. 2022 Aug 26;377(6609):eabm4247.

doi: 10.1126/science.abm4247. Epub 2022 Aug 26.

The genetic history of the Southern Arc: A bridge between West Asia and Europe

Iosif Lazaridis^#^{1

2}, Songül Alpaslan-Roodenberg^#^{2

3}, Ayşe Acar⁴, Ayşen Açıkkol⁵, Anagnostis Agelarakis⁶, Levon Aghikyan⁷, Uğur Akyüz⁸, Desislava Andreeva⁹, Gojko Andrijašević¹⁰, Dragana Antonović¹¹, Ian Armit¹², Alper Atmaca¹³, Pavel Avetisyan⁷, Ahmet İhsan Aytek¹⁴, Krum Bacvarov¹⁵, Ruben Badalyan⁷, Stefan Bakardzhiev¹⁶, Jacqueline Balen¹⁷, Lorenc Bejko¹⁸, Rebecca Bernardos², Andreas Bertsatos¹⁹, Hanifi Biber²⁰, Ahmet Bilir²¹, Mario Bodružić²², Michelle Bonogofsky²³, Clive Bonsall²⁴, Dušan Borić²⁵, Nikola Borovinić²⁶, Guillermo Bravo Morante³, Katharina Buttinger³, Kim Callan^{2

27}, Francesca Candilio²⁸, Mario Carić²⁹, Olivia Cheronet³, Stefan Chohadzhiev³⁰, Maria-Eleni Chovalopoulou¹⁹, Stella Chryssoulaki³¹, Ion Ciobanu^{32

33}, Natalija Čondić³⁴, Mihai Constantinescu³⁵, Emanuela Cristiani³⁶, Brendan J Culleton³⁷, Elizabeth Curtis^{2

27}, Jack Davis³⁸, Tatiana I Demcenco³⁹, Valentin Dergachev⁴⁰, Zafer Derin⁴¹, Sylvia Deskaj⁴², Seda Devejyan⁷, Vojislav Djordjević⁴³, Kellie Sara Duffett Carlson³, Laurie R Eccles⁴⁴, Nedko Elenski⁴⁵, Atilla Engin⁴⁶, Nihat Erdoğan⁴⁷, Sabiha Erir-Pazarcı⁴⁸, Daniel M Fernandes^{3

49}, Matthew Ferry^{2

27}, Suzanne Freilich³, Alin Frînculeasa⁵⁰, Michael L Galaty⁴², Beatriz Gamarra^{51

52

53}, Boris Gasparyan⁷, Bisserka Gaydarska⁵⁴, Elif Genç⁵⁵, Timur Gültekin⁵⁶, Serkan Gündüz⁵⁷, Tamás Hajdu⁵⁸, Volker Heyd⁵⁹, Suren Hobosyan⁷, Nelli Hovhannisyan⁶⁰, Iliya Iliev¹⁶, Lora Iliev^{2

27}, Stanislav Iliev⁶¹, İlkay İvgin⁶², Ivor Janković²⁹, Lence Jovanova⁶³, Panagiotis Karkanas⁶⁴, Berna Kavaz-Kındığılı⁶⁵, Esra Hilal Kaya⁶⁶, Denise Keating³, Douglas J Kennett^{37

67}, Seda Deniz Kesici⁶⁸, Anahit Khudaverdyan⁷, Krisztián Kiss^{58

69}, Sinan Kılıç²⁰, Paul Klostermann⁷⁰, Sinem Kostak Boca Negra Valdes⁶⁸, Saša Kovačević⁷¹, Marta Krenz-Niedbała⁷², Maja Krznarić Škrivanko⁷³, Rovena Kurti⁷⁴, Pasko Kuzman⁷⁵, Ann Marie Lawson^{2

27}, Catalin Lazar⁷⁶, Krassimir Leshtakov⁷⁷, Thomas E Levy⁷⁸, Ioannis Liritzis^{79

80}, Kirsi O Lorentz⁸¹, Sylwia Łukasik⁷², Matthew Mah^{2

27

82}, Swapan Mallick^{2

27}, Kirsten Mandl³, Kristine Martirosyan-Olshansky⁸³, Roger Matthews⁸⁴, Wendy Matthews⁸⁴, Kathleen McSweeney²⁴, Varduhi Melikyan⁷, Adam Micco², Megan Michel^{1

2

27}, Lidija Milašinović⁸⁵, Alissa Mittnik^{1

2

86}, Janet M Monge⁸⁷, Georgi Nekhrizov¹⁵, Rebecca Nicholls⁸⁸, Alexey G Nikitin⁸⁹, Vassil Nikolov¹⁵, Mario Novak²⁹, Iñigo Olalde^{2

90}, Jonas Oppenheimer^{2

27}, Anna Osterholtz⁹¹, Celal Özdemir¹³, Kadir Toykan Özdoğan³, Nurettin Öztürk⁶⁵, Nikos Papadimitriou⁹², Niki Papakonstantinou⁹³, Anastasia Papathanasiou⁹⁴, Lujana Paraman⁹⁵, Evgeny G Paskary⁹⁶, Nick Patterson^{1

82}, Ilian Petrakiev⁴⁵, Levon Petrosyan⁷, Vanya Petrova⁷⁷, Anna Philippa-Touchais⁹⁷, Ashot Piliposyan⁹⁸, Nada Pocuca Kuzman⁷⁵, Hrvoje Potrebica⁹⁹, Bianca Preda-Bălănică⁵⁹, Zrinka Premužić¹⁰⁰, T Douglas Price¹⁰¹, Lijun Qiu^{2

27}, Siniša Radović¹⁰², Kamal Raeuf Aziz¹⁰³, Petra Rajić Šikanjić²⁹, Kamal Rasheed Raheem¹⁰³, Sergei Razumov¹⁰⁴, Amy Richardson⁸⁴, Jacob Roodenberg¹⁰⁵, Rudenc Ruka⁷⁴, Victoria Russeva¹⁰⁶, Mustafa Şahin⁵⁷, Ayşegül Şarbak¹⁰⁷, Emre Savaş⁶⁸, Constanze Schattke³, Lynne Schepartz¹⁰⁸, Tayfun Selçuk⁶⁸, Ayla Sevim-Erol¹⁰⁹, Michel Shamoon-Pour¹¹⁰, Henry M Shephard¹¹¹, Athanasios Sideris¹¹², Angela Simalcsik^{32

113}, Hakob Simonyan¹¹⁴, Vitalij Sinika¹⁰⁴, Kendra Sirak², Ghenadie Sirbu¹¹⁵, Mario Šlaus¹¹⁶, Andrei Soficaru³⁵, Bilal Söğüt¹¹⁷, Arkadiusz Sołtysiak¹¹⁸, Çilem Sönmez-Sözer¹⁰⁹, Maria Stathi¹¹⁹, Martin Steskal¹²⁰, Kristin Stewardson^{2

27}, Sharon Stocker³⁸, Fadime Suata-Alpaslan¹²¹, Alexander Suvorov⁵⁹, Anna Szécsényi-Nagy¹²², Tamás Szeniczey⁵⁸, Nikolai Telnov¹⁰⁴, Strahil Temov¹²³, Nadezhda Todorova⁷⁷, Ulsi Tota^{74

124}, Gilles Touchais¹²⁵, Sevi Triantaphyllou⁹³, Atila Türker¹²⁶, Marina Ugarković⁷¹, Todor Valchev¹⁶, Fanica Veljanovska¹²³, Zlatko Videvski¹²³, Cristian Virag¹²⁷, Anna Wagner³, Sam Walsh¹²⁸, Piotr Włodarczak¹²⁹, J Noah Workman², Aram Yardumian^{130

131}, Evgenii Yarovoy¹³², Alper Yener Yavuz¹³³, Hakan Yılmaz²⁰, Fatma Zalzala^{2

27}, Anna Zettl³, Zhao Zhang², Rafet Çavuşoğlu²⁰, Nadin Rohland², Ron Pinhasi^{3

134}, David Reich^{1

2

27

82}, Ruben Davtyan⁷

Affiliations

¹ Department of Human Evolutionary Biology, Harvard University, Cambridge, MA 02138, USA.
² Department of Genetics, Harvard Medical School, Boston, MA 02115, USA.
³ Department of Evolutionary Anthropology, University of Vienna, 1030 Vienna, Austria.
⁴ Department of Anthropology, Faculty of Letters, Mardin Artuklu University, 47510 Artuklu, Mardin, Turkey.
⁵ Department of Anthropology, Faculty of Letters, Sivas Cumhuriyet University, 58140 Sivas, Turkey.
⁶ Department of History, Adelphi University, Garden City, NY 11530, USA.
⁷ Institute of Archaeology and Ethnography, NAS RA, 0025 Yerevan, Armenia.
⁸ Samsun Museum of Archeology and Ethnography, Kale Mahallesi, Merkez, İlkadım, 55030 Samsun, Turkey.
⁹ Iskra Museum of History, 6100 Kazanlak, Bulgaria.
¹⁰ Historical Musem in Kotor, 85330 Kotor, Montenegro.
¹¹ Institute of Archaeology, 11000 Belgrade, Serbia.
¹² Department of Archaeology, University of York, York YO1 7EP, UK.
¹³ Amasya Archaeology Museum, Mustafa Kemal Paşa Caddesi, 05000 Amasya, Turkey.
¹⁴ Department of Anthropology, Faculty of Arts and Science, Burdur Mehmet Akif University, 15100 Burdur, Turkey.
¹⁵ National Institute of Archaeology and Museum, Bulgarian Academy of Sciences, 1000 Sofia, Bulgaria.
¹⁶ Yambol Regional Historical Museum, 8600 Yambol, Bulgaria.
¹⁷ Archaeological Museum in Zagreb, 10000 Zagreb, Croatia.
¹⁸ Department of Archaeology and Heritage Studies, University of Tirana, 1010 Tirana, Albania.
¹⁹ Department of Animal and Human Physiology, Faculty of Biology, School of Sciences, National and Kapodistrian University of Athens, 10679 Athens, Greece.
²⁰ Department of Archaeology, Faculty of Humanities, Van Yüzüncü Yıl University, 65090 Tuşba, Van, Turkey.
²¹ Department of Archaeology, Faculty of Science and Letters, Düzce University, 81620 Düzce, Turkey.
²² Stratum Ltd., 21218 Seget Donji, Croatia.
²³ Independent Researcher, Berkeley, CA 94720, USA.
²⁴ School of History, Classics and Archaeology, University of Edinburgh, Edinburgh EH8 9AG, UK.
²⁵ The Italian Academy for Advanced Studies in America, Columbia University, New York, NY 10027, USA.
²⁶ Center for Conservation and Archaeology of Montenegro, 81250 Cetinje, Montenegro.
²⁷ Howard Hughes Medical Institute, Harvard Medical School, Boston, MA 02115, USA.
²⁸ Servizio di Bioarcheologia, Museo delle Civiltà, 00144 Rome, Italy.
²⁹ Centre for Applied Bioanthropology, Institute for Anthropological Research, 10000 Zagreb, Croatia.
³⁰ Department of Archaeology, University of Veliko Tarnovo "St. Cyril and St. Methodius," 5003 Veliko Tarnovo, Bulgaria.
³¹ Hellenic Ministry of Culture and Sports, Ephorate of Antiquities of Piraeus and the Islands, 10682 Piraeus, Greece.
³² "Orheiul Vechi" Cultural-Natural Reserve, Institute of Bioarchaeological and Ethnocultural Research, 3552 Butuceni, Moldova.
³³ National Archaeological Agency, 2012 Chișinău, Moldova.
³⁴ Archaeological Museum in Zadar, 23000 Zadar, Croatia.
³⁵ "Francisc I. Rainer" Institute of Anthropology, 050711 Bucharest, Romania.
³⁶ Department of Oral and Maxillo-Facial Sciences, Sapienza University of Rome, 00161 Rome, Italy.
³⁷ Institutes of Energy and the Environment, The Pennsylvania State University, University Park, PA 16802, USA.
³⁸ Department of Classics, University of Cincinnati, Cincinnati, OH 45221, USA.
³⁹ Independent Researcher, Aberystwyth SY23 4UH, UK.
⁴⁰ Center of Archaeology, Institute of Cultural Heritage, Academy of Science of Moldova, 2001 Chișinău, Moldova.
⁴¹ Department of Archaeology, Faculty of Letters, Ege University, 35100 Bornova-Izmir, Turkey.
⁴² Museum of Anthropological Archaeology, University of Michigan, Ann Arbor, MI 48109, USA.
⁴³ Narodni muzej Pančcevo, 26101 Pančevo, Serbia.
⁴⁴ Human Paleoecology and Isotope Geochemistry Lab, Department of Anthropology, The Pennsylvania State University, University Park, PA 16802, USA.
⁴⁵ Regional Museum of History - Veliko Tarnovo, 5000 Veliko Tarnovo, Bulgaria.
⁴⁶ Department of Archaeology, Faculty of Science and Letters, Gaziantep University, 27310 Gaziantep, Turkey.
⁴⁷ Mardin Archaeological Museum, Şar, Cumhuriyet Meydanı üstü, 47100 Artuklu, Mardin, Turkey.
⁴⁸ Muğla İl Kültür ve Turizm Müdürlüğü, 48000 Muğla, Turkey.
⁴⁹ Research Centre for Anthropology and Health (CIAS), Department of Life Sciences, University of Coimbra, 3000-456 Coimbra, Portugal.
⁵⁰ Prahova County Museum of History and Archaeology, 100042 Ploiești, Romania.
⁵¹ Institut Català de Paleoecologia Humana i Evolució Social, 43007 Tarragona, Spain.
⁵² Departament d'Història i Història de l'Art, Universitat Rovira i Virgili, 43002 Tarragona, Spain.
⁵³ School of Archaeology and Earth Institute, University College Dublin, Belfield, Dublin 4, Ireland.
⁵⁴ Department of Archaeology, Durham University, Durham DH1 3LE, UK.
⁵⁵ Department of Archaeology, Faculty of Science and Letters, Çukurova University, 01330 Balçalı-Sarıçam-Adana, Turkey.
⁵⁶ Department of Anthropology, Faculty of Humanities, Ankara University, 06100 Sıhhiye, Ankara, Turkey.
⁵⁷ Department of Archaeology, Faculty of Science and Letters, Bursa Uludağ University, 16059 Görükle, Bursa, Turkey.
⁵⁸ Department of Biological Anthropology, Institute of Biology, Eötvös Loránd University, 1053 Budapest, Hungary.
⁵⁹ Department of Cultures, University of Helsinki, 00100 Helsinki, Finland.
⁶⁰ Department of Ecology and Nature Protection, Yerevan State University, 0025 Yerevan, Armenia.
⁶¹ Regional Museum of History, 6300 Haskovo, Bulgaria.
⁶² Ministry of Culture and Tourism, İsmet İnönü Bulvarı, 06100 Emek, Ankara, Turkey.
⁶³ Museum of the City of Skopje, 1000 Skopje, North Macedonia.
⁶⁴ Malcolm H. Wiener Laboratory, American School of Classical Studies at Athens, 10676 Athens, Greece.
⁶⁵ Department of Archaeology, Faculty of Letters, Atatürk University, 25100 Erzurum, Turkey.
⁶⁶ Muğla Archaeological Museum and Yatağan Thermal Power Generation Company, Rescue Excavations, 48000 Muğla, Turkey.
⁶⁷ Department of Anthropology, University of California, Santa Barbara, Santa Barbara, CA 93106, USA.
⁶⁸ Bodrum Museum of Underwater Archeology, Çarşı Neighbourhood, 48400 Bodrum, Muğla, Turkey.
⁶⁹ Department of Anthropology, Hungarian Natural History Museum, 1117 Budapest, Hungary.
⁷⁰ Department of Anthropology, Natural History Museum Vienna, 1010 Vienna, Austria.
⁷¹ Institute of Archaeology, 10000 Zagreb, Croatia.
⁷² Faculty of Biology, Adam Mickiewicz University in Poznań, 61-614 Poznań, Poland.
⁷³ Municipal Museum Vinkovci, 32100 Vinkovci, Croatia.
⁷⁴ Prehistory Department, Albanian Institute of Archaeology, Academy of Albanian Studies, 1000 Tirana, Albania.
⁷⁵ National Museum in Ohrid, 6000 Ohrid, North Macedonia.
⁷⁶ ArchaeoSciences Division, Research Institute of the University of Bucharest, University of Bucharest, 050663 Bucharest, Romania.
⁷⁷ Department of Archaeology, St. Kliment Ohridski University of Sofia, 1504 Sofia, Bulgaria.
⁷⁸ Department of Anthropology, University of California, San Diego, La Jolla, CA 92093, USA.
⁷⁹ Key Research Institute of Yellow River Civilization and Sustainable Development and the Collaborative Innovation Center on Yellow River Civilization of Henan Province, Laboratory of Yellow River Cultural Heritage, Henan University, 475001 Kaifeng, China.
⁸⁰ European Academy of Sciences and Arts, 5020 Salzburg, Austria.
⁸¹ Science and Technology in Archaeology and Culture Research Center, The Cyprus Institute, 2121 Aglantzia, Nicosia, Cyprus.
⁸² Broad Institute of Harvard and MIT, Cambridge, MA 02142, USA.
⁸³ Cotsen Institute of Archaeology, University of California, Los Angeles, Los Angeles, CA 90095, USA.
⁸⁴ Department of Archaeology, University of Reading, Reading RG6 6AB, UK.
⁸⁵ National Museum of Kikinda, 23300 Kikinda, Serbia.
⁸⁶ Department of Archaeogenetics, Max Planck Institute for Evolutionary Anthropology, 04103 Leipzig, Germany.
⁸⁷ University of Pennsylvania Museum of Archaeology and Anthropology, Philadelphia, PA 19104, USA.
⁸⁸ School of Archaeological and Forensic Sciences, Faculty of Life Sciences, University of Bradford, Bradford BD7 1DP, UK.
⁸⁹ Department of Biology, Grand Valley State University, Allendale, MI 49401, USA.
⁹⁰ BIOMICs Research Group, University of the Basque Country UPV/EHU, 01006 Vitoria-Gasteiz, Spain.
⁹¹ Department of Anthropology and Middle Eastern Cultures, Mississippi State University, Mississippi State, MS 39762, USA.
⁹² Paul and Alexandra Canellopoulos Museum, 105-55 Athens, Greece.
⁹³ Faculty of Philosophy, School of History and Archaeology, Aristotle University of Thessaloniki, 54124 Thessaloniki, Greece.
⁹⁴ Ephorate of Paleoantropology and Speleology, Greek Ministry of Culture, 11636 Athens, Greece.
⁹⁵ Trogir Town Museum, 21220 Trogir, Croatia.
⁹⁶ Moldavian Historic-Geographical Society, 2044 Chișinău, Moldova.
⁹⁷ French School of Archaeology at Athens, 10680 Athens, Greece.
⁹⁸ Department of Armenian History, Armenian State Pedagogical University After Khachatur Abovyan, 0010 Yerevan, Armenia.
⁹⁹ Department of Archaeology, Faculty of Humanities and Social Sciences, University of Zagreb, 10000 Zagreb, Croatia.
¹⁰⁰ Independent Researcher, 10000 Zagreb, Croatia.
¹⁰¹ Laboratory for Archaeological Chemistry, University of Wisconsin-Madison, Madison, WI 53706, USA.
¹⁰² Institute for Quaternary Paleontology and Geology, Croatian Academy of Sciences and Arts, 10000 Zagreb, Croatia.
¹⁰³ Sulaymaniyah Directorate of Antiquities and Heritage, 46010 Sulaymaniyah, Iraq.
¹⁰⁴ Pridnestrovian University named after Taras Shevchenko, 3300 Tiraspol, Moldova.
¹⁰⁵ The Netherlands Institute for the Near East, 2311 Leiden, Netherlands.
¹⁰⁶ Institute of Experimental Morphology, Pathology and Archeology with Museum, Bulgarian Academy of Science, 1113 Sofia, Bulgaria.
¹⁰⁷ Department of Anthropology, Faculty of Science and Letters, Hitit University, 19040 Çorum, Turkey.
¹⁰⁸ School of Anatomical Sciences, The University of the Witwatersrand, 2193 Johannesburg, South Africa.
¹⁰⁹ Department of Anthropology, Faculty of Language and History - Geography, Ankara University, 06100 Sıhhiye, Ankara, Turkey.
¹¹⁰ Department of Anthropology, Binghamton University, Binghamton, NY 13902, USA.
¹¹¹ Archaeological Institute of America, Boston, MA 02108, USA.
¹¹² Institute of Classical Archaeology, Charles University, 11636 Prague, Czechia.
¹¹³ "Olga Necrasov" Centre of Anthropological Research, Romanian Academy Iași Branch, 2012 Iaşi Romania.
¹¹⁴ Scientific Research Center of the Historical and Cultural Heritage, 0010 Yerevan, Armenia.
¹¹⁵ Thracology Scientific Research Laboratory of the State University of Moldova, Department of Academic Management, Academy of Science of Moldova, 2009 Chișinău, Moldova.
¹¹⁶ Anthropological Center of the Croatian Academy of Sciences and Arts, 10000 Zagreb, Croatia.
¹¹⁷ Department of Archaeology, Faculty of Science and Arts, Pamukkale University, 20070 Denizli, Turkey.
¹¹⁸ Faculty of Archaeology, University of Warsaw, 00-927 Warszawa, Poland.
¹¹⁹ Ephorate of Antiquities of East Attica, Ministry of Culture and Sports, 10682 Athens, Greece.
¹²⁰ Austrian Archaeological Institute at the Austrian Academy of Sciences, 1190 Vienna, Austria.
¹²¹ Department of Anthropology, Faculty of Letters, Istanbul University, 34134 Istanbul, Turkey.
¹²² Institute of Archaeogenomics, Research Centre for the Humanities, Eötvös Loránd Research Network, 1097 Budapest, Hungary.
¹²³ Archaeology Museum of North Macedonia, 1000 Skopje, North Macedonia.
¹²⁴ Culture and Patrimony Department, University of Avignon, F-84029 Avignon, France.
¹²⁵ Department of the History of Art and Archaeology, Université Paris 1 Panthéon-Sorbonne, 75006 Paris, France.
¹²⁶ Department of Archaeology, Faculty of Science and Letters, Ondokuz Mayıs University, 55139 Atakum-Samsun, Turkey.
¹²⁷ Satu Mare County Museum, 440031 Satu Mare, Romania.
¹²⁸ School of Natural Sciences, University of Central Lancashire, Preston PR1 2HE, UK.
¹²⁹ Institute of Archaeology and Ethnology, Polish Academy of Sciences, 31-016 Kraków, Poland.
¹³⁰ Department of History and Social Sciences, Bryn Athyn College, Bryn Athyn, PA 19009, USA.
¹³¹ Penn Museum, University of Pennsylvania, Philadelphia, PA 19104, USA.
¹³² History of the Ancient World and Middle Ages Department, Moscow Region State University, Moscow Region, 141014 Mytishi, Russia.
¹³³ Department of Anthropology, Burdur Mehmet Akif Ersoy University, Istiklal Campus, 15100 Burdur, Turkey.
¹³⁴ Human Evolution and Archaeological Sciences, University of Vienna, 1030 Vienna, Austria.

^# Contributed equally.

PMID: 36007055
PMCID: PMC10064553
DOI: 10.1126/science.abm4247

The genetic history of the Southern Arc: A bridge between West Asia and Europe

Iosif Lazaridis et al. Science. 2022.

. 2022 Aug 26;377(6609):eabm4247.

doi: 10.1126/science.abm4247. Epub 2022 Aug 26.

Authors

Affiliations

¹ Department of Human Evolutionary Biology, Harvard University, Cambridge, MA 02138, USA.
² Department of Genetics, Harvard Medical School, Boston, MA 02115, USA.
³ Department of Evolutionary Anthropology, University of Vienna, 1030 Vienna, Austria.
⁴ Department of Anthropology, Faculty of Letters, Mardin Artuklu University, 47510 Artuklu, Mardin, Turkey.
⁵ Department of Anthropology, Faculty of Letters, Sivas Cumhuriyet University, 58140 Sivas, Turkey.
⁶ Department of History, Adelphi University, Garden City, NY 11530, USA.
⁷ Institute of Archaeology and Ethnography, NAS RA, 0025 Yerevan, Armenia.
⁸ Samsun Museum of Archeology and Ethnography, Kale Mahallesi, Merkez, İlkadım, 55030 Samsun, Turkey.
⁹ Iskra Museum of History, 6100 Kazanlak, Bulgaria.
¹⁰ Historical Musem in Kotor, 85330 Kotor, Montenegro.
¹¹ Institute of Archaeology, 11000 Belgrade, Serbia.
¹² Department of Archaeology, University of York, York YO1 7EP, UK.
¹³ Amasya Archaeology Museum, Mustafa Kemal Paşa Caddesi, 05000 Amasya, Turkey.
¹⁴ Department of Anthropology, Faculty of Arts and Science, Burdur Mehmet Akif University, 15100 Burdur, Turkey.
¹⁵ National Institute of Archaeology and Museum, Bulgarian Academy of Sciences, 1000 Sofia, Bulgaria.
¹⁶ Yambol Regional Historical Museum, 8600 Yambol, Bulgaria.
¹⁷ Archaeological Museum in Zagreb, 10000 Zagreb, Croatia.
¹⁸ Department of Archaeology and Heritage Studies, University of Tirana, 1010 Tirana, Albania.
¹⁹ Department of Animal and Human Physiology, Faculty of Biology, School of Sciences, National and Kapodistrian University of Athens, 10679 Athens, Greece.
²⁰ Department of Archaeology, Faculty of Humanities, Van Yüzüncü Yıl University, 65090 Tuşba, Van, Turkey.
²¹ Department of Archaeology, Faculty of Science and Letters, Düzce University, 81620 Düzce, Turkey.
²² Stratum Ltd., 21218 Seget Donji, Croatia.
²³ Independent Researcher, Berkeley, CA 94720, USA.
²⁴ School of History, Classics and Archaeology, University of Edinburgh, Edinburgh EH8 9AG, UK.
²⁵ The Italian Academy for Advanced Studies in America, Columbia University, New York, NY 10027, USA.
²⁶ Center for Conservation and Archaeology of Montenegro, 81250 Cetinje, Montenegro.
²⁷ Howard Hughes Medical Institute, Harvard Medical School, Boston, MA 02115, USA.
²⁸ Servizio di Bioarcheologia, Museo delle Civiltà, 00144 Rome, Italy.
²⁹ Centre for Applied Bioanthropology, Institute for Anthropological Research, 10000 Zagreb, Croatia.
³⁰ Department of Archaeology, University of Veliko Tarnovo "St. Cyril and St. Methodius," 5003 Veliko Tarnovo, Bulgaria.
³¹ Hellenic Ministry of Culture and Sports, Ephorate of Antiquities of Piraeus and the Islands, 10682 Piraeus, Greece.
³² "Orheiul Vechi" Cultural-Natural Reserve, Institute of Bioarchaeological and Ethnocultural Research, 3552 Butuceni, Moldova.
³³ National Archaeological Agency, 2012 Chișinău, Moldova.
³⁴ Archaeological Museum in Zadar, 23000 Zadar, Croatia.
³⁵ "Francisc I. Rainer" Institute of Anthropology, 050711 Bucharest, Romania.
³⁶ Department of Oral and Maxillo-Facial Sciences, Sapienza University of Rome, 00161 Rome, Italy.
³⁷ Institutes of Energy and the Environment, The Pennsylvania State University, University Park, PA 16802, USA.
³⁸ Department of Classics, University of Cincinnati, Cincinnati, OH 45221, USA.
³⁹ Independent Researcher, Aberystwyth SY23 4UH, UK.
⁴⁰ Center of Archaeology, Institute of Cultural Heritage, Academy of Science of Moldova, 2001 Chișinău, Moldova.
⁴¹ Department of Archaeology, Faculty of Letters, Ege University, 35100 Bornova-Izmir, Turkey.
⁴² Museum of Anthropological Archaeology, University of Michigan, Ann Arbor, MI 48109, USA.
⁴³ Narodni muzej Pančcevo, 26101 Pančevo, Serbia.
⁴⁴ Human Paleoecology and Isotope Geochemistry Lab, Department of Anthropology, The Pennsylvania State University, University Park, PA 16802, USA.
⁴⁵ Regional Museum of History - Veliko Tarnovo, 5000 Veliko Tarnovo, Bulgaria.
⁴⁶ Department of Archaeology, Faculty of Science and Letters, Gaziantep University, 27310 Gaziantep, Turkey.
⁴⁷ Mardin Archaeological Museum, Şar, Cumhuriyet Meydanı üstü, 47100 Artuklu, Mardin, Turkey.
⁴⁸ Muğla İl Kültür ve Turizm Müdürlüğü, 48000 Muğla, Turkey.
⁴⁹ Research Centre for Anthropology and Health (CIAS), Department of Life Sciences, University of Coimbra, 3000-456 Coimbra, Portugal.
⁵⁰ Prahova County Museum of History and Archaeology, 100042 Ploiești, Romania.
⁵¹ Institut Català de Paleoecologia Humana i Evolució Social, 43007 Tarragona, Spain.
⁵² Departament d'Història i Història de l'Art, Universitat Rovira i Virgili, 43002 Tarragona, Spain.
⁵³ School of Archaeology and Earth Institute, University College Dublin, Belfield, Dublin 4, Ireland.
⁵⁴ Department of Archaeology, Durham University, Durham DH1 3LE, UK.
⁵⁵ Department of Archaeology, Faculty of Science and Letters, Çukurova University, 01330 Balçalı-Sarıçam-Adana, Turkey.
⁵⁶ Department of Anthropology, Faculty of Humanities, Ankara University, 06100 Sıhhiye, Ankara, Turkey.
⁵⁷ Department of Archaeology, Faculty of Science and Letters, Bursa Uludağ University, 16059 Görükle, Bursa, Turkey.
⁵⁸ Department of Biological Anthropology, Institute of Biology, Eötvös Loránd University, 1053 Budapest, Hungary.
⁵⁹ Department of Cultures, University of Helsinki, 00100 Helsinki, Finland.
⁶⁰ Department of Ecology and Nature Protection, Yerevan State University, 0025 Yerevan, Armenia.
⁶¹ Regional Museum of History, 6300 Haskovo, Bulgaria.
⁶² Ministry of Culture and Tourism, İsmet İnönü Bulvarı, 06100 Emek, Ankara, Turkey.
⁶³ Museum of the City of Skopje, 1000 Skopje, North Macedonia.
⁶⁴ Malcolm H. Wiener Laboratory, American School of Classical Studies at Athens, 10676 Athens, Greece.
⁶⁵ Department of Archaeology, Faculty of Letters, Atatürk University, 25100 Erzurum, Turkey.
⁶⁶ Muğla Archaeological Museum and Yatağan Thermal Power Generation Company, Rescue Excavations, 48000 Muğla, Turkey.
⁶⁷ Department of Anthropology, University of California, Santa Barbara, Santa Barbara, CA 93106, USA.
⁶⁸ Bodrum Museum of Underwater Archeology, Çarşı Neighbourhood, 48400 Bodrum, Muğla, Turkey.
⁶⁹ Department of Anthropology, Hungarian Natural History Museum, 1117 Budapest, Hungary.
⁷⁰ Department of Anthropology, Natural History Museum Vienna, 1010 Vienna, Austria.
⁷¹ Institute of Archaeology, 10000 Zagreb, Croatia.
⁷² Faculty of Biology, Adam Mickiewicz University in Poznań, 61-614 Poznań, Poland.
⁷³ Municipal Museum Vinkovci, 32100 Vinkovci, Croatia.
⁷⁴ Prehistory Department, Albanian Institute of Archaeology, Academy of Albanian Studies, 1000 Tirana, Albania.
⁷⁵ National Museum in Ohrid, 6000 Ohrid, North Macedonia.
⁷⁶ ArchaeoSciences Division, Research Institute of the University of Bucharest, University of Bucharest, 050663 Bucharest, Romania.
⁷⁷ Department of Archaeology, St. Kliment Ohridski University of Sofia, 1504 Sofia, Bulgaria.
⁷⁸ Department of Anthropology, University of California, San Diego, La Jolla, CA 92093, USA.
⁷⁹ Key Research Institute of Yellow River Civilization and Sustainable Development and the Collaborative Innovation Center on Yellow River Civilization of Henan Province, Laboratory of Yellow River Cultural Heritage, Henan University, 475001 Kaifeng, China.
⁸⁰ European Academy of Sciences and Arts, 5020 Salzburg, Austria.
⁸¹ Science and Technology in Archaeology and Culture Research Center, The Cyprus Institute, 2121 Aglantzia, Nicosia, Cyprus.
⁸² Broad Institute of Harvard and MIT, Cambridge, MA 02142, USA.
⁸³ Cotsen Institute of Archaeology, University of California, Los Angeles, Los Angeles, CA 90095, USA.
⁸⁴ Department of Archaeology, University of Reading, Reading RG6 6AB, UK.
⁸⁵ National Museum of Kikinda, 23300 Kikinda, Serbia.
⁸⁶ Department of Archaeogenetics, Max Planck Institute for Evolutionary Anthropology, 04103 Leipzig, Germany.
⁸⁷ University of Pennsylvania Museum of Archaeology and Anthropology, Philadelphia, PA 19104, USA.
⁸⁸ School of Archaeological and Forensic Sciences, Faculty of Life Sciences, University of Bradford, Bradford BD7 1DP, UK.
⁸⁹ Department of Biology, Grand Valley State University, Allendale, MI 49401, USA.
⁹⁰ BIOMICs Research Group, University of the Basque Country UPV/EHU, 01006 Vitoria-Gasteiz, Spain.
⁹¹ Department of Anthropology and Middle Eastern Cultures, Mississippi State University, Mississippi State, MS 39762, USA.
⁹² Paul and Alexandra Canellopoulos Museum, 105-55 Athens, Greece.
⁹³ Faculty of Philosophy, School of History and Archaeology, Aristotle University of Thessaloniki, 54124 Thessaloniki, Greece.
⁹⁴ Ephorate of Paleoantropology and Speleology, Greek Ministry of Culture, 11636 Athens, Greece.
⁹⁵ Trogir Town Museum, 21220 Trogir, Croatia.
⁹⁶ Moldavian Historic-Geographical Society, 2044 Chișinău, Moldova.
⁹⁷ French School of Archaeology at Athens, 10680 Athens, Greece.
⁹⁸ Department of Armenian History, Armenian State Pedagogical University After Khachatur Abovyan, 0010 Yerevan, Armenia.
⁹⁹ Department of Archaeology, Faculty of Humanities and Social Sciences, University of Zagreb, 10000 Zagreb, Croatia.
¹⁰⁰ Independent Researcher, 10000 Zagreb, Croatia.
¹⁰¹ Laboratory for Archaeological Chemistry, University of Wisconsin-Madison, Madison, WI 53706, USA.
¹⁰² Institute for Quaternary Paleontology and Geology, Croatian Academy of Sciences and Arts, 10000 Zagreb, Croatia.
¹⁰³ Sulaymaniyah Directorate of Antiquities and Heritage, 46010 Sulaymaniyah, Iraq.
¹⁰⁴ Pridnestrovian University named after Taras Shevchenko, 3300 Tiraspol, Moldova.
¹⁰⁵ The Netherlands Institute for the Near East, 2311 Leiden, Netherlands.
¹⁰⁶ Institute of Experimental Morphology, Pathology and Archeology with Museum, Bulgarian Academy of Science, 1113 Sofia, Bulgaria.
¹⁰⁷ Department of Anthropology, Faculty of Science and Letters, Hitit University, 19040 Çorum, Turkey.
¹⁰⁸ School of Anatomical Sciences, The University of the Witwatersrand, 2193 Johannesburg, South Africa.
¹⁰⁹ Department of Anthropology, Faculty of Language and History - Geography, Ankara University, 06100 Sıhhiye, Ankara, Turkey.
¹¹⁰ Department of Anthropology, Binghamton University, Binghamton, NY 13902, USA.
¹¹¹ Archaeological Institute of America, Boston, MA 02108, USA.
¹¹² Institute of Classical Archaeology, Charles University, 11636 Prague, Czechia.
¹¹³ "Olga Necrasov" Centre of Anthropological Research, Romanian Academy Iași Branch, 2012 Iaşi Romania.
¹¹⁴ Scientific Research Center of the Historical and Cultural Heritage, 0010 Yerevan, Armenia.
¹¹⁵ Thracology Scientific Research Laboratory of the State University of Moldova, Department of Academic Management, Academy of Science of Moldova, 2009 Chișinău, Moldova.
¹¹⁶ Anthropological Center of the Croatian Academy of Sciences and Arts, 10000 Zagreb, Croatia.
¹¹⁷ Department of Archaeology, Faculty of Science and Arts, Pamukkale University, 20070 Denizli, Turkey.
¹¹⁸ Faculty of Archaeology, University of Warsaw, 00-927 Warszawa, Poland.
¹¹⁹ Ephorate of Antiquities of East Attica, Ministry of Culture and Sports, 10682 Athens, Greece.
¹²⁰ Austrian Archaeological Institute at the Austrian Academy of Sciences, 1190 Vienna, Austria.
¹²¹ Department of Anthropology, Faculty of Letters, Istanbul University, 34134 Istanbul, Turkey.
¹²² Institute of Archaeogenomics, Research Centre for the Humanities, Eötvös Loránd Research Network, 1097 Budapest, Hungary.
¹²³ Archaeology Museum of North Macedonia, 1000 Skopje, North Macedonia.
¹²⁴ Culture and Patrimony Department, University of Avignon, F-84029 Avignon, France.
¹²⁵ Department of the History of Art and Archaeology, Université Paris 1 Panthéon-Sorbonne, 75006 Paris, France.
¹²⁶ Department of Archaeology, Faculty of Science and Letters, Ondokuz Mayıs University, 55139 Atakum-Samsun, Turkey.
¹²⁷ Satu Mare County Museum, 440031 Satu Mare, Romania.
¹²⁸ School of Natural Sciences, University of Central Lancashire, Preston PR1 2HE, UK.
¹²⁹ Institute of Archaeology and Ethnology, Polish Academy of Sciences, 31-016 Kraków, Poland.
¹³⁰ Department of History and Social Sciences, Bryn Athyn College, Bryn Athyn, PA 19009, USA.
¹³¹ Penn Museum, University of Pennsylvania, Philadelphia, PA 19104, USA.
¹³² History of the Ancient World and Middle Ages Department, Moscow Region State University, Moscow Region, 141014 Mytishi, Russia.
¹³³ Department of Anthropology, Burdur Mehmet Akif Ersoy University, Istiklal Campus, 15100 Burdur, Turkey.
¹³⁴ Human Evolution and Archaeological Sciences, University of Vienna, 1030 Vienna, Austria.

^# Contributed equally.

PMID: 36007055
PMCID: PMC10064553
DOI: 10.1126/science.abm4247

Abstract

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.

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Figures

**Fig. 1.. Studied individuals and PCA analysis.**
(A) The geography of the Southern Arc; Sampling locations of previously published individuals (grey circle), new data on published individuals (pink square), and new individuals (yellow circle); convex hulls of individuals from each present-day country are also shown. (B) Timeline of studied individuals (random uniform jitter applied to the vertical dimension). (C) Principal components analysis of ancient individuals projected on modern West Eurasian variation. Country names represented by 3-letter ISO codes.

**Fig. 2:. The Anatolian heartland.**
Panels (A-E) show five components of ancestry in Anatolia from the Pottery Neolithic to the Roman/Byzantine period. Boxes in this and subsequent figures indicate the temporal extent (horizontal) and 95% confidence interval (±1.96s.e.) for each period; we also show the fit (solid line) and 5/95% (dotted lines) of the fit of a heteroskedastic Gaussian process (55) on the individuals without any assignment to populations, which allows us to appreciate the degree of variation in ancestry in each time period. The results show that across the peninsula the post-Neolithic period was characterized by expansion of Caucasus hunter-gatherer (CHG)-related ancestry (A) and dilution of northwest Anatolian-related ancestry (E). European hunter-gatherer-related ancestry from both the steppe/eastern Europe (B) and the Balkans (D) was insignificant until the last 3,000 years. A detailed look at the Chalcolithic/Bronze Age period (F) shows that populations there had ancestry intermediate between early farmers from western/central Anatolia (Barcın(10), Tepecik-Çiftlik(13), and Çatalhöyük(12)) and southeastern Anatolia (northern Mesopotamia at Mardin) on the other, the result of admixture between the preceding Neolithic populations, without discernible external influences (that would have elevated any of the five components above their Neolithic levels).

**Fig. 3:. Yamnaya origins and expansions.**
(A) The earliest inhabitants of the steppe (Eastern hunter-gatherers; EHG) were followed by Caucasus hunter-gatherer (CHG)-admixed populations by ~5000BCE and by Anatolian/Levantine-admixed populations by ~3000BCE with the emergence of the Yamnaya-Afanasievo genetic cluster. The proportion of Balkan hunter-gatherer-related ancestry (not shown) is 0.8±0.6% in the Yamnaya cluster and −0.5±0.5% in the Afanasievo. (B) The Yamnaya had nearly half their ancestry from CHG, higher than any Bronze Age Europeans from the Balkans, Italy, or central-northern Europe, but their CHG-EHG balance was equal, similar to the Corded Ware/Beaker clusters of central/northern Europe and contrasting with Southeastern and Mediterranean Europe where CHG was significantly higher than EHG. (95% confidence intervals of ±1.96s.e. are shown.)

**Fig. 4:. Genetic heterogeneity in Southeastern Europe following the Yamnaya expansion.**
Panels (A-E) show five components of ancestry in Southeastern Europe. The replacement of hunter-gatherer by early farmer ancestry (panels D and E) was followed by the rise of Caucasus hunter-gatherer (CHG) and Eastern hunter-gatherer (EHG) ancestry over the last 5,000 years (panels A and B) with Levantine ancestry being relatively unimportant and showing no discernible temporal pattern (C). In panel (F) we show a linear regression of population dates (using directly radiocarbon dated individuals for each population) on admixture times in generations; more recent populations have older admixture times, and the regression places admixture between populations related to the southeast European Neolithic and Yamnaya at 4853±205 years ago and the generation length at 28±4 years, virtually identical to its independent empirical estimation of 28 years.

**Fig. 5:. A genetic history of Armenia.**
Panels (A-D) show changes in the four components of ancestry. (A) Caucasus hunter-gatherer (CHG) is the most important component in all ages, rising to its maximum in the Kura-Araxes culture of the Early Bronze Age. (B) Eastern hunter-gatherer (EHG) ancestry first appears in the Chalcolithic at Areni Cave, disappears during the Kura-Araxes period, re-appears strongly in the Middle-Late Bronze Age period and decreases to ~1/3 of its peak value by ~2,000 years ago. (C, D) Levantine and Anatolian ancestry were present in all periods as minority components. Balkan hunter-gatherer ancestry (not shown) is <1% in all periods. (All individuals shown are from Armenia save for two Neolithic and a Chalcolithic individual previously published from Azerbaijan). (E) During the Middle-to-Late Bronze Age peak, Armenia had more EHG ancestry than its neighbors in West Asia (Anatolia, the Levant, and Iran), (F) C14-dated Bronze-to-Iron Age individuals from Armenia admixed 52.2±8.0 generations (1460±224 years) prior to their average date of 1119BCE, or ~2579BCE (mid-3rd millennium BCE), assuming a generation length of 28 years(56) (we use Early Bronze Age Armenia and Yamnaya cluster individuals from Russia as proxy sources).

**Fig. 6:. Y-chromosome links between the Southern Arc and the Eurasian steppe.**
(A) Phylogeny of haplogroup R-L389 (R1b1a1) with TMRCA estimates of yfull.com. (B) Caucasus hunter-gatherer (CHG)/Eastern hunter-gatherer (EHG) ancestral composition of R-L389 Y-chromosome individuals. (C) R-L389 individuals from the Southern Arc, representing a subset of the individuals plotted in panel B. Individuals more than 2,000 years old are shown.

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Comment in

Ancient genomes and West Eurasian history.
Arbuckle BS, Schwandt Z. Arbuckle BS, et al. Science. 2022 Aug 26;377(6609):922-923. doi: 10.1126/science.add9059. Epub 2022 Aug 25. Science. 2022. PMID: 36007032

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The genetic history of the Southern Arc: A bridge between West Asia and Europe

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