By sequencing 727 ancient individuals from the Southern Arc (Anatolia and its neighbors in Southeastern Europe and West Asia) over 10,000 years, we contextualize its Chalcolithic period and Bronze Age (about 5000 to 1000 BCE), when extensive gene flow entangled it with the Eurasian steppe. Two streams of migration transmitted Caucasus and Anatolian/Levantine ancestry northward, and the Yamnaya pastoralists, formed on the steppe, then spread southward into the Balkans and across the Caucasus into Armenia, where they left numerous patrilineal descendants. Anatolia was transformed by intra-West Asian gene flow, with negligible impact of the later Yamnaya migrations. This contrasts with all other regions where Indo-European languages were spoken, suggesting that the homeland of the Indo-Anatolian language family was in West Asia, with only secondary dispersals of non-Anatolian Indo-Europeans from the steppe.

Amasya Archaeology Museum Mustafa Kemal Paşa Caddesi 05000 Amasya Turkey

Anthropological Center of the Croatian Academy of Sciences and Arts 10000 Zagreb Croatia

Archaeological Institute of America Boston MA 02108 USA

Archaeological Museum in Zadar 23000 Zadar Croatia

Archaeological Museum in Zagreb 10000 Zagreb Croatia

Archaeology Museum of North Macedonia 1000 Skopje North Macedonia

ArchaeoSciences Division Research Institute of the University of Bucharest University of Bucharest 050663 Bucharest Romania

Austrian Archaeological Institute at the Austrian Academy of Sciences 1190 Vienna Austria

BIOMICs Research Group University of the Basque Country UPV EHU 01006 Vitoria Gasteiz Spain

Bodrum Museum of Underwater Archeology Çarşı Neighbourhood 48400 Bodrum Muğla Turkey

Broad Institute of Harvard and MIT Cambridge MA 02142 USA

Center for Conservation and Archaeology of Montenegro 81250 Cetinje Montenegro

Center of Archaeology Institute of Cultural Heritage Academy of Science of Moldova 2001 Chișinău Moldova

Centre for Applied Bioanthropology Institute for Anthropological Research 10000 Zagreb Croatia

Cotsen Institute of Archaeology University of California Los Angeles Los Angeles CA 90095 USA

Culture and Patrimony Department University of Avignon F 84029 Avignon France

Departament d'Història i Història de l'Art Universitat Rovira i Virgili 43002 Tarragona Spain

Department of Animal and Human Physiology Faculty of Biology School of Sciences National and Kapodistrian University of Athens 10679 Athens Greece

Department of Anthropology and Middle Eastern Cultures Mississippi State University Mississippi State MS 39762 USA

Department of Anthropology Binghamton University Binghamton NY 13902 USA

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Department of Anthropology Faculty of Language and History Geography Ankara University 06100 Sıhhiye Ankara Turkey

Department of Anthropology Faculty of Letters Istanbul University 34134 Istanbul Turkey

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Department of Archaeology and Heritage Studies University of Tirana 1010 Tirana Albania

Department of Archaeology Durham University Durham DH1 3LE UK

Department of Archaeology Faculty of Humanities and Social Sciences University of Zagreb 10000 Zagreb Croatia

Department of Archaeology Faculty of Humanities Van Yüzüncü Yıl University 65090 Tuşba Van Turkey

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Department of Armenian History Armenian State Pedagogical University After Khachatur Abovyan 0010 Yerevan Armenia

Department of Biological Anthropology Institute of Biology Eötvös Loránd University 1053 Budapest Hungary

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Department of Cultures University of Helsinki 00100 Helsinki Finland

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Hellenic Ministry of Culture and Sports Ephorate of Antiquities of Piraeus and the Islands 10682 Piraeus Greece

Historical Musem in Kotor 85330 Kotor Montenegro

History of the Ancient World and Middle Ages Department Moscow Region State University Moscow Region 141014 Mytishi Russia

Howard Hughes Medical Institute Harvard Medical School Boston MA 02115 USA

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Human Paleoecology and Isotope Geochemistry Lab Department of Anthropology The Pennsylvania State University University Park PA 16802 USA

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Institut Català de Paleoecologia Humana i Evolució Social 43007 Tarragona Spain

Institute for Quaternary Paleontology and Geology Croatian Academy of Sciences and Arts 10000 Zagreb Croatia

Institute of Archaeogenomics Research Centre for the Humanities Eötvös Loránd Research Network 1097 Budapest Hungary

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Ministry of Culture and Tourism İsmet İnönü Bulvarı 06100 Emek Ankara Turkey

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Muğla Archaeological Museum and Yatağan Thermal Power Generation Company Rescue Excavations 48000 Muğla Turkey

Muğla İl Kültür ve Turizm Müdürlüğü 48000 Muğla Turkey

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Museum of Anthropological Archaeology University of Michigan Ann Arbor MI 48109 USA

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National Archaeological Agency 2012 Chișinău Moldova

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Penn Museum University of Pennsylvania Philadelphia PA 19104 USA

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Prehistory Department Albanian Institute of Archaeology Academy of Albanian Studies 1000 Tirana Albania

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Research Centre for Anthropology and Health Department of Life Sciences University of Coimbra 3000 456 Coimbra Portugal

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The Genetic Origin of the Indo-Europeans

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