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- Iain Mathieson+,1,
- Songül Alpaslan Roodenberg1,
- Cosimo Posth2,3,
- Anna Szécsényi-Nagy4,
- Nadin Rohland1,
- Swapan Mallick1,5,
- Iñigo Olade1,
- Nasreen Broomandkhoshbacht1,5,
- Olivia Cheronet6,
- Daniel Fernandes6,7,
- Matthew Ferry1,5,
- Beatriz Gamarra6,
- Gloria González Fortes8,
- Wolfgang Haak2,9,
- Eadaoin Harney1,5,
- Ben Krause-Kyora2,
- Isil Kucukkalipci3,
- Megan Michel1,5,
- Alissa Mittnik3,
- Kathrin Nägele2,
- Mario Novak6,10,
- Jonas Oppenheimer1,5,
- Nick Patterson11,
- Saskia Pfrengle3,
- Kendra Sirak6,12,
- Kristin Stewardson1,5,
- Stefania Vai13,
- Stefan Alexandrov14,
- Kurt W. Alt15,16,17,
- Radian Andreescu18,
- Dragana Antonović19,
- Abigail Ash6,
- Nadezhda Atanassova20,
- Krum Bacvarov14,
- Mende Balázs Gusztáv4,
- Hervé Bocherens21,22,
- Michael Bolus23,
- Adina Boroneanţ24,
- Yavor Boyadzhiev14,
- Alicja Budnik25,
- Josip Burmaz26,
- Stefan Chohadzhiev27,
- Nicholas J. Conard28,22,
- Richard Cottiaux29,
- Maja Čuka30,
- Christophe Cupillard31,32,
- Dorothée G. Drucker22,
- Nedko Elenski33,
- Michael Francken34,
- Borislava Galabova35,
- Georgi Ganetovski36,
- Bernard Gely37,
- Tamás Hajdu38,
- Veneta Handzhyiska39,
- Katerina Harvati34,22,
- Thomas Higham40,
- Stanislav Iliev41,
- Ivor Janković10,42,
- Ivor Karavanić43,42,
- Douglas J. Kennett44,
- Darko Komšo30,
- Alexandra Kozak45,
- Damian Labuda46,
- Martina Lari13,
- Catalin Lazar47,
- Maleen Leppek48,
- Krassimir Leshtakov39,
- Domenico Lo Vetro49,50,
- Dženi Los26,
- Ivaylo Lozanov39,
- Maria Malina23,
- Fabio Martini50,49,
- Kath McSweeney51,
- Harald Meller17,
- Marko Menđušić52,
- Pavel Mirea53,
- Vyacheslav Moiseyev54,
- Vanya Petrova39,
- T. Douglas Price55,
- Angela Simalcsik56,
- Luca Sineo57,
- Mario Šlaus58,
- Vladimir Slavchev59,
- Petar Stanev33,
- Andrej Starović60,
- Tamás Szeniczey38,
- Sahra Talamo61,
- Maria Teschler-Nicola62,
- Corinne Thevenet63,
- Ivan Valchev39,
- Frédérique Valentin64,
- Sergey Vasilyev65,
- Fanica Veljanovska66,
- Svetlana Venelinova67,
- Elizaveta Veselovskaya65,
- Bence Viola68,69,
- Cristian Virag70,
- Joško Zaninović71,
- Steve Zauner72,
- Philipp W. Stockhammer48,2,
- Giulio Catalano57,
- Raiko Krauβ74,
- David Caramelli13,
- Gunita Zariņa75,
- Bisserka Gaydarska76,
- Malcolm Lillie77,
- Alexey G. Nikitin78,
- Inna Potekhina79,
- Anastasia Papathanasiou80,
- Dušan Borić81,
- Clive Bonsall51,
- Johannes Krause2,3,
- Ron Pinhasi*+,6,82 and
- David Reich*+,1,11,5
- 1Department of Genetics, Harvard Medical School, Boston 02115 MA USA
- 2Department of Archaeogenetics, Max Planck Institute for the Science of Human History, 07745 Jena, Germany
- 3Institute for Scientific Archaeology, University of Tuebingen, Germany
- 4Laboratory of Archaeogenetics, Institute of Archaeology, Research Centre for the Humanities, Hungarian Academy of Sciences, H-1097 Budapest, Hungary
- 5Howard Hughes Medical Institute, Harvard Medical School, Boston 02115 MA USA
- 6School of Archaeology and Earth Institute, University College Dublin, Belfield, Dublin 4, Republic of Ireland
- 7CIAS, Department of Life Sciences, University of Coimbra, 3000-456 Coimbra, Portugal
- 8Department of Life Sciences and Biotechnology, University of Ferrara, Via L. Borsari 46. Ferrara 44100 Italy
- 9Australian Centre for Ancient DNA, School of Biological Sciences, The University of Adelaide, SA-5005 Adelaide, Australia
- 10Institute for Anthropological Research, Ljudevita Gaja 32, 10000 Zagreb, Croatia
- 11Broad Institute of Harvard and MIT, Cambridge MA
- 12Department of Anthropology, Emory University, Atlanta, Georgia 30322, USA
- 13Dipartimento di Biologia, Università di Firenze, 50122 Florence, Italy
- 14National Institute of Archaeology and Museum, Bulgarian Academy of Sciences, 2 Saborna Str., BG-1000 Sofia, Bulgaria
- 15Danube Private University, A-3500 Krems, Austria
- 16Department of Biomedical Engineering and Integrative Prehistory and Archaeological Science, CH-4123 Basel-Allschwil, Switzerland
- 17State Office for Heritage Management and Archaeology Saxony-Anhalt and State Museum of Prehistory, D-06114 Halle, Germany
- 18Romanian National History Museum, Bucharest, Romania
- 19Institute of Archaeology, Belgrade, Serbia
- 20Institute of Experimental Morphology, Pathology and Anthropology with Museum, Bulgarian Academy of Sciences, Sofia, Bulgaria
- 21Department of Geosciences, Biogeology, Universität Tübingen, Hölderlinstr. 12, 72074 Tübingen, Germany
- 22Senckenberg Centre for Human Evolution and Palaeoenvironment, University of Tuebingen, 72072 Tuebingen, Germany
- 23Heidelberg Academy of Sciences and Humanities, Research Center “The Role of Culture in Early Expansions of Humans” at the University of Tuebingen, Rumelinstraβ 23, 72070 Tuebingen, Germany
- 24‘Vasile Pârvan’ Institute of Archaeology, Romanian Academy
- 25Human Biology Department, Cardinal Stefan Wyszy.ski University, Warsaw, Poland
- 26KADUCEJ d.o.o Papandopulova 27, 21000 Split, Croatia
- 27St. Cyril and Methodius University, Veliko Turnovo, Bulgaria
- 28Department of Early Prehistory and Quaternary Ecology, University of Tuebingen, Schloss HohenTuebingen, 72070 Tuebingen, Germany
- 29INRAP/UMR 8215 Trajectoires, 21 Allee de l’Universite, 92023 Nanterre, France
- 30Archaeological Museum of Istria, Carrarina 3, 52100 Pula, Croatia
- 31Service Regional de l’Archeologie de Bourgogne-Franche-Comte, 7 rue Charles Nodier, 25043 Besancon Cedex, France
- 32Laboratoire Chronoenvironnement, UMR 6249 du CNRS, UFR des Sciences et Techniques, 16 route de Gray, 25030 Besancon Cedex, France
- 33Regional Museum of History Veliko Tarnovo, Veliko Tarnovo, Bulgaria
- 34Institute for Archaeological Sciences, Paleoanthropology, University of Tuebingen, Rumelinstrase 23, 72070 Tuebingen, Germany
- 35Laboratory for human bio-archaeology, Bulgaria, 1202 Sofia, 42, George Washington str
- 36Regional Museum of History, Vratsa, Bulgaria
- 37Service Regional de l’Archeologie d’Auvergne-Rhone-Alpes, Le Grenier d’Abondance, 6 quai Saint-Vincent, 69283 Lyon Cedex 01, France
- 38Eotvos Lorand University, Faculty of Science, Institute of Biology, Department of Biological Anthropology, H-1117 Pazmany Peter setany 1/c. Budapest, Hungary
- 39Department of Archaeology, Sofia University St. Kliment Ohridski, Bulgaria
- 40Oxford Radiocarbon Accelerator Unit, Research Laboratory for Archaeology and the History of Art, University of Oxford, Dyson Perrins Building, South Parks Road, OX1 3QY Oxford, UK
- 41Regional Museum of History, Haskovo, Bulgaria
- 42Department of Anthropology, University of Wyoming, 1000 E. University Avenue, Laramie, WY 82071, USA
- 43Department of Archaeology, Faculty of Humanities and Social Sciences, University of Zagreb, Ivana Lu.i.a 3, 10000 Zagreb, Croatia
- 44Department of Anthropology and Institutes for Energy and the Environment, Pennsylvania State University, University Park, PA 16802
- 45Institute of Archaeology, National Academy of Sciences of Ukraine, Kyiv, Ukraine
- 46CHU Sainte-Justine Research Center, Pediatric Department, Universite de Montreal, Montreal, PQ, Canada, H3T 1C5
- 47National History Museum of Romania, Calea Victoriei, no. 12, 030026, Bucharest, Romania
- 48Institute for Pre-and Protohistoric Archaeology and the Archaeology of the Roman Provinces, Ludwig-Maximilians-University, Schellingstr. 12, 80799 Munich, Germany
- 49Museo e Istituto fiorentino di Preistoria, 50122 Florence, Italy
- 50Dipartimento SAGAS -Sezione di Archeologia e Antico Oriente, Universita degli Studi di Firenze, 50122 Florence, Italy
- 51School of History, Classics and Archaeology, University of Edinburgh, Edinburgh EH8 9AG, United Kingdom
- 52Conservation Department in.ibenik, Ministry of Culture of the Republic of Croatia, Jurja.ulinovi.a 1, 22000.ibenik, Croatia
- 53Teleorman County Museum, str. 1848, no. 1, 140033 Alexandria, Romania
- 54Peter the Great Museum of Anthropology and Ethnography (Kunstkamera) RAS, 199034 St. Petersburg, Russia
- 55University of Wisconsin, Madison WI, USA
- 56Olga Necrasov Centre for Anthropological Research, Romanian Academy. Ia.i Branch, Theodor Codrescu St. 2, P.C. 700481, Ia.i, Romania
- 57Dipartimento di Scienze e tecnologie biologiche, chimiche e farmaceutiche, Lab. of Anthropology,Universita degli studi di Palermo, Italy
- 58Anthropological Center, Croatian Academy of Sciences and Arts, 10000 Zagreb, Croatia
- 59Regional Historical Museum Varna, Maria Luiza Blvd. 41, BG-9000 Varna, Bulgaria
- 60National Museum in Belgrade, 1a Republic sq., Belgrade, Serbia
- 61Department of Human Evolution, Max Planck Institute for Evolutionary Anthropology, 04103 Leipzig, Germany
- 62Department of Anthropology, Natural History Museum Vienna, 1010 Vienna, Austria
- 63INRAP/UMR 8215 Trajectoires, 21 Allée de l’Universite, 92023 Nanterre, France
- 64CNRS/UMR 7041 ArScAn MAE, 21 Allée de l’Universite, 92023 Nanterre, France
- 65Institute of Ethnology and Anthropology, Russian Academy of Sciences, Leninsky Pr., 32a, Moscow, 119991, Russia
- 66Archaeological Museum of Macedonia, Skopje
- 67Regional museum of history, Shumen, Bulgaria
- 68Department of Anthropology, University of Toronto, Toronto, Ontario, M5S 2S2, Canada
- 69Institute of Archaeology & Ethnography, Siberian Branch, Russian Academy of Sciences, Lavrentiev Pr. 17, Novosibirsk 630090, Russia
- 70Satu Mare County Museum Archaeology Department,V. Lucaciu, nr.21, Satu Mare, Romania
- 71Municipal Museum Drni., Domovinskog rata 54, 22320 Drni., Croatia
- 72anthropol -Anthropologieservice, Schadenweilerstrase 80, 72379 Hechingen, Germany
- 73Dipartimento di Scienze e tecnologie biologiche, chimiche e farmaceutiche, Lab. of Anthropology,Universita degli studi di Palermo.Italy
- 74Institute for Prehistory, Early History and Medieval Archaeology, University of Tuebingen, Germany
- 75Institute of Latvian History, University of Latvia, Kalpaka Bulv.ris 4, R.ga 1050, Latvia
- 76Department of Archaeology, Durham University, UK
- 77School of Environmental Sciences: Geography, University of Hull, Hull HU6 7RX, UK
- 78Department of Biology, Grand Valley State University, Allendale, Michigan, USA
- 79Department of Bioarchaeology, Institute of Archaeology, National Academy of Sciences of Ukraine
- 80Ephorate of Paleoanthropology and Speleology, Athens, Greece
- 81Department of Archaeology, Cardiff University, Cardiff, UK
- 82Department of Anthropology, University of Vienna, Althanstrasse 14, 1090 Vienna, Austria
- ↵+Correspondence to I.M. (iain_mathieson{at}hms.harvard.edu) or D.R. (reich{at}genetics.med.harvard.edu) or R.P. (ron.pinhasi{at}ucd.ie)
↵* These authors contributed equally to the manuscript