Massive migration from the steppe was a source for Indo-European languages in Europe

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David Reich and colleagues generated genome-wide data from 69 Europeans who lived between 8,000 and 3,000 years ago. Their analyses reveal that closely related groups of early farmers — different from indigenous hunter-gatherers — appeared in Germany, Hungary and Spain at around 8,000 to 7,000 years ago. At the same time Russia was inhabited by a distinctive population of hunter-gatherers with high affinity to a 24,000-year-old Siberian. By 6,000 to 5,000 years ago, a resurgence of hunter-gatherer ancestry had occurred throughout much of Europe, apart from in Russia. Western and Eastern Europe came into contact about 4,500 years ago, leaving traces of steppe ancestry in present-day Europeans. In addition to providing new insights into Neolithic population dynamics, these analyses lend support to the theory of a steppe origin of at least some of the Indo-European languages of Europe. The reported findings are also consistent with a study of 101 Bronze Age genomes reported on \n page 167\n \n of this issue. We generated genome-wide data from 69 Europeans who lived between 8,000–3,000 years ago by enriching ancient DNA libraries for a target set of almost 400,000 polymorphisms. Enrichment of these positions decreases the sequencing required for genome-wide ancient DNA analysis by a median of around 250-fold, allowing us to study an order of magnitude more individuals than previous studies1,2,3,4,5,6,7,8 and to obtain new insights about the past. We show that the populations of Western and Far Eastern Europe followed opposite trajectories between 8,000–5,000 years ago. At the beginning of the Neolithic period in Europe, ∼8,000–7,000 years ago, closely related groups of early farmers appeared in Germany, Hungary and Spain, different from indigenous hunter-gatherers, whereas Russia was inhabited by a distinctive population of hunter-gatherers with high affinity to a ∼24,000-year-old Siberian6. By ∼6,000–5,000 years ago, farmers throughout much of Europe had more hunter-gatherer ancestry than their predecessors, but in Russia, the Yamnaya steppe herders of this time were descended not only from the preceding eastern European hunter-gatherers, but also from a population of Near Eastern ancestry. Western and Eastern Europe came into contact ∼4,500 years ago, as the Late Neolithic Corded Ware people from Germany traced ∼75% of their ancestry to the Yamnaya, documenting a massive migration into the heartland of Europe from its eastern periphery. This steppe ancestry persisted in all sampled central Europeans until at least ∼3,000 years ago, and is ubiquitous in present-day Europeans. These results provide support for a steppe origin9 of at least some of the Indo-European languages of Europe.","datePublished":"2015-03-02T00:00:00Z","dateModified":"2015-03-02T00:00:00Z","pageStart":"207","pageEnd":"211","sameAs":"https://doi.org/10.1038/nature14317","keywords":["Anthropology","Archaeology","Genome","Population genetics","Science","Humanities and Social 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Boston, USA","@type":"PostalAddress"},"@type":"Organization"},{"name":"Broad Institute of Harvard and MIT","address":{"name":"Broad Institute of Harvard and MIT, Cambridge, USA","@type":"PostalAddress"},"@type":"Organization"},{"name":"Howard Hughes Medical Institute, Harvard Medical School","address":{"name":"Howard Hughes Medical Institute, Harvard Medical School, Boston, USA","@type":"PostalAddress"},"@type":"Organization"}],"email":"reich@genetics.med.harvard.edu","@type":"Person"}],"isAccessibleForFree":false,"hasPart":{"isAccessibleForFree":false,"cssSelector":".main-content","@type":"WebPageElement"},"@type":"ScholarlyArticle"},"@context":"https://schema.org","@type":"WebPage"}</script> <link rel="canonical" href="https://www.nature.com/articles/nature14317"> <meta name="journal_id" content="41586"/> <meta name="dc.title" content="Massive migration from the steppe was a source for Indo-European languages in Europe"/> <meta name="dc.source" content="Nature 2015 522:7555"/> <meta name="dc.format" content="text/html"/> <meta name="dc.publisher" content="Nature Publishing Group"/> <meta name="dc.date" content="2015-03-02"/> <meta name="dc.type" content="OriginalPaper"/> <meta name="dc.language" content="En"/> <meta name="dc.copyright" content="2015 Springer Nature Limited"/> <meta name="dc.rights" content="2015 Springer Nature Limited"/> <meta name="dc.rightsAgent" content="journalpermissions@springernature.com"/> <meta name="dc.description" content="A genome-wide analysis of 69 ancient Europeans reveals the history of population migrations around the time that Indo-European languages arose in Europe, when there was a large migration into Europe from the Eurasian steppe in the east (providing a genetic ancestry still present in Europeans today); these findings support a ‘steppe origin’ hypothesis for how some Indo-European languages arose. David Reich and colleagues generated genome-wide data from 69 Europeans who lived between 8,000 and 3,000 years ago. Their analyses reveal that closely related groups of early farmers — different from indigenous hunter-gatherers — appeared in Germany, Hungary and Spain at around 8,000 to 7,000 years ago. At the same time Russia was inhabited by a distinctive population of hunter-gatherers with high affinity to a 24,000-year-old Siberian. By 6,000 to 5,000 years ago, a resurgence of hunter-gatherer ancestry had occurred throughout much of Europe, apart from in Russia. Western and Eastern Europe came into contact about 4,500 years ago, leaving traces of steppe ancestry in present-day Europeans. In addition to providing new insights into Neolithic population dynamics, these analyses lend support to the theory of a steppe origin of at least some of the Indo-European languages of Europe. The reported findings are also consistent with a study of 101 Bronze Age genomes reported on page 167 of this issue. We generated genome-wide data from 69 Europeans who lived between 8,000–3,000 years ago by enriching ancient DNA libraries for a target set of almost 400,000 polymorphisms. Enrichment of these positions decreases the sequencing required for genome-wide ancient DNA analysis by a median of around 250-fold, allowing us to study an order of magnitude more individuals than previous studies1,2,3,4,5,6,7,8 and to obtain new insights about the past. We show that the populations of Western and Far Eastern Europe followed opposite trajectories between 8,000–5,000 years ago. At the beginning of the Neolithic period in Europe, ∼8,000–7,000 years ago, closely related groups of early farmers appeared in Germany, Hungary and Spain, different from indigenous hunter-gatherers, whereas Russia was inhabited by a distinctive population of hunter-gatherers with high affinity to a ∼24,000-year-old Siberian6. By ∼6,000–5,000 years ago, farmers throughout much of Europe had more hunter-gatherer ancestry than their predecessors, but in Russia, the Yamnaya steppe herders of this time were descended not only from the preceding eastern European hunter-gatherers, but also from a population of Near Eastern ancestry. Western and Eastern Europe came into contact ∼4,500 years ago, as the Late Neolithic Corded Ware people from Germany traced ∼75% of their ancestry to the Yamnaya, documenting a massive migration into the heartland of Europe from its eastern periphery. This steppe ancestry persisted in all sampled central Europeans until at least ∼3,000 years ago, and is ubiquitous in present-day Europeans. These results provide support for a steppe origin9 of at least some of the Indo-European languages of Europe."/> <meta name="prism.issn" content="1476-4687"/> <meta name="prism.publicationName" content="Nature"/> <meta name="prism.publicationDate" content="2015-03-02"/> <meta name="prism.volume" content="522"/> <meta name="prism.number" content="7555"/> <meta name="prism.section" content="OriginalPaper"/> <meta name="prism.startingPage" content="207"/> <meta name="prism.endingPage" content="211"/> <meta name="prism.copyright" content="2015 Springer Nature Limited"/> <meta name="prism.rightsAgent" content="journalpermissions@springernature.com"/> <meta name="prism.url" content="https://www.nature.com/articles/nature14317"/> <meta name="prism.doi" content="doi:10.1038/nature14317"/> <meta name="citation_pdf_url" content="https://www.nature.com/articles/nature14317.pdf"/> <meta name="citation_fulltext_html_url" content="https://www.nature.com/articles/nature14317"/> <meta name="citation_journal_title" content="Nature"/> <meta name="citation_journal_abbrev" content="Nature"/> <meta name="citation_publisher" content="Nature Publishing Group"/> <meta name="citation_issn" content="1476-4687"/> <meta name="citation_title" content="Massive migration from the steppe was a source for Indo-European languages in Europe"/> <meta name="citation_volume" content="522"/> <meta name="citation_issue" content="7555"/> <meta name="citation_publication_date" content="2015/06"/> <meta name="citation_online_date" content="2015/03/02"/> <meta name="citation_firstpage" content="207"/> <meta name="citation_lastpage" content="211"/> <meta name="citation_article_type" content="Article"/> <meta name="citation_language" content="en"/> <meta name="dc.identifier" content="doi:10.1038/nature14317"/> <meta name="DOI" content="10.1038/nature14317"/> <meta name="size" content="165770"/> <meta name="citation_doi" content="10.1038/nature14317"/> <meta name="citation_springer_api_url" content="http://api.springer.com/xmldata/jats?q=doi:10.1038/nature14317&api_key="/> <meta name="description" content="A genome-wide analysis of 69 ancient Europeans reveals the history of population migrations around the time that Indo-European languages arose in Europe, when there was a large migration into Europe from the Eurasian steppe in the east (providing a genetic ancestry still present in Europeans today); these findings support a ‘steppe origin’ hypothesis for how some Indo-European languages arose. David Reich and colleagues generated genome-wide data from 69 Europeans who lived between 8,000 and 3,000 years ago. Their analyses reveal that closely related groups of early farmers — different from indigenous hunter-gatherers — appeared in Germany, Hungary and Spain at around 8,000 to 7,000 years ago. At the same time Russia was inhabited by a distinctive population of hunter-gatherers with high affinity to a 24,000-year-old Siberian. By 6,000 to 5,000 years ago, a resurgence of hunter-gatherer ancestry had occurred throughout much of Europe, apart from in Russia. Western and Eastern Europe came into contact about 4,500 years ago, leaving traces of steppe ancestry in present-day Europeans. In addition to providing new insights into Neolithic population dynamics, these analyses lend support to the theory of a steppe origin of at least some of the Indo-European languages of Europe. The reported findings are also consistent with a study of 101 Bronze Age genomes reported on page 167 of this issue. We generated genome-wide data from 69 Europeans who lived between 8,000–3,000 years ago by enriching ancient DNA libraries for a target set of almost 400,000 polymorphisms. Enrichment of these positions decreases the sequencing required for genome-wide ancient DNA analysis by a median of around 250-fold, allowing us to study an order of magnitude more individuals than previous studies1,2,3,4,5,6,7,8 and to obtain new insights about the past. We show that the populations of Western and Far Eastern Europe followed opposite trajectories between 8,000–5,000 years ago. At the beginning of the Neolithic period in Europe, ∼8,000–7,000 years ago, closely related groups of early farmers appeared in Germany, Hungary and Spain, different from indigenous hunter-gatherers, whereas Russia was inhabited by a distinctive population of hunter-gatherers with high affinity to a ∼24,000-year-old Siberian6. By ∼6,000–5,000 years ago, farmers throughout much of Europe had more hunter-gatherer ancestry than their predecessors, but in Russia, the Yamnaya steppe herders of this time were descended not only from the preceding eastern European hunter-gatherers, but also from a population of Near Eastern ancestry. Western and Eastern Europe came into contact ∼4,500 years ago, as the Late Neolithic Corded Ware people from Germany traced ∼75% of their ancestry to the Yamnaya, documenting a massive migration into the heartland of Europe from its eastern periphery. This steppe ancestry persisted in all sampled central Europeans until at least ∼3,000 years ago, and is ubiquitous in present-day Europeans. These results provide support for a steppe origin9 of at least some of the Indo-European languages of Europe."/> <meta name="dc.creator" content="Haak, Wolfgang"/> <meta name="dc.creator" content="Lazaridis, Iosif"/> <meta name="dc.creator" content="Patterson, Nick"/> <meta name="dc.creator" content="Rohland, Nadin"/> <meta name="dc.creator" content="Mallick, Swapan"/> <meta name="dc.creator" content="Llamas, Bastien"/> <meta name="dc.creator" content="Brandt, Guido"/> <meta name="dc.creator" content="Nordenfelt, Susanne"/> <meta name="dc.creator" content="Harney, Eadaoin"/> <meta name="dc.creator" content="Stewardson, Kristin"/> <meta name="dc.creator" content="Fu, Qiaomei"/> <meta name="dc.creator" content="Mittnik, Alissa"/> <meta name="dc.creator" content="Bánffy, Eszter"/> <meta name="dc.creator" content="Economou, Christos"/> <meta name="dc.creator" content="Francken, Michael"/> <meta name="dc.creator" content="Friederich, Susanne"/> <meta name="dc.creator" content="Pena, Rafael Garrido"/> <meta name="dc.creator" content="Hallgren, Fredrik"/> <meta name="dc.creator" content="Khartanovich, Valery"/> <meta name="dc.creator" content="Khokhlov, Aleksandr"/> <meta name="dc.creator" content="Kunst, Michael"/> <meta name="dc.creator" content="Kuznetsov, Pavel"/> <meta name="dc.creator" content="Meller, Harald"/> <meta name="dc.creator" content="Mochalov, Oleg"/> <meta name="dc.creator" content="Moiseyev, Vayacheslav"/> <meta name="dc.creator" content="Nicklisch, Nicole"/> <meta name="dc.creator" content="Pichler, Sandra L."/> <meta name="dc.creator" content="Risch, Roberto"/> <meta name="dc.creator" content="Rojo Guerra, Manuel A."/> <meta name="dc.creator" content="Roth, Christina"/> <meta name="dc.creator" content="Szécsényi-Nagy, Anna"/> <meta name="dc.creator" content="Wahl, Joachim"/> <meta name="dc.creator" content="Meyer, Matthias"/> <meta name="dc.creator" content="Krause, Johannes"/> <meta name="dc.creator" content="Brown, Dorcas"/> <meta name="dc.creator" content="Anthony, David"/> <meta name="dc.creator" content="Cooper, Alan"/> <meta name="dc.creator" content="Alt, Kurt Werner"/> <meta name="dc.creator" content="Reich, David"/> <meta name="dc.subject" content="Anthropology"/> <meta name="dc.subject" content="Archaeology"/> <meta name="dc.subject" content="Genome"/> <meta name="dc.subject" content="Population genetics"/> <meta name="citation_reference" content="citation_journal_title=Nature; citation_title=Genome sequence of a 45,000-year-old modern human from western Siberia; citation_author=Q Fu; citation_volume=514; citation_publication_date=2014; citation_pages=445-449; citation_id=CR1"/> <meta name="citation_reference" content="citation_journal_title=Nature Commun.; citation_title=Genome flux and stasis in a five millennium transect of European prehistory; citation_author=C Gamba; citation_volume=5; citation_publication_date=2014; citation_pages=5257; citation_id=CR2"/> <meta name="citation_reference" content="citation_journal_title=Nature Commun.; citation_title=New insights into the Tyrolean Iceman’s origin and phenotype as inferred by whole-genome sequencing; citation_author=A Keller; citation_volume=3; citation_publication_date=2012; citation_pages=698; citation_id=CR3"/> <meta name="citation_reference" content="citation_journal_title=Nature; citation_title=Ancient human genomes suggest three ancestral populations for present-day Europeans; citation_author=I Lazaridis; citation_volume=513; citation_publication_date=2014; citation_pages=409-413; citation_id=CR4"/> <meta name="citation_reference" content="citation_journal_title=Nature; citation_title=Derived immune and ancestral pigmentation alleles in a 7,000-year-old Mesolithic European; citation_author=I Olalde; citation_volume=507; citation_publication_date=2014; citation_pages=225-228; citation_id=CR5"/> <meta name="citation_reference" content="citation_journal_title=Nature; citation_title=Upper Palaeolithic Siberian genome reveals dual ancestry of Native Americans; citation_author=M Raghavan; citation_volume=505; citation_publication_date=2014; citation_pages=87-91; citation_id=CR6"/> <meta name="citation_reference" content="citation_journal_title=Science; citation_title=Genomic structure in Europeans dating back at least 36,200 years; citation_author=A Seguin-Orlando; citation_volume=346; citation_publication_date=2014; citation_pages=1113-1118; citation_id=CR7"/> <meta name="citation_reference" content="citation_journal_title=Science; citation_title=Genomic diversity and admixture differs for Stone-Age Scandinavian foragers and farmers; citation_author=P Skoglund; citation_volume=344; citation_publication_date=2014; citation_pages=747-750; citation_id=CR8"/> <meta name="citation_reference" content="citation_title=The Horse, the Wheel, and Language: How Bronze-Age Riders from the Eurasian Steppes Shaped the Modern World; citation_publication_date=2007; citation_id=CR9; citation_author=DW Anthony"/> <meta name="citation_reference" content="citation_journal_title=Proc. Natl Acad. Sci. USA; citation_title=DNA analysis of an early modern human from Tianyuan Cave, China; citation_author=Q Fu; citation_volume=110; citation_publication_date=2013; citation_pages=2223-2227; citation_id=CR10"/> <meta name="citation_reference" content="citation_journal_title=Phil. Trans. R. Soc. Lond. B; citation_title=Partial uracil–DNA–glycosylase treatment for screening of ancient DNA; citation_author=N Rohland, E Harney, S Mallick, S Nordenfelt, D Reich; citation_volume=370; citation_publication_date=2015; citation_pages=20130624; citation_id=CR11"/> <meta name="citation_reference" content="citation_journal_title=Genetics; citation_title=Ancient admixture in human history; citation_author=N Patterson; citation_volume=192; citation_publication_date=2012; citation_pages=1065-1093; citation_id=CR12"/> <meta name="citation_reference" content="citation_journal_title=Curr. Biol.; citation_title=A revised timescale for human evolution based on ancient mitochondrial genomes; citation_author=Q Fu; citation_volume=23; citation_publication_date=2013; citation_pages=553-559; citation_id=CR13"/> <meta name="citation_reference" content="citation_journal_title=Science; citation_title=Ancient DNA reveals key stages in the formation of central European mitochondrial genetic diversity; citation_author=G Brandt; citation_volume=342; citation_publication_date=2013; citation_pages=257-261; citation_id=CR14"/> <meta name="citation_reference" content="citation_journal_title=PLoS Genet.; citation_title=Ancient DNA reveals prehistoric gene-flow from Siberia in the complex human population history of North East Europe; citation_author=C Der Sarkissian; citation_volume=9; citation_publication_date=2013; citation_pages=e1003296; citation_id=CR15"/> <meta name="citation_reference" content="citation_journal_title=Nucleic Acids Res.; citation_title=Removal of deaminated cytosines and detection of in vivo methylation in ancient DNA; citation_author=AW Briggs; citation_volume=38; citation_publication_date=2010; citation_pages=e87; citation_doi=10.1093/nar/gkp1163; citation_id=CR16"/> <meta name="citation_reference" content="citation_journal_title=Proc. Natl Acad. Sci. USA; citation_title=Patterns of damage in genomic DNA sequences from a Neandertal; citation_author=AW Briggs; citation_volume=104; citation_publication_date=2007; citation_pages=14616-14621; citation_id=CR17"/> <meta name="citation_reference" content="citation_journal_title=Eur. J. Hum. Genet.; citation_title=A major Y-chromosome haplogroup R1b Holocene era founder effect in Central and Western Europe; citation_author=NM Myres; citation_volume=19; citation_publication_date=2011; citation_pages=95-101; citation_id=CR18"/> <meta name="citation_reference" content="citation_journal_title=Eur. J. Hum. Genet.; citation_title=The phylogenetic and geographic structure of Y-chromosome haplogroup R1a; citation_author=PA Underhill; citation_volume=23; citation_publication_date=2015; citation_pages=124-131; citation_id=CR19"/> <meta name="citation_reference" content="citation_journal_title=Science; citation_title=Origins and genetic legacy of Neolithic farmers and hunter-gatherers in Europe; citation_author=P Skoglund; citation_volume=336; citation_publication_date=2012; citation_pages=466-469; citation_id=CR20"/> <meta name="citation_reference" content="citation_title=Ancient Europe, 8000 B.C. to A.D. 1000: Encyclopedia of the Barbarian World; citation_publication_date=2003; citation_id=CR21; citation_author=J Czebreszuk; citation_author=PI Bogucki; citation_author=PJ Crabtree"/> <meta name="citation_reference" content="citation_journal_title=Mol. Biol. Evol.; citation_title=Efficient moment-based inference of admixture parameters and sources of gene flow; citation_author=M Lipson; citation_volume=30; citation_publication_date=2013; citation_pages=1788-1802; citation_id=CR22"/> <meta name="citation_reference" content="Szécsényi-Nagy, A. et al. Tracing the genetic origin of Europe’s first farmers reveals insights into their social organization. Preprint at bioRxiv http://dx.doi.org/10.1101/008664 (2014)"/> <meta name="citation_reference" content="citation_journal_title=PLoS Biol.; citation_title=Ancient DNA from European early Neolithic farmers reveals their Near Eastern affinities; citation_author=W Haak; citation_volume=8; citation_publication_date=2010; citation_pages=e1000536; citation_id=CR24"/> <meta name="citation_reference" content="citation_journal_title=Science; citation_title=A genetic atlas of human admixture history; citation_author=G Hellenthal; citation_volume=343; citation_publication_date=2014; citation_pages=747-751; citation_id=CR25"/> <meta name="citation_reference" content="citation_journal_title=PLoS Biol.; citation_title=The geography of recent genetic ancestry across Europe; citation_author=P Ralph, G Coop; citation_volume=11; citation_publication_date=2013; citation_pages=e1001555; citation_id=CR26"/> <meta name="citation_reference" content="citation_title=Archaeology and Language: The Puzzle of Indo-European Origins; citation_publication_date=1987; citation_id=CR27; citation_author=C Renfrew"/> <meta name="citation_reference" content="citation_title=First Farmers: The Origins of Agricultural Societies; citation_publication_date=2004; citation_id=CR28; citation_author=P Bellwood"/> <meta name="citation_reference" content="citation_journal_title=Sci. Am.; citation_title=The early history of Indo-European languages; citation_author=TV Gamkrelidze, VV Ivanov; citation_volume=262; citation_publication_date=1990; citation_pages=110-116; citation_id=CR29"/> <meta name="citation_reference" content="citation_title=In Search of the Indo-Europeans: Language, Archaeology and Myth; citation_publication_date=1991; citation_id=CR30; citation_author=JP Mallory"/> <meta name="citation_reference" content="citation_journal_title=Nucleic Acids Res.; citation_title=Double indexing overcomes inaccuracies in multiplex sequencing on the Illumina platform; citation_author=M Kircher, S Sawyer, M Meyer; citation_volume=40; citation_publication_date=2012; citation_pages=e3; citation_id=CR31"/> <meta name="citation_reference" content="citation_journal_title=Nature; citation_title=A mitochondrial genome sequence of a hominin from Sima de los Huesos; citation_author=M Meyer; citation_volume=505; citation_publication_date=2014; citation_pages=403-406; citation_id=CR32"/> <meta name="citation_reference" content="citation_journal_title=Phil. Trans. R. Soc. Lond. B; citation_title=Partial uracil–DNA–glycosylase treatment for screening of ancient DNA; citation_author=N Rohland, E Harney, S Mallick, S Nordenfelt, D Reich; citation_volume=370; citation_publication_date=2015; citation_pages=20130624; citation_id=CR33"/> <meta name="citation_reference" content="citation_journal_title=Genome Res.; citation_title=Cost-effective, high-throughput DNA sequencing libraries for multiplexed target capture; citation_author=N Rohland, D Reich; citation_volume=22; citation_publication_date=2012; citation_pages=939-946; citation_id=CR34"/> <meta name="citation_reference" content="citation_journal_title=Bioinformatics; citation_title=Fast and accurate short read alignment with Burrows–Wheeler transform; citation_author=H Li, R Durbin; citation_volume=25; citation_publication_date=2009; citation_pages=1754-1760; citation_id=CR35"/> <meta name="citation_reference" content="citation_journal_title=Am. J. Hum. Genet.; citation_title=A “Copernican” reassessment of the human mitochondrial DNA tree from its root; citation_author=DM Behar; citation_volume=90; citation_publication_date=2012; citation_pages=675-684; citation_id=CR36"/> <meta name="citation_reference" content="citation_journal_title=BMC Bioinformatics; citation_title=Kalign—an accurate and fast multiple sequence alignment algorithm; citation_author=T Lassmann, ELL Sonnhammer; citation_volume=6; citation_publication_date=2005; citation_pages=298; citation_id=CR37"/> <meta name="citation_reference" content="citation_journal_title=PLoS ONE; citation_title=Temporal patterns of nucleotide misincorporations and DNA fragmentation in ancient DNA; citation_author=S Sawyer, J Krause, K Guschanski, V Savolainen, S Pääbo; citation_volume=7; citation_publication_date=2012; citation_pages=e34131; citation_id=CR38"/> <meta name="citation_reference" content="citation_journal_title=Science; citation_title=A draft sequence of the Neandertal genome; citation_author=RE Green; citation_volume=328; citation_publication_date=2010; citation_pages=710-722; citation_id=CR39"/> <meta name="citation_reference" content="citation_journal_title=BMC Bioinformatics; citation_title=Enhancements to the ADMIXTURE algorithm for individual ancestry estimation; citation_author=DH Alexander, K Lange; citation_volume=12; citation_publication_date=2011; citation_pages=246; citation_id=CR40"/> <meta name="citation_reference" content="citation_journal_title=Genome Res.; citation_title=Fast model-based estimation of ancestry in unrelated individuals; citation_author=DH Alexander, J Novembre, K Lange; citation_volume=19; citation_publication_date=2009; citation_pages=1655-1664; citation_id=CR41"/> <meta name="citation_reference" content="citation_journal_title=Nature Genet.; citation_title=Principal component analysis of genetic data; citation_author=D Reich, AL Price, N Patterson; citation_volume=40; citation_publication_date=2008; citation_pages=491-492; citation_id=CR42"/> <meta name="citation_reference" content="citation_journal_title=Am. J. Hum. Genet.; citation_title=PLINK: a tool set for whole-genome association and population-based linkage analyses; citation_author=S Purcell; citation_volume=81; citation_publication_date=2007; citation_pages=559-575; citation_id=CR43"/> <meta name="citation_reference" content="citation_journal_title=J. Archaeol. Sci.; citation_title=Accurate sex identification of ancient human remains using DNA shotgun sequencing; citation_author=P Skoglund, J Storå, A Götherström, M Jakobsson; citation_volume=40; citation_publication_date=2013; citation_pages=4477-4482; citation_id=CR44"/> <meta name="citation_author" content="Haak, Wolfgang"/> <meta name="citation_author_institution" content="Australian Centre for Ancient DNA, School of Earth and Environmental Sciences & Environment Institute, University of Adelaide, Adelaide, Australia"/> <meta name="citation_author" content="Lazaridis, Iosif"/> <meta name="citation_author_institution" content="Department of Genetics, Harvard Medical School, Boston, USA"/> <meta name="citation_author_institution" content="Broad Institute of Harvard and MIT, Cambridge, USA"/> <meta name="citation_author" content="Patterson, Nick"/> <meta name="citation_author_institution" content="Broad Institute of Harvard and MIT, Cambridge, USA"/> <meta name="citation_author" 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"/> <meta name="citation_author" content="Krause, Johannes"/> <meta name="citation_author_institution" content="Institute for Archaeological Sciences, University of Tübingen, D-72070 Tübingen, Germany, "/> <meta name="citation_author_institution" content="Departments of Paleoanthropology and Archaeogenetics, Senckenberg Center for Human Evolution and Paleoenvironment, University of Tübingen, D-72070 Tübingen, Germany, "/> <meta name="citation_author_institution" content="Max Planck Institute for the Science of Human History, D-07745 Jena, Germany, "/> <meta name="citation_author" content="Brown, Dorcas"/> <meta name="citation_author_institution" content="Anthropology Department, Hartwick College, Oneonta, USA"/> <meta name="citation_author" content="Anthony, David"/> <meta name="citation_author_institution" content="Anthropology Department, Hartwick College, Oneonta, USA"/> <meta name="citation_author" content="Cooper, Alan"/> <meta name="citation_author_institution" content="Australian Centre for Ancient DNA, School of Earth and Environmental Sciences & Environment Institute, University of Adelaide, Adelaide, Australia"/> <meta name="citation_author" content="Alt, Kurt Werner"/> <meta name="citation_author_institution" content="Institute of Anthropology, Johannes Gutenberg University of Mainz, D-55128 Mainz, Germany, "/> <meta name="citation_author_institution" content="State Office for Heritage Management and Archaeology Saxony-Anhalt and State Museum of Prehistory, D-06114 Halle, Germany, "/> <meta name="citation_author_institution" content="Danube Private University, A-3500 Krems, Austria, "/> <meta name="citation_author_institution" content="Institute for Prehistory and Archaeological Science, University of Basel, CH-4003 Basel, Switzerland, "/> <meta name="citation_author" content="Reich, David"/> <meta name="citation_author_institution" content="Department of Genetics, Harvard Medical School, Boston, USA"/> <meta name="citation_author_institution" content="Broad Institute of Harvard and MIT, Cambridge, USA"/> <meta name="citation_author_institution" content="Howard Hughes Medical Institute, Harvard Medical School, Boston, USA"/> <meta name="access_endpoint" content="https://www.nature.com/platform/readcube-access"/> <meta name="twitter:site" content="@nature"/> <meta name="twitter:card" content="summary_large_image"/> <meta name="twitter:image:alt" content="Content cover image"/> <meta name="twitter:title" content="Massive migration from the steppe was a source for Indo-European languages in Europe"/> <meta name="twitter:description" content="Nature - David Reich and colleagues generated genome-wide data from 69 Europeans who lived between 8,000 and 3,000 years ago. Their analyses reveal that closely related groups of early farmers..."/> <meta name="twitter:image" content="https://media.springernature.com/full/springer-static/image/art%3A10.1038%2Fnature14317/MediaObjects/41586_2015_Article_BFnature14317_Fig1_HTML.jpg"/> <meta property="og:url" content="https://www.nature.com/articles/nature14317"/> <meta property="og:type" content="article"/> <meta property="og:site_name" content="Nature"/> <meta property="og:title" content="Massive migration from the steppe was a source for Indo-European languages in Europe - Nature"/> <meta property="og:description" content="A genome-wide analysis of 69 ancient Europeans reveals the history of population migrations around the time that Indo-European languages arose in Europe, when there was a large migration into Europe from the Eurasian steppe in the east (providing a genetic ancestry still present in Europeans today); these findings support a ‘steppe origin’ hypothesis for how some Indo-European languages arose."/> <meta property="og:image" content="https://media.springernature.com/m685/springer-static/image/art%3A10.1038%2Fnature14317/MediaObjects/41586_2015_Article_BFnature14317_Fig1_HTML.jpg"/> <script> window.eligibleForRa21 = 'true'; </script> </head> <body class="article-page"> <noscript><iframe src="https://www.googletagmanager.com/ns.html?id=GTM-MRVXSHQ" height="0" width="0" style="display:none;visibility:hidden"></iframe></noscript> <div class="position-relative cleared z-index-50 background-white" data-test="top-containers"> <a class="c-skip-link" href="#content">Skip to main content</a> <div class="c-grade-c-banner u-hide"> <div class="c-grade-c-banner__container"> <p>Thank you for visiting nature.com. 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itemscope itemtype="https://schema.org/ListItem"> <span itemprop="name">article</span><meta itemprop="position" content="3"></li> </ol> </div> </nav> </div> <div class="u-container u-mt-32 u-mb-32 u-clearfix" id="content" data-component="article-container" data-container-type="article"> <main class="c-article-main-column u-float-left js-main-column" data-track-component="article body"> <article lang="en"> <div class="c-article-header"> <header> <ul class="c-article-identifiers" data-test="article-identifier"> <li class="c-article-identifiers__item" data-test="article-category">Letter</li> <li class="c-article-identifiers__item">Published: <time datetime="2015-03-02">02 March 2015</time></li> </ul> <h1 class="c-article-title" data-test="article-title" data-article-title="">Massive migration from the steppe was a source for Indo-European languages in Europe</h1> <ul class="c-article-author-list c-article-author-list--long" data-test="authors-list" data-component-authors-activator="authors-list"><li class="c-article-author-list__item"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Wolfgang-Haak-Aff1" data-author-popup="auth-Wolfgang-Haak-Aff1" data-author-search="Haak, Wolfgang">Wolfgang Haak</a><sup class="u-js-hide"><a href="#Aff1">1</a></sup><sup class="u-js-hide"> <a href="#na1">na1</a></sup>, </li><li class="c-article-author-list__item"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Iosif-Lazaridis-Aff2-Aff3" data-author-popup="auth-Iosif-Lazaridis-Aff2-Aff3" data-author-search="Lazaridis, Iosif">Iosif Lazaridis</a><sup class="u-js-hide"><a href="#Aff2">2</a>,<a href="#Aff3">3</a></sup><sup class="u-js-hide"> <a href="#na1">na1</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Nick-Patterson-Aff3" data-author-popup="auth-Nick-Patterson-Aff3" data-author-search="Patterson, Nick">Nick Patterson</a><sup class="u-js-hide"><a href="#Aff3">3</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Nadin-Rohland-Aff2-Aff3" data-author-popup="auth-Nadin-Rohland-Aff2-Aff3" data-author-search="Rohland, Nadin">Nadin Rohland</a><sup class="u-js-hide"><a href="#Aff2">2</a>,<a href="#Aff3">3</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Swapan-Mallick-Aff2-Aff3-Aff4" data-author-popup="auth-Swapan-Mallick-Aff2-Aff3-Aff4" data-author-search="Mallick, Swapan">Swapan Mallick</a><sup class="u-js-hide"><a href="#Aff2">2</a>,<a href="#Aff3">3</a>,<a href="#Aff4">4</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Bastien-Llamas-Aff1" data-author-popup="auth-Bastien-Llamas-Aff1" data-author-search="Llamas, Bastien">Bastien Llamas</a><sup class="u-js-hide"><a href="#Aff1">1</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Guido-Brandt-Aff5" data-author-popup="auth-Guido-Brandt-Aff5" data-author-search="Brandt, Guido">Guido Brandt</a><sup class="u-js-hide"><a href="#Aff5">5</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Susanne-Nordenfelt-Aff2-Aff3" data-author-popup="auth-Susanne-Nordenfelt-Aff2-Aff3" data-author-search="Nordenfelt, Susanne">Susanne Nordenfelt</a><sup class="u-js-hide"><a href="#Aff2">2</a>,<a href="#Aff3">3</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Eadaoin-Harney-Aff2-Aff3-Aff4" data-author-popup="auth-Eadaoin-Harney-Aff2-Aff3-Aff4" data-author-search="Harney, Eadaoin">Eadaoin Harney</a><sup class="u-js-hide"><a href="#Aff2">2</a>,<a href="#Aff3">3</a>,<a href="#Aff4">4</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Kristin-Stewardson-Aff2-Aff3-Aff4" data-author-popup="auth-Kristin-Stewardson-Aff2-Aff3-Aff4" data-author-search="Stewardson, Kristin">Kristin Stewardson</a><sup class="u-js-hide"><a href="#Aff2">2</a>,<a href="#Aff3">3</a>,<a href="#Aff4">4</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Qiaomei-Fu-Aff2-Aff3-Aff6-Aff7" data-author-popup="auth-Qiaomei-Fu-Aff2-Aff3-Aff6-Aff7" data-author-search="Fu, Qiaomei">Qiaomei Fu</a><sup class="u-js-hide"><a href="#Aff2">2</a>,<a href="#Aff3">3</a>,<a href="#Aff6">6</a>,<a href="#Aff7">7</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Alissa-Mittnik-Aff8" data-author-popup="auth-Alissa-Mittnik-Aff8" data-author-search="Mittnik, Alissa">Alissa Mittnik</a><sup class="u-js-hide"><a href="#Aff8">8</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Eszter-B_nffy-Aff9-Aff10" data-author-popup="auth-Eszter-B_nffy-Aff9-Aff10" data-author-search="Bánffy, Eszter">Eszter Bánffy</a><sup class="u-js-hide"><a href="#Aff9">9</a>,<a href="#Aff10">10</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Christos-Economou-Aff11" data-author-popup="auth-Christos-Economou-Aff11" data-author-search="Economou, Christos">Christos Economou</a><sup class="u-js-hide"><a href="#Aff11">11</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Michael-Francken-Aff12" data-author-popup="auth-Michael-Francken-Aff12" data-author-search="Francken, Michael">Michael Francken</a><sup class="u-js-hide"><a href="#Aff12">12</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Susanne-Friederich-Aff13" data-author-popup="auth-Susanne-Friederich-Aff13" data-author-search="Friederich, Susanne">Susanne Friederich</a><sup class="u-js-hide"><a href="#Aff13">13</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Rafael_Garrido-Pena-Aff14" data-author-popup="auth-Rafael_Garrido-Pena-Aff14" data-author-search="Pena, Rafael Garrido">Rafael Garrido Pena</a><sup class="u-js-hide"><a href="#Aff14">14</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Fredrik-Hallgren-Aff15" data-author-popup="auth-Fredrik-Hallgren-Aff15" data-author-search="Hallgren, Fredrik">Fredrik Hallgren</a><sup class="u-js-hide"><a href="#Aff15">15</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Valery-Khartanovich-Aff16" data-author-popup="auth-Valery-Khartanovich-Aff16" data-author-search="Khartanovich, Valery">Valery Khartanovich</a><sup class="u-js-hide"><a href="#Aff16">16</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Aleksandr-Khokhlov-Aff17" data-author-popup="auth-Aleksandr-Khokhlov-Aff17" data-author-search="Khokhlov, Aleksandr">Aleksandr Khokhlov</a><sup class="u-js-hide"><a href="#Aff17">17</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Michael-Kunst-Aff18" data-author-popup="auth-Michael-Kunst-Aff18" data-author-search="Kunst, Michael">Michael Kunst</a><sup class="u-js-hide"><a href="#Aff18">18</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Pavel-Kuznetsov-Aff17" data-author-popup="auth-Pavel-Kuznetsov-Aff17" data-author-search="Kuznetsov, Pavel">Pavel Kuznetsov</a><sup class="u-js-hide"><a href="#Aff17">17</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Harald-Meller-Aff13" data-author-popup="auth-Harald-Meller-Aff13" data-author-search="Meller, Harald">Harald Meller</a><sup class="u-js-hide"><a href="#Aff13">13</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Oleg-Mochalov-Aff17" data-author-popup="auth-Oleg-Mochalov-Aff17" data-author-search="Mochalov, Oleg">Oleg Mochalov</a><sup class="u-js-hide"><a href="#Aff17">17</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Vayacheslav-Moiseyev-Aff16" data-author-popup="auth-Vayacheslav-Moiseyev-Aff16" data-author-search="Moiseyev, Vayacheslav">Vayacheslav Moiseyev</a><sup class="u-js-hide"><a href="#Aff16">16</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Nicole-Nicklisch-Aff5-Aff13-Aff19" data-author-popup="auth-Nicole-Nicklisch-Aff5-Aff13-Aff19" data-author-search="Nicklisch, Nicole">Nicole Nicklisch</a><sup class="u-js-hide"><a href="#Aff5">5</a>,<a href="#Aff13">13</a>,<a href="#Aff19">19</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Sandra_L_-Pichler-Aff20" data-author-popup="auth-Sandra_L_-Pichler-Aff20" data-author-search="Pichler, Sandra L.">Sandra L. Pichler</a><sup class="u-js-hide"><a href="#Aff20">20</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Roberto-Risch-Aff21" data-author-popup="auth-Roberto-Risch-Aff21" data-author-search="Risch, Roberto">Roberto Risch</a><sup class="u-js-hide"><a href="#Aff21">21</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Manuel_A_-Rojo_Guerra-Aff22" data-author-popup="auth-Manuel_A_-Rojo_Guerra-Aff22" data-author-search="Rojo Guerra, Manuel A.">Manuel A. Rojo Guerra</a><sup class="u-js-hide"><a href="#Aff22">22</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Christina-Roth-Aff5" data-author-popup="auth-Christina-Roth-Aff5" data-author-search="Roth, Christina">Christina Roth</a><sup class="u-js-hide"><a href="#Aff5">5</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Anna-Sz_cs_nyi_Nagy-Aff5-Aff9" data-author-popup="auth-Anna-Sz_cs_nyi_Nagy-Aff5-Aff9" data-author-search="Szécsényi-Nagy, Anna">Anna Szécsényi-Nagy</a><sup class="u-js-hide"><a href="#Aff5">5</a>,<a href="#Aff9">9</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Joachim-Wahl-Aff23" data-author-popup="auth-Joachim-Wahl-Aff23" data-author-search="Wahl, Joachim">Joachim Wahl</a><sup class="u-js-hide"><a href="#Aff23">23</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Matthias-Meyer-Aff6" data-author-popup="auth-Matthias-Meyer-Aff6" data-author-search="Meyer, Matthias">Matthias Meyer</a><sup class="u-js-hide"><a href="#Aff6">6</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Johannes-Krause-Aff8-Aff12-Aff24" data-author-popup="auth-Johannes-Krause-Aff8-Aff12-Aff24" data-author-search="Krause, Johannes">Johannes Krause</a><sup class="u-js-hide"><a href="#Aff8">8</a>,<a href="#Aff12">12</a>,<a href="#Aff24">24</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Dorcas-Brown-Aff25" data-author-popup="auth-Dorcas-Brown-Aff25" data-author-search="Brown, Dorcas">Dorcas Brown</a><sup class="u-js-hide"><a href="#Aff25">25</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-David-Anthony-Aff25" data-author-popup="auth-David-Anthony-Aff25" data-author-search="Anthony, David">David Anthony</a><sup class="u-js-hide"><a href="#Aff25">25</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Alan-Cooper-Aff1" data-author-popup="auth-Alan-Cooper-Aff1" data-author-search="Cooper, Alan">Alan Cooper</a><sup class="u-js-hide"><a href="#Aff1">1</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Kurt_Werner-Alt-Aff5-Aff13-Aff19-Aff20" data-author-popup="auth-Kurt_Werner-Alt-Aff5-Aff13-Aff19-Aff20" data-author-search="Alt, Kurt Werner">Kurt Werner Alt</a><sup class="u-js-hide"><a href="#Aff5">5</a>,<a href="#Aff13">13</a>,<a href="#Aff19">19</a>,<a href="#Aff20">20</a></sup> & </li><li class="c-article-author-list__show-more" aria-label="Show all 39 authors for this article" title="Show all 39 authors for this article">…</li><li class="c-article-author-list__item"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-David-Reich-Aff2-Aff3-Aff4" data-author-popup="auth-David-Reich-Aff2-Aff3-Aff4" data-author-search="Reich, David" data-corresp-id="c1">David Reich<svg width="16" height="16" focusable="false" role="img" aria-hidden="true" class="u-icon"><use xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="#icon-eds-i-mail-medium"></use></svg></a><sup class="u-js-hide"><a href="#Aff2">2</a>,<a href="#Aff3">3</a>,<a href="#Aff4">4</a></sup> </li></ul><button aria-expanded="false" class="c-article-author-list__button"><svg width="16" height="16" focusable="false" role="img" aria-hidden="true" class="u-icon"><use xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="#icon-eds-i-chevron-down-medium"></use></svg><span>Show authors</span></button> <p class="c-article-info-details" data-container-section="info"> <a data-test="journal-link" href="/" data-track="click" data-track-action="journal homepage" data-track-category="article body" data-track-label="link"><i data-test="journal-title">Nature</i></a> <b data-test="journal-volume"><span class="u-visually-hidden">volume</span> 522</b>, <span class="u-visually-hidden">pages </span>207–211 (<span data-test="article-publication-year">2015</span>)<a href="#citeas" class="c-article-info-details__cite-as u-hide-print" data-track="click" data-track-action="cite this article" data-track-label="link">Cite this article</a> </p> <div class="c-article-metrics-bar__wrapper u-clear-both"> <ul class="c-article-metrics-bar u-list-reset"> <li class=" c-article-metrics-bar__item" data-test="access-count"> <p class="c-article-metrics-bar__count">61k <span class="c-article-metrics-bar__label">Accesses</span></p> </li> <li class="c-article-metrics-bar__item" data-test="citation-count"> <p class="c-article-metrics-bar__count">1098 <span class="c-article-metrics-bar__label">Citations</span></p> </li> <li class="c-article-metrics-bar__item" data-test="altmetric-score"> <p class="c-article-metrics-bar__count">1166 <span class="c-article-metrics-bar__label">Altmetric</span></p> </li> <li class="c-article-metrics-bar__item"> <p class="c-article-metrics-bar__details"><a href="/articles/nature14317/metrics" data-track="click" data-track-action="view metrics" data-track-label="link" rel="nofollow">Metrics <span class="u-visually-hidden">details</span></a></p> </li> </ul> </div> </header> <div class="u-js-hide" data-component="article-subject-links"> <h3 class="c-article__sub-heading">Subjects</h3> <ul class="c-article-subject-list"> <li class="c-article-subject-list__subject"><a href="/subjects/anthropology" data-track="click" data-track-action="view subject" data-track-label="link">Anthropology</a></li><li class="c-article-subject-list__subject"><a href="/subjects/archaeology" data-track="click" data-track-action="view subject" data-track-label="link">Archaeology</a></li><li class="c-article-subject-list__subject"><a href="/subjects/genome" data-track="click" data-track-action="view subject" data-track-label="link">Genome</a></li><li class="c-article-subject-list__subject"><a href="/subjects/population-genetics" data-track="click" data-track-action="view subject" data-track-label="link">Population genetics</a></li> </ul> </div> </div> <div class="c-article-body"> <section aria-labelledby="Abs3" data-title="Abstract" lang="en"><div class="c-article-section" id="Abs3-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="Abs3">Abstract</h2><div class="c-article-section__content" id="Abs3-content"><p>We generated genome-wide data from 69 Europeans who lived between 8,000–3,000 years ago by enriching ancient DNA libraries for a target set of almost 400,000 polymorphisms. Enrichment of these positions decreases the sequencing required for genome-wide ancient DNA analysis by a median of around 250-fold, allowing us to study an order of magnitude more individuals than previous studies<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 1" title="Fu, Q. et al. Genome sequence of a 45,000-year-old modern human from western Siberia. Nature 514, 445–449 (2014)" href="/articles/nature14317#ref-CR1" id="ref-link-section-d123802842e1164">1</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2" title="Gamba, C. et al. Genome flux and stasis in a five millennium transect of European prehistory. Nature Commun. 5, 5257 (2014)" href="/articles/nature14317#ref-CR2" id="ref-link-section-d123802842e1167">2</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 3" title="Keller, A. et al. New insights into the Tyrolean Iceman’s origin and phenotype as inferred by whole-genome sequencing. Nature Commun. 3, 698 (2012)" href="/articles/nature14317#ref-CR3" id="ref-link-section-d123802842e1170">3</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 4" title="Lazaridis, I. et al. Ancient human genomes suggest three ancestral populations for present-day Europeans. Nature 513, 409–413 (2014)" href="/articles/nature14317#ref-CR4" id="ref-link-section-d123802842e1173">4</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 5" title="Olalde, I. et al. Derived immune and ancestral pigmentation alleles in a 7,000-year-old Mesolithic European. Nature 507, 225–228 (2014)" href="/articles/nature14317#ref-CR5" id="ref-link-section-d123802842e1176">5</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 6" title="Raghavan, M. et al. Upper Palaeolithic Siberian genome reveals dual ancestry of Native Americans. Nature 505, 87–91 (2014)" href="/articles/nature14317#ref-CR6" id="ref-link-section-d123802842e1179">6</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 7" title="Seguin-Orlando, A. et al. Genomic structure in Europeans dating back at least 36,200 years. Science 346, 1113–1118 (2014)" href="/articles/nature14317#ref-CR7" id="ref-link-section-d123802842e1183">7</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 8" title="Skoglund, P. et al. Genomic diversity and admixture differs for Stone-Age Scandinavian foragers and farmers. Science 344, 747–750 (2014)" href="/articles/nature14317#ref-CR8" id="ref-link-section-d123802842e1186">8</a></sup> and to obtain new insights about the past. We show that the populations of Western and Far Eastern Europe followed opposite trajectories between 8,000–5,000 years ago. At the beginning of the Neolithic period in Europe, <span class="stix">∼</span>8,000–7,000 years ago, closely related groups of early farmers appeared in Germany, Hungary and Spain, different from indigenous hunter-gatherers, whereas Russia was inhabited by a distinctive population of hunter-gatherers with high affinity to a <span class="stix">∼</span>24,000-year-old Siberian<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 6" title="Raghavan, M. et al. Upper Palaeolithic Siberian genome reveals dual ancestry of Native Americans. Nature 505, 87–91 (2014)" href="/articles/nature14317#ref-CR6" id="ref-link-section-d123802842e1190">6</a></sup>. By <span class="stix">∼</span>6,000–5,000 years ago, farmers throughout much of Europe had more hunter-gatherer ancestry than their predecessors, but in Russia, the Yamnaya steppe herders of this time were descended not only from the preceding eastern European hunter-gatherers, but also from a population of Near Eastern ancestry. Western and Eastern Europe came into contact <span class="stix">∼</span>4,500 years ago, as the Late Neolithic Corded Ware people from Germany traced <span class="stix">∼</span>75% of their ancestry to the Yamnaya, documenting a massive migration into the heartland of Europe from its eastern periphery. This steppe ancestry persisted in all sampled central Europeans until at least <span class="stix">∼</span>3,000 years ago, and is ubiquitous in present-day Europeans. These results provide support for a steppe origin<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 9" title="Anthony, D. W. The Horse, the Wheel, and Language: How Bronze-Age Riders from the Eurasian Steppes Shaped the Modern World (Princeton Univ. Press, 2007)" href="/articles/nature14317#ref-CR9" id="ref-link-section-d123802842e1194">9</a></sup> of at least some of the Indo-European languages of Europe.</p></div></div></section> <noscript> <div class="c-nature-box c-nature-box--side " data-component="entitlement-box"> <div class="js-access-button"> <a href="https://wayf.springernature.com?redirect_uri=https%3A%2F%2Fwww.nature.com%2Farticles%2Fnature14317" class="c-article__button" data-test="ra21"> <svg class="u-icon" width="18" height="18" aria-hidden="true" focusable="false"><use href="#icon-institution"></use></svg> <span class="c-article__button-text">Access through your institution</span> </a> </div> <div class="js-buy-button"> <a href="#access-options" class="c-article__button c-article__button--inverted" data-test="ra21"> <span>Buy or subscribe</span> </a> </div> </div> </noscript> <div class="js-context-bar-sticky-point-mobile" data-track-context="article body"> </div> <div class="c-article-access-provider u-mb-32 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class="c-article-section" id="accession-codes-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="accession-codes">Accession codes</h2><div class="c-article-section__content" id="accession-codes-content"> <h3 class="c-article__sub-heading">Primary accessions</h3> <h4 class="c-article__sub-heading c-article__sub-heading--small">European Nucleotide Archive</h4><ul class="c-article-data-availability-list u-list-reset"> <li class="c-article-data-availability-list__item"> <p> <a href="http://www.ebi.ac.uk/ena/data/view/PRJEB8448">PRJEB8448</a> </p> </li> </ul> <h3 class="c-article__sub-heading">Data deposits</h3> <p>The aligned sequences are available through the European Nucleotide Archive under accession number <a href="http://www.ebi.ac.uk/ena/data/view/PRJEB8448">PRJEB8448</a>. The Human Origins genotype dataset including ancient individuals can be found at (<a href="http://genetics.med.harvard.edu/reichlab/Reich_Lab/Datasets.html">http://genetics.med.harvard.edu/reichlab/Reich_Lab/Datasets.html</a>).</p> </div></div></section><div id="MagazineFulltextArticleBodySuffix"><section aria-labelledby="Bib1" data-title="References"><div class="c-article-section" id="Bib1-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="Bib1">References</h2><div class="c-article-section__content" id="Bib1-content"><div data-container-section="references"><ol class="c-article-references" data-track-component="outbound reference" data-track-context="references section"><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="1"><p class="c-article-references__text" id="ref-CR1">Fu, Q. et al. Genome sequence of a 45,000-year-old modern human from western Siberia. <i>Nature</i> <b>514</b>, 445–449 (2014)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="ads reference" data-track-action="ads reference" href="http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2014Natur.514..445F" aria-label="ADS reference 1">ADS</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:CAS:528:DC%2BC2cXhvVSmurvK" aria-label="CAS reference 1">CAS</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=25341783" aria-label="PubMed reference 1">PubMed</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed central reference" data-track-action="pubmed central reference" href="http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4753769" aria-label="PubMed Central reference 1">PubMed Central</a>  <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 1" href="http://scholar.google.com/scholar_lookup?&title=Genome%20sequence%20of%20a%2045%2C000-year-old%20modern%20human%20from%20western%20Siberia&journal=Nature&volume=514&pages=445-449&publication_year=2014&author=Fu%2CQ"> Google Scholar</a>  </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="2"><p class="c-article-references__text" id="ref-CR2">Gamba, C. et al. Genome flux and stasis in a five millennium transect of European prehistory. <i>Nature Commun.</i> <b>5</b>, 5257 (2014)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="ads reference" data-track-action="ads reference" href="http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2014NatCo...5.5257G" aria-label="ADS reference 2">ADS</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:CAS:528:DC%2BC2cXhvF2murjM" aria-label="CAS reference 2">CAS</a>  <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 2" href="http://scholar.google.com/scholar_lookup?&title=Genome%20flux%20and%20stasis%20in%20a%20five%20millennium%20transect%20of%20European%20prehistory&journal=Nature%20Commun.&volume=5&publication_year=2014&author=Gamba%2CC"> Google Scholar</a>  </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="3"><p class="c-article-references__text" id="ref-CR3">Keller, A. et al. New insights into the Tyrolean Iceman’s origin and phenotype as inferred by whole-genome sequencing. <i>Nature Commun.</i> <b>3</b>, 698 (2012)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="ads reference" data-track-action="ads reference" href="http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2012NatCo...3..698K" aria-label="ADS reference 3">ADS</a>  <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 3" href="http://scholar.google.com/scholar_lookup?&title=New%20insights%20into%20the%20Tyrolean%20Iceman%E2%80%99s%20origin%20and%20phenotype%20as%20inferred%20by%20whole-genome%20sequencing&journal=Nature%20Commun.&volume=3&publication_year=2012&author=Keller%2CA"> Google Scholar</a>  </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="4"><p class="c-article-references__text" id="ref-CR4">Lazaridis, I. et al. Ancient human genomes suggest three ancestral populations for present-day Europeans. <i>Nature</i> <b>513</b>, 409–413 (2014)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="ads reference" data-track-action="ads reference" href="http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2014Natur.513..409L" aria-label="ADS reference 4">ADS</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:CAS:528:DC%2BC2cXhsFOlu7nL" aria-label="CAS reference 4">CAS</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=25230663" aria-label="PubMed reference 4">PubMed</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed central reference" data-track-action="pubmed central reference" href="http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4170574" aria-label="PubMed Central reference 4">PubMed Central</a>  <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 4" href="http://scholar.google.com/scholar_lookup?&title=Ancient%20human%20genomes%20suggest%20three%20ancestral%20populations%20for%20present-day%20Europeans&journal=Nature&volume=513&pages=409-413&publication_year=2014&author=Lazaridis%2CI"> Google Scholar</a>  </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="5"><p class="c-article-references__text" id="ref-CR5">Olalde, I. et al. Derived immune and ancestral pigmentation alleles in a 7,000-year-old Mesolithic European. <i>Nature</i> <b>507</b>, 225–228 (2014)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="ads reference" data-track-action="ads reference" href="http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2014Natur.507..225O" aria-label="ADS reference 5">ADS</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:CAS:528:DC%2BC2cXktV2rsr8%3D" aria-label="CAS reference 5">CAS</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=24463515" aria-label="PubMed reference 5">PubMed</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed central reference" data-track-action="pubmed central reference" href="http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4269527" aria-label="PubMed Central reference 5">PubMed Central</a>  <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 5" href="http://scholar.google.com/scholar_lookup?&title=Derived%20immune%20and%20ancestral%20pigmentation%20alleles%20in%20a%207%2C000-year-old%20Mesolithic%20European&journal=Nature&volume=507&pages=225-228&publication_year=2014&author=Olalde%2CI"> Google Scholar</a>  </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="6"><p class="c-article-references__text" id="ref-CR6">Raghavan, M. et al. Upper Palaeolithic Siberian genome reveals dual ancestry of Native Americans. <i>Nature</i> <b>505</b>, 87–91 (2014)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="ads reference" data-track-action="ads reference" href="http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2014Natur.505...87R" aria-label="ADS reference 6">ADS</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=24256729" aria-label="PubMed reference 6">PubMed</a>  <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 6" href="http://scholar.google.com/scholar_lookup?&title=Upper%20Palaeolithic%20Siberian%20genome%20reveals%20dual%20ancestry%20of%20Native%20Americans&journal=Nature&volume=505&pages=87-91&publication_year=2014&author=Raghavan%2CM"> Google Scholar</a>  </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="7"><p class="c-article-references__text" id="ref-CR7">Seguin-Orlando, A. et al. Genomic structure in Europeans dating back at least 36,200 years. <i>Science</i> <b>346</b>, 1113–1118 (2014)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="ads reference" data-track-action="ads reference" href="http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2014Sci...346.1113S" aria-label="ADS reference 7">ADS</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:CAS:528:DC%2BC2cXhvFKqtLfK" aria-label="CAS reference 7">CAS</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=25378462" aria-label="PubMed reference 7">PubMed</a>  <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 7" href="http://scholar.google.com/scholar_lookup?&title=Genomic%20structure%20in%20Europeans%20dating%20back%20at%20least%2036%2C200%20years&journal=Science&volume=346&pages=1113-1118&publication_year=2014&author=Seguin-Orlando%2CA"> Google Scholar</a>  </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="8"><p class="c-article-references__text" id="ref-CR8">Skoglund, P. et al. Genomic diversity and admixture differs for Stone-Age Scandinavian foragers and farmers. <i>Science</i> <b>344</b>, 747–750 (2014)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="ads reference" data-track-action="ads reference" href="http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2014Sci...344..747S" aria-label="ADS reference 8">ADS</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:CAS:528:DC%2BC2cXotVahtLs%3D" aria-label="CAS reference 8">CAS</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=24762536" aria-label="PubMed reference 8">PubMed</a>  <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 8" href="http://scholar.google.com/scholar_lookup?&title=Genomic%20diversity%20and%20admixture%20differs%20for%20Stone-Age%20Scandinavian%20foragers%20and%20farmers&journal=Science&volume=344&pages=747-750&publication_year=2014&author=Skoglund%2CP"> Google Scholar</a>  </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="9"><p class="c-article-references__text" id="ref-CR9">Anthony, D. W. <i>The Horse, the Wheel, and Language: How Bronze-Age Riders from the Eurasian Steppes Shaped the Modern World</i> (Princeton Univ. Press, 2007)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 9" href="http://scholar.google.com/scholar_lookup?&title=The%20Horse%2C%20the%20Wheel%2C%20and%20Language%3A%20How%20Bronze-Age%20Riders%20from%20the%20Eurasian%20Steppes%20Shaped%20the%20Modern%20World&publication_year=2007&author=Anthony%2CDW"> Google Scholar</a>  </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="10"><p class="c-article-references__text" id="ref-CR10">Fu, Q. et al. DNA analysis of an early modern human from Tianyuan Cave, China. <i>Proc. Natl Acad. Sci. USA</i> <b>110</b>, 2223–2227 (2013)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="ads reference" data-track-action="ads reference" href="http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2013PNAS..110.2223F" aria-label="ADS reference 10">ADS</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:CAS:528:DC%2BC3sXjtVersLs%3D" aria-label="CAS reference 10">CAS</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=23341637" aria-label="PubMed reference 10">PubMed</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed central reference" data-track-action="pubmed central reference" href="http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3568306" aria-label="PubMed Central reference 10">PubMed Central</a>  <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 10" href="http://scholar.google.com/scholar_lookup?&title=DNA%20analysis%20of%20an%20early%20modern%20human%20from%20Tianyuan%20Cave%2C%20China&journal=Proc.%20Natl%20Acad.%20Sci.%20USA&volume=110&pages=2223-2227&publication_year=2013&author=Fu%2CQ"> Google Scholar</a>  </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="11"><p class="c-article-references__text" id="ref-CR11">Rohland, N., Harney, E., Mallick, S., Nordenfelt, S. & Reich, D. Partial uracil–DNA–glycosylase treatment for screening of ancient DNA. <i>Phil. Trans. R. Soc. Lond. B</i> <b>370</b>, 20130624 (2015)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 11" href="http://scholar.google.com/scholar_lookup?&title=Partial%20uracil%E2%80%93DNA%E2%80%93glycosylase%20treatment%20for%20screening%20of%20ancient%20DNA&journal=Phil.%20Trans.%20R.%20Soc.%20Lond.%20B&volume=370&publication_year=2015&author=Rohland%2CN&author=Harney%2CE&author=Mallick%2CS&author=Nordenfelt%2CS&author=Reich%2CD"> Google Scholar</a>  </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="12"><p class="c-article-references__text" id="ref-CR12">Patterson, N. et al. Ancient admixture in human history. <i>Genetics</i> <b>192</b>, 1065–1093 (2012)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=22960212" aria-label="PubMed reference 12">PubMed</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed central reference" data-track-action="pubmed central reference" href="http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3522152" aria-label="PubMed Central reference 12">PubMed Central</a>  <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 12" href="http://scholar.google.com/scholar_lookup?&title=Ancient%20admixture%20in%20human%20history&journal=Genetics&volume=192&pages=1065-1093&publication_year=2012&author=Patterson%2CN"> Google Scholar</a>  </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="13"><p class="c-article-references__text" id="ref-CR13">Fu, Q. et al. A revised timescale for human evolution based on ancient mitochondrial genomes. <i>Curr. Biol.</i> <b>23</b>, 553–559 (2013)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:CAS:528:DC%2BC3sXksFOlu7Y%3D" aria-label="CAS reference 13">CAS</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=23523248" aria-label="PubMed reference 13">PubMed</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed central reference" data-track-action="pubmed central reference" href="http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5036973" aria-label="PubMed Central reference 13">PubMed Central</a>  <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 13" href="http://scholar.google.com/scholar_lookup?&title=A%20revised%20timescale%20for%20human%20evolution%20based%20on%20ancient%20mitochondrial%20genomes&journal=Curr.%20Biol.&volume=23&pages=553-559&publication_year=2013&author=Fu%2CQ"> Google Scholar</a>  </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="14"><p class="c-article-references__text" id="ref-CR14">Brandt, G. et al. Ancient DNA reveals key stages in the formation of central European mitochondrial genetic diversity. <i>Science</i> <b>342</b>, 257–261 (2013)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="ads reference" data-track-action="ads reference" href="http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2013Sci...342..257B" aria-label="ADS reference 14">ADS</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:CAS:528:DC%2BC3sXhsFyns73I" aria-label="CAS reference 14">CAS</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=24115443" aria-label="PubMed reference 14">PubMed</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed central reference" data-track-action="pubmed central reference" href="http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4039305" aria-label="PubMed Central reference 14">PubMed Central</a>  <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 14" href="http://scholar.google.com/scholar_lookup?&title=Ancient%20DNA%20reveals%20key%20stages%20in%20the%20formation%20of%20central%20European%20mitochondrial%20genetic%20diversity&journal=Science&volume=342&pages=257-261&publication_year=2013&author=Brandt%2CG"> Google Scholar</a>  </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="15"><p class="c-article-references__text" id="ref-CR15">Der Sarkissian, C. et al. Ancient DNA reveals prehistoric gene-flow from Siberia in the complex human population history of North East Europe. <i>PLoS Genet.</i> <b>9</b>, e1003296 (2013)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:CAS:528:DC%2BC3sXkt1Wlt7w%3D" aria-label="CAS reference 15">CAS</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=23459685" aria-label="PubMed reference 15">PubMed</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed central reference" data-track-action="pubmed central reference" href="http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3573127" aria-label="PubMed Central reference 15">PubMed Central</a>  <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 15" href="http://scholar.google.com/scholar_lookup?&title=Ancient%20DNA%20reveals%20prehistoric%20gene-flow%20from%20Siberia%20in%20the%20complex%20human%20population%20history%20of%20North%20East%20Europe&journal=PLoS%20Genet.&volume=9&publication_year=2013&author=Der%20Sarkissian%2CC"> Google Scholar</a>  </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="16"><p class="c-article-references__text" id="ref-CR16">Briggs, A. W. et al. Removal of deaminated cytosines and detection of <i>in vivo</i> methylation in ancient DNA. <i>Nucleic Acids Res.</i> <b>38</b>, e87 (2010)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="10.1093/nar/gkp1163" data-track-item_id="10.1093/nar/gkp1163" data-track-value="article reference" data-track-action="article reference" href="https://doi.org/10.1093%2Fnar%2Fgkp1163" aria-label="Article reference 16" data-doi="10.1093/nar/gkp1163">Article</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=20028723" aria-label="PubMed reference 16">PubMed</a>  <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 16" href="http://scholar.google.com/scholar_lookup?&title=Removal%20of%20deaminated%20cytosines%20and%20detection%20of%20in%20vivo%20methylation%20in%20ancient%20DNA&journal=Nucleic%20Acids%20Res.&doi=10.1093%2Fnar%2Fgkp1163&volume=38&publication_year=2010&author=Briggs%2CAW"> Google Scholar</a>  </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="17"><p class="c-article-references__text" id="ref-CR17">Briggs, A. W. et al. Patterns of damage in genomic DNA sequences from a Neandertal. <i>Proc. Natl Acad. Sci. USA</i> <b>104</b>, 14616–14621 (2007)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="ads reference" data-track-action="ads reference" href="http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2007PNAS..10414616B" aria-label="ADS reference 17">ADS</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:CAS:528:DC%2BD2sXhtVOns7rF" aria-label="CAS reference 17">CAS</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=17715061" aria-label="PubMed reference 17">PubMed</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed central reference" data-track-action="pubmed central reference" href="http://www.ncbi.nlm.nih.gov/pmc/articles/PMC1976210" aria-label="PubMed Central reference 17">PubMed Central</a>  <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 17" href="http://scholar.google.com/scholar_lookup?&title=Patterns%20of%20damage%20in%20genomic%20DNA%20sequences%20from%20a%20Neandertal&journal=Proc.%20Natl%20Acad.%20Sci.%20USA&volume=104&pages=14616-14621&publication_year=2007&author=Briggs%2CAW"> Google Scholar</a>  </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="18"><p class="c-article-references__text" id="ref-CR18">Myres, N. M. et al. A major Y-chromosome haplogroup R1b Holocene era founder effect in Central and Western Europe. <i>Eur. J. Hum. Genet.</i> <b>19</b>, 95–101 (2011)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=20736979" aria-label="PubMed reference 18">PubMed</a>  <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 18" href="http://scholar.google.com/scholar_lookup?&title=A%20major%20Y-chromosome%20haplogroup%20R1b%20Holocene%20era%20founder%20effect%20in%20Central%20and%20Western%20Europe&journal=Eur.%20J.%20Hum.%20Genet.&volume=19&pages=95-101&publication_year=2011&author=Myres%2CNM"> Google Scholar</a>  </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="19"><p class="c-article-references__text" id="ref-CR19">Underhill, P. A. et al. The phylogenetic and geographic structure of Y-chromosome haplogroup R1a. <i>Eur. J. Hum. Genet.</i> <b>23</b>, 124–131 (2015)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=24667786" aria-label="PubMed reference 19">PubMed</a>  <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 19" href="http://scholar.google.com/scholar_lookup?&title=The%20phylogenetic%20and%20geographic%20structure%20of%20Y-chromosome%20haplogroup%20R1a&journal=Eur.%20J.%20Hum.%20Genet.&volume=23&pages=124-131&publication_year=2015&author=Underhill%2CPA"> Google Scholar</a>  </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="20"><p class="c-article-references__text" id="ref-CR20">Skoglund, P. et al. Origins and genetic legacy of Neolithic farmers and hunter-gatherers in Europe. <i>Science</i> <b>336</b>, 466–469 (2012)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="ads reference" data-track-action="ads reference" href="http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2012Sci...336..466S" aria-label="ADS reference 20">ADS</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:CAS:528:DC%2BC38XlvFGqsLc%3D" aria-label="CAS reference 20">CAS</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=22539720" aria-label="PubMed reference 20">PubMed</a>  <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 20" href="http://scholar.google.com/scholar_lookup?&title=Origins%20and%20genetic%20legacy%20of%20Neolithic%20farmers%20and%20hunter-gatherers%20in%20Europe&journal=Science&volume=336&pages=466-469&publication_year=2012&author=Skoglund%2CP"> Google Scholar</a>  </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="21"><p class="c-article-references__text" id="ref-CR21">Czebreszuk, J. in <i>Ancient Europe, 8000 B.C. to A.D. 1000: Encyclopedia of the Barbarian World</i> (eds Bogucki, P. I. & Crabtree, P. J. ) 467–475 (Charles Scribners & Sons, 2003)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 21" href="http://scholar.google.com/scholar_lookup?&title=Ancient%20Europe%2C%208000%20B.C.%20to%20A.D.%201000%3A%20Encyclopedia%20of%20the%20Barbarian%20World&pages=467-475&publication_year=2003&author=Czebreszuk%2CJ&author=Bogucki%2CPI&author=Crabtree%2CPJ"> Google Scholar</a>  </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="22"><p class="c-article-references__text" id="ref-CR22">Lipson, M. et al. Efficient moment-based inference of admixture parameters and sources of gene flow. <i>Mol. Biol. Evol.</i> <b>30</b>, 1788–1802 (2013)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:CAS:528:DC%2BC3sXhtFSrt7rI" aria-label="CAS reference 22">CAS</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=23709261" aria-label="PubMed reference 22">PubMed</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed central reference" data-track-action="pubmed central reference" href="http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3708505" aria-label="PubMed Central reference 22">PubMed Central</a>  <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 22" href="http://scholar.google.com/scholar_lookup?&title=Efficient%20moment-based%20inference%20of%20admixture%20parameters%20and%20sources%20of%20gene%20flow&journal=Mol.%20Biol.%20Evol.&volume=30&pages=1788-1802&publication_year=2013&author=Lipson%2CM"> Google Scholar</a>  </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="23"><p class="c-article-references__text" id="ref-CR23">Szécsényi-Nagy, A. et al. Tracing the genetic origin of Europe’s first farmers reveals insights into their social organization. Preprint at <i>bioRxiv</i><a href="https://doi.org/10.1101/008664" data-track="click_references" data-track-action="external reference" data-track-value="external reference" data-track-label="10.1101/008664">http://dx.doi.org/10.1101/008664</a> (2014)</p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="24"><p class="c-article-references__text" id="ref-CR24">Haak, W. et al. Ancient DNA from European early Neolithic farmers reveals their Near Eastern affinities. <i>PLoS Biol.</i> <b>8</b>, e1000536 (2010)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=21085689" aria-label="PubMed reference 24">PubMed</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed central reference" data-track-action="pubmed central reference" href="http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2976717" aria-label="PubMed Central reference 24">PubMed Central</a>  <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 24" href="http://scholar.google.com/scholar_lookup?&title=Ancient%20DNA%20from%20European%20early%20Neolithic%20farmers%20reveals%20their%20Near%20Eastern%20affinities&journal=PLoS%20Biol.&volume=8&publication_year=2010&author=Haak%2CW"> Google Scholar</a>  </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="25"><p class="c-article-references__text" id="ref-CR25">Hellenthal, G. et al. A genetic atlas of human admixture history. <i>Science</i> <b>343</b>, 747–751 (2014)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="ads reference" data-track-action="ads reference" href="http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2014Sci...343..747H" aria-label="ADS reference 25">ADS</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:CAS:528:DC%2BC2cXit12ks74%3D" aria-label="CAS reference 25">CAS</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=24531965" aria-label="PubMed reference 25">PubMed</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed central reference" data-track-action="pubmed central reference" href="http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4209567" aria-label="PubMed Central reference 25">PubMed Central</a>  <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 25" href="http://scholar.google.com/scholar_lookup?&title=A%20genetic%20atlas%20of%20human%20admixture%20history&journal=Science&volume=343&pages=747-751&publication_year=2014&author=Hellenthal%2CG"> Google Scholar</a>  </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="26"><p class="c-article-references__text" id="ref-CR26">Ralph, P. & Coop, G. The geography of recent genetic ancestry across Europe. <i>PLoS Biol.</i> <b>11</b>, e1001555 (2013)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:CAS:528:DC%2BC3sXptlyhu7g%3D" aria-label="CAS reference 26">CAS</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=23667324" aria-label="PubMed reference 26">PubMed</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed central reference" data-track-action="pubmed central reference" href="http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3646727" aria-label="PubMed Central reference 26">PubMed Central</a>  <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 26" href="http://scholar.google.com/scholar_lookup?&title=The%20geography%20of%20recent%20genetic%20ancestry%20across%20Europe&journal=PLoS%20Biol.&volume=11&publication_year=2013&author=Ralph%2CP&author=Coop%2CG"> Google Scholar</a>  </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="27"><p class="c-article-references__text" id="ref-CR27">Renfrew, C. <i>Archaeology and Language: The Puzzle of Indo-European Origins</i> (Pimlico, 1987)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 27" href="http://scholar.google.com/scholar_lookup?&title=Archaeology%20and%20Language%3A%20The%20Puzzle%20of%20Indo-European%20Origins&publication_year=1987&author=Renfrew%2CC"> Google Scholar</a>  </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="28"><p class="c-article-references__text" id="ref-CR28">Bellwood, P. <i>First Farmers: The Origins of Agricultural Societies</i> (Wiley-Blackwell, 2004)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 28" href="http://scholar.google.com/scholar_lookup?&title=First%20Farmers%3A%20The%20Origins%20of%20Agricultural%20Societies&publication_year=2004&author=Bellwood%2CP"> Google Scholar</a>  </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="29"><p class="c-article-references__text" id="ref-CR29">Gamkrelidze, T. V. & Ivanov, V. V. The early history of Indo-European languages. <i>Sci. Am.</i> <b>262</b>, 110–116 (1990)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 29" href="http://scholar.google.com/scholar_lookup?&title=The%20early%20history%20of%20Indo-European%20languages&journal=Sci.%20Am.&volume=262&pages=110-116&publication_year=1990&author=Gamkrelidze%2CTV&author=Ivanov%2CVV"> Google Scholar</a>  </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="30"><p class="c-article-references__text" id="ref-CR30">Mallory, J. P. <i>In Search of the Indo-Europeans: Language, Archaeology and Myth</i> (Thames and Hudson, 1991)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 30" href="http://scholar.google.com/scholar_lookup?&title=In%20Search%20of%20the%20Indo-Europeans%3A%20Language%2C%20Archaeology%20and%20Myth&publication_year=1991&author=Mallory%2CJP"> Google Scholar</a>  </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="31"><p class="c-article-references__text" id="ref-CR31">Kircher, M., Sawyer, S. & Meyer, M. Double indexing overcomes inaccuracies in multiplex sequencing on the Illumina platform. <i>Nucleic Acids Res.</i> <b>40</b>, e3 (2012)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:CAS:528:DC%2BC38Xit1GmtQ%3D%3D" aria-label="CAS reference 31">CAS</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=22021376" aria-label="PubMed reference 31">PubMed</a>  <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 31" href="http://scholar.google.com/scholar_lookup?&title=Double%20indexing%20overcomes%20inaccuracies%20in%20multiplex%20sequencing%20on%20the%20Illumina%20platform&journal=Nucleic%20Acids%20Res.&volume=40&publication_year=2012&author=Kircher%2CM&author=Sawyer%2CS&author=Meyer%2CM"> Google Scholar</a>  </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="32"><p class="c-article-references__text" id="ref-CR32">Meyer, M. et al. A mitochondrial genome sequence of a hominin from Sima de los Huesos. <i>Nature</i> <b>505</b>, 403–406 (2014)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="ads reference" data-track-action="ads reference" href="http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2014Natur.505..403M" aria-label="ADS reference 32">ADS</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:CAS:528:DC%2BC2cXotFaiuw%3D%3D" aria-label="CAS reference 32">CAS</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=24305051" aria-label="PubMed reference 32">PubMed</a>  <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 32" href="http://scholar.google.com/scholar_lookup?&title=A%20mitochondrial%20genome%20sequence%20of%20a%20hominin%20from%20Sima%20de%20los%20Huesos&journal=Nature&volume=505&pages=403-406&publication_year=2014&author=Meyer%2CM"> Google Scholar</a>  </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="33"><p class="c-article-references__text" id="ref-CR33">Rohland, N., Harney, E., Mallick, S., Nordenfelt, S. & Reich, D. Partial uracil–DNA–glycosylase treatment for screening of ancient DNA. <i>Phil. Trans. R. Soc. Lond. B</i> <b>370</b>, 20130624 (2015)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 33" href="http://scholar.google.com/scholar_lookup?&title=Partial%20uracil%E2%80%93DNA%E2%80%93glycosylase%20treatment%20for%20screening%20of%20ancient%20DNA&journal=Phil.%20Trans.%20R.%20Soc.%20Lond.%20B&volume=370&publication_year=2015&author=Rohland%2CN&author=Harney%2CE&author=Mallick%2CS&author=Nordenfelt%2CS&author=Reich%2CD"> Google Scholar</a>  </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="34"><p class="c-article-references__text" id="ref-CR34">Rohland, N. & Reich, D. Cost-effective, high-throughput DNA sequencing libraries for multiplexed target capture. <i>Genome Res.</i> <b>22</b>, 939–946 (2012)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:CAS:528:DC%2BC38Xms1antbk%3D" aria-label="CAS reference 34">CAS</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=22267522" aria-label="PubMed reference 34">PubMed</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed central reference" data-track-action="pubmed central reference" href="http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3337438" aria-label="PubMed Central reference 34">PubMed Central</a>  <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 34" href="http://scholar.google.com/scholar_lookup?&title=Cost-effective%2C%20high-throughput%20DNA%20sequencing%20libraries%20for%20multiplexed%20target%20capture&journal=Genome%20Res.&volume=22&pages=939-946&publication_year=2012&author=Rohland%2CN&author=Reich%2CD"> Google Scholar</a>  </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="35"><p class="c-article-references__text" id="ref-CR35">Li, H. & Durbin, R. Fast and accurate short read alignment with Burrows–Wheeler transform. <i>Bioinformatics</i> <b>25</b>, 1754–1760 (2009)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:CAS:528:DC%2BD1MXot1Cjtbo%3D" aria-label="CAS reference 35">CAS</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=19451168" aria-label="PubMed reference 35">PubMed</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed central reference" data-track-action="pubmed central reference" href="http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2705234" aria-label="PubMed Central reference 35">PubMed Central</a>  <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 35" href="http://scholar.google.com/scholar_lookup?&title=Fast%20and%20accurate%20short%20read%20alignment%20with%20Burrows%E2%80%93Wheeler%20transform&journal=Bioinformatics&volume=25&pages=1754-1760&publication_year=2009&author=Li%2CH&author=Durbin%2CR"> Google Scholar</a>  </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="36"><p class="c-article-references__text" id="ref-CR36">Behar, D. M. et al. A “Copernican” reassessment of the human mitochondrial DNA tree from its root. <i>Am. J. Hum. Genet.</i> <b>90</b>, 675–684 (2012)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:CAS:528:DC%2BC38XlsFGjsrw%3D" aria-label="CAS reference 36">CAS</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=22482806" aria-label="PubMed reference 36">PubMed</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed central reference" data-track-action="pubmed central reference" href="http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3322232" aria-label="PubMed Central reference 36">PubMed Central</a>  <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 36" href="http://scholar.google.com/scholar_lookup?&title=A%20%E2%80%9CCopernican%E2%80%9D%20reassessment%20of%20the%20human%20mitochondrial%20DNA%20tree%20from%20its%20root&journal=Am.%20J.%20Hum.%20Genet.&volume=90&pages=675-684&publication_year=2012&author=Behar%2CDM"> Google Scholar</a>  </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="37"><p class="c-article-references__text" id="ref-CR37">Lassmann, T. & Sonnhammer, E. L. L. Kalign—an accurate and fast multiple sequence alignment algorithm. <i>BMC Bioinformatics</i> <b>6</b>, 298 (2005)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=16343337" aria-label="PubMed reference 37">PubMed</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed central reference" data-track-action="pubmed central reference" href="http://www.ncbi.nlm.nih.gov/pmc/articles/PMC1325270" aria-label="PubMed Central reference 37">PubMed Central</a>  <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 37" href="http://scholar.google.com/scholar_lookup?&title=Kalign%E2%80%94an%20accurate%20and%20fast%20multiple%20sequence%20alignment%20algorithm&journal=BMC%20Bioinformatics&volume=6&publication_year=2005&author=Lassmann%2CT&author=Sonnhammer%2CELL"> Google Scholar</a>  </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="38"><p class="c-article-references__text" id="ref-CR38">Sawyer, S., Krause, J., Guschanski, K., Savolainen, V. & Pääbo, S. Temporal patterns of nucleotide misincorporations and DNA fragmentation in ancient DNA. <i>PLoS ONE</i> <b>7</b>, e34131 (2012)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="ads reference" data-track-action="ads reference" href="http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2012PLoSO...734131S" aria-label="ADS reference 38">ADS</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:CAS:528:DC%2BC38Xlslehu7k%3D" aria-label="CAS reference 38">CAS</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=22479540" aria-label="PubMed reference 38">PubMed</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed central reference" data-track-action="pubmed central reference" href="http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3316601" aria-label="PubMed Central reference 38">PubMed Central</a>  <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 38" href="http://scholar.google.com/scholar_lookup?&title=Temporal%20patterns%20of%20nucleotide%20misincorporations%20and%20DNA%20fragmentation%20in%20ancient%20DNA&journal=PLoS%20ONE&volume=7&publication_year=2012&author=Sawyer%2CS&author=Krause%2CJ&author=Guschanski%2CK&author=Savolainen%2CV&author=P%C3%A4%C3%A4bo%2CS"> Google Scholar</a>  </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="39"><p class="c-article-references__text" id="ref-CR39">Green, R. E. et al. A draft sequence of the Neandertal genome. <i>Science</i> <b>328</b>, 710–722 (2010)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="ads reference" data-track-action="ads reference" href="http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2010Sci...328..710G" aria-label="ADS reference 39">ADS</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:CAS:528:DC%2BC3cXlsFCitbo%3D" aria-label="CAS reference 39">CAS</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=20448178" aria-label="PubMed reference 39">PubMed</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed central reference" data-track-action="pubmed central reference" href="http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5100745" aria-label="PubMed Central reference 39">PubMed Central</a>  <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 39" href="http://scholar.google.com/scholar_lookup?&title=A%20draft%20sequence%20of%20the%20Neandertal%20genome&journal=Science&volume=328&pages=710-722&publication_year=2010&author=Green%2CRE"> Google Scholar</a>  </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="40"><p class="c-article-references__text" id="ref-CR40">Alexander, D. H. & Lange, K. Enhancements to the ADMIXTURE algorithm for individual ancestry estimation. <i>BMC Bioinformatics</i> <b>12</b>, 246 (2011)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=21682921" aria-label="PubMed reference 40">PubMed</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed central reference" data-track-action="pubmed central reference" href="http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3146885" aria-label="PubMed Central reference 40">PubMed Central</a>  <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 40" href="http://scholar.google.com/scholar_lookup?&title=Enhancements%20to%20the%20ADMIXTURE%20algorithm%20for%20individual%20ancestry%20estimation&journal=BMC%20Bioinformatics&volume=12&publication_year=2011&author=Alexander%2CDH&author=Lange%2CK"> Google Scholar</a>  </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="41"><p class="c-article-references__text" id="ref-CR41">Alexander, D. H., Novembre, J. & Lange, K. Fast model-based estimation of ancestry in unrelated individuals. <i>Genome Res.</i> <b>19</b>, 1655–1664 (2009)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:CAS:528:DC%2BD1MXhtFCjsLvL" aria-label="CAS reference 41">CAS</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=19648217" aria-label="PubMed reference 41">PubMed</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed central reference" data-track-action="pubmed central reference" href="http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2752134" aria-label="PubMed Central reference 41">PubMed Central</a>  <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 41" href="http://scholar.google.com/scholar_lookup?&title=Fast%20model-based%20estimation%20of%20ancestry%20in%20unrelated%20individuals&journal=Genome%20Res.&volume=19&pages=1655-1664&publication_year=2009&author=Alexander%2CDH&author=Novembre%2CJ&author=Lange%2CK"> Google Scholar</a>  </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="42"><p class="c-article-references__text" id="ref-CR42">Reich, D., Price, A. L. & Patterson, N. Principal component analysis of genetic data. <i>Nature Genet.</i> <b>40</b>, 491–492 (2008)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:CAS:528:DC%2BD1cXltFyksrc%3D" aria-label="CAS reference 42">CAS</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=18443580" aria-label="PubMed reference 42">PubMed</a>  <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 42" href="http://scholar.google.com/scholar_lookup?&title=Principal%20component%20analysis%20of%20genetic%20data&journal=Nature%20Genet.&volume=40&pages=491-492&publication_year=2008&author=Reich%2CD&author=Price%2CAL&author=Patterson%2CN"> Google Scholar</a>  </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="43"><p class="c-article-references__text" id="ref-CR43">Purcell, S. et al. PLINK: a tool set for whole-genome association and population-based linkage analyses. <i>Am. J. Hum. Genet.</i> <b>81</b>, 559–575 (2007)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:CAS:528:DC%2BD2sXhtVSqurrL" aria-label="CAS reference 43">CAS</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed reference" data-track-action="pubmed reference" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=17701901" aria-label="PubMed reference 43">PubMed</a>  <a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="pubmed central reference" data-track-action="pubmed central reference" href="http://www.ncbi.nlm.nih.gov/pmc/articles/PMC1950838" aria-label="PubMed Central reference 43">PubMed Central</a>  <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 43" href="http://scholar.google.com/scholar_lookup?&title=PLINK%3A%20a%20tool%20set%20for%20whole-genome%20association%20and%20population-based%20linkage%20analyses&journal=Am.%20J.%20Hum.%20Genet.&volume=81&pages=559-575&publication_year=2007&author=Purcell%2CS"> Google Scholar</a>  </p></li><li class="c-article-references__item js-c-reading-companion-references-item" data-counter="44"><p class="c-article-references__text" id="ref-CR44">Skoglund, P., Storå, J., Götherström, A. & Jakobsson, M. Accurate sex identification of ancient human remains using DNA shotgun sequencing. <i>J. Archaeol. Sci.</i> <b>40</b>, 4477–4482 (2013)</p><p class="c-article-references__links u-hide-print"><a data-track="click_references" rel="nofollow noopener" data-track-label="link" data-track-item_id="link" data-track-value="cas reference" data-track-action="cas reference" href="/articles/cas-redirect/1:CAS:528:DC%2BC3sXhslGjsr%2FL" aria-label="CAS reference 44">CAS</a>  <a data-track="click_references" data-track-action="google scholar reference" data-track-value="google scholar reference" data-track-label="link" data-track-item_id="link" rel="nofollow noopener" aria-label="Google Scholar reference 44" href="http://scholar.google.com/scholar_lookup?&title=Accurate%20sex%20identification%20of%20ancient%20human%20remains%20using%20DNA%20shotgun%20sequencing&journal=J.%20Archaeol.%20Sci.&volume=40&pages=4477-4482&publication_year=2013&author=Skoglund%2CP&author=Stor%C3%A5%2CJ&author=G%C3%B6therstr%C3%B6m%2CA&author=Jakobsson%2CM"> Google Scholar</a>  </p></li></ol><p class="c-article-references__download u-hide-print"><a data-track="click" data-track-action="download citation references" data-track-label="link" rel="nofollow" href="https://citation-needed.springer.com/v2/references/10.1038/nature14317?format=refman&flavour=references">Download references<svg width="16" height="16" focusable="false" role="img" aria-hidden="true" class="u-icon"><use xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="#icon-eds-i-download-medium"></use></svg></a></p></div></div></div></section></div><section data-title="Acknowledgements"><div class="c-article-section" id="Ack1-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="Ack1">Acknowledgements</h2><div class="c-article-section__content" id="Ack1-content"><p>We thank P. Bellwood, J. Burger, P. Heggarty, M. Lipson, C. Renfrew, J. Diamond, S.Pääbo, R. Pinhasi and P. Skoglund for critical comments, and the Initiative for the Science of the Human Past at Harvard for organizing a workshop around the issues touched on by this paper. We thank S. Pääbo for support for establishing the ancient DNA facilities in Boston, and P. Skoglund for detecting the presence of two related individuals in our data set. We thank L. Orlando, T. S. Korneliussen, and C. Gamba for help in obtaining data. We thank Agilent Technologies and G. Frommer for help in developing the capture reagents. We thank C. Der Sarkissian, G. Valverde, L. Papac and B. Nickel for wet laboratory support. We thank archaeologists V. Dresely, R. Ganslmeier, O. Balanvosky, J. Ignacio Royo Guillén, A. Osztás, V. Majerik, T. Paluch, K. Somogyi and V.Voicsek for sharing samples and discussion about archaeological context. This research was supported by an Australian Research Council grant to W.H. and B.L. (DP130102158), and German Research Foundation grants to K.W.A. (Al 287/7-1 and 7-3, Al 287/10-1 and Al 287/14-1) and to H.M. (Me 3245/1-1 and 1-3). D.R. was supported by US National Science Foundation HOMINID grant BCS-1032255, US National Institutes of Health grant GM100233, and the Howard Hughes Medical Institute.</p></div></div></section><section aria-labelledby="author-information" data-title="Author information"><div class="c-article-section" id="author-information-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="author-information">Author information</h2><div class="c-article-section__content" id="author-information-content"><span class="c-article-author-information__subtitle u-visually-hidden" id="author-notes">Author notes</span><ol class="c-article-author-information__list"><li class="c-article-author-information__item" id="na1"><p>Wolfgang Haak and Iosif Lazaridis: These authors contributed equally to this work.</p></li></ol><h3 class="c-article__sub-heading" id="affiliations">Authors and Affiliations</h3><ol class="c-article-author-affiliation__list"><li id="Aff1"><p class="c-article-author-affiliation__address">Australian Centre for Ancient DNA, School of Earth and Environmental Sciences & Environment Institute, University of Adelaide, Adelaide, 5005, South Australia, Australia</p><p class="c-article-author-affiliation__authors-list">Wolfgang Haak, Bastien Llamas & Alan Cooper</p></li><li id="Aff2"><p class="c-article-author-affiliation__address">Department of Genetics, Harvard Medical School, Boston, 02115, Massachusetts, USA</p><p class="c-article-author-affiliation__authors-list">Iosif Lazaridis, Nadin Rohland, Swapan Mallick, Susanne Nordenfelt, Eadaoin Harney, Kristin Stewardson, Qiaomei Fu & David Reich</p></li><li id="Aff3"><p class="c-article-author-affiliation__address">Broad Institute of Harvard and MIT, Cambridge, 02142, Massachusetts, USA</p><p class="c-article-author-affiliation__authors-list">Iosif Lazaridis, Nick Patterson, Nadin Rohland, Swapan Mallick, Susanne Nordenfelt, Eadaoin Harney, Kristin Stewardson, Qiaomei Fu & David Reich</p></li><li id="Aff4"><p class="c-article-author-affiliation__address">Howard Hughes Medical Institute, Harvard Medical School, Boston, 02115, Massachusetts, USA</p><p class="c-article-author-affiliation__authors-list">Swapan Mallick, Eadaoin Harney, Kristin Stewardson & David Reich</p></li><li id="Aff5"><p class="c-article-author-affiliation__address">Institute of Anthropology, Johannes Gutenberg University of Mainz, D-55128 Mainz, Germany, </p><p class="c-article-author-affiliation__authors-list">Guido Brandt, Nicole Nicklisch, Christina Roth, Anna Szécsényi-Nagy & Kurt Werner Alt</p></li><li id="Aff6"><p class="c-article-author-affiliation__address">Max Planck Institute for Evolutionary Anthropology, D-04103 Leipzig, Germany, </p><p class="c-article-author-affiliation__authors-list">Qiaomei Fu & Matthias Meyer</p></li><li id="Aff7"><p class="c-article-author-affiliation__address">Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, IVPP, CAS, Beijing 100049, China, </p><p class="c-article-author-affiliation__authors-list">Qiaomei Fu</p></li><li id="Aff8"><p class="c-article-author-affiliation__address">Institute for Archaeological Sciences, University of Tübingen, D-72070 Tübingen, Germany, </p><p class="c-article-author-affiliation__authors-list">Alissa Mittnik & Johannes Krause</p></li><li id="Aff9"><p class="c-article-author-affiliation__address">Institute of Archaeology, Research Centre for the Humanities, Hungarian Academy of Science, H-1014 Budapest, Hungary, </p><p class="c-article-author-affiliation__authors-list">Eszter Bánffy & Anna Szécsényi-Nagy</p></li><li id="Aff10"><p class="c-article-author-affiliation__address">Römisch Germanische Kommission (RGK) Frankfurt, D-60325 Frankfurt, Germany, </p><p class="c-article-author-affiliation__authors-list">Eszter Bánffy</p></li><li id="Aff11"><p class="c-article-author-affiliation__address">Archaeological Research Laboratory, Stockholm University, 114 18 Stockholm, Sweden, </p><p class="c-article-author-affiliation__authors-list">Christos Economou</p></li><li id="Aff12"><p class="c-article-author-affiliation__address">Departments of Paleoanthropology and Archaeogenetics, Senckenberg Center for Human Evolution and Paleoenvironment, University of Tübingen, D-72070 Tübingen, Germany, </p><p class="c-article-author-affiliation__authors-list">Michael Francken & Johannes Krause</p></li><li id="Aff13"><p class="c-article-author-affiliation__address">State Office for Heritage Management and Archaeology Saxony-Anhalt and State Museum of Prehistory, D-06114 Halle, Germany, </p><p class="c-article-author-affiliation__authors-list">Susanne Friederich, Harald Meller, Nicole Nicklisch & Kurt Werner Alt</p></li><li id="Aff14"><p class="c-article-author-affiliation__address">Departamento de Prehistoria y Arqueología, Facultad de Filosofía y Letras, Universidad Autónoma de Madrid, E-28049 Madrid, Spain, </p><p class="c-article-author-affiliation__authors-list">Rafael Garrido Pena</p></li><li id="Aff15"><p class="c-article-author-affiliation__address">The Cultural Heritage Foundation, Västerås 722 12, Sweden, </p><p class="c-article-author-affiliation__authors-list">Fredrik Hallgren</p></li><li id="Aff16"><p class="c-article-author-affiliation__address">Peter the Great Museum of Anthropology and Ethnography (Kunstkamera) RAS, St Petersburg 199034, Russia, </p><p class="c-article-author-affiliation__authors-list">Valery Khartanovich & Vayacheslav Moiseyev</p></li><li id="Aff17"><p class="c-article-author-affiliation__address">Volga State Academy of Social Sciences and Humanities, Samara 443099, Russia, </p><p class="c-article-author-affiliation__authors-list">Aleksandr Khokhlov, Pavel Kuznetsov & Oleg Mochalov</p></li><li id="Aff18"><p class="c-article-author-affiliation__address">Abteilung Madrid, Deutsches Archaeologisches Institut, E-28002 Madrid, Spain, </p><p class="c-article-author-affiliation__authors-list">Michael Kunst</p></li><li id="Aff19"><p class="c-article-author-affiliation__address">Danube Private University, A-3500 Krems, Austria, </p><p class="c-article-author-affiliation__authors-list">Nicole Nicklisch & Kurt Werner Alt</p></li><li id="Aff20"><p class="c-article-author-affiliation__address">Institute for Prehistory and Archaeological Science, University of Basel, CH-4003 Basel, Switzerland, </p><p class="c-article-author-affiliation__authors-list">Sandra L. Pichler & Kurt Werner Alt</p></li><li id="Aff21"><p class="c-article-author-affiliation__address">Departamento de Prehistòria, Universitat Autònoma de Barcelona, E-08193 Barcelona, Spain, </p><p class="c-article-author-affiliation__authors-list">Roberto Risch</p></li><li id="Aff22"><p class="c-article-author-affiliation__address">Departamento de Prehistòria y Arqueolgia, Universidad de Valladolid, E-47002 Valladolid, Spain, </p><p class="c-article-author-affiliation__authors-list">Manuel A. Rojo Guerra</p></li><li id="Aff23"><p class="c-article-author-affiliation__address">State Office for Cultural Heritage Management Baden-Württemberg, Osteology, D-78467 Konstanz, Germany, </p><p class="c-article-author-affiliation__authors-list">Joachim Wahl</p></li><li id="Aff24"><p class="c-article-author-affiliation__address">Max Planck Institute for the Science of Human History, D-07745 Jena, Germany, </p><p class="c-article-author-affiliation__authors-list">Johannes Krause</p></li><li id="Aff25"><p class="c-article-author-affiliation__address">Anthropology Department, Hartwick College, Oneonta, 13820, New York, USA</p><p class="c-article-author-affiliation__authors-list">Dorcas Brown & David Anthony</p></li></ol><div class="u-js-hide u-hide-print" data-test="author-info"><span class="c-article__sub-heading">Authors</span><ol class="c-article-authors-search u-list-reset"><li id="auth-Wolfgang-Haak-Aff1"><span class="c-article-authors-search__title u-h3 js-search-name">Wolfgang Haak</span><div class="c-article-authors-search__list"><div class="c-article-authors-search__item c-article-authors-search__list-item--left"><a href="/search?author=Wolfgang%20Haak" class="c-article-button" data-track="click" data-track-action="author link - 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W.H., E.B., C.E., M.F., S.F., R.G.P., F.H., V.K., A.K., M.K., P.K., H.M., O.M., V.M., N.N., S.L.P., R.R., M.A.R.G., C.R., A.S.-N., J.W., J.K., D.B., D.A., A.C., K.W.A. and D.R. assembled archaeological material, W.H., I.L., N.P., N.R., S.M., A.M. and D.R. analysed genetic data. I.L., N.P. and D.R. developed methods using <i>f</i> statistics for inferring admixture proportions. W.H., N.R., B.L., G.B., S.N., E.H., K.S. and A.M. performed wet laboratory ancient DNA work. I.L., N.R., S.M., B.L., Q.F., M.M. and D.R. developed the 390k capture reagent. W.H., I.L. and D.R. wrote the manuscript with help from all co-authors.</p><h3 class="c-article__sub-heading" id="corresponding-author">Corresponding author</h3><p id="corresponding-author-list">Correspondence to <a id="corresp-c1" href="mailto:reich@genetics.med.harvard.edu">David Reich</a>.</p></div></div></section><section data-title="Ethics declarations"><div class="c-article-section" id="ethics-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="ethics">Ethics declarations</h2><div class="c-article-section__content" id="ethics-content"> <h3 class="c-article__sub-heading">Competing interests</h3> <p>The authors declare no competing financial interests.</p> </div></div></section><section data-title="Extended data figures and tables"><div class="c-article-section" id="Sec10-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="Sec10">Extended data figures and tables</h2><div class="c-article-section__content" id="Sec10-content"><div data-test="supplementary-info"><div id="figshareContainer" class="c-article-figshare-container" data-test="figshare-container"></div><div class="c-article-supplementary__item js-c-reading-companion-figures-item" data-test="supp-item" id="Fig4"><h3 class="c-article-supplementary__title u-h3"><a class="print-link" data-track="click" data-track-action="view supplementary info" data-test="supp-info-link" data-track-label="extended data figure 1 outgroup f3 statistic f3(di" href="/articles/nature14317/figures/4" data-supp-info-image="//media.springernature.com/lw685/springer-static/image/art%3A10.1038%2Fnature14317/MediaObjects/41586_2015_Article_BFnature14317_Fig4_ESM.jpg">Extended Data Figure 1 Outgroup <i>f</i><sub>3</sub> statistic <i>f</i><sub>3</sub>(Dinka; <i>X</i>, <i>Y</i>), measuring the degree of shared drift among pairs of ancient individuals.</a></h3></div><div class="c-article-supplementary__item js-c-reading-companion-figures-item" data-test="supp-item" id="Fig5"><h3 class="c-article-supplementary__title u-h3"><a class="print-link" data-track="click" data-track-action="view supplementary info" data-test="supp-info-link" data-track-label="extended data figure 2 modelling corded ware as a " href="/articles/nature14317/figures/5" data-supp-info-image="//media.springernature.com/lw685/springer-static/image/art%3A10.1038%2Fnature14317/MediaObjects/41586_2015_Article_BFnature14317_Fig5_ESM.jpg">Extended Data Figure 2 Modelling Corded Ware as a mixture of <i>N</i> = 1, 2, or 3 ancestral populations.</a></h3><div class="c-article-supplementary__description" data-component="thumbnail-container"><p><b>a</b>, The left column shows a histogram of raw <i>f</i><sub>4</sub> statistic residuals and on the right <i>Z</i>-scores for the best-fitting (lowest squared 2-norm of the residuals, or <i>resnorm</i>) model at each <i>N</i>. <b>b</b>, The data on the left show <i>resnorm</i> and on the right show the maximum |<i>Z</i>| score change for different <i>N</i>. <b>c</b>, <i>resnorm</i> of different <i>N</i> = 2 models. The set of outgroups used in this analysis in the terminology of <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/nature14317#MOESM29">Supplementary Information</a> section 9 is ‘World Foci 15 + Ancients’.</p></div></div><div class="c-article-supplementary__item js-c-reading-companion-figures-item" data-test="supp-item" id="Fig6"><h3 class="c-article-supplementary__title u-h3"><a class="print-link" data-track="click" data-track-action="view supplementary info" data-test="supp-info-link" data-track-label="extended data figure 3 modelling europeans as mixt" href="/articles/nature14317/figures/6" data-supp-info-image="//media.springernature.com/lw685/springer-static/image/art%3A10.1038%2Fnature14317/MediaObjects/41586_2015_Article_BFnature14317_Fig6_ESM.jpg">Extended Data Figure 3 Modelling Europeans as mixtures of increasing complexity: <i>N</i> = 1 (EN), <i>N</i> = 2 (EN, WHG), <i>N</i> = 3 (EN, WHG, Yamnaya), <i>N</i> = 4 (EN, WHG, Yamnaya, Nganasan), <i>N</i> = 5 (EN, WHG, Yamnaya, Nganasan, BedouinB).</a></h3><div class="c-article-supplementary__description" data-component="thumbnail-container"><p>The residual norm of the fitted model (<a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/nature14317#MOESM29">Supplementary Information</a> section 9) and its changes are indicated.</p></div></div><div class="c-article-supplementary__item js-c-reading-companion-figures-item" data-test="supp-item" id="Fig7"><h3 class="c-article-supplementary__title u-h3"><a class="print-link" data-track="click" data-track-action="view supplementary info" data-test="supp-info-link" data-track-label="extended data figure 4 geographic distribution of " href="/articles/nature14317/figures/7" data-supp-info-image="//media.springernature.com/lw685/springer-static/image/art%3A10.1038%2Fnature14317/MediaObjects/41586_2015_Article_BFnature14317_Fig7_ESM.jpg">Extended Data Figure 4 Geographic distribution of archaeological cultures and graphic illustration of proposed population movements / turnovers discussed in the main text.</a></h3><div class="c-article-supplementary__description" data-component="thumbnail-container"><p><b>a</b>, Proposed routes of migration by early farmers into Europe <span class="stix">∼</span>9,000−7000 years ago. <b>b</b>, Resurgence of hunter-gatherer ancestry during the Middle Neolithic 7,000−5,000 years ago. <b>c</b>, Arrival of steppe ancestry in central Europe during the Late Neolithic <span class="stix">∼</span>4,500 years ago. White arrows indicate the two possible scenarios of the arrival of Indo-European language groups. Symbols of samples are identical to those in <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/nature14317#Fig1">Fig. 1</a>.</p></div></div><div class="c-article-supplementary__item" data-test="supp-item"><div class="c-article-table" data-test="inline-table" data-container-section="table" id="table-1"><figure><figcaption class="c-article-table__figcaption"><b id="Tab1" data-test="table-caption">Extended Data Table 1 Number of ancient Eurasian modern human samples screened in genome-wide studies to date</b></figcaption><div class="u-text-right u-hide-print"><a class="c-article__pill-button" data-test="table-link" data-track="click" data-track-action="view table" data-track-label="button" rel="nofollow" href="/articles/nature14317/tables/1" aria-label="Full size table 1"><span>Full size table</span><svg width="16" height="16" focusable="false" role="img" aria-hidden="true" class="u-icon"><use xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="#icon-eds-i-chevron-right-small"></use></svg></a></div></figure></div></div><div class="c-article-supplementary__item" data-test="supp-item"><div class="c-article-table" data-test="inline-table" data-container-section="table" id="table-2"><figure><figcaption class="c-article-table__figcaption"><b id="Tab2" data-test="table-caption">Extended Data Table 2 Summary of the archaeological context for the 69 newly reported samples</b></figcaption><div class="u-text-right u-hide-print"><a class="c-article__pill-button" data-test="table-link" data-track="click" data-track-action="view table" data-track-label="button" rel="nofollow" href="/articles/nature14317/tables/2" aria-label="Full size table 2"><span>Full size table</span><svg width="16" height="16" focusable="false" role="img" aria-hidden="true" class="u-icon"><use xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="#icon-eds-i-chevron-right-small"></use></svg></a></div></figure></div></div><div class="c-article-supplementary__item" data-test="supp-item"><div class="c-article-table" data-test="inline-table" data-container-section="table" id="table-3"><figure><figcaption class="c-article-table__figcaption"><b id="Tab3" data-test="table-caption">Extended Data Table 3 Pairwise <i>F</i><sub>ST</sub> for all ancient groups with ≥ 2 individuals, present-day Europeans with ≥ 10 individuals, and selected other groups</b></figcaption><div class="u-text-right u-hide-print"><a class="c-article__pill-button" data-test="table-link" data-track="click" data-track-action="view table" data-track-label="button" rel="nofollow" href="/articles/nature14317/tables/3" aria-label="Full size table 3"><span>Full size table</span><svg width="16" height="16" focusable="false" role="img" aria-hidden="true" class="u-icon"><use xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="#icon-eds-i-chevron-right-small"></use></svg></a></div></figure></div></div></div></div></div></section><section data-title="Supplementary information"><div class="c-article-section" id="Sec11-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="Sec11">Supplementary information</h2><div class="c-article-section__content" id="Sec11-content"><div data-test="supplementary-info"><div class="c-article-supplementary__item" data-test="supp-item" id="MOESM29"><h3 class="c-article-supplementary__title u-h3"><a class="print-link" data-track="click" data-track-action="view supplementary info" data-test="supp-info-link" data-track-label="supplementary information" href="https://static-content.springer.com/esm/art%3A10.1038%2Fnature14317/MediaObjects/41586_2015_BFnature14317_MOESM29_ESM.pdf" data-supp-info-image="">Supplementary Information</a></h3><div class="c-article-supplementary__description" data-component="thumbnail-container"><p>This file contains Supplementary Information sections 1-11, see contents page for more details (PDF 31143 kb)</p></div></div><div class="c-article-supplementary__item" data-test="supp-item" id="MOESM30"><h3 class="c-article-supplementary__title u-h3"><a class="print-link" data-track="click" data-track-action="view supplementary info" data-test="supp-info-link" data-track-label="supplementary data" href="https://static-content.springer.com/esm/art%3A10.1038%2Fnature14317/MediaObjects/41586_2015_BFnature14317_MOESM30_ESM.xlsx" data-supp-info-image="">Supplementary Data</a></h3><div class="c-article-supplementary__description" data-component="thumbnail-container"><p>This file contains Supplementary Data 1. 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id="citeas">Cite this article</h3><p class="c-bibliographic-information__citation">Haak, W., Lazaridis, I., Patterson, N. <i>et al.</i> Massive migration from the steppe was a source for Indo-European languages in Europe. <i>Nature</i> <b>522</b>, 207–211 (2015). https://doi.org/10.1038/nature14317</p><p class="c-bibliographic-information__download-citation u-hide-print"><a data-test="citation-link" data-track="click" data-track-action="download article citation" data-track-label="link" data-track-external="" rel="nofollow" href="https://citation-needed.springer.com/v2/references/10.1038/nature14317?format=refman&flavour=citation">Download citation<svg width="16" height="16" focusable="false" role="img" aria-hidden="true" class="u-icon"><use xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="#icon-eds-i-download-medium"></use></svg></a></p><ul class="c-bibliographic-information__list" data-test="publication-history"><li 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Massive migration from the steppe was a source for Indo-European languages in Europe | Nature