The Yamnaya archaeological complex appeared around 3300BCE across the steppes north of the Black and Caspian Seas, and by 3000BCE reached its maximal extent from Hungary in the west to Kazakhstan in the east. To localize the ancestral and geographical origins of the Yamnaya among the diverse Eneolithic people that preceded them, we studied ancient DNA data from 428 individuals of which 299 are reported for the first time, demonstrating three previously unknown Eneolithic genetic clines. First, a "Caucasus-Lower Volga" (CLV) Cline suffused with Caucasus hunter-gatherer (CHG) ancestry extended between a Caucasus Neolithic southern end in Neolithic Armenia, and a steppe northern end in Berezhnovka in the Lower Volga. Bidirectional gene flow across the CLV cline created admixed intermediate populations in both the north Caucasus, such as the Maikop people, and on the steppe, such as those at the site of Remontnoye north of the Manych depression. CLV people also helped form two major riverine clines by admixing with distinct groups of European hunter-gatherers. A "Volga Cline" was formed as Lower Volga people mixed with upriver populations that had more Eastern hunter-gatherer (EHG) ancestry, creating genetically hyper-variable populations as at Khvalynsk in the Middle Volga. A "Dnipro Cline" was formed as CLV people bearing both Caucasus Neolithic and Lower Volga ancestry moved west and acquired Ukraine Neolithic hunter-gatherer (UNHG) ancestry to establish the population of the Serednii Stih culture from which the direct ancestors of the Yamnaya themselves were formed around 4000BCE. This population grew rapidly after 3750-3350BCE, precipitating the expansion of people of the Yamnaya culture who totally displaced previous groups on the Volga and further east, while admixing with more sedentary groups in the west. CLV cline people with Lower Volga ancestry contributed four fifths of the ancestry of the Yamnaya, but also, entering Anatolia from the east, contributed at least a tenth of the ancestry of Bronze Age Central Anatolians, where the Hittite language, related to the Indo-European languages spread by the Yamnaya, was spoken. We thus propose that the final unity of the speakers of the "Proto-Indo-Anatolian" ancestral language of both Anatolian and Indo-European languages can be traced to CLV cline people sometime between 4400-4000 BCE.

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 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 of the Southern Federal University Rostov on Don Russia

Department of Archaeology Ethnography and Museology Altai State University Barnaul Russia

Department of Archaeology University of Szeged Szeged 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 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 4026 Debrecen Hungary

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

Fr 1 Rainer Institute of Anthropology University of Bucharest Bucharest Romania

Gavrilă Simion Eco Museum Research Institute Tulcea Romania

Hartwick College Dept of Anthropology USA

Howard Hughes Medical Institute Harvard Medical School Boston MA USA

Human Evolution and Archaeological Sciences University of Vienna Vienna Austria

Hungarian Natural History Museum Department of Anthropology Budapest Hungary

Ikerbasque Basque Foundation of Science Bilbao Spain

Independent Researcher 106 Federal Street 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 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 Republic of Kalmykia 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

Russian Academy of Sciences Institute of Ethnology and Anthropology Moscow Russia

Samara Regional Public Organization Historical ecological and cultural Association Povolzje

Samara State University of Social Sciences and Education Samara Russia

School of Archaeology University College 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 Republic of Khakassia Abakan Russia

State History Museum Department of Archaeology Moscow Russia

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

University of Oxford Faculty of History Oxford United Kingdom

Wellcome Centre for Human Genetics University of Oxford Oxford UK

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