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<!DOCTYPE html> <html lang="en" class="no-js"> <head> <meta charset="UTF-8"> <meta http-equiv="X-UA-Compatible" content="IE=edge"> <meta name="applicable-device" content="pc,mobile"> <meta name="viewport" content="width=device-width, initial-scale=1"> <title>Genes reveal traces of common recent demographic history for most of the Uralic-speaking populations | Genome Biology | Full Text</title> <meta name="citation_abstract" content="The genetic origins of Uralic speakers from across a vast territory in the temperate zone of North Eurasia have remained elusive. Previous studies have shown contrasting proportions of Eastern and Western Eurasian ancestry in their mitochondrial and Y chromosomal gene pools. While the maternal lineages reflect by and large the geographic background of a given Uralic-speaking population, the frequency of Y chromosomes of Eastern Eurasian origin is distinctively high among European Uralic speakers. The autosomal variation of Uralic speakers, however, has not yet been studied comprehensively. Here, we present a genome-wide analysis of 15 Uralic-speaking populations which cover all main groups of the linguistic family. We show that contemporary Uralic speakers are genetically very similar to their local geographical neighbours. However, when studying relationships among geographically distant populations, we find that most of the Uralic speakers and some of their neighbours share a genetic component of possibly Siberian origin. Additionally, we show that most Uralic speakers share significantly more genomic segments identity-by-descent with each other than with geographically equidistant speakers of other languages. We find that correlated genome-wide genetic and lexical distances among Uralic speakers suggest co-dispersion of genes and languages. Yet, we do not find long-range genetic ties between Estonians and Hungarians with their linguistic sisters that would distinguish them from their non-Uralic-speaking neighbours. We show that most Uralic speakers share a distinct ancestry component of likely Siberian origin, which suggests that the spread of Uralic languages involved at least some demic component."/> <meta name="journal_id" content="13059"/> <meta name="dc.title" content="Genes reveal traces of common recent demographic history for most of the Uralic-speaking populations"/> <meta name="dc.source" content="Genome Biology 2018 19:1"/> <meta name="dc.format" content="text/html"/> <meta name="dc.publisher" content="BioMed Central"/> <meta name="dc.date" content="2018-09-21"/> <meta name="dc.type" content="OriginalPaper"/> <meta name="dc.language" content="En"/> <meta name="dc.copyright" content="2018 The Author(s)."/> <meta name="dc.rights" content="2018 The Author(s)."/> <meta name="dc.rightsAgent" content="reprints@biomedcentral.com"/> <meta name="dc.description" content="The genetic origins of Uralic speakers from across a vast territory in the temperate zone of North Eurasia have remained elusive. Previous studies have shown contrasting proportions of Eastern and Western Eurasian ancestry in their mitochondrial and Y chromosomal gene pools. While the maternal lineages reflect by and large the geographic background of a given Uralic-speaking population, the frequency of Y chromosomes of Eastern Eurasian origin is distinctively high among European Uralic speakers. The autosomal variation of Uralic speakers, however, has not yet been studied comprehensively. Here, we present a genome-wide analysis of 15 Uralic-speaking populations which cover all main groups of the linguistic family. We show that contemporary Uralic speakers are genetically very similar to their local geographical neighbours. However, when studying relationships among geographically distant populations, we find that most of the Uralic speakers and some of their neighbours share a genetic component of possibly Siberian origin. Additionally, we show that most Uralic speakers share significantly more genomic segments identity-by-descent with each other than with geographically equidistant speakers of other languages. We find that correlated genome-wide genetic and lexical distances among Uralic speakers suggest co-dispersion of genes and languages. Yet, we do not find long-range genetic ties between Estonians and Hungarians with their linguistic sisters that would distinguish them from their non-Uralic-speaking neighbours. We show that most Uralic speakers share a distinct ancestry component of likely Siberian origin, which suggests that the spread of Uralic languages involved at least some demic component."/> <meta name="prism.issn" content="1474-760X"/> <meta name="prism.publicationName" content="Genome Biology"/> <meta name="prism.publicationDate" content="2018-09-21"/> <meta name="prism.volume" content="19"/> <meta name="prism.number" content="1"/> <meta name="prism.section" content="OriginalPaper"/> <meta name="prism.startingPage" content="1"/> <meta name="prism.endingPage" content="20"/> <meta name="prism.copyright" content="2018 The Author(s)."/> <meta name="prism.rightsAgent" content="reprints@biomedcentral.com"/> <meta name="prism.url" content="https://genomebiology.biomedcentral.com/articles/10.1186/s13059-018-1522-1"/> <meta name="prism.doi" content="doi:10.1186/s13059-018-1522-1"/> <meta name="citation_pdf_url" content="https://genomebiology.biomedcentral.com/counter/pdf/10.1186/s13059-018-1522-1"/> <meta name="citation_fulltext_html_url" content="https://genomebiology.biomedcentral.com/articles/10.1186/s13059-018-1522-1"/> <meta name="citation_journal_title" content="Genome Biology"/> <meta name="citation_journal_abbrev" content="Genome Biol"/> <meta name="citation_publisher" content="BioMed Central"/> <meta name="citation_issn" content="1474-760X"/> <meta name="citation_title" content="Genes reveal traces of common recent demographic history for most of the Uralic-speaking populations"/> <meta name="citation_volume" content="19"/> <meta name="citation_issue" content="1"/> <meta name="citation_publication_date" content="2018/12"/> <meta name="citation_online_date" content="2018/09/21"/> <meta name="citation_firstpage" content="1"/> <meta name="citation_lastpage" content="20"/> <meta name="citation_article_type" content="Research"/> <meta name="citation_fulltext_world_readable" content=""/> <meta name="citation_language" content="en"/> <meta name="dc.identifier" content="doi:10.1186/s13059-018-1522-1"/> <meta name="DOI" content="10.1186/s13059-018-1522-1"/> <meta name="size" content="331876"/> <meta name="citation_doi" content="10.1186/s13059-018-1522-1"/> <meta name="citation_springer_api_url" content="http://api.springer.com/xmldata/jats?q=doi:10.1186/s13059-018-1522-1&api_key="/> <meta name="description" content="The genetic origins of Uralic speakers from across a vast territory in the temperate zone of North Eurasia have remained elusive. Previous studies have shown contrasting proportions of Eastern and Western Eurasian ancestry in their mitochondrial and Y chromosomal gene pools. While the maternal lineages reflect by and large the geographic background of a given Uralic-speaking population, the frequency of Y chromosomes of Eastern Eurasian origin is distinctively high among European Uralic speakers. The autosomal variation of Uralic speakers, however, has not yet been studied comprehensively. Here, we present a genome-wide analysis of 15 Uralic-speaking populations which cover all main groups of the linguistic family. We show that contemporary Uralic speakers are genetically very similar to their local geographical neighbours. However, when studying relationships among geographically distant populations, we find that most of the Uralic speakers and some of their neighbours share a genetic component of possibly Siberian origin. Additionally, we show that most Uralic speakers share significantly more genomic segments identity-by-descent with each other than with geographically equidistant speakers of other languages. We find that correlated genome-wide genetic and lexical distances among Uralic speakers suggest co-dispersion of genes and languages. Yet, we do not find long-range genetic ties between Estonians and Hungarians with their linguistic sisters that would distinguish them from their non-Uralic-speaking neighbours. We show that most Uralic speakers share a distinct ancestry component of likely Siberian origin, which suggests that the spread of Uralic languages involved at least some demic component."/> <meta name="dc.creator" content="Tambets, Kristiina"/> <meta name="dc.creator" content="Yunusbayev, Bayazit"/> <meta name="dc.creator" content="Hudjashov, Georgi"/> <meta name="dc.creator" content="Ilumäe, Anne-Mai"/> <meta name="dc.creator" content="Rootsi, Siiri"/> <meta name="dc.creator" content="Honkola, Terhi"/> <meta name="dc.creator" content="Vesakoski, Outi"/> <meta name="dc.creator" content="Atkinson, Quentin"/> <meta name="dc.creator" content="Skoglund, Pontus"/> <meta name="dc.creator" content="Kushniarevich, Alena"/> <meta name="dc.creator" content="Litvinov, Sergey"/> <meta name="dc.creator" content="Reidla, Maere"/> <meta name="dc.creator" content="Metspalu, Ene"/> <meta name="dc.creator" content="Saag, Lehti"/> <meta name="dc.creator" content="Rantanen, Timo"/> <meta name="dc.creator" content="Karmin, Monika"/> <meta name="dc.creator" content="Parik, Jüri"/> <meta name="dc.creator" content="Zhadanov, Sergey I."/> <meta name="dc.creator" content="Gubina, Marina"/> <meta name="dc.creator" content="Damba, Larisa D."/> <meta name="dc.creator" content="Bermisheva, Marina"/> <meta name="dc.creator" content="Reisberg, Tuuli"/> <meta name="dc.creator" content="Dibirova, Khadizhat"/> <meta name="dc.creator" content="Evseeva, Irina"/> <meta name="dc.creator" content="Nelis, Mari"/> <meta name="dc.creator" content="Klovins, Janis"/> <meta name="dc.creator" content="Metspalu, Andres"/> <meta name="dc.creator" content="Esko, Tõnu"/> <meta name="dc.creator" content="Balanovsky, Oleg"/> <meta name="dc.creator" content="Balanovska, Elena"/> <meta name="dc.creator" content="Khusnutdinova, Elza K."/> <meta name="dc.creator" content="Osipova, Ludmila P."/> <meta name="dc.creator" content="Voevoda, Mikhail"/> <meta name="dc.creator" content="Villems, Richard"/> <meta name="dc.creator" content="Kivisild, Toomas"/> <meta name="dc.creator" content="Metspalu, Mait"/> <meta name="dc.subject" content="Animal Genetics and Genomics"/> <meta name="dc.subject" content="Human Genetics"/> <meta name="dc.subject" content="Plant Genetics and Genomics"/> <meta name="dc.subject" content="Microbial Genetics and Genomics"/> <meta name="dc.subject" content="Bioinformatics"/> <meta name="dc.subject" content="Evolutionary Biology"/> <meta name="citation_reference" content="citation_journal_title=PLoS Genet; citation_title=The genetic legacy of the expansion of Turkic-speaking nomads across Eurasia; citation_author=B Yunusbayev, M Metspalu, E Metspalu, A Valeev, S Litvinov; citation_volume=11; citation_publication_date=2015; citation_pages=e1005068; citation_doi=10.1371/journal.pgen.1005068; citation_id=CR1"/> <meta name="citation_reference" content="Haak W, Lazaridis I, Patterson N, Rohland N, Mallick S, et al. 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Previous studies have shown contrasting proportions of Eastern and Western Eurasian ancestry in their mitochondrial and Y chromosomal gene pools. While the maternal lineages reflect by and large the geographic background of a given Uralic-speaking population, the frequency of Y chromosomes of Eastern Eurasian origin is distinctively high among European Uralic speakers. The autosomal variation of Uralic speakers, however, has not yet been studied comprehensively. Results Here, we present a genome-wide analysis of 15 Uralic-speaking populations which cover all main groups of the linguistic family. We show that contemporary Uralic speakers are genetically very similar to their local geographical neighbours. However, when studying relationships among geographically distant populations, we find that most of the Uralic speakers and some of their neighbours share a genetic component of possibly Siberian origin. Additionally, we show that most Uralic speakers share significantly more genomic segments identity-by-descent with each other than with geographically equidistant speakers of other languages. We find that correlated genome-wide genetic and lexical distances among Uralic speakers suggest co-dispersion of genes and languages. Yet, we do not find long-range genetic ties between Estonians and Hungarians with their linguistic sisters that would distinguish them from their non-Uralic-speaking neighbours. Conclusions We show that most Uralic speakers share a distinct ancestry component of likely Siberian origin, which suggests that the spread of Uralic languages involved at least some demic component."/> <meta property="og:image" content="https://static-content.springer.com/image/art%3A10.1186%2Fs13059-018-1522-1/MediaObjects/13059_2018_1522_Fig1_HTML.png"/> <script type="application/ld+json">{"mainEntity":{"headline":"Genes reveal traces of common recent demographic history for most of the Uralic-speaking populations","description":"The genetic origins of Uralic speakers from across a vast territory in the temperate zone of North Eurasia have remained elusive. Previous studies have shown contrasting proportions of Eastern and Western Eurasian ancestry in their mitochondrial and Y chromosomal gene pools. While the maternal lineages reflect by and large the geographic background of a given Uralic-speaking population, the frequency of Y chromosomes of Eastern Eurasian origin is distinctively high among European Uralic speakers. The autosomal variation of Uralic speakers, however, has not yet been studied comprehensively. Here, we present a genome-wide analysis of 15 Uralic-speaking populations which cover all main groups of the linguistic family. We show that contemporary Uralic speakers are genetically very similar to their local geographical neighbours. However, when studying relationships among geographically distant populations, we find that most of the Uralic speakers and some of their neighbours share a genetic component of possibly Siberian origin. Additionally, we show that most Uralic speakers share significantly more genomic segments identity-by-descent with each other than with geographically equidistant speakers of other languages. 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class="c-article-title" data-test="article-title" data-article-title="">Genes reveal traces of common recent demographic history for most of the Uralic-speaking populations</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-Kristiina-Tambets-Aff1" data-author-popup="auth-Kristiina-Tambets-Aff1" data-author-search="Tambets, Kristiina" data-corresp-id="c1">Kristiina Tambets<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><span class="u-js-hide"> <a class="js-orcid" href="http://orcid.org/0000-0002-8173-6380"><span class="u-visually-hidden">ORCID: </span>orcid.org/0000-0002-8173-6380</a></span><sup class="u-js-hide"><a href="#Aff1">1</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-Bayazit-Yunusbayev-Aff1-Aff2" data-author-popup="auth-Bayazit-Yunusbayev-Aff1-Aff2" data-author-search="Yunusbayev, Bayazit">Bayazit Yunusbayev</a><sup class="u-js-hide"><a href="#Aff1">1</a>,<a href="#Aff2">2</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-Georgi-Hudjashov-Aff1-Aff3" data-author-popup="auth-Georgi-Hudjashov-Aff1-Aff3" data-author-search="Hudjashov, Georgi">Georgi Hudjashov</a><sup class="u-js-hide"><a href="#Aff1">1</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-Anne_Mai-Ilum_e-Aff1" data-author-popup="auth-Anne_Mai-Ilum_e-Aff1" data-author-search="Ilumäe, Anne-Mai">Anne-Mai Ilumäe</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-Siiri-Rootsi-Aff1" data-author-popup="auth-Siiri-Rootsi-Aff1" data-author-search="Rootsi, Siiri">Siiri Rootsi</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-Terhi-Honkola-Aff4-Aff5" data-author-popup="auth-Terhi-Honkola-Aff4-Aff5" data-author-search="Honkola, Terhi">Terhi Honkola</a><sup class="u-js-hide"><a href="#Aff4">4</a>,<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-Outi-Vesakoski-Aff4" data-author-popup="auth-Outi-Vesakoski-Aff4" data-author-search="Vesakoski, Outi">Outi Vesakoski</a><sup class="u-js-hide"><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-Quentin-Atkinson-Aff6-Aff7" data-author-popup="auth-Quentin-Atkinson-Aff6-Aff7" data-author-search="Atkinson, Quentin">Quentin Atkinson</a><sup class="u-js-hide"><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-Pontus-Skoglund-Aff8" data-author-popup="auth-Pontus-Skoglund-Aff8" data-author-search="Skoglund, Pontus">Pontus Skoglund</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-Alena-Kushniarevich-Aff1-Aff9" data-author-popup="auth-Alena-Kushniarevich-Aff1-Aff9" data-author-search="Kushniarevich, Alena">Alena Kushniarevich</a><sup class="u-js-hide"><a href="#Aff1">1</a>,<a href="#Aff9">9</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-Sergey-Litvinov-Aff1-Aff10" data-author-popup="auth-Sergey-Litvinov-Aff1-Aff10" data-author-search="Litvinov, Sergey">Sergey Litvinov</a><sup class="u-js-hide"><a href="#Aff1">1</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-Maere-Reidla-Aff1-Aff11" data-author-popup="auth-Maere-Reidla-Aff1-Aff11" data-author-search="Reidla, Maere">Maere Reidla</a><sup class="u-js-hide"><a href="#Aff1">1</a>,<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-Ene-Metspalu-Aff1" data-author-popup="auth-Ene-Metspalu-Aff1" data-author-search="Metspalu, Ene">Ene Metspalu</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-Lehti-Saag-Aff1-Aff11" data-author-popup="auth-Lehti-Saag-Aff1-Aff11" data-author-search="Saag, Lehti">Lehti Saag</a><sup class="u-js-hide"><a href="#Aff1">1</a>,<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-Timo-Rantanen-Aff12" data-author-popup="auth-Timo-Rantanen-Aff12" data-author-search="Rantanen, Timo">Timo Rantanen</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-Monika-Karmin-Aff1" data-author-popup="auth-Monika-Karmin-Aff1" data-author-search="Karmin, Monika">Monika Karmin</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-J_ri-Parik-Aff1-Aff11" data-author-popup="auth-J_ri-Parik-Aff1-Aff11" data-author-search="Parik, Jüri">Jüri Parik</a><sup class="u-js-hide"><a href="#Aff1">1</a>,<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-Sergey_I_-Zhadanov-Aff1-Aff13" data-author-popup="auth-Sergey_I_-Zhadanov-Aff1-Aff13" data-author-search="Zhadanov, Sergey I.">Sergey I. Zhadanov</a><sup class="u-js-hide"><a href="#Aff1">1</a>,<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-Marina-Gubina-Aff1-Aff14" data-author-popup="auth-Marina-Gubina-Aff1-Aff14" data-author-search="Gubina, Marina">Marina Gubina</a><sup class="u-js-hide"><a href="#Aff1">1</a>,<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-Larisa_D_-Damba-Aff1-Aff15" data-author-popup="auth-Larisa_D_-Damba-Aff1-Aff15" data-author-search="Damba, Larisa D.">Larisa D. Damba</a><sup class="u-js-hide"><a href="#Aff1">1</a>,<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-Marina-Bermisheva-Aff1-Aff10" data-author-popup="auth-Marina-Bermisheva-Aff1-Aff10" data-author-search="Bermisheva, Marina">Marina Bermisheva</a><sup class="u-js-hide"><a href="#Aff1">1</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-Tuuli-Reisberg-Aff1" data-author-popup="auth-Tuuli-Reisberg-Aff1" data-author-search="Reisberg, Tuuli">Tuuli Reisberg</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-Khadizhat-Dibirova-Aff1-Aff16" data-author-popup="auth-Khadizhat-Dibirova-Aff1-Aff16" data-author-search="Dibirova, Khadizhat">Khadizhat Dibirova</a><sup class="u-js-hide"><a href="#Aff1">1</a>,<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-Irina-Evseeva-Aff17-Aff18" data-author-popup="auth-Irina-Evseeva-Aff17-Aff18" data-author-search="Evseeva, Irina">Irina Evseeva</a><sup class="u-js-hide"><a href="#Aff17">17</a>,<a href="#Aff18">18</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-Mari-Nelis-Aff19" data-author-popup="auth-Mari-Nelis-Aff19" data-author-search="Nelis, Mari">Mari Nelis</a><sup class="u-js-hide"><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-Janis-Klovins-Aff20" data-author-popup="auth-Janis-Klovins-Aff20" data-author-search="Klovins, Janis">Janis Klovins</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-Andres-Metspalu-Aff19" data-author-popup="auth-Andres-Metspalu-Aff19" data-author-search="Metspalu, Andres">Andres Metspalu</a><sup class="u-js-hide"><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-T_nu-Esko-Aff19" data-author-popup="auth-T_nu-Esko-Aff19" data-author-search="Esko, Tõnu">Tõnu Esko</a><sup class="u-js-hide"><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-Oleg-Balanovsky-Aff16-Aff21" data-author-popup="auth-Oleg-Balanovsky-Aff16-Aff21" data-author-search="Balanovsky, Oleg">Oleg Balanovsky</a><sup class="u-js-hide"><a href="#Aff16">16</a>,<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-Elena-Balanovska-Aff16" data-author-popup="auth-Elena-Balanovska-Aff16" data-author-search="Balanovska, Elena">Elena Balanovska</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-Elza_K_-Khusnutdinova-Aff10-Aff22" data-author-popup="auth-Elza_K_-Khusnutdinova-Aff10-Aff22" data-author-search="Khusnutdinova, Elza K.">Elza K. Khusnutdinova</a><sup class="u-js-hide"><a href="#Aff10">10</a>,<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-Ludmila_P_-Osipova-Aff14-Aff23" data-author-popup="auth-Ludmila_P_-Osipova-Aff14-Aff23" data-author-search="Osipova, Ludmila P.">Ludmila P. Osipova</a><sup class="u-js-hide"><a href="#Aff14">14</a>,<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-Mikhail-Voevoda-Aff14-Aff23-Aff24" data-author-popup="auth-Mikhail-Voevoda-Aff14-Aff23-Aff24" data-author-search="Voevoda, Mikhail">Mikhail Voevoda</a><sup class="u-js-hide"><a href="#Aff14">14</a>,<a href="#Aff23">23</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-Richard-Villems-Aff1-Aff11" data-author-popup="auth-Richard-Villems-Aff1-Aff11" data-author-search="Villems, Richard">Richard Villems</a><sup class="u-js-hide"><a href="#Aff1">1</a>,<a href="#Aff11">11</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-Toomas-Kivisild-Aff1-Aff11-Aff25-Aff26" data-author-popup="auth-Toomas-Kivisild-Aff1-Aff11-Aff25-Aff26" data-author-search="Kivisild, Toomas">Toomas Kivisild</a><sup class="u-js-hide"><a href="#Aff1">1</a>,<a href="#Aff11">11</a>,<a href="#Aff25">25</a>,<a href="#Aff26">26</a></sup> & </li><li class="c-article-author-list__show-more" aria-label="Show all 36 authors for this article" title="Show all 36 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-Mait-Metspalu-Aff1" data-author-popup="auth-Mait-Metspalu-Aff1" data-author-search="Metspalu, Mait">Mait Metspalu</a><sup class="u-js-hide"><a href="#Aff1">1</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">Genome Biology</i></a> <b data-test="journal-volume"><span class="u-visually-hidden">volume</span> 19</b>, Article number: <span data-test="article-number">139</span> (<span data-test="article-publication-year">2018</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">45k <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">68 <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">99 <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/10.1186/s13059-018-1522-1/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> </div> <section aria-labelledby="Abs1" data-title="Abstract" lang="en"><div class="c-article-section" id="Abs1-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="Abs1">Abstract</h2><div class="c-article-section__content" id="Abs1-content"><h3 class="c-article__sub-heading" data-test="abstract-sub-heading">Background</h3><p>The genetic origins of Uralic speakers from across a vast territory in the temperate zone of North Eurasia have remained elusive. Previous studies have shown contrasting proportions of Eastern and Western Eurasian ancestry in their mitochondrial and Y chromosomal gene pools. While the maternal lineages reflect by and large the geographic background of a given Uralic-speaking population, the frequency of Y chromosomes of Eastern Eurasian origin is distinctively high among European Uralic speakers. The autosomal variation of Uralic speakers, however, has not yet been studied comprehensively.</p><h3 class="c-article__sub-heading" data-test="abstract-sub-heading">Results</h3><p>Here, we present a genome-wide analysis of 15 Uralic-speaking populations which cover all main groups of the linguistic family. We show that contemporary Uralic speakers are genetically very similar to their local geographical neighbours. However, when studying relationships among geographically distant populations, we find that most of the Uralic speakers and some of their neighbours share a genetic component of possibly Siberian origin. Additionally, we show that most Uralic speakers share significantly more genomic segments identity-by-descent with each other than with geographically equidistant speakers of other languages. We find that correlated genome-wide genetic and lexical distances among Uralic speakers suggest co-dispersion of genes and languages. Yet, we do not find long-range genetic ties between Estonians and Hungarians with their linguistic sisters that would distinguish them from their non-Uralic-speaking neighbours.</p><h3 class="c-article__sub-heading" data-test="abstract-sub-heading">Conclusions</h3><p>We show that most Uralic speakers share a distinct ancestry component of likely Siberian origin, which suggests that the spread of Uralic languages involved at least some demic component.</p></div></div></section> <section data-title="Background"><div class="c-article-section" id="Sec1-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="Sec1">Background</h2><div class="c-article-section__content" id="Sec1-content"><p>The linguistic landscape of North Eurasia is dominated by three language families—Turkic, Indo-European (IE) and Uralic. It has recently been shown that the spread of Turkic languages was mediated by gene flow from South Siberia [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 1" title="Yunusbayev B, Metspalu M, Metspalu E, Valeev A, Litvinov S, et al. The genetic legacy of the expansion of Turkic-speaking nomads across Eurasia. PLoS Genet. 2015;11:e1005068. https://doi.org/10.1371/journal.pgen.1005068 " href="/articles/10.1186/s13059-018-1522-1#ref-CR1" id="ref-link-section-d8349793e1355">1</a>]. Similarly, ancient DNA evidence of a major episode of gene flow from the Ponto-Caspian Steppe Belt to Central Europe and Central Asia during the Late Neolithic and Early Bronze Age (BA) has been interpreted as supporting the ‘Steppe Hypothesis’ of the spread of IE languages [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2" title="Haak W, Lazaridis I, Patterson N, Rohland N, Mallick S, et al. Massive migration from the steppe was a source for Indo-European languages in Europe. Nature. 2015; https://doi.org/10.1038/nature14317 " href="/articles/10.1186/s13059-018-1522-1#ref-CR2" id="ref-link-section-d8349793e1358">2</a>, <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 3" title="Allentoft ME, Sikora M, Sjögren K-G, Rasmussen S, Rasmussen M, et al. Population genomics of Bronze Age Eurasia. Nature. 2015;522:167–72. https://doi.org/10.1038/nature14507 " href="/articles/10.1186/s13059-018-1522-1#ref-CR3" id="ref-link-section-d8349793e1361">3</a>]. However, while historical linguists have some level of consensus over the origin and spread of the Uralic languages and archaeologists have views about the dynamics of material culture over the relevant time and space [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" title="Indreko R. Origin and area of settlement of the Fenno-Ugrian peoples. Science in Exile. Publication of the Scientific Quarterly “Scholar”. Heidelberg: Heidelberger Gutenberg-Druckerei GmbH; 1948. p. 3–24." href="#ref-CR4" id="ref-link-section-d8349793e1364">4</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" title="Setälä N. E (1926) Johdanto. In: Kannisto A, editor. Suomen suku I. Helsinki: Kustannusosakeyhtiö Otava." href="#ref-CR5" id="ref-link-section-d8349793e1364_1">5</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" title="Moora H. Die Vorzeit Estlands. Tartu: Akadeemiline Kooperatiiv; 1932. p. 21." href="#ref-CR6" id="ref-link-section-d8349793e1364_2">6</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" title="Carpelan C, Parpola A. Emergence, contacts and dispersal of Proto-Indo-European, Proto-Uralic and Proto-Aryan in archaeological perspective. In: Carpelan C, Parpola A, Koskikallio P, editors. Early contacts between Uralic and Indo-European: linguistic and archaeological considerations. Helsinki: Suomalais-Ugrilaisen Seuran Toimituksia. Mémoires de la Société Finno-Ougrienne; 2001. p. 55–150." href="#ref-CR7" id="ref-link-section-d8349793e1364_3">7</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 8" title="Kallio P. The language contact situation in prehistoric Northeastern Europe. In: Mailhammer R, Vennemann T, Olsen AB, editors. The linguistic roots of Europe: origin and development of European languages. Copenhagen: Museum Tusculanum Press; 2015. p. 77–102." href="/articles/10.1186/s13059-018-1522-1#ref-CR8" id="ref-link-section-d8349793e1367">8</a>], the genetic history of Uralic-speaking populations has remained poorly known.</p><p>The Uralic family contains 40–50 different languages [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" title="Abondolo D. In: Abondolo D, editor. The Uralic languages. New York/London: Routledge; 1998." href="#ref-CR9" id="ref-link-section-d8349793e1373">9</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" title="Lewis MP, Simons GF, Fennig CD. Ethnologue: languages of the world. 19th ed. Dallas: SIL International; 2016." href="#ref-CR10" id="ref-link-section-d8349793e1373_1">10</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 11" title="Salminen T. Europe and North Asia. In: Moseley C, editor. Encyclopedia of the world’s endangered languages. New York: Routledge; 2007. p. 211–82." href="/articles/10.1186/s13059-018-1522-1#ref-CR11" id="ref-link-section-d8349793e1376">11</a>] and covers a vast territory mainly from the shores of the Baltic Sea in Europe to the West Siberian Plain and the Taymyr Peninsula in Asia (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig1">1</a>a). According to the classical view, the Uralic languages derive from a protolanguage that split into two major branches—the Finno-Ugric (FU) and the Samoyed. The suggested age of the Uralic language family is 6,000–4,000 years before present (BP) (see e.g. [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" title="Janhunen J. Proto-Uralic — what, where, and when? In: Ylikoski J, editor. The Quasquicentennial of the Finno-Ugrian Society. Suomalais-Ugrilaisen Seuran Toimituksia. Mémoires de la Société Finno-Ougrienne. Helsinki: Tiedekirja; 2009. p. 57–78." href="#ref-CR12" id="ref-link-section-d8349793e1382">12</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" title="Honkola T, Vesakoski O, Korhonen K, Lehtinen J, Syrjänen K, et al. Cultural and climatic changes shape the evolutionary history of the Uralic languages. J Evol Biol. 2013; 26:1244–53. https://doi.org/10.1111/jeb.12107 " href="#ref-CR13" id="ref-link-section-d8349793e1382_1">13</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 14" title="Kallio P. Suomen kantakielten absoluuttista kronologiaa. Virittäjä. 2006;1:2–25." href="/articles/10.1186/s13059-018-1522-1#ref-CR14" id="ref-link-section-d8349793e1385">14</a>], cf. [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 15" title="Kulonen U-M. Kielitiede ja Suomen väeston juuret. In: Grünthal R, editor. Ennen muinoin: miten menneisyyttämme tutkitaan. Helsinki: Suomalaisen Kirjallisuuden Seura; 2002. p. 102–16." href="/articles/10.1186/s13059-018-1522-1#ref-CR15" id="ref-link-section-d8349793e1389">15</a>, <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 16" title="Syrjänen K, Honkola T, Korhonen K, Lehtinen J, Vesakoski O, et al. Shedding more light on language classification using basic vocabularies and phylogenetic methods. A case study of Uralic. Diachronica. 2013;30:323–52. https://doi.org/10.1075/dia.30.3.02syr " href="/articles/10.1186/s13059-018-1522-1#ref-CR16" id="ref-link-section-d8349793e1392">16</a>]). The most widely accepted hypotheses place the homeland of the Uralic language family into the watershed of river Volga and its tributaries Oka and Kama (see e.g. [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" title="Carpelan C. Essay on archaeology and languages in the western end of the Uralic zone. Tartu: Congressus Nonus Internationalis Fenno-Ugristarum. Pars I; 2000. p. 7–38." href="#ref-CR17" id="ref-link-section-d8349793e1395">17</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" title="Häkkinen J. Kantauralin ajoitus ja paikannus: perustelut puntarissa. J la Société Finno-Ougrienne. 2009;92:9–56." href="#ref-CR18" id="ref-link-section-d8349793e1395_1">18</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" title="Koivulehto J. The earliest contacts between Indo-European and Uralic speakers in the light of lexical loans. In: Carpelan C, editor. Early contacts between Uralic and Indo-European: linguistic and archaeological considerations. Helsinki: Suomalais-Ugrilaisen Seuran Toimituksia. Mémoires de la Société Finno-Ougrienne; 2001. p. 235–64." href="#ref-CR19" id="ref-link-section-d8349793e1395_2">19</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 20" title="Anthony DW. The horse, the wheel and language. How Bronze-Age riders from the Eurasian steppes shaped the modern world. Princeton: Princeton University Press; 2007. p. 93." href="/articles/10.1186/s13059-018-1522-1#ref-CR20" id="ref-link-section-d8349793e1398">20</a>] and references therein), while some scholars propose a Siberian homeland [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 12" title="Janhunen J. Proto-Uralic — what, where, and when? In: Ylikoski J, editor. The Quasquicentennial of the Finno-Ugrian Society. Suomalais-Ugrilaisen Seuran Toimituksia. Mémoires de la Société Finno-Ougrienne. Helsinki: Tiedekirja; 2009. p. 57–78." href="/articles/10.1186/s13059-018-1522-1#ref-CR12" id="ref-link-section-d8349793e1401">12</a>, <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 21" title="Napolskikh VV. Uralic fish-names and original home. Ural Jahrbücher, Neue Folge. 1993;12:35–57." href="/articles/10.1186/s13059-018-1522-1#ref-CR21" id="ref-link-section-d8349793e1404">21</a>, <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 22" title="Hajdú P. Über die alten Siedlungsraume der uralischen Sprachfamilie. Acta Linguist Hungaricae. 1964:47–83." href="/articles/10.1186/s13059-018-1522-1#ref-CR22" id="ref-link-section-d8349793e1408">22</a>]. The precursors of present-day FU languages gradually spread west towards the Baltic Sea (Proto-Finnic) [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 13" title="Honkola T, Vesakoski O, Korhonen K, Lehtinen J, Syrjänen K, et al. Cultural and climatic changes shape the evolutionary history of the Uralic languages. J Evol Biol. 2013; 26:1244–53. https://doi.org/10.1111/jeb.12107 " href="/articles/10.1186/s13059-018-1522-1#ref-CR13" id="ref-link-section-d8349793e1411">13</a>, <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 23" title="Lang V. Formation of Proto-Finnic - an archaeological scenario from the Bronze Age/Early Iron Age. In: Mantila H, Leinonen S, Brunni S, Palvianen S, Sivonen J, editors. Congressus Duodecimus Internationalis Fenno-Ugristarum. Oulu: University of Oulu; 2015. p. 63–84." href="/articles/10.1186/s13059-018-1522-1#ref-CR23" id="ref-link-section-d8349793e1414">23</a>], north-west (Proto-Saami) [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 24" title="Frog, Saarikivi J. De situ linguarum fennicarum aetatis ferreae, pars I. RMN Newsl. 2015;9:64–115." href="/articles/10.1186/s13059-018-1522-1#ref-CR24" id="ref-link-section-d8349793e1417">24</a>], north (Proto-Permian branch giving rise to Komi) [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 9" title="Abondolo D. In: Abondolo D, editor. The Uralic languages. New York/London: Routledge; 1998." href="/articles/10.1186/s13059-018-1522-1#ref-CR9" id="ref-link-section-d8349793e1420">9</a>], whereas some (Udmurt, Mordovian and Mari) remained in the Volga area. The precursors of the Ugric (Khanty, Mansi and Hungarian) and Samoyed languages (e.g. Nenets, Nganasan, Selkup), spoken today mostly to the east of the Ural Mountains, but also in Central Europe (Hungarian) and in Northeast (NE) Europe (Nenets), are thought to have descended from the easternmost varieties  of the Uralic proto-languages spoken in western [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 18" title="Häkkinen J. Kantauralin ajoitus ja paikannus: perustelut puntarissa. J la Société Finno-Ougrienne. 2009;92:9–56." href="/articles/10.1186/s13059-018-1522-1#ref-CR18" id="ref-link-section-d8349793e1423">18</a>] or eastern [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 12" title="Janhunen J. Proto-Uralic — what, where, and when? In: Ylikoski J, editor. The Quasquicentennial of the Finno-Ugrian Society. Suomalais-Ugrilaisen Seuran Toimituksia. Mémoires de la Société Finno-Ougrienne. Helsinki: Tiedekirja; 2009. p. 57–78." href="/articles/10.1186/s13059-018-1522-1#ref-CR12" id="ref-link-section-d8349793e1427">12</a>] side of the Ural Mountains. Their geographic range expansion occurred most likely as a combination of demic and cultural dispersal processes [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 9" title="Abondolo D. In: Abondolo D, editor. The Uralic languages. New York/London: Routledge; 1998." href="/articles/10.1186/s13059-018-1522-1#ref-CR9" id="ref-link-section-d8349793e1430">9</a>]. The proto-Hungarian spread southwest towards Central Europe during the first millennium AD, while the linguistic ancestors of Mansi and Khanty remained mostly in West Siberia [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 9" title="Abondolo D. In: Abondolo D, editor. The Uralic languages. New York/London: Routledge; 1998." href="/articles/10.1186/s13059-018-1522-1#ref-CR9" id="ref-link-section-d8349793e1433">9</a>]. The Samoyed languages reached the periphery of their present-day spread area in the Taimyr Peninsula as late as on sixteenth century AD [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 25" title="Popov A. The Nganasan: the material culture of the Tavgi Samoyeds. Bloomington: Indiana University Publications; 1966." href="/articles/10.1186/s13059-018-1522-1#ref-CR25" id="ref-link-section-d8349793e1436">25</a>, <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 26" title="Carpelan C. On Archaeological Aspects of Uralic, Finno-Ugric and Finnic Societies before AD 800. In: Nuorluoto J, editor. The Slavicization of the Russian North. Mechanisms and chronology. Slavica Helsingiensia; 2006. p. 78–92." href="/articles/10.1186/s13059-018-1522-1#ref-CR26" id="ref-link-section-d8349793e1439">26</a>]. Recent linguistic studies associate the diversification of the Uralic family with climatic and cultural changes [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 13" title="Honkola T, Vesakoski O, Korhonen K, Lehtinen J, Syrjänen K, et al. Cultural and climatic changes shape the evolutionary history of the Uralic languages. J Evol Biol. 2013; 26:1244–53. https://doi.org/10.1111/jeb.12107 " href="/articles/10.1186/s13059-018-1522-1#ref-CR13" id="ref-link-section-d8349793e1442">13</a>] that may have led to a considerable demographic changes since the Mesolithic times in their core areas and to further migrations towards the north and northwest.</p><div class="c-article-section__figure js-c-reading-companion-figures-item" data-test="figure" data-container-section="figure" id="figure-1" data-title="Fig. 1"><figure><figcaption><b id="Fig1" class="c-article-section__figure-caption" data-test="figure-caption-text">Fig. 1</b></figcaption><div class="c-article-section__figure-content"><div class="c-article-section__figure-item"><a class="c-article-section__figure-link" data-test="img-link" data-track="click" data-track-label="image" data-track-action="view figure" href="/articles/10.1186/s13059-018-1522-1/figures/1" rel="nofollow"><picture><source type="image/webp" srcset="//media.springernature.com/lw685/springer-static/image/art%3A10.1186%2Fs13059-018-1522-1/MediaObjects/13059_2018_1522_Fig1_HTML.png?as=webp"><img aria-describedby="Fig1" src="//media.springernature.com/lw685/springer-static/image/art%3A10.1186%2Fs13059-018-1522-1/MediaObjects/13059_2018_1522_Fig1_HTML.png" alt="figure 1" loading="lazy" width="685" height="659"></picture></a></div><div class="c-article-section__figure-description" data-test="bottom-caption" id="figure-1-desc"><p>Geographic distribution of the Uralic-speaking populations and the schematic tree of the Uralic languages. <b>a</b> The geographic spread of the Uralic-speaking populations. Colour coding corresponds to the respective language in panel <b>b</b>. <b>b</b> Schematic representation of the phylogeny of the Uralic languages. Pie diagrams indicate the relative share of West and East Eurasian mitochondrial (mtDNA) and Y chromosomal (Y) lineages. Data from Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM5">5</a>: Table S4 and Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM6">6</a>: Table S5</p></div></div><div class="u-text-right u-hide-print"><a class="c-article__pill-button" data-test="article-link" data-track="click" data-track-label="button" data-track-action="view figure" href="/articles/10.1186/s13059-018-1522-1/figures/1" data-track-dest="link:Figure1 Full size image" aria-label="Full size image figure 1" rel="nofollow"><span>Full size image</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><p>The question as which material cultures may have co-spread together with proto-Uralic and Uralic languages depends on the time estimates of the splits in the Uralic language tree. Deeper age estimates (6,000 BP) of the Uralic language tree suggest a connection between the spread of FU languages from the Volga River basin towards the Baltic Sea either with the expansion of the Neolithic culture of Combed Ware, e.g. [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 6" title="Moora H. Die Vorzeit Estlands. Tartu: Akadeemiline Kooperatiiv; 1932. p. 21." href="/articles/10.1186/s13059-018-1522-1#ref-CR6" id="ref-link-section-d8349793e1482">6</a>, <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 7" title="Carpelan C, Parpola A. Emergence, contacts and dispersal of Proto-Indo-European, Proto-Uralic and Proto-Aryan in archaeological perspective. In: Carpelan C, Parpola A, Koskikallio P, editors. Early contacts between Uralic and Indo-European: linguistic and archaeological considerations. Helsinki: Suomalais-Ugrilaisen Seuran Toimituksia. Mémoires de la Société Finno-Ougrienne; 2001. p. 55–150." href="/articles/10.1186/s13059-018-1522-1#ref-CR7" id="ref-link-section-d8349793e1485">7</a>, <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 17" title="Carpelan C. Essay on archaeology and languages in the western end of the Uralic zone. Tartu: Congressus Nonus Internationalis Fenno-Ugristarum. Pars I; 2000. p. 7–38." href="/articles/10.1186/s13059-018-1522-1#ref-CR17" id="ref-link-section-d8349793e1488">17</a>, <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 26" title="Carpelan C. On Archaeological Aspects of Uralic, Finno-Ugric and Finnic Societies before AD 800. In: Nuorluoto J, editor. The Slavicization of the Russian North. Mechanisms and chronology. Slavica Helsingiensia; 2006. p. 78–92." href="/articles/10.1186/s13059-018-1522-1#ref-CR26" id="ref-link-section-d8349793e1491">26</a>] or with the Neolithic Volosovo culture [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 7" title="Carpelan C, Parpola A. Emergence, contacts and dispersal of Proto-Indo-European, Proto-Uralic and Proto-Aryan in archaeological perspective. In: Carpelan C, Parpola A, Koskikallio P, editors. Early contacts between Uralic and Indo-European: linguistic and archaeological considerations. Helsinki: Suomalais-Ugrilaisen Seuran Toimituksia. Mémoires de la Société Finno-Ougrienne; 2001. p. 55–150." href="/articles/10.1186/s13059-018-1522-1#ref-CR7" id="ref-link-section-d8349793e1494">7</a>]. Younger age estimates support a link between the westward dispersion of Proto-Finno-Saamic and eastward dispersion of Proto-Samoyedic with a BA Sejma-Turbino (ST) cultural complex [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 14" title="Kallio P. Suomen kantakielten absoluuttista kronologiaa. Virittäjä. 2006;1:2–25." href="/articles/10.1186/s13059-018-1522-1#ref-CR14" id="ref-link-section-d8349793e1498">14</a>, <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 18" title="Häkkinen J. Kantauralin ajoitus ja paikannus: perustelut puntarissa. J la Société Finno-Ougrienne. 2009;92:9–56." href="/articles/10.1186/s13059-018-1522-1#ref-CR18" id="ref-link-section-d8349793e1501">18</a>, <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 27" title="Parpola A. The problem of Samoyed origins in the light of archaeology: On the formation and dispersal of East Uralic (Proto-Ugro-Samoyedic). In: Hyytiäinen T, Jalava L, Saarikivi J, Sandman E, editors. Per Urales ad Orientem, iter polyphonicum multilingue: Festskrift tillägnad Juha Janhunen på hans sextioårsdag den 12 februari 2012. Helsinki: Suomalais-Ugrilainen Seura; 2012. p. 287–98." href="/articles/10.1186/s13059-018-1522-1#ref-CR27" id="ref-link-section-d8349793e1504">27</a>, <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 28" title="Carpelan C. Käännekohtia Suomen esihistoriassa aikavälillä 5100-1000 eKr. In: Fogelberg P, editor. Pohjan poluilla. Suomalaisten juuret nykytutkimuksen mukaan. Helsinki: Suomen Tiedeseura; 1999. p. 249–80." href="/articles/10.1186/s13059-018-1522-1#ref-CR28" id="ref-link-section-d8349793e1507">28</a>] that mediated the diffusion of specific metal tools and weapons from the Altai Mountains over the Urals to Northern Europe or with the Netted Ware culture [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 23" title="Lang V. Formation of Proto-Finnic - an archaeological scenario from the Bronze Age/Early Iron Age. In: Mantila H, Leinonen S, Brunni S, Palvianen S, Sivonen J, editors. Congressus Duodecimus Internationalis Fenno-Ugristarum. Oulu: University of Oulu; 2015. p. 63–84." href="/articles/10.1186/s13059-018-1522-1#ref-CR23" id="ref-link-section-d8349793e1510">23</a>], which succeeded Volosovo culture in the west. It has been suggested that Proto-Uralic may have even served as the lingua franca of the merchants involved in the ST phenomenon [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 18" title="Häkkinen J. Kantauralin ajoitus ja paikannus: perustelut puntarissa. J la Société Finno-Ougrienne. 2009;92:9–56." href="/articles/10.1186/s13059-018-1522-1#ref-CR18" id="ref-link-section-d8349793e1513">18</a>]. All these scenarios imply that material culture of the Baltic Sea area in Europe was influenced by cultures spreading westward from the periphery of Europe and/or Siberia. Whether these dispersals involved the spread of both languages and people remains so far largely unknown.</p><p>Previous genetic studies have shown that demographic histories of Uralic-speaking populations inferred from maternally inherited mitochondrial (mtDNA) and paternally inherited Y chromosomes (chrY) are different. MtDNA studies of Uralic speakers suggest that the distribution of Western and Eastern Eurasian components is mostly determined by geography [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" title="Bermisheva M, Tambets K, Villems R, Khusnutdinova E. Diversity of mitochondrial DNA haplotypes in ethnic populations of the Volga-Ural region of Russia. Mol Biol. 2002;36:990–1001. https://doi.org/10.1023/A:1021677708482 " href="#ref-CR29" id="ref-link-section-d8349793e1519">29</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" title="Tambets K, Rootsi S, Kivisild T, Help H, Serk P, et al. The western and eastern roots of the Saami--the story of genetic “outliers” told by mitochondrial DNA and Y chromosomes. Am J Hum Genet. 2004;74:661–82. https://doi.org/10.1086/383203 " href="#ref-CR30" id="ref-link-section-d8349793e1519_1">30</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" title="Derbeneva OA, Starikovskaya EB, Wallace DC, Sukernik RI. Traces of early Eurasians in the Mansi of northwest Siberia revealed by mitochondrial DNA analysis. Am J Hum Genet. 2002;70:1009–114. https://doi.org/10.1086/339524 " href="#ref-CR31" id="ref-link-section-d8349793e1519_2">31</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 32" title="Derbeneva OA, Starikovskaya EB, Volod’ko NV, Wallace DC, Sukernik RI. Mitochondrial DNA variation in Kets and Nganasans and the early peoples of Northern Eurasia. Genetika. 2002;38:1554–60. https://doi.org/10.1023/A:1021111530654 " href="/articles/10.1186/s13059-018-1522-1#ref-CR32" id="ref-link-section-d8349793e1522">32</a>]. Thus, Western and Eastern Eurasian mtDNA lineages co-occur only in their contact zone in the Circum-Uralic region [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 29" title="Bermisheva M, Tambets K, Villems R, Khusnutdinova E. Diversity of mitochondrial DNA haplotypes in ethnic populations of the Volga-Ural region of Russia. Mol Biol. 2002;36:990–1001. https://doi.org/10.1023/A:1021677708482 " href="/articles/10.1186/s13059-018-1522-1#ref-CR29" id="ref-link-section-d8349793e1525">29</a>, <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 31" title="Derbeneva OA, Starikovskaya EB, Wallace DC, Sukernik RI. Traces of early Eurasians in the Mansi of northwest Siberia revealed by mitochondrial DNA analysis. Am J Hum Genet. 2002;70:1009–114. https://doi.org/10.1086/339524 " href="/articles/10.1186/s13059-018-1522-1#ref-CR31" id="ref-link-section-d8349793e1528">31</a>, <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 33" title="Pimenoff VN, Comas D, Palo JU, Vershubsky G, Kozlov A, et al. Northwest Siberian Khanty and Mansi in the junction of West and East Eurasian gene pools as revealed by uniparental markers. Eur J Hum Genet. 2008;16:1254–64. https://doi.org/10.1038/ejhg.2008.101 " href="/articles/10.1186/s13059-018-1522-1#ref-CR33" id="ref-link-section-d8349793e1531">33</a>]. In contrast, the spread of paternal lineages among Uralic speakers in Europe does not follow this pattern: up to one half of males belong to the pan-North Eurasian chrY haplogroup (hg) N3a, which is closely related to lineages found in Siberian and East Asian populations [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" title="Karmin M, Saag L, Vicente M, Sayres MAW, Järve M, et al. A recent bottleneck of Y chromosome diversity coincides with a global change in culture. Genome Res. 2015;25:459–66. https://doi.org/10.1101/gr.186684.114 " href="#ref-CR34" id="ref-link-section-d8349793e1535">34</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" title="Rootsi S, Zhivotovsky LA, Baldovic M, Kayser M, Kutuev IA, et al. A counter-clockwise northern route of the Y-chromosome haplogroup N from Southeast Asia towards Europe. Eur J Hum Genet. 2007;15:204–11. https://doi.org/10.1038/sj.ejhg.5201748 " href="#ref-CR35" id="ref-link-section-d8349793e1535_1">35</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 36" title="Ilumäe A-M, Reidla M, Chukhryaeva M, Järve M, Post H, et al. Human Y chromosome haplogroup N: a non-trivial time-resolved phylogeography that cuts across language families. Am J Hum Genet. 2016;99:163–73. https://doi.org/10.1016/j.ajhg.2016.05.025 " href="/articles/10.1186/s13059-018-1522-1#ref-CR36" id="ref-link-section-d8349793e1538">36</a>]. This hg is virtually absent or rare in Southern Europe and in IE-speaking Scandinavians [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 30" title="Tambets K, Rootsi S, Kivisild T, Help H, Serk P, et al. 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Mitochondrial DNA portrait of Latvians: towards the understanding of the genetic structure of Baltic-speaking populations. Ann Hum Genet. 2006;70:439–58. https://doi.org/10.1111/j.1469-1809.2005.00238.x " href="#ref-CR37" id="ref-link-section-d8349793e1547">37</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" title="Balanovsky O, Rootsi S, Pshenichnov A, Kivisild T, Churnosov M, et al. Two sources of the Russian patrilineal heritage in their Eurasian context. Am J Hum Genet. 2008;82:236–50. https://doi.org/10.1016/j.ajhg.2007.09.019 " href="#ref-CR38" id="ref-link-section-d8349793e1547_1">38</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" title="Fedorova SA, Reidla M, Metspalu E, Metspalu M, Rootsi S, et al. Autosomal and uniparental portraits of the native populations of Sakha (Yakutia): implications for the peopling of Northeast Eurasia. 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TRAMES A J Humanit Soc Sci. 2001;5:59–74." href="#ref-CR41" id="ref-link-section-d8349793e1547_4">41</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 42" title="Villems R, Adojaan M, Kivisild T, Metspalu E, Parik J, et al. Reconstruction of maternal lineages of Finno-Ugric speaking people and some remarks on their paternal inheritance. In: Wiik K, Julku K, editors. The roots of peoples and languages of Northern Eurasia I. Turku: Societas Historiae Fenno-Ugricae; 1998. p. 180–200." href="/articles/10.1186/s13059-018-1522-1#ref-CR42" id="ref-link-section-d8349793e1550">42</a>]. A recent study suggests that the high frequency of N3a lineages in Eastern and Northern Europe is due to a demic expansion from East Eurasia within the last 5000 years [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 35" title="Rootsi S, Zhivotovsky LA, Baldovic M, Kayser M, Kutuev IA, et al. A counter-clockwise northern route of the Y-chromosome haplogroup N from Southeast Asia towards Europe. Eur J Hum Genet. 2007;15:204–11. https://doi.org/10.1038/sj.ejhg.5201748 " href="/articles/10.1186/s13059-018-1522-1#ref-CR35" id="ref-link-section-d8349793e1554">35</a>, <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 36" title="Ilumäe A-M, Reidla M, Chukhryaeva M, Järve M, Post H, et al. Human Y chromosome haplogroup N: a non-trivial time-resolved phylogeography that cuts across language families. Am J Hum Genet. 2016;99:163–73. https://doi.org/10.1016/j.ajhg.2016.05.025 " href="/articles/10.1186/s13059-018-1522-1#ref-CR36" id="ref-link-section-d8349793e1557">36</a>]. It has also been suggested that certain hg N3a3`6 sub-branches may have co-spread with ST tools and possibly also FU languages [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 36" title="Ilumäe A-M, Reidla M, Chukhryaeva M, Järve M, Post H, et al. Human Y chromosome haplogroup N: a non-trivial time-resolved phylogeography that cuts across language families. Am J Hum Genet. 2016;99:163–73. https://doi.org/10.1016/j.ajhg.2016.05.025 " href="/articles/10.1186/s13059-018-1522-1#ref-CR36" id="ref-link-section-d8349793e1560">36</a>].</p><p>Our goal in this study was to test whether the Uralic-speaking peoples share recent common genetic ancestry in their genomes. Specifically, we tested whether the clear signal of migration between East Eurasia and Europe that is present in the distribution of paternal lineages could be also detected in the patterns of autosomal variation. It has been shown earlier that the genetic landscape of northern and northeastern European populations displays affinities with Siberia [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" title="Wong EHM, Khrunin A, Nichols L, Pushkarev D, Khokhrin D, et al. Reconstructing genetic history of Siberian and Northeastern European populations. Genome Res. 2017;27:1–14. https://doi.org/10.1101/gr.202945.115 " href="#ref-CR43" id="ref-link-section-d8349793e1567">43</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" title="Raghavan M, Skoglund P, Graf KE, Metspalu M, Albrechtsen A, et al. Upper Palaeolithic Siberian genome reveals dual ancestry of Native Americans. Nature. 2014;505:87–91. https://doi.org/10.1038/nature12736 " href="#ref-CR44" id="ref-link-section-d8349793e1567_1">44</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 45" title="Lazaridis I, Patterson N, Mittnik A, Renaud G, Mallick S, et al. Ancient human genomes suggest three ancestral populations for present-day Europeans. Nature. 2014;513 https://doi.org/10.1038/nature13673 " href="/articles/10.1186/s13059-018-1522-1#ref-CR45" id="ref-link-section-d8349793e1570">45</a>] and today the components of East Eurasian origin are seen most prominently among the Fennoscandian Saami [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 46" title="Guglielmino CR, Piazza A, Menozzi P, Cavalli-Sforza LL. Uralic genes in Europe. Am J Phys Anthr. 1990;83:57–68. https://doi.org/10.1002/ajpa.1330830107 " href="/articles/10.1186/s13059-018-1522-1#ref-CR46" id="ref-link-section-d8349793e1573">46</a>, <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 47" title="Huyghe JR, Fransen E, Hannula S, Van Laer L, Van Eyken E, et al. A genome-wide analysis of population structure in the Finnish Saami with implications for genetic association studies. Eur J Hum Genet. 2010;19:347–52. https://doi.org/10.1038/ejhg.2010.179 " href="/articles/10.1186/s13059-018-1522-1#ref-CR47" id="ref-link-section-d8349793e1576">47</a>], where they constitute about 13% of their genomes [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 47" title="Huyghe JR, Fransen E, Hannula S, Van Laer L, Van Eyken E, et al. A genome-wide analysis of population structure in the Finnish Saami with implications for genetic association studies. Eur J Hum Genet. 2010;19:347–52. https://doi.org/10.1038/ejhg.2010.179 " href="/articles/10.1186/s13059-018-1522-1#ref-CR47" id="ref-link-section-d8349793e1579">47</a>]. To this end, we generated a dataset of genome-wide genetic variation at over half a million genomic positions (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM1">1</a>: Table S1) for 15 Uralic-speaking populations (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM2">2</a>: Table S2), covering the main groups of the language family. We analysed this dataset in the context of relevant European and Asian populations.</p></div></div></section><section data-title="Results"><div class="c-article-section" id="Sec2-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="Sec2">Results</h2><div class="c-article-section__content" id="Sec2-content"><h3 class="c-article__sub-heading" id="Sec3">The population structure of Uralic speakers</h3><p>To contextualize the autosomal genetic diversity of Uralic speakers among other Eurasian populations (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM1">1</a>: Table S1), we first ran the principal component (PC) analysis (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig2">2</a>a, Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S1). The first two PCs (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig2">2</a>a, Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S1A) sketch the geography of the Eurasian populations along the East-West and North-South axes, respectively. The Uralic speakers, along with other populations speaking Slavic and Turkic languages, are scattered along the first PC axis in agreement with their geographic distribution (Figs. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig1">1</a> and <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig2">2</a>a) suggesting that geography is the main predictor of genetic affinity among the groups in the given area. Secondly, in support of this, we find that <i>F</i><sub>ST</sub>-distances between populations (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S2) decay in correlation with geographical distance (Pearson’s <i>r</i> = 0.77, <i>p</i> < 0.0001). On the UPGMA tree based on these <i>F</i><sub>ST</sub>-distances (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig2">2</a>b), the Uralic speakers cluster into several different groups close to their geographic neighbours.</p><div class="c-article-section__figure js-c-reading-companion-figures-item" data-test="figure" data-container-section="figure" id="figure-2" data-title="Fig. 2"><figure><figcaption><b id="Fig2" class="c-article-section__figure-caption" data-test="figure-caption-text">Fig. 2</b></figcaption><div class="c-article-section__figure-content"><div class="c-article-section__figure-item"><a class="c-article-section__figure-link" data-test="img-link" data-track="click" data-track-label="image" data-track-action="view figure" href="/articles/10.1186/s13059-018-1522-1/figures/2" rel="nofollow"><picture><source type="image/webp" srcset="//media.springernature.com/lw685/springer-static/image/art%3A10.1186%2Fs13059-018-1522-1/MediaObjects/13059_2018_1522_Fig2_HTML.png?as=webp"><img aria-describedby="Fig2" src="//media.springernature.com/lw685/springer-static/image/art%3A10.1186%2Fs13059-018-1522-1/MediaObjects/13059_2018_1522_Fig2_HTML.png" alt="figure 2" loading="lazy" width="685" height="867"></picture></a></div><div class="c-article-section__figure-description" data-test="bottom-caption" id="figure-2-desc"><p>Principal component analysis (PCA) and genetic distances of Uralic-speaking populations. <b>a</b> PCA (PC1 vs PC2) of the Uralic-speaking populations (highlighted, population abbreviations are as in Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM1">1</a>: Table S1). Values in brackets along the axes indicate the proportion of genetic variation explained by the components. <b>b</b> UPGMA tree of <i>F</i><sub>ST</sub> distances calculated based on autosomal genetic variation</p></div></div><div class="u-text-right u-hide-print"><a class="c-article__pill-button" data-test="article-link" data-track="click" data-track-label="button" data-track-action="view figure" href="/articles/10.1186/s13059-018-1522-1/figures/2" data-track-dest="link:Figure2 Full size image" aria-label="Full size image figure 2" rel="nofollow"><span>Full size image</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><p>We next used ADMIXTURE [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 48" title="Patterson N, Moorjani P, Luo Y, Mallick S, Rohland N, et al. Ancient admixture in human history. Genetics. 2012;192:1065–93. https://doi.org/10.1534/genetics.112.145037 " href="/articles/10.1186/s13059-018-1522-1#ref-CR48" id="ref-link-section-d8349793e1678">48</a>], which presents the individuals as composed of inferred genetic components in proportions that maximize Hardy-Weinberg and linkage equilibrium in the overall sample (see the ‘<a data-track="click" data-track-label="link" data-track-action="section anchor" href="/articles/10.1186/s13059-018-1522-1#Sec10">Methods</a>’ section for choice of presented <i>K</i>). Overall, and specifically at lower values of <i>K</i>, the genetic makeup of Uralic speakers resembles that of their geographic neighbours. The Saami and (a subset of) the Mansi serve as exceptions to that pattern being more similar to geographically more distant populations (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig3">3</a>a, Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: S3). However, starting from <i>K</i> = 9, ADMIXTURE identifies a genetic component (k9, magenta in Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig3">3</a>a, Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: S3), which is predominantly, although not exclusively, found in Uralic speakers. This component is also well visible on <i>K</i> = 10, which has the best cross-validation index among all tests (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: S3B). The spatial distribution of this component (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig3">3</a>b) shows a frequency peak among Ob-Ugric and Samoyed speakers as well as among neighbouring Kets (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig3">3</a>a). The proportion of k9 decreases rapidly from West Siberia towards east, south and west, constituting on average 40% of the genetic ancestry of FU speakers in Volga-Ural region (VUR) and 20% in their Turkic-speaking neighbours (Bashkirs, Tatars, Chuvashes; Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig3">3</a>a). The proportion of this component among the Saami in Northern Scandinavia is again similar to that of the VUR FU speakers, which is exceptional in the geographic context. It is also notable that North Russians, sampled from near the White Sea, differ from other Russians by sporting higher proportions of k9 (10–15%), which is similar to the values we observe in their Finnic-speaking neighbours. Notably, Estonians and Hungarians, who are geographically the westernmost Uralic speakers, virtually lack the k9 cluster membership.</p><div class="c-article-section__figure js-c-reading-companion-figures-item" data-test="figure" data-container-section="figure" id="figure-3" data-title="Fig. 3"><figure><figcaption><b id="Fig3" class="c-article-section__figure-caption" data-test="figure-caption-text">Fig. 3</b></figcaption><div class="c-article-section__figure-content"><div class="c-article-section__figure-item"><a class="c-article-section__figure-link" data-test="img-link" data-track="click" data-track-label="image" data-track-action="view figure" href="/articles/10.1186/s13059-018-1522-1/figures/3" rel="nofollow"><picture><source type="image/webp" srcset="//media.springernature.com/lw685/springer-static/image/art%3A10.1186%2Fs13059-018-1522-1/MediaObjects/13059_2018_1522_Fig3_HTML.png?as=webp"><img aria-describedby="Fig3" src="//media.springernature.com/lw685/springer-static/image/art%3A10.1186%2Fs13059-018-1522-1/MediaObjects/13059_2018_1522_Fig3_HTML.png" alt="figure 3" loading="lazy" width="685" height="474"></picture></a></div><div class="c-article-section__figure-description" data-test="bottom-caption" id="figure-3-desc"><p>Population structure of Uralic-speaking populations inferred from ADMIXTURE analysis on autosomal SNPs in Eurasian context. <b>a</b> Individual ancestry estimates for populations of interest for selected number of assumed ancestral populations (K3, K6, K9, K11). Ancestry components discussed in a main text (k2, k3, k5, k6, k9, k11) are indicated and have the same colours throughout. The names of the Uralic-speaking populations are indicated with blue (Finno-Ugric) or orange (Samoyedic). The full bar plot is presented in Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S3. <b>b</b> Frequency map of component k9</p></div></div><div class="u-text-right u-hide-print"><a class="c-article__pill-button" data-test="article-link" data-track="click" data-track-label="button" data-track-action="view figure" href="/articles/10.1186/s13059-018-1522-1/figures/3" data-track-dest="link:Figure3 Full size image" aria-label="Full size image figure 3" rel="nofollow"><span>Full size image</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><p>The mitochondrial gene pool of most of the Uralic speakers is comprised of typical West Eurasian mtDNAs (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig1">1</a>b). Only West Siberian Nenets and Nganasans have > 50% of Eastern Eurasian mtDNA variants (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM4">4</a>: Table S3 and Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM5">5</a>: Table S4). Contrary to that, a considerable amount of the chrY lineages of both West Siberian and European Uralic speakers belong to East Eurasian hg N (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig1">1</a>b, Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM6">6</a>: Table S5). The only exceptions to this pattern among the Uralic speakers are Hungarians and Selkups. Among Hungarians, hg N is virtually absent, while among Selkups, it is much less frequent (< 10%) than in other Samoyeds (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM6">6</a>: Table S5). The prevailing hg in the paternal pool of Selkups is hg Q, which they share with genetically similar Kets and South Siberians (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S11). Besides low frequencies of hg N Selkups share with East European populations also hg R1a-M458 (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM6">6</a>: Table S5). We performed correlation analysis to formally test whether the distribution of the k9 component (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig3">3</a>b) spatially overlaps with the spread of specific chrY hg N2 and N3 lineages that have been shown to be relevant in the context of Uralic speakers [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 36" title="Ilumäe A-M, Reidla M, Chukhryaeva M, Järve M, Post H, et al. Human Y chromosome haplogroup N: a non-trivial time-resolved phylogeography that cuts across language families. Am J Hum Genet. 2016;99:163–73. https://doi.org/10.1016/j.ajhg.2016.05.025 " href="/articles/10.1186/s13059-018-1522-1#ref-CR36" id="ref-link-section-d8349793e1780">36</a>]. We found a weak but significant correlation with the sub-hgs spread near the Ural Mountains, but not with those that reach up to Fennoscandia (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM7">7</a>: Table S6).</p><p>We also tested the different demographic histories of female and male lineages by comparing outgroup <i>f</i>3 results for autosomal and X chromosome (chrX) data for pairs of populations (Estonians, Udmurts or Khanty vs others) with high versus low probability to share their patrilineal ancestry in chrY hg N (see the ‘<a data-track="click" data-track-label="link" data-track-action="section anchor" href="/articles/10.1186/s13059-018-1522-1#Sec10">Methods’</a> section, Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S13). We found a minor but significant excess of autosomal affinity relative to chrX for pairs of populations that showed a higher than 10% chance of two randomly sampled males across the two groups sharing their chrY ancestry in hg N3-M178, compared to pairs of populations where such probability is lower than 5% (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S13).</p><p>In sum, these results suggest that most of the Uralic speakers may indeed share some level of genetic continuity via k9, which, however, also extends to the geographically close Turkic speakers.</p><h3 class="c-article__sub-heading" id="Sec4">Distilling the language-mediated excess of genetic continuity</h3><p>To test whether the common genetic substrate of Uralic speakers suggested by the k9 component also presents itself in the sharing patterns of derived alleles across the genome, we calculated <i>D</i>-statistics [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 49" title="Reich D, Thangaraj K, Patterson N, Price AL, Singh L. Reconstructing Indian population history. Nature. 2009;461:489–94. https://doi.org/10.1038/nature08365 " href="/articles/10.1186/s13059-018-1522-1#ref-CR49" id="ref-link-section-d8349793e1816">49</a>] as in Skoglund et al. [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 50" title="Skoglund P, Malmström H, Omrak A, Raghavan M, Valdiosera C, et al. Genomic diversity and admixture differs for Stone-Age Scandinavian foragers and farmers. Science. 2014;344:747–50. https://doi.org/10.1126/science.1253448 " href="/articles/10.1186/s13059-018-1522-1#ref-CR50" id="ref-link-section-d8349793e1819">50</a>] (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S4). We explored derived allele sharing patterns in a wide set of Eurasian populations contrasting sharing with the westernmost Uralic speakers (Saami, Finns, Estonians, Hungarians) on one hand and European populations (Swedes, Poles, French) on the other. We found that it is the admixture with the Siberians that makes the Western Uralic speakers different from the tested European populations (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S4A-F, H, J, L). Differentiating between Estonians and Finns, the Siberians share more derived alleles with Finns, while the geographic neighbours of Estonians (and Finns) share more alleles with Estonians (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S4M). Importantly, Estonians do not share more derived alleles with other Finnic, Saami, VUR FU or Ob-Ugric-speaking populations than Latvians (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S4O). The difference between Estonians and Latvians is instead manifested through significantly higher levels of shared drift between Estonians and Siberians on the one hand and Latvians and their immediate geographic neighbours on the other hand. None of the Uralic speakers, including linguistically close Khanty and Mansi, show significantly closer affinities to the Hungarians than any non-FU population from NE Europe (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S4R).</p><p>We next tested whether the Uralic-speaking populations share more identity-by-descent (IBD) [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 51" title="Browning BL, Browning SR. A fast, powerful method for detecting identity by descent. Am J Hum Genet. 2011;88:173–82. https://doi.org/10.1016/j.ajhg.2011.01.010 " href="/articles/10.1186/s13059-018-1522-1#ref-CR51" id="ref-link-section-d8349793e1841">51</a>] segments with geographically distant Uralic groups than their non-Uralic neighbours do (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig4">4</a> A–E, Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM8">8</a>: Table S7, Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM9">9</a>: Table S8, Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM10">10</a>: Table S9). High IBD sharing between Permic speakers and Khanty has been earlier reported in Triska et al. [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 52" title="Triska P, Chekanov N, Stepanov V, Khusnutdinova EK, Kumar GPA, et al. Between Lake Baikal and the Baltic Sea: genomic history of the gateway to Europe. BMC Genet. 2017;18(110) https://doi.org/10.1186/s12863-017-0578-3 " href="/articles/10.1186/s13059-018-1522-1#ref-CR52" id="ref-link-section-d8349793e1857">52</a>]. Indeed, Finnic speakers and Saami share more IBD segments with their distant linguistic relatives in VUR (Mari, Komi and Udmurts) and even with West Siberian Uralic speakers than NE Europeans in the control group (blue cells, Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig4">4</a> A). In addition, Saami and Karelians show a significant excess of IBD segment sharing with several non-Uralic peoples of Siberia (green cells, Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig4">4</a> A). Compared to their non-Uralic neighbours, the Samoyedic Nganasans share more IBD segments with all the tested Siberian Uralic speakers, most of the Uralic speakers from VUR, and even with Saami and Karelians from NE Europe (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig4">4</a> E). When Maris and Udmurts (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig4">4</a> C) are compared to their neighbouring Chuvashes, Tatars and Bashkirs, they display more shared IBD segments with Saami, Vepsians and North Russians in the west and specifically only with the Uralic-speaking populations to the east of the Ural Mountains. All the above-mentioned findings of IBD analyses attest to at least some degree of common genetic substrate among most of the analysed Uralic populations. Yet, we did not find any excess IBD sharing when Estonians (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig4">4</a> A), Hungarians (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig4">4</a> B) and Mordovians (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig4">4</a> D) were compared to the Uralic speakers from VUR and Siberia.</p><div class="c-article-section__figure js-c-reading-companion-figures-item" data-test="figure" data-container-section="figure" id="figure-4" data-title="Fig. 4"><figure><figcaption><b id="Fig4" class="c-article-section__figure-caption" data-test="figure-caption-text">Fig. 4</b></figcaption><div class="c-article-section__figure-content"><div class="c-article-section__figure-item"><a class="c-article-section__figure-link" data-test="img-link" data-track="click" data-track-label="image" data-track-action="view figure" href="/articles/10.1186/s13059-018-1522-1/figures/4" rel="nofollow"><picture><source type="image/webp" srcset="//media.springernature.com/lw685/springer-static/image/art%3A10.1186%2Fs13059-018-1522-1/MediaObjects/13059_2018_1522_Fig4_HTML.png?as=webp"><img aria-describedby="Fig4" src="//media.springernature.com/lw685/springer-static/image/art%3A10.1186%2Fs13059-018-1522-1/MediaObjects/13059_2018_1522_Fig4_HTML.png" alt="figure 4" loading="lazy" width="685" height="376"></picture></a></div><div class="c-article-section__figure-description" data-test="bottom-caption" id="figure-4-desc"><p>Share of ~ 1–2 cM identity-by-descent (IBD) segments within and between regional groups of Uralic speakers. For each Uralic-speaking population representing lines in this matrix, we performed permutation test to estimate if it shows higher IBD segment sharing with other population (listed in columns) as compared to their geographic control group. Empty rectangles indicate no excess IBD sharing, rectangles filled in blue indicate comparisons when statistically significant excess IBD sharing was detected between one Uralic-speaking population with another Uralic-speaking population (listed in columns), rectangles filled in green mark the comparisons when a Uralic-speaking population shows excess IBD sharing with a non-Uralic-speaking population. For each tested Uralic speaker (matrix rows) populations in the control group that were used to generate permuted samples are indicated using small circles. For example, the rectangle filled in blue for Vepsians and Komis (A) implies that the Uralic-speaking Vepsians share more IBD segments with the Uralic-speaking Komis than the geographic control group for Vepsians, i.e. populations indicated with small circles (Central and North Russians, Swedes, Latvians and Lithuanians). The rectangle filled in green for Vepsians and Dolgans shows that the Uralic-speaking Vepsians share more IBD segments with the non-Uralic-speaking Dolgans than the geographic control group</p></div></div><div class="u-text-right u-hide-print"><a class="c-article__pill-button" data-test="article-link" data-track="click" data-track-label="button" data-track-action="view figure" href="/articles/10.1186/s13059-018-1522-1/figures/4" data-track-dest="link:Figure4 Full size image" aria-label="Full size image figure 4" rel="nofollow"><span>Full size image</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><h4 class="c-article__sub-heading c-article__sub-heading--small" id="Sec5">fineSTRUCTURE. Globetrotter</h4><p>To study the fine-scale genetic structure of Uralic-speaking populations and to reconstruct past admixture events in their history, we used the haplotype-based approach implemented in fineSTRUCTURE [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 53" title="Lawson DJ, Hellenthal G, Myers S, Falush D. Inference of population structure using dense haplotype data. PLoS Genet. 2012;8:e1002453. https://doi.org/10.1371/journal.pgen.1002453 " href="/articles/10.1186/s13059-018-1522-1#ref-CR53" id="ref-link-section-d8349793e1906">53</a>] and GLOBETROTTER [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 54" title="Hellenthal G, Busby GBJ, Band G, Wilson JF, Capelli C, et al. A genetic atlas of human admixture history. Science (80- ). 2014;343:747–51. https://doi.org/10.1126/science.1243518 " href="/articles/10.1186/s13059-018-1522-1#ref-CR54" id="ref-link-section-d8349793e1909">54</a>]. fineSTRUCTURE clusters individuals into natural groups based on patterns of haplotype sharing similarity. The clusters identified in our sample largely correspond to self-identified ethnic groups while higher hierarchical clusters follow broader geographic proximity patterns (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figures S5-S6). In some cases, geographic and linguistic proximity co-vary and for example the clusters we name ‘Finnic’ and ‘Saami’ (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S5) consist of only Uralic speakers, while ‘Europe 1’ and ‘Europe 2’ encompass both Uralic and non-Uralic speakers. Here, Uralic-speaking Hungarians group together with Slavic speakers and Germanic-speaking Swedes in ‘Europe 2’ (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S5). Similarly, Uralic-speaking Estonians form a cluster with Baltic-speaking Latvians and Lithuanians (‘Europe 1’ in Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S5), which also includes Mordovians and Russians.</p> <h3 class="c-article__sub-heading" id="FPar1">Globetrotter full analysis</h3> <p>The natural clusters defined by fineSTRUCTURE were further used to study admixture history with Globetrotter analysis, which was applied in two different setups. First, we performed the ‘full’ analysis, where every recipient population could copy from any other donor group (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig5">5</a>a, Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM11">11</a>: Table S10). Populations of interest were clustered into three geographically defined groups: (1) European (blue palette, all studied Europeans except the easternmost VUR populations—Maris, Udmurts, Komis, Tatars, Bashkirs and Chuvashes); (2) VUR (green palette); and (3) West Siberian (magenta palette). Most of the inferred admixture events in this analysis were simple one-date events with high statistical support (third quartile of maxR2 fit scores of single date events = 0.91, Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM11">11</a>, Table S10). As expected, many events involved contacts between geographically close source populations. For example, the majority of admixture events in the European set (‘Europe 1’, ‘Europe 2’, ‘Finnic’ and ‘Saami’, Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig5">5</a>a) involve populations from within the set, and only two groups show traces of admixture from non-European sources: ‘Europe 2’ from Caucasus/Near Eastern groups and ‘Saami’ from the VUR (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig5">5</a>).</p><div class="c-article-section__figure js-c-reading-companion-figures-item" data-test="figure" data-container-section="figure" id="figure-5" data-title="Fig. 5"><figure><figcaption><b id="Fig5" class="c-article-section__figure-caption" data-test="figure-caption-text">Fig. 5</b></figcaption><div class="c-article-section__figure-content"><div class="c-article-section__figure-item"><a class="c-article-section__figure-link" data-test="img-link" data-track="click" data-track-label="image" data-track-action="view figure" href="/articles/10.1186/s13059-018-1522-1/figures/5" rel="nofollow"><picture><source type="image/webp" srcset="//media.springernature.com/lw685/springer-static/image/art%3A10.1186%2Fs13059-018-1522-1/MediaObjects/13059_2018_1522_Fig5_HTML.png?as=webp"><img aria-describedby="Fig5" src="//media.springernature.com/lw685/springer-static/image/art%3A10.1186%2Fs13059-018-1522-1/MediaObjects/13059_2018_1522_Fig5_HTML.png" alt="figure 5" loading="lazy" width="685" height="345"></picture></a></div><div class="c-article-section__figure-description" data-test="bottom-caption" id="figure-5-desc"><p>Circos plots of GLOBETROTTER (GT) results. The outer circle represents target groups for which GT inference was performed (wide segments) and additional surrogate populations, which were used to describe admixture in target populations (narrow segments). Geographic affiliation of target groups is colour-coded: blue—Europe (except populations from Volga-Ural region—Komis, Udmurts, Maris, Tatars, Chuvashes, Bashkirs); green—Volga-Ural region; and magenta—Western Siberia. Inner bar plots depict genetic composition of inferred sources of admixture in each of the target groups. A pair of sources is shown for a simple one-way admixture event between two populations, and an additional pair of sources for the less strongly signaled event is shown for a one-date multi-way admixture between more than two sources (marked as MW in the outer circle). In a simple one-date event, a pair of sources contributes 100% of the DNA of the target population. Surrogate populations in the inner bar plots are shaded according to the colour scheme given in the outer ring, and those contributing < 3% to mixing sources are coloured in grey. Point estimates and confidence intervals for the date of inferred admixture event are shown next to the cluster label. The details of the GT source groups are given in Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S5 and Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM11">11</a>: Table S10. <b>a</b> Results of ‘full’ analysis, where each cluster was allowed to copy from every other cluster. <b>b</b> Results of ‘regional’ analysis, where no copying between samples from the same geographical region was allowed. For example, in the ‘full’ analysis of the ‘Europe 1’ cluster, a simple one-date admixture event was detected. The first source population contributes 85% of the total DNA, including 76% from the ‘Europe 2’ surrogate; the second source contributes 15% and is dominated by the ‘Finnic’ cluster. The admixture took place around 1211 CE (95% CI: 1213–1412 CE). Abbreviations: C-Central; Cauc-Caucasus; E-East; N-North; S-South; Sib-Siberia; W-West.</p></div></div><div class="u-text-right u-hide-print"><a class="c-article__pill-button" data-test="article-link" data-track="click" data-track-label="button" data-track-action="view figure" href="/articles/10.1186/s13059-018-1522-1/figures/5" data-track-dest="link:Figure5 Full size image" aria-label="Full size image figure 5" rel="nofollow"><span>Full size image</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> <p>In the VUR (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig5">5</a>a), all populations except Maris and Udmurts show considerable admixture with other Europeans. In West Siberia, three out of four clusters display traces of admixture mostly with Siberian, and East and Central Asian donors. On the contrary, ‘Mansi’ has a distinctive admixture profile—nine Mansi samples cluster closely together with VUR populations (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S5; for ADMIXTURE profiles, see also Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig3">3</a>) and, similarly to them, show evidence for substantial recent admixture with Europeans around sixteenth to eighteenth century. This group is considered as an outlier here. The rest of the Mansi are clustered with Khanty people following the linguistic grouping of Ob-Ugric (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S5).</p> <h3 class="c-article__sub-heading" id="FPar2">Globetrotter regional analysis</h3> <p>The excess of admixture events between closely related geographical neighbours may mask traces of subtle genetic contacts with distantly related populations [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 54" title="Hellenthal G, Busby GBJ, Band G, Wilson JF, Capelli C, et al. A genetic atlas of human admixture history. Science (80- ). 2014;343:747–51. https://doi.org/10.1126/science.1243518 " href="/articles/10.1186/s13059-018-1522-1#ref-CR54" id="ref-link-section-d8349793e2001">54</a>]. Therefore, in the ‘regional’ analysis, we excluded neighbours from the set of possible surrogates, allowing copying only from donors with a different group label, if not specified differently (see Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM12">12</a>: Table S11 and note therein). Similarly to the ‘full’ analysis, ‘regional’ results have high statistical support (third quartile of maxR2 fit scores of single date events = 0.93, Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM11">11</a>: Table S10).</p> <p>In the European set, only the Uralic-speaking ‘Finnic’ and ‘Saami’ clusters have a detectable (more than 3%) amount of admixture with West Siberian sources (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig5">5</a>b), even if we leave aside the contribution from already admixed Western Siberian ‘Mansi’ cluster (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig5">5</a> and Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S5). In ‘Saami’, the Siberian influence is more notable as well as diverse: it is linked both with ‘Samoyed’ (consisting here of Nenets, Selkups and neighbouring Kets) and with West/Central Siberian (‘W-C-Sib’) clusters (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig5">5</a>b).</p> <p>The Uralic- and non-Uralic clusters from the VUR have different admixture histories. Turkic speakers (Bashkirs, Tatars and Chuvashes) contain three European donors (‘Europe1’, ‘Europe2’ and ‘W-S-Europe’; Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig5">5</a>b), while admixture in Uralic speakers displays mostly only one dominating European Eastern Baltic/Russian surrogate (‘Europe1’). In addition, Turkic speakers receive substantial genetic contribution from South Siberian/East Asian groups (‘E-Asia/S-Sib’ in Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig5">5</a>b), as was also shown earlier in Yunusbaev et al. [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 1" title="Yunusbayev B, Metspalu M, Metspalu E, Valeev A, Litvinov S, et al. The genetic legacy of the expansion of Turkic-speaking nomads across Eurasia. PLoS Genet. 2015;11:e1005068. https://doi.org/10.1371/journal.pgen.1005068 " href="/articles/10.1186/s13059-018-1522-1#ref-CR1" id="ref-link-section-d8349793e2034">1</a>]. This is not seen in the Uralic-speaking groups (Komis, Maris, Udmurts), who instead have both ‘Khanty-Mansi’ and ‘Samoyed’, i.e. Uralic-speaking Siberian donors. Contacts between Uralic speakers from Europe and West Siberia/VUR display mostly unidirectional east-to-west ‘donating’ pattern: for example, Komis are dominant surrogates for the ‘Finnic’ and ‘Saami’ groups, but the latter two do not contribute much to admixture events involving Komis. A similar trend was also seen in IBD analysis (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig4">4</a>).</p> <p>In the ‘regional’ analysis, admixture sources of West Siberian ‘Khanty-Mansi’ include Samoyeds and a range of VUR surrogates with a minor Central Asian/South Siberian component (‘C-Asia/S-Sib’). The ‘Samoyed’ cluster shows evidence for a complex one-date multiway admixture shaped by multiple regionally diverse surrogates dominated by West/Central Siberian and Khanty-Mansi groups. The ‘Samoyed’ cluster is, together with South and West Siberians, also a major contributor to an admixture event in a separate Samoyed-speaking group—Nganasans (‘Nganassan’, Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig5">5</a>b and Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S6, see Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM12">12</a>: Table S11 for details). The most distinct difference between Ob-Ugric (Khanty-Mansi) and Samoyedic speakers (Nenets, Selkups and Nganasans) is the presence of East Asian/South Siberian (‘E-Asia/S-Sib’, Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig5">5</a>) component in the latter.</p> <p>The time depth of the Globetrotter (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig5">5</a>b) inferred admixture events is relatively recent—500–1900 AD (see also complementary ALDER results, in Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM13">13</a>: Table S12 and Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S7)—and agrees broadly with the results reported in Busby et al. [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 55" title="Busby GBJ, Hellenthal G, Montinaro F, Tofanelli S, Bulayeva K, et al. The role of recent admixture in forming the contemporary west Eurasian genomic landscape. Curr Biol. 2015;25:2518–26. https://doi.org/10.1016/j.cub.2015.08.007 " href="/articles/10.1186/s13059-018-1522-1#ref-CR55" id="ref-link-section-d8349793e2068">55</a>]. A more detailed examination of the ALDER dates, however, reveals an interesting pattern. The admixture events detected in the Baltic Sea region and VUR Uralic speakers are the oldest (800–900 AD or older) followed by those in VUR Turkic speakers (<span class="stix">∼</span>1200–1300 AD), while the admixture dates for most of the Siberian populations (>1500 AD) are the most recent (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S7). The West Eurasian influx into West Siberia seen in modern genomes was thus very recent, while the East Eurasian influx into NE Europe seems to have taken place within the first millennium AD (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig5">5</a>b, Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S7).</p> <h3 class="c-article__sub-heading" id="Sec6">Affinities of the Uralic speakers with ancient Eurasians</h3><p>We next calculated outgroup <i>f</i>3-statistics [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 48" title="Patterson N, Moorjani P, Luo Y, Mallick S, Rohland N, et al. Ancient admixture in human history. Genetics. 2012;192:1065–93. https://doi.org/10.1534/genetics.112.145037 " href="/articles/10.1186/s13059-018-1522-1#ref-CR48" id="ref-link-section-d8349793e2094">48</a>] to estimate the extent of shared genetic drift between modern and ancient Eurasians (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM14">14</a>: Table S13, Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figures S8-S9). Consistent with previous reports [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 45" title="Lazaridis I, Patterson N, Mittnik A, Renaud G, Mallick S, et al. Ancient human genomes suggest three ancestral populations for present-day Europeans. Nature. 2014;513 https://doi.org/10.1038/nature13673 " href="/articles/10.1186/s13059-018-1522-1#ref-CR45" id="ref-link-section-d8349793e2103">45</a>, <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 50" title="Skoglund P, Malmström H, Omrak A, Raghavan M, Valdiosera C, et al. Genomic diversity and admixture differs for Stone-Age Scandinavian foragers and farmers. Science. 2014;344:747–50. https://doi.org/10.1126/science.1253448 " href="/articles/10.1186/s13059-018-1522-1#ref-CR50" id="ref-link-section-d8349793e2107">50</a>], we find that the NE European populations including the Uralic speakers share more drift with any European Mesolithic hunter-gatherer group than Central or Western Europeans (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S9A-C). Contrasting the genetic contribution of western hunter-gatherers (WHG) and eastern hunter-gatherers (EHG), we find that VUR Uralic speakers and the Saami share more drift with EHG. Conversely, WHG shares more drift with the Finnic and West European populations (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S9A). Interestingly, we see a similar pattern of excess of shared drift between VUR and EHG if we substitute WHG with the aDNA sample from the Yamnaya culture (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S9D). As reported before [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2" title="Haak W, Lazaridis I, Patterson N, Rohland N, Mallick S, et al. Massive migration from the steppe was a source for Indo-European languages in Europe. Nature. 2015; https://doi.org/10.1038/nature14317 " href="/articles/10.1186/s13059-018-1522-1#ref-CR2" id="ref-link-section-d8349793e2119">2</a>, <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 45" title="Lazaridis I, Patterson N, Mittnik A, Renaud G, Mallick S, et al. Ancient human genomes suggest three ancestral populations for present-day Europeans. Nature. 2014;513 https://doi.org/10.1038/nature13673 " href="/articles/10.1186/s13059-018-1522-1#ref-CR45" id="ref-link-section-d8349793e2122">45</a>], the genetic contribution of European early farmers decreases along an axis from Southern Europe towards the Ural Mountains (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig6">6</a>, Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S9E-F).</p><div class="c-article-section__figure js-c-reading-companion-figures-item" data-test="figure" data-container-section="figure" id="figure-6" data-title="Fig. 6"><figure><figcaption><b id="Fig6" class="c-article-section__figure-caption" data-test="figure-caption-text">Fig. 6</b></figcaption><div class="c-article-section__figure-content"><div class="c-article-section__figure-item"><a class="c-article-section__figure-link" data-test="img-link" data-track="click" data-track-label="image" data-track-action="view figure" href="/articles/10.1186/s13059-018-1522-1/figures/6" rel="nofollow"><picture><source type="image/webp" srcset="//media.springernature.com/lw685/springer-static/image/art%3A10.1186%2Fs13059-018-1522-1/MediaObjects/13059_2018_1522_Fig6_HTML.png?as=webp"><img aria-describedby="Fig6" src="//media.springernature.com/lw685/springer-static/image/art%3A10.1186%2Fs13059-018-1522-1/MediaObjects/13059_2018_1522_Fig6_HTML.png" alt="figure 6" loading="lazy" width="685" height="340"></picture></a></div><div class="c-article-section__figure-description" data-test="bottom-caption" id="figure-6-desc"><p>Proportions of ancestral components in studied European and Siberian populations and the tested qpGraph model. <b>a</b> The qpGraph model fitting the data for the tested populations. Colour codes for the terminal nodes: pink—modern populations (‘Population X’ refers to test population) and yellow—ancient populations (aDNA samples and their pools). Nodes coloured other than pink or yellow are hypothetical intermediate populations. We putatively named nodes which we used as admixture sources using the main recipient among known populations. The colours of intermediate nodes on the qpGraph model match those on the admixture proportions panel. <b>b</b> Admixture proportions (%) of ancestral components. We calculated the admixture proportions summing up the relative shares of a set of intermediate populations to explain the full spectrum of admixture components in the test population. We further did the same for the intermediate node CWC’ and present the proportions of the mixing three components in the stacked column bar of CWC’. Colour codes for ancestral components are as follows: dark green—Western hunter gatherer (WHG’); light green—Eastern hunter gatherer (EHG’); grey—European early farmer (LBK’); dark blue—carriers of Corded Ware culture (CWC’); and dark grey—Siberian. CWC’ consists of three sub-components: blue—Caucasian hunter-gatherer in Yamnaya (CHGinY’); light blue—Eastern hunter-gatherer in Yamnaya (EHGinY’); and light grey—Neolithic Levant (NeolL’)</p></div></div><div class="u-text-right u-hide-print"><a class="c-article__pill-button" data-test="article-link" data-track="click" data-track-label="button" data-track-action="view figure" href="/articles/10.1186/s13059-018-1522-1/figures/6" data-track-dest="link:Figure6 Full size image" aria-label="Full size image figure 6" rel="nofollow"><span>Full size image</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><p>We then used the <i>qpGraph</i> software [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 48" title="Patterson N, Moorjani P, Luo Y, Mallick S, Rohland N, et al. Ancient admixture in human history. Genetics. 2012;192:1065–93. https://doi.org/10.1534/genetics.112.145037 " href="/articles/10.1186/s13059-018-1522-1#ref-CR48" id="ref-link-section-d8349793e2162">48</a>] to test alternative demographic scenarios by trying to fit the genetic diversity observed in a range of the extant Finno-Ugric populations through a model involving the four basic European ancestral components: WHG, EHG, early farmers (LBK), steppe people of Yamnaya/Corded Ware culture (CWC) and a Siberian component (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig6">6</a>, Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S10). We chose the modern Nganasans to serve as a proxy for the latter component because we see least evidence for Western Eurasian admixture (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S3) among them. We also tested the Khantys for that proxy but the model did not fit (yielding <i>f</i>2-statistics, <i>Z</i>-score > 3). The only Uralic-speaking population that did not fit into the tested model with five ancestral components were Hungarians. The <i>qpGraph</i> estimates of the contributions from the Siberian component show that it is the main ancestry component in the West Siberian Uralic speakers and constitutes up to one third of the genomes of modern VUR and the Saami (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig6">6</a>). It drops, however, to less than 10% in most of NE Europe, to 5% in Estonians and close to zero in Latvians and Lithuanians. Indeed, Estonians show an excess of shared derived alleles with Nganasans compared to Latvians [<i>D</i>-statistic of the form D(Yorubas, Nganasans; Estonians, Latvians) = − 0.00263 (± 0.0008); <i>Z</i>-score = − 3.0691)] and Lithuanians [D(Yorubas, Nganasans; Estonians, Lithuanians) = − 0.00426 (± 0.0009); <i>Z</i>-score = − 5.6638)].</p><h3 class="c-article__sub-heading" id="Sec7">Correlation between linguistic, geographical and genetic data of Uralic speakers</h3><p>In order to determine whether and to what extent Uralic linguistic ancestry predicts genetic ancestry (see Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S12), we studied the correlations of genetic (autosomal, mtDNA and chrY, Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM15">15</a>: Table S14A-F), linguistic (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM15">15</a>: Table S14G) and geographical distances (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM15">15</a>: Table S14H) with Mantel [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 56" title="Mantel N. The detection of disease clustering and a generalized regression approach. Cancer Res. 1967;27:209–20. http://cancerres.aacrjournals.org/content/27/2_Part_1/209 " href="/articles/10.1186/s13059-018-1522-1#ref-CR56" id="ref-link-section-d8349793e2217">56</a>] and partial Mantel tests [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 57" title="Smouse PE, Long JC, Sokal RR. Multiple regression and correlation extensions of the mantel test of matrix correspondence. Syst Zool. 1986;35:627–32. https://doi.org/10.2307/2413122 " href="/articles/10.1186/s13059-018-1522-1#ref-CR57" id="ref-link-section-d8349793e2221">57</a>] (see the ‘M<a data-track="click" data-track-label="link" data-track-action="section anchor" href="/articles/10.1186/s13059-018-1522-1#Sec10">ethods</a>’ section for details). We used two types of autosomal distance matrices: <i>F</i><sub>ST</sub> distances and the fineSTRUCTURE coancestry based matrix using the data of shared chunk counts and two types of <i>F</i><sub>ST</sub>s with both mtDNA and chrY (six genetic distance matrices in total, Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM15">15</a>: Table S14).</p><p>Lexical distances between Uralic languages were significantly positively correlated with all types of genetic distances (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM16">16</a>: Table S15). Lexical distances also increased with geographical distances (<i>r</i> = 0.62, <i>p</i> = 0.001) as did all the genetic distances (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM16">16</a>: Table S15). When the effect of geographical distance was taken into account, lexical and autosomal distances still showed significant connections. For the fineSTRUCTURE-based distances, the correlation was twice stronger than for <i>F</i><sub>ST</sub>-based distances (<i>r</i> = 0.46, <i>p</i> = 0.001 vs <i>r</i> = 0.25, <i>p</i> = 0.01). This is consistent with the expectation that haplotype-based distances capture more recent signals of shared ancestry that are more relevant to recent history of language expansions. For mtDNA and chrY distances, correlation was not significant after correcting for geography. Thus, our findings indicate a clear relationship between autosomal genetic distances and lexical distances among Uralic-speaking populations, even when the effect of geographical distance is taken into account. The non-significant finding with respect to mtDNA and chrY data may reflect greater noise in these haploid loci. It is also worth noting that geographical distances significantly predict autosomal and chrY distances (but do not predict mtDNA distances) when keeping the lexical distance constant (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM16">16</a>: Table S15). This indicates that while lexical distance accounts for some of the variation in autosomal genetic distances between populations independent of geography, there remains genetic variation between groups that is attributable to geography independent of lexical distance—i.e. genetic variation is explained by a combination of lexical distance and geographic distance.</p></div></div></section><section data-title="Discussion"><div class="c-article-section" id="Sec8-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="Sec8">Discussion</h2><div class="c-article-section__content" id="Sec8-content"><p>In prehistoric times, with no written texts or alphabets, learning a language was only possible through direct contact and, hence, it is natural to expect that languages would often spread together with human migrations [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 58" title="Cavalli-Sforza LL, Menozzi P, Piazza A. The history and geography of human genes. Princeton: Princeton University Press. Xi; 1994. p. 541–18." href="/articles/10.1186/s13059-018-1522-1#ref-CR58" id="ref-link-section-d8349793e2282">58</a>]. There are several examples where indeed a large-scale migration of people has also resulted in the dispersal of new languages in a previously populated area [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" title="Tishkoff SA, Reed FA, Friedlaender FR, Ehret C, Ranciaro A, et al. The genetic structure and history of Africans and African Americans. Science (80- ). 2009;324:1035–44. https://doi.org/10.1126/science.1172257 " href="#ref-CR59" id="ref-link-section-d8349793e2285">59</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" title="Hudjashov G, Karafet TM, Lawson DJ, Downey S, Savina O, et al. Complex patterns of admixture across the Indonesian archipelago. Mol Biol Evol. 2017;34:2439–52. https://doi.org/10.1093/molbev/msx196 " href="#ref-CR60" id="ref-link-section-d8349793e2285_1">60</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 61" title="Leslie S, Winney B, Hellenthal G, Davison D, Boumertit A, et al. The fine-scale genetic structure of the British population. Nature. 2015;519:309–14. https://doi.org/10.1038/nature14230 " href="/articles/10.1186/s13059-018-1522-1#ref-CR61" id="ref-link-section-d8349793e2288">61</a>]. On the other hand, it has been shown that the spread of Turkic languages has at least in its westernmost reach been mediated by only a handful of migrants [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 1" title="Yunusbayev B, Metspalu M, Metspalu E, Valeev A, Litvinov S, et al. The genetic legacy of the expansion of Turkic-speaking nomads across Eurasia. PLoS Genet. 2015;11:e1005068. https://doi.org/10.1371/journal.pgen.1005068 " href="/articles/10.1186/s13059-018-1522-1#ref-CR1" id="ref-link-section-d8349793e2291">1</a>]. A more complex pattern of migration and admixture appears to be behind the ‘Central-East European’ supralinguistic genetic substratum characterising both East and West Slavic-IE speakers [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 62" title="Kushniarevich A, Utevska O, Chuhryaeva M, Agdzhoyan A, Dibirova K, et al. Genetic heritage of the Balto-Slavic speaking populations: a synthesis of autosomal, mitochondrial and Y-chromosomal data. PLoS One. 2015;10:e0135820. https://doi.org/10.1371/journal.pone.0135820 " href="/articles/10.1186/s13059-018-1522-1#ref-CR62" id="ref-link-section-d8349793e2294">62</a>].</p><p>Here, we have studied the genetic variation of 15 Uralic-speaking populations to reveal patterns that could correspond to the spatial distribution of the Uralic languages. Our analyses show that in the first approximation, the genetic diversity patterns of the Uralic speakers correspond to geography. Principal component and ADMIXTURE analyses suggest that the Uralic-speaking populations are genetically more similar to their neighbours than to geographically distant linguistic relatives. These analyses capture the broad-scale patterns of genetic variation arisen through the cumulative demographic processes in the population history of the continent. Importantly, ADMIXTURE analysis suggests a genetic component (k9) that is primarily present in most Uralic speakers (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig3">3</a>). Assuming that the spread of Uralic languages occurred within the past 5 kya, we next focused on haplotype sharing patterns between populations to concentrate on more recent demographic events. By inferring sharing of IBD segments between populations, we found the excess of shared IBD segments between most of the Uralic speakers (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig4">4</a>). This pattern is most notable for Uralic speakers in the Volga River basin who share more IBD segments with Uralic speakers both to the west and to the east of them than do their geographic neighbours.</p><p>Our Mantel test is consistent with the presence of common genetic substrate in most of the Uralic speakers (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM16">16</a>: Table S15): we found a significant association between autosomal genes and lexical variation that is independent of geographic proximity. This could be due to the legacy of ancient migrations shaping both genetic and linguistic diversity. Alternatively, the association could reflect a bias in gene flow between close linguistic relatives. The clear relationship between genetic and linguistic data does not extend to haploid markers, possibly because the latter are more prone to the effects of genetic drift. It has to be noted that this quantitative approach does not allow us to study the correlation between the spread of specific genetic and linguistic sub-lineages, for which a more precise case studies could provide information in the future.</p><p>We next used fineSTRUCTURE and Globetrotter approach to identify genetic clusters and admixture signals based on a wider spectrum of shared haplotypes. This approach does not depend on prior information on sample groupings and operates instead with data-driven natural groups defined by patterns of haplotype sharing. Most of the Finnic, Saami and VUR Uralic speakers form clusters in accordance with their self-reported linguistic affinity. These clusters are also distinct from the neighbouring Turkic speakers who form their own groupings. The exceptions here are for example the Mansi, who clearly form two clusters that differ in the extent of recent admixture with NE Europeans.</p><p>One of the notable observations that stands out in the fineSTRUCTURE analysis is that neither Hungarians nor Estonians or Mordovians form genetic clusters with other Uralic speakers but instead do so with a broad spectrum of geographically adjacent samples. Despite the documented history of the migration of Magyars [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 63" title="Róna-Tas A. Hungarians and Europe in the early middle ages: an introduction to early Hungarian history. Budapest: Central European University Press; 1999.https://doi.org/10.1017/S0067237800011413" href="/articles/10.1186/s13059-018-1522-1#ref-CR63" id="ref-link-section-d8349793e2319">63</a>] and their linguistic affinity to Khantys and Mansis, who today live east of the Ural Mountains, there is nothing in the present-day gene pool of the sampled Hungarians that we could tie specifically to other Uralic speakers. It is important to note here that our sample comes from the capital region. Given the complex history and ethnic makeup of Hungary, it is possible that a comprehensive sampling of the country could reveal genetic ties to the Ugric speakers. Furthermore, analyses of early medieval aDNA samples from Karos-Eperjesszög cemeteries in Hungary have revealed the presence of mtDNA haplogroups with East Asian provenance testifying for vestiges of a real migration of people from the east [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 64" title="Neparáczki E, Juhász Z, Pamjav H, Fehér T, Csányi B, et al. Genetic structure of the early Hungarian conquerors inferred from mtDNA haplotypes and Y-chromosome haplogroups in a small cemetery. Mol Gen Genomics. 2017;292:201–14. https://doi.org/10.1007/s00438-016-1267-z " href="/articles/10.1186/s13059-018-1522-1#ref-CR64" id="ref-link-section-d8349793e2322">64</a>].</p><p>Perhaps even more surprisingly, we found that Estonians, who show close affinities in IBD analysis to neighbouring Finnic speakers and Saami, do not share an excess of IBD segments with the VUR or Siberian Uralic speakers. This is even more striking considering that the immediate neighbours—Finns, Vepsians and Karelians—do. In this context, it is important to remind that the limited (5%, Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig6">6</a>) East Eurasian impact in the autosomal gene pool of modern Estonians contrasts with the fact that more than 30% of Estonian (but not Hungarian) men carry chrY N3 that has an East Eurasian origin and is very frequent among NE European Uralic speakers [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 36" title="Ilumäe A-M, Reidla M, Chukhryaeva M, Järve M, Post H, et al. Human Y chromosome haplogroup N: a non-trivial time-resolved phylogeography that cuts across language families. Am J Hum Genet. 2016;99:163–73. https://doi.org/10.1016/j.ajhg.2016.05.025 " href="/articles/10.1186/s13059-018-1522-1#ref-CR36" id="ref-link-section-d8349793e2331">36</a>]. However, the spread of chrY hg N3 is not language group specific as it shows similar frequencies in Baltic-speaking Latvians and Lithuanians, and in North Russians, who in all our analyses are very similar to Finnic-speakers. The latter, however, are believed to have either significantly admixed with their Uralic-speaking neighbours or have undergone a language shift from Uralic to Indo-European [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 38" title="Balanovsky O, Rootsi S, Pshenichnov A, Kivisild T, Churnosov M, et al. Two sources of the Russian patrilineal heritage in their Eurasian context. Am J Hum Genet. 2008;82:236–50. https://doi.org/10.1016/j.ajhg.2007.09.019 " href="/articles/10.1186/s13059-018-1522-1#ref-CR38" id="ref-link-section-d8349793e2334">38</a>].</p><p>Saami stand out from other NE European populations by drawing up to 30% of their autosomal ancestry from Asian genetic components (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig3">3</a>). They also display long-range genetic affinities with both the Uralic- and non-Uralic-speaking Siberians (Figs. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig4">4</a> and <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig5">5</a>). This is probably because the ancestors of the modern Saami (a) have lived in isolation from Southern and Eastern European gene flow and (b) have had more contacts with Nordic peoples on both sides of the Ural Mountains, driven by the similar life-style of the Arctic people. Curiously enough, the mtDNA heritage of Saami can be considered as predominantly (< 90%) Western Eurasian [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 30" title="Tambets K, Rootsi S, Kivisild T, Help H, Serk P, et al. The western and eastern roots of the Saami--the story of genetic “outliers” told by mitochondrial DNA and Y chromosomes. Am J Hum Genet. 2004;74:661–82. https://doi.org/10.1086/383203 " href="/articles/10.1186/s13059-018-1522-1#ref-CR30" id="ref-link-section-d8349793e2349">30</a>].</p><p>With some exceptions such as Estonians, Hungarians and Mordovians, both IBD sharing and Globetrotter results suggest that there are detectable inter-regional haplotype sharing ties between Uralic speakers from West Siberia and VUR, and between NE European Uralic speakers and VUR. In other words, there is a fragmented pattern of haplotype sharing between populations but no unifying signal of sharing that unite all the studied Uralic speakers.</p><p>Recent aDNA studies have shown that extant European populations draw ancestry form three main migration waves during the Upper Palaeolithic, the Neolithic and Early Bronze Age [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2" title="Haak W, Lazaridis I, Patterson N, Rohland N, Mallick S, et al. Massive migration from the steppe was a source for Indo-European languages in Europe. Nature. 2015; https://doi.org/10.1038/nature14317 " href="/articles/10.1186/s13059-018-1522-1#ref-CR2" id="ref-link-section-d8349793e2358">2</a>, <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 3" title="Allentoft ME, Sikora M, Sjögren K-G, Rasmussen S, Rasmussen M, et al. Population genomics of Bronze Age Eurasia. Nature. 2015;522:167–72. https://doi.org/10.1038/nature14507 " href="/articles/10.1186/s13059-018-1522-1#ref-CR3" id="ref-link-section-d8349793e2361">3</a>, <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 45" title="Lazaridis I, Patterson N, Mittnik A, Renaud G, Mallick S, et al. Ancient human genomes suggest three ancestral populations for present-day Europeans. Nature. 2014;513 https://doi.org/10.1038/nature13673 " href="/articles/10.1186/s13059-018-1522-1#ref-CR45" id="ref-link-section-d8349793e2364">45</a>]. The more detailed reconstructions concerning NE Europe up to the Corded Ware culture agree broadly with this scenario and reveal regional differences [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" title="Mittnik A, Wang CC, Pfrengle S, Daubaras M, Zariņa G, et al. The genetic prehistory of the Baltic Sea region. Nat. Commun. 2018;9:442. https://doi.org/10.1038/s41467-018-02825-9  " href="#ref-CR65" id="ref-link-section-d8349793e2367">65</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" title="Saag L, Varul L, Scheib CL, Stenderup J, Allentoft ME, et al. Extensive farming in Estonia started through a sex-biased migration from the steppe. Curr Biol. 2017;27:2185–93. https://doi.org/10.1016/j.cub.2017.06.022 " href="#ref-CR66" id="ref-link-section-d8349793e2367_1">66</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 67" title="Jones ER, Zarina G, Moiseyev V, Lightfoot E, Nigst PR, et al. The Neolithic transition in the Baltic was not driven by admixture with early European farmers. Curr Biol. 2017;27:576–82. https://doi.org/10.1016/j.cub.2016.12.060 " href="/articles/10.1186/s13059-018-1522-1#ref-CR67" id="ref-link-section-d8349793e2370">67</a>]. However, to explain the demographic history of extant NE European populations, we need to invoke a novel genetic component in Europe—the Siberian. The geographic distribution of the main part of this component is likely associated with the spread of Uralic speakers but gene flow from Siberian sources in historic and modern Uralic speakers has been more complex, as revealed also by a recent study of ancient DNA from Fennoscandia and Northwest Russia [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 68" title="Lamnidis TC, Majander K, Jeong C, Salmela E, Wessman A, et al. (2018) Ancient Fennoscandian genomes reveal origin and spread of Siberian ancestry in Europe. bioRxiv. https://doi.org/10.1101/285437 ." href="/articles/10.1186/s13059-018-1522-1#ref-CR68" id="ref-link-section-d8349793e2374">68</a>]. Thus, the Siberian component we introduce here is not the perfect but still the current best candidate for the genetic counterpart in the spread of Uralic languages. On the westernmost reach of Uralic speakers, the extent of this shared ancestry is, however, small on the genome level and is significantly sex-specific in its nature. The shared ancestry is clearly pronounced in chrY, with Uralic speakers showing distinctively high (29% on average) frequency of hg N3-M178. The tested Uralic-speaking populations show marginally, though significantly, higher affinity to populations with high frequency of N3-M178 in the autosomal loci than predicted from their X chromosomal similarity and their comparison to other populations where N3-M178 is infrequent or absent. These sex-specific differences which are widely spread among Uralic speakers today may trace their origins back to the time of the shared population history of the Uralic populations and reflect complex socio-cultural factors amplified by small effective population sizes, potentially including examples such as male-specific <i>elite dominance</i> and/or cultural inheritance of male reproductive fitness [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 34" title="Karmin M, Saag L, Vicente M, Sayres MAW, Järve M, et al. A recent bottleneck of Y chromosome diversity coincides with a global change in culture. Genome Res. 2015;25:459–66. https://doi.org/10.1101/gr.186684.114 " href="/articles/10.1186/s13059-018-1522-1#ref-CR34" id="ref-link-section-d8349793e2380">34</a>, <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 69" title="Zeng TC, Aw AJ, Feldman MW. Cultural hitchhiking and competition between patrilineal kin groups explain the post-Neolithic Y-chromosome bottleneck. Nat Commun. 2018;9:2077. https://doi.org/10.1038/s41467-018-04375-6 " href="/articles/10.1186/s13059-018-1522-1#ref-CR69" id="ref-link-section-d8349793e2383">69</a>] during the time of their dispersal and admixture with neighbouring groups.</p><p>Understanding the interplay between the cultural and demographic processes leading to these observations will, no doubt, motivate future studies, especially those that will be done in the field of ancient genomics.</p></div></div></section><section data-title="Conclusions"><div class="c-article-section" id="Sec9-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="Sec9">Conclusions</h2><div class="c-article-section__content" id="Sec9-content"><p>Here, we present for the first time the comparison of genome-wide genetic variation of nearly all extant Uralic-speaking populations from Europe and Siberia. We show that (1) the Uralic speakers are genetically most similar to their geographical neighbours; (2) nevertheless, most Uralic speakers along with some of their geographic neighbours share a distinct ancestry component of likely Siberian origin. Furthermore, (3) most geographically distant Uralic speaking populations share more genomic IBD segments with each other than with equidistant populations speaking other languages and (4) there is a positive correlation between linguistic and genetic data of the Uralic speakers. This suggests that the spread of the Uralic languages was at least to some degree associated with movement of people. Moreover, the discovery of the Siberian component shows that the three known major components of genetic diversity in Europe (European hunter-gatherers, early Neolithic farmers and the Early Bronze Age steppe people) are not enough to explain the extant genetic diversity in (northeast) Europe.</p></div></div></section><section data-title="Methods"><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">Methods</h2><div class="c-article-section__content" id="Sec10-content"><h3 class="c-article__sub-heading" id="Sec11">Linguistic background and geographical location of the samples</h3><p>Approximately 20.5 million people speak Uralic languages today [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 10" title="Lewis MP, Simons GF, Fennig CD. Ethnologue: languages of the world. 19th ed. Dallas: SIL International; 2016." href="/articles/10.1186/s13059-018-1522-1#ref-CR10" id="ref-link-section-d8349793e2409">10</a>] (see details in Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM2">2</a>: Table S2), and only three of the Uralic languages—Hungarian, Finnish and Estonian (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig1">1</a>a, Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM2">2</a>: Table S2)—are not listed as endangered in the <i>UNESCO Atlas of the World’s Languages in Danger</i> [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 70" title="Moseley C, editor. Atlas of the world’s languages in danger. 3rd ed. Paris: UNESCO Publishing; 2010." href="/articles/10.1186/s13059-018-1522-1#ref-CR70" id="ref-link-section-d8349793e2425">70</a>]<i>.</i> In this study, DNA samples of a total of 15 Uralic-speaking populations from Europe and Western Siberia were collected from the present-day spread area of corresponding Uralic languages (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig1">1</a>, Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM2">2</a>: Table S2). The population affiliations of the samples were derived from the reported self-identity of the volunteers. We assume that these affiliations reflect also the language they speak as their mother tongue. The samples used here do not encompass all extant Uralic languages, but represent examples of each of the main branches of the family, and cover the whole distribution area.</p><p>Due to the small sample sizes and genetic homogeneity revealed by the genetic profiles on the ADMIXTURE plot of some of the studied populations, we pooled the samples of Erzas and Mokshas together as Mordovians and the samples of Permyak and Zyryan Komis as Komi. The heterogeneous Mansi population was divided into two to three subsets, according to the proportions of Eastern and Western Eurasian ancestry components in their genetic profiles. The Finnish group consists of Finns and Ingrian Finns who have been analysed separately in the analyses of PCA, <i>F</i><sub>ST</sub> distance calculations, ADMIXTURE and <i>D</i>-statistics.</p><p>DNA of the samples was extracted from whole blood according to the phenol/chloroform method [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 71" title="Miller SA, Dykes DD, Polesky HF. A simple salting out procedure for extracting DNA from human nucleated cells. Nucleic Acids Res. 1988;16:1215. https://doi.org/10.1093/nar/16.3.1215 " href="/articles/10.1186/s13059-018-1522-1#ref-CR71" id="ref-link-section-d8349793e2450">71</a>]. DNA concentrations were determined with spectrophotometry (NanoDrop, Wilmington).</p><p>A total of 286 samples of Uralic-speaking individuals, of those 121 genotyped in this study, were analysed in the context of 1514 Eurasian samples (including 14 samples published for the first time) based on whole genome single nucleotide polymorphisms (SNPs) (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM1">1</a>: Table S1). All these samples, together with the larger sample set of Uralic speakers, were characterized for mtDNA and chrY markers.</p><h3 class="c-article__sub-heading" id="Sec12">Population structure and admixture</h3><p>A total of 135 samples from this study were genotyped using the Illumina 610K, 650K, 660K or 1M SNPs arrays (Human610-Quad, HumanHap650Y, Human660W-Quad or HumanOmni1-Quad BeadChip DNA Analysis BeadChip Kits) and analysed for the whole-genome variation together with published genotype data (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM1">1</a>: Table S1). We used the software PLINK 1.05 [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 72" title="Purcell S, Neale B, Todd-Brown K, Thomas L, Ferreira MA, et al. PLINK: a tool set for whole-genome association and population-based linkage analyses. Am J Hum Genet. 2007;81:559–75. https://doi.org/10.1086/519795 " href="/articles/10.1186/s13059-018-1522-1#ref-CR72" id="ref-link-section-d8349793e2470">72</a>] to filter the dataset and to include only SNPs of autosomal chromosomes with minor allele frequency > 0.1% and genotyping success > 97%. We excluded SNPs in strong linkage disequilibrium (LD) (pairwise genotypic correlation <i>r</i><sup>2</sup> > 0.4) in a window of 200 SNPs (sliding the window by 25 SNPs at a time), due to the possible effect of background LD on PCA and structure-like analysis. To exclude possible close relative pairs (first and second degree) among the individuals, the software KING v1.4 [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 73" title="Manichaikul A, Mychaleckyj JC, Rich SS, Daly K, Sale M, et al. Robust relationship inference in genome-wide association studies. Bioinformatics. 2010;26:2867–73. https://doi.org/10.1093/bioinformatics/btq559 " href="/articles/10.1186/s13059-018-1522-1#ref-CR73" id="ref-link-section-d8349793e2477">73</a>] was applied to the entire dataset and the resulting data were confirmed by REAP v.1.2 [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 74" title="Thornton T, Tang H, Hoffmann TJ, Ochs-Balcom HM, Caan BJ, et al. Estimating kinship in admixed populations. Am J Hum Genet. 2012;91:122–38. https://doi.org/10.1016/j.ajhg.2012.05.024 " href="/articles/10.1186/s13059-018-1522-1#ref-CR74" id="ref-link-section-d8349793e2480">74</a>]. The samples (populations and no. of individuals) used for different analysis are given in Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM1">1</a>: Table S1.</p><h4 class="c-article__sub-heading c-article__sub-heading--small" id="Sec13">PCA and <i>F</i> <sub>ST</sub> </h4><p>PCA (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig2">2</a>a and Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S1) was carried out with the smartpca program of the EIGENSOFT package [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 75" title="Patterson N, Price AL, Reich D. Population structure and eigenanalysis. PLoS Genet. 2006;2:e190. https://doi.org/10.1371/journal.pgen.0020190 " href="/articles/10.1186/s13059-018-1522-1#ref-CR75" id="ref-link-section-d8349793e2506">75</a>], using 171,454 SNPs. Mean pairwise <i>F</i><sub>ST</sub> values between populations and regional population groups for 303,671 autosomal SNPs (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S2) were calculated with the method of Weir and Cockerman [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 76" title="Weir BS, Cockerham CC. Estimating F-statistics for the analysis of population structure. Evolution (N Y). 1984;38:1358–70. https://doi.org/10.2307/2408641 " href="/articles/10.1186/s13059-018-1522-1#ref-CR76" id="ref-link-section-d8349793e2517">76</a>] as in Metspalu et al. [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 77" title="Metspalu M, Romero IG, Yunusbayev B, Chaubey G, Mallick CB, et al. Shared and unique components of human population structure and genome-wide signals of positive selection in South Asia. Am J Hum Genet. 2011;89:731–44. https://doi.org/10.1016/j.ajhg.2011.11.010 " href="/articles/10.1186/s13059-018-1522-1#ref-CR77" id="ref-link-section-d8349793e2520">77</a>]. Only populations with <i>n</i> > 4 were included in <i>F</i><sub>ST</sub> calculations. The UPGMA tree that visualizes the clustering based on the genetic distances of studied population was built with MEGA7 [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 78" title="Kumar S, Stecher G, Tamura K. MEGA7: molecular evolutionary genetics analysis version 7.0 for bigger datasets. Mol Biol Evol. 2016;33:1870–4. https://doi.org/10.1093/molbev/msw054 " href="/articles/10.1186/s13059-018-1522-1#ref-CR78" id="ref-link-section-d8349793e2530">78</a>] (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig2">2</a>b).</p><h4 class="c-article__sub-heading c-article__sub-heading--small" id="Sec14">ADMIXTURE</h4><p>The population structure was analysed using the software ADMIXTURE [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 79" title="Alexander DH, Novembre J, Lange K. Fast model-based estimation of ancestry in unrelated individuals. Genome Res. 2009;19:1655–64. https://doi.org/10.1101/gr.094052.109 " href="/articles/10.1186/s13059-018-1522-1#ref-CR79" id="ref-link-section-d8349793e2545">79</a>] implementing a structure-like [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 80" title="Pritchard JK, Stephens M, Donnelly P. Inference of population structure using multilocus genotype data. Genetics. 2000;155:945–59. http://www.genetics.org/content/155/2/945 " href="/articles/10.1186/s13059-018-1522-1#ref-CR80" id="ref-link-section-d8349793e2548">80</a>] model-based maximum likelihood clustering algorithm (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig3">3</a> and Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S3). The final dataset of ADMIXTURE analysis of Uralic-speaking populations in the general Eurasian background consisted of 181,005 SNPs and 1800 individuals from 111 populations (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM1">1</a>: Table S1). We ran ADMIXTURE 100 times for each assumed number of ancestral populations (<i>K</i>) from <i>K</i> = 3 to <i>K</i> = 20 to observe the deviation of the results between individual runs (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S3A). According to a low level of variation in log likelihood scores (LLs < 1) within the top 10% fraction of runs with the highest LLs [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 81" title="Rasmussen M, Li Y, Lindgreen S, Pedersen JS, Albrechtsen A, et al. Ancient human genome sequence of an extinct Palaeo-Eskimo. Nature. 2010;463:757–62. https://doi.org/10.1038/nature08835 " href="/articles/10.1186/s13059-018-1522-1#ref-CR81" id="ref-link-section-d8349793e2573">81</a>], we assume that usable results were at <i>K</i> = 3 to <i>K</i> = 18 and the best fit <i>K</i> value appears on K10 level (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S3). We use letter <i>k</i> to refer to the specific components in the genetic profiles of individuals/populations. The frequency of k9 component from Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM7">7</a>: Table S6B was spatially mapped (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig3">3</a>b) with Surfer software (v7, Golden Software, Inc.).</p><h4 class="c-article__sub-heading c-article__sub-heading--small" id="Sec15"> <i>D</i>-statistics</h4><p>We calculated <i>D</i>-statistics [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 49" title="Reich D, Thangaraj K, Patterson N, Price AL, Singh L. Reconstructing Indian population history. Nature. 2009;461:489–94. https://doi.org/10.1038/nature08365 " href="/articles/10.1186/s13059-018-1522-1#ref-CR49" id="ref-link-section-d8349793e2613">49</a>] (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S4A-R) from tests of four populations in the form of (A, B; C, D), where A is an outgroup, B is a test population, C is an Uralic-speaking population and D is a non-Uralic-speaking population, for a suggested tree-like population history as in Skoglund et al. [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 50" title="Skoglund P, Malmström H, Omrak A, Raghavan M, Valdiosera C, et al. Genomic diversity and admixture differs for Stone-Age Scandinavian foragers and farmers. Science. 2014;344:747–50. https://doi.org/10.1126/science.1253448 " href="/articles/10.1186/s13059-018-1522-1#ref-CR50" id="ref-link-section-d8349793e2619">50</a>]. The test provides information on whether or not the test population (B) shares more derived alleles with one population from a pair (C, D) than is expected from the process of incomplete lineage sorting without admixture, indicating a recent gene flow between B and C or B and D. If <i>D</i> < 0, a test population (B) shares more derived alleles with the Uralic-speaking population (C) compared to the non-Uralic-speaking population (D); if <i>D</i> > 0, a test population (B) shares more derived alleles with the non-Uralic-speaking population (D) compared to the Uralic-speaking population (C). We used Yorubas as an outgroup (A) and one out of four westernmost Uralic-speaking populations: Saami from Sweden, Finns, Estonians and Hungarians; and one out of three European non-Uralic speakers: French (representing West Europeans), Poles (representing East Europeans) and Swedes (representing North Europeans) as a fixed pair (C, D). We ran the <i>D</i>-statistics test (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S4A-L) with a list of European and Siberian populations (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM1">1</a>: Table S1) used as the test population (B). We also ran the test using the local geographical neighbours of European Uralic speakers (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S4M-R). The null hypothesis was that there is no excess share of derived alleles between the Uralic-speaking populations and test population B (<i>D</i> = 0). Only the <i>D</i>-values with |<i>Z</i>-score| > 3 were considered significant.</p><h4 class="c-article__sub-heading c-article__sub-heading--small" id="Sec16">Analysis of IBD segment sharing</h4><p>We used the fastIBD algorithm implemented in the BEAGLE 3.3 software [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 51" title="Browning BL, Browning SR. A fast, powerful method for detecting identity by descent. Am J Hum Genet. 2011;88:173–82. https://doi.org/10.1016/j.ajhg.2011.01.010 " href="/articles/10.1186/s13059-018-1522-1#ref-CR51" id="ref-link-section-d8349793e2660">51</a>] to detect chromosomal tracts (> 1 cM in length) that are IBD between pairs of individuals (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig4">4</a>, Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM8">8</a>: Table S7, Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM9">9</a>: Table S8, Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM10">10</a>: Table S9). IBD tracts with a fastIBD score of 1e−10 from ten independent runs were further post-processed using the algorithm developed by Ralph and Coop 2013 [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 82" title="Ralph P, Coop G. The geography of recent genetic ancestry across Europe. PLoS Biol. 2013;11:e1001555. https://doi.org/10.1371/journal.pbio.1001555 " href="/articles/10.1186/s13059-018-1522-1#ref-CR82" id="ref-link-section-d8349793e2676">82</a>]. This algorithm removes spurius gaps, breaks introduced into long blocks of IBD by low marker density and phasing switch-error and performs final IBD tract calls. IBD tracts were first sorted into bins (classes) based on their length: 1–2, 2–3, 3–4 and 4–5 cM. Within each bin, we computed average length of IBD (sum of all tracts divided by sample size) between randomly chosen pairs of subsamples from two populations. We then tested whether Uralic speakers from different regions demonstrate more IBD sharing between each other. For this, we split populations in our dataset into three regional groups: 1—Baltic-Scandinavian; 2—Eastern European-Volga; and 3—Siberian. We then computed average IBD sharing between Uralic speakers from two different regions. Next, within each region, for each tested Uralic-speaking population, we selected non-Uralic-speaking populations that are geographically close to them, as a control group. In order, to assess, for example, whether Finns from the Baltic-Scandinavian region have higher than expected IBD sharing with Udmurts, we compared observed IBD sharing with values characteristic of the control group populations. Namely, we took multiple random samples from the pooled set of control for Finns and computed IBD sharing with Udmurts and compared it with the observed value. Given no recent shared ancestry between Finns and Udmurts due to linguistic relatedness, Finns are expected to show the same level of IBD sharing as their control group. IBD sharing values higher than background were counted to compute a <i>p</i> value. We note that for some populations, we do not have an appropriate geographic control group to carry out this kind of permutation test. Nevertheless, most of the Uralic peoples tested show higher IBD sharing with distant Uralic speakers compared to their regional non-Uralic control, and this suggests higher number of shared ancestors between Uralic speakers within the past dozens of generations.</p><h4 class="c-article__sub-heading c-article__sub-heading--small" id="Sec17">fineSTRUCTURE and GLOBETROTTER</h4><p>While ADMIXTURE uses independent unlinked SNPs for reconstructing individual ancestries, fineSTRUCTURE is a much more powerful approach which infers fine-scale population structure from haplotype data. Each individual is presented as a matrix of non-recombining genomic chunks received from a set of multiple donor individuals. The patterns of similarities between these copying matrices are then used to cluster individuals into genetic groups using the Bayesian approach (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig5">5</a>, Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figures S5-S6). This multistage process included the following steps:</p><ol class="u-list-style-none"> <li> <span class="u-custom-list-number">(a)</span> <p>First, we phased the data with SHAPEIT v.2 [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 83" title="Delaneau O, Zagury J-F, Marchini J. Improved whole-chromosome phasing for disease and population genetic studies. Nat Methods. 2013;10:5–6. https://doi.org/10.1038/nmeth.2307 " href="/articles/10.1186/s13059-018-1522-1#ref-CR83" id="ref-link-section-d8349793e2706">83</a>], using the HapMap phase II b37 genetic map;</p> </li> <li> <span class="u-custom-list-number">(b)</span> <p>We performed population assignments (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM1">1</a>: Table S1) to genetic groups (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S5) using fineSTRUCTURE v.2 [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 53" title="Lawson DJ, Hellenthal G, Myers S, Falush D. Inference of population structure using dense haplotype data. PLoS Genet. 2012;8:e1002453. https://doi.org/10.1371/journal.pgen.1002453 " href="/articles/10.1186/s13059-018-1522-1#ref-CR53" id="ref-link-section-d8349793e2726">53</a>]. We estimated initial Ne and θ parameters using 10% of the samples and 10 Expectation-Maximization steps of the algorithm. Next, we described each individual recipient chromosome as a mixture of genetic chunks from the set of all other individuals (donors) using the estimated demographic parameters;</p> </li> <li> <span class="u-custom-list-number">(c)</span> <p>We used a matrix of the copying vectors generated in the previous step to cluster the individuals using the Bayesian algorithm. We performed two parallel runs and assessed convergence between them using Gelman-Rubin statistics, as implemented in the software. Convergence was reached after 35 million MCMC iterations, including the first three million iterations, which we discarded as burn-in;</p> </li> <li> <span class="u-custom-list-number">(d)</span> <p>Finally, we performed the tree-building step using default settings and used the run with the highest observed posterior likelihood to cluster the samples into genetic groups. We inspected the population dendrogram manually and assigned samples to individual groups. We excluded a few outlying samples, which showed evidence for very recent genetic admixture or incorrect population identification, from further admixture inference with Globetrotter (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S5, Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM1">1</a>: Table S1).</p> </li> </ol><p>After assigning individual samples into natural genetic groups, we performed two types of Globetrotter analysis (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig5">5</a>), following the guidelines as in Hellenthal et al. [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 54" title="Hellenthal G, Busby GBJ, Band G, Wilson JF, Capelli C, et al. A genetic atlas of human admixture history. Science (80- ). 2014;343:747–51. https://doi.org/10.1126/science.1243518 " href="/articles/10.1186/s13059-018-1522-1#ref-CR54" id="ref-link-section-d8349793e2766">54</a>]. First, in the ‘full’ analysis (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM11">11</a>: Table S10), we allowed the recipient individual to copy from every donor population, except from its own population label (self-copying). Second, in the ‘regional’ analysis (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM11">11</a>: Table S10), we grouped the genetic clusters identified by fineSTRUCTURE into three geographic regions: Europe, VUR and Western Siberia. We allowed no self-copying within regional groups (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM12">12</a>: Table S11 and see Note therein). In both analyses, we used additional donors from outside of the populations of interest to describe genetic ancestry, but we did not perform admixture inference for them. These included ‘W-S-Europe’, ‘Cauc/N-East’, ‘N-Cauc/C-Asia’, ‘C-Asia/S-Sib’, "E-Asia/S-Sib", ‘Far East’, ‘W-C-Sib’, ‘Sib 1’ and ‘Sib 2’ (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S5, Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM12">12</a>: Table S11). For all, except the ‘Nganassan’ group, we grouped all donors from Western and Central Siberia to form the ‘W-C-Sib’ cluster. For the admixture analysis of Nganasan samples, we split the ‘W-C-Sib’ cluster into ‘Sib 1’, ‘Sib 2’ and ‘Nganassan’ and excluded the latter from the generation of copying vectors to deny self-copying as explained above (see Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM12">12</a>: Table S11 for details).</p><h4 class="c-article__sub-heading c-article__sub-heading--small" id="Sec18">ALDER analysis</h4><p>We used a method based on the decay of admixture linkage disequilibrium (LD) implemented in the ALDER v1.03 software [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 84" title="Loh P-R, Lipson M, Patterson N, Moorjani P, Pickrell JK, et al. Inferring admixture histories of human populations using linkage disequilibrium. Genetics. 2013;193:1233–54. https://doi.org/10.1534/genetics.112.147330 " href="/articles/10.1186/s13059-018-1522-1#ref-CR84" id="ref-link-section-d8349793e2796">84</a>] to test and date the admixture signal in contemporary Northern Eurasian populations (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S7). We tested all population triplets in our dataset with pre-set ALDER v1.03 parameters (‘multiple admixture tests’ mode). We report the admixed populations with their pairs of reference populations and their inferred admixture timeframe that passed all the pre-test steps had significant <i>p</i> values and highest two-reference weighted LD curve amplitude, presenting only triplets with consistent LD decay rates if possible (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM13">13</a>: Table S12). Exceptions include Finns, Swedish Saami, Vepsians and Khanty whose decay time constants for all reference populations disagree by more than 25% (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM13">13</a>: Table S12), which may stem from bottlenecks in their demographic history [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 84" title="Loh P-R, Lipson M, Patterson N, Moorjani P, Pickrell JK, et al. Inferring admixture histories of human populations using linkage disequilibrium. Genetics. 2013;193:1233–54. https://doi.org/10.1534/genetics.112.147330 " href="/articles/10.1186/s13059-018-1522-1#ref-CR84" id="ref-link-section-d8349793e2812">84</a>].</p><h4 class="c-article__sub-heading c-article__sub-heading--small" id="Sec19">Outgroup <i>f</i>3-statistics</h4><p>We performed <i>f</i>3 analysis of our modern and published ancient human genotyping data with the AdmixTools v3.0 software package [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 48" title="Patterson N, Moorjani P, Luo Y, Mallick S, Rohland N, et al. Ancient admixture in human history. Genetics. 2012;192:1065–93. https://doi.org/10.1534/genetics.112.145037 " href="/articles/10.1186/s13059-018-1522-1#ref-CR48" id="ref-link-section-d8349793e2830">48</a>]. The outgroup <i>f</i>3-statistic (outgroup; X, Y) is a function of shared branch length between X and Y in the absence of admixture with the outgroup [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 44" title="Raghavan M, Skoglund P, Graf KE, Metspalu M, Albrechtsen A, et al. Upper Palaeolithic Siberian genome reveals dual ancestry of Native Americans. Nature. 2014;505:87–91. https://doi.org/10.1038/nature12736 " href="/articles/10.1186/s13059-018-1522-1#ref-CR44" id="ref-link-section-d8349793e2836">44</a>]. We used Yorubas as an outgroup to non-African populations and computed <i>f</i>3-statistics in the form of (Yorubas; ancient group, modern group) to investigate the shared history of a set of 47 European, Siberian and East Asian populations, including the Uralic speakers and 216 ancient genomes (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM1">1</a>: Table S1). We first prepared our modern dataset by excluding all positions with less than 3% genotyping success rate, and A/T and C/G polymorphisms to minimize potential strand mismatch problems. We extracted genotype information of 522,274 SNPs, which passed the filtering criteria, from the ancient DNA dataset of Mathieson et al. [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 85" title="Mathieson I, Lazaridis I, Rohland N, Mallick S, Patterson N, et al. Genome-wide patterns of selection in 230 ancient Eurasians. Nature. 2015;528:499–503. https://doi.org/10.1038/nature16152 " href="/articles/10.1186/s13059-018-1522-1#ref-CR85" id="ref-link-section-d8349793e2846">85</a>]. We divided ancient samples further into groups according to their cultural background as in the source article [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 85" title="Mathieson I, Lazaridis I, Rohland N, Mallick S, Patterson N, et al. Genome-wide patterns of selection in 230 ancient Eurasians. Nature. 2015;528:499–503. https://doi.org/10.1038/nature16152 " href="/articles/10.1186/s13059-018-1522-1#ref-CR85" id="ref-link-section-d8349793e2849">85</a>] (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM1">1</a>: Table S1) and merged the modern and ancient datasets. We performed an outgroup <i>f</i>3 test on all pairwise combinations between ancient and modern groups (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S8 and Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM14">14</a>: Table S13).</p><p>To allow for chrX versus autosomes comparison, outgroup <i>f</i>3 statistics of the form <i>f</i>3(Yorubas; test population, comparison population) were computed with Uralic-speaking populations and their geographical neighbours as test populations, and all European and Siberian populations from the EBC Illumina dataset as comparison populations. The analyses (data shown for Estonians, Udmurts and Khantys in Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S13) were run both using the same autosomal SNPs as for qpGraph (see below, Illumina chip dataset filtered by missing genotypes and minor allele frequency; 511,602 SNPs) and also all chrX positions available in the filtered dataset (12,547 SNPs). Since all individuals inherit half of their autosomal material from their father but only females inherit their chrX from their father, then in this comparison, chrX data gives more information about the female and autosomal data about the male ancestors of a population. Considering that chrY hg N3-M178 has a distinctively high frequency in Uralic-speaking populations, we used a summary statistic p(M178-coA), the probability for a pair of men sampled from two different populations to share their chrY ancestry in hg N3-M178 (calculated by multiplying hg frequencies for compared population pairs, data from Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM6">6</a>: Table S5). This data was added to plots opposing the chrX and autosomal outgroup <i>f</i>3 results of the above-mentioned test populations to see whether those results also reflect the differences observed between chrY and mtDNA affinities among the populations are reflected also in the chrX and autosomal data (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S13). The significance of the slope and interception of the regression lines of high (> 10%) and low (< 5%) M178-co-ancestry groups under a linear model was tested with ANOVA in R, using the car package [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 86" title="Fox J, Weisberg S. An R companion to applied regression. 2nd ed: Sage Publications; 2011. https://socialsciences.mcmaster.ca/jfox/Books/Companion/ " href="/articles/10.1186/s13059-018-1522-1#ref-CR86" id="ref-link-section-d8349793e2887">86</a>].</p><h4 class="c-article__sub-heading c-article__sub-heading--small" id="Sec20">qpGraph</h4><p>We ran the <i>qpGraph</i> software v6.5 of the AdmixTools v4.1 package [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 48" title="Patterson N, Moorjani P, Luo Y, Mallick S, Rohland N, et al. Ancient admixture in human history. Genetics. 2012;192:1065–93. https://doi.org/10.1534/genetics.112.145037 " href="/articles/10.1186/s13059-018-1522-1#ref-CR48" id="ref-link-section-d8349793e2901">48</a>] on a merged dataset of modern and ancient data. To merge the two datasets, we extracted the 511,602 SNPs present in the quality filtered Illumina chip data from a dataset containing ancient samples from Lazaridis et al. [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 87" title="Lazaridis I, Nadel D, Rollefson G, Merrett DC, Rohland N, et al. Genomic insights into the origin of farming in the ancient near east. Nature. 2016;536:419–24. https://doi.org/10.1038/nature19310 " href="/articles/10.1186/s13059-018-1522-1#ref-CR87" id="ref-link-section-d8349793e2904">87</a>], Jones et al. [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 67" title="Jones ER, Zarina G, Moiseyev V, Lightfoot E, Nigst PR, et al. The Neolithic transition in the Baltic was not driven by admixture with early European farmers. Curr Biol. 2017;27:576–82. https://doi.org/10.1016/j.cub.2016.12.060 " href="/articles/10.1186/s13059-018-1522-1#ref-CR67" id="ref-link-section-d8349793e2907">67</a>] and Saag et al. [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 66" title="Saag L, Varul L, Scheib CL, Stenderup J, Allentoft ME, et al. Extensive farming in Estonia started through a sex-biased migration from the steppe. Curr Biol. 2017;27:2185–93. https://doi.org/10.1016/j.cub.2017.06.022 " href="/articles/10.1186/s13059-018-1522-1#ref-CR66" id="ref-link-section-d8349793e2910">66</a>] resulting in a genotyping rate of 0.4. Only samples with at least 100,000 SNPs covered were used in the analysis. We used qpGraph with default settings, with Yorubas as an outgroup, with the useallsnps = YES option, retaining 362,380 SNPs. We were able to fit the demographic model with our data (<i>f</i>2-statistics`|<i>Z</i>-score| < 3) when we modelled ancient and modern European populations through several admixture events shown in Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig6">6</a> (see Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S10 for details). Of the tested Uralic-speaking populations, only Hungarians did not fit into the model.</p><h3 class="c-article__sub-heading" id="Sec21">MtDNA and Y chromosomal variation</h3><p>We present new genotype data of 1578 mtDNAs and 994 chrY of Uralic-speaking individuals, which include also all those individuals genotyped for autosomal markers. MtDNA hgs were determined by genotyping the variation of the first hypervariable segment (HVS-I) and coding region markers of mtDNA (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM4">4</a>: Table S3). The PCR-amplified probes were examined by RFLP or direct sequencing. The classification of mtDNA hgs follows the present nomenclature of the Global Human Mitochondrial DNA Phylogenetic Tree (mtDNA tree Build 17) [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 88" title="Van Oven M, Kayser M. Updated comprehensive phylogenetic tree of global human mitochondrial DNA variation. Hum Mutat. 2009;30:386–94. https://doi.org/10.1002/humu.20921 " href="/articles/10.1186/s13059-018-1522-1#ref-CR88" id="ref-link-section-d8349793e2938">88</a>]. The samples studied for chrY variation were genotyped for 18 NRY SNP markers at minimal, analysed by PCR/AFLP, PCR/RFLP or PCR/sequencing. The hg designation follows common nomenclature [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 34" title="Karmin M, Saag L, Vicente M, Sayres MAW, Järve M, et al. A recent bottleneck of Y chromosome diversity coincides with a global change in culture. Genome Res. 2015;25:459–66. https://doi.org/10.1101/gr.186684.114 " href="/articles/10.1186/s13059-018-1522-1#ref-CR34" id="ref-link-section-d8349793e2941">34</a>, <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 89" title="Karafet TM, Mendez FL, Meilerman MB, Underhill PA, Zegura SL, et al. New binary polymorphisms reshape and increase resolution of the human Y chromosomal haplogroup tree. Genome Res. 2008;18:830–8. https://doi.org/10.1101/gr.7172008 " href="/articles/10.1186/s13059-018-1522-1#ref-CR89" id="ref-link-section-d8349793e2944">89</a>, <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 90" title="Van Oven M, Van Geystelen A, Kayser M, Decorte R, Larmuseau MHD. Seeing the wood for the trees: a minimal reference phylogeny for the human Y chromosome. Hum Mutat. 2014;35:187–91. https://doi.org/10.1002/humu.22468 " href="/articles/10.1186/s13059-018-1522-1#ref-CR90" id="ref-link-section-d8349793e2947">90</a>]. The hg frequencies for mtDNA and chrY were calculated and presented in a context of published data of 12,157 mtDNA (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM5">5</a>: Table S4) and 9730 chrY (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM6">6</a>: Table S5). A subset of the samples from the chrY hg Q was analysed for markers M346 and L54 [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 91" title="Dulik MC, Zhadanov SI, Osipova LP, Askapuli A, Gau L, et al. Mitochondrial DNA and Y chromosome variation provides evidence for a recent common ancestry between Native Americans and indigenous Altaians. Am J Hum Genet. 2012;90:229–46. https://doi.org/10.1016/j.ajhg.2011.12.014 " href="/articles/10.1186/s13059-018-1522-1#ref-CR91" id="ref-link-section-d8349793e2957">91</a>] (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM17">17</a>: Table S16, Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S11) and a subset of the Selkup samples from hg R1a for marker M458.</p><h3 class="c-article__sub-heading" id="Sec22">Correlation analysis</h3><h4 class="c-article__sub-heading c-article__sub-heading--small" id="Sec23">Linguistic data and lexical distances</h4><p>To measure linguistic distances, we first created a Uralic family tree (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM3">3</a>: Figure S12A) by using Uralic basic vocabulary data and cognate coding as described in Syrjänen et al. [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 16" title="Syrjänen K, Honkola T, Korhonen K, Lehtinen J, Vesakoski O, et al. Shedding more light on language classification using basic vocabularies and phylogenetic methods. A case study of Uralic. Diachronica. 2013;30:323–52. https://doi.org/10.1075/dia.30.3.02syr " href="/articles/10.1186/s13059-018-1522-1#ref-CR16" id="ref-link-section-d8349793e2981">16</a>] and Lehtinen et al. [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 92" title="Lehtinen J, Honkola T, Korhonen K, Syrjänen K, Wahlberg N, et al. Behind family trees: secondary connections in Uralic language networks. Lang Dyn Chang. 2014;4:189–221." href="/articles/10.1186/s13059-018-1522-1#ref-CR92" id="ref-link-section-d8349793e2984">92</a>] with extension to Nganasan (data collected by BEDLAN). We used only the languages matching the ethnic identity of the individuals sampled for the genetic analyses (16 languages in total). We chose North Saami language to represent the genetic sample of Swedish Saami as that population has been sampled between the speaker areas of Lule Saami and North Saami (we do not have linguistic data on Lule Saami). We used Kildin Saami language to represent the genetic sample of Kola Saami as the sample has been collected from the classic distribution range of Kildin Saami.</p><p>Our linguistic data comprises of basic vocabulary data referring to meanings (words) that are universal, maximally resistant to borrowings and temporally stable. The Uralic basic vocabulary data and cognate assessments were achieved from the available literature. The data was collected by one single person (Jyri Lehtinen) which ensured equal quality of the data throughout the languages. In total, we had 226 meanings based on the meanings listed in Swadesh 100 and 200 lists [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 93" title="Swadesh M. Lexicostatistic dating of prehistoric ethnic contacts. P Am Philos Soc. 1952;96:452–63." href="/articles/10.1186/s13059-018-1522-1#ref-CR93" id="ref-link-section-d8349793e2990">93</a>, <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 94" title="Swadesh M. Towards greater accuracy in lexicostatistic dating. Int J Am Linguist. 1955;21:121–37. https://doi.org/10.1086/464321 " href="/articles/10.1186/s13059-018-1522-1#ref-CR94" id="ref-link-section-d8349793e2993">94</a>] and the Leipzig-Jakarta list [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 95" title="Tadmor U. Loanwords in the world’s languages: findings and results. In: Haspelmath M, Tadmor U, editors. Loanwords in the World’s languages: a comparative handbook. Berlin: Walter de Gruyter; 2009. p. 55–75." href="/articles/10.1186/s13059-018-1522-1#ref-CR95" id="ref-link-section-d8349793e2996">95</a>]. We used the whole data without extracting the known loan words, as Lehtinen et al. [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 92" title="Lehtinen J, Honkola T, Korhonen K, Syrjänen K, Wahlberg N, et al. Behind family trees: secondary connections in Uralic language networks. Lang Dyn Chang. 2014;4:189–221." href="/articles/10.1186/s13059-018-1522-1#ref-CR92" id="ref-link-section-d8349793e2999">92</a>] concludes that the loan words do not mess the evolutionary signal of the Uralic tree, but add information of the horizontal transfer of lexical material. The linguistic data was coded into binary form according to the cognacy relationships, i.e. whether the words for a meaning in two languages shared a common origin (=1) or not (=0). The phylogenetic tree was made with the MrBayes software [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 96" title="Huelsenbeck JP, Ronquist F. MRBAYES: Bayesian inference of phylogeny. Bioinformatics. 2001;17:754–5. https://doi.org/10.1093/bioinformatics/17.8.754 " href="/articles/10.1186/s13059-018-1522-1#ref-CR96" id="ref-link-section-d8349793e3002">96</a>] by following the settings in Syrjänen et al. [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 16" title="Syrjänen K, Honkola T, Korhonen K, Lehtinen J, Vesakoski O, et al. Shedding more light on language classification using basic vocabularies and phylogenetic methods. A case study of Uralic. Diachronica. 2013;30:323–52. https://doi.org/10.1075/dia.30.3.02syr " href="/articles/10.1186/s13059-018-1522-1#ref-CR16" id="ref-link-section-d8349793e3006">16</a>]. The produced phylogeny resembles the ones in Syrjänen et al. [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 16" title="Syrjänen K, Honkola T, Korhonen K, Lehtinen J, Vesakoski O, et al. Shedding more light on language classification using basic vocabularies and phylogenetic methods. A case study of Uralic. Diachronica. 2013;30:323–52. https://doi.org/10.1075/dia.30.3.02syr " href="/articles/10.1186/s13059-018-1522-1#ref-CR16" id="ref-link-section-d8349793e3009">16</a>] and Lehtinen et al. [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 92" title="Lehtinen J, Honkola T, Korhonen K, Syrjänen K, Wahlberg N, et al. Behind family trees: secondary connections in Uralic language networks. Lang Dyn Chang. 2014;4:189–221." href="/articles/10.1186/s13059-018-1522-1#ref-CR92" id="ref-link-section-d8349793e3012">92</a>], has a well-supported structure following the outcomes in the earlier Uralic literature and is better resolved than many of the recent trees made with traditional linguistic methods without objective computational analyses or large data behind (see Syrjänen et al. [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 16" title="Syrjänen K, Honkola T, Korhonen K, Lehtinen J, Vesakoski O, et al. Shedding more light on language classification using basic vocabularies and phylogenetic methods. A case study of Uralic. Diachronica. 2013;30:323–52. https://doi.org/10.1075/dia.30.3.02syr " href="/articles/10.1186/s13059-018-1522-1#ref-CR16" id="ref-link-section-d8349793e3015">16</a>] for review). The in-depth presentation of the language data, analyses, the comparison between trees, networks and earlier suggestions of the Uralic tree are given in Lehtinen et al. [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 92" title="Lehtinen J, Honkola T, Korhonen K, Syrjänen K, Wahlberg N, et al. Behind family trees: secondary connections in Uralic language networks. Lang Dyn Chang. 2014;4:189–221." href="/articles/10.1186/s13059-018-1522-1#ref-CR92" id="ref-link-section-d8349793e3018">92</a>] and Syrjänen et al. [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 16" title="Syrjänen K, Honkola T, Korhonen K, Lehtinen J, Vesakoski O, et al. Shedding more light on language classification using basic vocabularies and phylogenetic methods. A case study of Uralic. Diachronica. 2013;30:323–52. https://doi.org/10.1075/dia.30.3.02syr " href="/articles/10.1186/s13059-018-1522-1#ref-CR16" id="ref-link-section-d8349793e3021">16</a>], respectively. Some early criticism of the use of Swadesh list data by geneticists concerned the use of distance-based tree-building techniques known as ‘lexicostatistics’. Over the last decade, the application of modern Bayesian phylogenetic methods to linguistic data has allowed researchers to overcome these problems [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 97" title="Atkinson QD, Gray RD. How old is the Indo-European language family? Illumination or more moths to the flame? In: Clackson J, Forster P, Renfrew C, editors. Phylogenetic methods and the prehistory of languages. Cambridge: MacDonald Institute; 2006. p. 91–109." href="/articles/10.1186/s13059-018-1522-1#ref-CR97" id="ref-link-section-d8349793e3025">97</a>, <a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 98" title="Greenhill S, Currie T, Gray R. Does horizontal transmission invalidate cultural phylogenies? Proc Biol Sci. 2009;276:22992306. https://doi.org/10.1098/rspb.2008.1944 " href="/articles/10.1186/s13059-018-1522-1#ref-CR98" id="ref-link-section-d8349793e3028">98</a>]. We calculated pairwise linguistic distances between the tips of the phylogenetic tree (i.e. branch lengths) with the ‘ape’ package [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 99" title="Paradis E, Claude J, Strimmer K. APE: analyses of phylogenetics and evolution in R language. Bioinformatics. 2004;20:289–90. https://doi.org/10.1093/bioinformatics/btg412 " href="/articles/10.1186/s13059-018-1522-1#ref-CR99" id="ref-link-section-d8349793e3031">99</a>] in R [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 100" title="R-Core-Team. R: a language and environment for statistical computing. Vienna: R Foundation for statistical computing; 2016. https://www.r-project.org/ " href="/articles/10.1186/s13059-018-1522-1#ref-CR100" id="ref-link-section-d8349793e3034">100</a>]. Geographical distances were calculated as great-circle-distances (haversine) between genetic sampling locations with the ‘geosphere’ package in R [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 101" title="Hijmans RJ (2016) Geosphere: spherical trigonometry. R package. https://CRAN.R-project.org/package=geosphere ." href="/articles/10.1186/s13059-018-1522-1#ref-CR101" id="ref-link-section-d8349793e3037">101</a>].</p><p>To assess correlation between genetic, linguistic (lexical) and geographic distances for the Uralic-speaking populations, we employed the Mantel test [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 56" title="Mantel N. The detection of disease clustering and a generalized regression approach. Cancer Res. 1967;27:209–20. http://cancerres.aacrjournals.org/content/27/2_Part_1/209 " href="/articles/10.1186/s13059-018-1522-1#ref-CR56" id="ref-link-section-d8349793e3043">56</a>] using Pearson’s correlation coefficient. To test whether statistically significant associations between linguistic and genetic affiliations reflect the same events in population history or parallel but separate isolation by distance processes, partial correlations keeping geography constant were performed [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 102" title="Nettle D, Harriss L. Genetic and linguistic affinities between human populations in Eurasia and West Africa. Hum Biol. 2003;75:331–44. https://doi.org/10.1353/hub.2003.0048 " href="/articles/10.1186/s13059-018-1522-1#ref-CR102" id="ref-link-section-d8349793e3046">102</a>]. Analyses were performed using the ‘vegan’ package [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 103" title="Oksanen J, Kindt R, Legendre P, O’Hara B, Stevens MHH, et al. The vegan package. Community Ecol Packag. 2007;10:631–7. https://CRAN.R-project.org/package=vegan " href="/articles/10.1186/s13059-018-1522-1#ref-CR103" id="ref-link-section-d8349793e3049">103</a>] in R. Statistical significance was assessed using 1000 random permutations. We applied both of these tests to four types of genetic matrices. We used (1) Slatkin’s linearized Ф<sub>ST</sub> [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 104" title="Slatkin M. A measure of population subdivision based on microsatellite allele frequencies. Genetics. 1995;139:457–62. http://www.genetics.org/content/139/1/457 " href="/articles/10.1186/s13059-018-1522-1#ref-CR104" id="ref-link-section-d8349793e3054">104</a>] values calculated based on both mtDNA and chrY hg frequencies and (2) conventional F<sub>ST</sub> for both mtDNA and chrY [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 105" title="Excoffier L, Lischer HEL. Arlequin suite ver 3.5: a new series of programs to perform population genetics analyses under Linux and Windows. Mol Ecol Resour. 2010;10:564–7. https://doi.org/10.1111/j.1755-0998.2010.02847.x " href="/articles/10.1186/s13059-018-1522-1#ref-CR105" id="ref-link-section-d8349793e3060">105</a>], (3) Weir and Cockerham [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 76" title="Weir BS, Cockerham CC. Estimating F-statistics for the analysis of population structure. Evolution (N Y). 1984;38:1358–70. https://doi.org/10.2307/2408641 " href="/articles/10.1186/s13059-018-1522-1#ref-CR76" id="ref-link-section-d8349793e3063">76</a>] pairwise average F<sub>ST</sub> for autosomal SNPs and (4) total variation distances (TVD) [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 61" title="Leslie S, Winney B, Hellenthal G, Davison D, Boumertit A, et al. The fine-scale genetic structure of the British population. Nature. 2015;519:309–14. https://doi.org/10.1038/nature14230 " href="/articles/10.1186/s13059-018-1522-1#ref-CR61" id="ref-link-section-d8349793e3068">61</a>] between group pairs of fineSTRUCTURE chunkcount matrix (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM15">15</a>: Table S14). The calculations of geographic distances between populations were performed by using approximate latitude and longitude data for the sample sites (Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM15">15</a>: Table S14I). The results of the Mantel test are presented in Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM16">16</a>: Table S15.</p><h4 class="c-article__sub-heading c-article__sub-heading--small" id="Sec24">Correlation between autosomal ADMIXTURE component k9 l and chrY hg N sublineages</h4><p>Pearson correlation coefficients between two variables—the frequency of the k9 ancestral component (Fig.<a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/10.1186/s13059-018-1522-1#Fig3">3</a>b) and the frequency of chrY hg N sub-hgs in European, Volga-Uralic, Siberian and Central Asian populations —were calculated in R [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 100" title="R-Core-Team. R: a language and environment for statistical computing. Vienna: R Foundation for statistical computing; 2016. https://www.r-project.org/ " href="/articles/10.1186/s13059-018-1522-1#ref-CR100" id="ref-link-section-d8349793e3092">100</a>] using <i>cor.test()</i>. Results are presented in Additional file <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/10.1186/s13059-018-1522-1#MOESM7">7</a>: Table S6.</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">Yunusbayev B, Metspalu M, Metspalu E, Valeev A, Litvinov S, et al. 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Genes reflect the common recent history of the Uralic-speaking populations. Gene Expression Omnibus: NCBI gene expression and hybridization array data repository. <a href="https://www.ncbi.nlm.nih.gov/geo/query/acc.cgi?acc=GSE108646" data-track="click_references" data-track-action="external reference" data-track-value="external reference" data-track-label="https://www.ncbi.nlm.nih.gov/geo/query/acc.cgi?acc=GSE108646">https://www.ncbi.nlm.nih.gov/geo/query/acc.cgi?acc=GSE108646</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/10.1186/s13059-018-1522-1?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 the individuals who provided DNA samples for this study. We are very thankful for late Prof. Lars Beckman for doing the anthropological research and sharing the samples of the Swedish Saami. We are grateful to Dr. Garreth Hellenthal and Dr. Daniel Lawson for the scripts and the advice provided for the ChromoPainter and fineSTRUCTURE analysis. We would like to thank Erwan Pennarun and Dr. Mari Järve for technical assistance, Viljo Soo for his help in autosomal genotyping and Prof. Karl Pajusalu and Dr. Miina Norvik for their advice on linguistics. All calculations were carried out at the High Performance Computing Center of the University of Tartu.</p> <h3 class="c-article__sub-heading" id="FPar3">Funding</h3> <p>This work was supported by the European Union through the European Regional Development Fund with projects 2014-2020.4.01.15-0012, 2014-2020.4.01.16-0271, 2014-2020.4.01.16-0125 and by the Estonian Basic Research Grant IUT24 to the Estonian Biocentre. Estonian Personal Research grants PUT1217, PUT1339 and PRG243 support KT, AK and MM respectively. TH and OV are grateful to Kone Foundation (Finland) for funding the work of the BEDLAN team. LPO was supported by Russian Federation State Research Project No. 0324-2018-0016. Russian Foundation for Basic Research grant no 16-06-00303 supported EB. NEFREX grant funded by the European Union (People Marie Curie Actions; International Research Staff Exchange Scheme; call FP7-PEOPLE-2012-IRSES-number 318979) supported GH, QA, MK and MM. None of the funding bodies played a role in study design or data collection, analysis, or interpretation.</p> <h3 class="c-article__sub-heading" id="FPar4">Availability of data and materials</h3> <p>The dataset generated and analysed during the current study is available in the NCBI-GEO repository [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 106" title="Edgar R, Domrachev M, Lash AE. Gene expression omnibus: NCBI gene expression and hybridization array data repository. Nucleic Acids Res. 2002;30:207–10. 
 https://doi.org/10.1093/nar/30.1.207
 
 
 " href="/articles/10.1186/s13059-018-1522-1#ref-CR106" id="ref-link-section-d8349793e3130">106</a>] and is available through the accession no. GSE108646 [<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 107" title="Tambets, K., Metspalu, M., Villems R. Genes reflect the common recent history of the Uralic-speaking populations. Gene Expression Omnibus: NCBI gene expression and hybridization array data repository. 
 https://www.ncbi.nlm.nih.gov/geo/query/acc.cgi?acc=GSE108646
 
 ." href="/articles/10.1186/s13059-018-1522-1#ref-CR107" id="ref-link-section-d8349793e3133">107</a>]: <a href="https://www.ncbi.nlm.nih.gov/geo/query/acc.cgi?acc=GSE108646">https://www.ncbi.nlm.nih.gov/geo/query/acc.cgi?acc=GSE108646</a>.</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"><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">Estonian Biocentre, Institute of Genomics, University of Tartu, Riia 23b, 51010, Tartu, Estonia</p><p class="c-article-author-affiliation__authors-list">Kristiina Tambets, Bayazit Yunusbayev, Georgi Hudjashov, Anne-Mai Ilumäe, Siiri Rootsi, Alena Kushniarevich, Sergey Litvinov, Maere Reidla, Ene Metspalu, Lehti Saag, Monika Karmin, Jüri Parik, Sergey I. Zhadanov, Marina Gubina, Larisa D. Damba, Marina Bermisheva, Tuuli Reisberg, Khadizhat Dibirova, Richard Villems, Toomas Kivisild & Mait Metspalu</p></li><li id="Aff2"><p class="c-article-author-affiliation__address">Ufa Scientific Center of RAS, Ufa, 450054, Russia</p><p class="c-article-author-affiliation__authors-list">Bayazit Yunusbayev</p></li><li id="Aff3"><p class="c-article-author-affiliation__address">Statistics and Bioinformatics Group, Institute of Fundamental Sciences, Massey University, Palmerston North, 4442, New Zealand</p><p class="c-article-author-affiliation__authors-list">Georgi Hudjashov</p></li><li id="Aff4"><p class="c-article-author-affiliation__address">Department of Biology, University of Turku, 20014, Turku, Finland</p><p class="c-article-author-affiliation__authors-list">Terhi Honkola & Outi Vesakoski</p></li><li id="Aff5"><p class="c-article-author-affiliation__address">Institute of Estonian and General Linguistics, University of Tartu, 51014, Tartu, Estonia</p><p class="c-article-author-affiliation__authors-list">Terhi Honkola</p></li><li id="Aff6"><p class="c-article-author-affiliation__address">School of Psychology, University of Auckland, Auckland, 1142, New Zealand</p><p class="c-article-author-affiliation__authors-list">Quentin Atkinson</p></li><li id="Aff7"><p class="c-article-author-affiliation__address">Department of Linguistic and Cultural Evolution, Max Planck Institute for the Science of Human History, D-07745, Jena, Germany</p><p class="c-article-author-affiliation__authors-list">Quentin Atkinson</p></li><li id="Aff8"><p class="c-article-author-affiliation__address">The Francis Crick Institute, 1 Midland Road, London, NW1 1AT, UK</p><p class="c-article-author-affiliation__authors-list">Pontus Skoglund</p></li><li id="Aff9"><p class="c-article-author-affiliation__address">Institute of Genetics and Cytology of the National Academy of Sciences of Belarus, Minsk, 220072, Republic of Belarus</p><p class="c-article-author-affiliation__authors-list">Alena Kushniarevich</p></li><li id="Aff10"><p class="c-article-author-affiliation__address">Institute of Biochemistry and Genetics, Ufa Scientific Center of RAS, Ufa, 450054, Russia</p><p class="c-article-author-affiliation__authors-list">Sergey Litvinov, Marina Bermisheva & Elza K. Khusnutdinova</p></li><li id="Aff11"><p class="c-article-author-affiliation__address">Department of Evolutionary Biology, Institute of Molecular and Cell Biology, University of Tartu, 51010, Tartu, Estonia</p><p class="c-article-author-affiliation__authors-list">Maere Reidla, Lehti Saag, Jüri Parik, Richard Villems & Toomas Kivisild</p></li><li id="Aff12"><p class="c-article-author-affiliation__address">Department of Geography and Geology, University of Turku, 20014, Turku, Finland</p><p class="c-article-author-affiliation__authors-list">Timo Rantanen</p></li><li id="Aff13"><p class="c-article-author-affiliation__address">Department of Radiology, The Mount Sinai Medical Center, New York, NY, 10029, USA</p><p class="c-article-author-affiliation__authors-list">Sergey I. Zhadanov</p></li><li id="Aff14"><p class="c-article-author-affiliation__address">Institute of Cytology and Genetics, Siberian Branch of RAS, Novosibirsk, 630090, Russia</p><p class="c-article-author-affiliation__authors-list">Marina Gubina, Ludmila P. Osipova & Mikhail Voevoda</p></li><li id="Aff15"><p class="c-article-author-affiliation__address">Research Institute of Medical and Social Problems and Control of the Healthcare Department of Tuva Republic, Kyzyl, 667003, Russia</p><p class="c-article-author-affiliation__authors-list">Larisa D. Damba</p></li><li id="Aff16"><p class="c-article-author-affiliation__address">Research Centre for Medical Genetics, Russian Academy of Medical Sciences, Moscow, 115478, Russia</p><p class="c-article-author-affiliation__authors-list">Khadizhat Dibirova, Oleg Balanovsky & Elena Balanovska</p></li><li id="Aff17"><p class="c-article-author-affiliation__address">Northern State Medical University, Arkhangelsk, 163000, Russia</p><p class="c-article-author-affiliation__authors-list">Irina Evseeva</p></li><li id="Aff18"><p class="c-article-author-affiliation__address">Anthony Nolan, London, NW3 2NU, UK</p><p class="c-article-author-affiliation__authors-list">Irina Evseeva</p></li><li id="Aff19"><p class="c-article-author-affiliation__address">Research Centre of Estonian Genome Center, Institute of Genomics, University of Tartu, 51010, Tartu, Estonia</p><p class="c-article-author-affiliation__authors-list">Mari Nelis, Andres Metspalu & Tõnu Esko</p></li><li id="Aff20"><p class="c-article-author-affiliation__address">Latvian Biomedical Research and Study Centre, Riga, LV-1067, Latvia</p><p class="c-article-author-affiliation__authors-list">Janis Klovins</p></li><li id="Aff21"><p class="c-article-author-affiliation__address">Vavilov Institute for General Genetics, RAS, Moscow, 119991, Russia</p><p class="c-article-author-affiliation__authors-list">Oleg Balanovsky</p></li><li id="Aff22"><p class="c-article-author-affiliation__address">Department of Genetics and Fundamental Medicine, Bashkir State University, Ufa, 450054, Russia</p><p class="c-article-author-affiliation__authors-list">Elza K. Khusnutdinova</p></li><li id="Aff23"><p class="c-article-author-affiliation__address">Novosibirsk State University, 2 Pirogova Str, Novosibirsk, 630090, Russia</p><p class="c-article-author-affiliation__authors-list">Ludmila P. Osipova & Mikhail Voevoda</p></li><li id="Aff24"><p class="c-article-author-affiliation__address">Institute of Internal Medicine, Siberian Branch of Russian Academy of Medical Sciences, Novosibirsk, 630090, Russia</p><p class="c-article-author-affiliation__authors-list">Mikhail Voevoda</p></li><li id="Aff25"><p class="c-article-author-affiliation__address">Department of Archaeology, University of Cambridge, Cambridge, CB2 1QH, UK</p><p class="c-article-author-affiliation__authors-list">Toomas Kivisild</p></li><li id="Aff26"><p class="c-article-author-affiliation__address">Department of Human Genetics, KU Leuven, Leuven, 3000, Belgium</p><p class="c-article-author-affiliation__authors-list">Toomas Kivisild</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-Kristiina-Tambets-Aff1"><span class="c-article-authors-search__title u-h3 js-search-name">Kristiina Tambets</span><div class="c-article-authors-search__list"><div class="c-article-authors-search__item c-article-authors-search__list-item--left"><a href="https://www.biomedcentral.com/search?query=author%23Kristiina%20Tambets" class="c-article-button" data-track="click" data-track-action="author link - 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BY designed the study, performed analysis, interpreted results and wrote the paper. GH performed analysis, interpreted results and wrote the paper. A-MI performed experiments for Y chr data, performed analysis, interpreted results and helped to write the paper. SR performed the experiments for Y chr data, interpreted results and helped to write the paper. TH and OV compiled and provided linguistic datasets, performed analysis, interpreted results and wrote the paper. QA performed correlation analysis of genetic, geographic and linguistic datasets, interpreted the results and wrote the paper. PS and AK performed analysis, interpreted results and helped to write the paper. SL performed the analysis and interpreted the results. MR performed the experiments for mtDNA and Y chr data, performed the analysis, interpreted the results and helped to write the paper. EM, JP and KD performed the experiments for mtDNA, Y chr and autosomal data and interpreted the results. LS analysed the autosomal data, interpreted the results and helped to write the paper. TiR compiled and analysed the linguistic map and geographical data of the sample. MK interpreted the results and contributed to the writing of the paper. SIZ performed the experiments for the mtDNA data and interpreted the results. MG conducted the anthropological research and collection of Nenets, Ket and Selkup samples, performed the experiments for mtDNA and Y chr data and interpreted results. LDD conducted the anthropological research and collection of Khanty and Mansi samples, performed experiments for mtDNA and Y chr data and interpreted the results. MB performed the experiments for the mtDNA and Y chr data and interpreted the results. TR performed the experiments for the mtDNA data, analysed the autosomal data and interpreted the results. IE performed the experiments for mtDNA and interpreted the results. MN performed the experiments for the autosomal data and interpreted the results. JK conducted the anthropological research and the collection of Latvian samples. AM and TE provided samples from the collection of Estonian Genome Centre and interpreted the results. OB and EB conducted the anthropological research and collection of Kola Saami samples, performed the experiments for the mtDNA, Y chr and autosomal data and interpreted the results. EKK conducted anthropological research and collection of the samples of Volga-Ural region samples and interpreted the results. LPO conducted the anthropological research and collection of Nenets, Ket and Selkup samples. MV conducted the anthropological research and collection of Khanty and Mansi samples. RV designed the study, interpreted the results and wrote the paper. TK and MM designed the study, analysed the data, interpreted the results and wrote the paper. All authors read and approved the final manuscript.</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:ktambets@ebc.ee">Kristiina Tambets</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" id="FPar5">Ethics approval and consent to participate</h3> <p>DNA samples were obtained from unrelated volunteers, all donors provided informed consent and all experiments were performed in accordance with the relevant guidelines and regulations of the involved countries. The research has been approved by the Research Ethics Commitees of the University of Tartu and the Russian Academy of Sciences (approval nos. 228/M-40, 252/M-17, 17146-9217)<i>.</i> Experimental methods of the study comply with the Helsinki Declaration.</p> <h3 class="c-article__sub-heading" id="FPar6">Consent for publication</h3> <p>Not applicable</p> <h3 class="c-article__sub-heading" id="FPar7">Competing interests</h3> <p>The authors declare that they have no competing interests.</p> <h3 class="c-article__sub-heading" id="FPar8">Publisher’s Note</h3> <p>Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.</p> </div></div></section><section data-title="Additional files"><div class="c-article-section" id="Sec25-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="Sec25">Additional files</h2><div class="c-article-section__content" id="Sec25-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" data-test="supp-item" id="MOESM1"><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="additional file 1:" href="https://static-content.springer.com/esm/art%3A10.1186%2Fs13059-018-1522-1/MediaObjects/13059_2018_1522_MOESM1_ESM.xlsx" data-supp-info-image="">Additional file 1:</a></h3><div class="c-article-supplementary__description" data-component="thumbnail-container"><p><b>Table S1.</b> Autosomal data used in the study. (XLSX 28 kb)</p></div></div><div class="c-article-supplementary__item" data-test="supp-item" id="MOESM2"><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="additional file 2:" href="https://static-content.springer.com/esm/art%3A10.1186%2Fs13059-018-1522-1/MediaObjects/13059_2018_1522_MOESM2_ESM.pdf" data-supp-info-image="">Additional file 2:</a></h3><div class="c-article-supplementary__description" data-component="thumbnail-container"><p><b>Table S2.</b> Ethnolinguistic characteristics of studied Uralic populations. (PDF 15 kb)</p></div></div><div class="c-article-supplementary__item" data-test="supp-item" id="MOESM3"><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="additional file 3:" href="https://static-content.springer.com/esm/art%3A10.1186%2Fs13059-018-1522-1/MediaObjects/13059_2018_1522_MOESM3_ESM.pdf" data-supp-info-image="">Additional file 3:</a></h3><div class="c-article-supplementary__description" data-component="thumbnail-container"><p><b>Figure S1.</b> Principal component analysis of the studied Uralic-speaking populations in Eurasian context. <b>Figure S2.</b> F<sub>ST</sub> distances of the studied Uralic-speaking populations in the global context. <b>Figure S3.</b> ADMIXTURE analysis. <b>Figure S4.</b> <i>D</i>-statistics. <b>Figure S5.</b> fineSTRUCTURE tree. <b>Figure S6.</b> fineSTRUCTURE heatmap. <b>Figure S7.</b> Admixture dates of ALDER. <b>Figure S8.</b> Outgroup <i>f</i>3 test between modern and ancient samples. <b>Figure S9.</b> Outgroup <i>f</i>3 results plotted pairwise against each other. <b>Figure S10.</b> Admixture graphs. <b>Figure S11.</b> Phylogenetic network of Y chromosome hg Q. <b>Figure S12.</b> Trees constructed from linguistic and genetic data of Uralic peoples. <b>Figure S13.</b> Comparison of autosomal and X chromosomal outgroup <i>f</i>3 statistics for Estonians, Udmurts and Khanty. (PDF 11550 kb)</p></div></div><div class="c-article-supplementary__item" data-test="supp-item" id="MOESM4"><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="additional file 4:" href="https://static-content.springer.com/esm/art%3A10.1186%2Fs13059-018-1522-1/MediaObjects/13059_2018_1522_MOESM4_ESM.xlsx" data-supp-info-image="">Additional file 4:</a></h3><div class="c-article-supplementary__description" data-component="thumbnail-container"><p><b>Table S3.</b> Mitochondrial DNA data of Uralic speakers. (XLSX 541 kb)</p></div></div><div class="c-article-supplementary__item" data-test="supp-item" id="MOESM5"><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="additional file 5:" href="https://static-content.springer.com/esm/art%3A10.1186%2Fs13059-018-1522-1/MediaObjects/13059_2018_1522_MOESM5_ESM.xlsx" data-supp-info-image="">Additional file 5:</a></h3><div class="c-article-supplementary__description" data-component="thumbnail-container"><p><b>Table S4.</b> Mitochondrial DNA haplogroup frequencies in Eurasia. (XLSX 71 kb)</p></div></div><div class="c-article-supplementary__item" data-test="supp-item" id="MOESM6"><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="additional file 6:" href="https://static-content.springer.com/esm/art%3A10.1186%2Fs13059-018-1522-1/MediaObjects/13059_2018_1522_MOESM6_ESM.xlsx" data-supp-info-image="">Additional file 6:</a></h3><div class="c-article-supplementary__description" data-component="thumbnail-container"><p><b>Table S5.</b> Y chromosome haplogroup frequencies in Eurasia. (XLSX 22 kb)</p></div></div><div class="c-article-supplementary__item" data-test="supp-item" id="MOESM7"><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="additional file 7:" href="https://static-content.springer.com/esm/art%3A10.1186%2Fs13059-018-1522-1/MediaObjects/13059_2018_1522_MOESM7_ESM.xlsx" data-supp-info-image="">Additional file 7:</a></h3><div class="c-article-supplementary__description" data-component="thumbnail-container"><p><b>Table S6.</b> Correlation analysis of k9 ancestral component and chromosome Y haplogroup N sub-haplogroups. (XLSX 16 kb)</p></div></div><div class="c-article-supplementary__item" data-test="supp-item" id="MOESM8"><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="additional file 8:" href="https://static-content.springer.com/esm/art%3A10.1186%2Fs13059-018-1522-1/MediaObjects/13059_2018_1522_MOESM8_ESM.xlsx" data-supp-info-image="">Additional file 8:</a></h3><div class="c-article-supplementary__description" data-component="thumbnail-container"><p><b>Table S7.</b> Average identity-by-descent-segment share in Northeast European populations. (XLSX 50 kb)</p></div></div><div class="c-article-supplementary__item" data-test="supp-item" id="MOESM9"><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="additional file 9:" href="https://static-content.springer.com/esm/art%3A10.1186%2Fs13059-018-1522-1/MediaObjects/13059_2018_1522_MOESM9_ESM.xlsx" data-supp-info-image="">Additional file 9:</a></h3><div class="c-article-supplementary__description" data-component="thumbnail-container"><p><b>Table S8.</b> Average identity-by-descent-segment share in Central and East European populations. (XLSX 44 kb)</p></div></div><div class="c-article-supplementary__item" data-test="supp-item" id="MOESM10"><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="additional file 10:" href="https://static-content.springer.com/esm/art%3A10.1186%2Fs13059-018-1522-1/MediaObjects/13059_2018_1522_MOESM10_ESM.xlsx" data-supp-info-image="">Additional file 10:</a></h3><div class="c-article-supplementary__description" data-component="thumbnail-container"><p><b>Table S9.</b> Average identity-by-descent-segment share in Siberian populations. (XLSX 41 kb)</p></div></div><div class="c-article-supplementary__item" data-test="supp-item" id="MOESM11"><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="additional file 11:" href="https://static-content.springer.com/esm/art%3A10.1186%2Fs13059-018-1522-1/MediaObjects/13059_2018_1522_MOESM11_ESM.xlsx" data-supp-info-image="">Additional file 11:</a></h3><div class="c-article-supplementary__description" data-component="thumbnail-container"><p><b>Table S10.</b> Globetrotter results. (XLSX 48 kb)</p></div></div><div class="c-article-supplementary__item" data-test="supp-item" id="MOESM12"><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="additional file 12:" href="https://static-content.springer.com/esm/art%3A10.1186%2Fs13059-018-1522-1/MediaObjects/13059_2018_1522_MOESM12_ESM.pdf" data-supp-info-image="">Additional file 12:</a></h3><div class="c-article-supplementary__description" data-component="thumbnail-container"><p><b>Table S11.</b> Details of Globetrotter regional analysis. (PDF 50 kb)</p></div></div><div class="c-article-supplementary__item" data-test="supp-item" id="MOESM13"><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="additional file 13:" href="https://static-content.springer.com/esm/art%3A10.1186%2Fs13059-018-1522-1/MediaObjects/13059_2018_1522_MOESM13_ESM.xlsx" data-supp-info-image="">Additional file 13:</a></h3><div class="c-article-supplementary__description" data-component="thumbnail-container"><p><b>Table S12.</b> ALDER vs Globetrotter dates. (XLSX 210 kb)</p></div></div><div class="c-article-supplementary__item" data-test="supp-item" id="MOESM14"><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="additional file 14:" href="https://static-content.springer.com/esm/art%3A10.1186%2Fs13059-018-1522-1/MediaObjects/13059_2018_1522_MOESM14_ESM.xlsx" data-supp-info-image="">Additional file 14:</a></h3><div class="c-article-supplementary__description" data-component="thumbnail-container"><p><b>Table S13.</b> Outgroup <i>f</i>3 statistic. (XLSX 305 kb)</p></div></div><div class="c-article-supplementary__item" data-test="supp-item" id="MOESM15"><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="additional file 15:" href="https://static-content.springer.com/esm/art%3A10.1186%2Fs13059-018-1522-1/MediaObjects/13059_2018_1522_MOESM15_ESM.xlsx" data-supp-info-image="">Additional file 15:</a></h3><div class="c-article-supplementary__description" data-component="thumbnail-container"><p><b>Table S14.</b> Data matrices for correlation analysis and geographical coordinates of studied populations. (XLSX 38 kb)</p></div></div><div class="c-article-supplementary__item" data-test="supp-item" id="MOESM16"><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="additional file 16:" href="https://static-content.springer.com/esm/art%3A10.1186%2Fs13059-018-1522-1/MediaObjects/13059_2018_1522_MOESM16_ESM.xlsx" data-supp-info-image="">Additional file 16:</a></h3><div class="c-article-supplementary__description" data-component="thumbnail-container"><p><b>Table S15.</b> Mantel test results. (XLSX 11 kb)</p></div></div><div class="c-article-supplementary__item" data-test="supp-item" id="MOESM17"><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="additional file 17:" href="https://static-content.springer.com/esm/art%3A10.1186%2Fs13059-018-1522-1/MediaObjects/13059_2018_1522_MOESM17_ESM.xlsx" data-supp-info-image="">Additional file 17:</a></h3><div class="c-article-supplementary__description" data-component="thumbnail-container"><p><b>Table S16.</b> Y chromosomal hg Q STR data. 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id="citeas">Cite this article</h3><p class="c-bibliographic-information__citation">Tambets, K., Yunusbayev, B., Hudjashov, G. <i>et al.</i> Genes reveal traces of common recent demographic history for most of the Uralic-speaking populations. <i>Genome Biol</i> <b>19</b>, 139 (2018). https://doi.org/10.1186/s13059-018-1522-1</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/10.1186/s13059-018-1522-1?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 class="c-bibliographic-information__list-item"><p>Received<span class="u-hide">: </span><span class="c-bibliographic-information__value"><time datetime="2018-01-12">12 January 2018</time></span></p></li><li class="c-bibliographic-information__list-item"><p>Accepted<span class="u-hide">: </span><span class="c-bibliographic-information__value"><time datetime="2018-09-03">03 September 2018</time></span></p></li><li class="c-bibliographic-information__list-item"><p>Published<span class="u-hide">: </span><span class="c-bibliographic-information__value"><time datetime="2018-09-21">21 September 2018</time></span></p></li><li class="c-bibliographic-information__list-item c-bibliographic-information__list-item--full-width"><p><abbr title="Digital Object Identifier">DOI</abbr><span class="u-hide">: </span><span class="c-bibliographic-information__value">https://doi.org/10.1186/s13059-018-1522-1</span></p></li></ul><div data-component="share-box"><div class="c-article-share-box u-display-none" 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