The Yamnaya archaeological complex appeared around 3300 BC across the steppes north of the Black and Caspian Seas, and by 3000 BC it reached its maximal extent, ranging from Hungary in the west to Kazakhstan in the east. To localize Yamnaya origins among the preceding Eneolithic people, we assembled ancient DNA from 435 individuals, demonstrating three genetic clines. A Caucasus-lower Volga (CLV) cline suffused with Caucasus hunter-gatherer1 ancestry extended between a Caucasus Neolithic southern end and a northern end at Berezhnovka along the lower Volga river. Bidirectional gene flow created intermediate populations, such as the north Caucasus Maikop people, and those at Remontnoye on the steppe. The Volga cline was formed as CLV people mixed with upriver populations of Eastern hunter-gatherer2 ancestry, creating hypervariable groups, including one at Khvalynsk. The Dnipro cline was formed when CLV people moved west, mixing with people with Ukraine Neolithic hunter-gatherer ancestry3 along the Dnipro and Don rivers to establish Serednii Stih groups, from whom Yamnaya ancestors formed around 4000 BC and grew rapidly after 3750-3350 BC. The CLV people contributed around four-fifths of the ancestry of the Yamnaya and, entering Anatolia, probably from the east, at least one-tenth of the ancestry of Bronze Age central Anatolians, who spoke Hittite4,5. We therefore propose that the final unity of the speakers of 'proto-Indo-Anatolian', the language ancestral to both Anatolian and Indo-European people, occurred in CLV people some time between 4400 BC and 4000 BC.

5 F Voino Yasenetsky Krasnoyarsk State Medical University Krasnoyarsk Russia

Azov History Archaeology and Palaeontology Museum Reserve Azov Russia

BIOMICs Research Group Department of Zoology and Animal Cell Biology University of the Basque Country UPV EHU Vitoria Gasteiz Spain

Broad Institute of Harvard and MIT Cambridge MA USA

Center for Stone Age Archeology Institute of History and Archaeology Ural Branch of the Russian Academy of Sciences Ekaterinburg Russia

Center of Human Ecology Institute of Ethnology and Anthropology Russian Academy of Science Moscow Russia

Centre for Applied Bioanthropology Institute for Anthropological Research Zagreb Croatia

Centre for Egyptological Studies of the Russian Academy of Sciences Russian Academy of Sciences Moscow Russia

Damjanich János Museum Szolnok Hungary

Department of Anthropology Hungarian Natural History Museum Hungarian National Museum Public Collection Centre Budapest Hungary

Department of Archaeogenetics Max Planck Institute for Evolutionary Anthropology Leipzig Germany

Department of Archaeology and Heritage Faculty of Humanities University of Primorska Koper Slovenia

Department of Archaeology and History of the Ancient World Southern Federal University Rostov on Don Russia

Department of Archaeology Ethnography and Museology Altai State University Barnaul Russia

Department of Archaeology State History Museum Moscow Russia

Department of Archaeology University of Szeged Szeged Hungary

Department of Biological Anthropology Institute of Biology Eötvös Loránd University Budapest Hungary

Department of Biological Anthropology Institute of Biology University of Szeged Szeged Hungary

Department of Biology and Ecology Faculty of Science University of Ostrava Ostrava Czechia

Department of Evolutionary Anthropology University of Vienna Vienna Austria

Department of General History Historical and Literary Institute of the State University of Education Ministry of Education Moscow Moscow Russia

Department of Genetics Harvard Medical School Boston MA USA

Department of Geography Faculty of Humanities University Valahia of Târgoviște Târgovişte Romania

Department of History of the Institute of Humanities Ural Federal University Ekaterinburg Russia

Department of Human Evolutionary Biology Harvard University Cambridge MA USA

Department of Mountain and Highland Archaeology Institute of Archaeology and Ethnology Polish Academy of Science Kraków Poland

Department of Regional Studies of Russia National and State Confessional Relations Altai State University Barnaul Russia

Department of Statistics University of Oxford Oxford UK

Déri Museum Debrecen Hungary

Faculty of History University of Oxford Oxford UK

Federal Research Centre Fundamentals of Biotechnology of the Russian Academy of Sciences Moscow Russia

Fr 1 Rainer Institute of Anthropology University of Bucharest Bucharest Romania

Gavrilă Simion Eco Museum Research Institute Tulcea Romania

Hartwick College Department of Anthropology Oneonta NY USA

Historical Ecological and Cultural Association Povolzje Samara Regional Public Organization Samara Russia

Howard Hughes Medical Institute Harvard Medical School Boston MA USA

Human Evolution and Archaeological Sciences University of Vienna Vienna Austria

Ikerbasque Basque Foundation of Science Bilbao Spain

Independent researcher Philadelphia PA USA

Institute for the History of Material Culture Russian Academy of Sciences St Petersburg Russia

Institute of Archaeogenomics HUN REN Research Centre for the Humanities Budapest Hungary

Institute of Archaeology HUN REN Research Centre for the Humanities Budapest Hungary

Institute of Archaeology named after A Kh Margulan Almaty Kazakhstan

Institute of Archeology named after A Kh Khalikov Tatarstan Academy of Sciences Kazan Russia

Institute of Ethnology and Anthropology Russian Academy of Sciences Moscow Russia

Institute of History Archaeology and Ethnography Dagestan Branch of the Russian Academy of Science Makhachkala Dagestan Russia

Institute of Parasitology Biology Centre of the Czech Academy of Sciences České Budějovice Czechia

Institute of Plant and Animal Ecology Urals Branch of the Russian Academy of Sciences Yekaterinburg Russia

Kalmyk Scientific Centre of the Russian Academy of Sciences Elista Russia

Laboratory of Ancient and Medieval Archaeology of Eurasia Altai State University Barnaul Russia

Museo delle Civiltà Italian Ministry of Culture Rome Italy

Museum of Vojvodina Novi Sad Serbia

National Agency for Archaeology Chișinău Republic of Moldova

National Research Tomsk State University Tomsk Russia

Olga Necrasov Centre for Anthropological Research Romanian Academy Iași Branch Iași Romania

Orheiul Vechi Cultural Natural Reserve Institute of Bioarchaeological and Ethnocultural Research Chișinău Republic of Moldova

Peter the Great Museum of Anthropology and Ethnography Department of Physical Anthropology St Petersburg Russia

Prahova County Museum of History and Archaeology Ploiești Romania

Research Institute and Museum of Anthropology Moscow Russia

Research Institute GAUK RO Don Heritage Rostov on Don Russia

Samara State University of Social Sciences and Education Samara Russia

School of Archaeology University College Dublin Dublin Ireland

Slovak National Museum Archaeological Museum Bratislava Slovak Republic

Slovak National Museum Natural History Museum Bratislava Slovak Republic

State Autonomous Cultural Institution of the Republic of Khakassia Khakassian National Museum of Local Lore named after L R Kyzlasova Abakan Russia

Tyumen Scientific Center of the Siberian Branch of Russian Academy of Sciences Institute of Problems of Northern Development Tyumen Russia

Wellcome Centre for Human Genetics University of Oxford Oxford UK

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