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    The genomic formation of South and Central Asia

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    The genetic formation of Central and South Asian populations has been unclear because of an absence of ancient DNA. To address this gap, we generated genome-wide data from 362 ancient individuals, including the first from eastern Iran, Turan (Uzbekistan, Turkmenistan, and Tajikistan), Bronze Age Kazakhstan, and South Asia. Our data reveal a complex set of genetic sources that ultimately combined to form the ancestry of South Asians today. We document a southward spread of genetic ancestry from the Eurasian Steppe, correlating with the archaeologically known expansion of pastoralist sites from the Steppe to Turan in the Middle Bronze Age (2300-1500 BCE). These Steppe communities mixed genetically with peoples of the Bactria Margiana Archaeological Complex (BMAC) whom they encountered in Turan (primarily descendants of earlier agriculturalists of Iran), but there is no evidence that the main BMAC population contributed genetically to later South Asians. Instead, Steppe communities integrated farther south throughout the 2nd millennium BCE, and we show that they mixed with a more southern population that we document at multiple sites as outlier individuals exhibiting a distinctive mixture of ancestry related to Iranian agriculturalists and South Asian hunter-gathers. We call this group Indus Periphery because they were found at sites in cultural contact with the Indus Valley Civilization (IVC) and along its northern fringe, and also because they were genetically similar to post-IVC groups in the Swat Valley of Pakistan. By co-analyzing ancient DNA and genomic data from diverse present-day South Asians, we show that Indus Periphery-related people are the single most important source of ancestry in South Asia —} consistent with the idea that the Indus Periphery individuals are providing us with the first direct look at the ancestry of peoples of the IVC {— and we develop a model for the formation of present-day South Asians in terms of the temporally and geographically proximate sources of Indus Periphery-related, Steppe, and local South Asian hunter-gatherer-related ancestry. Our results show how ancestry from the Steppe genetically linked Europe and South Asia in the Bronze Age, and identifies the populations that almost certainly were responsible for spreading Indo-European languages across much of Eurasia

    The formation of human populations in South and Central Asia

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    By sequencing 523 ancient humans, we show that the primary source of ancestry in modern South Asians is a prehistoric genetic gradient between people related to early hunter-gatherers of Iran and Southeast Asia. After the Indus Valley Civilization's decline, its people mixed with individuals in the southeast to form one of the two main ancestral populations of South Asia, whose direct descendants live in southern India. Simultaneously, they mixed with descendants of Steppe pastoralists who, starting around 4000 years ago, spread via Central Asia to form the other main ancestral population. The Steppe ancestry in South Asia has the same profile as that in Bronze Age Eastern Europe, tracking a movement of people that affected both regions and that likely spread the distinctive features shared between Indo-Iranian and Balto-Slavic languages.N.P. carried out this work while a fellow at the Radcliffe Institute for Advanced Study at Harvard University. P.M. was supported by a Burroughs Wellcome Fund CASI award. N.N. is supported by a NIGMS (GM007753) fellowship. T.C. and A.D. were supported by the Russian Science Foundation (project 14-50-00036). T.M.S. was supported by the Russian Foundation for Basic Research (grant 18-09-00779) “Anthropological and archaeological aspects of ethnogenesis of the population of the southern part of Western and Central Siberia in the Neolithic and Early Bronze Age.” D.P., S.S., and D.L. were supported by European Research Council ERC-2011-AdG 295733 grant (Langelin). O.M. was supported by a grant from the Ministry of Education and Sciences of the Russian Federation No. 33.1907, 2017/Π4 “Traditional and innovational models of a development of ancient Volga population”. A.E. was supported by a grant from the Ministry of Education and Sciences of the Russian Federation No. 33.5494, 2017/BP “Borderlands of cultural worlds (Southern Urals from Antiquity to Early Modern period).” Radiocarbon dating work supported by the NSF Archaeometry program BCS-1460369 to D.Ken. and B.J.C. and by the NSF Archaeology program BCS-1725067 to D.Ken. K.Th. was supported by NCP fund (MLP0117) of the Council of Scientific and Industrial Research (CSIR), Government of India, New Delhi. N.Bo., A.N., and M.Z. were supported by the Max Planck Society. D.Re. is an Investigator of the Howard Hughes Medical Institute, and his ancient DNA laboratory work was supported by National Science Foundation HOMINID grant BCS-1032255, by National Institutes of Health grant GM100233, by an Allen Discovery Center grant, and by grant 61220 from the John Templeton Foundation
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