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<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">Ancient human genomes suggest three ancestral populations for present-day
Europeans</title>
<author><name sortKey="Lazaridis, Iosif" sort="Lazaridis, Iosif" uniqKey="Lazaridis I" first="Iosif" last="Lazaridis">Iosif Lazaridis</name>
<affiliation><nlm:aff id="A1">Department of Genetics, Harvard Medical School, Boston, MA, 02115, USA</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A2">Broad Institute of Harvard and MIT, Cambridge, MA, 02142, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Patterson, Nick" sort="Patterson, Nick" uniqKey="Patterson N" first="Nick" last="Patterson">Nick Patterson</name>
<affiliation><nlm:aff id="A2">Broad Institute of Harvard and MIT, Cambridge, MA, 02142, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Mittnik, Alissa" sort="Mittnik, Alissa" uniqKey="Mittnik A" first="Alissa" last="Mittnik">Alissa Mittnik</name>
<affiliation><nlm:aff id="A3">Institute for Archaeological Sciences, University of Tübingen, Tübingen, 72074, Germany</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Renaud, Gabriel" sort="Renaud, Gabriel" uniqKey="Renaud G" first="Gabriel" last="Renaud">Gabriel Renaud</name>
<affiliation><nlm:aff id="A4">Max Planck Institute for Evolutionary Anthropology, Leipzig, 04103, Germany</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Mallick, Swapan" sort="Mallick, Swapan" uniqKey="Mallick S" first="Swapan" last="Mallick">Swapan Mallick</name>
<affiliation><nlm:aff id="A1">Department of Genetics, Harvard Medical School, Boston, MA, 02115, USA</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A2">Broad Institute of Harvard and MIT, Cambridge, MA, 02142, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Kirsanow, Karola" sort="Kirsanow, Karola" uniqKey="Kirsanow K" first="Karola" last="Kirsanow">Karola Kirsanow</name>
<affiliation><nlm:aff id="A5">Johannes Gutenberg University Mainz, Institute of Anthropology, Mainz, D-55128, Germany</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Sudmant, Peter H" sort="Sudmant, Peter H" uniqKey="Sudmant P" first="Peter H." last="Sudmant">Peter H. Sudmant</name>
<affiliation><nlm:aff id="A6">Department of Genome Sciences, University of Washington, Seattle, WA, 98195, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Schraiber, Joshua G" sort="Schraiber, Joshua G" uniqKey="Schraiber J" first="Joshua G." last="Schraiber">Joshua G. Schraiber</name>
<affiliation><nlm:aff id="A7">Department of Integrative Biology, University of California, Berkeley, CA, 94720-3140, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Castellano, Sergi" sort="Castellano, Sergi" uniqKey="Castellano S" first="Sergi" last="Castellano">Sergi Castellano</name>
<affiliation><nlm:aff id="A4">Max Planck Institute for Evolutionary Anthropology, Leipzig, 04103, Germany</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Lipson, Mark" sort="Lipson, Mark" uniqKey="Lipson M" first="Mark" last="Lipson">Mark Lipson</name>
<affiliation><nlm:aff id="A8">Department of Mathematics and Computer Science and Artificial Intelligence Laboratory, Massachusetts Institute of Technology, Cambridge, MA 02139, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Berger, Bonnie" sort="Berger, Bonnie" uniqKey="Berger B" first="Bonnie" last="Berger">Bonnie Berger</name>
<affiliation><nlm:aff id="A2">Broad Institute of Harvard and MIT, Cambridge, MA, 02142, USA</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A8">Department of Mathematics and Computer Science and Artificial Intelligence Laboratory, Massachusetts Institute of Technology, Cambridge, MA 02139, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Economou, Christos" sort="Economou, Christos" uniqKey="Economou C" first="Christos" last="Economou">Christos Economou</name>
<affiliation><nlm:aff id="A9">Archaeological Research Laboratory, Stockholm University, 114 18, Sweden</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Bollongino, Ruth" sort="Bollongino, Ruth" uniqKey="Bollongino R" first="Ruth" last="Bollongino">Ruth Bollongino</name>
<affiliation><nlm:aff id="A5">Johannes Gutenberg University Mainz, Institute of Anthropology, Mainz, D-55128, Germany</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Fu, Qiaomei" sort="Fu, Qiaomei" uniqKey="Fu Q" first="Qiaomei" last="Fu">Qiaomei Fu</name>
<affiliation><nlm:aff id="A1">Department of Genetics, Harvard Medical School, Boston, MA, 02115, USA</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A4">Max Planck Institute for Evolutionary Anthropology, Leipzig, 04103, Germany</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A10">Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, IVPP, CAS, Beijing, 100049, China</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Bos, Kirsten I" sort="Bos, Kirsten I" uniqKey="Bos K" first="Kirsten I." last="Bos">Kirsten I. Bos</name>
<affiliation><nlm:aff id="A3">Institute for Archaeological Sciences, University of Tübingen, Tübingen, 72074, Germany</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Nordenfelt, Susanne" sort="Nordenfelt, Susanne" uniqKey="Nordenfelt S" first="Susanne" last="Nordenfelt">Susanne Nordenfelt</name>
<affiliation><nlm:aff id="A1">Department of Genetics, Harvard Medical School, Boston, MA, 02115, USA</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A2">Broad Institute of Harvard and MIT, Cambridge, MA, 02142, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Li, Heng" sort="Li, Heng" uniqKey="Li H" first="Heng" last="Li">Heng Li</name>
<affiliation><nlm:aff id="A1">Department of Genetics, Harvard Medical School, Boston, MA, 02115, USA</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A2">Broad Institute of Harvard and MIT, Cambridge, MA, 02142, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="De Filippo, Cesare" sort="De Filippo, Cesare" uniqKey="De Filippo C" first="Cesare" last="De Filippo">Cesare De Filippo</name>
<affiliation><nlm:aff id="A4">Max Planck Institute for Evolutionary Anthropology, Leipzig, 04103, Germany</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Prufer, Kay" sort="Prufer, Kay" uniqKey="Prufer K" first="Kay" last="Prüfer">Kay Prüfer</name>
<affiliation><nlm:aff id="A4">Max Planck Institute for Evolutionary Anthropology, Leipzig, 04103, Germany</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Sawyer, Susanna" sort="Sawyer, Susanna" uniqKey="Sawyer S" first="Susanna" last="Sawyer">Susanna Sawyer</name>
<affiliation><nlm:aff id="A4">Max Planck Institute for Evolutionary Anthropology, Leipzig, 04103, Germany</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Posth, Cosimo" sort="Posth, Cosimo" uniqKey="Posth C" first="Cosimo" last="Posth">Cosimo Posth</name>
<affiliation><nlm:aff id="A3">Institute for Archaeological Sciences, University of Tübingen, Tübingen, 72074, Germany</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Haak, Wolfgang" sort="Haak, Wolfgang" uniqKey="Haak W" first="Wolfgang" last="Haak">Wolfgang Haak</name>
<affiliation><nlm:aff id="A11">Australian Centre for Ancient DNA, School of Earth and Environmental Sciences, University of Adelaide, Adelaide, South Australia, SA 5005, Australia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Hallgren, Fredrik" sort="Hallgren, Fredrik" uniqKey="Hallgren F" first="Fredrik" last="Hallgren">Fredrik Hallgren</name>
<affiliation><nlm:aff id="A12">The Cultural Heritage Foundation, Västerås, 722 12, Sweden</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Fornander, Elin" sort="Fornander, Elin" uniqKey="Fornander E" first="Elin" last="Fornander">Elin Fornander</name>
<affiliation><nlm:aff id="A12">The Cultural Heritage Foundation, Västerås, 722 12, Sweden</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Rohland, Nadin" sort="Rohland, Nadin" uniqKey="Rohland N" first="Nadin" last="Rohland">Nadin Rohland</name>
<affiliation><nlm:aff id="A1">Department of Genetics, Harvard Medical School, Boston, MA, 02115, USA</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A2">Broad Institute of Harvard and MIT, Cambridge, MA, 02142, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Delsate, Dominique" sort="Delsate, Dominique" uniqKey="Delsate D" first="Dominique" last="Delsate">Dominique Delsate</name>
<affiliation><nlm:aff id="A13">National Museum of Natural History, L-2160, Luxembourg</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A14">National Center of Archaeological Research, National Museum of History and Art, L-2345, Luxembourg</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Francken, Michael" sort="Francken, Michael" uniqKey="Francken M" first="Michael" last="Francken">Michael Francken</name>
<affiliation><nlm:aff id="A15">Department of Paleoanthropology, Senckenberg Center for Human Evolution and Paleoenvironment, University of Tübingen, Tübingen, D-72070, Germany</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Guinet, Jean Michel" sort="Guinet, Jean Michel" uniqKey="Guinet J" first="Jean-Michel" last="Guinet">Jean-Michel Guinet</name>
<affiliation><nlm:aff id="A13">National Museum of Natural History, L-2160, Luxembourg</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Wahl, Joachim" sort="Wahl, Joachim" uniqKey="Wahl J" first="Joachim" last="Wahl">Joachim Wahl</name>
<affiliation><nlm:aff id="A16">State Office for Cultural Heritage Management Baden-Württemberg, Osteology, Konstanz, D-78467, Germany</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Ayodo, George" sort="Ayodo, George" uniqKey="Ayodo G" first="George" last="Ayodo">George Ayodo</name>
<affiliation><nlm:aff id="A17">Center for Global Health and Child Development, Kisumu, 40100, Kenya</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Babiker, Hamza A" sort="Babiker, Hamza A" uniqKey="Babiker H" first="Hamza A." last="Babiker">Hamza A. Babiker</name>
<affiliation><nlm:aff id="A18">Institutes of Evolution, Immunology and Infection Research, School of Biological Sciences, University of Edinburgh, Edinburgh, EH9 3JT, UK</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A19">Biochemistry Department, Faculty of Medicine, Sultan Qaboos University, Alkhod, Muscat, 123, Oman</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Bailliet, Graciela" sort="Bailliet, Graciela" uniqKey="Bailliet G" first="Graciela" last="Bailliet">Graciela Bailliet</name>
<affiliation><nlm:aff id="A20">Laboratorio de Genética Molecular Poblacional, Instituto Multidisciplinario de Biología Celular (IMBICE), CCT-CONICET & CICPBA, La Plata, B1906APO, Argentina</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Balanovska, Elena" sort="Balanovska, Elena" uniqKey="Balanovska E" first="Elena" last="Balanovska">Elena Balanovska</name>
<affiliation><nlm:aff id="A21">Research Centre for Medical Genetics, Moscow, 115478, Russia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Balanovsky, Oleg" sort="Balanovsky, Oleg" uniqKey="Balanovsky O" first="Oleg" last="Balanovsky">Oleg Balanovsky</name>
<affiliation><nlm:aff id="A21">Research Centre for Medical Genetics, Moscow, 115478, Russia</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A22">Vavilov Institute for General Genetics, Moscow, 119991, Russia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Barrantes, Ramiro" sort="Barrantes, Ramiro" uniqKey="Barrantes R" first="Ramiro" last="Barrantes">Ramiro Barrantes</name>
<affiliation><nlm:aff id="A23">Escuela de Biología, Universidad de Costa Rica, San José, 2060, Costa Rica</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Bedoya, Gabriel" sort="Bedoya, Gabriel" uniqKey="Bedoya G" first="Gabriel" last="Bedoya">Gabriel Bedoya</name>
<affiliation><nlm:aff id="A24">Institute of Biology, Research group GENMOL, Universidad de Antioquia, Medellín, Colombia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Ben Ami, Haim" sort="Ben Ami, Haim" uniqKey="Ben Ami H" first="Haim" last="Ben-Ami">Haim Ben-Ami</name>
<affiliation><nlm:aff id="A25">Rambam Health Care Campus, Haifa, 31096, Israel</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Bene, Judit" sort="Bene, Judit" uniqKey="Bene J" first="Judit" last="Bene">Judit Bene</name>
<affiliation><nlm:aff id="A26">Department of Medical Genetics and Szentagothai Research Center, University of Pécs, Pécs, H-7624 Hungary</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Berrada, Fouad" sort="Berrada, Fouad" uniqKey="Berrada F" first="Fouad" last="Berrada">Fouad Berrada</name>
<affiliation><nlm:aff id="A27">Al Akhawayn University in Ifrane (AUI), School of Science and Engineering, Ifrane, 53000, Morocco</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Bravi, Claudio M" sort="Bravi, Claudio M" uniqKey="Bravi C" first="Claudio M." last="Bravi">Claudio M. Bravi</name>
<affiliation><nlm:aff id="A20">Laboratorio de Genética Molecular Poblacional, Instituto Multidisciplinario de Biología Celular (IMBICE), CCT-CONICET & CICPBA, La Plata, B1906APO, Argentina</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Brisighelli, Francesca" sort="Brisighelli, Francesca" uniqKey="Brisighelli F" first="Francesca" last="Brisighelli">Francesca Brisighelli</name>
<affiliation><nlm:aff id="A28">Forensic Genetics Laboratory, Institute of Legal Medicine, Università Cattolica del Sacro Cuore, Rome, 00168, Italy</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Busby, George B J" sort="Busby, George B J" uniqKey="Busby G" first="George B. J." last="Busby">George B. J. Busby</name>
<affiliation><nlm:aff id="A29">Department of Zoology, University of Oxford, Oxford, OX1 3PS, UK</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A30">Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford, OX3 7BN, UK</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Cali, Francesco" sort="Cali, Francesco" uniqKey="Cali F" first="Francesco" last="Cali">Francesco Cali</name>
<affiliation><nlm:aff id="A31">Laboratorio di Genetica Molecolare, IRCCS Associazione Oasi Maria SS, Troina, 94018, Italy</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Churnosov, Mikhail" sort="Churnosov, Mikhail" uniqKey="Churnosov M" first="Mikhail" last="Churnosov">Mikhail Churnosov</name>
<affiliation><nlm:aff id="A32">Belgorod State University, Belgorod, 308015, Russia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Cole, David E C" sort="Cole, David E C" uniqKey="Cole D" first="David E. C." last="Cole">David E. C. Cole</name>
<affiliation><nlm:aff id="A33">Department of Laboratory Medicine and Pathobiology, University of Toronto, Toronto, Ontario, M5G 1L5, Canada</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Corach, Daniel" sort="Corach, Daniel" uniqKey="Corach D" first="Daniel" last="Corach">Daniel Corach</name>
<affiliation><nlm:aff id="A34">Servicio de Huellas Digitales Genéticas, School of Pharmacy and Biochemistry, Universidad de Buenos Aires, 1113 CABA, Argentina</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Damba, Larissa" sort="Damba, Larissa" uniqKey="Damba L" first="Larissa" last="Damba">Larissa Damba</name>
<affiliation><nlm:aff id="A35">Institute of Cytology and Genetics, Siberian Branch of Russian Academy of Sciences, Novosibirsk, 630090, Russia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Van Driem, George" sort="Van Driem, George" uniqKey="Van Driem G" first="George" last="Van Driem">George Van Driem</name>
<affiliation><nlm:aff id="A36">Institute of Linguistics, University of Bern, Bern, CH-3012, Switzerland</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Dryomov, Stanislav" sort="Dryomov, Stanislav" uniqKey="Dryomov S" first="Stanislav" last="Dryomov">Stanislav Dryomov</name>
<affiliation><nlm:aff id="A37">Laboratory of Human Molecular Genetics, Institute of Molecular and Cellular Biology, Russian Academy of Science, Siberian Branch, Novosibirsk, 630090, Russia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Dugoujon, Jean Michel" sort="Dugoujon, Jean Michel" uniqKey="Dugoujon J" first="Jean-Michel" last="Dugoujon">Jean-Michel Dugoujon</name>
<affiliation><nlm:aff id="A38">Anthropologie Moléculaire et Imagerie de Synthèse, CNRS UMR 5288, Université Paul Sabatier Toulouse III, Toulouse, 31000, France</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Fedorova, Sardana A" sort="Fedorova, Sardana A" uniqKey="Fedorova S" first="Sardana A." last="Fedorova">Sardana A. Fedorova</name>
<affiliation><nlm:aff id="A39">Yakut Research Center of Complex Medical Problems and North-Eastern Federal University, Yakutsk, 677010, Russia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Romero, Irene Gallego" sort="Romero, Irene Gallego" uniqKey="Romero I" first="Irene Gallego" last="Romero">Irene Gallego Romero</name>
<affiliation><nlm:aff id="A40">Department of Human Genetics, University of Chicago, Chicago, IL, 60637, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Gubina, Marina" sort="Gubina, Marina" uniqKey="Gubina M" first="Marina" last="Gubina">Marina Gubina</name>
<affiliation><nlm:aff id="A35">Institute of Cytology and Genetics, Siberian Branch of Russian Academy of Sciences, Novosibirsk, 630090, Russia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Hammer, Michael" sort="Hammer, Michael" uniqKey="Hammer M" first="Michael" last="Hammer">Michael Hammer</name>
<affiliation><nlm:aff id="A41">ARL Division of Biotechnology, University of Arizona, Tucson, AZ, 85721, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Henn, Brenna M" sort="Henn, Brenna M" uniqKey="Henn B" first="Brenna M." last="Henn">Brenna M. Henn</name>
<affiliation><nlm:aff id="A42">Department of Ecology and Evolution, Stony Brook University, Stony Brook, NY, 11794, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Hervig, Tor" sort="Hervig, Tor" uniqKey="Hervig T" first="Tor" last="Hervig">Tor Hervig</name>
<affiliation><nlm:aff id="A43">Department of Clinical Science, University of Bergen, Bergen, 5021, Norway</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Hodoglugil, Ugur" sort="Hodoglugil, Ugur" uniqKey="Hodoglugil U" first="Ugur" last="Hodoglugil">Ugur Hodoglugil</name>
<affiliation><nlm:aff id="A44">NextBio, part of Illumina, Santa Clara, CA, USA 95050</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Jha, Aashish R" sort="Jha, Aashish R" uniqKey="Jha A" first="Aashish R." last="Jha">Aashish R. Jha</name>
<affiliation><nlm:aff id="A40">Department of Human Genetics, University of Chicago, Chicago, IL, 60637, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Karachanak Yankova, Sena" sort="Karachanak Yankova, Sena" uniqKey="Karachanak Yankova S" first="Sena" last="Karachanak-Yankova">Sena Karachanak-Yankova</name>
<affiliation><nlm:aff id="A45">Dept. of Medical Genetics, National Human Genome Center, Medical University Sofia, Sofia, 1431, Bulgaria</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Khusainova, Rita" sort="Khusainova, Rita" uniqKey="Khusainova R" first="Rita" last="Khusainova">Rita Khusainova</name>
<affiliation><nlm:aff id="A46">Institute of Biochemistry and Genetics, Ufa Research Centre, Russian Academy of Sciences, Ufa, 450054, Russia</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A47">Department of Genetics and Fundamental Medicine, Bashkir State University, Ufa, 450074, Russia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Khusnutdinova, Elza" sort="Khusnutdinova, Elza" uniqKey="Khusnutdinova E" first="Elza" last="Khusnutdinova">Elza Khusnutdinova</name>
<affiliation><nlm:aff id="A46">Institute of Biochemistry and Genetics, Ufa Research Centre, Russian Academy of Sciences, Ufa, 450054, Russia</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A47">Department of Genetics and Fundamental Medicine, Bashkir State University, Ufa, 450074, Russia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Kittles, Rick" sort="Kittles, Rick" uniqKey="Kittles R" first="Rick" last="Kittles">Rick Kittles</name>
<affiliation><nlm:aff id="A48">College of Medicine, University of Illinois at Chicago, Chicago, IL, 60607, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Kivisild, Toomas" sort="Kivisild, Toomas" uniqKey="Kivisild T" first="Toomas" last="Kivisild">Toomas Kivisild</name>
<affiliation><nlm:aff id="A49">Division of Biological Anthropology, University of Cambridge, Cambridge, United Kingdom CB2 1QH</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Klitz, William" sort="Klitz, William" uniqKey="Klitz W" first="William" last="Klitz">William Klitz</name>
<affiliation><nlm:aff id="A7">Department of Integrative Biology, University of California, Berkeley, CA, 94720-3140, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Ku Inskas, Vaidutis" sort="Ku Inskas, Vaidutis" uniqKey="Ku Inskas V" first="Vaidutis" last="Ku Inskas">Vaidutis Ku Inskas</name>
<affiliation><nlm:aff id="A50">Department of Human and Medical Genetics, Vilnius University, Vilnius, LT-08661, Lithuania</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Kushniarevich, Alena" sort="Kushniarevich, Alena" uniqKey="Kushniarevich A" first="Alena" last="Kushniarevich">Alena Kushniarevich</name>
<affiliation><nlm:aff id="A51">Estonian Biocentre, Evolutionary Biology group, Tartu, 51010, Estonia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Laredj, Leila" sort="Laredj, Leila" uniqKey="Laredj L" first="Leila" last="Laredj">Leila Laredj</name>
<affiliation><nlm:aff id="A52">Translational Medicine and Neurogenetics, Institut de Génétique et de Biologie Moléculaire et Cellulaire, Illkirch, 67404, France</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Litvinov, Sergey" sort="Litvinov, Sergey" uniqKey="Litvinov S" first="Sergey" last="Litvinov">Sergey Litvinov</name>
<affiliation><nlm:aff id="A46">Institute of Biochemistry and Genetics, Ufa Research Centre, Russian Academy of Sciences, Ufa, 450054, Russia</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A47">Department of Genetics and Fundamental Medicine, Bashkir State University, Ufa, 450074, Russia</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A51">Estonian Biocentre, Evolutionary Biology group, Tartu, 51010, Estonia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Loukidis, Theologos" sort="Loukidis, Theologos" uniqKey="Loukidis T" first="Theologos" last="Loukidis">Theologos Loukidis</name>
</author>
<author><name sortKey="Mahley, Robert W" sort="Mahley, Robert W" uniqKey="Mahley R" first="Robert W." last="Mahley">Robert W. Mahley</name>
<affiliation><nlm:aff id="A53">Gladstone Institutes, San Francisco, CA, 94158, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Melegh, Bela" sort="Melegh, Bela" uniqKey="Melegh B" first="Béla" last="Melegh">Béla Melegh</name>
<affiliation><nlm:aff id="A26">Department of Medical Genetics and Szentagothai Research Center, University of Pécs, Pécs, H-7624 Hungary</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Metspalu, Ene" sort="Metspalu, Ene" uniqKey="Metspalu E" first="Ene" last="Metspalu">Ene Metspalu</name>
<affiliation><nlm:aff id="A54">Department of Evolutionary Biology, University of Tartu, Tartu, 51010, Estonia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Molina, Julio" sort="Molina, Julio" uniqKey="Molina J" first="Julio" last="Molina">Julio Molina</name>
<affiliation><nlm:aff id="A55">Centro de Investigaciones Biomédicas de Guatemala, Ciudad de Guatemala, Guatemala</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Mountain, Joanna" sort="Mountain, Joanna" uniqKey="Mountain J" first="Joanna" last="Mountain">Joanna Mountain</name>
<affiliation><nlm:aff id="A56">Research Department, 23andMe, Inc. Mountain View, CA, 94043, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="N Kk L J Rvi, Klemetti" sort="N Kk L J Rvi, Klemetti" uniqKey="N Kk L J Rvi K" first="Klemetti" last="N Kk L J Rvi">Klemetti N Kk L J Rvi</name>
<affiliation><nlm:aff id="A57">Cultural Anthropology Program, University of Oulu, Oulu, 90014, Finland</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Nesheva, Desislava" sort="Nesheva, Desislava" uniqKey="Nesheva D" first="Desislava" last="Nesheva">Desislava Nesheva</name>
<affiliation><nlm:aff id="A45">Dept. of Medical Genetics, National Human Genome Center, Medical University Sofia, Sofia, 1431, Bulgaria</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Nyambo, Thomas" sort="Nyambo, Thomas" uniqKey="Nyambo T" first="Thomas" last="Nyambo">Thomas Nyambo</name>
<affiliation><nlm:aff id="A58">Department of Biochemistry, Muhimbili University of Health and Allied Sciences, Dar es Salaam, Tanzania</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Osipova, Ludmila" sort="Osipova, Ludmila" uniqKey="Osipova L" first="Ludmila" last="Osipova">Ludmila Osipova</name>
<affiliation><nlm:aff id="A35">Institute of Cytology and Genetics, Siberian Branch of Russian Academy of Sciences, Novosibirsk, 630090, Russia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Parik, Juri" sort="Parik, Juri" uniqKey="Parik J" first="Jüri" last="Parik">Jüri Parik</name>
<affiliation><nlm:aff id="A54">Department of Evolutionary Biology, University of Tartu, Tartu, 51010, Estonia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Platonov, Fedor" sort="Platonov, Fedor" uniqKey="Platonov F" first="Fedor" last="Platonov">Fedor Platonov</name>
<affiliation><nlm:aff id="A59">Research Institute of Health, North-Eastern Federal University, Yakutsk, 677000, Russia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Posukh, Olga" sort="Posukh, Olga" uniqKey="Posukh O" first="Olga" last="Posukh">Olga Posukh</name>
<affiliation><nlm:aff id="A35">Institute of Cytology and Genetics, Siberian Branch of Russian Academy of Sciences, Novosibirsk, 630090, Russia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Romano, Valentino" sort="Romano, Valentino" uniqKey="Romano V" first="Valentino" last="Romano">Valentino Romano</name>
<affiliation><nlm:aff id="A60">Dipartimento di Fisica e Chimica, Università di Palermo, Palermo, 90128, Italy</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Rothhammer, Francisco" sort="Rothhammer, Francisco" uniqKey="Rothhammer F" first="Francisco" last="Rothhammer">Francisco Rothhammer</name>
<affiliation><nlm:aff id="A61">Instituto de Alta Investigación, Universidad de Tarapacá, Arica, Chile</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A62">Programa de Genética Humana ICBM Facultad de Medicina Universidad de Chile, Santiago, Chile</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A63">Centro de Investigaciones del Hombre en el Desierto, Arica, Chile</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Rudan, Igor" sort="Rudan, Igor" uniqKey="Rudan I" first="Igor" last="Rudan">Igor Rudan</name>
<affiliation><nlm:aff id="A64">Centre for Population Health Sciences, The University of Edinburgh Medical School, Edinburgh, Scotland, EH8 9AG, UK</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Ruizbakiev, Ruslan" sort="Ruizbakiev, Ruslan" uniqKey="Ruizbakiev R" first="Ruslan" last="Ruizbakiev">Ruslan Ruizbakiev</name>
<affiliation><nlm:aff id="A65">Deceased: formerly of the Institute of Immunology, Academy of Science, Tashkent, 70000, Uzbekistan</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Sahakyan, Hovhannes" sort="Sahakyan, Hovhannes" uniqKey="Sahakyan H" first="Hovhannes" last="Sahakyan">Hovhannes Sahakyan</name>
<affiliation><nlm:aff id="A51">Estonian Biocentre, Evolutionary Biology group, Tartu, 51010, Estonia</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A66">Laboratory of Ethnogenomics, Institute of Molecular Biology, National Academy of Sciences of Armenia, Yerevan, 0014, Armenia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Sajantila, Antti" sort="Sajantila, Antti" uniqKey="Sajantila A" first="Antti" last="Sajantila">Antti Sajantila</name>
<affiliation><nlm:aff id="A67">Department of Forensic Medicine, Hjelt Institute, University of Helsinki, Helsinki, 00014, Finland</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A68">Institute of Applied Genetics, Department of Molecular and Medical Genetics, University of North Texas Health Science Center, Fort Worth, Texas 76107, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Salas, Antonio" sort="Salas, Antonio" uniqKey="Salas A" first="Antonio" last="Salas">Antonio Salas</name>
<affiliation><nlm:aff id="A69">Unidade 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</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Starikovskaya, Elena B" sort="Starikovskaya, Elena B" uniqKey="Starikovskaya E" first="Elena B." last="Starikovskaya">Elena B. Starikovskaya</name>
<affiliation><nlm:aff id="A37">Laboratory of Human Molecular Genetics, Institute of Molecular and Cellular Biology, Russian Academy of Science, Siberian Branch, Novosibirsk, 630090, Russia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Tarekegn, Ayele" sort="Tarekegn, Ayele" uniqKey="Tarekegn A" first="Ayele" last="Tarekegn">Ayele Tarekegn</name>
<affiliation><nlm:aff id="A70">Research Fellow, Henry Stewart Group, Russell House, London WC1A 2HN, UK</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Toncheva, Draga" sort="Toncheva, Draga" uniqKey="Toncheva D" first="Draga" last="Toncheva">Draga Toncheva</name>
<affiliation><nlm:aff id="A45">Dept. of Medical Genetics, National Human Genome Center, Medical University Sofia, Sofia, 1431, Bulgaria</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Turdikulova, Shahlo" sort="Turdikulova, Shahlo" uniqKey="Turdikulova S" first="Shahlo" last="Turdikulova">Shahlo Turdikulova</name>
<affiliation><nlm:aff id="A71">Institute of Bioorganic Chemistry Academy of Sciences Republic of Uzbekistan, Tashkent, 100125, Uzbekistan</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Uktveryte, Ingrida" sort="Uktveryte, Ingrida" uniqKey="Uktveryte I" first="Ingrida" last="Uktveryte">Ingrida Uktveryte</name>
<affiliation><nlm:aff id="A50">Department of Human and Medical Genetics, Vilnius University, Vilnius, LT-08661, Lithuania</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Utevska, Olga" sort="Utevska, Olga" uniqKey="Utevska O" first="Olga" last="Utevska">Olga Utevska</name>
<affiliation><nlm:aff id="A72">Department of Genetics and Cytology, V.N. Karazin Kharkiv National University, Kharkiv, 61077, Ukraine</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Vasquez, Rene" sort="Vasquez, Rene" uniqKey="Vasquez R" first="René" last="Vasquez">René Vasquez</name>
<affiliation><nlm:aff id="A73">Instituto Boliviano de Biología de la Altura, Universidad Autonoma Tomás Frías, Potosí, Bolivia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Villena, Mercedes" sort="Villena, Mercedes" uniqKey="Villena M" first="Mercedes" last="Villena">Mercedes Villena</name>
<affiliation><nlm:aff id="A73">Instituto Boliviano de Biología de la Altura, Universidad Autonoma Tomás Frías, Potosí, Bolivia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Voevoda, Mikhail" sort="Voevoda, Mikhail" uniqKey="Voevoda M" first="Mikhail" last="Voevoda">Mikhail Voevoda</name>
<affiliation><nlm:aff id="A35">Institute of Cytology and Genetics, Siberian Branch of Russian Academy of Sciences, Novosibirsk, 630090, Russia</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A74">Inst. of Internal Medicine, Siberian Branch of Russian Acad. of Medical Sciences, Novosibirsk, 630089, Russia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Winkler, Cheryl" sort="Winkler, Cheryl" uniqKey="Winkler C" first="Cheryl" last="Winkler">Cheryl Winkler</name>
<affiliation><nlm:aff id="A75">Basic Research Laboratory, NCI, NIH, Frederick National Laboratory, Leidos Biomedical, Inc., Frederick, MD 21702, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Yepiskoposyan, Levon" sort="Yepiskoposyan, Levon" uniqKey="Yepiskoposyan L" first="Levon" last="Yepiskoposyan">Levon Yepiskoposyan</name>
<affiliation><nlm:aff id="A66">Laboratory of Ethnogenomics, Institute of Molecular Biology, National Academy of Sciences of Armenia, Yerevan, 0014, Armenia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Zalloua, Pierre" sort="Zalloua, Pierre" uniqKey="Zalloua P" first="Pierre" last="Zalloua">Pierre Zalloua</name>
<affiliation><nlm:aff id="A76">Lebanese American University, School of Medicine, Beirut, 13-5053, Lebanon</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A77">Harvard School of Public Health, Boston, 02115, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Zemunik, Tatijana" sort="Zemunik, Tatijana" uniqKey="Zemunik T" first="Tatijana" last="Zemunik">Tatijana Zemunik</name>
<affiliation><nlm:aff id="A78">Department of Medical Biology, University of Split, School of Medicine, Split, 21000, Croatia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Cooper, Alan" sort="Cooper, Alan" uniqKey="Cooper A" first="Alan" last="Cooper">Alan Cooper</name>
<affiliation><nlm:aff id="A11">Australian Centre for Ancient DNA, School of Earth and Environmental Sciences, University of Adelaide, Adelaide, South Australia, SA 5005, Australia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Capelli, Cristian" sort="Capelli, Cristian" uniqKey="Capelli C" first="Cristian" last="Capelli">Cristian Capelli</name>
<affiliation><nlm:aff id="A29">Department of Zoology, University of Oxford, Oxford, OX1 3PS, UK</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Thomas, Mark G" sort="Thomas, Mark G" uniqKey="Thomas M" first="Mark G." last="Thomas">Mark G. Thomas</name>
<affiliation><nlm:aff id="A79">Department of Genetics, Evolution and Environment, University College London, WC1E 6BT, UK</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Ruiz Linares, Andres" sort="Ruiz Linares, Andres" uniqKey="Ruiz Linares A" first="Andres" last="Ruiz-Linares">Andres Ruiz-Linares</name>
<affiliation><nlm:aff id="A79">Department of Genetics, Evolution and Environment, University College London, WC1E 6BT, UK</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Tishkoff, Sarah A" sort="Tishkoff, Sarah A" uniqKey="Tishkoff S" first="Sarah A." last="Tishkoff">Sarah A. Tishkoff</name>
<affiliation><nlm:aff id="A80">Department of Biology and Genetics. University of Pennsylvania, Philadelphia, Pennsylvania, 19104, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Singh, Lalji" sort="Singh, Lalji" uniqKey="Singh L" first="Lalji" last="Singh">Lalji Singh</name>
<affiliation><nlm:aff id="A81">CSIR-Centre for Cellular and Molecular Biology, Hyderabad, 500 007, India</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Thangaraj, Kumarasamy" sort="Thangaraj, Kumarasamy" uniqKey="Thangaraj K" first="Kumarasamy" last="Thangaraj">Kumarasamy Thangaraj</name>
<affiliation><nlm:aff id="A81">CSIR-Centre for Cellular and Molecular Biology, Hyderabad, 500 007, India</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Villems, Richard" sort="Villems, Richard" uniqKey="Villems R" first="Richard" last="Villems">Richard Villems</name>
<affiliation><nlm:aff id="A51">Estonian Biocentre, Evolutionary Biology group, Tartu, 51010, Estonia</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A54">Department of Evolutionary Biology, University of Tartu, Tartu, 51010, Estonia</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A82">Estonian Academy of Sciences, Tallinn, 10130, Estonia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Comas, David" sort="Comas, David" uniqKey="Comas D" first="David" last="Comas">David Comas</name>
<affiliation><nlm:aff id="A83">Institut de Biologia Evolutiva (CSIC-UPF), Departament de Ciències Experimentals i de la Salut, Universitat Pompeu Fabra, Barcelona, 08003, Spain</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Sukernik, Rem" sort="Sukernik, Rem" uniqKey="Sukernik R" first="Rem" last="Sukernik">Rem Sukernik</name>
<affiliation><nlm:aff id="A37">Laboratory of Human Molecular Genetics, Institute of Molecular and Cellular Biology, Russian Academy of Science, Siberian Branch, Novosibirsk, 630090, Russia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Metspalu, Mait" sort="Metspalu, Mait" uniqKey="Metspalu M" first="Mait" last="Metspalu">Mait Metspalu</name>
<affiliation><nlm:aff id="A51">Estonian Biocentre, Evolutionary Biology group, Tartu, 51010, Estonia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Meyer, Matthias" sort="Meyer, Matthias" uniqKey="Meyer M" first="Matthias" last="Meyer">Matthias Meyer</name>
<affiliation><nlm:aff id="A4">Max Planck Institute for Evolutionary Anthropology, Leipzig, 04103, Germany</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Eichler, Evan E" sort="Eichler, Evan E" uniqKey="Eichler E" first="Evan E." last="Eichler">Evan E. Eichler</name>
<affiliation><nlm:aff id="A6">Department of Genome Sciences, University of Washington, Seattle, WA, 98195, USA</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A84">Howard Hughes Medical Institute, University of Washington, Seattle, WA, USA 98195</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Burger, Joachim" sort="Burger, Joachim" uniqKey="Burger J" first="Joachim" last="Burger">Joachim Burger</name>
<affiliation><nlm:aff id="A5">Johannes Gutenberg University Mainz, Institute of Anthropology, Mainz, D-55128, Germany</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Slatkin, Montgomery" sort="Slatkin, Montgomery" uniqKey="Slatkin M" first="Montgomery" last="Slatkin">Montgomery Slatkin</name>
<affiliation><nlm:aff id="A7">Department of Integrative Biology, University of California, Berkeley, CA, 94720-3140, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="P Bo, Svante" sort="P Bo, Svante" uniqKey="P Bo S" first="Svante" last="P Bo">Svante P Bo</name>
<affiliation><nlm:aff id="A4">Max Planck Institute for Evolutionary Anthropology, Leipzig, 04103, Germany</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Kelso, Janet" sort="Kelso, Janet" uniqKey="Kelso J" first="Janet" last="Kelso">Janet Kelso</name>
<affiliation><nlm:aff id="A4">Max Planck Institute for Evolutionary Anthropology, Leipzig, 04103, Germany</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Reich, David" sort="Reich, David" uniqKey="Reich D" first="David" last="Reich">David Reich</name>
<affiliation><nlm:aff id="A1">Department of Genetics, Harvard Medical School, Boston, MA, 02115, USA</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A2">Broad Institute of Harvard and MIT, Cambridge, MA, 02142, USA</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A85">Howard Hughes Medical Institute, Harvard Medical School, Boston, MA, 02115, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Krause, Johannes" sort="Krause, Johannes" uniqKey="Krause J" first="Johannes" last="Krause">Johannes Krause</name>
<affiliation><nlm:aff id="A3">Institute for Archaeological Sciences, University of Tübingen, Tübingen, 72074, Germany</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A86">Senckenberg Centre for Human Evolution and Palaeoenvironment, University of Tübingen, 72070, Germany</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A87">Max Planck Institut für Geschichte und Naturwissenschaften, Jena, 07745, Germany</nlm:aff>
</affiliation>
</author>
</titleStmt>
<publicationStmt><idno type="wicri:source">PMC</idno>
<idno type="pmid">25230663</idno>
<idno type="pmc">4170574</idno>
<idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4170574</idno>
<idno type="RBID">PMC:4170574</idno>
<idno type="doi">10.1038/nature13673</idno>
<date when="2014">2014</date>
<idno type="wicri:Area/Pmc/Corpus">002D67</idno>
<idno type="wicri:explorRef" wicri:stream="Pmc" wicri:step="Corpus" wicri:corpus="PMC">002D67</idno>
</publicationStmt>
<sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">Ancient human genomes suggest three ancestral populations for present-day
Europeans</title>
<author><name sortKey="Lazaridis, Iosif" sort="Lazaridis, Iosif" uniqKey="Lazaridis I" first="Iosif" last="Lazaridis">Iosif Lazaridis</name>
<affiliation><nlm:aff id="A1">Department of Genetics, Harvard Medical School, Boston, MA, 02115, USA</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A2">Broad Institute of Harvard and MIT, Cambridge, MA, 02142, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Patterson, Nick" sort="Patterson, Nick" uniqKey="Patterson N" first="Nick" last="Patterson">Nick Patterson</name>
<affiliation><nlm:aff id="A2">Broad Institute of Harvard and MIT, Cambridge, MA, 02142, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Mittnik, Alissa" sort="Mittnik, Alissa" uniqKey="Mittnik A" first="Alissa" last="Mittnik">Alissa Mittnik</name>
<affiliation><nlm:aff id="A3">Institute for Archaeological Sciences, University of Tübingen, Tübingen, 72074, Germany</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Renaud, Gabriel" sort="Renaud, Gabriel" uniqKey="Renaud G" first="Gabriel" last="Renaud">Gabriel Renaud</name>
<affiliation><nlm:aff id="A4">Max Planck Institute for Evolutionary Anthropology, Leipzig, 04103, Germany</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Mallick, Swapan" sort="Mallick, Swapan" uniqKey="Mallick S" first="Swapan" last="Mallick">Swapan Mallick</name>
<affiliation><nlm:aff id="A1">Department of Genetics, Harvard Medical School, Boston, MA, 02115, USA</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A2">Broad Institute of Harvard and MIT, Cambridge, MA, 02142, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Kirsanow, Karola" sort="Kirsanow, Karola" uniqKey="Kirsanow K" first="Karola" last="Kirsanow">Karola Kirsanow</name>
<affiliation><nlm:aff id="A5">Johannes Gutenberg University Mainz, Institute of Anthropology, Mainz, D-55128, Germany</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Sudmant, Peter H" sort="Sudmant, Peter H" uniqKey="Sudmant P" first="Peter H." last="Sudmant">Peter H. Sudmant</name>
<affiliation><nlm:aff id="A6">Department of Genome Sciences, University of Washington, Seattle, WA, 98195, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Schraiber, Joshua G" sort="Schraiber, Joshua G" uniqKey="Schraiber J" first="Joshua G." last="Schraiber">Joshua G. Schraiber</name>
<affiliation><nlm:aff id="A7">Department of Integrative Biology, University of California, Berkeley, CA, 94720-3140, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Castellano, Sergi" sort="Castellano, Sergi" uniqKey="Castellano S" first="Sergi" last="Castellano">Sergi Castellano</name>
<affiliation><nlm:aff id="A4">Max Planck Institute for Evolutionary Anthropology, Leipzig, 04103, Germany</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Lipson, Mark" sort="Lipson, Mark" uniqKey="Lipson M" first="Mark" last="Lipson">Mark Lipson</name>
<affiliation><nlm:aff id="A8">Department of Mathematics and Computer Science and Artificial Intelligence Laboratory, Massachusetts Institute of Technology, Cambridge, MA 02139, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Berger, Bonnie" sort="Berger, Bonnie" uniqKey="Berger B" first="Bonnie" last="Berger">Bonnie Berger</name>
<affiliation><nlm:aff id="A2">Broad Institute of Harvard and MIT, Cambridge, MA, 02142, USA</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A8">Department of Mathematics and Computer Science and Artificial Intelligence Laboratory, Massachusetts Institute of Technology, Cambridge, MA 02139, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Economou, Christos" sort="Economou, Christos" uniqKey="Economou C" first="Christos" last="Economou">Christos Economou</name>
<affiliation><nlm:aff id="A9">Archaeological Research Laboratory, Stockholm University, 114 18, Sweden</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Bollongino, Ruth" sort="Bollongino, Ruth" uniqKey="Bollongino R" first="Ruth" last="Bollongino">Ruth Bollongino</name>
<affiliation><nlm:aff id="A5">Johannes Gutenberg University Mainz, Institute of Anthropology, Mainz, D-55128, Germany</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Fu, Qiaomei" sort="Fu, Qiaomei" uniqKey="Fu Q" first="Qiaomei" last="Fu">Qiaomei Fu</name>
<affiliation><nlm:aff id="A1">Department of Genetics, Harvard Medical School, Boston, MA, 02115, USA</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A4">Max Planck Institute for Evolutionary Anthropology, Leipzig, 04103, Germany</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A10">Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, IVPP, CAS, Beijing, 100049, China</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Bos, Kirsten I" sort="Bos, Kirsten I" uniqKey="Bos K" first="Kirsten I." last="Bos">Kirsten I. Bos</name>
<affiliation><nlm:aff id="A3">Institute for Archaeological Sciences, University of Tübingen, Tübingen, 72074, Germany</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Nordenfelt, Susanne" sort="Nordenfelt, Susanne" uniqKey="Nordenfelt S" first="Susanne" last="Nordenfelt">Susanne Nordenfelt</name>
<affiliation><nlm:aff id="A1">Department of Genetics, Harvard Medical School, Boston, MA, 02115, USA</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A2">Broad Institute of Harvard and MIT, Cambridge, MA, 02142, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Li, Heng" sort="Li, Heng" uniqKey="Li H" first="Heng" last="Li">Heng Li</name>
<affiliation><nlm:aff id="A1">Department of Genetics, Harvard Medical School, Boston, MA, 02115, USA</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A2">Broad Institute of Harvard and MIT, Cambridge, MA, 02142, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="De Filippo, Cesare" sort="De Filippo, Cesare" uniqKey="De Filippo C" first="Cesare" last="De Filippo">Cesare De Filippo</name>
<affiliation><nlm:aff id="A4">Max Planck Institute for Evolutionary Anthropology, Leipzig, 04103, Germany</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Prufer, Kay" sort="Prufer, Kay" uniqKey="Prufer K" first="Kay" last="Prüfer">Kay Prüfer</name>
<affiliation><nlm:aff id="A4">Max Planck Institute for Evolutionary Anthropology, Leipzig, 04103, Germany</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Sawyer, Susanna" sort="Sawyer, Susanna" uniqKey="Sawyer S" first="Susanna" last="Sawyer">Susanna Sawyer</name>
<affiliation><nlm:aff id="A4">Max Planck Institute for Evolutionary Anthropology, Leipzig, 04103, Germany</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Posth, Cosimo" sort="Posth, Cosimo" uniqKey="Posth C" first="Cosimo" last="Posth">Cosimo Posth</name>
<affiliation><nlm:aff id="A3">Institute for Archaeological Sciences, University of Tübingen, Tübingen, 72074, Germany</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Haak, Wolfgang" sort="Haak, Wolfgang" uniqKey="Haak W" first="Wolfgang" last="Haak">Wolfgang Haak</name>
<affiliation><nlm:aff id="A11">Australian Centre for Ancient DNA, School of Earth and Environmental Sciences, University of Adelaide, Adelaide, South Australia, SA 5005, Australia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Hallgren, Fredrik" sort="Hallgren, Fredrik" uniqKey="Hallgren F" first="Fredrik" last="Hallgren">Fredrik Hallgren</name>
<affiliation><nlm:aff id="A12">The Cultural Heritage Foundation, Västerås, 722 12, Sweden</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Fornander, Elin" sort="Fornander, Elin" uniqKey="Fornander E" first="Elin" last="Fornander">Elin Fornander</name>
<affiliation><nlm:aff id="A12">The Cultural Heritage Foundation, Västerås, 722 12, Sweden</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Rohland, Nadin" sort="Rohland, Nadin" uniqKey="Rohland N" first="Nadin" last="Rohland">Nadin Rohland</name>
<affiliation><nlm:aff id="A1">Department of Genetics, Harvard Medical School, Boston, MA, 02115, USA</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A2">Broad Institute of Harvard and MIT, Cambridge, MA, 02142, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Delsate, Dominique" sort="Delsate, Dominique" uniqKey="Delsate D" first="Dominique" last="Delsate">Dominique Delsate</name>
<affiliation><nlm:aff id="A13">National Museum of Natural History, L-2160, Luxembourg</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A14">National Center of Archaeological Research, National Museum of History and Art, L-2345, Luxembourg</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Francken, Michael" sort="Francken, Michael" uniqKey="Francken M" first="Michael" last="Francken">Michael Francken</name>
<affiliation><nlm:aff id="A15">Department of Paleoanthropology, Senckenberg Center for Human Evolution and Paleoenvironment, University of Tübingen, Tübingen, D-72070, Germany</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Guinet, Jean Michel" sort="Guinet, Jean Michel" uniqKey="Guinet J" first="Jean-Michel" last="Guinet">Jean-Michel Guinet</name>
<affiliation><nlm:aff id="A13">National Museum of Natural History, L-2160, Luxembourg</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Wahl, Joachim" sort="Wahl, Joachim" uniqKey="Wahl J" first="Joachim" last="Wahl">Joachim Wahl</name>
<affiliation><nlm:aff id="A16">State Office for Cultural Heritage Management Baden-Württemberg, Osteology, Konstanz, D-78467, Germany</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Ayodo, George" sort="Ayodo, George" uniqKey="Ayodo G" first="George" last="Ayodo">George Ayodo</name>
<affiliation><nlm:aff id="A17">Center for Global Health and Child Development, Kisumu, 40100, Kenya</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Babiker, Hamza A" sort="Babiker, Hamza A" uniqKey="Babiker H" first="Hamza A." last="Babiker">Hamza A. Babiker</name>
<affiliation><nlm:aff id="A18">Institutes of Evolution, Immunology and Infection Research, School of Biological Sciences, University of Edinburgh, Edinburgh, EH9 3JT, UK</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A19">Biochemistry Department, Faculty of Medicine, Sultan Qaboos University, Alkhod, Muscat, 123, Oman</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Bailliet, Graciela" sort="Bailliet, Graciela" uniqKey="Bailliet G" first="Graciela" last="Bailliet">Graciela Bailliet</name>
<affiliation><nlm:aff id="A20">Laboratorio de Genética Molecular Poblacional, Instituto Multidisciplinario de Biología Celular (IMBICE), CCT-CONICET & CICPBA, La Plata, B1906APO, Argentina</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Balanovska, Elena" sort="Balanovska, Elena" uniqKey="Balanovska E" first="Elena" last="Balanovska">Elena Balanovska</name>
<affiliation><nlm:aff id="A21">Research Centre for Medical Genetics, Moscow, 115478, Russia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Balanovsky, Oleg" sort="Balanovsky, Oleg" uniqKey="Balanovsky O" first="Oleg" last="Balanovsky">Oleg Balanovsky</name>
<affiliation><nlm:aff id="A21">Research Centre for Medical Genetics, Moscow, 115478, Russia</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A22">Vavilov Institute for General Genetics, Moscow, 119991, Russia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Barrantes, Ramiro" sort="Barrantes, Ramiro" uniqKey="Barrantes R" first="Ramiro" last="Barrantes">Ramiro Barrantes</name>
<affiliation><nlm:aff id="A23">Escuela de Biología, Universidad de Costa Rica, San José, 2060, Costa Rica</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Bedoya, Gabriel" sort="Bedoya, Gabriel" uniqKey="Bedoya G" first="Gabriel" last="Bedoya">Gabriel Bedoya</name>
<affiliation><nlm:aff id="A24">Institute of Biology, Research group GENMOL, Universidad de Antioquia, Medellín, Colombia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Ben Ami, Haim" sort="Ben Ami, Haim" uniqKey="Ben Ami H" first="Haim" last="Ben-Ami">Haim Ben-Ami</name>
<affiliation><nlm:aff id="A25">Rambam Health Care Campus, Haifa, 31096, Israel</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Bene, Judit" sort="Bene, Judit" uniqKey="Bene J" first="Judit" last="Bene">Judit Bene</name>
<affiliation><nlm:aff id="A26">Department of Medical Genetics and Szentagothai Research Center, University of Pécs, Pécs, H-7624 Hungary</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Berrada, Fouad" sort="Berrada, Fouad" uniqKey="Berrada F" first="Fouad" last="Berrada">Fouad Berrada</name>
<affiliation><nlm:aff id="A27">Al Akhawayn University in Ifrane (AUI), School of Science and Engineering, Ifrane, 53000, Morocco</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Bravi, Claudio M" sort="Bravi, Claudio M" uniqKey="Bravi C" first="Claudio M." last="Bravi">Claudio M. Bravi</name>
<affiliation><nlm:aff id="A20">Laboratorio de Genética Molecular Poblacional, Instituto Multidisciplinario de Biología Celular (IMBICE), CCT-CONICET & CICPBA, La Plata, B1906APO, Argentina</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Brisighelli, Francesca" sort="Brisighelli, Francesca" uniqKey="Brisighelli F" first="Francesca" last="Brisighelli">Francesca Brisighelli</name>
<affiliation><nlm:aff id="A28">Forensic Genetics Laboratory, Institute of Legal Medicine, Università Cattolica del Sacro Cuore, Rome, 00168, Italy</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Busby, George B J" sort="Busby, George B J" uniqKey="Busby G" first="George B. J." last="Busby">George B. J. Busby</name>
<affiliation><nlm:aff id="A29">Department of Zoology, University of Oxford, Oxford, OX1 3PS, UK</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A30">Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford, OX3 7BN, UK</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Cali, Francesco" sort="Cali, Francesco" uniqKey="Cali F" first="Francesco" last="Cali">Francesco Cali</name>
<affiliation><nlm:aff id="A31">Laboratorio di Genetica Molecolare, IRCCS Associazione Oasi Maria SS, Troina, 94018, Italy</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Churnosov, Mikhail" sort="Churnosov, Mikhail" uniqKey="Churnosov M" first="Mikhail" last="Churnosov">Mikhail Churnosov</name>
<affiliation><nlm:aff id="A32">Belgorod State University, Belgorod, 308015, Russia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Cole, David E C" sort="Cole, David E C" uniqKey="Cole D" first="David E. C." last="Cole">David E. C. Cole</name>
<affiliation><nlm:aff id="A33">Department of Laboratory Medicine and Pathobiology, University of Toronto, Toronto, Ontario, M5G 1L5, Canada</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Corach, Daniel" sort="Corach, Daniel" uniqKey="Corach D" first="Daniel" last="Corach">Daniel Corach</name>
<affiliation><nlm:aff id="A34">Servicio de Huellas Digitales Genéticas, School of Pharmacy and Biochemistry, Universidad de Buenos Aires, 1113 CABA, Argentina</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Damba, Larissa" sort="Damba, Larissa" uniqKey="Damba L" first="Larissa" last="Damba">Larissa Damba</name>
<affiliation><nlm:aff id="A35">Institute of Cytology and Genetics, Siberian Branch of Russian Academy of Sciences, Novosibirsk, 630090, Russia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Van Driem, George" sort="Van Driem, George" uniqKey="Van Driem G" first="George" last="Van Driem">George Van Driem</name>
<affiliation><nlm:aff id="A36">Institute of Linguistics, University of Bern, Bern, CH-3012, Switzerland</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Dryomov, Stanislav" sort="Dryomov, Stanislav" uniqKey="Dryomov S" first="Stanislav" last="Dryomov">Stanislav Dryomov</name>
<affiliation><nlm:aff id="A37">Laboratory of Human Molecular Genetics, Institute of Molecular and Cellular Biology, Russian Academy of Science, Siberian Branch, Novosibirsk, 630090, Russia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Dugoujon, Jean Michel" sort="Dugoujon, Jean Michel" uniqKey="Dugoujon J" first="Jean-Michel" last="Dugoujon">Jean-Michel Dugoujon</name>
<affiliation><nlm:aff id="A38">Anthropologie Moléculaire et Imagerie de Synthèse, CNRS UMR 5288, Université Paul Sabatier Toulouse III, Toulouse, 31000, France</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Fedorova, Sardana A" sort="Fedorova, Sardana A" uniqKey="Fedorova S" first="Sardana A." last="Fedorova">Sardana A. Fedorova</name>
<affiliation><nlm:aff id="A39">Yakut Research Center of Complex Medical Problems and North-Eastern Federal University, Yakutsk, 677010, Russia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Romero, Irene Gallego" sort="Romero, Irene Gallego" uniqKey="Romero I" first="Irene Gallego" last="Romero">Irene Gallego Romero</name>
<affiliation><nlm:aff id="A40">Department of Human Genetics, University of Chicago, Chicago, IL, 60637, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Gubina, Marina" sort="Gubina, Marina" uniqKey="Gubina M" first="Marina" last="Gubina">Marina Gubina</name>
<affiliation><nlm:aff id="A35">Institute of Cytology and Genetics, Siberian Branch of Russian Academy of Sciences, Novosibirsk, 630090, Russia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Hammer, Michael" sort="Hammer, Michael" uniqKey="Hammer M" first="Michael" last="Hammer">Michael Hammer</name>
<affiliation><nlm:aff id="A41">ARL Division of Biotechnology, University of Arizona, Tucson, AZ, 85721, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Henn, Brenna M" sort="Henn, Brenna M" uniqKey="Henn B" first="Brenna M." last="Henn">Brenna M. Henn</name>
<affiliation><nlm:aff id="A42">Department of Ecology and Evolution, Stony Brook University, Stony Brook, NY, 11794, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Hervig, Tor" sort="Hervig, Tor" uniqKey="Hervig T" first="Tor" last="Hervig">Tor Hervig</name>
<affiliation><nlm:aff id="A43">Department of Clinical Science, University of Bergen, Bergen, 5021, Norway</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Hodoglugil, Ugur" sort="Hodoglugil, Ugur" uniqKey="Hodoglugil U" first="Ugur" last="Hodoglugil">Ugur Hodoglugil</name>
<affiliation><nlm:aff id="A44">NextBio, part of Illumina, Santa Clara, CA, USA 95050</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Jha, Aashish R" sort="Jha, Aashish R" uniqKey="Jha A" first="Aashish R." last="Jha">Aashish R. Jha</name>
<affiliation><nlm:aff id="A40">Department of Human Genetics, University of Chicago, Chicago, IL, 60637, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Karachanak Yankova, Sena" sort="Karachanak Yankova, Sena" uniqKey="Karachanak Yankova S" first="Sena" last="Karachanak-Yankova">Sena Karachanak-Yankova</name>
<affiliation><nlm:aff id="A45">Dept. of Medical Genetics, National Human Genome Center, Medical University Sofia, Sofia, 1431, Bulgaria</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Khusainova, Rita" sort="Khusainova, Rita" uniqKey="Khusainova R" first="Rita" last="Khusainova">Rita Khusainova</name>
<affiliation><nlm:aff id="A46">Institute of Biochemistry and Genetics, Ufa Research Centre, Russian Academy of Sciences, Ufa, 450054, Russia</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A47">Department of Genetics and Fundamental Medicine, Bashkir State University, Ufa, 450074, Russia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Khusnutdinova, Elza" sort="Khusnutdinova, Elza" uniqKey="Khusnutdinova E" first="Elza" last="Khusnutdinova">Elza Khusnutdinova</name>
<affiliation><nlm:aff id="A46">Institute of Biochemistry and Genetics, Ufa Research Centre, Russian Academy of Sciences, Ufa, 450054, Russia</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A47">Department of Genetics and Fundamental Medicine, Bashkir State University, Ufa, 450074, Russia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Kittles, Rick" sort="Kittles, Rick" uniqKey="Kittles R" first="Rick" last="Kittles">Rick Kittles</name>
<affiliation><nlm:aff id="A48">College of Medicine, University of Illinois at Chicago, Chicago, IL, 60607, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Kivisild, Toomas" sort="Kivisild, Toomas" uniqKey="Kivisild T" first="Toomas" last="Kivisild">Toomas Kivisild</name>
<affiliation><nlm:aff id="A49">Division of Biological Anthropology, University of Cambridge, Cambridge, United Kingdom CB2 1QH</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Klitz, William" sort="Klitz, William" uniqKey="Klitz W" first="William" last="Klitz">William Klitz</name>
<affiliation><nlm:aff id="A7">Department of Integrative Biology, University of California, Berkeley, CA, 94720-3140, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Ku Inskas, Vaidutis" sort="Ku Inskas, Vaidutis" uniqKey="Ku Inskas V" first="Vaidutis" last="Ku Inskas">Vaidutis Ku Inskas</name>
<affiliation><nlm:aff id="A50">Department of Human and Medical Genetics, Vilnius University, Vilnius, LT-08661, Lithuania</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Kushniarevich, Alena" sort="Kushniarevich, Alena" uniqKey="Kushniarevich A" first="Alena" last="Kushniarevich">Alena Kushniarevich</name>
<affiliation><nlm:aff id="A51">Estonian Biocentre, Evolutionary Biology group, Tartu, 51010, Estonia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Laredj, Leila" sort="Laredj, Leila" uniqKey="Laredj L" first="Leila" last="Laredj">Leila Laredj</name>
<affiliation><nlm:aff id="A52">Translational Medicine and Neurogenetics, Institut de Génétique et de Biologie Moléculaire et Cellulaire, Illkirch, 67404, France</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Litvinov, Sergey" sort="Litvinov, Sergey" uniqKey="Litvinov S" first="Sergey" last="Litvinov">Sergey Litvinov</name>
<affiliation><nlm:aff id="A46">Institute of Biochemistry and Genetics, Ufa Research Centre, Russian Academy of Sciences, Ufa, 450054, Russia</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A47">Department of Genetics and Fundamental Medicine, Bashkir State University, Ufa, 450074, Russia</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A51">Estonian Biocentre, Evolutionary Biology group, Tartu, 51010, Estonia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Loukidis, Theologos" sort="Loukidis, Theologos" uniqKey="Loukidis T" first="Theologos" last="Loukidis">Theologos Loukidis</name>
</author>
<author><name sortKey="Mahley, Robert W" sort="Mahley, Robert W" uniqKey="Mahley R" first="Robert W." last="Mahley">Robert W. Mahley</name>
<affiliation><nlm:aff id="A53">Gladstone Institutes, San Francisco, CA, 94158, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Melegh, Bela" sort="Melegh, Bela" uniqKey="Melegh B" first="Béla" last="Melegh">Béla Melegh</name>
<affiliation><nlm:aff id="A26">Department of Medical Genetics and Szentagothai Research Center, University of Pécs, Pécs, H-7624 Hungary</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Metspalu, Ene" sort="Metspalu, Ene" uniqKey="Metspalu E" first="Ene" last="Metspalu">Ene Metspalu</name>
<affiliation><nlm:aff id="A54">Department of Evolutionary Biology, University of Tartu, Tartu, 51010, Estonia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Molina, Julio" sort="Molina, Julio" uniqKey="Molina J" first="Julio" last="Molina">Julio Molina</name>
<affiliation><nlm:aff id="A55">Centro de Investigaciones Biomédicas de Guatemala, Ciudad de Guatemala, Guatemala</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Mountain, Joanna" sort="Mountain, Joanna" uniqKey="Mountain J" first="Joanna" last="Mountain">Joanna Mountain</name>
<affiliation><nlm:aff id="A56">Research Department, 23andMe, Inc. Mountain View, CA, 94043, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="N Kk L J Rvi, Klemetti" sort="N Kk L J Rvi, Klemetti" uniqKey="N Kk L J Rvi K" first="Klemetti" last="N Kk L J Rvi">Klemetti N Kk L J Rvi</name>
<affiliation><nlm:aff id="A57">Cultural Anthropology Program, University of Oulu, Oulu, 90014, Finland</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Nesheva, Desislava" sort="Nesheva, Desislava" uniqKey="Nesheva D" first="Desislava" last="Nesheva">Desislava Nesheva</name>
<affiliation><nlm:aff id="A45">Dept. of Medical Genetics, National Human Genome Center, Medical University Sofia, Sofia, 1431, Bulgaria</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Nyambo, Thomas" sort="Nyambo, Thomas" uniqKey="Nyambo T" first="Thomas" last="Nyambo">Thomas Nyambo</name>
<affiliation><nlm:aff id="A58">Department of Biochemistry, Muhimbili University of Health and Allied Sciences, Dar es Salaam, Tanzania</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Osipova, Ludmila" sort="Osipova, Ludmila" uniqKey="Osipova L" first="Ludmila" last="Osipova">Ludmila Osipova</name>
<affiliation><nlm:aff id="A35">Institute of Cytology and Genetics, Siberian Branch of Russian Academy of Sciences, Novosibirsk, 630090, Russia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Parik, Juri" sort="Parik, Juri" uniqKey="Parik J" first="Jüri" last="Parik">Jüri Parik</name>
<affiliation><nlm:aff id="A54">Department of Evolutionary Biology, University of Tartu, Tartu, 51010, Estonia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Platonov, Fedor" sort="Platonov, Fedor" uniqKey="Platonov F" first="Fedor" last="Platonov">Fedor Platonov</name>
<affiliation><nlm:aff id="A59">Research Institute of Health, North-Eastern Federal University, Yakutsk, 677000, Russia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Posukh, Olga" sort="Posukh, Olga" uniqKey="Posukh O" first="Olga" last="Posukh">Olga Posukh</name>
<affiliation><nlm:aff id="A35">Institute of Cytology and Genetics, Siberian Branch of Russian Academy of Sciences, Novosibirsk, 630090, Russia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Romano, Valentino" sort="Romano, Valentino" uniqKey="Romano V" first="Valentino" last="Romano">Valentino Romano</name>
<affiliation><nlm:aff id="A60">Dipartimento di Fisica e Chimica, Università di Palermo, Palermo, 90128, Italy</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Rothhammer, Francisco" sort="Rothhammer, Francisco" uniqKey="Rothhammer F" first="Francisco" last="Rothhammer">Francisco Rothhammer</name>
<affiliation><nlm:aff id="A61">Instituto de Alta Investigación, Universidad de Tarapacá, Arica, Chile</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A62">Programa de Genética Humana ICBM Facultad de Medicina Universidad de Chile, Santiago, Chile</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A63">Centro de Investigaciones del Hombre en el Desierto, Arica, Chile</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Rudan, Igor" sort="Rudan, Igor" uniqKey="Rudan I" first="Igor" last="Rudan">Igor Rudan</name>
<affiliation><nlm:aff id="A64">Centre for Population Health Sciences, The University of Edinburgh Medical School, Edinburgh, Scotland, EH8 9AG, UK</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Ruizbakiev, Ruslan" sort="Ruizbakiev, Ruslan" uniqKey="Ruizbakiev R" first="Ruslan" last="Ruizbakiev">Ruslan Ruizbakiev</name>
<affiliation><nlm:aff id="A65">Deceased: formerly of the Institute of Immunology, Academy of Science, Tashkent, 70000, Uzbekistan</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Sahakyan, Hovhannes" sort="Sahakyan, Hovhannes" uniqKey="Sahakyan H" first="Hovhannes" last="Sahakyan">Hovhannes Sahakyan</name>
<affiliation><nlm:aff id="A51">Estonian Biocentre, Evolutionary Biology group, Tartu, 51010, Estonia</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A66">Laboratory of Ethnogenomics, Institute of Molecular Biology, National Academy of Sciences of Armenia, Yerevan, 0014, Armenia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Sajantila, Antti" sort="Sajantila, Antti" uniqKey="Sajantila A" first="Antti" last="Sajantila">Antti Sajantila</name>
<affiliation><nlm:aff id="A67">Department of Forensic Medicine, Hjelt Institute, University of Helsinki, Helsinki, 00014, Finland</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A68">Institute of Applied Genetics, Department of Molecular and Medical Genetics, University of North Texas Health Science Center, Fort Worth, Texas 76107, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Salas, Antonio" sort="Salas, Antonio" uniqKey="Salas A" first="Antonio" last="Salas">Antonio Salas</name>
<affiliation><nlm:aff id="A69">Unidade 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</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Starikovskaya, Elena B" sort="Starikovskaya, Elena B" uniqKey="Starikovskaya E" first="Elena B." last="Starikovskaya">Elena B. Starikovskaya</name>
<affiliation><nlm:aff id="A37">Laboratory of Human Molecular Genetics, Institute of Molecular and Cellular Biology, Russian Academy of Science, Siberian Branch, Novosibirsk, 630090, Russia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Tarekegn, Ayele" sort="Tarekegn, Ayele" uniqKey="Tarekegn A" first="Ayele" last="Tarekegn">Ayele Tarekegn</name>
<affiliation><nlm:aff id="A70">Research Fellow, Henry Stewart Group, Russell House, London WC1A 2HN, UK</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Toncheva, Draga" sort="Toncheva, Draga" uniqKey="Toncheva D" first="Draga" last="Toncheva">Draga Toncheva</name>
<affiliation><nlm:aff id="A45">Dept. of Medical Genetics, National Human Genome Center, Medical University Sofia, Sofia, 1431, Bulgaria</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Turdikulova, Shahlo" sort="Turdikulova, Shahlo" uniqKey="Turdikulova S" first="Shahlo" last="Turdikulova">Shahlo Turdikulova</name>
<affiliation><nlm:aff id="A71">Institute of Bioorganic Chemistry Academy of Sciences Republic of Uzbekistan, Tashkent, 100125, Uzbekistan</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Uktveryte, Ingrida" sort="Uktveryte, Ingrida" uniqKey="Uktveryte I" first="Ingrida" last="Uktveryte">Ingrida Uktveryte</name>
<affiliation><nlm:aff id="A50">Department of Human and Medical Genetics, Vilnius University, Vilnius, LT-08661, Lithuania</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Utevska, Olga" sort="Utevska, Olga" uniqKey="Utevska O" first="Olga" last="Utevska">Olga Utevska</name>
<affiliation><nlm:aff id="A72">Department of Genetics and Cytology, V.N. Karazin Kharkiv National University, Kharkiv, 61077, Ukraine</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Vasquez, Rene" sort="Vasquez, Rene" uniqKey="Vasquez R" first="René" last="Vasquez">René Vasquez</name>
<affiliation><nlm:aff id="A73">Instituto Boliviano de Biología de la Altura, Universidad Autonoma Tomás Frías, Potosí, Bolivia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Villena, Mercedes" sort="Villena, Mercedes" uniqKey="Villena M" first="Mercedes" last="Villena">Mercedes Villena</name>
<affiliation><nlm:aff id="A73">Instituto Boliviano de Biología de la Altura, Universidad Autonoma Tomás Frías, Potosí, Bolivia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Voevoda, Mikhail" sort="Voevoda, Mikhail" uniqKey="Voevoda M" first="Mikhail" last="Voevoda">Mikhail Voevoda</name>
<affiliation><nlm:aff id="A35">Institute of Cytology and Genetics, Siberian Branch of Russian Academy of Sciences, Novosibirsk, 630090, Russia</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A74">Inst. of Internal Medicine, Siberian Branch of Russian Acad. of Medical Sciences, Novosibirsk, 630089, Russia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Winkler, Cheryl" sort="Winkler, Cheryl" uniqKey="Winkler C" first="Cheryl" last="Winkler">Cheryl Winkler</name>
<affiliation><nlm:aff id="A75">Basic Research Laboratory, NCI, NIH, Frederick National Laboratory, Leidos Biomedical, Inc., Frederick, MD 21702, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Yepiskoposyan, Levon" sort="Yepiskoposyan, Levon" uniqKey="Yepiskoposyan L" first="Levon" last="Yepiskoposyan">Levon Yepiskoposyan</name>
<affiliation><nlm:aff id="A66">Laboratory of Ethnogenomics, Institute of Molecular Biology, National Academy of Sciences of Armenia, Yerevan, 0014, Armenia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Zalloua, Pierre" sort="Zalloua, Pierre" uniqKey="Zalloua P" first="Pierre" last="Zalloua">Pierre Zalloua</name>
<affiliation><nlm:aff id="A76">Lebanese American University, School of Medicine, Beirut, 13-5053, Lebanon</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A77">Harvard School of Public Health, Boston, 02115, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Zemunik, Tatijana" sort="Zemunik, Tatijana" uniqKey="Zemunik T" first="Tatijana" last="Zemunik">Tatijana Zemunik</name>
<affiliation><nlm:aff id="A78">Department of Medical Biology, University of Split, School of Medicine, Split, 21000, Croatia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Cooper, Alan" sort="Cooper, Alan" uniqKey="Cooper A" first="Alan" last="Cooper">Alan Cooper</name>
<affiliation><nlm:aff id="A11">Australian Centre for Ancient DNA, School of Earth and Environmental Sciences, University of Adelaide, Adelaide, South Australia, SA 5005, Australia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Capelli, Cristian" sort="Capelli, Cristian" uniqKey="Capelli C" first="Cristian" last="Capelli">Cristian Capelli</name>
<affiliation><nlm:aff id="A29">Department of Zoology, University of Oxford, Oxford, OX1 3PS, UK</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Thomas, Mark G" sort="Thomas, Mark G" uniqKey="Thomas M" first="Mark G." last="Thomas">Mark G. Thomas</name>
<affiliation><nlm:aff id="A79">Department of Genetics, Evolution and Environment, University College London, WC1E 6BT, UK</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Ruiz Linares, Andres" sort="Ruiz Linares, Andres" uniqKey="Ruiz Linares A" first="Andres" last="Ruiz-Linares">Andres Ruiz-Linares</name>
<affiliation><nlm:aff id="A79">Department of Genetics, Evolution and Environment, University College London, WC1E 6BT, UK</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Tishkoff, Sarah A" sort="Tishkoff, Sarah A" uniqKey="Tishkoff S" first="Sarah A." last="Tishkoff">Sarah A. Tishkoff</name>
<affiliation><nlm:aff id="A80">Department of Biology and Genetics. University of Pennsylvania, Philadelphia, Pennsylvania, 19104, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Singh, Lalji" sort="Singh, Lalji" uniqKey="Singh L" first="Lalji" last="Singh">Lalji Singh</name>
<affiliation><nlm:aff id="A81">CSIR-Centre for Cellular and Molecular Biology, Hyderabad, 500 007, India</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Thangaraj, Kumarasamy" sort="Thangaraj, Kumarasamy" uniqKey="Thangaraj K" first="Kumarasamy" last="Thangaraj">Kumarasamy Thangaraj</name>
<affiliation><nlm:aff id="A81">CSIR-Centre for Cellular and Molecular Biology, Hyderabad, 500 007, India</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Villems, Richard" sort="Villems, Richard" uniqKey="Villems R" first="Richard" last="Villems">Richard Villems</name>
<affiliation><nlm:aff id="A51">Estonian Biocentre, Evolutionary Biology group, Tartu, 51010, Estonia</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A54">Department of Evolutionary Biology, University of Tartu, Tartu, 51010, Estonia</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A82">Estonian Academy of Sciences, Tallinn, 10130, Estonia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Comas, David" sort="Comas, David" uniqKey="Comas D" first="David" last="Comas">David Comas</name>
<affiliation><nlm:aff id="A83">Institut de Biologia Evolutiva (CSIC-UPF), Departament de Ciències Experimentals i de la Salut, Universitat Pompeu Fabra, Barcelona, 08003, Spain</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Sukernik, Rem" sort="Sukernik, Rem" uniqKey="Sukernik R" first="Rem" last="Sukernik">Rem Sukernik</name>
<affiliation><nlm:aff id="A37">Laboratory of Human Molecular Genetics, Institute of Molecular and Cellular Biology, Russian Academy of Science, Siberian Branch, Novosibirsk, 630090, Russia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Metspalu, Mait" sort="Metspalu, Mait" uniqKey="Metspalu M" first="Mait" last="Metspalu">Mait Metspalu</name>
<affiliation><nlm:aff id="A51">Estonian Biocentre, Evolutionary Biology group, Tartu, 51010, Estonia</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Meyer, Matthias" sort="Meyer, Matthias" uniqKey="Meyer M" first="Matthias" last="Meyer">Matthias Meyer</name>
<affiliation><nlm:aff id="A4">Max Planck Institute for Evolutionary Anthropology, Leipzig, 04103, Germany</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Eichler, Evan E" sort="Eichler, Evan E" uniqKey="Eichler E" first="Evan E." last="Eichler">Evan E. Eichler</name>
<affiliation><nlm:aff id="A6">Department of Genome Sciences, University of Washington, Seattle, WA, 98195, USA</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A84">Howard Hughes Medical Institute, University of Washington, Seattle, WA, USA 98195</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Burger, Joachim" sort="Burger, Joachim" uniqKey="Burger J" first="Joachim" last="Burger">Joachim Burger</name>
<affiliation><nlm:aff id="A5">Johannes Gutenberg University Mainz, Institute of Anthropology, Mainz, D-55128, Germany</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Slatkin, Montgomery" sort="Slatkin, Montgomery" uniqKey="Slatkin M" first="Montgomery" last="Slatkin">Montgomery Slatkin</name>
<affiliation><nlm:aff id="A7">Department of Integrative Biology, University of California, Berkeley, CA, 94720-3140, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="P Bo, Svante" sort="P Bo, Svante" uniqKey="P Bo S" first="Svante" last="P Bo">Svante P Bo</name>
<affiliation><nlm:aff id="A4">Max Planck Institute for Evolutionary Anthropology, Leipzig, 04103, Germany</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Kelso, Janet" sort="Kelso, Janet" uniqKey="Kelso J" first="Janet" last="Kelso">Janet Kelso</name>
<affiliation><nlm:aff id="A4">Max Planck Institute for Evolutionary Anthropology, Leipzig, 04103, Germany</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Reich, David" sort="Reich, David" uniqKey="Reich D" first="David" last="Reich">David Reich</name>
<affiliation><nlm:aff id="A1">Department of Genetics, Harvard Medical School, Boston, MA, 02115, USA</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A2">Broad Institute of Harvard and MIT, Cambridge, MA, 02142, USA</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A85">Howard Hughes Medical Institute, Harvard Medical School, Boston, MA, 02115, USA</nlm:aff>
</affiliation>
</author>
<author><name sortKey="Krause, Johannes" sort="Krause, Johannes" uniqKey="Krause J" first="Johannes" last="Krause">Johannes Krause</name>
<affiliation><nlm:aff id="A3">Institute for Archaeological Sciences, University of Tübingen, Tübingen, 72074, Germany</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A86">Senckenberg Centre for Human Evolution and Palaeoenvironment, University of Tübingen, 72070, Germany</nlm:aff>
</affiliation>
<affiliation><nlm:aff id="A87">Max Planck Institut für Geschichte und Naturwissenschaften, Jena, 07745, Germany</nlm:aff>
</affiliation>
</author>
</analytic>
<series><title level="j">Nature</title>
<idno type="ISSN">0028-0836</idno>
<idno type="eISSN">1476-4687</idno>
<imprint><date when="2014">2014</date>
</imprint>
</series>
</biblStruct>
</sourceDesc>
</fileDesc>
<profileDesc><textClass></textClass>
</profileDesc>
</teiHeader>
<front><div type="abstract" xml:lang="en"><p id="P1">We sequenced the genomes of a ~7,000 year old farmer from Germany and eight
~8,000 year old hunter-gatherers from Luxembourg and Sweden. We analyzed these and other
ancient genomes<sup><xref rid="R1" ref-type="bibr">1</xref>
–<xref rid="R4" ref-type="bibr">4</xref>
</sup>
with 2,345 contemporary humans to show that most
present 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) related to Upper Paleolithic Siberians<sup><xref rid="R3" ref-type="bibr">3</xref>
</sup>
, who 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”
population that split prior to the diversification of other non-African lineages.</p>
</div>
</front>
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<pmc article-type="research-article"><pmc-dir>properties open_access</pmc-dir>
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<front><journal-meta><journal-id journal-id-type="nlm-journal-id">0410462</journal-id>
<journal-id journal-id-type="pubmed-jr-id">6011</journal-id>
<journal-id journal-id-type="nlm-ta">Nature</journal-id>
<journal-id journal-id-type="iso-abbrev">Nature</journal-id>
<journal-title-group><journal-title>Nature</journal-title>
</journal-title-group>
<issn pub-type="ppub">0028-0836</issn>
<issn pub-type="epub">1476-4687</issn>
</journal-meta>
<article-meta><article-id pub-id-type="pmid">25230663</article-id>
<article-id pub-id-type="pmc">4170574</article-id>
<article-id pub-id-type="doi">10.1038/nature13673</article-id>
<article-id pub-id-type="manuscript">NIHMS613260</article-id>
<article-categories><subj-group subj-group-type="heading"><subject>Article</subject>
</subj-group>
</article-categories>
<title-group><article-title>Ancient human genomes suggest three ancestral populations for present-day
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<contrib contrib-type="author"><name><surname>Economou</surname>
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<given-names>Cesare</given-names>
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<contrib contrib-type="author"><name><surname>Wahl</surname>
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<contrib contrib-type="author"><name><surname>Barrantes</surname>
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<contrib contrib-type="author"><name><surname>Ben-Ami</surname>
<given-names>Haim</given-names>
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<xref ref-type="aff" rid="A25">25</xref>
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<contrib contrib-type="author"><name><surname>Bene</surname>
<given-names>Judit</given-names>
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<contrib contrib-type="author"><name><surname>Berrada</surname>
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<given-names>Claudio M.</given-names>
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<xref ref-type="aff" rid="A20">20</xref>
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<contrib contrib-type="author"><name><surname>Brisighelli</surname>
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<contrib contrib-type="author"><name><surname>Busby</surname>
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<xref ref-type="aff" rid="A30">30</xref>
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<contrib contrib-type="author"><name><surname>Cali</surname>
<given-names>Francesco</given-names>
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<contrib contrib-type="author"><name><surname>Churnosov</surname>
<given-names>Mikhail</given-names>
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<given-names>David E. C.</given-names>
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<given-names>Larissa</given-names>
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</contrib>
<contrib contrib-type="author"><name><surname>van Driem</surname>
<given-names>George</given-names>
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<given-names>Stanislav</given-names>
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<given-names>Jean-Michel</given-names>
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<given-names>Sardana A.</given-names>
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<contrib contrib-type="author"><name><surname>Romero</surname>
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<contrib contrib-type="author"><name><surname>Gubina</surname>
<given-names>Marina</given-names>
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<contrib contrib-type="author"><name><surname>Hammer</surname>
<given-names>Michael</given-names>
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<given-names>Elza</given-names>
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<given-names>Rick</given-names>
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<given-names>Toomas</given-names>
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<given-names>Vaidutis</given-names>
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</contrib>
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<given-names>Theologos</given-names>
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</name>
<xref ref-type="aff" rid="A64">65</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Ruizbakiev</surname>
<given-names>Ruslan</given-names>
</name>
<xref ref-type="aff" rid="A65">66</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Sahakyan</surname>
<given-names>Hovhannes</given-names>
</name>
<xref ref-type="aff" rid="A51">51</xref>
<xref ref-type="aff" rid="A66">67</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Sajantila</surname>
<given-names>Antti</given-names>
</name>
<xref ref-type="aff" rid="A67">68</xref>
<xref ref-type="aff" rid="A68">69</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Salas</surname>
<given-names>Antonio</given-names>
</name>
<xref ref-type="aff" rid="A69">70</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Starikovskaya</surname>
<given-names>Elena B.</given-names>
</name>
<xref ref-type="aff" rid="A37">37</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Tarekegn</surname>
<given-names>Ayele</given-names>
</name>
<xref ref-type="aff" rid="A70">71</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Toncheva</surname>
<given-names>Draga</given-names>
</name>
<xref ref-type="aff" rid="A45">45</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Turdikulova</surname>
<given-names>Shahlo</given-names>
</name>
<xref ref-type="aff" rid="A71">72</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Uktveryte</surname>
<given-names>Ingrida</given-names>
</name>
<xref ref-type="aff" rid="A50">50</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Utevska</surname>
<given-names>Olga</given-names>
</name>
<xref ref-type="aff" rid="A72">73</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Vasquez</surname>
<given-names>René</given-names>
</name>
<xref ref-type="aff" rid="A73">74</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Villena</surname>
<given-names>Mercedes</given-names>
</name>
<xref ref-type="aff" rid="A73">74</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Voevoda</surname>
<given-names>Mikhail</given-names>
</name>
<xref ref-type="aff" rid="A35">35</xref>
<xref ref-type="aff" rid="A74">75</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Winkler</surname>
<given-names>Cheryl</given-names>
</name>
<xref ref-type="aff" rid="A75">76</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Yepiskoposyan</surname>
<given-names>Levon</given-names>
</name>
<xref ref-type="aff" rid="A66">67</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Zalloua</surname>
<given-names>Pierre</given-names>
</name>
<xref ref-type="aff" rid="A76">77</xref>
<xref ref-type="aff" rid="A77">78</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Zemunik</surname>
<given-names>Tatijana</given-names>
</name>
<xref ref-type="aff" rid="A78">79</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Cooper</surname>
<given-names>Alan</given-names>
</name>
<xref ref-type="aff" rid="A11">11</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Capelli</surname>
<given-names>Cristian</given-names>
</name>
<xref ref-type="aff" rid="A29">29</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Thomas</surname>
<given-names>Mark G.</given-names>
</name>
<xref ref-type="aff" rid="A79">80</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Ruiz-Linares</surname>
<given-names>Andres</given-names>
</name>
<xref ref-type="aff" rid="A79">80</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Tishkoff</surname>
<given-names>Sarah A.</given-names>
</name>
<xref ref-type="aff" rid="A80">81</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Singh</surname>
<given-names>Lalji</given-names>
</name>
<xref ref-type="aff" rid="A81">82</xref>
<xref rid="FN3" ref-type="author-notes">83</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Thangaraj</surname>
<given-names>Kumarasamy</given-names>
</name>
<xref ref-type="aff" rid="A81">82</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Villems</surname>
<given-names>Richard</given-names>
</name>
<xref ref-type="aff" rid="A51">51</xref>
<xref ref-type="aff" rid="A54">55</xref>
<xref ref-type="aff" rid="A82">84</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Comas</surname>
<given-names>David</given-names>
</name>
<xref ref-type="aff" rid="A83">85</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Sukernik</surname>
<given-names>Rem</given-names>
</name>
<xref ref-type="aff" rid="A37">37</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Metspalu</surname>
<given-names>Mait</given-names>
</name>
<xref ref-type="aff" rid="A51">51</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Meyer</surname>
<given-names>Matthias</given-names>
</name>
<xref ref-type="aff" rid="A4">4</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Eichler</surname>
<given-names>Evan E.</given-names>
</name>
<xref ref-type="aff" rid="A6">6</xref>
<xref ref-type="aff" rid="A84">86</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Burger</surname>
<given-names>Joachim</given-names>
</name>
<xref ref-type="aff" rid="A5">5</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Slatkin</surname>
<given-names>Montgomery</given-names>
</name>
<xref ref-type="aff" rid="A7">7</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Pääbo</surname>
<given-names>Svante</given-names>
</name>
<xref ref-type="aff" rid="A4">4</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Kelso</surname>
<given-names>Janet</given-names>
</name>
<xref ref-type="aff" rid="A4">4</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Reich</surname>
<given-names>David</given-names>
</name>
<xref ref-type="aff" rid="A1">1</xref>
<xref ref-type="aff" rid="A2">2</xref>
<xref ref-type="aff" rid="A85">87</xref>
<xref rid="FN1" ref-type="author-notes">†</xref>
</contrib>
<contrib contrib-type="author"><name><surname>Krause</surname>
<given-names>Johannes</given-names>
</name>
<xref ref-type="aff" rid="A3">3</xref>
<xref ref-type="aff" rid="A86">88</xref>
<xref ref-type="aff" rid="A87">90</xref>
<xref rid="FN1" ref-type="author-notes">†</xref>
</contrib>
</contrib-group>
<aff id="A1"><label>1</label>
Department of Genetics, Harvard Medical School, Boston, MA, 02115, USA</aff>
<aff id="A2"><label>2</label>
Broad Institute of Harvard and MIT, Cambridge, MA, 02142, USA</aff>
<aff id="A3"><label>3</label>
Institute for Archaeological Sciences, University of Tübingen, Tübingen, 72074, Germany</aff>
<aff id="A4"><label>4</label>
Max Planck Institute for Evolutionary Anthropology, Leipzig, 04103, Germany</aff>
<aff id="A5"><label>5</label>
Johannes Gutenberg University Mainz, Institute of Anthropology, Mainz, D-55128, Germany</aff>
<aff id="A6"><label>6</label>
Department of Genome Sciences, University of Washington, Seattle, WA, 98195, USA</aff>
<aff id="A7"><label>7</label>
Department of Integrative Biology, University of California, Berkeley, CA, 94720-3140, USA</aff>
<aff id="A8"><label>8</label>
Department of Mathematics and Computer Science and Artificial Intelligence Laboratory, Massachusetts Institute of Technology, Cambridge, MA 02139, USA</aff>
<aff id="A9"><label>9</label>
Archaeological Research Laboratory, Stockholm University, 114 18, Sweden</aff>
<aff id="A10"><label>10</label>
Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, IVPP, CAS, Beijing, 100049, China</aff>
<aff id="A11"><label>11</label>
Australian Centre for Ancient DNA, School of Earth and Environmental Sciences, University of Adelaide, Adelaide, South Australia, SA 5005, Australia</aff>
<aff id="A12"><label>12</label>
The Cultural Heritage Foundation, Västerås, 722 12, Sweden</aff>
<aff id="A13"><label>13</label>
National Museum of Natural History, L-2160, Luxembourg</aff>
<aff id="A14"><label>14</label>
National Center of Archaeological Research, National Museum of History and Art, L-2345, Luxembourg</aff>
<aff id="A15"><label>15</label>
Department of Paleoanthropology, Senckenberg Center for Human Evolution and Paleoenvironment, University of Tübingen, Tübingen, D-72070, Germany</aff>
<aff id="A16"><label>16</label>
State Office for Cultural Heritage Management Baden-Württemberg, Osteology, Konstanz, D-78467, Germany</aff>
<aff id="A17"><label>17</label>
Center for Global Health and Child Development, Kisumu, 40100, Kenya</aff>
<aff id="A18"><label>18</label>
Institutes of Evolution, Immunology and Infection Research, School of Biological Sciences, University of Edinburgh, Edinburgh, EH9 3JT, UK</aff>
<aff id="A19"><label>19</label>
Biochemistry Department, Faculty of Medicine, Sultan Qaboos University, Alkhod, Muscat, 123, Oman</aff>
<aff id="A20"><label>20</label>
Laboratorio de Genética Molecular Poblacional, Instituto Multidisciplinario de Biología Celular (IMBICE), CCT-CONICET & CICPBA, La Plata, B1906APO, Argentina</aff>
<aff id="A21"><label>21</label>
Research Centre for Medical Genetics, Moscow, 115478, Russia</aff>
<aff id="A22"><label>22</label>
Vavilov Institute for General Genetics, Moscow, 119991, Russia</aff>
<aff id="A23"><label>23</label>
Escuela de Biología, Universidad de Costa Rica, San José, 2060, Costa Rica</aff>
<aff id="A24"><label>24</label>
Institute of Biology, Research group GENMOL, Universidad de Antioquia, Medellín, Colombia</aff>
<aff id="A25"><label>25</label>
Rambam Health Care Campus, Haifa, 31096, Israel</aff>
<aff id="A26"><label>26</label>
Department of Medical Genetics and Szentagothai Research Center, University of Pécs, Pécs, H-7624 Hungary</aff>
<aff id="A27"><label>27</label>
Al Akhawayn University in Ifrane (AUI), School of Science and Engineering, Ifrane, 53000, Morocco</aff>
<aff id="A28"><label>28</label>
Forensic Genetics Laboratory, Institute of Legal Medicine, Università Cattolica del Sacro Cuore, Rome, 00168, Italy</aff>
<aff id="A29"><label>29</label>
Department of Zoology, University of Oxford, Oxford, OX1 3PS, UK</aff>
<aff id="A30"><label>30</label>
Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford, OX3 7BN, UK</aff>
<aff id="A31"><label>31</label>
Laboratorio di Genetica Molecolare, IRCCS Associazione Oasi Maria SS, Troina, 94018, Italy</aff>
<aff id="A32"><label>32</label>
Belgorod State University, Belgorod, 308015, Russia</aff>
<aff id="A33"><label>33</label>
Department of Laboratory Medicine and Pathobiology, University of Toronto, Toronto, Ontario, M5G 1L5, Canada</aff>
<aff id="A34"><label>34</label>
Servicio de Huellas Digitales Genéticas, School of Pharmacy and Biochemistry, Universidad de Buenos Aires, 1113 CABA, Argentina</aff>
<aff id="A35"><label>35</label>
Institute of Cytology and Genetics, Siberian Branch of Russian Academy of Sciences, Novosibirsk, 630090, Russia</aff>
<aff id="A36"><label>36</label>
Institute of Linguistics, University of Bern, Bern, CH-3012, Switzerland</aff>
<aff id="A37"><label>37</label>
Laboratory of Human Molecular Genetics, Institute of Molecular and Cellular Biology, Russian Academy of Science, Siberian Branch, Novosibirsk, 630090, Russia</aff>
<aff id="A38"><label>38</label>
Anthropologie Moléculaire et Imagerie de Synthèse, CNRS UMR 5288, Université Paul Sabatier Toulouse III, Toulouse, 31000, France</aff>
<aff id="A39"><label>39</label>
Yakut Research Center of Complex Medical Problems and North-Eastern Federal University, Yakutsk, 677010, Russia</aff>
<aff id="A40"><label>40</label>
Department of Human Genetics, University of Chicago, Chicago, IL, 60637, USA</aff>
<aff id="A41"><label>41</label>
ARL Division of Biotechnology, University of Arizona, Tucson, AZ, 85721, USA</aff>
<aff id="A42"><label>42</label>
Department of Ecology and Evolution, Stony Brook University, Stony Brook, NY, 11794, USA</aff>
<aff id="A43"><label>43</label>
Department of Clinical Science, University of Bergen, Bergen, 5021, Norway</aff>
<aff id="A44"><label>44</label>
NextBio, part of Illumina, Santa Clara, CA, USA 95050</aff>
<aff id="A45"><label>45</label>
Dept. of Medical Genetics, National Human Genome Center, Medical University Sofia, Sofia, 1431, Bulgaria</aff>
<aff id="A46"><label>46</label>
Institute of Biochemistry and Genetics, Ufa Research Centre, Russian Academy of Sciences, Ufa, 450054, Russia</aff>
<aff id="A47"><label>47</label>
Department of Genetics and Fundamental Medicine, Bashkir State University, Ufa, 450074, Russia</aff>
<aff id="A48"><label>48</label>
College of Medicine, University of Illinois at Chicago, Chicago, IL, 60607, USA</aff>
<aff id="A49"><label>49</label>
Division of Biological Anthropology, University of Cambridge, Cambridge, United Kingdom CB2 1QH</aff>
<aff id="A50"><label>50</label>
Department of Human and Medical Genetics, Vilnius University, Vilnius, LT-08661, Lithuania</aff>
<aff id="A51"><label>51</label>
Estonian Biocentre, Evolutionary Biology group, Tartu, 51010, Estonia</aff>
<aff id="A52"><label>52</label>
Translational Medicine and Neurogenetics, Institut de Génétique et de Biologie Moléculaire et Cellulaire, Illkirch, 67404, France</aff>
<aff id="A53"><label>54</label>
Gladstone Institutes, San Francisco, CA, 94158, USA</aff>
<aff id="A54"><label>55</label>
Department of Evolutionary Biology, University of Tartu, Tartu, 51010, Estonia</aff>
<aff id="A55"><label>56</label>
Centro de Investigaciones Biomédicas de Guatemala, Ciudad de Guatemala, Guatemala</aff>
<aff id="A56"><label>57</label>
Research Department, 23andMe, Inc. Mountain View, CA, 94043, USA</aff>
<aff id="A57"><label>58</label>
Cultural Anthropology Program, University of Oulu, Oulu, 90014, Finland</aff>
<aff id="A58"><label>59</label>
Department of Biochemistry, Muhimbili University of Health and Allied Sciences, Dar es Salaam, Tanzania</aff>
<aff id="A59"><label>60</label>
Research Institute of Health, North-Eastern Federal University, Yakutsk, 677000, Russia</aff>
<aff id="A60"><label>61</label>
Dipartimento di Fisica e Chimica, Università di Palermo, Palermo, 90128, Italy</aff>
<aff id="A61"><label>62</label>
Instituto de Alta Investigación, Universidad de Tarapacá, Arica, Chile</aff>
<aff id="A62"><label>63</label>
Programa de Genética Humana ICBM Facultad de Medicina Universidad de Chile, Santiago, Chile</aff>
<aff id="A63"><label>64</label>
Centro de Investigaciones del Hombre en el Desierto, Arica, Chile</aff>
<aff id="A64"><label>65</label>
Centre for Population Health Sciences, The University of Edinburgh Medical School, Edinburgh, Scotland, EH8 9AG, UK</aff>
<aff id="A65"><label>66</label>
Deceased: formerly of the Institute of Immunology, Academy of Science, Tashkent, 70000, Uzbekistan</aff>
<aff id="A66"><label>67</label>
Laboratory of Ethnogenomics, Institute of Molecular Biology, National Academy of Sciences of Armenia, Yerevan, 0014, Armenia</aff>
<aff id="A67"><label>68</label>
Department of Forensic Medicine, Hjelt Institute, University of Helsinki, Helsinki, 00014, Finland</aff>
<aff id="A68"><label>69</label>
Institute of Applied Genetics, Department of Molecular and Medical Genetics, University of North Texas Health Science Center, Fort Worth, Texas 76107, USA</aff>
<aff id="A69"><label>70</label>
Unidade 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</aff>
<aff id="A70"><label>71</label>
Research Fellow, Henry Stewart Group, Russell House, London WC1A 2HN, UK</aff>
<aff id="A71"><label>72</label>
Institute of Bioorganic Chemistry Academy of Sciences Republic of Uzbekistan, Tashkent, 100125, Uzbekistan</aff>
<aff id="A72"><label>73</label>
Department of Genetics and Cytology, V.N. Karazin Kharkiv National University, Kharkiv, 61077, Ukraine</aff>
<aff id="A73"><label>74</label>
Instituto Boliviano de Biología de la Altura, Universidad Autonoma Tomás Frías, Potosí, Bolivia</aff>
<aff id="A74"><label>75</label>
Inst. of Internal Medicine, Siberian Branch of Russian Acad. of Medical Sciences, Novosibirsk, 630089, Russia</aff>
<aff id="A75"><label>76</label>
Basic Research Laboratory, NCI, NIH, Frederick National Laboratory, Leidos Biomedical, Inc., Frederick, MD 21702, USA</aff>
<aff id="A76"><label>77</label>
Lebanese American University, School of Medicine, Beirut, 13-5053, Lebanon</aff>
<aff id="A77"><label>78</label>
Harvard School of Public Health, Boston, 02115, USA</aff>
<aff id="A78"><label>79</label>
Department of Medical Biology, University of Split, School of Medicine, Split, 21000, Croatia</aff>
<aff id="A79"><label>80</label>
Department of Genetics, Evolution and Environment, University College London, WC1E 6BT, UK</aff>
<aff id="A80"><label>81</label>
Department of Biology and Genetics. University of Pennsylvania, Philadelphia, Pennsylvania, 19104, USA</aff>
<aff id="A81"><label>82</label>
CSIR-Centre for Cellular and Molecular Biology, Hyderabad, 500 007, India</aff>
<aff id="A82"><label>84</label>
Estonian Academy of Sciences, Tallinn, 10130, Estonia</aff>
<aff id="A83"><label>85</label>
Institut de Biologia Evolutiva (CSIC-UPF), Departament de Ciències Experimentals i de la Salut, Universitat Pompeu Fabra, Barcelona, 08003, Spain</aff>
<aff id="A84"><label>86</label>
Howard Hughes Medical Institute, University of Washington, Seattle, WA, USA 98195</aff>
<aff id="A85"><label>87</label>
Howard Hughes Medical Institute, Harvard Medical School, Boston, MA, 02115, USA</aff>
<aff id="A86"><label>88</label>
Senckenberg Centre for Human Evolution and Palaeoenvironment, University of Tübingen, 72070, Germany</aff>
<aff id="A87"><label>90</label>
Max Planck Institut für Geschichte und Naturwissenschaften, Jena, 07745, Germany</aff>
<author-notes><corresp id="FN1"><label>†</label>
Correspondence and requests for materials should be addressed to
David Reich (<email>reich@genetics.med.harvard.edu</email>
) or Johannes Krause
(<email>johannes.krause@uni-tuebingen.de</email>
)</corresp>
<fn id="FN2" fn-type="current-aff"><label>53</label>
<p>Currently employed by AMGEN; 33 Kazantzaki Str, Ilioupolis 16342, Athens, Greece</p>
</fn>
<fn id="FN3" fn-type="present-address"><label>83</label>
<p>Present address: Banaras Hindu University, Varanasi, 221 005, India</p>
</fn>
</author-notes>
<pub-date pub-type="nihms-submitted"><day>6</day>
<month>8</month>
<year>2014</year>
</pub-date>
<pub-date pub-type="ppub"><day>18</day>
<month>9</month>
<year>2014</year>
</pub-date>
<pub-date pub-type="pmc-release"><day>18</day>
<month>3</month>
<year>2015</year>
</pub-date>
<volume>513</volume>
<issue>7518</issue>
<fpage>409</fpage>
<lpage>413</lpage>
<pmc-comment>elocation-id from pubmed: 10.1038/nature13673</pmc-comment>
<permissions><license xlink:href="http://www.nature.com/authors/editorial_policies/license.html#terms"><license-p>Users may view, print, copy, and download text and data-mine the content in such documents, for the purposes of academic research, subject always to the full Conditions of use:<ext-link ext-link-type="uri" xlink:href="http://www.nature.com/authors/editorial_policies/license.html#terms">http://www.nature.com/authors/editorial_policies/license.html#terms</ext-link>
</license-p>
</license>
</permissions>
<abstract><p id="P1">We sequenced the genomes of a ~7,000 year old farmer from Germany and eight
~8,000 year old hunter-gatherers from Luxembourg and Sweden. We analyzed these and other
ancient genomes<sup><xref rid="R1" ref-type="bibr">1</xref>
–<xref rid="R4" ref-type="bibr">4</xref>
</sup>
with 2,345 contemporary humans to show that most
present 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) related to Upper Paleolithic Siberians<sup><xref rid="R3" ref-type="bibr">3</xref>
</sup>
, who 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”
population that split prior to the diversification of other non-African lineages.</p>
</abstract>
</article-meta>
</front>
<body><p id="P2">Near Eastern migrants played a major role in the introduction of agriculture to
Europe, as ancient DNA indicates that early European farmers were distinct from European
hunter-gatherers<sup><xref rid="R4" ref-type="bibr">4</xref>
,<xref rid="R5" ref-type="bibr">5</xref>
</sup>
and close to present-day Near Easterners<sup><xref rid="R4" ref-type="bibr">4</xref>
,<xref rid="R6" ref-type="bibr">6</xref>
</sup>
. However, modelling
present-day Europeans as a mixture of these two ancestral populations<sup><xref rid="R4" ref-type="bibr">4</xref>
</sup>
does not account for the fact that they are also admixed
with a population related to Native Americans<sup><xref rid="R7" ref-type="bibr">7</xref>
,<xref rid="R8" ref-type="bibr">8</xref>
</sup>
. To clarify the prehistory of
Europe, we sequenced nine ancient genomes (<xref rid="F1" ref-type="fig">Fig. 1A</xref>
; <xref rid="F3" ref-type="fig">Extended Data Fig. 1</xref>
): “Stuttgart” (19-fold
coverage), a ~7,000 year old skeleton found in Germany in the context of artifacts from the
first widespread farming culture of central Europe, the <italic>Linearbandkeramik</italic>
;
“Loschbour” (22-fold), an ~8,000 year old skeleton from the Loschbour rock
shelter in Luxembourg, discovered in the context of hunter-gatherer artifacts (SI1; SI2); and
seven ~8,000 year old samples (0.01–2.4-fold) from a hunter-gatherer burial in Motala,
Sweden (the highest coverage individual was “Motala12”).</p>
<p id="P3">Sequence reads from all samples revealed >20% C→T and G→A
deamination-derived mismatches at the ends of the molecules that are characteristic of ancient
DNA<sup><xref rid="R9" ref-type="bibr">9</xref>
,<xref rid="R10" ref-type="bibr">10</xref>
</sup>
(SI3). We estimate nuclear contamination rates to be 0.3% for
Stuttgart and 0.4% for Loschbour (SI3), and mitochondrial (mtDNA) contamination rates
to be 0.3% for Stuttgart, 0.4% for Loschbour, and 0.01–5% for
the Motala individuals (SI3). Stuttgart has mtDNA haplogroup T2, typical of Neolithic
Europeans<sup><xref rid="R11" ref-type="bibr">11</xref>
</sup>
, and Loschbour and all
Motala individuals have the U5 or U2 haplogroups, typical of hunter-gatherers<sup><xref rid="R5" ref-type="bibr">5</xref>
,<xref rid="R9" ref-type="bibr">9</xref>
</sup>
(SI4).
Stuttgart is female, while Loschbour and five Motala individuals are male (SI5) and belong to
Y-chromosome haplogroup I, suggesting that this was common in pre-agricultural Europeans
(SI5).</p>
<p id="P4">We carried out large-scale sequencing of libraries prepared with uracil DNA
glycosylase (UDG), which removes deaminated cytosines, thus reducing errors arising from
ancient DNA damage (SI3). The ancient individuals had indistinguishable levels of Neanderthal
ancestry when compared to each other (~2%) and to present-day Eurasians (SI6). The
heterozygosity of Stuttgart (0.00074) is at the high end of present-day Europeans, while that
of Loschbour (0.00048) is lower than in any present humans (SI2), reflecting a strong
bottleneck in Loschbour’s ancestors as the genetic data show that he was not recently
inbred (<xref rid="F4" ref-type="fig">Extended Data Fig. 2</xref>
). High copy numbers for the
salivary amylase gene (<italic>AMY1)</italic>
have been associated with a high starch
diet<sup><xref rid="R12" ref-type="bibr">12</xref>
</sup>
; our data are consistent with
this finding in that the ancient hunter gatherers La Braña (from Iberia)<sup><xref rid="R2" ref-type="bibr">2</xref>
</sup>
, Motala12, and Loschbour had 5, 6 and 13 copies
respectively, whereas the Stuttgart farmer had 16 (SI7). Both Loschbour and Stuttgart had dark
hair (>99% probability); and Loschbour, like La Braña and Motala12, likely
had blue or intermediate-colored eyes (>75%) while Stuttgart likely had brown eyes
(>99%) (SI8). Neither Loschbour nor La Braña carries the skin-lightening
allele in <italic>SLC24A5</italic>
that is homozygous in Stuttgart and nearly fixed in
Europeans today<sup><xref rid="R2" ref-type="bibr">2</xref>
</sup>
, but Motala12 carries at
least one copy of the derived allele, showing that this allele was present in Europe prior to
the advent of agriculture.</p>
<p id="P5">We compared the ancient genomes to 2,345 present-day humans from 203 populations
genotyped at 594,924 autosomal single nucleotide polymorphisms (SNPs) with the Human Origins
array<sup><xref rid="R8" ref-type="bibr">8</xref>
</sup>
(SI9) (<xref rid="T2" ref-type="table">Extended Data Table 1</xref>
). We used ADMIXTURE<sup><xref rid="R13" ref-type="bibr">13</xref>
</sup>
to identify 59 “West Eurasian” populations
that cluster with Europe and the Near East (SI9 and <xref rid="F5" ref-type="fig">Extended
Data Fig. 3</xref>
). Principal component analysis (PCA)<sup><xref rid="R14" ref-type="bibr">14</xref>
</sup>
(SI10) (<xref rid="F1" ref-type="fig">Fig. 1B</xref>
) indicates a
discontinuity between the Near East and Europe, with each showing north-south clines bridged
only by a few populations of mainly Mediterranean origin. We projected<sup><xref rid="R15" ref-type="bibr">15</xref>
</sup>
the newly sequenced and previously published<sup><xref rid="R1" ref-type="bibr">1</xref>
–<xref rid="R4" ref-type="bibr">4</xref>
</sup>
ancient genomes onto the first two principal components (PCs) (<xref rid="F1" ref-type="fig">Fig. 1B</xref>
). Upper Paleolithic hunter-gatherers<sup><xref rid="R3" ref-type="bibr">3</xref>
</sup>
from Siberia like the MA1 (Mal’ta) individual project at the
northern end of the PCA, suggesting an “Ancient North Eurasian”
meta-population (ANE). European hunter-gatherers from Spain<sup><xref rid="R2" ref-type="bibr">2</xref>
</sup>
, Luxembourg, and Sweden<sup><xref rid="R4" ref-type="bibr">4</xref>
</sup>
fall beyond present-day Europeans in the direction of European differentiation from the Near
East, and form a “West European Hunter-Gatherer” (WHG) cluster including
Loschbour and La Braña<sup><xref rid="R2" ref-type="bibr">2</xref>
</sup>
, and a
“Scandinavian Hunter-Gatherer” (SHG) cluster including the Motala individuals
and ~5,000 year old hunter-gatherers from the Pitted Ware Culture<sup><xref rid="R4" ref-type="bibr">4</xref>
</sup>
. An “Early European Farmer” (EEF) cluster
includes Stuttgart, the ~5,300 year old Tyrolean Iceman<sup><xref rid="R1" ref-type="bibr">1</xref>
</sup>
and a ~5,000 year old Swedish farmer<sup><xref rid="R4" ref-type="bibr">4</xref>
</sup>
.</p>
<p id="P6">Patterns observed in PCA may be affected by sample composition (SI10) and their
interpretation in terms of admixture events is not straightforward, so we rely on formal
analysis of <italic>f</italic>
-statistics<sup><xref rid="R8" ref-type="bibr">8</xref>
</sup>
to
document mixture of at least three source populations in the ancestry of present Europeans. We
began by computing all possible statistics of the form <italic>f<sub>3</sub>
(Test;
Ref<sub>1</sub>
, Ref<sub>2</sub>
)</italic>
(SI11), which if significantly negative show
unambiguously<sup><xref rid="R8" ref-type="bibr">8</xref>
</sup>
that <italic>Test</italic>
is admixed between populations anciently related to <italic>Ref<sub>1</sub>
</italic>
and
<italic>Ref<sub>2</sub>
</italic>
(we choose <italic>Ref<sub>1</sub>
</italic>
and
<italic>Ref<sub>2</sub>
</italic>
from 5 ancient and 192 present populations). The lowest
<italic>f<sub>3</sub>
</italic>
-statistics for Europeans are negative (93% are
>4 standard errors below 0), with most showing strong support for at least one ancient
individual being one of the references (SI11). Europeans almost always have their lowest
<italic>f<sub>3</sub>
</italic>
with either (EEF, ANE) or (WHG, Near East) (SI11, <xref rid="T1" ref-type="table">Table 1</xref>
, <xref rid="T2" ref-type="table">Extended Data
Table 1</xref>
), which would not be expected if there were just two ancient sources of
ancestry (in which case the best references for all Europeans would be similar). The lowest
<italic>f<sub>3</sub>
</italic>
-statistic for Near Easterners always takes Stuttgart as one
of the reference populations, consistent with a Near Eastern origin for Stuttgart’s
ancestors (<xref rid="T1" ref-type="table">Table 1</xref>
). We also computed the statistic
<italic>f<sub>4</sub>
(Test, Stuttgart; MA1, Chimp),</italic>
which measures whether
<italic>MA1</italic>
shares more alleles with a <italic>Test</italic>
population or with
Stuttgart. This statistic is significantly positive (<xref rid="F6" ref-type="fig">Extended
Data Fig. 4</xref>
, <xref rid="T2" ref-type="table">Extended Data Table 1</xref>
) if
<italic>Test</italic>
is nearly any present-day West Eurasian population, showing that
MA1-related ancestry has increased since the time of early farmers like Stuttgart (the
analogous statistic using Native Americans instead of MA1 is correlated but smaller in
magnitude (<xref rid="F7" ref-type="fig">Extended Data Fig. 5</xref>
), indicating that MA1 is
a better surrogate than the Native Americans who were first used to document ANE ancestry in
Europe<sup><xref rid="R7" ref-type="bibr">7</xref>
,<xref rid="R8" ref-type="bibr">8</xref>
</sup>
). The analogous statistic <italic>f<sub>4</sub>
(Test, Stuttgart; Loschbour,
Chimp)</italic>
is nearly always positive in Europeans and negative in Near Easterners,
indicating that Europeans have more ancestry from populations related to Loschbour than do
Near Easterners (<xref rid="F6" ref-type="fig">Extended Data Fig. 4</xref>
, <xref rid="T2" ref-type="table">Extended Data Table 1</xref>
). <xref rid="T3" ref-type="table">Extended
Data Table 2</xref>
documents the robustness of key
<italic>f<sub>4</sub>
-</italic>
statistics by recomputing them using transversion polymorphisms
not affected by ancient DNA damage, and also using whole-genome sequencing data not affected
by SNP ascertainment bias. <xref rid="F8" ref-type="fig">Extended Data Fig. 6</xref>
shows the
geographic gradients in the degree of allele sharing of present-day West Eurasians (as
measured by <italic>f<sub>4</sub>
</italic>
-statistics) with Stuttgart (EEF), Loschbour (WHG)
and MA1 (ANE).</p>
<p id="P7">To determine the minimum number of source populations needed to explain the data for
many European populations taken together, we studied the matrix of all possible statistics of
the form <italic>f<sub>4</sub>
(Test<sub>base</sub>
, Test<sub>i</sub>
; O<sub>base</sub>
,
O<sub>j</sub>
)</italic>
(SI12). <italic>Test<sub>base</sub>
</italic>
is a reference
European population, <italic>Test<sub>i</sub>
</italic>
is the set of all other European
<italic>Test</italic>
populations, <italic>O<sub>base</sub>
</italic>
is a reference
outgroup, and <italic>O<sub>j</sub>
</italic>
is the set of other outgroups (ancient DNA
samples, Onge, Karitiana, and Mbuti). The rank of the <italic>(i, j)</italic>
matrix reflects
the minimum number of sources that contributed to the <italic>Test</italic>
populations<sup><xref rid="R16" ref-type="bibr">16</xref>
,<xref rid="R17" ref-type="bibr">17</xref>
</sup>
. For a pool of individuals from 23 <italic>Test</italic>
populations
representing most present-day European groups, this analysis rejects descent from just two
sources (P<10<sup>−12</sup>
by a Hotelling T-test<sup><xref rid="R17" ref-type="bibr">17</xref>
</sup>
). However, three source populations are consistent with
the data after excluding the Spanish who have evidence for African admixture<sup><xref rid="R18" ref-type="bibr">18</xref>
–<xref rid="R20" ref-type="bibr">20</xref>
</sup>
(P=0.019, not significant after multiple-hypothesis correction),
consistent with the results from ADMIXTURE (SI9), PCA (<xref rid="F1" ref-type="fig">Fig.
1B</xref>
, SI10) and <italic>f</italic>
-statistics (<xref rid="T2" ref-type="table">Extended
Data Table 1</xref>
, <xref rid="F8" ref-type="fig">Extended Data Fig. 6</xref>
, SI11, SI12).
We caution that the finding of three sources could be consistent with a larger number of
mixture events. Moreover, the source populations may themselves have been mixed. Indeed, the
positive <italic>f<sub>4</sub>
(Stuttgart, Test; Loschbour, Chimp)</italic>
statistics obtained
when <italic>Test</italic>
is Near Eastern (<xref rid="T2" ref-type="table">Extended Data
Table 1</xref>
) imply that the EEF had some WHG-related ancestry, which was greater than
0% and as high as 45% (SI13).</p>
<p id="P8">We used the ADMIXTUREGRAPH software<sup><xref rid="R8" ref-type="bibr">8</xref>
,<xref rid="R15" ref-type="bibr">15</xref>
</sup>
to fit a model (a tree structure augmented by
admixture events) to the data, exploring models relating the three ancient populations
(Stuttgart, Loschbour, and MA1) to two eastern non-Africans (Onge and Karitiana) and
sub-Saharan Africans (Mbuti). We found no models that fit the data with 0 or 1 admixture
events, but did find a model that fit with 2 admixture events (SI14). The successful model
(<xref rid="F2" ref-type="fig">Fig. 2A</xref>
) confirms the existence of MA1-related
admixture in Native Americans<sup><xref rid="R3" ref-type="bibr">3</xref>
</sup>
, but includes
the novel inference that Stuttgart is partially (44 ± 10%) derived from a
lineage that split prior to the separation of eastern non-Africans from the common ancestor of
WHG and ANE. The existence of such “Basal Eurasian” admixture into Stuttgart
provides a simple explanation for our finding that diverse eastern non-African populations
share significantly more alleles with ancient European and Upper Paleolithic Siberian
hunter-gatherers than with Stuttgart (that is, <italic>f<sub>4</sub>
(Eastern non-African,
Chimp; Hunter-gatherer, Stuttgart)</italic>
is significantly positive), but that
hunter-gatherers appear to be equally related to most eastern groups (SI14). We verified the
robustness of the model by reanalyzing the data using the unsupervised MixMapper<sup><xref rid="R7" ref-type="bibr">7</xref>
</sup>
(SI15) and TreeMix<sup><xref rid="R21" ref-type="bibr">21</xref>
</sup>
software (SI16), which both identified the same admixture
events. The ANE/WHG split must have occurred >24,000 years ago (as it must predate the age
of MA1<sup><xref rid="R3" ref-type="bibr">3</xref>
</sup>
), and the WHG/Eastern non-African
split must have occurred >40,000 years ago (as it must predate the Tianyuan<sup><xref rid="R22" ref-type="bibr">22</xref>
</sup>
individual from China which clusters with Asians
to the exclusion of Europeans). The Basal Eurasian split must be even older, and might be
related to early settlement of the Levant<sup><xref rid="R23" ref-type="bibr">23</xref>
</sup>
or Arabia<sup><xref rid="R24" ref-type="bibr">24</xref>
,<xref rid="R25" ref-type="bibr">25</xref>
</sup>
prior to the diversification of most Eurasians, or more recent gene flow
from Africa<sup><xref rid="R26" ref-type="bibr">26</xref>
</sup>
. However, the Basal Eurasian
population shares much of the genetic drift common to non-African populations after their
separation from Africans, and thus does not appear to represent gene flow between sub-Saharan
Africans and the ancestors of non-Africans after the out-of-Africa bottleneck (SI14).</p>
<p id="P9">Fitting present-day Europeans into the model, we find that few populations can be fit
as 2-way mixtures, but nearly all are compatible with 3-way mixtures of ANE/EEF/WHG (SI14).
The mixture proportions from the fitted model (<xref rid="F2" ref-type="fig">Fig. 2B</xref>
;
<xref rid="T4" ref-type="table">Extended Data Table 3</xref>
) are encouragingly consistent
with those obtained from a separate method that relates European populations to diverse
outgroups using <italic>f<sub>4</sub>
</italic>
-statistics, assuming only that MA1 is an
unmixed descendent of ANE, Loschbour of WHG, and Stuttgart of EEF (SI17). We infer that EEF
ancestry in Europe today ranges from ~30% in the Baltic region to ~90% in the
Mediterranean, consistent with patterns of identity-by-descent (IBD) sharing<sup><xref rid="R27" ref-type="bibr">27</xref>
,<xref rid="R28" ref-type="bibr">28</xref>
</sup>
(SI18)
and shared haplotype analysis (chromosome painting)<sup><xref rid="R29" ref-type="bibr">29</xref>
</sup>
(SI19) in which Loschbour shares more segments with northern Europeans
and Stuttgart with southern Europeans. Southern Europeans inherited their European
hunter-gatherer ancestry mostly via EEF ancestors (<xref rid="F8" ref-type="fig">Extended Data
Fig. 6</xref>
), while Northern Europeans acquired up to 50% of WHG ancestry above
and beyond the WHG-related ancestry which they received through their EEF ancestors. Europeans
have a larger proportion of WHG than ANE ancestry in general. By contrast, in the Near East
there is no detectable WHG ancestry, but up to ~29% ANE in the Caucasus (SI14). A
striking feature of these findings is that ANE ancestry is inferred to be present in nearly
all Europeans today (with a maximum of ~20%), but was absent in both farmers and
hunter-gatherers from central/western Europe during the Neolithic transition. At the same
time, we infer that ANE ancestry was not completely absent from the larger European region at
that time: we find that it was present in ~8,000 years old Scandinavian hunter-gatherers,
since MA1 shares more alleles with Motala12 (SHG) than with Loschbour, and Motala12 fits as a
mixture of 81% WHG and 19% ANE (SI14).</p>
<p id="P10">Two sets of European populations are poor fits for the model. Sicilians, Maltese,
and Ashkenazi Jews have EEF estimates of >100% consistent with their having more
Near Eastern ancestry than can be explained via EEF admixture (SI17). They also cannot be
jointly fit with other Europeans (SI14), and they fall in the gap between European and Near
Easterners (<xref rid="F1" ref-type="fig">Fig. 1B</xref>
). Finns, Mordovians and Russians
(from the northwest of Russia) also do not fit (SI14; <xref rid="T4" ref-type="table">Extended
Data Table 3</xref>
) due to East Eurasian gene flow into the ancestors of these northeastern
European populations. These populations (and Chuvash and Saami) are more related to East
Asians than can be explained by ANE admixture (<xref rid="F9" ref-type="fig">Extended Data
Fig. 7</xref>
), likely reflecting a separate stream of Siberian gene flow into northeastern
Europe (SI14).</p>
<p id="P11">Several questions will be important to address in future ancient DNA work. Where and
when did the Near Eastern farmers admix with European hunter-gatherers to produce the EEF? How
did the ancestors of present-day Europeans first acquire their ANE ancestry? Discontinuity in
central Europe during the late Neolithic (~4,500 years ago) associated with the appearance of
mtDNA types absent in earlier farmers and hunter-gatherers<sup><xref rid="R30" ref-type="bibr">30</xref>
</sup>
raises the possibility that ANE ancestry may have also appeared at this
time. Finally, it is important to study ancient genome sequences from the Near East to provide
insights into the history of the Basal Eurasians.</p>
<sec id="S1" specific-use="web-only"><title>Online Methods</title>
<sec id="S2"><title>Archeological context, sampling and DNA extraction</title>
<p id="P12">The Loschbour sample stems from a male skeleton excavated in 1935 at the
Loschbour rock shelter in Heffingen, Luxembourg. The skeleton was AMS radiocarbon dated to
7,205 ± 50 years before present (OxA-7738; 6,220-5,990 cal BC)<sup><xref rid="R31" ref-type="bibr">31</xref>
</sup>
. At the Palaeogenetics Laboratory in Mainz, material
for DNA extraction was sampled from tooth 16 (an upper right M1 molar) after irradiation
with UV-light, surface removal, and pulverization in a mixer mill. DNA extraction took
place in the palaeogenetics facilities in the Institute for Archaeological Sciences at the
University of Tübingen. Three extracts were made in total, one from 80 mg of
powder using an established silica based protocol<sup><xref rid="R32" ref-type="bibr">32</xref>
</sup>
and two additional extracts from 90 mg of powder each with a protocol
optimized for the recovery of short DNA molecules<sup><xref rid="R33" ref-type="bibr">33</xref>
</sup>
.</p>
<p id="P13">The Stuttgart sample was taken from a female skeleton excavated in 1982 at the
site Viesenhäuser Hof, Stuttgart-Mühlhausen, Germany. It was attributed to
the Linearbandkeramik (5,500-4,800 BC) through associated pottery artifacts and the
chronology was corroborated by radiocarbon dating of the stratigraphy<sup><xref rid="R34" ref-type="bibr">34</xref>
</sup>
. Both sampling and DNA extraction took place in the
Institute for Archaeological Sciences at the University of Tübingen. Tooth 47 (a
lower right M2 molar) was removed and material from the inner part was sampled with a
sterile dentistry drill. An extract was made using 40 mg of bone powder<sup><xref rid="R33" ref-type="bibr">33</xref>
</sup>
.</p>
<p id="P14">The Motala individuals were recovered from the site of Kanaljorden in the town
of Motala, Östergötland, Sweden, excavated between 2009 and 2013. The
human remains at this site are represented by several adult skulls and one infant
skeleton. All individuals are part of a ritual deposition at the bottom of a small lake.
Direct radiocarbon dates on the remains range between 7,013 ± 76 and 6,701
± 64 BP (6,361-5,516 cal BC), corresponding to the late Middle Mesolithic of
Scandinavia. Samples were taken from the teeth of the nine best preserved skulls, as well
as a femur and tibia. Bone powder was removed from the inner parts of the teeth or bones
with a sterile dentistry drill. DNA from 100 mg of bone powder was extracted<sup><xref rid="R35" ref-type="bibr">35</xref>
</sup>
in the ancient DNA laboratory of the
Archaeological Research Laboratory, Stockholm.</p>
</sec>
<sec id="S3"><title>Library preparation</title>
<p id="P15">Illumina sequencing libraries were prepared using either double- or
single-stranded library preparation protocols<sup><xref rid="R36" ref-type="bibr">36</xref>
,<xref rid="R37" ref-type="bibr">37</xref>
</sup>
(SI1). For high-coverage
shotgun sequencing libraries, a DNA repair step with Uracil-DNA-glycosylase (UDG) and
endonuclease VIII (endo VIII) treatment was included in order to remove uracil
residues<sup><xref rid="R38" ref-type="bibr">38</xref>
</sup>
. Size fractionation on a
PAGE gel was also performed in order to remove longer DNA molecules that are more likely
to be contaminants<sup><xref rid="R37" ref-type="bibr">37</xref>
</sup>
. Positive and blank
controls were carried along during every step of library preparation.</p>
</sec>
<sec id="S4"><title>Shotgun sequencing and read processing</title>
<p id="P16">All non-UDG-treated libraries were sequenced either on an Illumina Genome
Analyzer IIx with 2×76 + 7 cycles for the Loschbour and Motala libraries,
or on an Illumina MiSeq with 2×150 + 8 + 8 cycles for the
Stuttgart library. We followed the manufacturer’s protocol for multiplex
sequencing. Raw overlapping forward and reverse reads were merged and filtered for
quality<sup><xref rid="R39" ref-type="bibr">39</xref>
</sup>
and mapped to the human
reference genome (hg19/GRCh37/1000Genomes) using the Burrows-Wheeler Aligner
(BWA)<sup><xref rid="R40" ref-type="bibr">40</xref>
</sup>
(SI2). For deeper
sequencing, UDG-treated libraries of Loschbour were sequenced on 3 Illumina HiSeq 2000
lanes with 50-bp single-end reads, 8 Illumina HiSeq 2000 lanes of 100-bp paired-end reads
and 8 Illumina HiSeq 2500 lanes of 101-bp paired-end reads. The UDG-treated library for
Stuttgart was sequenced on 8 HiSeq 2000 lanes and 101-bp paired-end reads. The UDG-treated
libraries for Motala were sequenced on 8 HiSeq 2000 lanes of 100-bp paired-end reads, with
4 lanes each for two pools (one of 3 individuals and one of 4 individuals). We also
sequenced an additional 8 HiSeq 2000 lanes for Motala12, the Motala sample with the
highest percentage of endogenous human DNA. For the Loschbour and Stuttgart high coverage
individuals, diploid genotype calls were obtained using the Genome Analysis Toolkit
(GATK)<sup><xref rid="R41" ref-type="bibr">41</xref>
</sup>
.</p>
</sec>
<sec id="S5"><title>Enrichment of mitochondrial DNA and sequencing</title>
<p id="P17">To test for DNA preservation and mtDNA contamination non-UDG-treated libraries
of Loschbour and all Motala samples were enriched for human mitochondrial DNA using a
bead-based capture approach with present-day human DNA as bait<sup><xref rid="R42" ref-type="bibr">42</xref>
</sup>
. UDG-treatment was omitted in order to allow
characterization of damage patterns typical for ancient DNA<sup><xref rid="R10" ref-type="bibr">10</xref>
</sup>
. The captured libraries were sequenced on an Illumina
Genome Analyzer IIx platform with 2 × 76 + 7 cycles and the resulting
reads were merged and quality filtered<sup><xref rid="R39" ref-type="bibr">39</xref>
</sup>
. The sequences were mapped to the Reconstructed Sapiens Reference
Sequence, RSRS<sup><xref rid="R43" ref-type="bibr">43</xref>
</sup>
, using a custom
iterative mapping assembler, MIA<sup><xref rid="R44" ref-type="bibr">44</xref>
</sup>
(SI4).</p>
</sec>
<sec id="S6"><title>Contamination estimates</title>
<p id="P18">We assessed if the sequences had the characteristics of authentic ancient DNA
using four approaches. First we searched for evidence of contamination by determining
whether the sequences mapping to the mitochondrial genome were consistent with deriving
from more than one individual<sup><xref rid="R44" ref-type="bibr">44</xref>
,<xref rid="R45" ref-type="bibr">45</xref>
</sup>
. Second, for the high-coverage Loschbour and
Stuttgart genomes, we used a maximum-likelihood-based estimate of autosomal contamination
that uses variation at sites that are fixed in the 1000 Genomes data to estimate error,
heterozygosity and contamination<sup><xref rid="R46" ref-type="bibr">46</xref>
</sup>
simultaneously. Third, we estimated contamination based on the rate of polymorphic sites
on the X chromosome of the male Loschbour individual<sup><xref rid="R47" ref-type="bibr">47</xref>
</sup>
(SI3) Fourth, we analyzed non-UDG treated reads mapping to the RSRS
to search for aDNA-typical damage patterns resulting in C→T changes at the
5′-end of the molecule<sup><xref rid="R10" ref-type="bibr">10</xref>
</sup>
(SI3).</p>
</sec>
<sec id="S7"><title>Phylogenetic analysis of the mitochondrial genomes</title>
<p id="P19">All nine complete mitochondrial genomes that fulfilled the criteria of
authenticity were assigned to haplogroups using Haplofind<sup><xref rid="R48" ref-type="bibr">48</xref>
</sup>
. A Maximum Parsimony tree including present day humans
and previously published ancient mtDNA sequences was generated with MEGA<sup><xref rid="R49" ref-type="bibr">49</xref>
</sup>
. The effect of branch shortening due to a
lower number of substitutions in ancient lineages was studied by calculating the
nucleotide edit distance to the root for all haplogroup R sequences (SI4).</p>
</sec>
<sec id="S8"><title>Sex determination and Y-chromosome analysis</title>
<p id="P20">We assessed the sex of all sequenced individuals by using the ratio of (chrY) to
(chrY+chrX) aligned reads<sup><xref rid="R50" ref-type="bibr">50</xref>
</sup>
. We
downloaded a list of Y-chromosome SNPs curated by the International Society of Genetic
Genealogy (ISOGG, <ext-link ext-link-type="uri" xlink:href="http://www.isogg.org">http://www.isogg.org</ext-link>
) v. 9.22 (accessed Feb. 18, 2014) and determined the
state of the ancient individuals at positions where a single allele was observed and
MAPQ≥30. We excluded C/G or A/T SNPs due to uncertainty about the polarity of the
mutation in the database. The ancient individuals were assigned haplogroups based on their
derived state (SI5). We also used BEAST v1.7.51<sup><xref rid="R51" ref-type="bibr">51</xref>
</sup>
to assess the phylogenetic position of Loschbour using 623 males from
around the world with 2,799 variant sites across 500kb of non-recombining Y-chromosome
sequence<sup><xref rid="R52" ref-type="bibr">52</xref>
</sup>
(SI5).</p>
</sec>
<sec id="S9"><title>Estimation of Neanderthal admixture</title>
<p id="P21">We estimate Neanderthal admixture in ancient individuals with the
<italic>f<sub>4</sub>
-</italic>
ratio or <italic>S</italic>
-statistic<sup><xref rid="R8" ref-type="bibr">8</xref>
,<xref rid="R53" ref-type="bibr">53</xref>
,<xref rid="R54" ref-type="bibr">54</xref>
</sup>
<italic>α̂</italic>
=
<italic>f<sub>4</sub>
</italic>
(<italic>Altai</italic>
, <italic>Denisova; Test</italic>
,
<italic>Yoruba</italic>
)/<italic>f<sub>4</sub>
</italic>
(<italic>Altai</italic>
,
<italic>Denisova; Vindija</italic>
, <italic>Yoruba</italic>
) which uses whole genome
data from Altai, a high coverage (52×) Neanderthal genome sequence<sup><xref rid="R55" ref-type="bibr">55</xref>
</sup>
, Denisova, a high coverage
sequence<sup><xref rid="R37" ref-type="bibr">37</xref>
</sup>
from another archaic
human population (31×), and Vindija, a low coverage (1.3×) Neanderthal
genome from a mixture of three Neanderthal individuals from Vindija Cave in
Croatia<sup><xref rid="R53" ref-type="bibr">53</xref>
</sup>
.</p>
</sec>
<sec id="S10" sec-type="background"><title>Inference of demographic history and inbreeding</title>
<p id="P22">We used the Pairwise Sequentially Markovian Coalescent (PSMC)<sup><xref rid="R56" ref-type="bibr">56</xref>
</sup>
to infer the size of the ancestral
population of Stuttgart and Loschbour. This analysis requires high quality diploid
genotype calls and cannot be performed in the low-coverage Motala samples. To determine
whether the low effective population size inferred for Loschbour is due to recent
inbreeding, we plotted the time-to-most-recent common ancestor (TMRCA) along each of
chr1-22 to detect runs of low TMRCA.</p>
</sec>
<sec id="S11"><title>Analysis of segmental duplications and copy number variants</title>
<p id="P23">We built read-depth based copy number maps for the Loschbour, Stuttgart and
Motala12 genomes in addition to the Denisova and Altai Neanderthal genome and 25 deeply
sequenced modern genomes<sup><xref rid="R55" ref-type="bibr">55</xref>
</sup>
(SI7). We
built these maps by aligning reads, subdivided into their non-overlapping 36-bp
constituents, against the reference genome using the mrsFAST aligner<sup><xref rid="R57" ref-type="bibr">57</xref>
</sup>
, and renormalizing read-depth for local GC content. We
estimated copy numbers in windows of 500 unmasked base pairs slid at 100 bp intervals
across the genome. We called copy number variants using a scale space filter algorithm. We
genotyped variants of interest and compared the genotypes to those from individuals
sequenced as part of the 1000 Genomes Project<sup><xref rid="R58" ref-type="bibr">58</xref>
</sup>
.</p>
</sec>
<sec id="S12"><title>Phenotypic inference</title>
<p id="P24">We inferred likely phenotypes (SI8) by analyzing DNA polymorphism data in the
VCF format<sup><xref rid="R59" ref-type="bibr">59</xref>
</sup>
using VCFtools (<ext-link ext-link-type="uri" xlink:href="http://vcftoools.sourceforge.net/">http://vcftoools.sourceforge.net/</ext-link>
). For the Loschbour and Stuttgart
individuals, we included data from sites not flagged as LowQuality, with genotype quality
(GQ) of ≥30, and SNP quality (QUAL) of ≥50. For Motala12, which is of
lower coverage, we included sites having at least 2× coverage and that passed
visual inspection of the local alignment using samtools tview (<ext-link ext-link-type="uri" xlink:href="http://samtools.sourceforge.net">http://samtools.sourceforge.net</ext-link>
)<sup><xref rid="R60" ref-type="bibr">60</xref>
</sup>
</p>
</sec>
<sec id="S13"><title>Human Origins dataset curation</title>
<p id="P25">The Human Origins array consists of 14 panels of SNPs for which the
ascertainment is well known<sup><xref rid="R8" ref-type="bibr">8</xref>
,<xref rid="R61" ref-type="bibr">61</xref>
</sup>
. All population genetics analysis were carried out on
a set of 594,924 autosomal SNPs, after restricting to sites that had >90%
completeness across 7 different batches of sequencing, and that had >97.5%
concordance with at least one of two subsets of samples for which whole genome sequencing
data was also available. The total dataset consists of 2,722 individuals, which we
filtered to 2,345 individuals (203 populations) after removing outlier individuals or
relatives based on visual inspection of PCA plots<sup><xref rid="R14" ref-type="bibr">14</xref>
,<xref rid="R62" ref-type="bibr">62</xref>
</sup>
or model-based clustering
analysis<sup><xref rid="R13" ref-type="bibr">13</xref>
</sup>
. Whole genome amplified
(WGA) individuals were not used in analysis, except for a Saami individual who we included
because of the special interest of this population for Northeastern European population
history (<xref rid="F9" ref-type="fig">Extended Data Fig. 7</xref>
).</p>
</sec>
<sec id="S14"><title>ADMIXTURE analysis</title>
<p id="P26">We merged all Human Origins genotype data with whole genome sequencing data from
Loschbour, Stuttgart, MA1, Motala12, Motala_merge, and LaBrana. We then thinned the
resulting dataset to remove SNPs in linkage-disequilibrium with PLINK 1.07<sup><xref rid="R63" ref-type="bibr">63</xref>
</sup>
, using a window size of 200 SNPs advanced by
25 SNPs and an r<sup><xref rid="R2" ref-type="bibr">2</xref>
</sup>
threshold of 0.4. We
ran ADMIXTURE 1.23<sup><xref rid="R13" ref-type="bibr">13</xref>
,<xref rid="R64" ref-type="bibr">64</xref>
</sup>
for 100 replicates with different starting random
seeds, default 5-fold cross-validation, and varying the number of ancestral populations K
between 2 and 20. We assessed clustering quality using CLUMPP<sup><xref rid="R65" ref-type="bibr">65</xref>
</sup>
. We used the ADMIXTURE results to identify a set of 59
“West Eurasian” (European/Near Eastern) populations based on values of a
“West Eurasian” ancestral population at K=3 (SI9). We also
identified 15 populations for use as “non-West Eurasian outgroups” based
on their having at least 10 individuals and no evidence of European or Near Eastern
admixture at K=11, the lowest K for which Near Eastern/European-maximized
ancestral populations appeared consistently across all 100 replicates.</p>
</sec>
<sec id="S15"><title>Principal Components Analysis</title>
<p id="P27">We used <italic>smartpca</italic>
<sup><xref rid="R14" ref-type="bibr">14</xref>
</sup>
(version: 10210) from EIGENSOFT<sup><xref rid="R62" ref-type="bibr">62</xref>
,<xref rid="R66" ref-type="bibr">66</xref>
</sup>
5.0.1 to carry out
Principal Components Analysis (PCA) (SI10). We performed PCA on a subset on individuals
and then projected others using the <italic>lsqproject: YES</italic>
option that gives an
unbiased inference of the position of samples even in the presence of missing data
(especially important for ancient DNA).</p>
</sec>
<sec id="S16"><title><italic>f<sub>3</sub>
</italic>
-statistics</title>
<p id="P28">We use the <italic>f<sub>3</sub>
</italic>
-statistic<sup>8</sup>
<inline-formula><mml:math id="M1" overflow="scroll"><mml:msub><mml:mi>f</mml:mi>
<mml:mn>3</mml:mn>
</mml:msub>
<mml:mo stretchy="false">(</mml:mo>
<mml:mi mathvariant="italic">Test</mml:mi>
<mml:mo>;</mml:mo>
<mml:mi>R</mml:mi>
<mml:mi>e</mml:mi>
<mml:msub><mml:mi>f</mml:mi>
<mml:mn>1</mml:mn>
</mml:msub>
<mml:mi>R</mml:mi>
<mml:mi>e</mml:mi>
<mml:msub><mml:mi>f</mml:mi>
<mml:mn>2</mml:mn>
</mml:msub>
<mml:mo stretchy="false">)</mml:mo>
<mml:mo>=</mml:mo>
<mml:mstyle scriptlevel="1"><mml:mfrac><mml:mn>1</mml:mn>
<mml:mi>N</mml:mi>
</mml:mfrac>
</mml:mstyle>
<mml:msubsup><mml:mo>∑</mml:mo>
<mml:mrow><mml:mi>i</mml:mi>
<mml:mo>=</mml:mo>
<mml:mn>1</mml:mn>
</mml:mrow>
<mml:mi>N</mml:mi>
</mml:msubsup>
<mml:mrow><mml:mo stretchy="false">(</mml:mo>
<mml:msub><mml:mi>t</mml:mi>
<mml:mi>i</mml:mi>
</mml:msub>
<mml:mo>-</mml:mo>
<mml:msub><mml:mi>r</mml:mi>
<mml:mrow><mml:mn>1</mml:mn>
<mml:mo>,</mml:mo>
<mml:mi>i</mml:mi>
</mml:mrow>
</mml:msub>
<mml:mo stretchy="false">)</mml:mo>
<mml:mspace width="0.16667em"></mml:mspace>
<mml:mo stretchy="false">(</mml:mo>
<mml:msub><mml:mi>t</mml:mi>
<mml:mi>i</mml:mi>
</mml:msub>
<mml:mo>-</mml:mo>
<mml:msub><mml:mi>r</mml:mi>
<mml:mrow><mml:mn>2</mml:mn>
<mml:mo>,</mml:mo>
<mml:mi>i</mml:mi>
</mml:mrow>
</mml:msub>
<mml:mo stretchy="false">)</mml:mo>
</mml:mrow>
</mml:math>
</inline-formula>
, where <italic>t<sub>i</sub>
</italic>
,
<italic>r</italic>
<sub>1,i</sub>
and <italic>r</italic>
<sub>2,i</sub>
are the allele
frequencies for the <italic>i</italic>
<sup>th</sup>
SNP in populations <italic>Test,
Ref</italic>
<sub>1</sub>
<italic>, Ref</italic>
<sub>2</sub>
<italic>,</italic>
respectively, to determine if there is evidence that the <italic>Test</italic>
population
is derived from admixture of populations related to <italic>Ref<sub>1</sub>
</italic>
and
<italic>Ref<sub>2</sub>
</italic>
(SI11). A significantly negative statistic provides
unambiguous evidence of mixture in the <italic>Test</italic>
population<sup><xref rid="R8" ref-type="bibr">8</xref>
</sup>
. We allow <italic>Ref<sub>1</sub>
</italic>
and
<italic>Ref<sub>2</sub>
</italic>
to be any Human Origins population with 4 or more
individuals, or Loschbour, Stuttgart, MA1, Motala12, LaBrana. We assess significance of
the <italic>f<sub>3</sub>
</italic>
-statistics using a block jackknife<sup><xref rid="R67" ref-type="bibr">67</xref>
</sup>
and a block size of 5cM. We report significance as the
number of standard errors by which the statistic differs from zero (Z-score). We also
perform an analysis in which we constrain the reference populations to be (i) EEF
(Stuttgart) and WHG (Loschbour or LaBrana), (ii) EEF and a Near Eastern population, (iii)
EEF and ANE (MA1), or (iv) any two present-day populations, and compute a Z<sub>diff</sub>
score between the lowest <italic>f<sub>3</sub>
</italic>
-statistic observed in the dataset,
and the <italic>f<sub>3</sub>
-</italic>
statistic observed for the specified pair.</p>
</sec>
<sec id="S17"><title><italic>f<sub>4</sub>
</italic>
-statistics</title>
<p id="P29">We analyze <italic>f<sub>4</sub>
</italic>
-statistics<sup><xref rid="R8" ref-type="bibr">8</xref>
</sup>
of the form <inline-formula><mml:math id="M2" overflow="scroll"><mml:msub><mml:mi>f</mml:mi>
<mml:mn>4</mml:mn>
</mml:msub>
<mml:mo stretchy="false">(</mml:mo>
<mml:mi>A</mml:mi>
<mml:mo>,</mml:mo>
<mml:mi>B</mml:mi>
<mml:mo>;</mml:mo>
<mml:mi>C</mml:mi>
<mml:mo>,</mml:mo>
<mml:mi>D</mml:mi>
<mml:mo stretchy="false">)</mml:mo>
<mml:mo>=</mml:mo>
<mml:mstyle scriptlevel="1"><mml:mfrac><mml:mn>1</mml:mn>
<mml:mi>N</mml:mi>
</mml:mfrac>
</mml:mstyle>
<mml:msubsup><mml:mo>∑</mml:mo>
<mml:mrow><mml:mi>i</mml:mi>
<mml:mo>=</mml:mo>
<mml:mn>1</mml:mn>
</mml:mrow>
<mml:mi>N</mml:mi>
</mml:msubsup>
<mml:mrow><mml:mo stretchy="false">(</mml:mo>
<mml:msub><mml:mi>a</mml:mi>
<mml:mi>i</mml:mi>
</mml:msub>
<mml:mo>-</mml:mo>
<mml:msub><mml:mi>b</mml:mi>
<mml:mi>i</mml:mi>
</mml:msub>
<mml:mo stretchy="false">)</mml:mo>
<mml:mspace width="0.16667em"></mml:mspace>
<mml:mo stretchy="false">(</mml:mo>
<mml:msub><mml:mi>c</mml:mi>
<mml:mi>i</mml:mi>
</mml:msub>
<mml:mo>-</mml:mo>
<mml:msub><mml:mi>d</mml:mi>
<mml:mi>i</mml:mi>
</mml:msub>
<mml:mo stretchy="false">)</mml:mo>
</mml:mrow>
</mml:math>
</inline-formula>
to assess if populations A, B are consistent with forming a
clade in an unrooted tree with respect to C, D. If they form a clade, the allele frequency
differences between the two pairs should be uncorrelated and the statistic has an expected
value of 0. We set the outgroup <italic>D</italic>
to be a sub-Saharan African population
or Chimpanzee. We systematically tried all possible combinations of the ancient samples or
15 “non-West Eurasian outgroups” identified by ADMIXTURE analysis as A, B,
C to determine their genetic affinities (SI14). Setting A as a present-day test population
and B as either Stuttgart or BedouinB, we documented relatedness to C=(Loschbour
or MA1) or C=(MA1 and Karitiana) or C=(MA1 or Han) (<xref rid="F6" ref-type="fig">Extended Data Figs. 4</xref>
, <xref rid="F7" ref-type="fig">5</xref>
,
<xref rid="F9" ref-type="fig">7</xref>
). Setting C as a test population and (A, B) a
pair from (Loschbour, Stuttgart, MA1) we documented differential relatedness to ancient
populations (<xref rid="F8" ref-type="fig">Extended Data Fig. 6</xref>
). We computed
<italic>D</italic>
-statistics<sup><xref rid="R53" ref-type="bibr">53</xref>
</sup>
using
transversion polymorphisms in whole genome sequence data<sup><xref rid="R55" ref-type="bibr">55</xref>
</sup>
to confirm robustness to ascertainment and ancient DNA
damage (<xref rid="T3" ref-type="table">Extended Data Table 2</xref>
).</p>
</sec>
<sec id="S18"><title>Minimum number of source populations for Europeans</title>
<p id="P30">We used <italic>qpWave</italic>
<sup><xref rid="R16" ref-type="bibr">16</xref>
,<xref rid="R17" ref-type="bibr">17</xref>
</sup>
to study the minimum number
of source populations for a designated set of Europeans (SI12). We use
<italic>f<sub>4</sub>
</italic>
-statistics of the form <italic>X(l, r) =
f<sub>4</sub>
(l<sub>0</sub>
, l; r<sub>0</sub>
, r)</italic>
where
<italic>l<sub>0</sub>
</italic>
,<italic>r<sub>0</sub>
</italic>
are arbitrarily chosen
“base” populations, and <italic>l</italic>
, <italic>r</italic>
are other
populations from two sets <italic>L</italic>
and <italic>R</italic>
respectively. If
<italic>X(l, r)</italic>
has rank <italic>r</italic>
and there were <italic>n</italic>
waves of immigration into <italic>R</italic>
with no back-migration from
<italic>R</italic>
to <italic>L</italic>
, then <italic>r+1</italic>
≤
<italic>n</italic>
. We set <italic>L</italic>
to include <italic>Stuttgart, Loschbour,
MA1, Onge, Karitiana, Mbuti</italic>
and <italic>R</italic>
to include 23 modern
European populations who fit the model of SI14 and had admixture proportions within the
interval [0,1] for the method with minimal modeling assumptions
(SI17).</p>
</sec>
<sec id="S19"><title>Admixture proportions for Stuttgart in the absence of a Near Eastern ancient
genome</title>
<p id="P31">We used Loschbour and BedouinB as surrogates for “Unknown
hunter-gatherer” and Near Eastern (NE) farmer populations that contributed to
Stuttgart (SI13). Ancient Near Eastern ancestry in Stuttgart is estimated by the
<italic>f<sub>4</sub>
</italic>
-ratio<sup><xref rid="R8" ref-type="bibr">8</xref>
,<xref rid="R15" ref-type="bibr">15</xref>
</sup>
<italic>f<sub>4</sub>
(Outgroup, X; Loschbour, Stuttgart)/f<sub>4</sub>
(Outgroup, X;
Loschbour, NE)</italic>
. A complication is that BedouinB is a mixture of NE and African
ancestry. We therefore subtracted<sup><xref rid="R17" ref-type="bibr">17</xref>
</sup>
the
effects of African ancestry using estimates of the BedouinB African admixture proportion
from ADMIXTURE (SI9) or ALDER<sup><xref rid="R68" ref-type="bibr">68</xref>
</sup>
.</p>
</sec>
<sec id="S20"><title>Admixture graph modeling</title>
<p id="P32">We used ADMIXTUREGRAPH<sup><xref rid="R8" ref-type="bibr">8</xref>
</sup>
(version 3110) to model population relationships between Loschbour, Stuttgart, Onge, and
Karitiana using Mbuti as an African outgroup. We assessed model fit using a block
jackknife of differences between estimated and fitted <italic>f</italic>
-statistics for
the set of included populations (we expressed the fit as a Z score). We determined that a
model failed if |Z|>3 for at least one <italic>f</italic>
-statistic. A basic tree model
failed and we manually amended the model to test all possible models with a single
admixture event, which also failed. Further manual amendment to include 2 admixture events
resulted in 8 successful models, only one of which could be amended to also fit MA1 as an
additional constraint. We successfully fit both the Iceman and LaBrana into this model as
simple clades and Motala12 as a 2-way mixture. We also fit present-day West Eurasians as
clades, 2-way mixtures, or 3-way mixtures in this basic model, achieving a successful fit
for a larger number of European populations (n=26) as 3-way mixtures. We estimated
the individual admixture proportions from the fitted model parameters. To test if fitted
parameters for different populations are consistent with each other, we jointly fit all
pairs of populations <italic>A</italic>
and <italic>B</italic>
by modifying ADMIXTUREGRAPH
to add a large constant (10,000) to the variance term <italic>f<sub>3</sub>
(A<sub>0</sub>
,
A, B).</italic>
By doing this, we can safely ignore recent gene flow within Europe that
affects statistics that include both <italic>A</italic>
and <italic>B</italic>
.</p>
</sec>
<sec id="S21"><title>Ancestry estimates from <italic>f<sub>4</sub>
</italic>
-ratios</title>
<p id="P33">We estimate EEF ancestry using the
<italic>f<sub>4</sub>
</italic>
-ratio<sup><xref rid="R8" ref-type="bibr">8</xref>
,<xref rid="R15" ref-type="bibr">15</xref>
</sup>
<italic>f<sub>4</sub>
(Mbuti, Onge; Loschbour,
European)</italic>
/<italic>f<sub>4</sub>
(Mbuti, Onge; Loschbour, Stuttgart),</italic>
which produces consistent results with ADMIXTUREGRAPH (SI14). We use
<italic>f<sub>4</sub>
(Stuttgart, Loschbour; Onge
MA1)</italic>
/<italic>f<sub>4</sub>
(Mbuti, MA1; Onge, Loschbour)</italic>
to estimate
Basal Eurasian admixture into Stuttgart. We use <italic>f<sub>4</sub>
(Stuttgart,
Loschbour; Onge Karitiana)</italic>
/<italic>f<sub>4</sub>
(Stuttgart, Loschbour; Onge
MA1)</italic>
to estimate ANE mixture in Karitiana (<xref rid="F2" ref-type="fig">Fig.
2B</xref>
). We use <italic>f<sub>4</sub>
(Test, Stuttgart; Karitiana,
Onge)</italic>
/<italic>f<sub>4</sub>
(MA1, Stuttgart; Karitiana, Onge)</italic>
to
lower bound ANE mixture into North Caucasian populations.</p>
</sec>
<sec id="S22"><title><italic>MixMapper</italic>
analysis</title>
<p id="P34">We carried out <italic>MixMapper</italic>
2.0<sup><xref rid="R7" ref-type="bibr">7</xref>
</sup>
analysis, a semi-supervised admixture graph fitting technique. First,
we infer a scaffold tree of populations without strong evidence of mixture relative to
each other (Mbuti, Onge, Loschbour and MA1). We do not include European populations in the
scaffold as all had significantly negative <italic>f<sub>3</sub>
</italic>
-statistics
indicating admixture. We then ran <italic>MixMapper</italic>
to infer the relatedness of
the other ancient and present-day samples, fitting them onto the scaffold as 2- or 3-way
mixtures. The uncertainty in all parameter estimates is measured by block bootstrap
resampling of the SNP set (100 replicates with 50 blocks).</p>
</sec>
<sec id="S23"><title><italic>TreeMix</italic>
analysis</title>
<p id="P35">We applied <italic>TreeMix</italic>
<sup><xref rid="R21" ref-type="bibr">21</xref>
</sup>
to Loschbour, Stuttgart, Motala12, and MA1<sup><xref rid="R3" ref-type="bibr">3</xref>
</sup>
, LaBrana<sup><xref rid="R2" ref-type="bibr">2</xref>
</sup>
and the Iceman<sup><xref rid="R1" ref-type="bibr">1</xref>
</sup>
, along
with the present-day samples of Karitiana, Onge and Mbuti. We restricted the analysis to
265,521 Human Origins array sites after excluding any SNPs where there were no-calls in
any of the studied individuals. The tree was rooted with Mbuti and standard errors were
estimated using blocks of 500 SNPs. We repeated the analysis on whole-genome sequence
data, rooting with Chimp and replacing Onge with Dai since we did not have Onge whole
genome sequence data<sup><xref rid="R55" ref-type="bibr">55</xref>
</sup>
. We varied the
number of migration events (<italic>m</italic>
) between 0 and 5.</p>
</sec>
<sec id="S24"><title>Inferring admixture proportions with minimal modeling assumptions</title>
<p id="P36">We devised a method to infer ancestry proportions from three ancestral
populations (EEF, WHG, and ANE) without strong phylogenetic assumptions (SI17). We rely on
15 “non-West Eurasian” outgroups and study <italic>f<sub>4</sub>
(European,
Stuttgart; O<sub>1</sub>
, O<sub>2</sub>
)</italic>
which equals
<italic>αβ f<sub>4</sub>
(Loschbour, Stuttgart; O<sub>1</sub>
,
O<sub>2</sub>
)</italic>
+ <italic>α(1−β)
f<sub>4</sub>
(MA1, Stuttgart; O<sub>1</sub>
, O<sub>2</sub>
)</italic>
if
<italic>European</italic>
has 1−<italic>a</italic>
ancestry from EEF and
<italic>β</italic>
, 1−<italic>β</italic>
ancestry from WHG and
ANE respectively. This defines a system of <inline-formula><mml:math id="M3" overflow="scroll"><mml:mo>(</mml:mo>
<mml:mtable><mml:mtr><mml:mtd><mml:mn>15</mml:mn>
</mml:mtd>
</mml:mtr>
<mml:mtr><mml:mtd><mml:mn>2</mml:mn>
</mml:mtd>
</mml:mtr>
</mml:mtable>
<mml:mo>)</mml:mo>
<mml:mo>=</mml:mo>
<mml:mn>105</mml:mn>
</mml:math>
</inline-formula>
equations with unknowns
<italic>αβ</italic>
,
<italic>α</italic>
(1−<italic>β</italic>
), which we solve with
least squares implemented in the function <italic>lsfit</italic>
in <italic>R</italic>
to
obtain estimates of <italic>α</italic>
and <italic>β</italic>
. We repeated
this computation 22 times dropping one chromosome at a time<sup><xref rid="R20" ref-type="bibr">20</xref>
</sup>
to obtain block jackknife<sup><xref rid="R67" ref-type="bibr">67</xref>
</sup>
estimates of the ancestry proportions and standard
errors, with block size equal to the number of SNPs per chromosome. We assessed
consistency of the inferred admixture proportions with those derived from the
ADMIXTUREGRAPH model based on the number of standard errors between the two (<xref rid="T2" ref-type="table">Extended Data Table 1</xref>
).</p>
</sec>
<sec id="S25"><title>Haplotype-based analyses</title>
<p id="P37">We used RefinedIBD from BEAGLE 4<sup><xref rid="R27" ref-type="bibr">27</xref>
</sup>
with the settings <italic>ibdtrim</italic>
=20 and
<italic>ibdwindow</italic>
=25 to study IBD sharing between Loschbour and
Stuttgart and populations from the POPRES dataset<sup><xref rid="R69" ref-type="bibr">69</xref>
</sup>
. We kept all IBD tracts spanning at least 0.5 centimorgans (cM) and
with a LOD score >3 (SI18). We also used ChromoPainter<sup><xref rid="R29" ref-type="bibr">29</xref>
</sup>
to study haplotype sharing between Loschbour and
Stuttgart and present-day West Eurasian populations (SI19). We identified 495,357 SNPs
that were complete in all individuals and phased the data using Beagle 4<sup><xref rid="R27" ref-type="bibr">27</xref>
</sup>
with parameters
<italic>phase-its</italic>
=50 and <italic>impute-its</italic>
=10. We did
not keep sites with missing data to avoid imputing modern alleles into the ancient
individuals. We used both unlinked (-k 1000) and linked modes (estimating -n and -M by
sampling 10% of individuals). We combined ChromoPainter output for chromosomes
1-22 using ChromoCombine<sup><xref rid="R29" ref-type="bibr">29</xref>
</sup>
. We carried
out a PCA of the co-ancestry matrix using fineSTRUCTURE<sup><xref rid="R29" ref-type="bibr">29</xref>
</sup>
.</p>
</sec>
</sec>
<sec id="S26"><title>Extended Data</title>
<fig id="F3" orientation="portrait" position="anchor"><label>Extended Data Figure 1</label>
<caption><p id="P39">Photographs of analyzed ancient samples.</p>
<p id="P40">(A) Loschbour skull; (B) Stuttgart skull, missing the lower right M2 we
sampled; (C) excavation at Kanaljorden in Motala, Sweden; (D) Motala 1 in situ.</p>
</caption>
<graphic xlink:href="nihms613260f3"></graphic>
</fig>
<fig id="F4" orientation="portrait" position="anchor"><label>Extended Data Figure 2</label>
<caption><p id="P41">Pairwise Sequential Markovian Coalescent (PSMC) analysis.</p>
<p id="P42">(A) Inference of population size as a function of time, showing a very small
recent population size over the most recent period in the ancestry of Loschbour (at
least the last 5–10 thousand years). (B) Inferred time since the most recent
common ancestor from the PSMC for chromosomes 20, 21, 22 (top to bottom); Stuttgart is
plotted on top and Loschbour at bottom.</p>
</caption>
<graphic xlink:href="nihms613260f4"></graphic>
</fig>
<fig id="F5" orientation="portrait" position="anchor"><label>Extended Data Figure 3</label>
<caption><p id="P43">ADMIXTURE analysis (K=2 to K=20).</p>
<p id="P44">Ancient samples (Loschbour, Stuttgart, Motala_merge, Motala12, MA1, and
LaBrana) are at left.</p>
</caption>
<graphic xlink:href="nihms613260f5"></graphic>
</fig>
<fig id="F6" orientation="portrait" position="anchor"><label>Extended Data Figure 4</label>
<caption><p id="P45">ANE ancestry is present in both Europe and the Near East but WHG ancestry is
restricted to Europe, which cannot be due to a single admixture event.</p>
<p id="P46">(<italic>x-</italic>
axis) We computed the statistic
<italic>f<sub>4</sub>
(Test, Stuttgart; MA1, Chimp)</italic>
, which measures where MA1
shares more alleles with a test population than with Stuttgart. It is positive for most
European and Near Eastern populations, consistent with ANE (MA1-related) gene flow into
both regions. (<italic>y-</italic>
axis) We computed the statistic
<italic>f<sub>4</sub>
(Test, Stuttgart; Loschbour, Chimp),</italic>
which measures
whether Loschbour shares more alleles with a test sample than with Stuttgart. Only
European populations show positive values of this statistic, providing evidence of WHG
(Loschbour-related) admixture only in Europeans.</p>
</caption>
<graphic xlink:href="nihms613260f6"></graphic>
</fig>
<fig id="F7" orientation="portrait" position="anchor"><label>Extended Data Figure 5</label>
<caption><p id="P47">MA1 is the best surrogate for ANE for which we have data.</p>
<p id="P48">Europeans share more alleles with MA1 than with Karitiana, as we see from the
fact that in a plot of <italic>f<sub>4</sub>
(Test, BedouinB; MA1, Chimp)</italic>
and
<italic>f<sub>4</sub>
(Test, BedouinB; Karitiana, Chimp)</italic>
, the European cline
deviates in the direction of MA1, rather than Karitiana (the slope is >1 and European
populations are above the line indicating equality of these two statistics).</p>
</caption>
<graphic xlink:href="nihms613260f7"></graphic>
</fig>
<fig id="F8" orientation="portrait" position="anchor"><label>Extended Data Figure 6</label>
<caption><p id="P49">The differential relatedness of West Eurasians to Stuttgart (EEF), Loschbour
(WHG), and MA1 (ANE) cannot be explained by two-way mixture.</p>
<p id="P50">We plot on a West Eurasian map the statistic <italic>f<sub>4</sub>
(Test,
Chimp; A<sub>1</sub>
, A<sub>2</sub>
)</italic>
, where <italic>A<sub>1</sub>
</italic>
and <italic>A<sub>2</sub>
</italic>
are a pair of the three ancient samples representing
the three ancestral populations of Europe. (A) In both Europe and the Near
East/Caucasus, populations from the south have more relatedness to Stuttgart than those
from the north where ANE influence is also important. (B) Northern European populations
share more alleles with Loschbour than with Stuttgart, as they have additional WHG
ancestry beyond what was already present in EEF. (C) We observe a striking contrast
between Europe west of the Caucasus and the Near East in degree of relatedness to WHG.
In Europe, there is a much higher degree of allele sharing with Loschbour than with MA1,
which we ascribe to the 60–80% WHG/(WHG+ANE) ratio in most
Europeans that we report in SI14. In contrast, the Near East has no appreciable WHG
ancestry but some ANE ancestry, especially in the northern Caucasus. (Jewish populations
are marked with a square in this figure to assist in interpretation as their ancestry is
often anomalous for their geographic regions.)</p>
</caption>
<graphic xlink:href="nihms613260f8"></graphic>
</fig>
<fig id="F9" orientation="portrait" position="anchor"><label>Extended Data Figure 7</label>
<caption><p id="P51">Evidence for Siberian gene flow into far northeastern Europe.</p>
<p id="P52">Some northeastern European populations (Chuvash, Finnish, Russian, Mordovian,
Saami) share more alleles with Han Chinese than with other Europeans who are arrayed in
a cline from Stuttgart to Lithuanians/Estonians in a plot of <italic>f<sub>4</sub>
(Test,
BedouinB; Han, Mbuti)</italic>
against <italic>f<sub>4</sub>
(Test, BedouinB; MA1,
Mbuti)</italic>
.</p>
</caption>
<graphic xlink:href="nihms613260f9"></graphic>
</fig>
<table-wrap id="T2" position="anchor" orientation="portrait"><label>Extended Data Table 1</label>
<caption><p id="P53">West Eurasians genotyped on the Human Origins array and key
<italic>f</italic>
-statistics.</p>
</caption>
<table frame="box" rules="groups"><thead><tr><th valign="middle" align="left" rowspan="1" colspan="1"></th>
<th colspan="3" valign="middle" align="center" rowspan="1">Sampling Location</th>
<th colspan="4" valign="middle" align="center" rowspan="1">Lowest <italic>f<sub>3</sub>
(X;
Ref<sub>1</sub>
, Ref<sub>2</sub>
)</italic>
</th>
<th colspan="5" valign="middle" align="center" rowspan="1">Lowest <italic>f<sub>3</sub>
(X; EEF,
WHG)</italic>
<break></break>
(Z<0 and Zdiff<3 reported)</th>
<th colspan="5" valign="middle" align="center" rowspan="1">Lowest <italic>f<sub>3</sub>
(X; Near
East, WHG)</italic>
<break></break>
(Z<0 and Zdiff<3 reported)</th>
<th colspan="5" valign="middle" align="center" rowspan="1">Lowest <italic>f<sub>3</sub>
(X; EEF,
ANE)</italic>
<break></break>
(Z<0 and Zdiff<3 reported)</th>
<th colspan="2" valign="middle" align="center" rowspan="1"><italic>f<sub>4</sub>
(Stuttgart,
X;</italic>
<break></break>
<italic>Loschbour, Chimp)</italic>
</th>
<th colspan="2" valign="middle" align="center" rowspan="1"><italic>f<sub>4</sub>
(Stuttgart,
X;</italic>
<break></break>
<italic>MA1, Chimp)</italic>
</th>
</tr>
<tr><th valign="bottom" align="left" rowspan="1" colspan="1">X</th>
<th valign="bottom" align="right" rowspan="1" colspan="1">N</th>
<th valign="bottom" align="right" rowspan="1" colspan="1">Lat.</th>
<th valign="bottom" align="right" rowspan="1" colspan="1">Long.</th>
<th valign="bottom" align="left" rowspan="1" colspan="1">Ref<sub>1</sub>
</th>
<th valign="bottom" align="left" rowspan="1" colspan="1">Ref<sub>2</sub>
</th>
<th valign="bottom" align="left" rowspan="1" colspan="1">statistic</th>
<th valign="bottom" align="right" rowspan="1" colspan="1">Z</th>
<th valign="bottom" align="left" rowspan="1" colspan="1">Ref<sub>1</sub>
</th>
<th valign="bottom" align="left" rowspan="1" colspan="1">Ref<sub>2</sub>
</th>
<th valign="bottom" align="center" rowspan="1" colspan="1">statistic</th>
<th valign="bottom" align="center" rowspan="1" colspan="1">Z</th>
<th valign="bottom" align="center" rowspan="1" colspan="1">Zdiff</th>
<th valign="bottom" align="left" rowspan="1" colspan="1">Ref<sub>1</sub>
</th>
<th valign="bottom" align="left" rowspan="1" colspan="1">Ref<sub>2</sub>
</th>
<th valign="bottom" align="center" rowspan="1" colspan="1">statistic</th>
<th valign="bottom" align="right" rowspan="1" colspan="1">Z</th>
<th valign="bottom" align="center" rowspan="1" colspan="1">Zdiff</th>
<th valign="bottom" align="left" rowspan="1" colspan="1">Ref<sub>1</sub>
</th>
<th valign="bottom" align="left" rowspan="1" colspan="1">Ref<sub>2</sub>
</th>
<th valign="bottom" align="center" rowspan="1" colspan="1">statistic</th>
<th valign="bottom" align="right" rowspan="1" colspan="1">Z</th>
<th valign="bottom" align="right" rowspan="1" colspan="1">Zdiff</th>
<th valign="bottom" align="right" rowspan="1" colspan="1">statistic</th>
<th valign="bottom" align="right" rowspan="1" colspan="1">Z</th>
<th valign="bottom" align="right" rowspan="1" colspan="1">statistic</th>
<th valign="bottom" align="right" rowspan="1" colspan="1">Z</th>
</tr>
</thead>
<tbody><tr><td align="left" valign="top" rowspan="1" colspan="1">Abkhasian</td>
<td align="right" valign="top" rowspan="1" colspan="1">9</td>
<td align="right" valign="top" rowspan="1" colspan="1">43</td>
<td align="right" valign="top" rowspan="1" colspan="1">41.02</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0053</td>
<td align="right" valign="top" rowspan="1" colspan="1">−2.9</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1">Georgian</td>
<td align="left" valign="top" rowspan="1" colspan="1">LaB</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0004</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.5</td>
<td align="center" valign="top" rowspan="1" colspan="1">2.6</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0053</td>
<td align="right" valign="top" rowspan="1" colspan="1">−2.9</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0020</td>
<td align="right" valign="top" rowspan="1" colspan="1">4.2</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0023</td>
<td align="right" valign="top" rowspan="1" colspan="1">−4.7</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Adygei</td>
<td align="right" valign="top" rowspan="1" colspan="1">17</td>
<td align="right" valign="top" rowspan="1" colspan="1">44</td>
<td align="right" valign="top" rowspan="1" colspan="1">39</td>
<td align="left" valign="top" rowspan="1" colspan="1">Piapoco</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0073</td>
<td align="right" valign="top" rowspan="1" colspan="1">−5.9</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0067</td>
<td align="right" valign="top" rowspan="1" colspan="1">−4.1</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.3</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0013</td>
<td align="right" valign="top" rowspan="1" colspan="1">2.6</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0029</td>
<td align="right" valign="top" rowspan="1" colspan="1">−6.0</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Albanian</td>
<td align="right" valign="top" rowspan="1" colspan="1">6</td>
<td align="right" valign="top" rowspan="1" colspan="1">41.33</td>
<td align="right" valign="top" rowspan="1" colspan="1">19.83</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0121</td>
<td align="right" valign="top" rowspan="1" colspan="1">−7.0</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1">Iraqi_Jew</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0090</td>
<td align="right" valign="top" rowspan="1" colspan="1">−9.1</td>
<td align="center" valign="top" rowspan="1" colspan="1">1.7</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0121</td>
<td align="right" valign="top" rowspan="1" colspan="1">−7.0</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0009</td>
<td align="right" valign="top" rowspan="1" colspan="1">−1.8</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0027</td>
<td align="right" valign="top" rowspan="1" colspan="1">−5.4</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Armenian</td>
<td align="right" valign="top" rowspan="1" colspan="1">10</td>
<td align="right" valign="top" rowspan="1" colspan="1">40.19</td>
<td align="right" valign="top" rowspan="1" colspan="1">44.55</td>
<td align="left" valign="top" rowspan="1" colspan="1">GujaratiC</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0070</td>
<td align="right" valign="top" rowspan="1" colspan="1">−8.2</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0068</td>
<td align="right" valign="top" rowspan="1" colspan="1">−4.1</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.1</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0022</td>
<td align="right" valign="top" rowspan="1" colspan="1">4.5</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0016</td>
<td align="right" valign="top" rowspan="1" colspan="1">−3.3</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Ashkenazi_Jew</td>
<td align="right" valign="top" rowspan="1" colspan="1">7</td>
<td align="right" valign="top" rowspan="1" colspan="1">52.23</td>
<td align="right" valign="top" rowspan="1" colspan="1">21.02</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0057</td>
<td align="right" valign="top" rowspan="1" colspan="1">−3.4</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1">Iraqi_Jew</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0042</td>
<td align="right" valign="top" rowspan="1" colspan="1">−4.7</td>
<td align="center" valign="top" rowspan="1" colspan="1">1.0</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0057</td>
<td align="right" valign="top" rowspan="1" colspan="1">−3.4</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0008</td>
<td align="right" valign="top" rowspan="1" colspan="1">1.7</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0010</td>
<td align="right" valign="top" rowspan="1" colspan="1">−2.0</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Balkar</td>
<td align="right" valign="top" rowspan="1" colspan="1">10</td>
<td align="right" valign="top" rowspan="1" colspan="1">43.48</td>
<td align="right" valign="top" rowspan="1" colspan="1">43.62</td>
<td align="left" valign="top" rowspan="1" colspan="1">Piapoco</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0113</td>
<td align="right" valign="top" rowspan="1" colspan="1">−8.9</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0092</td>
<td align="right" valign="top" rowspan="1" colspan="1">−5.5</td>
<td align="right" valign="top" rowspan="1" colspan="1">1.1</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0014</td>
<td align="right" valign="top" rowspan="1" colspan="1">2.9</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0027</td>
<td align="right" valign="top" rowspan="1" colspan="1">−5.6</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Basque</td>
<td align="right" valign="top" rowspan="1" colspan="1">29</td>
<td align="right" valign="top" rowspan="1" colspan="1">43.04</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.65</td>
<td align="left" valign="top" rowspan="1" colspan="1">Iraqi_Jew</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0083</td>
<td align="right" valign="top" rowspan="1" colspan="1">−10.3</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0061</td>
<td align="center" valign="top" rowspan="1" colspan="1">−3.8</td>
<td align="center" valign="top" rowspan="1" colspan="1">1.3</td>
<td align="left" valign="top" rowspan="1" colspan="1">Iraqi_Jew</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0083</td>
<td align="right" valign="top" rowspan="1" colspan="1">−10.3</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.0</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0041</td>
<td align="right" valign="top" rowspan="1" colspan="1">−2.4</td>
<td align="right" valign="top" rowspan="1" colspan="1">2.2</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0034</td>
<td align="right" valign="top" rowspan="1" colspan="1">−7.2</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0032</td>
<td align="right" valign="top" rowspan="1" colspan="1">−6.7</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">BedouinA</td>
<td align="right" valign="top" rowspan="1" colspan="1">25</td>
<td align="right" valign="top" rowspan="1" colspan="1">31</td>
<td align="right" valign="top" rowspan="1" colspan="1">35</td>
<td align="left" valign="top" rowspan="1" colspan="1">Esan</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0162</td>
<td align="right" valign="top" rowspan="1" colspan="1">−18.2</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0062</td>
<td align="right" valign="top" rowspan="1" colspan="1">13.0</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0026</td>
<td align="right" valign="top" rowspan="1" colspan="1">5.4</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">BedouinB</td>
<td align="right" valign="top" rowspan="1" colspan="1">19</td>
<td align="right" valign="top" rowspan="1" colspan="1">31</td>
<td align="right" valign="top" rowspan="1" colspan="1">35</td>
<td align="left" valign="top" rowspan="1" colspan="1">Esan</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">0.0089</td>
<td align="right" valign="top" rowspan="1" colspan="1">7.8</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0046</td>
<td align="right" valign="top" rowspan="1" colspan="1">9.3</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0019</td>
<td align="right" valign="top" rowspan="1" colspan="1">3.9</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Belarusian</td>
<td align="right" valign="top" rowspan="1" colspan="1">10</td>
<td align="right" valign="top" rowspan="1" colspan="1">53.92</td>
<td align="right" valign="top" rowspan="1" colspan="1">28.01</td>
<td align="left" valign="top" rowspan="1" colspan="1">Georgian</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0133</td>
<td align="right" valign="top" rowspan="1" colspan="1">−17.6</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1">Georgian</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0133</td>
<td align="right" valign="top" rowspan="1" colspan="1">−17.6</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.0</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0102</td>
<td align="right" valign="top" rowspan="1" colspan="1">−6.1</td>
<td align="right" valign="top" rowspan="1" colspan="1">1.9</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0035</td>
<td align="right" valign="top" rowspan="1" colspan="1">−6.9</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0042</td>
<td align="right" valign="top" rowspan="1" colspan="1">−8.6</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Bergamo</td>
<td align="right" valign="top" rowspan="1" colspan="1">12</td>
<td align="right" valign="top" rowspan="1" colspan="1">46</td>
<td align="right" valign="top" rowspan="1" colspan="1">10</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0106</td>
<td align="right" valign="top" rowspan="1" colspan="1">−6.2</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0068</td>
<td align="center" valign="top" rowspan="1" colspan="1">−4.2</td>
<td align="center" valign="top" rowspan="1" colspan="1">1.7</td>
<td align="left" valign="top" rowspan="1" colspan="1">Iraqi_Jew</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0100</td>
<td align="right" valign="top" rowspan="1" colspan="1">−11.9</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.3</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0106</td>
<td align="right" valign="top" rowspan="1" colspan="1">−6.2</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0018</td>
<td align="right" valign="top" rowspan="1" colspan="1">−3.9</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0028</td>
<td align="right" valign="top" rowspan="1" colspan="1">−5.8</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Bulgarian</td>
<td align="right" valign="top" rowspan="1" colspan="1">10</td>
<td align="right" valign="top" rowspan="1" colspan="1">42.16</td>
<td align="right" valign="top" rowspan="1" colspan="1">24.74</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0130</td>
<td align="right" valign="top" rowspan="1" colspan="1">−8.2</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">LaB</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0074</td>
<td align="center" valign="top" rowspan="1" colspan="1">−4.5</td>
<td align="center" valign="top" rowspan="1" colspan="1">2.8</td>
<td align="left" valign="top" rowspan="1" colspan="1">Iraqi_Jew</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0106</td>
<td align="right" valign="top" rowspan="1" colspan="1">−12.4</td>
<td align="center" valign="top" rowspan="1" colspan="1">1.5</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0130</td>
<td align="right" valign="top" rowspan="1" colspan="1">−8.2</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0012</td>
<td align="right" valign="top" rowspan="1" colspan="1">−2.5</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0028</td>
<td align="right" valign="top" rowspan="1" colspan="1">−5.9</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Chechen</td>
<td align="right" valign="top" rowspan="1" colspan="1">9</td>
<td align="right" valign="top" rowspan="1" colspan="1">43.33</td>
<td align="right" valign="top" rowspan="1" colspan="1">45.65</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0056</td>
<td align="right" valign="top" rowspan="1" colspan="1">−3.2</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1">Georgian</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0002</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.3</td>
<td align="center" valign="top" rowspan="1" colspan="1">2.8</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0056</td>
<td align="right" valign="top" rowspan="1" colspan="1">−3.2</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0011</td>
<td align="right" valign="top" rowspan="1" colspan="1">2.3</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0031</td>
<td align="right" valign="top" rowspan="1" colspan="1">−6.2</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Croatian</td>
<td align="right" valign="top" rowspan="1" colspan="1">10</td>
<td align="right" valign="top" rowspan="1" colspan="1">43.51</td>
<td align="right" valign="top" rowspan="1" colspan="1">16.45</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0114</td>
<td align="right" valign="top" rowspan="1" colspan="1">−6.7</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0065</td>
<td align="center" valign="top" rowspan="1" colspan="1">−3.8</td>
<td align="center" valign="top" rowspan="1" colspan="1">2.1</td>
<td align="left" valign="top" rowspan="1" colspan="1">Iraqi_Jew</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0112</td>
<td align="right" valign="top" rowspan="1" colspan="1">−13.0</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.2</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0114</td>
<td align="right" valign="top" rowspan="1" colspan="1">−6.7</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0023</td>
<td align="right" valign="top" rowspan="1" colspan="1">−4.7</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0035</td>
<td align="right" valign="top" rowspan="1" colspan="1">−7.4</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Cypriot</td>
<td align="right" valign="top" rowspan="1" colspan="1">8</td>
<td align="right" valign="top" rowspan="1" colspan="1">35.13</td>
<td align="right" valign="top" rowspan="1" colspan="1">33.43</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0057</td>
<td align="right" valign="top" rowspan="1" colspan="1">−3.2</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1">Yemenite_Jew</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0013</td>
<td align="right" valign="top" rowspan="1" colspan="1">−1.5</td>
<td align="center" valign="top" rowspan="1" colspan="1">2.5</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0057</td>
<td align="right" valign="top" rowspan="1" colspan="1">−3.2</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0019</td>
<td align="right" valign="top" rowspan="1" colspan="1">3.9</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0012</td>
<td align="right" valign="top" rowspan="1" colspan="1">−2.5</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Czech</td>
<td align="right" valign="top" rowspan="1" colspan="1">10</td>
<td align="right" valign="top" rowspan="1" colspan="1">50.1</td>
<td align="right" valign="top" rowspan="1" colspan="1">14.4</td>
<td align="left" valign="top" rowspan="1" colspan="1">Georgian</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0137</td>
<td align="right" valign="top" rowspan="1" colspan="1">−17.9</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0088</td>
<td align="center" valign="top" rowspan="1" colspan="1">−5.3</td>
<td align="center" valign="top" rowspan="1" colspan="1">3.0</td>
<td align="left" valign="top" rowspan="1" colspan="1">Georgian</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0137</td>
<td align="right" valign="top" rowspan="1" colspan="1">−17.9</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.0</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0121</td>
<td align="right" valign="top" rowspan="1" colspan="1">−7.2</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.9</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0032</td>
<td align="right" valign="top" rowspan="1" colspan="1">−6.6</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0040</td>
<td align="right" valign="top" rowspan="1" colspan="1">−8.2</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Druze</td>
<td align="right" valign="top" rowspan="1" colspan="1">39</td>
<td align="right" valign="top" rowspan="1" colspan="1">32</td>
<td align="right" valign="top" rowspan="1" colspan="1">35</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0024</td>
<td align="right" valign="top" rowspan="1" colspan="1">−1.5</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0024</td>
<td align="right" valign="top" rowspan="1" colspan="1">−1.5</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0028</td>
<td align="right" valign="top" rowspan="1" colspan="1">5.9</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0006</td>
<td align="right" valign="top" rowspan="1" colspan="1">−1.3</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">English</td>
<td align="right" valign="top" rowspan="1" colspan="1">10</td>
<td align="right" valign="top" rowspan="1" colspan="1">50.75</td>
<td align="right" valign="top" rowspan="1" colspan="1">−2.09</td>
<td align="left" valign="top" rowspan="1" colspan="1">Iraqi_Jew</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0129</td>
<td align="right" valign="top" rowspan="1" colspan="1">−14.8</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0090</td>
<td align="center" valign="top" rowspan="1" colspan="1">−5.5</td>
<td align="center" valign="top" rowspan="1" colspan="1">2.2</td>
<td align="left" valign="top" rowspan="1" colspan="1">Iraqi_Jew</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0129</td>
<td align="right" valign="top" rowspan="1" colspan="1">−14.8</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.0</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0125</td>
<td align="right" valign="top" rowspan="1" colspan="1">−7.4</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.1</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0032</td>
<td align="right" valign="top" rowspan="1" colspan="1">−6.5</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0041</td>
<td align="right" valign="top" rowspan="1" colspan="1">−8.5</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Estonian</td>
<td align="right" valign="top" rowspan="1" colspan="1">10</td>
<td align="right" valign="top" rowspan="1" colspan="1">58.54</td>
<td align="right" valign="top" rowspan="1" colspan="1">24.89</td>
<td align="left" valign="top" rowspan="1" colspan="1">Abkhasian</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0124</td>
<td align="right" valign="top" rowspan="1" colspan="1">−15.1</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1">Abkhasian</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0124</td>
<td align="right" valign="top" rowspan="1" colspan="1">−15.1</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.0</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0094</td>
<td align="right" valign="top" rowspan="1" colspan="1">−5.6</td>
<td align="right" valign="top" rowspan="1" colspan="1">1.9</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0043</td>
<td align="right" valign="top" rowspan="1" colspan="1">−8.5</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0051</td>
<td align="right" valign="top" rowspan="1" colspan="1">−10.1</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Finnish</td>
<td align="right" valign="top" rowspan="1" colspan="1">7</td>
<td align="right" valign="top" rowspan="1" colspan="1">60.2</td>
<td align="right" valign="top" rowspan="1" colspan="1">24.9</td>
<td align="left" valign="top" rowspan="1" colspan="1">Abkhasian</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0102</td>
<td align="right" valign="top" rowspan="1" colspan="1">−11.3</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1">Abkhasian</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0102</td>
<td align="right" valign="top" rowspan="1" colspan="1">−11.3</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.0</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0078</td>
<td align="right" valign="top" rowspan="1" colspan="1">−4.4</td>
<td align="right" valign="top" rowspan="1" colspan="1">1.4</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0035</td>
<td align="right" valign="top" rowspan="1" colspan="1">−6.9</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0045</td>
<td align="right" valign="top" rowspan="1" colspan="1">−9.1</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">French</td>
<td align="right" valign="top" rowspan="1" colspan="1">25</td>
<td align="right" valign="top" rowspan="1" colspan="1">46</td>
<td align="right" valign="top" rowspan="1" colspan="1">2</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0131</td>
<td align="right" valign="top" rowspan="1" colspan="1">−8.4</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0098</td>
<td align="center" valign="top" rowspan="1" colspan="1">−6.3</td>
<td align="center" valign="top" rowspan="1" colspan="1">1.5</td>
<td align="left" valign="top" rowspan="1" colspan="1">Iraqi_Jew</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0129</td>
<td align="right" valign="top" rowspan="1" colspan="1">−16.8</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.2</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0131</td>
<td align="right" valign="top" rowspan="1" colspan="1">−8.4</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0027</td>
<td align="right" valign="top" rowspan="1" colspan="1">−5.6</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0036</td>
<td align="right" valign="top" rowspan="1" colspan="1">−7.7</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">French_South</td>
<td align="right" valign="top" rowspan="1" colspan="1">7</td>
<td align="right" valign="top" rowspan="1" colspan="1">43.44</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.62</td>
<td align="left" valign="top" rowspan="1" colspan="1">Iraqi_Jew</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0095</td>
<td align="right" valign="top" rowspan="1" colspan="1">−9.5</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">LaB</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0089</td>
<td align="center" valign="top" rowspan="1" colspan="1">−5.0</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.3</td>
<td align="left" valign="top" rowspan="1" colspan="1">Iraqi_Jew</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0095</td>
<td align="right" valign="top" rowspan="1" colspan="1">−9.5</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.0</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0086</td>
<td align="right" valign="top" rowspan="1" colspan="1">−4.8</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.4</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0030</td>
<td align="right" valign="top" rowspan="1" colspan="1">−6.2</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0031</td>
<td align="right" valign="top" rowspan="1" colspan="1">−6.2</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Georgian</td>
<td align="right" valign="top" rowspan="1" colspan="1">10</td>
<td align="right" valign="top" rowspan="1" colspan="1">42.5</td>
<td align="right" valign="top" rowspan="1" colspan="1">41.85</td>
<td align="left" valign="top" rowspan="1" colspan="1">GujaratiC</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0036</td>
<td align="right" valign="top" rowspan="1" colspan="1">−4.0</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0036</td>
<td align="right" valign="top" rowspan="1" colspan="1">−2.1</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.2</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0020</td>
<td align="right" valign="top" rowspan="1" colspan="1">4.2</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0019</td>
<td align="right" valign="top" rowspan="1" colspan="1">−3.9</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Georgian_Jew</td>
<td align="right" valign="top" rowspan="1" colspan="1">7</td>
<td align="right" valign="top" rowspan="1" colspan="1">41.72</td>
<td align="right" valign="top" rowspan="1" colspan="1">44.78</td>
<td align="left" valign="top" rowspan="1" colspan="1">GujaratiC</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0009</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.9</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0002</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.1</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.3</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0022</td>
<td align="right" valign="top" rowspan="1" colspan="1">4.3</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0017</td>
<td align="right" valign="top" rowspan="1" colspan="1">−3.4</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Greek</td>
<td align="right" valign="top" rowspan="1" colspan="1">20</td>
<td align="right" valign="top" rowspan="1" colspan="1">39.84</td>
<td align="right" valign="top" rowspan="1" colspan="1">23.17</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0118</td>
<td align="right" valign="top" rowspan="1" colspan="1">−7.4</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1">Iraqi_Jew</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0080</td>
<td align="right" valign="top" rowspan="1" colspan="1">−11.1</td>
<td align="center" valign="top" rowspan="1" colspan="1">2.3</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0118</td>
<td align="right" valign="top" rowspan="1" colspan="1">−7.4</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0004</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.9</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0026</td>
<td align="right" valign="top" rowspan="1" colspan="1">−5.6</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Hungarian</td>
<td align="right" valign="top" rowspan="1" colspan="1">20</td>
<td align="right" valign="top" rowspan="1" colspan="1">47.49</td>
<td align="right" valign="top" rowspan="1" colspan="1">19.08</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0133</td>
<td align="right" valign="top" rowspan="1" colspan="1">−8.4</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0087</td>
<td align="center" valign="top" rowspan="1" colspan="1">−5.6</td>
<td align="center" valign="top" rowspan="1" colspan="1">2.2</td>
<td align="left" valign="top" rowspan="1" colspan="1">Iraqi_Jew</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0127</td>
<td align="right" valign="top" rowspan="1" colspan="1">−15.9</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.4</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0133</td>
<td align="right" valign="top" rowspan="1" colspan="1">−8.4</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0025</td>
<td align="right" valign="top" rowspan="1" colspan="1">−5.3</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0037</td>
<td align="right" valign="top" rowspan="1" colspan="1">−7.8</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Icelandic</td>
<td align="right" valign="top" rowspan="1" colspan="1">12</td>
<td align="right" valign="top" rowspan="1" colspan="1">64.13</td>
<td align="right" valign="top" rowspan="1" colspan="1">−21.93</td>
<td align="left" valign="top" rowspan="1" colspan="1">Abkhasian</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0121</td>
<td align="right" valign="top" rowspan="1" colspan="1">−15.6</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0078</td>
<td align="center" valign="top" rowspan="1" colspan="1">−4.8</td>
<td align="center" valign="top" rowspan="1" colspan="1">2.7</td>
<td align="left" valign="top" rowspan="1" colspan="1">Abkhasian</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0121</td>
<td align="right" valign="top" rowspan="1" colspan="1">−15.6</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.0</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0097</td>
<td align="right" valign="top" rowspan="1" colspan="1">−5.9</td>
<td align="right" valign="top" rowspan="1" colspan="1">1.5</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0038</td>
<td align="right" valign="top" rowspan="1" colspan="1">−7.7</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0043</td>
<td align="right" valign="top" rowspan="1" colspan="1">−8.9</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Iranian</td>
<td align="right" valign="top" rowspan="1" colspan="1">8</td>
<td align="right" valign="top" rowspan="1" colspan="1">35.59</td>
<td align="right" valign="top" rowspan="1" colspan="1">51.46</td>
<td align="left" valign="top" rowspan="1" colspan="1">Piapoco</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0094</td>
<td align="right" valign="top" rowspan="1" colspan="1">−7.2</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0087</td>
<td align="right" valign="top" rowspan="1" colspan="1">−5.2</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.4</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0031</td>
<td align="right" valign="top" rowspan="1" colspan="1">6.3</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0016</td>
<td align="right" valign="top" rowspan="1" colspan="1">−3.2</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Iranian_Jew</td>
<td align="right" valign="top" rowspan="1" colspan="1">9</td>
<td align="right" valign="top" rowspan="1" colspan="1">35.7</td>
<td align="right" valign="top" rowspan="1" colspan="1">51.42</td>
<td align="left" valign="top" rowspan="1" colspan="1">GujaratiC</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0018</td>
<td align="right" valign="top" rowspan="1" colspan="1">−2.0</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0012</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.6</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.2</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0028</td>
<td align="right" valign="top" rowspan="1" colspan="1">5.7</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0011</td>
<td align="right" valign="top" rowspan="1" colspan="1">−2.2</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Iraqi_Jew</td>
<td align="right" valign="top" rowspan="1" colspan="1">6</td>
<td align="right" valign="top" rowspan="1" colspan="1">33.33</td>
<td align="right" valign="top" rowspan="1" colspan="1">44.42</td>
<td align="left" valign="top" rowspan="1" colspan="1">Vishwabrahmin</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0026</td>
<td align="right" valign="top" rowspan="1" colspan="1">−2.6</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0009</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.5</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.9</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0030</td>
<td align="right" valign="top" rowspan="1" colspan="1">6.1</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0005</td>
<td align="right" valign="top" rowspan="1" colspan="1">−1.0</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Jordanian</td>
<td align="right" valign="top" rowspan="1" colspan="1">9</td>
<td align="right" valign="top" rowspan="1" colspan="1">32.05</td>
<td align="right" valign="top" rowspan="1" colspan="1">35.91</td>
<td align="left" valign="top" rowspan="1" colspan="1">Esan</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0145</td>
<td align="right" valign="top" rowspan="1" colspan="1">−14.3</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0048</td>
<td align="right" valign="top" rowspan="1" colspan="1">9.6</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0014</td>
<td align="right" valign="top" rowspan="1" colspan="1">2.8</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Kumyk</td>
<td align="right" valign="top" rowspan="1" colspan="1">8</td>
<td align="right" valign="top" rowspan="1" colspan="1">43.25</td>
<td align="right" valign="top" rowspan="1" colspan="1">46.58</td>
<td align="left" valign="top" rowspan="1" colspan="1">Piapoco</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0111</td>
<td align="right" valign="top" rowspan="1" colspan="1">−8.2</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0109</td>
<td align="right" valign="top" rowspan="1" colspan="1">−6.5</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.1</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0015</td>
<td align="right" valign="top" rowspan="1" colspan="1">3.1</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0028</td>
<td align="right" valign="top" rowspan="1" colspan="1">−5.7</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Lebanese</td>
<td align="right" valign="top" rowspan="1" colspan="1">8</td>
<td align="right" valign="top" rowspan="1" colspan="1">33.82</td>
<td align="right" valign="top" rowspan="1" colspan="1">35.57</td>
<td align="left" valign="top" rowspan="1" colspan="1">Esan</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0105</td>
<td align="right" valign="top" rowspan="1" colspan="1">−9.4</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0068</td>
<td align="right" valign="top" rowspan="1" colspan="1">−3.9</td>
<td align="right" valign="top" rowspan="1" colspan="1">1.9</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0038</td>
<td align="right" valign="top" rowspan="1" colspan="1">7.7</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0002</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.4</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Lezgin</td>
<td align="right" valign="top" rowspan="1" colspan="1">9</td>
<td align="right" valign="top" rowspan="1" colspan="1">42.12</td>
<td align="right" valign="top" rowspan="1" colspan="1">48.18</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0100</td>
<td align="right" valign="top" rowspan="1" colspan="1">−6.0</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0100</td>
<td align="right" valign="top" rowspan="1" colspan="1">−6.0</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0013</td>
<td align="right" valign="top" rowspan="1" colspan="1">2.7</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0037</td>
<td align="right" valign="top" rowspan="1" colspan="1">−7.5</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Libyan_Jew</td>
<td align="right" valign="top" rowspan="1" colspan="1">9</td>
<td align="right" valign="top" rowspan="1" colspan="1">32.92</td>
<td align="right" valign="top" rowspan="1" colspan="1">13.18</td>
<td align="left" valign="top" rowspan="1" colspan="1">Esan</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0051</td>
<td align="right" valign="top" rowspan="1" colspan="1">−4.4</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.0000</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0</td>
<td align="right" valign="top" rowspan="1" colspan="1">2.7</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0030</td>
<td align="right" valign="top" rowspan="1" colspan="1">6.2</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0004</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.9</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Lithuanian</td>
<td align="right" valign="top" rowspan="1" colspan="1">10</td>
<td align="right" valign="top" rowspan="1" colspan="1">54.9</td>
<td align="right" valign="top" rowspan="1" colspan="1">23.92</td>
<td align="left" valign="top" rowspan="1" colspan="1">Abkhasian</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0119</td>
<td align="right" valign="top" rowspan="1" colspan="1">−14.9</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1">Abkhasian</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0119</td>
<td align="right" valign="top" rowspan="1" colspan="1">−14.9</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.0</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0069</td>
<td align="right" valign="top" rowspan="1" colspan="1">−3.9</td>
<td align="right" valign="top" rowspan="1" colspan="1">2.8</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0045</td>
<td align="right" valign="top" rowspan="1" colspan="1">−9.0</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0048</td>
<td align="right" valign="top" rowspan="1" colspan="1">−9.9</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Maltese</td>
<td align="right" valign="top" rowspan="1" colspan="1">8</td>
<td align="right" valign="top" rowspan="1" colspan="1">35.94</td>
<td align="right" valign="top" rowspan="1" colspan="1">14.38</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0086</td>
<td align="right" valign="top" rowspan="1" colspan="1">−4.9</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1">Yemenite_Jew</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0051</td>
<td align="right" valign="top" rowspan="1" colspan="1">−6.0</td>
<td align="center" valign="top" rowspan="1" colspan="1">2.0</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0086</td>
<td align="right" valign="top" rowspan="1" colspan="1">−4.9</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0013</td>
<td align="right" valign="top" rowspan="1" colspan="1">2.7</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0011</td>
<td align="right" valign="top" rowspan="1" colspan="1">−2.3</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Mordovian</td>
<td align="right" valign="top" rowspan="1" colspan="1">10</td>
<td align="right" valign="top" rowspan="1" colspan="1">54.18</td>
<td align="right" valign="top" rowspan="1" colspan="1">45.18</td>
<td align="left" valign="top" rowspan="1" colspan="1">Abkhasian</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0115</td>
<td align="right" valign="top" rowspan="1" colspan="1">−14.4</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1">Abkhasian</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0115</td>
<td align="right" valign="top" rowspan="1" colspan="1">−14.4</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.0</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0113</td>
<td align="right" valign="top" rowspan="1" colspan="1">−6.6</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.3</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0028</td>
<td align="right" valign="top" rowspan="1" colspan="1">−5.5</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0044</td>
<td align="right" valign="top" rowspan="1" colspan="1">−9.0</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Moroccan_Jew</td>
<td align="right" valign="top" rowspan="1" colspan="1">6</td>
<td align="right" valign="top" rowspan="1" colspan="1">34.02</td>
<td align="right" valign="top" rowspan="1" colspan="1">−6.84</td>
<td align="left" valign="top" rowspan="1" colspan="1">Esan</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0062</td>
<td align="right" valign="top" rowspan="1" colspan="1">−5.2</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1">Yemenite_Jew</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0021</td>
<td align="right" valign="top" rowspan="1" colspan="1">−2.2</td>
<td align="center" valign="top" rowspan="1" colspan="1">2.9</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0032</td>
<td align="right" valign="top" rowspan="1" colspan="1">−1.7</td>
<td align="right" valign="top" rowspan="1" colspan="1">1.4</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0021</td>
<td align="right" valign="top" rowspan="1" colspan="1">4.3</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0001</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.1</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">North_Ossetian</td>
<td align="right" valign="top" rowspan="1" colspan="1">10</td>
<td align="right" valign="top" rowspan="1" colspan="1">43.02</td>
<td align="right" valign="top" rowspan="1" colspan="1">44.65</td>
<td align="left" valign="top" rowspan="1" colspan="1">Piapoco</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0093</td>
<td align="right" valign="top" rowspan="1" colspan="1">−7.2</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0076</td>
<td align="right" valign="top" rowspan="1" colspan="1">−4.4</td>
<td align="right" valign="top" rowspan="1" colspan="1">1.0</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0014</td>
<td align="right" valign="top" rowspan="1" colspan="1">2.9</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0028</td>
<td align="right" valign="top" rowspan="1" colspan="1">−5.6</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Norwegian</td>
<td align="right" valign="top" rowspan="1" colspan="1">11</td>
<td align="right" valign="top" rowspan="1" colspan="1">60.36</td>
<td align="right" valign="top" rowspan="1" colspan="1">5.36</td>
<td align="left" valign="top" rowspan="1" colspan="1">Georgian</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0120</td>
<td align="right" valign="top" rowspan="1" colspan="1">−14.8</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1">Georgian</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0120</td>
<td align="right" valign="top" rowspan="1" colspan="1">−14.8</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.0</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0093</td>
<td align="right" valign="top" rowspan="1" colspan="1">−5.4</td>
<td align="right" valign="top" rowspan="1" colspan="1">1.4</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0035</td>
<td align="right" valign="top" rowspan="1" colspan="1">−7.3</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0042</td>
<td align="right" valign="top" rowspan="1" colspan="1">−8.7</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Orcadian</td>
<td align="right" valign="top" rowspan="1" colspan="1">13</td>
<td align="right" valign="top" rowspan="1" colspan="1">59</td>
<td align="right" valign="top" rowspan="1" colspan="1">−3</td>
<td align="left" valign="top" rowspan="1" colspan="1">Armenian</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0102</td>
<td align="right" valign="top" rowspan="1" colspan="1">−13.4</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0059</td>
<td align="center" valign="top" rowspan="1" colspan="1">−3.6</td>
<td align="center" valign="top" rowspan="1" colspan="1">2.5</td>
<td align="left" valign="top" rowspan="1" colspan="1">Armenian</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0102</td>
<td align="right" valign="top" rowspan="1" colspan="1">−13.4</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.0</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0098</td>
<td align="right" valign="top" rowspan="1" colspan="1">−5.9</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.5</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0032</td>
<td align="right" valign="top" rowspan="1" colspan="1">−6.7</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0042</td>
<td align="right" valign="top" rowspan="1" colspan="1">−8.6</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Palestinian</td>
<td align="right" valign="top" rowspan="1" colspan="1">38</td>
<td align="right" valign="top" rowspan="1" colspan="1">32</td>
<td align="right" valign="top" rowspan="1" colspan="1">35</td>
<td align="left" valign="top" rowspan="1" colspan="1">Esan</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0120</td>
<td align="right" valign="top" rowspan="1" colspan="1">−13.2</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0047</td>
<td align="right" valign="top" rowspan="1" colspan="1">10.2</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0014</td>
<td align="right" valign="top" rowspan="1" colspan="1">3.1</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Russian</td>
<td align="right" valign="top" rowspan="1" colspan="1">22</td>
<td align="right" valign="top" rowspan="1" colspan="1">61</td>
<td align="right" valign="top" rowspan="1" colspan="1">40</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chukchi</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0119</td>
<td align="right" valign="top" rowspan="1" colspan="1">−11.3</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1">Abkhasian</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0119</td>
<td align="right" valign="top" rowspan="1" colspan="1">−17.1</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.0</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0106</td>
<td align="right" valign="top" rowspan="1" colspan="1">−6.6</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.8</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0030</td>
<td align="right" valign="top" rowspan="1" colspan="1">−6.2</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0046</td>
<td align="right" valign="top" rowspan="1" colspan="1">−9.4</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Sardinian</td>
<td align="right" valign="top" rowspan="1" colspan="1">27</td>
<td align="right" valign="top" rowspan="1" colspan="1">40</td>
<td align="right" valign="top" rowspan="1" colspan="1">9</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">LaB</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0044</td>
<td align="right" valign="top" rowspan="1" colspan="1">−2.6</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">LaB</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0044</td>
<td align="center" valign="top" rowspan="1" colspan="1">−2.6</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.0</td>
<td align="left" valign="top" rowspan="1" colspan="1">Iraqi_Jew</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0033</td>
<td align="right" valign="top" rowspan="1" colspan="1">−4.2</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.0</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0035</td>
<td align="right" valign="top" rowspan="1" colspan="1">−2.1</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.3</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0016</td>
<td align="right" valign="top" rowspan="1" colspan="1">−3.4</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0015</td>
<td align="right" valign="top" rowspan="1" colspan="1">−3.3</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Saudi</td>
<td align="right" valign="top" rowspan="1" colspan="1">8</td>
<td align="right" valign="top" rowspan="1" colspan="1">18.49</td>
<td align="right" valign="top" rowspan="1" colspan="1">42.52</td>
<td align="left" valign="top" rowspan="1" colspan="1">Kgalagadi</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0042</td>
<td align="right" valign="top" rowspan="1" colspan="1">−3.6</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0042</td>
<td align="right" valign="top" rowspan="1" colspan="1">8.6</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0015</td>
<td align="right" valign="top" rowspan="1" colspan="1">3.1</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Scottish</td>
<td align="right" valign="top" rowspan="1" colspan="1">4</td>
<td align="right" valign="top" rowspan="1" colspan="1">56.04</td>
<td align="right" valign="top" rowspan="1" colspan="1">−3.94</td>
<td align="left" valign="top" rowspan="1" colspan="1">Iraqi_Jew</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0103</td>
<td align="right" valign="top" rowspan="1" colspan="1">−8.3</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1">Iraqi_Jew</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0103</td>
<td align="right" valign="top" rowspan="1" colspan="1">−8.3</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.0</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0090</td>
<td align="right" valign="top" rowspan="1" colspan="1">−4.7</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.7</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0034</td>
<td align="right" valign="top" rowspan="1" colspan="1">−6.4</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0045</td>
<td align="right" valign="top" rowspan="1" colspan="1">−8.7</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Sicilian</td>
<td align="right" valign="top" rowspan="1" colspan="1">11</td>
<td align="right" valign="top" rowspan="1" colspan="1">37.59</td>
<td align="right" valign="top" rowspan="1" colspan="1">13.77</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0108</td>
<td align="right" valign="top" rowspan="1" colspan="1">−6.5</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1">Yemenite_Jew</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0066</td>
<td align="right" valign="top" rowspan="1" colspan="1">−8.1</td>
<td align="center" valign="top" rowspan="1" colspan="1">2.4</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0108</td>
<td align="right" valign="top" rowspan="1" colspan="1">−6.5</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0006</td>
<td align="right" valign="top" rowspan="1" colspan="1">1.3</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0015</td>
<td align="right" valign="top" rowspan="1" colspan="1">−3.2</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Spanish</td>
<td align="right" valign="top" rowspan="1" colspan="1">53</td>
<td align="right" valign="top" rowspan="1" colspan="1">40.43</td>
<td align="right" valign="top" rowspan="1" colspan="1">−2.83</td>
<td align="left" valign="top" rowspan="1" colspan="1">Iraqi_Jew</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0126</td>
<td align="right" valign="top" rowspan="1" colspan="1">−17.8</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0104</td>
<td align="center" valign="top" rowspan="1" colspan="1">−6.8</td>
<td align="center" valign="top" rowspan="1" colspan="1">1.4</td>
<td align="left" valign="top" rowspan="1" colspan="1">Iraqi_Jew</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0126</td>
<td align="right" valign="top" rowspan="1" colspan="1">−17.8</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.0</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0120</td>
<td align="right" valign="top" rowspan="1" colspan="1">−7.6</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.3</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0019</td>
<td align="right" valign="top" rowspan="1" colspan="1">−4.2</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0024</td>
<td align="right" valign="top" rowspan="1" colspan="1">−5.2</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Spanish_North</td>
<td align="right" valign="top" rowspan="1" colspan="1">5</td>
<td align="right" valign="top" rowspan="1" colspan="1">42.8</td>
<td align="right" valign="top" rowspan="1" colspan="1">−2.7</td>
<td align="left" valign="top" rowspan="1" colspan="1">Iraqi_Jew</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0112</td>
<td align="right" valign="top" rowspan="1" colspan="1">−9.9</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0102</td>
<td align="center" valign="top" rowspan="1" colspan="1">−5.4</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.5</td>
<td align="left" valign="top" rowspan="1" colspan="1">Iraqi_Jew</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0112</td>
<td align="right" valign="top" rowspan="1" colspan="1">−9.9</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.0</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0082</td>
<td align="right" valign="top" rowspan="1" colspan="1">−4.4</td>
<td align="right" valign="top" rowspan="1" colspan="1">1.3</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0035</td>
<td align="right" valign="top" rowspan="1" colspan="1">−6.9</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0032</td>
<td align="right" valign="top" rowspan="1" colspan="1">−6.4</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Syrian</td>
<td align="right" valign="top" rowspan="1" colspan="1">8</td>
<td align="right" valign="top" rowspan="1" colspan="1">35.13</td>
<td align="right" valign="top" rowspan="1" colspan="1">36.87</td>
<td align="left" valign="top" rowspan="1" colspan="1">Esan</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0101</td>
<td align="right" valign="top" rowspan="1" colspan="1">−8.7</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0044</td>
<td align="right" valign="top" rowspan="1" colspan="1">8.6</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0012</td>
<td align="right" valign="top" rowspan="1" colspan="1">2.4</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Tunisian_Jew</td>
<td align="right" valign="top" rowspan="1" colspan="1">7</td>
<td align="right" valign="top" rowspan="1" colspan="1">36.8</td>
<td align="right" valign="top" rowspan="1" colspan="1">10.18</td>
<td align="left" valign="top" rowspan="1" colspan="1">Gambian</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0026</td>
<td align="right" valign="top" rowspan="1" colspan="1">−2.0</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0026</td>
<td align="right" valign="top" rowspan="1" colspan="1">5.2</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0002</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.5</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Turkish</td>
<td align="right" valign="top" rowspan="1" colspan="1">56</td>
<td align="right" valign="top" rowspan="1" colspan="1">39.22</td>
<td align="right" valign="top" rowspan="1" colspan="1">32.66</td>
<td align="left" valign="top" rowspan="1" colspan="1">Piapoco</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0129</td>
<td align="right" valign="top" rowspan="1" colspan="1">−11.3</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0106</td>
<td align="right" valign="top" rowspan="1" colspan="1">−6.9</td>
<td align="right" valign="top" rowspan="1" colspan="1">1.3</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0018</td>
<td align="right" valign="top" rowspan="1" colspan="1">3.8</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0019</td>
<td align="right" valign="top" rowspan="1" colspan="1">−4.0</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Turkish_Jew</td>
<td align="right" valign="top" rowspan="1" colspan="1">8</td>
<td align="right" valign="top" rowspan="1" colspan="1">41.02</td>
<td align="right" valign="top" rowspan="1" colspan="1">28.95</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0075</td>
<td align="right" valign="top" rowspan="1" colspan="1">−4.3</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1">Yemenite_Jew</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0049</td>
<td align="right" valign="top" rowspan="1" colspan="1">−5.8</td>
<td align="center" valign="top" rowspan="1" colspan="1">1.4</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0075</td>
<td align="right" valign="top" rowspan="1" colspan="1">−4.3</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0017</td>
<td align="right" valign="top" rowspan="1" colspan="1">3.6</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0006</td>
<td align="right" valign="top" rowspan="1" colspan="1">−1.3</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Tuscan</td>
<td align="right" valign="top" rowspan="1" colspan="1">8</td>
<td align="right" valign="top" rowspan="1" colspan="1">43</td>
<td align="right" valign="top" rowspan="1" colspan="1">11</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0109</td>
<td align="right" valign="top" rowspan="1" colspan="1">−6.4</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0055</td>
<td align="center" valign="top" rowspan="1" colspan="1">−3.2</td>
<td align="center" valign="top" rowspan="1" colspan="1">2.3</td>
<td align="left" valign="top" rowspan="1" colspan="1">Iraqi_Jew</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0092</td>
<td align="right" valign="top" rowspan="1" colspan="1">−10.1</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.9</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0109</td>
<td align="right" valign="top" rowspan="1" colspan="1">−6.4</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0011</td>
<td align="right" valign="top" rowspan="1" colspan="1">−2.2</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0024</td>
<td align="right" valign="top" rowspan="1" colspan="1">−5.0</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Ukrainian</td>
<td align="right" valign="top" rowspan="1" colspan="1">9</td>
<td align="right" valign="top" rowspan="1" colspan="1">50.29</td>
<td align="right" valign="top" rowspan="1" colspan="1">31.56</td>
<td align="left" valign="top" rowspan="1" colspan="1">Georgian</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0134</td>
<td align="right" valign="top" rowspan="1" colspan="1">−16.7</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1">Georgian</td>
<td align="left" valign="top" rowspan="1" colspan="1">Los</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0134</td>
<td align="right" valign="top" rowspan="1" colspan="1">−16.7</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.0</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0114</td>
<td align="right" valign="top" rowspan="1" colspan="1">−6.6</td>
<td align="right" valign="top" rowspan="1" colspan="1">1.3</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0032</td>
<td align="right" valign="top" rowspan="1" colspan="1">−6.4</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0041</td>
<td align="right" valign="top" rowspan="1" colspan="1">−8.5</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Yemenite_Jew</td>
<td align="right" valign="top" rowspan="1" colspan="1">8</td>
<td align="right" valign="top" rowspan="1" colspan="1">15.35</td>
<td align="right" valign="top" rowspan="1" colspan="1">44.2</td>
<td align="left" valign="top" rowspan="1" colspan="1">Esan</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stu</td>
<td align="left" valign="top" rowspan="1" colspan="1">−0.0027</td>
<td align="right" valign="top" rowspan="1" colspan="1">−2.4</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0046</td>
<td align="right" valign="top" rowspan="1" colspan="1">9.1</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0013</td>
<td align="right" valign="top" rowspan="1" colspan="1">2.6</td>
</tr>
</tbody>
</table>
<table-wrap-foot><fn id="TFN3"><p id="P54">Note: <italic>Zdiff</italic>
is the number of standard errors of the
difference between the lowest <italic>f<sub>3</sub>
</italic>
-statistic over all
reference pairs and the lowest <italic>f<sub>3</sub>
</italic>
-statistic for a subset
of reference pairs.</p>
</fn>
<fn id="TFN1"><p id="P55">Abbreviations used: Stu: Stuttgart; Los: Loschbour; LaB: LaBrana.</p>
</fn>
</table-wrap-foot>
</table-wrap>
<table-wrap id="T3" position="anchor" orientation="portrait"><label>Extended Data Table 2</label>
<caption><p id="P56">Confirmation of key findings on transversions and on whole genome sequence
data.</p>
</caption>
<table frame="box" rules="groups"><thead><tr><th align="left" valign="middle" rowspan="4" colspan="1">Interpretation</th>
<th colspan="8" align="center" valign="bottom" rowspan="1">D(A, B; C, D) on Human Origins genotype
data</th>
<th colspan="6" align="center" valign="bottom" rowspan="1">D(A, B; C, D) on whole genome sequence
data transversions</th>
</tr>
<tr><th colspan="14" valign="bottom" align="center" rowspan="1"><hr></hr>
</th>
</tr>
<tr><th align="left" valign="bottom" rowspan="2" colspan="1">A</th>
<th align="left" valign="bottom" rowspan="2" colspan="1">B</th>
<th align="left" valign="bottom" rowspan="2" colspan="1">C</th>
<th align="left" valign="bottom" rowspan="2" colspan="1">D</th>
<th colspan="2" align="center" valign="bottom" rowspan="1">594,924 SNPs</th>
<th colspan="2" align="center" valign="bottom" rowspan="1">110,817 transversions</th>
<th align="left" valign="bottom" rowspan="2" colspan="1">A</th>
<th align="left" valign="bottom" rowspan="2" colspan="1">B</th>
<th align="left" valign="bottom" rowspan="2" colspan="1">C</th>
<th align="left" valign="bottom" rowspan="2" colspan="1">D</th>
<th align="right" valign="bottom" rowspan="2" colspan="1">statistic</th>
<th align="right" valign="bottom" rowspan="2" colspan="1">Z</th>
</tr>
<tr><th align="center" valign="bottom" rowspan="1" colspan="1">statistic</th>
<th align="right" valign="bottom" rowspan="1" colspan="1">Z</th>
<th align="right" valign="bottom" rowspan="1" colspan="1">statistic</th>
<th align="right" valign="bottom" rowspan="1" colspan="1">Z</th>
</tr>
</thead>
<tbody><tr><td align="left" valign="middle" rowspan="1" colspan="1">Stuttgart has Near Eastern ancestry</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stuttgart</td>
<td align="left" valign="top" rowspan="1" colspan="1">Armenian</td>
<td align="left" valign="top" rowspan="1" colspan="1">Loschbour</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.0219</td>
<td align="right" valign="top" rowspan="1" colspan="1">4.5</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0189</td>
<td align="right" valign="top" rowspan="1" colspan="1">2.9</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
</tr>
<tr><td colspan="15" valign="bottom" align="left" rowspan="1"><hr></hr>
</td>
</tr>
<tr><td align="left" valign="middle" rowspan="2" colspan="1">Europeans have more WHG-related ancestry
than Stuttgart</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stuttgart</td>
<td align="left" valign="top" rowspan="1" colspan="1">French</td>
<td align="left" valign="top" rowspan="1" colspan="1">Loschbour</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0266</td>
<td align="right" valign="top" rowspan="1" colspan="1">−5.7</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.031</td>
<td align="right" valign="top" rowspan="1" colspan="1">−5.0</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stuttgart</td>
<td align="left" valign="top" rowspan="1" colspan="1">French2</td>
<td align="left" valign="top" rowspan="1" colspan="1">Loschbour</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.03</td>
<td align="right" valign="top" rowspan="1" colspan="1">−4.7</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Lithuanian</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stuttgart</td>
<td align="left" valign="top" rowspan="1" colspan="1">Loschbour</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.0446</td>
<td align="right" valign="top" rowspan="1" colspan="1">9.1</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0477</td>
<td align="right" valign="top" rowspan="1" colspan="1">7.2</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
</tr>
<tr><td colspan="15" valign="bottom" align="left" rowspan="1"><hr></hr>
</td>
</tr>
<tr><td align="left" valign="middle" rowspan="2" colspan="1">West Eurasians have more ANE-related
ancestry than Stuttgart</td>
<td align="left" valign="top" rowspan="1" colspan="1">French</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stuttgart</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.0367</td>
<td align="right" valign="top" rowspan="1" colspan="1">7.7</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0386</td>
<td align="right" valign="top" rowspan="1" colspan="1">5.5</td>
<td align="left" valign="top" rowspan="1" colspan="1">French2</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stuttgart</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.037</td>
<td align="right" valign="top" rowspan="1" colspan="1">6.4</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Lezgin</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stuttgart</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.0372</td>
<td align="right" valign="top" rowspan="1" colspan="1">7.6</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0409</td>
<td align="right" valign="top" rowspan="1" colspan="1">5.6</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
</tr>
<tr><td colspan="15" valign="bottom" align="left" rowspan="1"><hr></hr>
</td>
</tr>
<tr><td align="left" valign="middle" rowspan="1" colspan="1">MA1 is a better surrogate of ANE ancestry than
Karitiana</td>
<td align="left" valign="top" rowspan="1" colspan="1">French</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="left" valign="top" rowspan="1" colspan="1">Karitiana</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.0207</td>
<td align="right" valign="top" rowspan="1" colspan="1">4.5</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0214</td>
<td align="right" valign="top" rowspan="1" colspan="1">2.8</td>
<td align="left" valign="top" rowspan="1" colspan="1">French2</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="left" valign="top" rowspan="1" colspan="1">Karitiana2</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.026</td>
<td align="right" valign="top" rowspan="1" colspan="1">3.8</td>
</tr>
<tr><td colspan="15" valign="bottom" align="left" rowspan="1"><hr></hr>
</td>
</tr>
<tr><td align="left" valign="middle" rowspan="9" colspan="1">Eastern non-Africans closer to
WHG/ANE/SHG than to EEF</td>
<td align="left" valign="top" rowspan="1" colspan="1">Loschbour</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stuttgart</td>
<td align="left" valign="top" rowspan="1" colspan="1">Onge</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.0196</td>
<td align="right" valign="top" rowspan="1" colspan="1">3.5</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0202</td>
<td align="right" valign="top" rowspan="1" colspan="1">2.5</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Loschbour</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stuttgart</td>
<td align="left" valign="top" rowspan="1" colspan="1">Papuan</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.0142</td>
<td align="right" valign="top" rowspan="1" colspan="1">2.6</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0127</td>
<td align="right" valign="top" rowspan="1" colspan="1">1.5</td>
<td align="left" valign="top" rowspan="1" colspan="1">Loschbour</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stuttgart</td>
<td align="left" valign="top" rowspan="1" colspan="1">Papuan2</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.017</td>
<td align="right" valign="top" rowspan="1" colspan="1">2.7</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Loschbour</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stuttgart</td>
<td align="left" valign="top" rowspan="1" colspan="1">Dai</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.0164</td>
<td align="right" valign="top" rowspan="1" colspan="1">3.2</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.021</td>
<td align="right" valign="top" rowspan="1" colspan="1">2.8</td>
<td align="left" valign="top" rowspan="1" colspan="1">Loschbour</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stuttgart</td>
<td align="left" valign="top" rowspan="1" colspan="1">Dai2</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.018</td>
<td align="right" valign="top" rowspan="1" colspan="1">2.9</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stuttgart</td>
<td align="left" valign="top" rowspan="1" colspan="1">Papuan</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.0139</td>
<td align="right" valign="top" rowspan="1" colspan="1">2.2</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0103</td>
<td align="right" valign="top" rowspan="1" colspan="1">1.0</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stuttgart</td>
<td align="left" valign="top" rowspan="1" colspan="1">Papuan2</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.018</td>
<td align="right" valign="top" rowspan="1" colspan="1">2.8</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stuttgart</td>
<td align="left" valign="top" rowspan="1" colspan="1">Dai</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.0174</td>
<td align="right" valign="top" rowspan="1" colspan="1">3.0</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.016</td>
<td align="right" valign="top" rowspan="1" colspan="1">1.7</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stuttgart</td>
<td align="left" valign="top" rowspan="1" colspan="1">Dai2</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.028</td>
<td align="right" valign="top" rowspan="1" colspan="1">4.3</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Motala12</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stuttgart</td>
<td align="left" valign="top" rowspan="1" colspan="1">Papuan</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.0182</td>
<td align="right" valign="top" rowspan="1" colspan="1">3.2</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.011</td>
<td align="right" valign="top" rowspan="1" colspan="1">1.1</td>
<td align="left" valign="top" rowspan="1" colspan="1">Motala12</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stuttgart</td>
<td align="left" valign="top" rowspan="1" colspan="1">Papuan2</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.023</td>
<td align="right" valign="top" rowspan="1" colspan="1">3.7</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Motala12</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stuttgart</td>
<td align="left" valign="top" rowspan="1" colspan="1">Dai</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.0156</td>
<td align="right" valign="top" rowspan="1" colspan="1">2.8</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0149</td>
<td align="right" valign="top" rowspan="1" colspan="1">1.6</td>
<td align="left" valign="top" rowspan="1" colspan="1">Motala12</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stuttgart</td>
<td align="left" valign="top" rowspan="1" colspan="1">Dai2</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.02</td>
<td align="right" valign="top" rowspan="1" colspan="1">3.2</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">LaBrana</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stuttgart</td>
<td align="left" valign="top" rowspan="1" colspan="1">Papuan</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.0123</td>
<td align="right" valign="top" rowspan="1" colspan="1">2.3</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0101</td>
<td align="right" valign="top" rowspan="1" colspan="1">1.1</td>
<td align="left" valign="top" rowspan="1" colspan="1">LaBrana</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stuttgart</td>
<td align="left" valign="top" rowspan="1" colspan="1">Papuan2</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.02</td>
<td align="right" valign="top" rowspan="1" colspan="1">3.2</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">LaBrana</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stuttgart</td>
<td align="left" valign="top" rowspan="1" colspan="1">Dai</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.0149</td>
<td align="right" valign="top" rowspan="1" colspan="1">2.9</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0228</td>
<td align="right" valign="top" rowspan="1" colspan="1">2.5</td>
<td align="left" valign="top" rowspan="1" colspan="1">LaBrana</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stuttgart</td>
<td align="left" valign="top" rowspan="1" colspan="1">Dai2</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.024</td>
<td align="right" valign="top" rowspan="1" colspan="1">3.7</td>
</tr>
<tr><td colspan="15" valign="bottom" align="left" rowspan="1"><hr></hr>
</td>
</tr>
<tr><td align="left" valign="middle" rowspan="1" colspan="1">Native Americans closer to ANE than to WHG</td>
<td align="left" valign="top" rowspan="1" colspan="1">Karitiana</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="left" valign="top" rowspan="1" colspan="1">Loschbour</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.0467</td>
<td align="right" valign="top" rowspan="1" colspan="1">7.1</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0467</td>
<td align="right" valign="top" rowspan="1" colspan="1">4.4</td>
<td align="left" valign="top" rowspan="1" colspan="1">Karitiana2</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="left" valign="top" rowspan="1" colspan="1">Loschbour</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.052</td>
<td align="right" valign="top" rowspan="1" colspan="1">7.1</td>
</tr>
<tr><td colspan="15" valign="bottom" align="left" rowspan="1"><hr></hr>
</td>
</tr>
<tr><td align="left" valign="middle" rowspan="2" colspan="1">West Eurasians closer to Native Americans
than to other Eastern non-Africans</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stuttgart</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="left" valign="top" rowspan="1" colspan="1">Karitiana</td>
<td align="left" valign="top" rowspan="1" colspan="1">Papuan</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.0559</td>
<td align="right" valign="top" rowspan="1" colspan="1">10.9</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0474</td>
<td align="right" valign="top" rowspan="1" colspan="1">6.6</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stuttgart</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="left" valign="top" rowspan="1" colspan="1">Karitiana2</td>
<td align="left" valign="top" rowspan="1" colspan="1">Papuan2</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.052</td>
<td align="right" valign="top" rowspan="1" colspan="1">7.6</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Stuttgart</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="left" valign="top" rowspan="1" colspan="1">Karitiana</td>
<td align="left" valign="top" rowspan="1" colspan="1">Onge</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.0237</td>
<td align="right" valign="top" rowspan="1" colspan="1">5.1</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0179</td>
<td align="right" valign="top" rowspan="1" colspan="1">2.6</td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="left" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
</tr>
<tr><td colspan="15" valign="bottom" align="left" rowspan="1"><hr></hr>
</td>
</tr>
<tr><td align="left" valign="middle" rowspan="6" colspan="1">Ancient Eurasian hunter-gatherers equally
related to Eastern non-Africans other than Native Americans</td>
<td align="left" valign="top" rowspan="1" colspan="1">Loschbour</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="left" valign="top" rowspan="1" colspan="1">Dai</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0015</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.2</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0016</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.2</td>
<td align="left" valign="top" rowspan="1" colspan="1">Loschbour</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="left" valign="top" rowspan="1" colspan="1">Dai2</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.013</td>
<td align="right" valign="top" rowspan="1" colspan="1">−1.9</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Loschbour</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="left" valign="top" rowspan="1" colspan="1">Papuan</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.0002</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0012</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.1</td>
<td align="left" valign="top" rowspan="1" colspan="1">Loschbour</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="left" valign="top" rowspan="1" colspan="1">Papuan2</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.003</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.4</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Loschbour</td>
<td align="left" valign="top" rowspan="1" colspan="1">Motala12</td>
<td align="left" valign="top" rowspan="1" colspan="1">Dai</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.0024</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.4</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.009</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.9</td>
<td align="left" valign="top" rowspan="1" colspan="1">Loschbour</td>
<td align="left" valign="top" rowspan="1" colspan="1">Motala12</td>
<td align="left" valign="top" rowspan="1" colspan="1">Dai2</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.002</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.3</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Loschbour</td>
<td align="left" valign="top" rowspan="1" colspan="1">Motala12</td>
<td align="left" valign="top" rowspan="1" colspan="1">Papuan</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0028</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.4</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0046</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.5</td>
<td align="left" valign="top" rowspan="1" colspan="1">Loschbour</td>
<td align="left" valign="top" rowspan="1" colspan="1">Motala12</td>
<td align="left" valign="top" rowspan="1" colspan="1">Papuan2</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.004</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.6</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="left" valign="top" rowspan="1" colspan="1">Motala12</td>
<td align="left" valign="top" rowspan="1" colspan="1">Dai</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.0026</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.4</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0047</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.4</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="left" valign="top" rowspan="1" colspan="1">Motala12</td>
<td align="left" valign="top" rowspan="1" colspan="1">Dai2</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.01</td>
<td align="right" valign="top" rowspan="1" colspan="1">1.5</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="left" valign="top" rowspan="1" colspan="1">Motala12</td>
<td align="left" valign="top" rowspan="1" colspan="1">Papuan</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0047</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.7</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.001</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.1</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="left" valign="top" rowspan="1" colspan="1">Motala12</td>
<td align="left" valign="top" rowspan="1" colspan="1">Papuan2</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.004</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.5</td>
</tr>
<tr><td colspan="15" valign="bottom" align="left" rowspan="1"><hr></hr>
</td>
</tr>
<tr><td align="left" valign="middle" rowspan="3" colspan="1">LaBrana and Loschbour are a clade</td>
<td align="left" valign="top" rowspan="1" colspan="1">LaBrana</td>
<td align="left" valign="top" rowspan="1" colspan="1">Loschbour</td>
<td align="left" valign="top" rowspan="1" colspan="1">Dai</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0028</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.5</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0024</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.3</td>
<td align="left" valign="top" rowspan="1" colspan="1">LaBrana</td>
<td align="left" valign="top" rowspan="1" colspan="1">Loschbour</td>
<td align="left" valign="top" rowspan="1" colspan="1">Dai2</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.007</td>
<td align="right" valign="top" rowspan="1" colspan="1">1.1</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">LaBrana</td>
<td align="left" valign="top" rowspan="1" colspan="1">Loschbour</td>
<td align="left" valign="top" rowspan="1" colspan="1">Papuan</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0031</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.5</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0012</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.1</td>
<td align="left" valign="top" rowspan="1" colspan="1">LaBrana</td>
<td align="left" valign="top" rowspan="1" colspan="1">Loschbour</td>
<td align="left" valign="top" rowspan="1" colspan="1">Papuan2</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.002</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.3</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">LaBrana</td>
<td align="left" valign="top" rowspan="1" colspan="1">Loschbour</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.006</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.8</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0101</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.7</td>
<td align="left" valign="top" rowspan="1" colspan="1">LaBrana</td>
<td align="left" valign="top" rowspan="1" colspan="1">Loschbour</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.005</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.7</td>
</tr>
<tr><td colspan="15" valign="bottom" align="left" rowspan="1"><hr></hr>
</td>
</tr>
<tr><td align="left" valign="middle" rowspan="2" colspan="1">SHG closer to ANE than to WHG</td>
<td align="left" valign="top" rowspan="1" colspan="1">Motala12</td>
<td align="left" valign="top" rowspan="1" colspan="1">Loschbour</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.0425</td>
<td align="right" valign="top" rowspan="1" colspan="1">5.3</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0353</td>
<td align="right" valign="top" rowspan="1" colspan="1">2.6</td>
<td align="left" valign="top" rowspan="1" colspan="1">Motala12</td>
<td align="left" valign="top" rowspan="1" colspan="1">Loschbour</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.042</td>
<td align="right" valign="top" rowspan="1" colspan="1">5.9</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Motala12</td>
<td align="left" valign="top" rowspan="1" colspan="1">LaBrana</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.0465</td>
<td align="right" valign="top" rowspan="1" colspan="1">5.8</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.0347</td>
<td align="right" valign="top" rowspan="1" colspan="1">2.4</td>
<td align="left" valign="top" rowspan="1" colspan="1">Motala12</td>
<td align="left" valign="top" rowspan="1" colspan="1">LaBrana</td>
<td align="left" valign="top" rowspan="1" colspan="1">MA1</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.038</td>
<td align="right" valign="top" rowspan="1" colspan="1">5.4</td>
</tr>
<tr><td colspan="15" valign="bottom" align="left" rowspan="1"><hr></hr>
</td>
</tr>
<tr><td align="left" valign="middle" rowspan="1" colspan="1">LaBrana and Loschbour equally related to
Stuttgart</td>
<td align="left" valign="top" rowspan="1" colspan="1">LaBrana</td>
<td align="left" valign="top" rowspan="1" colspan="1">Loschbour</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stuttgart</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="center" valign="top" rowspan="1" colspan="1">−0.0176</td>
<td align="right" valign="top" rowspan="1" colspan="1">−2.6</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.0106</td>
<td align="right" valign="top" rowspan="1" colspan="1">−1.0</td>
<td align="left" valign="top" rowspan="1" colspan="1">LaBrana</td>
<td align="left" valign="top" rowspan="1" colspan="1">Loschbour</td>
<td align="left" valign="top" rowspan="1" colspan="1">Stuttgart</td>
<td align="left" valign="top" rowspan="1" colspan="1">Chimp</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.012</td>
<td align="right" valign="top" rowspan="1" colspan="1">−1.8</td>
</tr>
</tbody>
</table>
</table-wrap>
<table-wrap id="T4" position="anchor" orientation="portrait"><label>Extended Data Table 3</label>
<caption><p id="P57">Admixture proportions for European populations. The estimates from the model
with minimal assumptions are from SI17. The estimates from the full modeling are from
SI14 either by single population analysis or co-fitting population pairs and averaging
over fits (these averages are the results plotted in <xref rid="F2" ref-type="fig">Fig.
2B</xref>
). Populations that do not fit the models are not reported.</p>
</caption>
<table frame="box" rules="groups"><thead><tr><th valign="bottom" rowspan="3" align="left" colspan="1"></th>
<th colspan="3" valign="bottom" align="center" rowspan="1">Full modeling of<break></break>
population
relationships<break></break>
(individual fits)</th>
<th colspan="6" valign="bottom" align="center" rowspan="1">Full modeling of<break></break>
population
relationships<break></break>
(averaged fits)</th>
<th colspan="3" valign="bottom" align="center" rowspan="1">Modeling of
population<break></break>
relationships with<break></break>
minimal assumptions</th>
<th colspan="3" valign="bottom" align="center" rowspan="1">Model-based (averaged)<break></break>
- Model
with minimal<break></break>
assumptions (Z-score)</th>
</tr>
<tr><th valign="bottom" align="right" rowspan="1" colspan="1">EEF</th>
<th valign="bottom" align="right" rowspan="1" colspan="1">WHG</th>
<th valign="bottom" align="right" rowspan="1" colspan="1">ANE</th>
<th colspan="2" valign="bottom" align="center" rowspan="1">EEF</th>
<th colspan="2" valign="bottom" align="center" rowspan="1">WHG</th>
<th colspan="2" valign="bottom" align="center" rowspan="1">ANE</th>
<th valign="bottom" align="right" rowspan="1" colspan="1">EEF</th>
<th valign="bottom" align="right" rowspan="1" colspan="1">WHG</th>
<th valign="bottom" align="center" rowspan="1" colspan="1">ANE</th>
<th valign="bottom" align="right" rowspan="1" colspan="1">EEF</th>
<th valign="bottom" align="right" rowspan="1" colspan="1">WHG</th>
<th valign="bottom" align="right" rowspan="1" colspan="1">ANE</th>
</tr>
<tr><th valign="bottom" align="right" rowspan="1" colspan="1"></th>
<th valign="bottom" align="right" rowspan="1" colspan="1"></th>
<th valign="bottom" align="right" rowspan="1" colspan="1"></th>
<th valign="bottom" align="center" rowspan="1" colspan="1">Mean</th>
<th valign="bottom" align="center" rowspan="1" colspan="1">Range</th>
<th valign="bottom" align="right" rowspan="1" colspan="1">Mean</th>
<th valign="bottom" align="center" rowspan="1" colspan="1">Range</th>
<th valign="bottom" align="right" rowspan="1" colspan="1">Mean</th>
<th valign="bottom" align="center" rowspan="1" colspan="1">Range</th>
<th valign="bottom" align="right" rowspan="1" colspan="1"></th>
<th valign="bottom" align="right" rowspan="1" colspan="1"></th>
<th valign="bottom" align="center" rowspan="1" colspan="1"></th>
<th valign="bottom" align="right" rowspan="1" colspan="1"></th>
<th valign="bottom" align="right" rowspan="1" colspan="1"></th>
<th valign="bottom" align="right" rowspan="1" colspan="1"></th>
</tr>
</thead>
<tbody><tr><td align="left" valign="top" rowspan="1" colspan="1">Albanian</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.781</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.092</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.127</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.781</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.772–0.819</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.082</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.032–0.098</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.137</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.129–0.158</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.595 ± 0.112</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.353 ± 0.150</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.052 ± 0.049</td>
<td align="right" valign="top" rowspan="1" colspan="1">1.658</td>
<td align="right" valign="top" rowspan="1" colspan="1">−1.807</td>
<td align="right" valign="top" rowspan="1" colspan="1">1.741</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Ashkenazi_Jew</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.931</td>
<td align="right" valign="top" rowspan="1" colspan="1">0</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.069</td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1">0.938 ± 0.146</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.021 ± 0.185</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.083 ± 0.049</td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Basque</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.593</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.293</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.114</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.569</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.527–0.616</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.335</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.255–0.392</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.096</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.076–0.129</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.569 ± 0.091</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.315 ± 0.124</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.115 ± 0.041</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.001</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.165</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.472</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Belarusian</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.418</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.431</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.151</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.426</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.397–0.464</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.408</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.338–0.443</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.167</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.150–0.199</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.272 ± 0.094</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.554 ± 0.131</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.174 ± 0.047</td>
<td align="right" valign="top" rowspan="1" colspan="1">1.637</td>
<td align="right" valign="top" rowspan="1" colspan="1">−1.118</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.158</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Bergamo</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.715</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.177</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.108</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.721</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.704–0.793</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.163</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.061–0.189</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.117</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.104–0.147</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.644 ± 0.125</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.248 ± 0.170</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.108 ± 0.053</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.615</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.503</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.162</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Bulgarian</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.712</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.147</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.141</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.718</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.707–0.778</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.132</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.047–0.151</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.151</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.138–0.175</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.556 ± 0.110</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.328 ± 0.143</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.116 ± 0.043</td>
<td align="right" valign="top" rowspan="1" colspan="1">1.469</td>
<td align="right" valign="top" rowspan="1" colspan="1">−1.372</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.804</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Croatian</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.561</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.293</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.145</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.564</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.548–0.586</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.285</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.242–0.310</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.151</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.137–0.172</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.453 ± 0.122</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.407 ± 0.159</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.140 ± 0.046</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.911</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.768</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.238</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Czech</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.495</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.338</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.167</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.489</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.460–0.531</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.348</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.273–0.382</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.163</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.145–0.196</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.402 ± 0.117</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.400 ± 0.162</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.198 ± 0.050</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.744</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.322</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.698</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">English</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.495</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.364</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.141</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.503</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.476–0.536</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.353</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.296–0.382</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.144</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.130–0.169</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.475 ± 0.091</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.357 ± 0.125</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.168 ± 0.043</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.304</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.028</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.561</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Estonian</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.322</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.495</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.183</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.323</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.293–0.345</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.49</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.451–0.520</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.187</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.172–0.205</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.072 ± 0.121</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.778 ± 0.176</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.150 ± 0.064</td>
<td align="right" valign="top" rowspan="1" colspan="1">2.070</td>
<td align="right" valign="top" rowspan="1" colspan="1">−1.636</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.584</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">French</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.554</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.311</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.135</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.563</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.537–0.601</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.297</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.230–0.328</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.14</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.126–0.169</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.498 ± 0.097</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.359 ± 0.127</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.142 ± 0.039</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.672</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.487</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.060</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">French_South</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.675</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.195</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.13</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.636</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.589–0.738</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.256</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.111–0.323</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.108</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.088–0.151</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.636 ± 0.116</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.225 ± 0.165</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.140 ± 0.057</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.003</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.189</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.558</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Greek</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.792</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.058</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.151</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.791</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.780–0.816</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.048</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.019–0.060</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.161</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.150–0.171</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.658 ± 0.098</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.255 ± 0.127</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.086 ± 0.039</td>
<td align="right" valign="top" rowspan="1" colspan="1">1.357</td>
<td align="right" valign="top" rowspan="1" colspan="1">−1.627</td>
<td align="right" valign="top" rowspan="1" colspan="1">1.915</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Hungarian</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.558</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.264</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.179</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.548</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.520–0.590</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.279</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.199–0.313</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.174</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.156–0.210</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.391 ± 0.109</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.454 ± 0.153</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.155 ± 0.050</td>
<td align="right" valign="top" rowspan="1" colspan="1">1.437</td>
<td align="right" valign="top" rowspan="1" colspan="1">−1.145</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.371</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Icelandic</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.394</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.456</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.15</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.409</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.386–0.424</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.448</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.409–0.473</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.143</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.126–0.170</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.342 ± 0.102</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.476 ± 0.137</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.182 ± 0.045</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.654</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.204</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.861</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Lithuanian</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.364</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.464</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.172</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.352</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.327–0.384</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.488</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.433–0.527</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.16</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.135–0.184</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.248 ± 0.117</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.548 ± 0.163</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.205 ± 0.052</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.886</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.367</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.864</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Maltese</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.932</td>
<td align="right" valign="top" rowspan="1" colspan="1">0</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.068</td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1">1.298 ± 0.185</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.509 ± 0.248</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.211 ± 0.079</td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Norwegian</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.411</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.428</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.161</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.417</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.388–0.438</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.423</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.383–0.450</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.16</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.140–0.181</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.273 ± 0.115</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.557 ± 0.161</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.170 ± 0.055</td>
<td align="right" valign="top" rowspan="1" colspan="1">1.252</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.831</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.185</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Orcadian</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.457</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.385</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.158</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.465</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.439–0.493</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.378</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.329–0.403</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.157</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.140–0.179</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.395 ± 0.088</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.437 ± 0.122</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.168 ± 0.041</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.798</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.487</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.264</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Sardinian</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.817</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.175</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.008</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.818</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.791–0.874</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.141</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.058–0.182</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.041</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.026–0.068</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.883 ± 0.128</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.075 ± 0.166</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.042 ± 0.048</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.510</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.400</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.024</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Scottish</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.39</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.428</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.182</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.408</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.387–0.424</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.421</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.384–0.448</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.171</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.149–0.201</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.286 ± 0.112</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.532 ± 0.156</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.182 ± 0.053</td>
<td align="right" valign="top" rowspan="1" colspan="1">1.091</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.712</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.210</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Sicilian</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.903</td>
<td align="right" valign="top" rowspan="1" colspan="1">0</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.097</td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1">1.012 ± 0.149</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.131 ± 0.199</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.119 ± 0.060</td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Spanish</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.809</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.068</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.123</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.759</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.736–0.804</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.126</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.066–0.170</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.115</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.091–0.151</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.856 ± 0.126</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.015 ± 0.165</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.160 ± 0.049</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.769</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.855</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.922</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Spanish_North</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.713</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.125</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.163</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.612</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.561–0.660</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.292</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.214–0.365</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.096</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.072–0.126</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.581 ± 0.120</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.298 ± 0.158</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.121 ± 0.046</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.254</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.038</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.533</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Tuscan</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.746</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.136</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.118</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.751</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.737–0.806</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.123</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.047–0.145</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.126</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.114–0.150</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.734 ± 0.118</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.153 ± 0.160</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.113 ± 0.054</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.141</td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.188</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.249</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Ukrainian</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.462</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.387</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.151</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.463</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.445–0.491</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.376</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.322–0.399</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.16</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.148–0.187</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.259 ± 0.123</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.596 ± 0.173</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.145 ± 0.057</td>
<td align="right" valign="top" rowspan="1" colspan="1">1.661</td>
<td align="right" valign="top" rowspan="1" colspan="1">−1.269</td>
<td align="right" valign="top" rowspan="1" colspan="1">0.269</td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Finnish</td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.299 ± 0.204</td>
<td align="right" valign="top" rowspan="1" colspan="1">1.194 ± 0.296</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.105 ± 0.105</td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Mordovian</td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.255 ± 0.173</td>
<td align="right" valign="top" rowspan="1" colspan="1">1.151 ± 0.246</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.104 ± 0.090</td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
</tr>
<tr><td align="left" valign="top" rowspan="1" colspan="1">Russian</td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="center" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1">−0.303 ± 0.211</td>
<td align="right" valign="top" rowspan="1" colspan="1">1.230 ± 0.301</td>
<td align="center" valign="top" rowspan="1" colspan="1">0.072 ± 0.106</td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
<td align="right" valign="top" rowspan="1" colspan="1"></td>
</tr>
</tbody>
</table>
</table-wrap>
</sec>
<sec sec-type="supplementary-material" id="S29"><title>Supplementary Material</title>
<supplementary-material content-type="local-data" id="SD1"><media xlink:href="NIHMS613260-supplement-supplement_1.pdf" orientation="portrait" xlink:type="simple" id="d37e8281" position="anchor"></media>
</supplementary-material>
</sec>
</body>
<back><ack id="S27"><p>We are grateful to Cynthia Beall, Neil Bradman, Amha Gebremedhin, Damian Labuda, Mari Nelis
and Anna Di Rienzo for sharing DNA samples; to Detlef Weigel, Christa Lanz, Verena
Schünemann, Peter Bauer and Olaf Riess for support and access to DNA sequencing
facilities; to Philip Johnson for advice on contamination estimation; to Garrett Hellenthal
for help with the ChromoPainter software; and to Pontus Skoglund for sharing graphics
software. We thank Kenneth Nordtvedt for alerting us to newly discovered Y-chromosome SNPs.
We downloaded the POPRES data from dbGaP at <ext-link ext-link-type="uri" xlink:href="http://www.ncbi.nlm.nih.gov/projects/gap/cgi-bin/study.cgi?study_id=phs000145.v4.p2">http://www.ncbi.nlm.nih.gov/projects/gap/cgi-bin/study.cgi?study_id=phs000145.v4.p2</ext-link>
through dbGaP accession number phs000145.v1.p2. We thank all the volunteers who donated DNA.
We thank the staff of the Unità Operativa Complessa di Medicina Trasfusionale,
Azienda Ospedaliera Umberto I, Siracusa, Italy for assistance in sample collection; and The
National Laboratory for the Genetics of Israeli Populations for facilitating access to DNA.
We thank colleagues at the Applied Genomics at the Children’s Hospital of
Philadelphia, especially Hakon Hakonarson, Cecilia Kim, Kelly Thomas, and Cuiping Hou, for
genotyping samples on the Human Origins array. JKr is grateful for support from DFG grant
# KR 4015/1-1, the Carl-Zeiss Foundation and the Baden Württemberg
Foundation. SP, GR, QF, CF, KP, SC and JKe acknowledge support from the Presidential
Innovation Fund of the Max Planck Society. GR was supported by an NSERC fellowship. JGS
acknowledges use of the Extreme Science and Engineering Discovery Environment (XSEDE), which
is supported by NSF grant number OCI-1053575. EB and OB were supported by RFBR grants
13-06-00670, 13-04-01711, 13-04-90420 and by the Molecular and Cell Biology Program of the
Presidium, Russian Academy of Sciences. BM was supported by grants OTKA 73430 and 103983.
ASaj was supported by a Finnish Professorpool (Paulo Foundation) Grant. The Lithuanian
sampling was supported by the LITGEN project (VP1-3.1-ŠMM-07-K-01-013), funded by
the European Social Fund under the Global Grant Measure. AS was supported by Spanish grants
SAF2011-26983 and EM 2012/045. OU was supported by Ukrainian SFFS grant F53.4/071. SAT was
supported by NIH Pioneer Award 8DP1ES022577-04 and NSF HOMINID award BCS-0827436. KT was
supported by an Indian CSIR Network Project (GENESIS: BSC0121). LS was supported by an
Indian CSIR Bhatnagar Fellowship. RV, MM, JP and EM were supported by the European Union
Regional Development Fund through the Centre of Excellence in Genomics to the Estonian
Biocentre and University of Tartu and by an Estonian Basic Research grant SF0270177As08. MM
was additionally supported by Estonian Science Foundation grant #8973. JGS and MS
were supported by NIH grant GM40282. PHS and EEE were supported by NIH grants HG004120 and
HG002385. DR and NP were supported by NSF HOMINID award BCS-1032255 and NIH grant GM100233.
DR and EEE are Howard Hughes Medical Institute investigators. This project has been funded
in part with federal funds from the National Cancer Institute, National Institutes of
Health, under contract HHSN26120080001E. The content of this publication does not
necessarily reflect the views or policies of the Department of Health and Human Services,
nor does mention of trade names, commercial products, or organizations imply endorsement by
the U.S. Government. This Research was supported in part by the Intramural Research Program
of the NIH, National Cancer Institute, Center for Cancer Research.</p>
</ack>
<fn-group><fn id="FN4"><p><xref rid="SD1" ref-type="supplementary-material">Supplementary Information</xref>
is
linked to the online version of the paper at <ext-link ext-link-type="uri" xlink:href="www.nature.com/nature">www.nature.com/nature</ext-link>
. The fully public
version of the Human Origins dataset can be found at <ext-link ext-link-type="uri" xlink:href="http://genetics.med.harvard.edu/reichlab/Reich_Lab/Datasets.html">http://genetics.med.harvard.edu/reichlab/Reich_Lab/Datasets.html</ext-link>
. The full
version of the dataset (including additional samples) is available to researchers who send
a signed letter to DR indicating that they will abide by specified usage conditions
(SI9).</p>
</fn>
<fn id="FN5" fn-type="con"><p><bold>Author contributions</bold>
</p>
<p>BB, EEE, JBu, MS, SP, JKe, DR and JKr supervised the study. IL, NP, AM, GR, SM, KK, PHS,
JGS, SC, ML, QF, HL, CdF, KP, WH, MMet, MMey and DR analyzed genetic data. FH, EF, DD, MF,
J-MG, JW, AC and JKr obtained human remains. AM, CE, RBo, KB, SS, CP, NR and JKr processed
ancient DNA. IL, NP, SN, NR, GA, HAB, GBa, EB, OB, RBa, GBe, HB-A, JBe, FBe, CMB, FBr,
GBJB, FC, MC, DECC, DCor, LD, GvD, SD, J-MD, SAF, IGR, MG, MH, BH, TH, UH, ARJ, SK-Y, RKh,
EK, RKi, TK, WK, VK, AK, LL, SL, TL, RWM, BM, EM, JMol, JMou, KN, DN, TN, LO, JP, FP, OLP,
VR, FR, IR, RR, HS, ASaj, ASal, EBS, ATar, DT, ST, IU, OU, RVa, MVi, MVo, CW, LY, PZ, TZ,
CC, MGT, AR-L, SAT, LS, KT, RVi, DCom, RS, MMet, SP and DR assembled the genotyping
dataset. IL, NP, DR and JKr wrote the manuscript with help from all co-authors.</p>
</fn>
<fn id="FN6"><p>The aligned sequences are available through the European Nucleotide Archive under
accession number PRJEB6272.</p>
</fn>
<fn id="FN7" fn-type="conflict"><p>The authors declare competing financial interests: UH is an employee of Illumina, TL is
an employee of AMGEN, and JM is an employee of 23andMe.</p>
</fn>
</fn-group>
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<floats-group><fig id="F1" orientation="portrait" position="float"><label>Figure 1</label>
<caption><title>Map of West Eurasian populations and Principal Component Analysis</title>
<p>(a) Geographical locations of analyzed samples, with color coding matching the PCA. We
show all sampling locations for each population, which results in multiple points for some
(e.g., Spain). (b) PCA on all present-day West Eurasians, with ancient and selected
eastern non-African samples projected. European hunter-gatherers fall beyond present-day
Europeans in the direction of European differentiation from the Near East. Stuttgart
clusters with other Neolithic Europeans and present-day Sardinians. MA1 falls outside the
variation of present-day West Eurasians in the direction of southern-northern
differentiation along dimension 2.</p>
</caption>
<graphic xlink:href="nihms613260f1a"></graphic>
<graphic xlink:href="nihms613260f1b"></graphic>
</fig>
<fig id="F2" orientation="portrait" position="float"><label>Figure 2</label>
<caption><title>Modeling of West Eurasian population history</title>
<p>(a) A three-way mixture model that is a fit to the data for many populations. Present-day
samples are colored in blue, ancient in red, and reconstructed ancestral populations in
green. Solid lines represent descent without mixture, and dashed lines represent
admixture. We print mixture proportions and one standard error for the two mixtures
relating the highly divergent ancestral populations. (We do not print the estimate for the
“European” population as it varies depending on the population). (b) We
plot the proportions of ancestry from each of three inferred ancestral populations (EEF,
ANE and WHG).</p>
</caption>
<graphic xlink:href="nihms613260f2a"></graphic>
<graphic xlink:href="nihms613260f2b"></graphic>
</fig>
<table-wrap id="T1" position="float" orientation="portrait"><label>Table 1</label>
<caption><p>Lowest <italic>f<sub>3</sub>
</italic>
-statistics for each West Eurasian population</p>
</caption>
<table frame="box" rules="rows"><thead><tr><th valign="middle" align="left" rowspan="1" colspan="1"><italic>Ref<sub>1</sub>
</italic>
</th>
<th valign="middle" align="left" rowspan="1" colspan="1"><italic>Ref<sub>2</sub>
</italic>
</th>
<th valign="middle" align="left" rowspan="1" colspan="1">Target for which these two references give the lowest
<italic>f<sub>3</sub>
(X; Ref<sub>1</sub>
, Ref<sub>2</sub>
)</italic>
</th>
</tr>
</thead>
<tbody><tr><td align="left" valign="middle" rowspan="1" colspan="1"><bold>WHG</bold>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1"><bold>EEF</bold>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Sardinian***</td>
</tr>
<tr><td align="left" valign="middle" rowspan="1" colspan="1"><bold>WHG</bold>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1"><bold>Near East</bold>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Basque, Belarusian, Czech, English, Estonian, Finnish,
French_South, Icelandic, Lithuanian, Mordovian, Norwegian, Orcadian, Scottish,
Spanish, Spanish_North, Ukrainian</td>
</tr>
<tr><td align="left" valign="middle" rowspan="1" colspan="1"><bold>WHG</bold>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1"><bold>Siberian</bold>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Russian</td>
</tr>
<tr><td align="left" valign="middle" rowspan="1" colspan="1"><bold>EEF</bold>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1"><bold>ANE</bold>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Abkhasian***, Albanian,
Ashkenazi_Jew****, Bergamo, Bulgarian,
Chechen****, Croatian,
Cypriot****, Druze**, French, Greek,
Hungarian, Lezgin, Maltese, Sicilian, Turkish_Jew, Tuscan</td>
</tr>
<tr><td align="left" valign="middle" rowspan="1" colspan="1"><bold>EEF</bold>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1"><bold>Native American</bold>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Adygei, Balkar, Iranian, Kumyk, North_Ossetian,
Turkish</td>
</tr>
<tr><td align="left" valign="middle" rowspan="1" colspan="1"><bold>EEF</bold>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1"><bold>African</bold>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">BedouinA, BedouinB<sup>†</sup>
, Jordanian,
Lebanese, Libyan_Jew, Moroccan_Jew, Palestinian,
Saudi****, Syrian,
Tunisian_Jew***, Yemenite_Jew***</td>
</tr>
<tr><td align="left" valign="middle" rowspan="1" colspan="1"><bold>EEF</bold>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1"><bold>South Asian</bold>
</td>
<td align="left" valign="middle" rowspan="1" colspan="1">Armenian, Georgian****,
Georgian_Jew*, Iranian_Jew***,
Iraqi_Jew***</td>
</tr>
</tbody>
</table>
<table-wrap-foot><fn id="TFN2"><p>Note: WHG = Loschbour or LaBraña; EEF=Stuttgart;
ANE=MA1; Native American=Piapoco; African=Esan, Gambian, or
Kgalagadi; South Asian=GujaratiC or Vishwabrahmin. Statistics are negative with
Z<-4 unless otherwise noted: <sup>†</sup>
(positive) or *,
**, ***, ****,
to indicate Z less than 0, −1, −2, and −3 respectively. The
complete list of statistics can be found in <xref rid="T2" ref-type="table">Extended
Data Table 1</xref>
.</p>
</fn>
</table-wrap-foot>
</table-wrap>
</floats-group>
</pmc>
</record>
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