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Ancient human genomes suggest three ancestral populations for present-day Europeans

Iosif Lazaridis, Nick Patterson, Alissa Mittnik, Gabriel Renaud, Swapan Mallick, Karola Kirsanow, Peter H. Sudmant, Joshua G. Schraiber, Sergi Castellano, Mark Lipson, Bonnie Berger, Christos Economou, Ruth Bollongino, Qiaomei Fu, Kirsten I. Bos, Susanne Nordenfelt, Heng Li, Cesare de Filippo, Kay Prüfer, Susanna Sawyer, Cosimo Posth, Wolfgang Haak, Fredrik Hallgren, Elin Fornander, Nadin Rohland, Dominique Delsate, Michael Francken, Jean-Michel Guinet, Joachim Wahl, George Ayodo, Hamza A. Babiker, Graciela Bailliet, Elena Balanovska, Oleg Balanovsky, Ramiro Barrantes, Gabriel Bedoya, Haim Ben-Ami, Judit Bene, Fouad Berrada, Claudio M. Bravi, Francesca Brisighelli, George Busby, Francesco Cali, Mikhail Churnosov, David E. C. Cole, Daniel Corach, Larissa Damba, George van Driem, Stanislav Dryomov, Jean-Michel Dugoujon, Sardana A. Fedorova, Irene Gallego Romero, Marina Gubina, Michael Hammer, Brenna Henn, Tor Hervig, Ugur Hodoglugil, Aashish R. Jha, Sena Karachanak-Yankova, Rita Khusainova, Elza Khusnutdinova, Rick Kittles, Toomas Kivisild, William Klitz, Vaidutis Kučinskas, Alena Kushniarevich, Leila Laredj, Sergey Litvinov, Theologos Loukidis, Robert W. Mahley, Béla Melegh, Ene Metspalu, Julio Molina, Joanna Mountain, Klemetti Näkkäläjärvi, Desislava Nesheva, Thomas Nyambo, Ludmila Osipova, Jüri Parik, Fedor Platonov, Olga Posukh, Valentino Romano, Francisco Rothhammer, Igor Rudan, Ruslan Ruizbakiev, Hovhannes Sahakyan, Antti Sajantila, Antonio Salas, Elena B. Starikovskaya, Ayele Tarekegn, Draga Toncheva, Shahlo Turdikulova, Ingrida Uktveryte, Olga Utevska, René Vasquez, Mercedes Villena, Mikhail Voevoda, Cheryl Winkler, Levon Yepiskoposyan, Pierre Zalloua, Tatijana Zemunik, Alan Cooper, Cristian Capelli, Mark G. Thomas, Andres Ruiz-Linares, Sarah A. Tishkoff, Lalji Singh, Kumarasamy Thangaraj, Richard Villems, David Comas, Rem Sukernik, Mait Metspalu, Matthias Meyer, Evan E. Eichler, Joachim Burger, Montgomery Slatkin, Svante Pääbo, Janet Kelso, David Reich, Johannes Krause
doi: https://doi.org/10.1101/001552
Iosif Lazaridis
1Department of Genetics, Harvard Medical School, Boston, MA, 02115, USA
2Broad Institute of Harvard and MIT, Cambridge, MA, 02142, USA
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Nick Patterson
2Broad Institute of Harvard and MIT, Cambridge, MA, 02142, USA
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Alissa Mittnik
3Institute for Archaeological Sciences, University of Tübingen, Tübingen, 72074, Germany
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Gabriel Renaud
4Max Planck Institute for Evolutionary Anthropology, Leipzig, 04103, Germany
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Swapan Mallick
1Department of Genetics, Harvard Medical School, Boston, MA, 02115, USA
2Broad Institute of Harvard and MIT, Cambridge, MA, 02142, USA
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Karola Kirsanow
5Johannes Gutenberg University Mainz, Institute of Anthropology, Mainz, D-55128, Germany
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Peter H. Sudmant
6Department of Genome Sciences, University of Washington, Seattle, WA, 98195, USA
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Joshua G. Schraiber
7Department of Integrative Biology, University of California, Berkeley, CA, 94720-3140, USA
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Sergi Castellano
4Max Planck Institute for Evolutionary Anthropology, Leipzig, 04103, Germany
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Mark Lipson
8Department of Mathematics and Computer Science and Artificial Intelligence Laboratory, Massachusetts Institute of Technology, Cambridge, MA 02139, USA
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Bonnie Berger
2Broad Institute of Harvard and MIT, Cambridge, MA, 02142, USA
8Department of Mathematics and Computer Science and Artificial Intelligence Laboratory, Massachusetts Institute of Technology, Cambridge, MA 02139, USA
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Christos Economou
9Archaeological Research Laboratory, Stockholm University, 114 18, Sweden
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Ruth Bollongino
5Johannes Gutenberg University Mainz, Institute of Anthropology, Mainz, D-55128, Germany
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Qiaomei Fu
1Department of Genetics, Harvard Medical School, Boston, MA, 02115, USA
4Max Planck Institute for Evolutionary Anthropology, Leipzig, 04103, Germany
10Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, IVPP, CAS, Beijing, 100049, China
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Kirsten I. Bos
3Institute for Archaeological Sciences, University of Tübingen, Tübingen, 72074, Germany
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Susanne Nordenfelt
1Department of Genetics, Harvard Medical School, Boston, MA, 02115, USA
2Broad Institute of Harvard and MIT, Cambridge, MA, 02142, USA
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Heng Li
1Department of Genetics, Harvard Medical School, Boston, MA, 02115, USA
2Broad Institute of Harvard and MIT, Cambridge, MA, 02142, USA
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Cesare de Filippo
4Max Planck Institute for Evolutionary Anthropology, Leipzig, 04103, Germany
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Kay Prüfer
4Max Planck Institute for Evolutionary Anthropology, Leipzig, 04103, Germany
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Susanna Sawyer
4Max Planck Institute for Evolutionary Anthropology, Leipzig, 04103, Germany
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Cosimo Posth
3Institute for Archaeological Sciences, University of Tübingen, Tübingen, 72074, Germany
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Wolfgang Haak
11Australian Centre for Ancient DNA, School of Earth and Environmental Sciences, University of Adelaide, Adelaide, South Australia, SA 5005, Australia
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Fredrik Hallgren
12The Cultural Heritage Foundation, Västerås, 722 12, Sweden
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Elin Fornander
12The Cultural Heritage Foundation, Västerås, 722 12, Sweden
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Nadin Rohland
1Department of Genetics, Harvard Medical School, Boston, MA, 02115, USA
2Broad Institute of Harvard and MIT, Cambridge, MA, 02142, USA
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Dominique Delsate
13National Museum of Natural History, L-2160, Luxembourg
14National Center of Archaeological Research, National Museum of History and Art, L-2345, Luxembourg
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Michael Francken
15Department of Paleoanthropology, Senckenberg Center for Human Evolution and Paleoenvironment, University of Tübingen, Tübingen, D-72070, Germany
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Jean-Michel Guinet
13National Museum of Natural History, L-2160, Luxembourg
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Joachim Wahl
16State Office for Cultural Heritage Management Baden-Württemberg, Osteology, Konstanz, D-78467, Germany
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George Ayodo
17Center for Global Health and Child Development, Kisumu, 40100, Kenya
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Hamza A. Babiker
18Institutes of Evolution, Immunology and Infection Research, School of Biological Sciences, University of Edinburgh, Edinburgh, EH9 3JT, UK
19Biochemistry Department, Faculty of Medicine, Sultan Qaboos University, Alkhod, Muscat, 123, Oman
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Graciela Bailliet
20Laboratorio de Genética Molecular Poblacional, Instituto Multidisciplinario de Biología Celular (IMBICE), CCT-CONICET & CICPBA, La Plata, B1906APO, Argentina
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Elena Balanovska
21Research Centre for Medical Genetics, Moscow, 115478, Russia
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Oleg Balanovsky
21Research Centre for Medical Genetics, Moscow, 115478, Russia
22Vavilov Institute for General Genetics, Moscow, 119991, Russia
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Ramiro Barrantes
23Escuela de Biología, Universidad de Costa Rica, San José, 2060, Costa Rica
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Gabriel Bedoya
24Institute of Biology, Research group GENMOL, Universidad de Antioquia, Medellín, Colombia
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Haim Ben-Ami
25Rambam Health Care Campus, Haifa, 31096, Israel
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Judit Bene
26Department of Medical Genetics and Szentagothai Research Center, University of Pécs, Pécs, H-7624 Hungary
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Fouad Berrada
27Al Akhawayn University in Ifrane (AUI), School of Science and Engineering, Ifrane, 53000, Morocco
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Claudio M. Bravi
20Laboratorio de Genética Molecular Poblacional, Instituto Multidisciplinario de Biología Celular (IMBICE), CCT-CONICET & CICPBA, La Plata, B1906APO, Argentina
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Francesca Brisighelli
28Forensic Genetics Laboratory, Institute of Legal Medicine, Università Cattolica del Sacro Cuore, Rome, 00168, Italy
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George Busby
29Department of Zoology, University of Oxford, Oxford, OX1 3PS, UK
30Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford, OX3 7BN, UK
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Francesco Cali
31Laboratorio di Genetica Molecolare, IRCCS Associazione Oasi Maria SS, Troina, 94018, Italy
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Mikhail Churnosov
32Belgorod State University, Belgorod, 308015, Russia
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David E. C. Cole
33Department of Laboratory Medicine and Pathobiology, University of Toronto, Toronto, Ontario, M5G 1L5, Canada
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Daniel Corach
34Servicio de Huellas Digitales Genéticas, School of Pharmacy and Biochemistry, Universidad de Buenos Aires, 1113 CABA, Argentina
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Larissa Damba
35Institute of Cytology and Genetics, Siberian Branch of Russian Academy of Sciences, Novosibirsk, 630090, Russia
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George van Driem
36Institute of Linguistics, University of Bern, Bern, CH-3012, Switzerland
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Stanislav Dryomov
37Laboratory of Human Molecular Genetics, Institute of Molecular and Cellular Biology, Russian Academy of Science, Siberian Branch, Novosibirsk, 630090, Russia
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Jean-Michel Dugoujon
38Anthropologie Moléculaire et Imagerie de Synthèse, CNRS UMR 5288, Université Paul Sabatier Toulouse III, Toulouse, 31000, France
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Sardana A. Fedorova
39Yakut Research Center of Complex Medical Problems and North-Eastern Federal University, Yakutsk, 677010, Russia
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Irene Gallego Romero
40Department of Human Genetics, University of Chicago, Chicago, IL, 60637, USA
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Marina Gubina
35Institute of Cytology and Genetics, Siberian Branch of Russian Academy of Sciences, Novosibirsk, 630090, Russia
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Michael Hammer
41ARL Division of Biotechnology, University of Arizona, Tucson, AZ, 85721, USA
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Brenna Henn
42Department of Ecology and Evolution, Stony Brook University, Stony Brook, NY, 11794, USA
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Tor Hervig
43Department of Clinical Science, University of Bergen, Bergen, 5021, Norway
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Ugur Hodoglugil
44NextBio, part of Illumina, Santa Clara, CA, USA 95050
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Aashish R. Jha
40Department of Human Genetics, University of Chicago, Chicago, IL, 60637, USA
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Sena Karachanak-Yankova
45Dept. of Medical Genetics, National Human Genome Center, Medical University Sofia, Sofia, 1431, Bulgaria
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Rita Khusainova
46Institute of Biochemistry and Genetics, Ufa Research Centre, Russian Academy of Sciences, Ufa, 450054, Russia
47Department of Genetics and Fundamental Medicine, Bashkir State University, Ufa, 450074, Russia
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Elza Khusnutdinova
46Institute of Biochemistry and Genetics, Ufa Research Centre, Russian Academy of Sciences, Ufa, 450054, Russia
47Department of Genetics and Fundamental Medicine, Bashkir State University, Ufa, 450074, Russia
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Rick Kittles
48College of Medicine, University of Illinois at Chicago, Chicago, IL, 60607, USA
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Toomas Kivisild
49Division of Biological Anthropology, University of Cambridge, Cambridge, United Kingdom CB2 1QH
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William Klitz
50School of Public Health, University of California, Berkeley, CA, 94720, USA
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Vaidutis Kučinskas
51Department of Human and Medical Genetics, Vilnius University, Vilnius, LT-08661, Lithuania
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Alena Kushniarevich
52Estonian Biocentre, Evolutionary Biology group, Tartu, 51010, Estonia
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Leila Laredj
53Translational Medicine and Neurogenetics, Institut de Génétique et de Biologie Moléculaire et Cellulaire, Illkirch, 67404, France
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Sergey Litvinov
46Institute of Biochemistry and Genetics, Ufa Research Centre, Russian Academy of Sciences, Ufa, 450054, Russia
47Department of Genetics and Fundamental Medicine, Bashkir State University, Ufa, 450074, Russia
52Estonian Biocentre, Evolutionary Biology group, Tartu, 51010, Estonia
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Theologos Loukidis
54Currently employed by AMGEN; 33 Kazantzaki Str, Ilioupolis 16342, Athens, Greece
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Robert W. Mahley
55Gladstone Institutes, San Francisco, CA, 94158, USA
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Béla Melegh
26Department of Medical Genetics and Szentagothai Research Center, University of Pécs, Pécs, H-7624 Hungary
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Ene Metspalu
56Department of Evolutionary Biology, University of Tartu, Tartu, 51010, Estonia
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Julio Molina
57Centro de Investigaciones Biomédicas de Guatemala, Ciudad de Guatemala, Guatemala
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Joanna Mountain
58Research Department, 23andMe, Inc. Mountain View, CA, 94043, USA
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Klemetti Näkkäläjärvi
59Cultural Anthropology Program, University of Oulu, Oulu, 90014, Finland
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Desislava Nesheva
45Dept. of Medical Genetics, National Human Genome Center, Medical University Sofia, Sofia, 1431, Bulgaria
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Thomas Nyambo
60Department of Biochemistry, Muhimbili University of Health and Allied Sciences, Dar es Salaam, Tanzania
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Ludmila Osipova
35Institute of Cytology and Genetics, Siberian Branch of Russian Academy of Sciences, Novosibirsk, 630090, Russia
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Jüri Parik
56Department of Evolutionary Biology, University of Tartu, Tartu, 51010, Estonia
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Fedor Platonov
61Research Institute of Health, North-Eastern Federal University, Yakutsk, 677000, Russia
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Olga Posukh
35Institute of Cytology and Genetics, Siberian Branch of Russian Academy of Sciences, Novosibirsk, 630090, Russia
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Valentino Romano
62Dipartimento di Fisica e Chimica, Università di Palermo, Palermo, 90128, Italy
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Francisco Rothhammer
63Instituto de Alta Investigación, Universidad de Tarapacá, Arica, Chile
64Programa de Genética Humana ICBM Facultad de Medicina Universidad de Chile, Santiago, Chile
65Centro de Investigaciones del Hombre en el Desierto, Arica, Chile
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Igor Rudan
66Centre for Population Health Sciences, The University of Edinburgh Medical School, Edinburgh, Scotland, EH8 9AG, UK
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Ruslan Ruizbakiev
67Deceased: formerly of the Institute of Immunology, Academy of Science, Tashkent, 70000, Uzbekistan
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Hovhannes Sahakyan
52Estonian Biocentre, Evolutionary Biology group, Tartu, 51010, Estonia
68Laboratory of Ethnogenomics, Institute of Molecular Biology, National Academy of Sciences of Armenia, Yerevan, 0014, Armenia
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Antti Sajantila
69Department of Forensic Medicine, Hjelt Institute, University of Helsinki, Helsinki, 00014, Finland
70Institute of Applied Genetics, Department of Molecular and Medical Genetics, University of North Texas Health Science Center, Fort Worth, Texas 76107, USA
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Antonio Salas
71Unidade de Xenética, Departamento de Anatomía Patolóxica e Ciencias Forenses, and Instituto de Ciencias Forenses, Grupo de Medicina Xenómica (GMX), Facultade de Medicina, Universidade de Santiago de Compostela, Galcia, 15872, Spain
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Elena B. Starikovskaya
37Laboratory of Human Molecular Genetics, Institute of Molecular and Cellular Biology, Russian Academy of Science, Siberian Branch, Novosibirsk, 630090, Russia
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Ayele Tarekegn
72Research Fellow, Henry Stewart Group, Russell House, London WC1A 2HN, UK
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Draga Toncheva
45Dept. of Medical Genetics, National Human Genome Center, Medical University Sofia, Sofia, 1431, Bulgaria
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Shahlo Turdikulova
73Institute of Bioorganic Chemistry Academy of Sciences Republic of Uzbekistan, Tashkent, 100125, Uzbekistan
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Ingrida Uktveryte
51Department of Human and Medical Genetics, Vilnius University, Vilnius, LT-08661, Lithuania
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Olga Utevska
74Department of Genetics and Cytology, V.N. Karazin Kharkiv National University, Kharkiv, 61077, Ukraine
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René Vasquez
75Instituto Boliviano de Biología de la Altura, Universidad Autonoma Tomás Frías, Potosí, Bolivia
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Mercedes Villena
75Instituto Boliviano de Biología de la Altura, Universidad Autonoma Tomás Frías, Potosí, Bolivia
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Mikhail Voevoda
35Institute of Cytology and Genetics, Siberian Branch of Russian Academy of Sciences, Novosibirsk, 630090, Russia
76Inst. of Internal Medicine, Siberian Branch of Russian Acad. of Medical Sciences, Novosibirsk, 630089, Russia
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Cheryl Winkler
77Molecular Genetics Epidemiology Section, Frederick Natoinal Lab for Cancer Research, NCI, NIH, Frederick, MD 21702, USA
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Levon Yepiskoposyan
68Laboratory of Ethnogenomics, Institute of Molecular Biology, National Academy of Sciences of Armenia, Yerevan, 0014, Armenia
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Pierre Zalloua
78Lebanese American University, School of Medicine, Beirut, 13-5053, Lebanon
79Harvard School of Public Health, Boston, 02115, USA
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Tatijana Zemunik
80Department of Medical Biology, University of Split, School of Medicine, Split, 21000, Croatia
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Alan Cooper
11Australian Centre for Ancient DNA, School of Earth and Environmental Sciences, University of Adelaide, Adelaide, South Australia, SA 5005, Australia
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Cristian Capelli
29Department of Zoology, University of Oxford, Oxford, OX1 3PS, UK
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Mark G. Thomas
81Department of Genetics, Evolution and Environment, University College London, WC1E 6BT, UK
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Andres Ruiz-Linares
81Department of Genetics, Evolution and Environment, University College London, WC1E 6BT, UK
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Sarah A. Tishkoff
82Department of Biology and Genetics. University of Pennsylvania, Philadelphia, Pennsylvania, 19104, USA
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Lalji Singh
83CSIR-Centre for Cellular and Molecular Biology, Hyderabad, 500 007, India
84Present address: Banaras Hindu University, Varanasi, 221 005, India
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Kumarasamy Thangaraj
83CSIR-Centre for Cellular and Molecular Biology, Hyderabad, 500 007, India
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Richard Villems
52Estonian Biocentre, Evolutionary Biology group, Tartu, 51010, Estonia
56Department of Evolutionary Biology, University of Tartu, Tartu, 51010, Estonia
85Estonian Academy of Sciences, Tallinn, 10130, Estonia
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David Comas
86Institut de Biologia Evolutiva (CSIC-UPF), Departament de Ciències Experimentals i de la Salut, Universitat Pompeu Fabra, Barcelona, 08003, Spain
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Rem Sukernik
37Laboratory of Human Molecular Genetics, Institute of Molecular and Cellular Biology, Russian Academy of Science, Siberian Branch, Novosibirsk, 630090, Russia
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Mait Metspalu
52Estonian Biocentre, Evolutionary Biology group, Tartu, 51010, Estonia
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Matthias Meyer
4Max Planck Institute for Evolutionary Anthropology, Leipzig, 04103, Germany
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Evan E. Eichler
6Department of Genome Sciences, University of Washington, Seattle, WA, 98195, USA
87Howard Hughes Medical Institute, University of Washington, Seattle, WA, USA 98195
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Joachim Burger
5Johannes Gutenberg University Mainz, Institute of Anthropology, Mainz, D-55128, Germany
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Montgomery Slatkin
7Department of Integrative Biology, University of California, Berkeley, CA, 94720-3140, USA
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Svante Pääbo
4Max Planck Institute for Evolutionary Anthropology, Leipzig, 04103, Germany
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Janet Kelso
4Max Planck Institute for Evolutionary Anthropology, Leipzig, 04103, Germany
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David Reich
1Department of Genetics, Harvard Medical School, Boston, MA, 02115, USA
2Broad Institute of Harvard and MIT, Cambridge, MA, 02142, USA
88Howard Hughes Medical Institute, Harvard Medical School, Boston, MA, 02115, USA
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Johannes Krause
3Institute for Archaeological Sciences, University of Tübingen, Tübingen, 72074, Germany
89Senckenberg Centre for Human Evolution and Palaeoenvironment, University of Tübingen, 72070, Germany
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Abstract

We sequenced genomes from a ∼7,000 year old early farmer from Stuttgart in Germany, an ∼8,000 year old hunter-gatherer from Luxembourg, and seven ∼8,000 year old hunter-gatherers from southern Sweden. We analyzed these data together with other ancient genomes and 2,345 contemporary humans to show that the great majority of present-day Europeans derive from at least three highly differentiated populations: West European Hunter-Gatherers (WHG), who contributed ancestry to all Europeans but not to Near Easterners; Ancient North Eurasians (ANE), who were most closely related to Upper Paleolithic Siberians and contributed to both Europeans and Near Easterners; and Early European Farmers (EEF), who were mainly of Near Eastern origin but also harbored WHG-related ancestry. We model these populations’ deep relationships and show that EEF had ∼44% ancestry from a “Basal Eurasian” lineage that split prior to the diversification of all other non-African lineages.

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Ancient human genomes suggest three ancestral populations for present-day Europeans
Iosif Lazaridis, Nick Patterson, Alissa Mittnik, Gabriel Renaud, Swapan Mallick, Karola Kirsanow, Peter H. Sudmant, Joshua G. Schraiber, Sergi Castellano, Mark Lipson, Bonnie Berger, Christos Economou, Ruth Bollongino, Qiaomei Fu, Kirsten I. Bos, Susanne Nordenfelt, Heng Li, Cesare de Filippo, Kay Prüfer, Susanna Sawyer, Cosimo Posth, Wolfgang Haak, Fredrik Hallgren, Elin Fornander, Nadin Rohland, Dominique Delsate, Michael Francken, Jean-Michel Guinet, Joachim Wahl, George Ayodo, Hamza A. Babiker, Graciela Bailliet, Elena Balanovska, Oleg Balanovsky, Ramiro Barrantes, Gabriel Bedoya, Haim Ben-Ami, Judit Bene, Fouad Berrada, Claudio M. Bravi, Francesca Brisighelli, George Busby, Francesco Cali, Mikhail Churnosov, David E. C. Cole, Daniel Corach, Larissa Damba, George van Driem, Stanislav Dryomov, Jean-Michel Dugoujon, Sardana A. Fedorova, Irene Gallego Romero, Marina Gubina, Michael Hammer, Brenna Henn, Tor Hervig, Ugur Hodoglugil, Aashish R. Jha, Sena Karachanak-Yankova, Rita Khusainova, Elza Khusnutdinova, Rick Kittles, Toomas Kivisild, William Klitz, Vaidutis Kučinskas, Alena Kushniarevich, Leila Laredj, Sergey Litvinov, Theologos Loukidis, Robert W. Mahley, Béla Melegh, Ene Metspalu, Julio Molina, Joanna Mountain, Klemetti Näkkäläjärvi, Desislava Nesheva, Thomas Nyambo, Ludmila Osipova, Jüri Parik, Fedor Platonov, Olga Posukh, Valentino Romano, Francisco Rothhammer, Igor Rudan, Ruslan Ruizbakiev, Hovhannes Sahakyan, Antti Sajantila, Antonio Salas, Elena B. Starikovskaya, Ayele Tarekegn, Draga Toncheva, Shahlo Turdikulova, Ingrida Uktveryte, Olga Utevska, René Vasquez, Mercedes Villena, Mikhail Voevoda, Cheryl Winkler, Levon Yepiskoposyan, Pierre Zalloua, Tatijana Zemunik, Alan Cooper, Cristian Capelli, Mark G. Thomas, Andres Ruiz-Linares, Sarah A. Tishkoff, Lalji Singh, Kumarasamy Thangaraj, Richard Villems, David Comas, Rem Sukernik, Mait Metspalu, Matthias Meyer, Evan E. Eichler, Joachim Burger, Montgomery Slatkin, Svante Pääbo, Janet Kelso, David Reich, Johannes Krause
bioRxiv 001552; doi: https://doi.org/10.1101/001552
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Ancient human genomes suggest three ancestral populations for present-day Europeans
Iosif Lazaridis, Nick Patterson, Alissa Mittnik, Gabriel Renaud, Swapan Mallick, Karola Kirsanow, Peter H. Sudmant, Joshua G. Schraiber, Sergi Castellano, Mark Lipson, Bonnie Berger, Christos Economou, Ruth Bollongino, Qiaomei Fu, Kirsten I. Bos, Susanne Nordenfelt, Heng Li, Cesare de Filippo, Kay Prüfer, Susanna Sawyer, Cosimo Posth, Wolfgang Haak, Fredrik Hallgren, Elin Fornander, Nadin Rohland, Dominique Delsate, Michael Francken, Jean-Michel Guinet, Joachim Wahl, George Ayodo, Hamza A. Babiker, Graciela Bailliet, Elena Balanovska, Oleg Balanovsky, Ramiro Barrantes, Gabriel Bedoya, Haim Ben-Ami, Judit Bene, Fouad Berrada, Claudio M. Bravi, Francesca Brisighelli, George Busby, Francesco Cali, Mikhail Churnosov, David E. C. Cole, Daniel Corach, Larissa Damba, George van Driem, Stanislav Dryomov, Jean-Michel Dugoujon, Sardana A. Fedorova, Irene Gallego Romero, Marina Gubina, Michael Hammer, Brenna Henn, Tor Hervig, Ugur Hodoglugil, Aashish R. Jha, Sena Karachanak-Yankova, Rita Khusainova, Elza Khusnutdinova, Rick Kittles, Toomas Kivisild, William Klitz, Vaidutis Kučinskas, Alena Kushniarevich, Leila Laredj, Sergey Litvinov, Theologos Loukidis, Robert W. Mahley, Béla Melegh, Ene Metspalu, Julio Molina, Joanna Mountain, Klemetti Näkkäläjärvi, Desislava Nesheva, Thomas Nyambo, Ludmila Osipova, Jüri Parik, Fedor Platonov, Olga Posukh, Valentino Romano, Francisco Rothhammer, Igor Rudan, Ruslan Ruizbakiev, Hovhannes Sahakyan, Antti Sajantila, Antonio Salas, Elena B. Starikovskaya, Ayele Tarekegn, Draga Toncheva, Shahlo Turdikulova, Ingrida Uktveryte, Olga Utevska, René Vasquez, Mercedes Villena, Mikhail Voevoda, Cheryl Winkler, Levon Yepiskoposyan, Pierre Zalloua, Tatijana Zemunik, Alan Cooper, Cristian Capelli, Mark G. Thomas, Andres Ruiz-Linares, Sarah A. Tishkoff, Lalji Singh, Kumarasamy Thangaraj, Richard Villems, David Comas, Rem Sukernik, Mait Metspalu, Matthias Meyer, Evan E. Eichler, Joachim Burger, Montgomery Slatkin, Svante Pääbo, Janet Kelso, David Reich, Johannes Krause
bioRxiv 001552; doi: https://doi.org/10.1101/001552

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