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The genetic prehistory of the Greater Caucasus | bioRxiv

<!DOCTYPE html> <html lang="en" dir="ltr" xmlns="http://www.w3.org/1999/xhtml" xmlns:mml="http://www.w3.org/1998/Math/MathML"> <head prefix="og: http://ogp.me/ns# article: http://ogp.me/ns/article# book: http://ogp.me/ns/book#" > <!--[if IE]><![endif]--> <link rel="dns-prefetch" href="//d33xdlntwy0kbs.cloudfront.net" /> <link rel="dns-prefetch" href="//www.google.com" /> <link rel="dns-prefetch" href="//scholar.google.com" /> <link rel="dns-prefetch" href="//www.googletagmanager.com" /> <meta http-equiv="Content-Type" content="text/html; charset=utf-8" /> <link rel="shortcut icon" href="https://www.biorxiv.org/sites/default/files/images/favicon.ico" type="image/vnd.microsoft.icon" /> <meta name="viewport" content="width=device-width, initial-scale=1" /> <link rel="alternate" type="application/pdf" title="Full Text (PDF)" href="/content/10.1101/322347v1.full.pdf" /> <link rel="alternate" type="text/plain" title="Full Text (Plain)" href="/content/10.1101/322347v1.full.txt" /> <meta name="type" content="article" /> <meta name="category" content="article" /> <meta name="HW.identifier" content="/biorxiv/early/2018/05/16/322347.atom" /> <meta name="HW.pisa" content="biorxiv;322347v1" /> <meta name="DC.Format" content="text/html" /> <meta name="DC.Language" content="en" /> <meta name="DC.Title" content="The genetic prehistory of the Greater Caucasus" /> <meta name="DC.Identifier" content="10.1101/322347" /> <meta name="DC.Date" content="2018-05-16" /> <meta name="DC.Publisher" content="Cold Spring Harbor Laboratory" /> <meta name="DC.Rights" content="© 2018, Posted by Cold Spring Harbor Laboratory. This pre-print is available under a Creative Commons License (Attribution-NonCommercial-NoDerivs 4.0 International), CC BY-NC-ND 4.0, as described at http://creativecommons.org/licenses/by-nc-nd/4.0/" /> <meta name="DC.AccessRights" content="restricted" /> <meta name="DC.Description" content="Archaeogenetic studies have described the formation of Eurasian ‘steppe ancestry’ as a mixture of Eastern and Caucasus hunter-gatherers. However, it remains unclear when and where this ancestry arose and whether it was related to a horizon of cultural innovations in the 4th millennium BCE that subsequently facilitated the advance of pastoral societies likely linked to the dispersal of Indo-European languages. To address this, we generated genome-wide SNP data from 45 prehistoric individuals along a 3000-year temporal transect in the North Caucasus. We observe a genetic separation between the groups of the Caucasus and those of the adjacent steppe. The Caucasus groups are genetically similar to contemporaneous populations south of it, suggesting that – unlike today – the Caucasus acted as a bridge rather than an insurmountable barrier to human movement. The steppe groups from Yamnaya and subsequent pastoralist cultures show evidence for previously undetected farmer-related ancestry from different contact zones, while Steppe Maykop individuals harbour additional Upper Palaeolithic Siberian and Native American related ancestry." /> <meta name="DC.Contributor" content="Chuan-Chao Wang" /> <meta name="DC.Contributor" content="Sabine Reinhold" /> <meta name="DC.Contributor" content="Alexey Kalmykov" /> <meta name="DC.Contributor" content="Antje Wissgott" /> <meta name="DC.Contributor" content="Guido Brandt" /> <meta name="DC.Contributor" content="Choongwon Jeong" /> <meta name="DC.Contributor" content="Olivia Cheronet" /> <meta name="DC.Contributor" content="Matthew Ferry" /> <meta name="DC.Contributor" content="Eadaoin Harney" /> <meta name="DC.Contributor" content="Denise Keating" /> <meta name="DC.Contributor" content="Swapan Mallick" /> <meta name="DC.Contributor" content="Nadin Rohland" /> <meta name="DC.Contributor" content="Kristin Stewardson" /> <meta name="DC.Contributor" content="Anatoly R. Kantorovich" /> <meta name="DC.Contributor" content="Vladimir E. Maslov" /> <meta name="DC.Contributor" content="Vladimira G. Petrenko" /> <meta name="DC.Contributor" content="Vladimir R. Erlikh" /> <meta name="DC.Contributor" content="Biaslan Ch. Atabiev" /> <meta name="DC.Contributor" content="Rabadan G. Magomedov" /> <meta name="DC.Contributor" content="Philipp L. Kohl" /> <meta name="DC.Contributor" content="Kurt W. Alt" /> <meta name="DC.Contributor" content="Sandra L. Pichler" /> <meta name="DC.Contributor" content="Claudia Gerling" /> <meta name="DC.Contributor" content="Harald Meller" /> <meta name="DC.Contributor" content="Benik Vardanyan" /> <meta name="DC.Contributor" content="Larisa Yeganyan" /> <meta name="DC.Contributor" content="Alexey D. Rezepkin" /> <meta name="DC.Contributor" content="Dirk Mariaschk" /> <meta name="DC.Contributor" content="Natalia Berezina" /> <meta name="DC.Contributor" content="Julia Gresky" /> <meta name="DC.Contributor" content="Katharina Fuchs" /> <meta name="DC.Contributor" content="Corina Knipper" /> <meta name="DC.Contributor" content="Stephan Schiffels" /> <meta name="DC.Contributor" content="Elena Balanovska" /> <meta name="DC.Contributor" content="Oleg Balanovsky" /> <meta name="DC.Contributor" content="Iain Mathieson" /> <meta name="DC.Contributor" content="Thomas Higham" /> <meta name="DC.Contributor" content="Yakov B. Berezin" /> <meta name="DC.Contributor" content="Alexandra Buzhilova" /> <meta name="DC.Contributor" content="Viktor Trifonov" /> <meta name="DC.Contributor" content="Ron Pinhasi" /> <meta name="DC.Contributor" content="Andrej B. Belinskiy" /> <meta name="DC.Contributor" content="David Reich" /> <meta name="DC.Contributor" content="Svend Hansen" /> <meta name="DC.Contributor" content="Johannes Krause" /> <meta name="DC.Contributor" content="Wolfgang Haak" /> <meta name="article:published_time" content="2018-05-16" /> <meta name="article:section" content="New Results" /> <meta name="citation_title" content="The genetic prehistory of the Greater Caucasus" /> <meta name="citation_abstract" lang="en" content="&lt;h3&gt;Abstract&lt;/h3&gt; &lt;p&gt;Archaeogenetic studies have described the formation of Eurasian ‘steppe ancestry’ as a mixture of Eastern and Caucasus hunter-gatherers. However, it remains unclear when and where this ancestry arose and whether it was related to a horizon of cultural innovations in the 4&lt;sup&gt;th&lt;/sup&gt; millennium BCE that subsequently facilitated the advance of pastoral societies likely linked to the dispersal of Indo-European languages. To address this, we generated genome-wide SNP data from 45 prehistoric individuals along a 3000-year temporal transect in the North Caucasus. We observe a genetic separation between the groups of the Caucasus and those of the adjacent steppe. The Caucasus groups are genetically similar to contemporaneous populations south of it, suggesting that – unlike today – the Caucasus acted as a bridge rather than an insurmountable barrier to human movement. The steppe groups from Yamnaya and subsequent pastoralist cultures show evidence for previously undetected farmer-related ancestry from different contact zones, while Steppe Maykop individuals harbour additional Upper Palaeolithic Siberian and Native American related ancestry.&lt;/p&gt;" /> <meta name="citation_journal_title" content="bioRxiv" /> <meta name="citation_publisher" content="Cold Spring Harbor Laboratory" /> <meta name="citation_publication_date" content="2018/01/01" /> <meta name="citation_mjid" content="biorxiv;322347v1" /> <meta name="citation_id" content="322347v1" /> <meta name="citation_public_url" content="https://www.biorxiv.org/content/10.1101/322347v1" /> <meta name="citation_abstract_html_url" content="https://www.biorxiv.org/content/10.1101/322347v1.abstract" /> <meta name="citation_full_html_url" content="https://www.biorxiv.org/content/10.1101/322347v1.full" /> <meta name="citation_pdf_url" 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Early Levallois technology and the Lower to Middle Paleolithic transition in the Southern Caucasus. Science 345, 1609–1613 (2014)." /> <meta name="citation_reference" content="Pinhasi R, et al. New chronology for the Middle Palaeolithic of the southern Caucasussuggests early demise of Neanderthals in this region. Journalof Human Evolution 63, 770–780 (2012)." /> <meta name="citation_reference" content="Lordkipanidze D, et al. A complete skull from Dmanisi, Georgia, and the evolutionary biology of early Homo. Science 342, 326–331 (2013)." /> <meta name="citation_reference" content="Yunusbayev B, et al. The caucasus as an asymmetric semipermeable barrier to ancient human migrations. Molecular Biology and Evolution 29, 359–365 (2012)." /> <meta name="citation_reference" content="Balanovsky O, et al. Parallel evolution of genes and languages in the Caucasus region. Mol Biol Evol 28, 2905–2920 (2011)." /> <meta name="citation_reference" content="Orth A, et al. [Polytypic species Mus musculus in Transcaucasia]. C R Acad Sci III319, 435–441 (1996)." /> <meta name="citation_reference" content="Manceau V, Despres L, Bouvet J, Taberlet P. Systematics of the genus Capra inferred from mitochondrial DNA sequence data. Mol Phylogenet Evol 13, 504–510 (1999)." /> <meta name="citation_reference" content="Seddon JM, Santucci F, Reeve N, Hewitt GM. Caucasus Mountains divide postulated postglacial colonization routes in the white-breasted hedgehog, Erinaceus concolor. Journal of Evolutionary Biology 15, 463–467 (2002)." /> <meta name="citation_reference" content="Helwing B, et al. The Kura projects: New research on the later prehistory of the southern Caucasus. In: Archaologie in Iran und Turan (ed^(eds). Dietrich Reimer Verlag (2017)." /> <meta name="citation_reference" content="Stein GJ. The Development of Indigenous Social Complexity in Late Chalcolithic Upper Mesopotamia in the 5th-4th Millennia BC - An Initial Assessment. Origini 34, 125–151 (2014)." /> <meta name="citation_reference" content="citation_title=The Cambridge World Prehistory;citation_year=2014" /> <meta name="citation_reference" content="Kohl P. The Making of Bronze Age Eurasia Cambridge University Press (2007)." /> <meta name="citation_reference" content="Librado P, et al. The Evolutionary Origin and Genetic Makeup of Domestic Horses. Genetics 204, 423–434 (2016)." /> <meta name="citation_reference" content="Benecke N, Becker C, Kuchelmann HC. Finding the Woolly Sheep. Metaanalyses of archaeozoological data from Southwest-Asia and Southeast-Europe. e-Forschungsberichte 1, 12–18 (2017)." /> <meta name="citation_reference" content="citation_title=Appropriating innovations: entangled knowledge in Eurasia 5000-1500 BCE, Papers of the Conference;citation_year=2017" /> <meta name="citation_reference" content="Anthony DW, Brown DR. The Secondary Products Revolution, Horse-Riding, and Mounted Warfare. J World Prehist 24, 131–160 (2011)." /> <meta name="citation_reference" content="Anthony DW. The Horse, the Wheel, and Language: How Bronze-Age Riders from the Eurasian Steppes Shaped theModern World. Princeton University Press (2007)." /> <meta name="citation_reference" content="Frachetti MD. Multiregional Emergence of Mobile Pastoralism and Nonuniform Institutional Complexity across Eurasia. Current Anthropology 53, 2–38 (2012)." /> <meta name="citation_reference" content="Haak W, et al. Massive migration from the steppe was a source for Indo-European languages in Europe. Nature 522, 207–211 (2015)." /> <meta name="citation_reference" content="Allentoft ME, et al. Population genomics of Bronze Age Eurasia. Nature 522, 167–172 (2015)." /> <meta name="citation_reference" content="Narasimhan VM, et al. The Genomic Formation of South and Central Asia. bioRxiv, (2018)." /> <meta name="citation_reference" content="Govedarica B. Zeptertrager, Herrscher der Steppen: Die frühen Ockergräber des älteren Äneolithikums im karpatenbalkanischen Gebiet und im Steppenraum Sudost- ünd Osteuropas. von Zabern (2004)." /> <meta name="citation_reference" content="Lazaridis I, et al. Genomic insights into the origin of farming in the ancient Near East. Nature 536, 419–424 (2016)." /> <meta name="citation_reference" content="Mathieson I, et al. The genomic history of southeastern Europe. Nature, (2018)." /> <meta name="citation_reference" content="Günther T, Jakobsson M. Genes mirror migrations and cultures in prehistoric Europe-a population genomic perspective. Curr Opin Genet Dev 41, 115–123 (2016)." /> <meta name="citation_reference" content="Lazaridis I. et al. Ancient human genomes suggest three ancestral populations for present-day Europeans. Nature 513, 409–413 (2014)." /> <meta name="citation_reference" content="Mathieson I, et al. Genome-wide patterns of selection in 230 ancient Eurasians. Nature 528, 499–503 (2015)." /> <meta name="citation_reference" content="Jones ER, et al. Upper Palaeolithic genomes reveal deep roots of modem Eurasians. NatCommun 6, 8912 (2015)." /> <meta name="citation_reference" content="Broushaki F, et al. Early Neolithic genomes from the eastern Fertile Crescent. Science 353, 499–503 (2016)." /> <meta name="citation_reference" content="Schonberg A, Theunert C, Li M, Stoneking M, Nasidze I. High-throughput sequencing of complete human mtDNA genomes from the Caucasus and West Asia: high diversity and demographic inferences. Eur JHum Genet 19, 988–994 (2011)." /> <meta name="citation_reference" content="Stein G. The Development of Indigenous Social Complexity in Late Chalcolithic Upper Mesopotamia in the 5th-4th Millennia BC: An Initial Assesment. Origini, 125–151 (2012)." /> <meta name="citation_reference" content="Fu Q, et al. An early modern human from Romania with a recent Neanderthal ancestor. Nature 524, 216–219 (2015)." /> <meta name="citation_reference" content="Mallick S, et al. The Simons Genome Diversity Project: 300 genomes from 142 diverse populations. Nature 538, 201–206 (2016)." /> <meta name="citation_reference" content="Lipson M, et al. Parallel palaeogenomic transects reveal complex genetic history of early European farmers. Nature 551, 368–372 (2017)." /> <meta name="citation_reference" content="Olalde I, et al. The Beaker phenomenon and the genomic transformation of northwest Europe. Nature, (2018)." /> <meta name="citation_reference" content="Raghavan M, et al. Upper Palaeolithic Siberian genome reveals dual ancestry of Native Americans. Nature 505, 87–91 (2014)." /> <meta name="citation_reference" content="Fu Q, et al. The genetic history of Ice Age Europe. Nature 534, 200–205 (2016)." /> <meta name="citation_reference" content="Kilinc GM, et al. The Demographic Development of the First Farmers in Anatolia. Curr Biol 26, 2659–2666 (2016)." /> <meta name="citation_reference" content="Omrak A, et al. Genomic Evidence Establishes Anatolia as the Source of the European Neolithic Gene Pool. Curr Biol 26, 270–275 (2016)." /> <meta name="citation_reference" content="Gallego-Llorente M, et al. The genetics of an early Neolithic pastoralist from the Zagros, Iran. Sci Rep 6, 31326 (2016)." /> <meta name="citation_reference" content="Hofmanova Z, et al. Early farmers from across Europe directly descended from Neolithic Aegeans. Proc Natl Acad Sci U S A 113, 6886–6891 (2016)." /> <meta name="citation_reference" content="Olalde I, et al. Derived immune and ancestral pigmentation alleles in a 7,000-year-old Mesolithic European. Nature 507, 225–228 (2014)." /> <meta name="citation_reference" content="Gamba C, et al. Genome flux and stasis in a five millennium transect of European prehistory. Nat Commun 5, 5257 (2014)." /> <meta name="citation_reference" content="Patterson N, et al. Ancient admixture in human history. Genetics 192, 10651093 (2012)." /> <meta name="citation_reference" content="Alexander DH, Novembre J, Lange K. Fast model-based estimation of ancestry in unrelated individuals. Genome Res 19, 1655–1664 (2009)." /> <meta name="citation_reference" content="Loh P-R, et al. Inferring admixture histories of human populations using linkage disequilibrium. Genetics 193, 1233–1254 (2013)." /> <meta name="citation_reference" content="Moorjani P, Sankararaman S, Fu Q, Przeworski M, Patterson N, Reich D. A genetic method for dating ancient genomes provides a direct estimate of human generation interval in the last 45,000 years. Proc Natl Acad Sci U S A 113, 5652–5657 (2016)." /> <meta name="citation_reference" content="Forsyth J. The Caucasus. A history. Cambridge University Press (2013)." /> <meta name="citation_reference" content="citation_title=Adaptaciya kulturpaleolita - eneolita k izmeniyam prirodnoy sredy na Severo-Zapadnom Kavkaze;citation_year=2009" /> <meta name="citation_reference" content="Gorelik A. Zu kaukasischen und vorderasiatischen Einflussen bei der Neolithisierung im unteren Donbecken. Eurasia Antiqua 20, 143–170 (2014[2017])." /> <meta name="citation_reference" content="Pitskhelauri K. Uruk migrants in the Caucasus. Bull Georg Natl Acad Sci 6, (2012)." /> <meta name="citation_reference" content="Sherratt A. Economy and society in prehistoric Europe: Changing perspectives. Princeton University Press (1997)." /> <meta name="citation_reference" content="citation_title=Western Anatolia before Troy. Proto-Urbanisation in the 4th Millenium BC?;citation_year=2014" /> <meta name="citation_reference" content="citation_title=Appropriating Innovation. Entangeled knowledge in Eurasia, 5000-1500 BCE;citation_year=2017" /> <meta name="citation_reference" content="Shishlina N. Reconstruction of the Bronze Age of the Caspian steppes: Life styles and life ways of pastoral nomads. Archaeopress [u.a.] (2008)." /> <meta name="citation_reference" content="citation_title=Von Majkop bis Trialeti. Gewinnung und Verbreitung von Metallen und Obsidian in Kaukasien im 4.-2. Jt. v. Chr.;citation_year=2010" /> <meta name="citation_reference" content="Szmyt M. Fourth-third millennium BC stone cist graves between the Carpathians and Crimea. An outline of issues. Baltic-Pontic Studies 19, 107–147 (2014)." /> <meta name="citation_reference" content="citation_title=At the Northern Frontier of Near Eastern Archaeology: Recent Research on Caucasia and Anatolia in the Bronze Age;citation_year=2017" /> <meta name="citation_reference" content="Rassamakin JJ. Die nordpontische Steppe in der Kupferzeit: Gräber aus der Mitte des 5. Jts. bis Ende des 4. Jt. v. Chr. von Zabern (2004)." /> <meta name="citation_reference" content="Trifonov VA. Zapadne predeli rasprostraneniya maykopskoy kultury. Izvestiya Samarskogo Nauchnogo Centra Rossiyskoy Akademii Nauk 16, 276–284 (2014)." /> <meta name="citation_reference" content="citation_title=Transitions to the bronze age;citation_year=2013" /> <meta name="citation_reference" content="Furholt M. Upending a ‘Totality’: Re-evaluating Corded Ware Variability in Late Neolithic Europe. P Prehist Soc 80, 67–86 (2014)." /> <meta name="citation_reference" content="Furholt M. Massive Migrations? The Impact of Recent aDNA Studies on our View of Third MillenniumEurope. European Journal of Archaeology, 1–33 (2017)." /> <meta name="citation_reference" content="Mallory JP. In Search of the Indo-Europeans: Language, Archaeology and Myth. Thames and Hudson (1991)." /> <meta name="citation_reference" content="Dabney J, et al. Complete mitochondrial genome sequence of a Middle Pleistocene cave bear reconstructed from ultrashort DNA fragments. Proc Natl Acad Sci USA 110, 15758–15763 (2013)." /> <meta name="citation_reference" content="Rohland N, Harney E, Mallick S, Nordenfelt S, Reich D. Partial uracil-DNA-glycosylase treatment for screening of ancient DNA. Philos Trans R Soc Lond B Biol Sci 370, 20130624 (2015)." /> <meta name="citation_reference" content="Renaud G, Stenzel U, Kelso J. leeHom: adaptor trimming and merging for Illumina sequencing reads]. Nucleic Acids Res 42, e141 (2014)." /> <meta name="citation_reference" content="Peltzer A, et al. EAGER: efficient ancient genome reconstruction. Genome Biol 17, 60 (2016)." /> <meta name="citation_reference" content="Li H, Durbin R. Fast and accurate short read alignment with Burrows-Wheeler transform. Bioinformatics 25, (2009)." /> <meta name="citation_reference" content="Korneliussen TS, Albrechtsen A, Nielsen R. ANGSD: Analysis of Next Generation Sequencing Data. BMC Bioinformatics 15, 1–13 (2014)." /> <meta name="citation_reference" content="Jeong C, et al. Characterizing the genetic history of admixture across inner Eurasia. bioRxiv, (2018)." /> <meta name="citation_reference" content="Purcell S, et al. PLINK: a tool set for whole-genome association and population-based linkage analyses. Am J Hum Genet 81, 559–575 (2007)." /> <meta name="citation_reference" content="Li H, et al. The Sequence Alignment/Map format and SAMtools. Bioinformatics 25, 2078–2079 (2009)." /> <meta name="citation_reference" content="Kearse M, et al. Geneious Basic: an integrated and extendable desktop software platform for the organization and analysis of sequence data. Bioinformatics 28, 1647–1649 (2012)." /> <meta name="citation_reference" content="Weissensteiner H, et al. HaploGrep 2: mitochondrial haplogroup classification in the era of high-throughput sequencing. Nucleic Acids Res 44, W58–63 (2016)." /> <meta name="citation_reference" content="Fu Q, et al. A revised timescale for human evolution based on ancient mitochondrial genomes. Curr Biol 23, 553–559 (2013)." /> <meta name="citation_reference" content="Lipatov M, Sanjeev K, Patro R, Veeramah K. Maximum Likelihood Estimation of Biological Relatedness from Low Coverage Sequencing Data. bioRxiv, (2015)." /> <meta name="citation_reference" content="Monroy Kuhn JM, Jakobsson M, Günther T. Estimating genetic kin relationships in prehistoric populations. bioRxiv, (2017)." /> <meta name="twitter:title" content="The genetic prehistory of the Greater Caucasus" /> <meta name="twitter:site" content="@biorxivpreprint" /> <meta name="twitter:card" content="summary" /> <meta name="twitter:image" content="https://www.biorxiv.org/sites/default/files/images/biorxiv_logo_homepage7-5-small.png" /> <meta name="twitter:description" content="Archaeogenetic studies have described the formation of Eurasian ‘steppe ancestry’ as a mixture of Eastern and Caucasus hunter-gatherers. However, it remains unclear when and where this ancestry arose and whether it was related to a horizon of cultural innovations in the 4th millennium BCE that subsequently facilitated the advance of pastoral societies likely linked to the dispersal of Indo-European languages. To address this, we generated genome-wide SNP data from 45 prehistoric individuals along a 3000-year temporal transect in the North Caucasus. We observe a genetic separation between the groups of the Caucasus and those of the adjacent steppe. The Caucasus groups are genetically similar to contemporaneous populations south of it, suggesting that – unlike today – the Caucasus acted as a bridge rather than an insurmountable barrier to human movement. The steppe groups from Yamnaya and subsequent pastoralist cultures show evidence for previously undetected farmer-related ancestry from different contact zones, while Steppe Maykop individuals harbour additional Upper Palaeolithic Siberian and Native American related ancestry." /> <meta name="og-title" property="og:title" content="The genetic prehistory of the Greater Caucasus" /> <meta name="og-url" property="og:url" content="https://www.biorxiv.org/content/10.1101/322347v1" /> <meta name="og-site-name" property="og:site_name" content="bioRxiv" /> <meta name="og-description" property="og:description" content="Archaeogenetic studies have described the formation of Eurasian ‘steppe ancestry’ as a mixture of Eastern and Caucasus hunter-gatherers. However, it remains unclear when and where this ancestry arose and whether it was related to a horizon of cultural innovations in the 4th millennium BCE that subsequently facilitated the advance of pastoral societies likely linked to the dispersal of Indo-European languages. To address this, we generated genome-wide SNP data from 45 prehistoric individuals along a 3000-year temporal transect in the North Caucasus. We observe a genetic separation between the groups of the Caucasus and those of the adjacent steppe. The Caucasus groups are genetically similar to contemporaneous populations south of it, suggesting that – unlike today – the Caucasus acted as a bridge rather than an insurmountable barrier to human movement. The steppe groups from Yamnaya and subsequent pastoralist cultures show evidence for previously undetected farmer-related ancestry from different contact zones, while Steppe Maykop individuals harbour additional Upper Palaeolithic Siberian and Native American related ancestry." /> <meta name="og-type" property="og:type" content="article" /> <meta name="og-image" property="og:image" content="https://www.biorxiv.org/sites/default/files/images/biorxiv_logo_homepage7-5-small.png" /> <meta name="citation_date" content="2018-05-16" /> <link rel="alternate" type="application/vnd.ms-powerpoint" title="Powerpoint" href="/content/10.1101/322347v1.ppt" /> <meta name="description" content="bioRxiv - the preprint server for biology, operated by Cold Spring Harbor Laboratory, a research and educational institution" /> <meta name="generator" content="Drupal 7 (http://drupal.org)" /> <link rel="canonical" href="https://www.biorxiv.org/content/10.1101/322347v1" /> <link rel="shortlink" 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id="zone-content" class="zone zone-content clearfix container-30"> <div class="grid-28 suffix-1 prefix-1 region region-content" id="region-content"> <div class="region-inner region-content-inner"> <a id="main-content"></a> <div class="block block-system block-main block-system-main odd block-without-title" id="block-system-main"> <div class="block-inner clearfix"> <div class="content clearfix"> <div class="panel-display panels-960-layout jcore-2col-layout" > <div class="panel-row-wrapper clearfix"> <div class="main-content-wrapper grid-17 suffix-1 alpha"> <div class="panel-panel panel-region-content"> <div class="inside"><div class="panel-pane pane-highwire-article-citation" > <div class="pane-content"> <div class="highwire-article-citation highwire-citation-type-highwire-article node98922" data-node-nid="98922" id="node-98922--21510241855" data-pisa="biorxiv;322347v1" data-pisa-master="biorxiv;322347" data-apath="/biorxiv/early/2018/05/16/322347.atom" data-hw-author-tooltip-instance="highwire_author_tooltip"><div class="highwire-cite highwire-cite-highwire-article highwire-citation-biorxiv-article-top clearfix has-author-tooltip" > <span class="biorxiv-article-type"> New Results </span> <h1 class="highwire-cite-title" id="page-title">The genetic prehistory of the Greater Caucasus</h1> <div class="highwire-cite-authors" ><span class="highwire-citation-authors"><span class="highwire-citation-author first" data-delta="0"><span class="nlm-given-names">Chuan-Chao</span> <span class="nlm-surname">Wang</span></span>, <span class="highwire-citation-author" data-delta="1"><span class="nlm-given-names">Sabine</span> <span class="nlm-surname">Reinhold</span></span>, <span class="highwire-citation-author" data-delta="2"><span class="nlm-given-names">Alexey</span> <span class="nlm-surname">Kalmykov</span></span>, <span class="highwire-citation-author" data-delta="3"><span class="nlm-given-names">Antje</span> <span class="nlm-surname">Wissgott</span></span>, <span class="highwire-citation-author" data-delta="4"><span class="nlm-given-names">Guido</span> <span class="nlm-surname">Brandt</span></span>, <span class="highwire-citation-author" data-delta="5"><span class="nlm-given-names">Choongwon</span> <span class="nlm-surname">Jeong</span></span>, <span class="highwire-citation-author" data-delta="6"><span class="nlm-given-names">Olivia</span> <span class="nlm-surname">Cheronet</span></span>, <span class="highwire-citation-author" data-delta="7"><span class="nlm-given-names">Matthew</span> <span class="nlm-surname">Ferry</span></span>, <span class="highwire-citation-author" data-delta="8"><span class="nlm-given-names">Eadaoin</span> <span class="nlm-surname">Harney</span></span>, <span class="highwire-citation-author" data-delta="9"><span class="nlm-given-names">Denise</span> <span class="nlm-surname">Keating</span></span>, <span class="highwire-citation-author" data-delta="10"><span class="nlm-given-names">Swapan</span> <span class="nlm-surname">Mallick</span></span>, <span class="highwire-citation-author" data-delta="11"><span class="nlm-given-names">Nadin</span> <span class="nlm-surname">Rohland</span></span>, <span class="highwire-citation-author" data-delta="12"><span class="nlm-given-names">Kristin</span> <span class="nlm-surname">Stewardson</span></span>, <span class="highwire-citation-author" data-delta="13"><span class="nlm-given-names">Anatoly R.</span> <span class="nlm-surname">Kantorovich</span></span>, <span class="highwire-citation-author" data-delta="14"><span class="nlm-given-names">Vladimir E.</span> <span class="nlm-surname">Maslov</span></span>, <span class="highwire-citation-author" data-delta="15"><span class="nlm-given-names">Vladimira G.</span> <span class="nlm-surname">Petrenko</span></span>, <span class="highwire-citation-author" data-delta="16"><span class="nlm-given-names">Vladimir R.</span> <span class="nlm-surname">Erlikh</span></span>, <span class="highwire-citation-author" data-delta="17"><span class="nlm-given-names">Biaslan Ch.</span> <span class="nlm-surname">Atabiev</span></span>, <span class="highwire-citation-author" data-delta="18"><span class="nlm-given-names">Rabadan G.</span> <span class="nlm-surname">Magomedov</span></span>, <span class="highwire-citation-author" data-delta="19"><span class="nlm-given-names">Philipp L.</span> <span class="nlm-surname">Kohl</span></span>, <span class="highwire-citation-author" data-delta="20"><span class="nlm-given-names">Kurt W.</span> <span class="nlm-surname">Alt</span></span>, <span class="highwire-citation-author" data-delta="21"><span class="nlm-given-names">Sandra L.</span> <span class="nlm-surname">Pichler</span></span>, <span class="highwire-citation-author" data-delta="22"><span class="nlm-given-names">Claudia</span> <span class="nlm-surname">Gerling</span></span>, <span class="highwire-citation-author" data-delta="23"><span class="nlm-given-names">Harald</span> <span class="nlm-surname">Meller</span></span>, <span class="highwire-citation-author" data-delta="24"><span class="nlm-given-names">Benik</span> <span class="nlm-surname">Vardanyan</span></span>, <span class="highwire-citation-author" data-delta="25"><span class="nlm-given-names">Larisa</span> <span class="nlm-surname">Yeganyan</span></span>, <span class="highwire-citation-author" data-delta="26"><span class="nlm-given-names">Alexey D.</span> <span class="nlm-surname">Rezepkin</span></span>, <span class="highwire-citation-author" data-delta="27"><span class="nlm-given-names">Dirk</span> <span class="nlm-surname">Mariaschk</span></span>, <span class="highwire-citation-author" data-delta="28"><span class="nlm-given-names">Natalia</span> <span class="nlm-surname">Berezina</span></span>, <span class="highwire-citation-author" data-delta="29"><span class="nlm-given-names">Julia</span> <span class="nlm-surname">Gresky</span></span>, <span class="highwire-citation-author" data-delta="30"><span class="nlm-given-names">Katharina</span> <span class="nlm-surname">Fuchs</span></span>, <span class="highwire-citation-author" data-delta="31"><span class="nlm-given-names">Corina</span> <span class="nlm-surname">Knipper</span></span>, <span class="highwire-citation-author" data-delta="32"><span class="nlm-given-names">Stephan</span> <span class="nlm-surname">Schiffels</span></span>, <span class="highwire-citation-author" data-delta="33"><span class="nlm-given-names">Elena</span> <span class="nlm-surname">Balanovska</span></span>, <span class="highwire-citation-author" data-delta="34"><span class="nlm-given-names">Oleg</span> <span class="nlm-surname">Balanovsky</span></span>, <span class="highwire-citation-author" data-delta="35"><span class="nlm-given-names">Iain</span> <span class="nlm-surname">Mathieson</span></span>, <span class="highwire-citation-author" data-delta="36"><span class="nlm-given-names">Thomas</span> <span class="nlm-surname">Higham</span></span>, <span class="highwire-citation-author" data-delta="37"><span class="nlm-given-names">Yakov B.</span> <span class="nlm-surname">Berezin</span></span>, <span class="highwire-citation-author" data-delta="38"><span class="nlm-given-names">Alexandra</span> <span class="nlm-surname">Buzhilova</span></span>, <span class="highwire-citation-author" data-delta="39"><span class="nlm-given-names">Viktor</span> <span class="nlm-surname">Trifonov</span></span>, <span class="highwire-citation-author" data-delta="40"><span class="nlm-given-names">Ron</span> <span class="nlm-surname">Pinhasi</span></span>, <span class="highwire-citation-author" data-delta="41"><span class="nlm-given-names">Andrej B.</span> <span class="nlm-surname">Belinskiy</span></span>, <span class="highwire-citation-author" data-delta="42"><span class="nlm-given-names">David</span> <span class="nlm-surname">Reich</span></span>, <span class="highwire-citation-author" data-delta="43"><span class="nlm-given-names">Svend</span> <span class="nlm-surname">Hansen</span></span>, <span class="highwire-citation-author" data-delta="44"><span class="nlm-given-names">Johannes</span> <span class="nlm-surname">Krause</span></span>, <span class="highwire-citation-author" data-delta="45"><span class="nlm-given-names">Wolfgang</span> <span class="nlm-surname">Haak</span></span></span></div> <div class="highwire-cite-metadata" ><span class="highwire-cite-metadata-doi highwire-cite-metadata"><span class="label">doi:</span> https://doi.org/10.1101/322347 </span></div> </div> <div id="hw-article-author-popups-node-98922--21510241855" style="display: none;"><div class="author-tooltip-0"><div class="author-tooltip-name">Chuan-Chao Wang </div><div class="author-tooltip-affiliation"><span class="author-tooltip-text"><div class='author-affiliation'><span class='nlm-sup'>1</span><span class='nlm-institution'>Department of Archaeogenetics, Max-Planck Institute for the Science of Human History</span>, Kahlaische Strasse 10, D-07745 Jena, <span class='nlm-country'>Germany</span></div><div class='author-affiliation'><span class='nlm-sup'>2</span><span class='nlm-institution'>Department of Anthropology and Ethnology, Xiamen University 361005 Xiamen</span>, <span class='nlm-country'>China</span></div></span></div><ul class="author-tooltip-find-more"><li class="author-tooltip-gs-link first"><a href="/lookup/google-scholar?link_type=googlescholar&amp;gs_type=author&amp;author%5B0%5D=Chuan-Chao%2BWang%2B" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on Google Scholar</a></li><li class="author-tooltip-pubmed-link"><a href="/lookup/external-ref?access_num=Wang%20C&amp;link_type=AUTHORSEARCH" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on PubMed</a></li><li class="author-site-search-link"><a href="/search/author1%3AChuan-Chao%2BWang%2B" rel="nofollow" class="" data-icon-position="" data-hide-link-title="0">Search for this author on this site</a></li><li class="author-corresp-email-link last"><span>For correspondence: <a href="/cdn-cgi/l/email-protection#e68e87878da6958e8ec88b9681c88283" class="" data-icon-position="" data-hide-link-title="0"><span class="__cf_email__" data-cfemail="9bf3fafaf0dbe8f3f3b5f6ebfcb5fffe">[email&#160;protected]</span></a> <a href="/cdn-cgi/l/email-protection#5d362f3c282e381d2e353573302d3a733938" class="" data-icon-position="" data-hide-link-title="0"><span class="__cf_email__" data-cfemail="fe958c9f8b8d9bbe8d9696d0938e99d09a9b">[email&#160;protected]</span></a> <a href="/cdn-cgi/l/email-protection#2552444b42655d48500b4041500b464b" class="" data-icon-position="" data-hide-link-title="0"><span class="__cf_email__" data-cfemail="0176606f6641796c742f6465742f626f">[email&#160;protected]</span></a> <a href="/cdn-cgi/l/email-protection#deada8bbb0baf0b6bfb0adbbb09ebabfb7b0adaaf0babb" class="" data-icon-position="" data-hide-link-title="0"><span class="__cf_email__" data-cfemail="6e1d180b000a40060f001d0b002e0a0f07001d1a400a0b">[email&#160;protected]</span></a></span></li></ul></div><div class="author-tooltip-1"><div class="author-tooltip-name">Sabine Reinhold </div><div class="author-tooltip-affiliation"><span class="author-tooltip-text"><div class='author-affiliation'><span class='nlm-sup'>3</span><span class='nlm-institution'>German Archaeological Institute, Eurasia Department</span>, Im Dol 2-6, D-14195 Berlin, <span class='nlm-country'>Germany</span></div></span></div><ul class="author-tooltip-find-more"><li class="author-tooltip-gs-link first"><a href="/lookup/google-scholar?link_type=googlescholar&amp;gs_type=author&amp;author%5B0%5D=Sabine%2BReinhold%2B" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on Google Scholar</a></li><li class="author-tooltip-pubmed-link"><a href="/lookup/external-ref?access_num=Reinhold%20S&amp;link_type=AUTHORSEARCH" target="_blank" class="" 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History</span>, Kahlaische Strasse 10, D-07745 Jena, <span class='nlm-country'>Germany</span></div></span></div><ul class="author-tooltip-find-more"><li class="author-tooltip-gs-link first"><a href="/lookup/google-scholar?link_type=googlescholar&amp;gs_type=author&amp;author%5B0%5D=Guido%2BBrandt%2B" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on Google Scholar</a></li><li class="author-tooltip-pubmed-link"><a href="/lookup/external-ref?access_num=Brandt%20G&amp;link_type=AUTHORSEARCH" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on PubMed</a></li><li class="author-site-search-link last"><a href="/search/author1%3AGuido%2BBrandt%2B" rel="nofollow" class="" data-icon-position="" data-hide-link-title="0">Search for this author on this site</a></li></ul></div><div class="author-tooltip-5"><div class="author-tooltip-name">Choongwon Jeong </div><div class="author-tooltip-affiliation"><span class="author-tooltip-text"><div class='author-affiliation'><span class='nlm-sup'>1</span><span class='nlm-institution'>Department of Archaeogenetics, Max-Planck Institute for the Science of Human History</span>, Kahlaische Strasse 10, D-07745 Jena, <span class='nlm-country'>Germany</span></div></span></div><ul class="author-tooltip-find-more"><li class="author-tooltip-gs-link first"><a href="/lookup/google-scholar?link_type=googlescholar&amp;gs_type=author&amp;author%5B0%5D=Choongwon%2BJeong%2B" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on Google Scholar</a></li><li class="author-tooltip-pubmed-link"><a href="/lookup/external-ref?access_num=Jeong%20C&amp;link_type=AUTHORSEARCH" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on PubMed</a></li><li class="author-site-search-link last"><a href="/search/author1%3AChoongwon%2BJeong%2B" rel="nofollow" class="" data-icon-position="" 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href="/lookup/google-scholar?link_type=googlescholar&amp;gs_type=author&amp;author%5B0%5D=Olivia%2BCheronet%2B" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on Google Scholar</a></li><li class="author-tooltip-pubmed-link"><a href="/lookup/external-ref?access_num=Cheronet%20O&amp;link_type=AUTHORSEARCH" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on PubMed</a></li><li class="author-site-search-link last"><a href="/search/author1%3AOlivia%2BCheronet%2B" rel="nofollow" class="" data-icon-position="" data-hide-link-title="0">Search for this author on this site</a></li></ul></div><div class="author-tooltip-7"><div class="author-tooltip-name">Matthew Ferry </div><div class="author-tooltip-affiliation"><span class="author-tooltip-text"><div class='author-affiliation'><span class='nlm-sup'>8</span><span class='nlm-institution'>Department of Genetics, Harvard Medical School</span>, Boston 02115 MA, <span class='nlm-country'>USA</span></div><div class='author-affiliation'><span class='nlm-sup'>9</span><span class='nlm-institution'>Howard Hughes Medical Institute, Harvard Medical School</span>, Boston 02115 MA, <span class='nlm-country'>USA</span></div></span></div><ul class="author-tooltip-find-more"><li class="author-tooltip-gs-link first"><a href="/lookup/google-scholar?link_type=googlescholar&amp;gs_type=author&amp;author%5B0%5D=Matthew%2BFerry%2B" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on Google Scholar</a></li><li class="author-tooltip-pubmed-link"><a href="/lookup/external-ref?access_num=Ferry%20M&amp;link_type=AUTHORSEARCH" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on PubMed</a></li><li class="author-site-search-link last"><a href="/search/author1%3AMatthew%2BFerry%2B" rel="nofollow" class="" data-icon-position="" data-hide-link-title="0">Search for this author on this 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href="/lookup/google-scholar?link_type=googlescholar&amp;gs_type=author&amp;author%5B0%5D=Eadaoin%2BHarney%2B" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on Google Scholar</a></li><li class="author-tooltip-pubmed-link"><a href="/lookup/external-ref?access_num=Harney%20E&amp;link_type=AUTHORSEARCH" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on PubMed</a></li><li class="author-site-search-link last"><a href="/search/author1%3AEadaoin%2BHarney%2B" rel="nofollow" class="" data-icon-position="" data-hide-link-title="0">Search for this author on this site</a></li></ul></div><div class="author-tooltip-9"><div class="author-tooltip-name">Denise Keating </div><div class="author-tooltip-affiliation"><span class="author-tooltip-text"><div class='author-affiliation'><span class='nlm-sup'>5</span><span class='nlm-institution'>Earth Institute, University College Dublin</span>,Dublin 4, <span class='nlm-country'>Ireland</span></div><div class='author-affiliation'><span class='nlm-sup'>7</span><span class='nlm-institution'>School of Archaeology, University College Dublin</span>, Dublin 4, <span class='nlm-country'>Ireland</span></div></span></div><ul class="author-tooltip-find-more"><li class="author-tooltip-gs-link first"><a href="/lookup/google-scholar?link_type=googlescholar&amp;gs_type=author&amp;author%5B0%5D=Denise%2BKeating%2B" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on Google Scholar</a></li><li class="author-tooltip-pubmed-link"><a href="/lookup/external-ref?access_num=Keating%20D&amp;link_type=AUTHORSEARCH" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on PubMed</a></li><li class="author-site-search-link last"><a href="/search/author1%3ADenise%2BKeating%2B" rel="nofollow" class="" data-icon-position="" data-hide-link-title="0">Search for this author on this 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class='nlm-country'>USA</span></div><div class='author-affiliation'><span class='nlm-sup'>11</span><span class='nlm-institution'>Broad Institute of Harvard and MIT, Cambridge 02142 MA</span>,<span class='nlm-country'>USA</span></div></span></div><ul class="author-tooltip-find-more"><li class="author-tooltip-gs-link first"><a href="/lookup/google-scholar?link_type=googlescholar&amp;gs_type=author&amp;author%5B0%5D=Nadin%2BRohland%2B" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on Google Scholar</a></li><li class="author-tooltip-pubmed-link"><a href="/lookup/external-ref?access_num=Rohland%20N&amp;link_type=AUTHORSEARCH" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on PubMed</a></li><li class="author-site-search-link last"><a href="/search/author1%3ANadin%2BRohland%2B" rel="nofollow" class="" data-icon-position="" data-hide-link-title="0">Search for this author on this site</a></li></ul></div><div 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class="author-tooltip-text"><div class='author-affiliation'><span class='nlm-sup'>3</span><span class='nlm-institution'>German Archaeological Institute, Eurasia Department</span>, Im Dol 2-6, D-14195 Berlin, <span class='nlm-country'>Germany</span></div></span></div><ul class="author-tooltip-find-more"><li class="author-tooltip-gs-link first"><a href="/lookup/google-scholar?link_type=googlescholar&amp;gs_type=author&amp;author%5B0%5D=Svend%2BHansen%2B" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on Google Scholar</a></li><li class="author-tooltip-pubmed-link"><a href="/lookup/external-ref?access_num=Hansen%20S&amp;link_type=AUTHORSEARCH" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on PubMed</a></li><li class="author-site-search-link"><a href="/search/author1%3ASvend%2BHansen%2B" rel="nofollow" class="" data-icon-position="" data-hide-link-title="0">Search for this author on this site</a></li><li class="author-corresp-email-link last"><span>For correspondence: <a href="/cdn-cgi/l/email-protection#85ede4e4eec5f6ededabe8f5e2abe1e0" class="" data-icon-position="" data-hide-link-title="0"><span class="__cf_email__" data-cfemail="8ce4edede7ccffe4e4a2e1fceba2e8e9">[email&#160;protected]</span></a> <a href="/cdn-cgi/l/email-protection#5e352c3f2b2d3b1e2d363670332e39703a3b" class="" data-icon-position="" data-hide-link-title="0"><span class="__cf_email__" data-cfemail="a6cdd4c7d3d5c3e6d5cece88cbd6c188c2c3">[email&#160;protected]</span></a> <a href="/cdn-cgi/l/email-protection#6215030c05221a0f174c0706174c010c" class="" data-icon-position="" data-hide-link-title="0"><span class="__cf_email__" data-cfemail="3b4c5a555c7b43564e155e5f4e155855">[email&#160;protected]</span></a> <a href="/cdn-cgi/l/email-protection#cbb8bdaea5afe5a3aaa5b8aea58bafaaa2a5b8bfe5afae" class="" data-icon-position="" data-hide-link-title="0"><span class="__cf_email__" 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class='nlm-institution'>Department of Archaeogenetics, Max-Planck Institute for the Science of Human History</span>, Kahlaische Strasse 10, D-07745 Jena, <span class='nlm-country'>Germany</span></div><div class='author-affiliation'><span class='nlm-sup'>36</span><span class='nlm-institution'>School of Biological Sciences, The University of Adelaide</span>, Adelaide 5005, <span class='nlm-country'>Australia</span></div></span></div><ul class="author-tooltip-find-more"><li class="author-tooltip-gs-link first"><a href="/lookup/google-scholar?link_type=googlescholar&amp;gs_type=author&amp;author%5B0%5D=Wolfgang%2BHaak%2B" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on Google Scholar</a></li><li class="author-tooltip-pubmed-link"><a href="/lookup/external-ref?access_num=Haak%20W&amp;link_type=AUTHORSEARCH" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on PubMed</a></li><li 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However, it remains unclear when and where this ancestry arose and whether it was related to a horizon of cultural innovations in the 4<sup>th</sup> millennium BCE that subsequently facilitated the advance of pastoral societies likely linked to the dispersal of Indo-European languages. To address this, we generated genome-wide SNP data from 45 prehistoric individuals along a 3000-year temporal transect in the North Caucasus. We observe a genetic separation between the groups of the Caucasus and those of the adjacent steppe. The Caucasus groups are genetically similar to contemporaneous populations south of it, suggesting that – unlike today – the Caucasus acted as a bridge rather than an insurmountable barrier to human movement. The steppe groups from Yamnaya and subsequent pastoralist cultures show evidence for previously undetected farmer-related ancestry from different contact zones, while Steppe Maykop individuals harbour additional Upper Palaeolithic Siberian and Native American related ancestry.</p></div><span class="highwire-journal-article-marker-end"></span></div><span class="related-urls"></span></div></div> </div> </div> <div class="panel-separator"></div><div class="panel-pane pane-biorxiv-copyright" > <div class="pane-content"> <div class="field field-name-field-highwire-copyright field-type-text field-label-inline clearfix"><div class="field-label">Copyright&nbsp;</div><div class="field-items"><div class="field-item even">The copyright holder for this preprint is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity.<span class="license-type"> It is made available under a <a href="http://creativecommons.org/licenses/by-nc-nd/4.0/" class="" 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<div class="inside"><div class="panel-pane pane-highwire-article-citation" > <div class="pane-content"> <div class="highwire-article-citation highwire-citation-type-highwire-article node98922--3" data-node-nid="98922" id="node-98922--4265447575" data-pisa="biorxiv;322347v1" data-pisa-master="biorxiv;322347" data-seqnum="98922" data-apath="/biorxiv/early/2018/05/16/322347.atom"><div class="highwire-cite highwire-cite-highwire-article highwire-citation-biorxiv-article-pap-list clearfix" > <div class="highwire-cite-title" > <div class="highwire-cite-title">The genetic prehistory of the Greater Caucasus</div> </div> <div class="highwire-cite-authors" ><span class="highwire-citation-authors"><span class="highwire-citation-author first" data-delta="0"><span class="nlm-given-names">Chuan-Chao</span> <span class="nlm-surname">Wang</span></span>, <span class="highwire-citation-author" data-delta="1"><span class="nlm-given-names">Sabine</span> <span class="nlm-surname">Reinhold</span></span>, 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class="highwire-citation-author" data-delta="35"><span class="nlm-given-names">Iain</span> <span class="nlm-surname">Mathieson</span></span>, <span class="highwire-citation-author" data-delta="36"><span class="nlm-given-names">Thomas</span> <span class="nlm-surname">Higham</span></span>, <span class="highwire-citation-author" data-delta="37"><span class="nlm-given-names">Yakov B.</span> <span class="nlm-surname">Berezin</span></span>, <span class="highwire-citation-author" data-delta="38"><span class="nlm-given-names">Alexandra</span> <span class="nlm-surname">Buzhilova</span></span>, <span class="highwire-citation-author" data-delta="39"><span class="nlm-given-names">Viktor</span> <span class="nlm-surname">Trifonov</span></span>, <span class="highwire-citation-author" data-delta="40"><span class="nlm-given-names">Ron</span> <span class="nlm-surname">Pinhasi</span></span>, <span class="highwire-citation-author" data-delta="41"><span class="nlm-given-names">Andrej B.</span> <span 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class="panel-pane pane-minipanel-dialog-link pane-biorxiv-cite-tool" > <div class="pane-content"> <div class='minipanel-dialog-wrapper'><div class='minipanel-dialog-link-link'><a href="/" oncontextmenu="javascript: return false;" class="minipanel-dialog-link-trigger link-icon" title="Citation Tools"><span class="icon-globe"></span> <span class="title">Citation Tools</span></a></div><div class='minipanel-dialog-link-mini' style='display:none'><div class="panel-display panel-1col clearfix" id="mini-panel-biorxiv_cite_tool"> <div class="panel-panel panel-col"> <div><div class="panel-pane pane-highwire-citation-export" > <div class="pane-content"> <div class="highwire-citation-export"> <div class="highwire-citation-info"> <div class="highwire-article-citation highwire-citation-type-highwire-article cite-tool-node98922--3" data-node-nid="98922" id="citation-node-98922--4648975520" data-pisa="biorxiv;322347v1" data-pisa-master="biorxiv;322347" data-seqnum="98922" data-apath="/biorxiv/early/2018/05/16/322347.atom"><div class="highwire-cite highwire-cite-highwire-article highwire-citation-biorxiv-article-pap-list clearfix" > <div class="highwire-cite-title" > <div class="highwire-cite-title">The genetic prehistory of the Greater Caucasus</div> </div> <div class="highwire-cite-authors" ><span class="highwire-citation-authors"><span class="highwire-citation-author first" data-delta="0"><span class="nlm-given-names">Chuan-Chao</span> <span class="nlm-surname">Wang</span></span>, <span class="highwire-citation-author" data-delta="1"><span class="nlm-given-names">Sabine</span> <span class="nlm-surname">Reinhold</span></span>, <span class="highwire-citation-author" data-delta="2"><span class="nlm-given-names">Alexey</span> <span class="nlm-surname">Kalmykov</span></span>, <span class="highwire-citation-author" data-delta="3"><span class="nlm-given-names">Antje</span> <span class="nlm-surname">Wissgott</span></span>, <span class="highwire-citation-author" data-delta="4"><span class="nlm-given-names">Guido</span> <span class="nlm-surname">Brandt</span></span>, <span class="highwire-citation-author" data-delta="5"><span class="nlm-given-names">Choongwon</span> <span class="nlm-surname">Jeong</span></span>, <span class="highwire-citation-author" data-delta="6"><span class="nlm-given-names">Olivia</span> <span class="nlm-surname">Cheronet</span></span>, <span class="highwire-citation-author" data-delta="7"><span class="nlm-given-names">Matthew</span> <span class="nlm-surname">Ferry</span></span>, <span class="highwire-citation-author" data-delta="8"><span class="nlm-given-names">Eadaoin</span> <span class="nlm-surname">Harney</span></span>, <span class="highwire-citation-author" data-delta="9"><span class="nlm-given-names">Denise</span> <span class="nlm-surname">Keating</span></span>, <span class="highwire-citation-author" data-delta="10"><span class="nlm-given-names">Swapan</span> <span class="nlm-surname">Mallick</span></span>, <span class="highwire-citation-author" data-delta="11"><span class="nlm-given-names">Nadin</span> <span class="nlm-surname">Rohland</span></span>, <span class="highwire-citation-author" data-delta="12"><span class="nlm-given-names">Kristin</span> <span class="nlm-surname">Stewardson</span></span>, <span class="highwire-citation-author" data-delta="13"><span class="nlm-given-names">Anatoly R.</span> <span class="nlm-surname">Kantorovich</span></span>, <span class="highwire-citation-author" data-delta="14"><span class="nlm-given-names">Vladimir E.</span> <span class="nlm-surname">Maslov</span></span>, <span class="highwire-citation-author" data-delta="15"><span class="nlm-given-names">Vladimira G.</span> <span class="nlm-surname">Petrenko</span></span>, <span class="highwire-citation-author" data-delta="16"><span class="nlm-given-names">Vladimir R.</span> <span class="nlm-surname">Erlikh</span></span>, <span class="highwire-citation-author" data-delta="17"><span class="nlm-given-names">Biaslan Ch.</span> <span class="nlm-surname">Atabiev</span></span>, <span class="highwire-citation-author" data-delta="18"><span class="nlm-given-names">Rabadan G.</span> <span class="nlm-surname">Magomedov</span></span>, <span class="highwire-citation-author" data-delta="19"><span class="nlm-given-names">Philipp L.</span> <span class="nlm-surname">Kohl</span></span>, <span class="highwire-citation-author" data-delta="20"><span class="nlm-given-names">Kurt W.</span> <span class="nlm-surname">Alt</span></span>, <span class="highwire-citation-author" data-delta="21"><span class="nlm-given-names">Sandra L.</span> <span class="nlm-surname">Pichler</span></span>, <span class="highwire-citation-author" data-delta="22"><span class="nlm-given-names">Claudia</span> <span class="nlm-surname">Gerling</span></span>, <span class="highwire-citation-author" data-delta="23"><span class="nlm-given-names">Harald</span> <span class="nlm-surname">Meller</span></span>, <span class="highwire-citation-author" data-delta="24"><span class="nlm-given-names">Benik</span> <span class="nlm-surname">Vardanyan</span></span>, <span class="highwire-citation-author" data-delta="25"><span class="nlm-given-names">Larisa</span> <span class="nlm-surname">Yeganyan</span></span>, <span class="highwire-citation-author" data-delta="26"><span class="nlm-given-names">Alexey D.</span> <span class="nlm-surname">Rezepkin</span></span>, <span class="highwire-citation-author" data-delta="27"><span class="nlm-given-names">Dirk</span> <span class="nlm-surname">Mariaschk</span></span>, <span class="highwire-citation-author" data-delta="28"><span class="nlm-given-names">Natalia</span> <span class="nlm-surname">Berezina</span></span>, <span class="highwire-citation-author" data-delta="29"><span class="nlm-given-names">Julia</span> <span class="nlm-surname">Gresky</span></span>, <span class="highwire-citation-author" data-delta="30"><span 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class="highwire-citation-author" data-delta="37"><span class="nlm-given-names">Yakov B.</span> <span class="nlm-surname">Berezin</span></span>, <span class="highwire-citation-author" data-delta="38"><span class="nlm-given-names">Alexandra</span> <span class="nlm-surname">Buzhilova</span></span>, <span class="highwire-citation-author" data-delta="39"><span class="nlm-given-names">Viktor</span> <span class="nlm-surname">Trifonov</span></span>, <span class="highwire-citation-author" data-delta="40"><span class="nlm-given-names">Ron</span> <span class="nlm-surname">Pinhasi</span></span>, <span class="highwire-citation-author" data-delta="41"><span class="nlm-given-names">Andrej B.</span> <span class="nlm-surname">Belinskiy</span></span>, <span class="highwire-citation-author" data-delta="42"><span class="nlm-given-names">David</span> <span class="nlm-surname">Reich</span></span>, <span class="highwire-citation-author" data-delta="43"><span class="nlm-given-names">Svend</span> <span 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