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Research Article

The formation of human populations in South and Central Asia

Vagheesh M. Narasimhan https://orcid.org/0000-0001-8651-8844 [email protected], Nick Patterson https://orcid.org/0000-0003-0302-684X [email protected], Priya Moorjani, Nadin Rohland, Rebecca Bernardos https://orcid.org/0000-0003-4625-3727, Swapan Mallick https://orcid.org/0000-0002-4531-4439, Iosif Lazaridis, Nathan Nakatsuka https://orcid.org/0000-0002-5091-4914, Iñigo Olalde, Mark Lipson https://orcid.org/0000-0001-5346-9329, Alexander M. Kim https://orcid.org/0000-0001-5894-8062, Luca M. Olivieri https://orcid.org/0000-0003-0484-7332, Alfredo Coppa, Massimo Vidale https://orcid.org/0000-0002-0543-6511, James Mallory, Vyacheslav Moiseyev, Egor Kitov https://orcid.org/0000-0002-0159-3288, Janet Monge https://orcid.org/0000-0002-4364-6296, Nicole Adamski https://orcid.org/0000-0002-2268-1359, Neel Alex, Nasreen Broomandkhoshbacht https://orcid.org/0000-0001-5350-1608, Francesca Candilio https://orcid.org/0000-0002-4668-1361, Kimberly Callan https://orcid.org/0000-0003-3170-8514, Olivia Cheronet https://orcid.org/0000-0001-6760-1204, Brendan J. Culleton, Matthew Ferry https://orcid.org/0000-0001-7319-0085, Daniel Fernandes https://orcid.org/0000-0002-7434-6552, Suzanne Freilich https://orcid.org/0000-0003-0472-6311, Beatriz Gamarra https://orcid.org/0000-0003-3764-497X, Daniel Gaudio https://orcid.org/0000-0002-2009-1574, Mateja Hajdinjak, Éadaoin Harney, Thomas K. Harper, Denise Keating, Ann Marie Lawson https://orcid.org/0000-0003-0990-2329, Matthew Mah https://orcid.org/0000-0001-8987-6436, Kirsten Mandl https://orcid.org/0000-0002-2078-0775, Megan Michel, Mario Novak https://orcid.org/0000-0002-4567-8742, Jonas Oppenheimer https://orcid.org/0000-0001-7973-6173, Niraj Rai, Kendra Sirak, Viviane Slon https://orcid.org/0000-0001-6485-7159, Kristin Stewardson, Fatma Zalzala https://orcid.org/0000-0002-8981-1277, Zhao Zhang https://orcid.org/0000-0002-7562-7559, Gaziz Akhatov, Anatoly N. Bagashev https://orcid.org/0000-0003-2420-6276, Alessandra Bagnera https://orcid.org/0000-0001-8717-7738, Bauryzhan Baitanayev https://orcid.org/0000-0002-2013-0473, Julio Bendezu-Sarmiento, Arman A. Bissembaev, Gian Luca Bonora https://orcid.org/0000-0001-5170-206X, Temirlan T. Chargynov, Tatiana Chikisheva https://orcid.org/0000-0003-1985-1369, Petr K. Dashkovskiy, Anatoly Derevianko https://orcid.org/0000-0003-1156-8331, Miroslav Dobeš, Katerina Douka, Nadezhda Dubova https://orcid.org/0000-0002-4340-1037, Meiram N. Duisengali, Dmitry Enshin, Andrey Epimakhov https://orcid.org/0000-0002-0141-1026, Alexey V. Fribus https://orcid.org/0000-0003-3208-0319, Dorian Fuller https://orcid.org/0000-0002-4859-080X, Alexander Goryachev, Andrey Gromov https://orcid.org/0000-0002-3263-3801, Sergey P. Grushin https://orcid.org/0000-0002-5404-6632, Bryan Hanks https://orcid.org/0000-0002-9740-8162, Margaret Judd https://orcid.org/0000-0002-7409-5597, Erlan Kazizov https://orcid.org/0000-0002-8301-1287, Aleksander Khokhlov https://orcid.org/0000-0003-0442-9616, Aleksander P. Krygin, Elena Kupriyanova, Pavel Kuznetsov https://orcid.org/0000-0002-4898-5350, Donata Luiselli https://orcid.org/0000-0003-2105-2478, Farhod Maksudov https://orcid.org/0000-0002-8492-0634, Aslan M. Mamedov, Talgat B. Mamirov, Christopher Meiklejohn, Deborah C. Merrett https://orcid.org/0000-0001-9539-7271, Roberto Micheli, Oleg Mochalov, Samariddin Mustafokulov https://orcid.org/0000-0002-1700-2497, Ayushi Nayak, Davide Pettener https://orcid.org/0000-0003-1068-2466, Richard Potts https://orcid.org/0000-0001-6008-0100, Dmitry Razhev, Marina Rykun, Stefania Sarno, Tatyana M. Savenkova, Kulyan Sikhymbaeva https://orcid.org/0000-0003-0340-2622, Sergey M. Slepchenko, Oroz A. Soltobaev https://orcid.org/0000-0003-0554-2485, Nadezhda Stepanova https://orcid.org/0000-0003-4017-5641, Svetlana Svyatko, Kubatbek Tabaldiev https://orcid.org/0000-0002-6679-8030, Maria Teschler-Nicola, Alexey A. Tishkin https://orcid.org/0000-0002-7769-136X, Vitaly V. Tkachev https://orcid.org/0000-0002-6355-1745, Sergey Vasilyev https://orcid.org/0000-0003-0128-6568, Petr Velemínský https://orcid.org/0000-0003-3691-7817, Dmitriy Voyakin, Antonina Yermolayeva https://orcid.org/0000-0003-0354-0304, Muhammad Zahir, Valery S. Zubkov, Alisa Zubova, Vasant S. Shinde, Carles Lalueza-Fox https://orcid.org/0000-0002-1730-5914, Matthias Meyer https://orcid.org/0000-0002-4760-558X, David Anthony https://orcid.org/0000-0001-9122-0313, Nicole Boivin, Kumarasamy Thangaraj https://orcid.org/0000-0002-0075-0106, Douglas J. Kennett, Michael Frachetti https://orcid.org/0000-0001-6906-4334 [email protected], Ron Pinhasi https://orcid.org/0000-0003-1629-8131 [email protected], and David Reich https://orcid.org/0000-0002-7037-5292 [email protected]
Science6 Sep 2019Vol 365, Issue 6457DOI: 10.1126/science.aat7487

Ancient human movements through Asia

Ancient DNA has allowed us to begin tracing the history of human movements across the globe. Narasimhan et al. identify a complex pattern of human migrations and admixture events in South and Central Asia by performing genetic analysis of more than 500 people who lived over the past 8000 years (see the Perspective by Schaefer and Shapiro). They establish key phases in the population prehistory of Eurasia, including the spread of farming peoples from the Near East, with movements both westward and eastward. The people known as the Yamnaya in the Bronze Age also moved both westward and eastward from a focal area located north of the Black Sea. The overall patterns of genetic clines reflect similar and parallel patterns in South Asia and Europe.
Science, this issue p. eaat7487; see also p. 981

Structured Abstract

RATIONALE

To elucidate the extent to which the major cultural transformations of farming, pastoralism, and shifts in the distribution of languages in Eurasia were accompanied by movement of people, we report genome-wide ancient DNA data from 523 individuals spanning the last 8000 years, mostly from Central Asia and northernmost South Asia.

RESULTS

The movement of people following the advent of farming resulted in genetic gradients across Eurasia that can be modeled as mixtures of seven deeply divergent populations. A key gradient formed in southwestern Asia beginning in the Neolithic and continuing into the Bronze Age, with more Anatolian farmer–related ancestry in the west and more Iranian farmer–related ancestry in the east. This cline extended to the desert oases of Central Asia and was the primary source of ancestry in peoples of the Bronze Age Bactria Margiana Archaeological Complex (BMAC). This supports the idea that the archaeologically documented dispersal of domesticates was accompanied by the spread of people from multiple centers of domestication.
The main population of the BMAC carried no ancestry from Steppe pastoralists and did not contribute substantially to later South Asians. However, Steppe pastoralist ancestry appeared in outlier individuals at BMAC sites by the turn of the second millennium BCE around the same time as it appeared on the southern Steppe. Using data from ancient individuals from the Swat Valley of northernmost South Asia, we show that Steppe ancestry then integrated further south in the first half of the second millennium BCE, contributing up to 30% of the ancestry of modern groups in South Asia. 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 unique features shared between Indo-Iranian and Balto-Slavic languages.
The primary ancestral population of modern South Asians is a mixture of people related to early Holocene populations of Iran and South Asia that we detect in outlier individuals from two sites in cultural contact with the Indus Valley Civilization (IVC), making it plausible that it was characteristic of the IVC. After the IVC’s decline, this population mixed with northwestern groups with Steppe ancestry to form the “Ancestral North Indians” (ANI) and also mixed with southeastern groups to form the “Ancestral South Indians” (ASI), whose direct descendants today live in tribal groups in southern India. Mixtures of these two post-IVC groups—the ANI and ASI—drive the main gradient of genetic variation in South Asia today.

CONCLUSION

Earlier work recorded massive population movement from the Eurasian Steppe into Europe early in the third millennium BCE, likely spreading Indo-European languages. We reveal a parallel series of events leading to the spread of Steppe ancestry to South Asia, thereby documenting movements of people that were likely conduits for the spread of Indo-European languages.
The Bronze Age spread of Yamnaya Steppe pastoralist ancestry into two subcontinents—Europe and South Asia.
Pie charts reflect the proportion of Yamnaya ancestry, and dates reflect the earliest available ancient DNA with Yamnaya ancestry in each region. Ancient DNA has not yet been found for the ANI and ASI, so for these the range is inferred statistically.

Abstract

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.
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Supplementary Material

Summary

Materials and Methods
Figs. S1 to S61
Tables S1 to S93
Genotypes for Newly Reported Individuals
References (92226)

Resources

File (aat7487_narasimhan_sm.pdf)
File (aat7487_tables1-5.xlsx)

References and Notes

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Volume 365Issue 64576 September 2019
PubMed: 31488661

History

Received: 30 March 2018
Revision received: 19 February 2019
Accepted: 30 July 2019

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Acknowledgments

We acknowledge the people past and present whose samples we analyzed in this study. We thank O. Uberti for the design of Fig. 3 and the print summary figure. We thank the Minusinsk Regional Museum of N. M. Martyanov for sharing skeletal samples. We thank O. Ismagulov, A. Ismagulova, and A. Kunbolot Toktonsunovich for facilitating access to skeletal material. We thank the Department of Archaeology and Museums, Government of Pakistan, the Directorate of Archaeology and Museums, Government of Khyber-Pakhtunkhwa Province (Pakistan), and the ‘S. Sergi’ Museum of the Department of Environmental Biology, Sapienza University (Rome) for facilitating access to the materials from Swat excavated by the Italian Archaeological Mission (now ISMEO). Funding: 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. Author contributions: N.P., P.M., N.Ro., M.Me., N.Bo., K.Th., D.Ken., M.Fr., R.Pi., and D.Re. supervised the study. A.Ki., L.O., A.C., M.V., J.Ma., V.M., E.Ki., J.Mo., G.A., A.Baga., A.Bagn., B.B., J.B., A.Biss., G.B., T.Cha., T.Chi., P.D., A.D., M.Do., K.D., N.D., M.Du., D.E., A.E., A.F., D.Fu., A.Go., A.Gr., S.G., B.H., M.J., E.Ka., A.Kh., A.Kr., E.Ku., P.K., D.L., F.M., A. M., T.M., C.M., D.M., R.M., O.M., S.Mu., A.N., D.P., R.Po., D.Ra., M.R., S.Sa., T.S., K.Sik., S.Sl., O.S., N.S., S.Sv., K.Ta., M.T., A.T., V.T., S.V., P.V., D.V., A.Y., M.Z., V.Z, A.Z., V.Sh., C.L., D.A., N.Bo., M.Fr., and R.Pi. provided samples and assembled archaeological and anthropological information. V.N., N.P., P.M., N.Ro., R.B., S.Ma., I.L., N.N., I.O., M.L., N.Ad., N.A., N.Br., F.C., O.C., B.J.C., M.Fe., D.Fe., S.F., B.Ga., D.G., M.H., E.H., T.H., D.Kea., A.L., M.Ma., K.M., M.Mi., M.N., J.O., N.Ra., K.Sir., V.Sl., K.Ste., Z.Z., M.Me., and D.Re. performed ancient DNA laboratory work or radiocarbon laboratory work or data processing work. V.N., N.P., P.M., I.O., N.Al., S.M., and D.R. analyzed genetic data. V.N., N.P., and D.R. wrote the manuscript with input from all coauthors. Competing interests: The authors declare no competing interests. Data and materials availability: All newly reported sequencing data are available from the European Nucleotide Archive, accession number PRJEB32466, and the software for dating admixture events in ancient samples is available from Zenodo (90). Our online data visualizer is available at https://public.tableau.com/profile/vagheesh#!/vizhome/TheFormationofHumanPopulationsinSouthandCentralAsia/AncientDNA.

Authors

Affiliations

Department of Genetics, Harvard Medical School, Boston, MA 02115, USA.
Broad Institute of Harvard and MIT, Cambridge, MA 02142, USA.
Radcliffe Institute for Advanced Study, Harvard University, Cambridge, MA 02138, USA.
Priya Moorjani
Department of Molecular and Cell Biology, University of California, Berkeley, CA 94720, USA.
Center for Computational Biology, University of California, Berkeley, CA 94720, USA.
Nadin Rohland
Department of Genetics, Harvard Medical School, Boston, MA 02115, USA.
Broad Institute of Harvard and MIT, Cambridge, MA 02142, USA.
Department of Genetics, Harvard Medical School, Boston, MA 02115, USA.
Department of Genetics, Harvard Medical School, Boston, MA 02115, USA.
Broad Institute of Harvard and MIT, Cambridge, MA 02142, USA.
Howard Hughes Medical Institute, Harvard Medical School, Boston, MA 02115, USA.
Iosif Lazaridis
Department of Genetics, Harvard Medical School, Boston, MA 02115, USA.
Department of Genetics, Harvard Medical School, Boston, MA 02115, USA.
Harvard-MIT Division of Health Sciences and Technology, Harvard Medical School, Boston, MA 02115, USA.
Iñigo Olalde
Department of Genetics, Harvard Medical School, Boston, MA 02115, USA.
Department of Genetics, Harvard Medical School, Boston, MA 02115, USA.
Department of Genetics, Harvard Medical School, Boston, MA 02115, USA.
Department of Anthropology, Harvard University, Cambridge, MA 02138, USA.
ISMEO - International Association of Mediterranean and Oriental Studies, Italian Archaeological Mission in Pakistan, 19200 Saidu Sharif (Swat), Pakistan.
Alfredo Coppa
Department of Environmental Biology, Sapienza University, Rome 00185, Italy.
ISMEO - International Association of Mediterranean and Oriental Studies, Italian Archaeological Mission in Pakistan, 19200 Saidu Sharif (Swat), Pakistan.
Department of Cultural Heritage: Archaeology and History of Art, Cinema and Music, University of Padua, Padua 35139, Italy.
James Mallory
School of Natural and Built Environment, Queen’s University Belfast, Belfast BT7 1NN, Northern Ireland, UK.
Vyacheslav Moiseyev
Peter the Great Museum of Anthropology and Ethnography (Kunstkamera), Russian Academy of Science, St. Petersburg 199034, Russia.
Center of Physical Anthropology, Institute of Ethnology and Anthropology, Russian Academy of Sciences, Moscow 119991, Russia.
A.Kh. Margulan Institute of Archaeology, Almaty 050010, Kazakhstan.
Al-Farabi Kazakh National University, Almaty 050040, Kazakhstan.
University of Pennsylvania Museum of Archaeology and Anthropology, Philadelphia, PA 19104, USA.
Department of Genetics, Harvard Medical School, Boston, MA 02115, USA.
Howard Hughes Medical Institute, Harvard Medical School, Boston, MA 02115, USA.
Neel Alex
Department of Electrical Engineering and Computer Science, University of California, Berkeley, CA 94720, USA.
Nasreen Broomandkhoshbacht https://orcid.org/0000-0001-5350-1608
Department of Genetics, Harvard Medical School, Boston, MA 02115, USA.
Howard Hughes Medical Institute, Harvard Medical School, Boston, MA 02115, USA.
Present address: Department of Anthropology, University of California, Santa Cruz, CA 95064, USA.
Earth Institute, University College Dublin, Dublin 4, Ireland.
Soprintendenza Archeologia, Belle Arti e Paesaggio per la Città Metropolitana di Cagliari e le Province di Oristano e Sud Sardegna, Cagliari 09124, Italy.
Department of Genetics, Harvard Medical School, Boston, MA 02115, USA.
Howard Hughes Medical Institute, Harvard Medical School, Boston, MA 02115, USA.
Earth Institute, University College Dublin, Dublin 4, Ireland.
School of Archaeology, University College Dublin, Dublin 4, Ireland.
Department of Evolutionary Anthropology, University of Vienna, 1090 Vienna, Austria.
Brendan J. Culleton
Institutes of Energy and the Environment, Pennsylvania State University, University Park, PA 16802, USA.
Department of Genetics, Harvard Medical School, Boston, MA 02115, USA.
Howard Hughes Medical Institute, Harvard Medical School, Boston, MA 02115, USA.
Earth Institute, University College Dublin, Dublin 4, Ireland.
School of Archaeology, University College Dublin, Dublin 4, Ireland.
Department of Evolutionary Anthropology, University of Vienna, 1090 Vienna, Austria.
CIAS, Department of Life Sciences, University of Coimbra, Coimbra 3000-456, Portugal.
Department of Evolutionary Anthropology, University of Vienna, 1090 Vienna, Austria.
Earth Institute, University College Dublin, Dublin 4, Ireland.
School of Archaeology, University College Dublin, Dublin 4, Ireland.
Catalan Institute of Human Paleoecology and Social Evolution (IPHES), Tarragona 43007, Spain.
Present address: Institut Català de Paleoecologia Humana i Evolució Social (IPHES), 43007 Tarragona, Spain.
Present address: Universitat Rovira i Virgili (URV), Àrea de Prehistòria, 43002 Tarragona, Spain.
Earth Institute, University College Dublin, Dublin 4, Ireland.
School of Archaeology, University College Dublin, Dublin 4, Ireland.
Mateja Hajdinjak
Max Planck Institute for Evolutionary Anthropology, Leipzig 04103, Germany.
Éadaoin Harney
Department of Genetics, Harvard Medical School, Boston, MA 02115, USA.
Howard Hughes Medical Institute, Harvard Medical School, Boston, MA 02115, USA.
Department of Organismic and Evolutionary Biology, Harvard University, Cambridge, MA 02138, USA.
Thomas K. Harper
Department of Anthropology, Pennsylvania State University, University Park, PA 16802, USA.
Denise Keating
Earth Institute, University College Dublin, Dublin 4, Ireland.
Department of Genetics, Harvard Medical School, Boston, MA 02115, USA.
Howard Hughes Medical Institute, Harvard Medical School, Boston, MA 02115, USA.
Department of Genetics, Harvard Medical School, Boston, MA 02115, USA.
Broad Institute of Harvard and MIT, Cambridge, MA 02142, USA.
Howard Hughes Medical Institute, Harvard Medical School, Boston, MA 02115, USA.
Department of Evolutionary Anthropology, University of Vienna, 1090 Vienna, Austria.
Megan Michel
Department of Genetics, Harvard Medical School, Boston, MA 02115, USA.
Howard Hughes Medical Institute, Harvard Medical School, Boston, MA 02115, USA.
Present address: Max Planck-Harvard Research Center for the Archaeoscience of the Ancient Mediterranean, Cambridge, MA 02138, USA.
Present address: Department of Human Evolutionary Biology, Harvard University, Cambridge MA, 02138, USA.
Earth Institute, University College Dublin, Dublin 4, Ireland.
Institute for Anthropological Research, Zagreb 10000, Croatia.
Department of Genetics, Harvard Medical School, Boston, MA 02115, USA.
Howard Hughes Medical Institute, Harvard Medical School, Boston, MA 02115, USA.
Present address: Department of Biomolecular Engineering, University of California, Santa Cruz, CA 95064, USA.
Niraj Rai
CSIR-Centre for Cellular and Molecular Biology, Hyderabad 500 007, India.
Birbal Sahni Institute of Palaeosciences, Lucknow 226007, India.
Kendra Sirak
Department of Genetics, Harvard Medical School, Boston, MA 02115, USA.
Earth Institute, University College Dublin, Dublin 4, Ireland.
Department of Anthropology, Emory University, Atlanta, GA 30322, USA.
Max Planck Institute for Evolutionary Anthropology, Leipzig 04103, Germany.
Kristin Stewardson
Department of Genetics, Harvard Medical School, Boston, MA 02115, USA.
Howard Hughes Medical Institute, Harvard Medical School, Boston, MA 02115, USA.
Department of Genetics, Harvard Medical School, Boston, MA 02115, USA.
Howard Hughes Medical Institute, Harvard Medical School, Boston, MA 02115, USA.
Department of Genetics, Harvard Medical School, Boston, MA 02115, USA.
Gaziz Akhatov
A.Kh. Margulan Institute of Archaeology, Almaty 050010, Kazakhstan.
Tyumen Scientific Centre SB RAS, Institute of the Problems of Northern Development, Tyumen 625003, Russia.
ISMEO - International Association of Mediterranean and Oriental Studies, Italian Archaeological Mission in Pakistan, 19200 Saidu Sharif (Swat), Pakistan.
A.Kh. Margulan Institute of Archaeology, Almaty 050010, Kazakhstan.
Julio Bendezu-Sarmiento
CNRS-EXT500, Directeur de la Delegation Archaologique Francaise en Afghanistan (DAFA), Embassy of France in Kabul, Afghanistan.
Arman A. Bissembaev
A.Kh. Margulan Institute of Archaeology, Almaty 050010, Kazakhstan.
Aktobe Regional Historical Museum, Aktobe 030006, Kazakhstan.
Archaeology of Asia Department, ISMEO - International Association of Mediterranean and Oriental Studies, Rome RM00186, Italy.
Temirlan T. Chargynov
Kyrgyz National University, 720033 Bishkek, Kyrgyzstan.
Institute of Archaeology and Ethnography, Siberian Branch, Russian Academy of Sciences, Novosibirsk 630090, Russia.
Petr K. Dashkovskiy
Department of Political History, National and State-Confessional Relations, Altai State University, Barnaul 656049, Russia.
Institute of Archaeology and Ethnography, Siberian Branch, Russian Academy of Sciences, Novosibirsk 630090, Russia.
Miroslav Dobeš
Institute of Archaeology, Czech Academy of Sciences, Prague 118 01, Czech Republic.
Katerina Douka
Department of Archaeology, Max Planck Institute for the Science of Human History, Jena 07745, Germany.
Oxford Radiocarbon Accelerator Unit, Research Laboratory for Archaeology and the History of Art, University of Oxford, Oxford OX1 3QY, UK.
Center of Physical Anthropology, Institute of Ethnology and Anthropology, Russian Academy of Sciences, Moscow 119991, Russia.
Meiram N. Duisengali
Aktobe Regional Historical Museum, Aktobe 030006, Kazakhstan.
Dmitry Enshin
Tyumen Scientific Centre SB RAS, Institute of the Problems of Northern Development, Tyumen 625003, Russia.
Institute of History and Archaeology, Ural Branch RAS, Yekaterinburg 620990, Russia.
South Ural State University, Chelyabinsk 454080, Russia.
Department of Archaeology, Kemerovo State University, Kemerovo 650043, Russia.
Institute of Archaeology, University College London, London WC1H 0PY, UK.
School of Cultural Heritage, Northwest University, Shanxi, 710069, China.
Alexander Goryachev
Tyumen Scientific Centre SB RAS, Institute of the Problems of Northern Development, Tyumen 625003, Russia.
Peter the Great Museum of Anthropology and Ethnography (Kunstkamera), Russian Academy of Science, St. Petersburg 199034, Russia.
Department of Archaeology, Ethnography and Museology, Altai State University, Barnaul 656049, Russia.
Department of Anthropology, University of Pittsburgh, Pittsburgh, PA 15260, USA.
Department of Anthropology, University of Pittsburgh, Pittsburgh, PA 15260, USA.
A.Kh. Margulan Institute of Archaeology, Almaty 050010, Kazakhstan.
Samara State University of Social Sciences and Education, Samara 443099, Russia.
Aleksander P. Krygin
West Kazakhstan Regional Center for History and Archaeology, Uralsk 090000, Kazakhstan.
Elena Kupriyanova
Scientific and Educational Center of Study on the Problem of Nature and Man, Chelyabinsk State University, Chelyabinsk 454021, Russia.
Samara State University of Social Sciences and Education, Samara 443099, Russia.
Department of Cultural Heritage, University of Bologna, 48121 Ravenna, Italy.
Institute for Archaeological Research, Uzbekistan Academy of Sciences, Samarkand 140151, Uzbekistan.
Aslan M. Mamedov
Center for Research, Restoration and Protection of Historical and Cultural Heritage of Aktobe Region, Aktobe 030007, Kazakhstan.
Talgat B. Mamirov
A.Kh. Margulan Institute of Archaeology, Almaty 050010, Kazakhstan.
Christopher Meiklejohn
Department of Anthropology, University of Winnipeg, Winnipeg, MB R3B 2E9, Canada.
Department of Archaeology, Simon Fraser University, Burnaby, BC V5A 1S6, Canada.
Roberto Micheli
ISMEO - International Association of Mediterranean and Oriental Studies, Italian Archaeological Mission in Pakistan, 19200 Saidu Sharif (Swat), Pakistan.
MiBAC – Ministero per i Beni e le Attività Culturali - Soprintendenza Archeologia, belle arti e paesaggio del Friuli Venezia Giulia, 34135 Trieste, Italy.
Oleg Mochalov
Samara State University of Social Sciences and Education, Samara 443099, Russia.
Institute for Archaeological Research, Uzbekistan Academy of Sciences, Samarkand 140151, Uzbekistan.
Afrosiab Museum, Samarkand 140151, Uzbekistan.
Ayushi Nayak
Department of Archaeology, Max Planck Institute for the Science of Human History, Jena 07745, Germany.
Department of Biological, Geological and Environmental Sciences, Alma Mater Studiorum – University of Bologna, Bologna 40126, Italy.
Human Origins Program, National Museum of Natural History, Smithsonian Institution, Washington, DC 20013, USA.
Dmitry Razhev
Tyumen Scientific Centre SB RAS, Institute of the Problems of Northern Development, Tyumen 625003, Russia.
Marina Rykun
National Research Tomsk State University, Tomsk 634050, Russia.
Stefania Sarno
Department of Biological, Geological and Environmental Sciences, Alma Mater Studiorum – University of Bologna, Bologna 40126, Italy.
Tatyana M. Savenkova
F. Voino-Yasenetsky Krasnoyarsk State Medical University, Krasnoyarsk 660022, Russia.
Central State Museum Republic of Kazakhstan, Samal-1 Microdistrict, Almaty 050010, Kazakhstan.
Sergey M. Slepchenko
Tyumen Scientific Centre SB RAS, Institute of the Problems of Northern Development, Tyumen 625003, Russia.
Kyrgyz National University, 720033 Bishkek, Kyrgyzstan.
Institute of Archaeology and Ethnography, Siberian Branch, Russian Academy of Sciences, Novosibirsk 630090, Russia.
Svetlana Svyatko
Peter the Great Museum of Anthropology and Ethnography (Kunstkamera), Russian Academy of Science, St. Petersburg 199034, Russia.
CHRONO Centre for Climate, the Environment, and Chronology, Queen's University of Belfast, Belfast BT7 1NN, Northern Ireland, UK.
Kyrgyz-Turkish Manas University, Bishkek, Kyrgyzstan.
Maria Teschler-Nicola
Department of Evolutionary Anthropology, University of Vienna, 1090 Vienna, Austria.
Department of Anthropology, Natural History Museum Vienna, 1010 Vienna, Austria.
Department of Archaeology, Ethnography and Museology, The Laboratory of Interdisciplinary Studies in Archaeology of Western Siberia and Altai, Altai State University, Barnaul 656049, Russia.
Institute of Steppe, Ural Branch RAS, Orenburg 460000, Russia.
Center of Physical Anthropology, Institute of Ethnology and Anthropology, Russian Academy of Sciences, Moscow 119991, Russia.
Center for Egyptological Studies RAS, Moscow 119991, Russia.
Department of Anthropology, National Museum, Prague 115 79, Czech Republic.
Dmitriy Voyakin
A.Kh. Margulan Institute of Archaeology, Almaty 050010, Kazakhstan.
Archaeological Expertise LLP, Almaty 050060, Kazakhstan.
A.Kh. Margulan Institute of Archaeology, Almaty 050010, Kazakhstan.
Muhammad Zahir
Department of Archaeology, Max Planck Institute for the Science of Human History, Jena 07745, Germany.
Department of Archaeology, Hazara University, Mansehra 21300, Pakistan.
Valery S. Zubkov
N.F. Katanov Khakassia State University, Abakan 655017, Russia.
Alisa Zubova
Peter the Great Museum of Anthropology and Ethnography (Kunstkamera), Russian Academy of Science, St. Petersburg 199034, Russia.
Vasant S. Shinde
Department of Archaeology, Deccan College Post-Graduate and Research Institute, Pune 411006, India.
Institute of Evolutionary Biology, CSIC-Universitat Pompeu Fabra, Barcelona 08003, Spain.
Max Planck Institute for Evolutionary Anthropology, Leipzig 04103, Germany.
Anthropology Department, Hartwick College, Oneonta, NY 13820, USA.
Nicole Boivin
Department of Archaeology, Max Planck Institute for the Science of Human History, Jena 07745, Germany.
CSIR-Centre for Cellular and Molecular Biology, Hyderabad 500 007, India.
Douglas J. Kennett
Institutes of Energy and the Environment, Pennsylvania State University, University Park, PA 16802, USA.
Department of Anthropology, Pennsylvania State University, University Park, PA 16802, USA.
Department of Anthropology, University of California, Santa Barbara, CA 93106, USA.
Department of Anthropology, Washington University in St. Louis, St. Louis, MO 63112, USA.
Spatial Analysis, Interpretation, and Exploration Laboratory, Washington University in St. Louis, St. Louis, MO 63112, USA.
Earth Institute, University College Dublin, Dublin 4, Ireland.
Department of Evolutionary Anthropology, University of Vienna, 1090 Vienna, Austria.
Department of Genetics, Harvard Medical School, Boston, MA 02115, USA.
Broad Institute of Harvard and MIT, Cambridge, MA 02142, USA.
Howard Hughes Medical Institute, Harvard Medical School, Boston, MA 02115, USA.
Max Planck-Harvard Research Center for the Archaeoscience of the Ancient Mediterranean, Cambridge, MA 02138, USA.

Notes

*
These authors contributed equally to this work.
†Corresponding author. Email: [email protected] (V.M.N.); [email protected] (N.P.); [email protected] (M.Fr.); [email protected] (R.Pi.); [email protected] (D.Re.)
These authors contributed equally to this work.

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Science
Volume 365|Issue 6457
6 September 2019
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Received:30 March 2018
Accepted:30 July 2019
Published in print:6 September 2019
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