. 2022 Aug 26;377(6609):940-951.

doi: 10.1126/science.abq0755. Epub 2022 Aug 25.

A genetic probe into the ancient and medieval history of Southern Europe and West Asia

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, 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: 36007020
PMCID: PMC10019558
DOI: 10.1126/science.abq0755

A genetic probe into the ancient and medieval history of Southern Europe and West Asia

Iosif Lazaridis et al. Science. 2022.

. 2022 Aug 26;377(6609):940-951.

doi: 10.1126/science.abq0755. Epub 2022 Aug 25.

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, 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: 36007020
PMCID: PMC10019558
DOI: 10.1126/science.abq0755

Abstract

Literary and archaeological sources have preserved a rich history of Southern Europe and West Asia since the Bronze Age that can be complemented by genetics. Mycenaean period elites in Greece did not differ from the general population and included both people with some steppe ancestry and others, like the Griffin Warrior, without it. Similarly, people in the central area of the Urartian Kingdom around Lake Van lacked the steppe ancestry characteristic of the kingdom's northern provinces. Anatolia exhibited extraordinary continuity down to the Roman and Byzantine periods, with its people serving as the demographic core of much of the Roman Empire, including the city of Rome itself. During medieval times, migrations associated with Slavic and Turkic speakers profoundly affected the region.

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Conflict of interest statement

Competing interests: The authors declare that they have no competing interests.

Figures

**Fig. 1:. Genetic heterogeneity in the Aegean.**
(A) A map of Aegean sites. (B) Timeline of Aegean individuals, with vertical jitter added to distinguish contemporaneous individuals. Ancestry changes of five components (C-G) show an increase of Caucasus hunter-gatherer (CHG) and Eastern European hunter-gatherer (EHG) ancestry over time and a dilution of Anatolian Neolithic ancestry. During the Minoan and Mycenaean periods of the Bronze Age (H) Eastern European hunter-gatherer ancestry was variable, absent in Minoan individuals of Crete and present in most, but not all Mycenaean individuals of the mainland.

**Fig. 2:. The Kingdom of Urartu and its neighbors.**
Panels (A-D) show comparisons of ancestry in four ancestral components (SRB_Iron_Gates_HG, the 5^th component of the model of (1) is negligible). This analysis shows a stark contrast between Armenia and the other populations in terms of Eastern European hunter-gatherer ancestry (B), and between Van and Assyrian Mesopotamia in terms of Levantine ancestry (C). When unlabeled individuals are ordered in increasing Eastern European hunter-gatherer ancestry, Assyrian Mesopotamia, Van lack this ancestry (except an outlier individual from Van), while individuals from Armenia mostly possess it, and those from Hasanlu have a limited range from zero Eastern European hunter-gatherer ancestry to a maximum level that is less than that seen in Armenia.

**Fig. 3:. The Roman Empire East and West.**
(A) The Imperial period Romans from the vicinity of the city of Rome in Central Italy resembled Roman/Byzantine Anatolians in their average admixture proportions (95% confidence interval (C.I.) of ±1.96 standard errors shown as boxes and a heteroskedastic Gaussian process is fitted to unlabeled Italian and Anatolian individuals; dashed lines indicate 5% and 95% quantiles). (B) P-values of the Baringhaus-Franz multivariate two-sample test(47) for pairs of populations indicate that Imperial Romans can be drawn from the same distribution as Roman/Byzantine ones (p=0.19), but are significantly different (p≤2.16e-03) from all other periods of Italy. (C) Hierarchical clustering of raw ancestry estimates of diverse individuals shows overlapping distributions of Imperial Roman and Anatolian Roman/Byzantine individuals (black) without knowledge of their ancestry labels and differentiated from the distributions of SE Europe, Armenia, and the Levant.

**Fig. 4:. Central Asian Turkic admixture in Anatolia.**
(A) Individuals from Çapalıbağ (1300-1650CE) and present-day Turkish individuals are intermediate between Byzantine Anatolia and 500-1500CE Central Asians along a global principal components analysis distinguishing West from East Eurasians (left-to-right on the horizontal dimension; noise added on the vertical dimension to distinguish points). (B) 2-way unsupervised ADMIXTURE analysis of “eastern” ancestry: Byzantine: (0%), present-day Turkish (9%), Çapalıbağ (18%), Central Asian individuals differ between 100% (in Mongolia) to 43% (some ancient populations of Kazakhstan and Kyrgyzstan). (C) Individuals from Çapalıbağ in Turkey admixed 12.2±1.4 generations (342±39 years) prior to their time using Byzantine Anatolians and Central Asians (from 500-1500CE) as sources. (D) Present-day Turkish people genotyped on the Human Origins array (35) admixed 30.6±1.9 generations ago (857±53 years) using the same sources as in (D).

**Fig. 5:. Byzantine and Medieval Southeastern Europe.**
We sort admixture proportions of Anatolian Neolithic ancestry to investigate the dilution of this ancestry in present-day populations from Southeastern Europe. Roman/Medieval/Byzantine-era individuals are indicated in bold. During the Bronze Age the range of this ancestry was immense as observed in (1), but present-day people from the Balkans have less of this ancestry than was the case from the Bronze Age through the Iron Age and down to the classical antiquity (Ancient). Medieval/Byzantine people from the Balkans were diverse, with some (right) continuing the ancient pattern of high Anatolian Neolithic ancestry, several (middle) overlapping with the range of present-day people, and some (left) having as little such ancestry as present-day Balto-Slavic people from Eastern Europe.

**Fig. 6:. Pigmentation in West Eurasia.**
We show the temporal distribution of genetically predicted Eye (A), Skin (B), and Hair (C) color in West Eurasians of the last 16,000 years; each point represents an individual, with the top row for each subphenotype corresponding to Southern Arc and the bottom row corresponding to northern, central and western Europeans and people of the Eurasian steppe. Panel (D) shows composite phenotypes of all three aspects of pigmentation using the same color scheme as A-C and denoted as eye color (circle), hair color (top), and skin color (bottom) in the composite phenotype symbols. The modal phenotype of West Eurasians had brown eyes, intermediate skin pigmentation, and brown hair, with the highest prevalence (Fisher’s exact test) of low pigmentation outside the Southern Arc (in the rest of Europe and the Eurasian steppe).

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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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References

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A genetic probe into the ancient and medieval history of Southern Europe and West Asia

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A genetic probe into the ancient and medieval history of Southern Europe and West Asia

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