Department of Biology, , Universidad Autónoma de Madrid, ; Madrid, Spain
                [90 ]GRID grid.424195.f, ISNI 0000 0001 2106 6832, Eurasia Department of the German Archaeological Institute, ; Berlin, Germany
                [91 ]GRID grid.11348.3f, ISNI 0000 0001 0942 1117, Evolutionary Adaptive Genomics, Institute of Biochemistry and Biology, Faculty of Mathematics and Science, , University of Potsdam, ; Potsdam, Germany
                [92 ]GRID grid.418779.4, ISNI 0000 0001 0708 0355, Department of Evolutionary Genetics, , Leibniz-Institute for Zoo and Wildlife Research, ; Berlin, Germany
                [93 ]GRID grid.7468.d, ISNI 0000 0001 2248 7639, Albrecht Daniel Thaer-Institute, Faculty of Life Sciences, , Humboldt University Berlin, ; Berlin, Germany
                [94 ]GRID grid.5252.0, ISNI 0000 0004 1936 973X, ArchaeoBioCenter and Institute of Palaeoanatomy, Domestication Research and the History of Veterinary Medicine, , LMU Munich, ; Munich, Germany
                [95 ]GRID grid.452781.d, ISNI 0000 0001 2203 6205, SNSB, State Collection of Anthropology and Palaeoanatomy, ; Munich, Germany
                [96 ]GRID grid.415877.8, ISNI 0000 0001 2254 1834, ArchaeoZOOlogy in Siberia and Central Asia—ZooSCAn International Research Laboratory, Institute of Archeology and Ethnography of the Siberian Branch of the RAS, ; Novosibirsk, Russia
                [97 ]GRID grid.426493.e, ISNI 0000 0004 1800 742X, Department of Eastern European and Siberian Archaeology, , State Hermitage Museum, ; St Petersburg, Russia
                [98 ]GRID grid.482778.6, ISNI 0000 0001 2197 0186, Paleoecology Laboratory, Institute of Plant and Animal Ecology, Ural Branch of the Russian Academy of Sciences, ; Ekaterinburg, Russia
                [99 ]GRID grid.439287.3, ISNI 0000 0001 2314 7601, Zoological Institute, Russian Academy of Sciences, ; St Petersburg, Russia
                [100 ]GRID grid.6441.7, ISNI 0000 0001 2243 2806, Department of Archaeology, History Faculty, , Vilnius University, ; Vilnius, Lithuania
                [101 ]GRID grid.443586.8, Laboratory for Archaeological Research, Faculty of History and Law, , Kostanay State University, ; Kostanay, Kazakhstan
                [102 ]Department of History and Archaeology, Surgut Governmental University, Surgut, Russia
                [103 ]Saryarka Archaeological Institute, Buketov Karaganda University, Karaganda, Kazakhstan
                [104 ]Toraighyrov University, Joint Research Center for Archeological Studies, Pavlodar, Kazakhstan
                [105 ]GRID grid.55380.3b, ISNI 0000 0004 0398 5415, Faculty of History, L. N. Gumilev Eurasian National University, ; Nur-Sultan, Kazakhstan
                [106 ]GRID grid.77184.3d, ISNI 0000 0000 8887 5266, Institute of Archaeology and Steppe Civilizations, , Al-Farabi Kazakh National University, ; Almaty, Kazakhstan
                [107 ]GRID grid.10698.36, ISNI 0000000122483208, Department of Anthropology, Alumni Building, , University of North Carolina at Chapel Hill, ; Chapel Hill, NC USA
                [108 ]Nasledie Cultural Heritage Unit, Stavropol, Russia
                [109 ]Research Center for the Preservation of Cultural Heritage, Saratov, Russia
                [110 ]GRID grid.445790.b, ISNI 0000 0001 2218 2982, Department of Russian History and Archaeology, , Samara State University of Social Sciences and Education, ; Samara, Russia
                [111 ]GRID grid.440724.1, ISNI 0000 0000 9958 5862, Russian and Foreign History Department, , South Ural State University, ; Chelyabinsk, Russia
                [112 ]GRID grid.465317.2, ISNI 0000 0001 2224 8785, South Ural Department, , Institute of History and Archaeology, Ural Branch of the Russian Academy of Sciences, ; Ekaterinburg, Russia
                [113 ]Archaeological School, Chuvash State Institute of Humanities, Cheboksary, Russia
                [114 ]GRID grid.412761.7, ISNI 0000 0004 0645 736X, Department of History of the Institute of Humanities, , Ural Federal University, ; Ekaterinburg, Russia
                [115 ]GRID grid.38142.3c, ISNI 000000041936754X, Department of Human Evolutionary Biology, , Harvard University, ; Cambridge, MA USA
                [116 ]GRID grid.418410.8, ISNI 0000 0001 0115 6427, Anthropology Faculty, , Hartwick College, ; Oneonta NY, USA
                [117 ]GRID grid.5254.6, ISNI 0000 0001 0674 042X, Department of Nordic Studies and Linguistics, , University of Copenhagen, ; Copenhagen, Denmark
                [118 ]GRID grid.5132.5, ISNI 0000 0001 2312 1970, Leiden University Center for Linguistics, , Leiden University, ; Leiden, The Netherlands
                [119 ]GRID grid.8761.8, ISNI 0000 0000 9919 9582, Department of Historical Studies, , University of Gothenburg, ; Gothenburg, Sweden
                [120 ]GRID grid.452548.a, ISNI 0000 0000 9817 5300, Present Address: Lundbeck Foundation GeoGenetics Centre, ; Copenhagen, Denmark
                [121 ]GRID grid.440522.5, ISNI 0000 0004 0478 6450, Department of Biotechnology, , Abdul Wali Khan University, ; Mardan, Pakistan
                Author information
                http://orcid.org/0000-0002-0811-8754
                http://orcid.org/0000-0003-1718-3010
                http://orcid.org/0000-0003-2637-3131
                http://orcid.org/0000-0002-3404-3397
                http://orcid.org/0000-0001-7618-625X
                http://orcid.org/0000-0001-8865-0937
                http://orcid.org/0000-0002-9786-0693
                http://orcid.org/0000-0003-2011-2141
                http://orcid.org/0000-0002-4226-3218
                http://orcid.org/0000-0001-5672-2756
                http://orcid.org/0000-0003-4159-0695
                http://orcid.org/0000-0002-6310-4056
                http://orcid.org/0000-0002-6998-8454
                http://orcid.org/0000-0002-3785-6342
                http://orcid.org/0000-0002-0836-7814
                http://orcid.org/0000-0002-5768-0698
                http://orcid.org/0000-0003-0015-8142
                http://orcid.org/0000-0001-6606-8516
                http://orcid.org/0000-0001-7824-2155
                http://orcid.org/0000-0002-1649-336X
                http://orcid.org/0000-0001-9886-0372
                http://orcid.org/0000-0001-6194-0475
                http://orcid.org/0000-0002-4315-7257
                http://orcid.org/0000-0002-5236-1444
                http://orcid.org/0000-0003-3630-9459
                http://orcid.org/0000-0001-9144-3920
                http://orcid.org/0000-0003-2475-2007
                http://orcid.org/0000-0001-7249-9953
                http://orcid.org/0000-0003-0894-2628
                http://orcid.org/0000-0002-2773-7454
                http://orcid.org/0000-0003-1901-1001
                http://orcid.org/0000-0001-6951-0509
                http://orcid.org/0000-0002-5445-5516
                http://orcid.org/0000-0002-8107-6300
                http://orcid.org/0000-0002-1888-2713
                http://orcid.org/0000-0002-3708-0476
                http://orcid.org/0000-0001-7562-3454
                http://orcid.org/0000-0003-3360-089X
                http://orcid.org/0000-0003-3936-1850
                Article
                4018
                10.1038/s41586-021-04018-9
                8550961
                34671162
                84a4ee77-36e6-4cd2-a6fb-69f005338eee
                © The Author(s) 2021

                Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. The images or other third party material in this article are included in the article’s Creative Commons license, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons license and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this license, visit http://creativecommons.org/licenses/by/4.0/.

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                : 4 May 2021
                : 10 September 2021
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                © The Author(s), under exclusive licence to Springer Nature Limited 2021

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                evolutionary genetics,population genetics
                Uncategorized
                evolutionary genetics, population genetics

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