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Genetic history of Europe - Wikipedia
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class="vector-toc-list"> <li id="toc-Palaeolithic" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Palaeolithic"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.1</span> <span>Palaeolithic</span> </div> </a> <ul id="toc-Palaeolithic-sublist" class="vector-toc-list"> <li id="toc-Upper_Paleolithic" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Upper_Paleolithic"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.1.1</span> <span>Upper Paleolithic</span> </div> </a> <ul id="toc-Upper_Paleolithic-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Last_Glacial_Maximum" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Last_Glacial_Maximum"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.1.2</span> <span>Last Glacial Maximum</span> </div> </a> <ul id="toc-Last_Glacial_Maximum-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Mesolithic" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Mesolithic"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.2</span> <span>Mesolithic</span> </div> </a> <ul id="toc-Mesolithic-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Neolithic" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Neolithic"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.3</span> <span>Neolithic</span> </div> </a> <ul id="toc-Neolithic-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Bronze_Age" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Bronze_Age"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.4</span> <span>Bronze Age</span> </div> </a> <ul id="toc-Bronze_Age-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Recent_history" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Recent_history"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.5</span> <span>Recent history</span> </div> </a> <ul id="toc-Recent_history-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Genetics_of_modern_European_populations" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Genetics_of_modern_European_populations"> <div class="vector-toc-text"> <span class="vector-toc-numb">2</span> <span>Genetics of modern European populations</span> </div> </a> <button aria-controls="toc-Genetics_of_modern_European_populations-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle Genetics of modern European populations subsection</span> </button> <ul id="toc-Genetics_of_modern_European_populations-sublist" class="vector-toc-list"> <li id="toc-Patrilineal_studies" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Patrilineal_studies"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.1</span> <span>Patrilineal studies</span> </div> </a> <ul id="toc-Patrilineal_studies-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Matrilineal_studies" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Matrilineal_studies"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.2</span> <span>Matrilineal studies</span> </div> </a> <ul id="toc-Matrilineal_studies-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-European_population_sub-structure" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#European_population_sub-structure"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.3</span> <span>European population sub-structure</span> </div> </a> <ul id="toc-European_population_sub-structure-sublist" class="vector-toc-list"> <li id="toc-Autosomal_DNA" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Autosomal_DNA"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.3.1</span> <span>Autosomal DNA</span> </div> </a> <ul id="toc-Autosomal_DNA-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Autosomal_genetic_distances_(Fst)_based_on_SNPs_(2009)" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Autosomal_genetic_distances_(Fst)_based_on_SNPs_(2009)"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.3.2</span> <span>Autosomal genetic distances (Fst) based on SNPs (2009)</span> </div> </a> <ul id="toc-Autosomal_genetic_distances_(Fst)_based_on_SNPs_(2009)-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> </ul> </li> <li id="toc-History_of_research" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#History_of_research"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>History of research</span> </div> </a> <button aria-controls="toc-History_of_research-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle History of research subsection</span> </button> <ul id="toc-History_of_research-sublist" class="vector-toc-list"> <li id="toc-Classical_genetic_markers_(by_proxy)" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Classical_genetic_markers_(by_proxy)"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.1</span> <span>Classical genetic markers (by proxy)</span> </div> </a> <ul id="toc-Classical_genetic_markers_(by_proxy)-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Direct_DNA_analysis" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Direct_DNA_analysis"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2</span> <span>Direct DNA analysis</span> </div> </a> <ul id="toc-Direct_DNA_analysis-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-See_also" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#See_also"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>See also</span> </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>References</span> </div> </a> <button aria-controls="toc-References-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle References subsection</span> </button> <ul id="toc-References-sublist" class="vector-toc-list"> <li id="toc-Inline_citations" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Inline_citations"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.1</span> <span>Inline citations</span> </div> </a> <ul id="toc-Inline_citations-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Sources_referenced" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Sources_referenced"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.2</span> <span>Sources referenced</span> </div> </a> <ul id="toc-Sources_referenced-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Further_reading" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Further_reading"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>Further reading</span> </div> </a> <ul id="toc-Further_reading-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>External links</span> </div> </a> <ul id="toc-External_links-sublist" class="vector-toc-list"> </ul> </li> </ul> </div> </div> </nav> </div> </div> <div class="mw-content-container"> <main id="content" class="mw-body"> <header class="mw-body-header vector-page-titlebar"> <nav aria-label="Contents" class="vector-toc-landmark"> <div id="vector-page-titlebar-toc" class="vector-dropdown vector-page-titlebar-toc vector-button-flush-left" > <input type="checkbox" id="vector-page-titlebar-toc-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-vector-page-titlebar-toc" class="vector-dropdown-checkbox " aria-label="Toggle the table of contents" > <label id="vector-page-titlebar-toc-label" for="vector-page-titlebar-toc-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--icon-only " aria-hidden="true" ><span class="vector-icon mw-ui-icon-listBullet mw-ui-icon-wikimedia-listBullet"></span> <span class="vector-dropdown-label-text">Toggle the table of contents</span> </label> <div class="vector-dropdown-content"> <div id="vector-page-titlebar-toc-unpinned-container" class="vector-unpinned-container"> </div> </div> </div> </nav> <h1 id="firstHeading" class="firstHeading mw-first-heading"><span class="mw-page-title-main">Genetic history of Europe</span></h1> <div id="p-lang-btn" class="vector-dropdown mw-portlet mw-portlet-lang" > <input type="checkbox" id="p-lang-btn-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-p-lang-btn" class="vector-dropdown-checkbox mw-interlanguage-selector" aria-label="Go to an article in another language. Available in 13 languages" > <label id="p-lang-btn-label" for="p-lang-btn-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--action-progressive mw-portlet-lang-heading-13" aria-hidden="true" ><span class="vector-icon mw-ui-icon-language-progressive mw-ui-icon-wikimedia-language-progressive"></span> <span class="vector-dropdown-label-text">13 languages</span> </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D8%A7%D9%84%D8%AA%D8%A7%D8%B1%D9%8A%D8%AE_%D8%A7%D9%84%D8%AC%D9%8A%D9%86%D9%8A_%D9%84%D8%A3%D9%88%D8%B1%D9%88%D8%A8%D8%A7" title="التاريخ الجيني لأوروبا – Arabic" lang="ar" hreflang="ar" data-title="التاريخ الجيني لأوروبا" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Hist%C3%B2ria_gen%C3%A8tica_d%27Europa" title="Història genètica d'Europa – Catalan" lang="ca" hreflang="ca" data-title="Història genètica d'Europa" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target"><span>Català</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Historia_gen%C3%A9tica_de_Europa" title="Historia genética de Europa – Spanish" lang="es" hreflang="es" data-title="Historia genética de Europa" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-eu mw-list-item"><a href="https://eu.wikipedia.org/wiki/Europako_historia_genetikoa" title="Europako historia genetikoa – Basque" lang="eu" hreflang="eu" data-title="Europako historia genetikoa" data-language-autonym="Euskara" data-language-local-name="Basque" class="interlanguage-link-target"><span>Euskara</span></a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/Histoire_g%C3%A9n%C3%A9tique_des_populations_europ%C3%A9ennes" title="Histoire génétique des populations européennes – French" lang="fr" hreflang="fr" data-title="Histoire génétique des populations européennes" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-hy mw-list-item"><a href="https://hy.wikipedia.org/wiki/%D4%B5%D5%BE%D6%80%D5%B8%D5%BA%D5%A1%D5%B5%D5%AB_%D5%A3%D5%A5%D5%B6%D5%A5%D5%BF%D5%AB%D5%AF%D5%A1%D5%AF%D5%A1%D5%B6_%D5%BA%D5%A1%D5%BF%D5%B4%D5%B8%D6%82%D5%A9%D5%B5%D5%B8%D6%82%D5%B6" title="Եվրոպայի գենետիկական պատմություն – Armenian" lang="hy" hreflang="hy" data-title="Եվրոպայի գենետիկական պատմություն" data-language-autonym="Հայերեն" data-language-local-name="Armenian" class="interlanguage-link-target"><span>Հայերեն</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Storia_genetica_dell%27Europa" title="Storia genetica dell'Europa – Italian" lang="it" hreflang="it" data-title="Storia genetica dell'Europa" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target"><span>Italiano</span></a></li><li class="interlanguage-link interwiki-mk mw-list-item"><a href="https://mk.wikipedia.org/wiki/%D0%93%D0%B5%D0%BD%D0%B5%D1%82%D1%81%D0%BA%D0%B0_%D0%B8%D1%81%D1%82%D0%BE%D1%80%D0%B8%D1%98%D0%B0_%D0%BD%D0%B0_%D0%95%D0%B2%D1%80%D0%BE%D0%BF%D0%B0" title="Генетска историја на Европа – Macedonian" lang="mk" hreflang="mk" data-title="Генетска историја на Европа" data-language-autonym="Македонски" data-language-local-name="Macedonian" class="interlanguage-link-target"><span>Македонски</span></a></li><li class="interlanguage-link interwiki-pt mw-list-item"><a href="https://pt.wikipedia.org/wiki/Hist%C3%B3ria_gen%C3%A9tica_da_Europa" title="História genética da Europa – Portuguese" lang="pt" hreflang="pt" data-title="História genética da Europa" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target"><span>Português</span></a></li><li class="interlanguage-link interwiki-ru mw-list-item"><a href="https://ru.wikipedia.org/wiki/%D0%93%D0%B5%D0%BD%D0%B5%D1%82%D0%B8%D1%87%D0%B5%D1%81%D0%BA%D0%B0%D1%8F_%D0%B8%D1%81%D1%82%D0%BE%D1%80%D0%B8%D1%8F_%D0%95%D0%B2%D1%80%D0%BE%D0%BF%D1%8B" title="Генетическая история Европы – Russian" lang="ru" hreflang="ru" data-title="Генетическая история Европы" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/Eurooppalaisten_alkuper%C3%A4" title="Eurooppalaisten alkuperä – Finnish" lang="fi" hreflang="fi" data-title="Eurooppalaisten alkuperä" data-language-autonym="Suomi" data-language-local-name="Finnish" class="interlanguage-link-target"><span>Suomi</span></a></li><li class="interlanguage-link interwiki-ta mw-list-item"><a href="https://ta.wikipedia.org/wiki/%E0%AE%90%E0%AE%B0%E0%AF%8B%E0%AE%AA%E0%AF%8D%E0%AE%AA%E0%AE%BF%E0%AE%AF_%E0%AE%AE%E0%AE%B0%E0%AE%AA%E0%AE%BF%E0%AE%AF%E0%AE%B2%E0%AF%8D_%E0%AE%B5%E0%AE%B0%E0%AE%B2%E0%AE%BE%E0%AE%B1%E0%AF%81" title="ஐரோப்பிய மரபியல் வரலாறு – Tamil" lang="ta" hreflang="ta" data-title="ஐரோப்பிய மரபியல் வரலாறு" data-language-autonym="தமிழ்" data-language-local-name="Tamil" class="interlanguage-link-target"><span>தமிழ்</span></a></li><li class="interlanguage-link interwiki-zh mw-list-item"><a href="https://zh.wikipedia.org/wiki/%E6%AD%90%E6%B4%B2%E9%81%BA%E5%82%B3%E5%8F%B2" title="歐洲遺傳史 – Chinese" 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decoding="async" width="400" height="300" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/2b/European_genetic_structure_%28based_on_SNPs%29_PC_analysis.png/600px-European_genetic_structure_%28based_on_SNPs%29_PC_analysis.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/2b/European_genetic_structure_%28based_on_SNPs%29_PC_analysis.png/800px-European_genetic_structure_%28based_on_SNPs%29_PC_analysis.png 2x" data-file-width="2067" data-file-height="1552" /></a><figcaption> The European genetic structure today (based on 273,464 SNPs). Three levels of structure as revealed by PC analysis are shown: A) inter-continental; B) intra-continental; and C) inside a single country (Estonia), where median values of the PC1&2 are shown. D) European map illustrating the origin of sample and population size. CEU – Utah residents with ancestry from Northern and Western Europe, CHB – Han Chinese from Beijing, JPT – Japanese from Tokyo, and YRI – Yoruba from Ibadan, Nigeria.<sup id="cite_ref-FleischerNelis2009_1-0" class="reference"><a href="#cite_note-FleischerNelis2009-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </figcaption></figure> <p>The <b>genetic history of Europe</b> includes information around the formation, <a href="/wiki/Ethnogenesis" title="Ethnogenesis">ethnogenesis</a>, and other <a href="/wiki/DNA" title="DNA">DNA</a>-specific information about <a href="/wiki/Population" title="Population">populations</a> <a href="/wiki/Indigenous_peoples" title="Indigenous peoples">indigenous</a>, or living in <a href="/wiki/Europe" title="Europe">Europe</a>. </p><p><a href="/wiki/European_early_modern_humans" class="mw-redirect" title="European early modern humans">European early modern human</a> (EEMH) lineages between 40 and 26 ka (<a href="/wiki/Aurignacian" title="Aurignacian">Aurignacian</a>) were still part of a large Western Eurasian "meta-population", related to Central and Western Asian populations.<sup id="cite_ref-Seguin-Orlando2014_2-0" class="reference"><a href="#cite_note-Seguin-Orlando2014-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> Divergence into genetically distinct sub-populations within Western Eurasia is a result of increased <a href="/wiki/Recent_human_evolution" title="Recent human evolution">selection pressure</a> and <a href="/wiki/Founder_effect" title="Founder effect">founder effects</a> during the <a href="/wiki/Last_Glacial_Maximum" title="Last Glacial Maximum">Last Glacial Maximum</a> (LGM, <a href="/wiki/Gravettian" title="Gravettian">Gravettian</a>).<sup id="cite_ref-LGMpressures_3-0" class="reference"><a href="#cite_note-LGMpressures-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> </p><p>By the end of the LGM, after 20 ka, A Western European lineage, dubbed <a href="/wiki/West_European_hunter-gatherer" class="mw-redirect" title="West European hunter-gatherer">west European hunter-gatherer</a> (WHG) emerged from the <a href="/wiki/Solutrean" title="Solutrean">Solutrean</a> <a href="/wiki/Last_Glacial_Maximum_refugia" title="Last Glacial Maximum refugia">refugium</a> during the <a href="/wiki/European_Mesolithic" class="mw-redirect" title="European Mesolithic">European Mesolithic</a>.<sup id="cite_ref-Jones2015_4-0" class="reference"><a href="#cite_note-Jones2015-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> These mesolithic hunter-gatherer cultures are subsequently replaced in the <a href="/wiki/Neolithic_Revolution" title="Neolithic Revolution">Neolithic Revolution</a> as a result of the arrival of <a href="/wiki/Early_European_Farmers" title="Early European Farmers">Early European Farmer</a> (EEF) lineages derived from mesolithic populations of West Asia (<a href="/wiki/Mesolithic_Anatolia" class="mw-redirect" title="Mesolithic Anatolia">Anatolia</a> and the <a href="/wiki/Mesolithic_Caucasus" class="mw-redirect" title="Mesolithic Caucasus">Caucasus</a>).<sup id="cite_ref-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> In the <a href="/wiki/European_Bronze_Age" class="mw-redirect" title="European Bronze Age">European Bronze Age</a>, there were again substantial population replacements in parts of Europe by the intrusion of <a href="/wiki/Western_Steppe_Herders" title="Western Steppe Herders">Western Steppe Herder</a> (WSH) lineages from the <a href="/wiki/Pontic%E2%80%93Caspian_steppe" title="Pontic–Caspian steppe">Pontic–Caspian steppes</a>, arising from admixture between <a href="/wiki/Eastern_Hunter_Gatherers" class="mw-redirect" title="Eastern Hunter Gatherers">Eastern Hunter Gatherers</a> (EHG) and peoples related to Near Easterners. These Bronze Age population replacements are associated with the <a href="/wiki/Bell_Beaker_culture" title="Bell Beaker culture">Bell Beaker</a> and <a href="/wiki/Corded_Ware_culture" title="Corded Ware culture">Corded Ware</a> cultures archaeologically and with the <a href="/wiki/Indo-European_expansion" class="mw-redirect" title="Indo-European expansion">Indo-European expansion</a> linguistically.<sup id="cite_ref-6" class="reference"><a href="#cite_note-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Lazaridis2014_7-0" class="reference"><a href="#cite_note-Lazaridis2014-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> </p><p>As a result of the population movements during the Mesolithic to Bronze Age, modern European populations are distinguished by differences in WHG, EEF and <a href="/wiki/Ancient_North_Eurasian" title="Ancient North Eurasian">Ancient North Eurasian</a> (ANE) ancestry.<sup id="cite_ref-8" class="reference"><a href="#cite_note-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-ann_9-0" class="reference"><a href="#cite_note-ann-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Natgeo_10-0" class="reference"><a href="#cite_note-Natgeo-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> Admixture rates varied geographically; in the late Neolithic, WHG ancestry in farmers in Hungary was at around 10%, in Germany around 25% and in Iberia as high as 50%.<sup id="cite_ref-11" class="reference"><a href="#cite_note-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> The contribution of EEF is more significant in Mediterranean Europe, and declines towards northern and northeastern Europe, where WHG ancestry is stronger; the <a href="/wiki/Sardinians" class="mw-redirect" title="Sardinians">Sardinians</a> are considered to be the closest European group to the population of the EEF. </p><p>Ethnogenesis of the modern <a href="/wiki/Ethnic_groups_of_Europe" class="mw-redirect" title="Ethnic groups of Europe">ethnic groups of Europe</a> in the <a href="/wiki/History_of_Europe" title="History of Europe">historical period</a> is associated with numerous admixture events, primarily those associated with the <a href="/wiki/Migration_period" class="mw-redirect" title="Migration period">Migration period</a> and the <a href="/wiki/Decline_of_the_Roman_Empire" class="mw-redirect" title="Decline of the Roman Empire">decline of the Roman Empire</a>, associated with the <a href="/wiki/Germanic_languages" title="Germanic languages">Germanic</a>, <a href="/wiki/Viking_expansion" title="Viking expansion">Norse</a>, and <a href="/wiki/Early_Slavs" title="Early Slavs">Slavic</a> expansions </p><p>Research into the genetic history of Europe became possible in the second half of the 20th century, but did not yield results with high resolution before the 1990s. In the 1990s, preliminary results became possible, but they remained mostly limited to studies of <a href="/wiki/Mitochondrial_DNA" title="Mitochondrial DNA">mitochondrial</a> and <a href="/wiki/Y_chromosome" title="Y chromosome">Y-chromosomal</a> lineages. <a href="/wiki/Autosome" title="Autosome">Autosomal DNA</a> became more easily accessible in the 2000s, and since the mid-2010s, results of previously unattainable resolution, many of them based on full-genome analysis of ancient DNA, have been published at an accelerated pace.<sup id="cite_ref-Dutchen_2015_12-0" class="reference"><a href="#cite_note-Dutchen_2015-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Fu_2016_13-0" class="reference"><a href="#cite_note-Fu_2016-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Prehistory">Prehistory</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Genetic_history_of_Europe&action=edit&section=1" title="Edit section: Prehistory"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1237032888/mw-parser-output/.tmulti">.mw-parser-output .tmulti .multiimageinner{display:flex;flex-direction:column}.mw-parser-output .tmulti .trow{display:flex;flex-direction:row;clear:left;flex-wrap:wrap;width:100%;box-sizing:border-box}.mw-parser-output .tmulti .tsingle{margin:1px;float:left}.mw-parser-output .tmulti .theader{clear:both;font-weight:bold;text-align:center;align-self:center;background-color:transparent;width:100%}.mw-parser-output .tmulti .thumbcaption{background-color:transparent}.mw-parser-output .tmulti .text-align-left{text-align:left}.mw-parser-output .tmulti .text-align-right{text-align:right}.mw-parser-output .tmulti .text-align-center{text-align:center}@media all and (max-width:720px){.mw-parser-output .tmulti .thumbinner{width:100%!important;box-sizing:border-box;max-width:none!important;align-items:center}.mw-parser-output .tmulti .trow{justify-content:center}.mw-parser-output .tmulti .tsingle{float:none!important;max-width:100%!important;box-sizing:border-box;text-align:center}.mw-parser-output .tmulti .tsingle .thumbcaption{text-align:left}.mw-parser-output .tmulti .trow>.thumbcaption{text-align:center}}@media screen{html.skin-theme-clientpref-night .mw-parser-output .tmulti .multiimageinner img{background-color:white}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .tmulti .multiimageinner img{background-color:white}}</style><div class="thumb tmulti tright"><div class="thumbinner multiimageinner" style="width:332px;max-width:332px"><div class="trow"><div class="tsingle" style="width:330px;max-width:330px"><div class="thumbimage" style="height:154px;overflow:hidden"><span typeof="mw:File"><a href="/wiki/File:Neanderthal_distribution.jpg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/9/91/Neanderthal_distribution.jpg/328px-Neanderthal_distribution.jpg" decoding="async" width="328" height="154" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/91/Neanderthal_distribution.jpg/492px-Neanderthal_distribution.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/91/Neanderthal_distribution.jpg/656px-Neanderthal_distribution.jpg 2x" data-file-width="1693" data-file-height="795" /></a></span></div><div class="thumbcaption text-align-center">Distribution of the <a href="/wiki/Neanderthals" class="mw-redirect" title="Neanderthals">Neanderthals</a>, and main sites</div></div></div></div></div> <p>Due to natural selection, the percentage of Neanderthal DNA in ancient Europeans gradually decreased over time. From 45,000 BP to 7,000 BP, the percentage dropped from around 3–6% to 2%.<sup id="cite_ref-Fu_2016_13-1" class="reference"><a href="#cite_note-Fu_2016-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> The removal of Neanderthal-derived alleles occurred more frequently around genes than other parts of the genome.<sup id="cite_ref-Fu_2016_13-2" class="reference"><a href="#cite_note-Fu_2016-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Palaeolithic">Palaeolithic</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Genetic_history_of_Europe&action=edit&section=2" title="Edit section: Palaeolithic"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1236090951">.mw-parser-output .hatnote{font-style:italic}.mw-parser-output div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Peopling_of_Europe" class="mw-redirect" title="Peopling of Europe">Peopling of Europe</a>, <a href="/wiki/Archaic_human_admixture_with_modern_humans#Neanderthals" class="mw-redirect" title="Archaic human admixture with modern humans">Archaic human admixture with modern humans § Neanderthals</a>, and <a href="/wiki/Paleolithic_Europe" title="Paleolithic Europe">Paleolithic Europe</a></div> <p><a href="/wiki/Neanderthal" title="Neanderthal">Neanderthals</a> inhabited much of Europe and western Asia from as far back as 130,000 years ago. They existed in Europe as late as 30,000 years ago. They were eventually replaced by <a href="/wiki/Anatomically_modern_human" class="mw-redirect" title="Anatomically modern human">anatomically modern humans</a> (AMH; sometimes known as <a href="/wiki/Cro-Magnon" title="Cro-Magnon">Cro-Magnons</a>), who began to appear in Europe circa 40,000 years ago. Given that the two hominid species likely coexisted in Europe, anthropologists have long wondered whether the two interacted.<sup id="cite_ref-sciencemag1525_14-0" class="reference"><a href="#cite_note-sciencemag1525-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> The question was resolved only in 2010, when it was established that Eurasian populations exhibit Neanderthal admixture, estimated at 1.5–2.1% on average.<sup id="cite_ref-pruf13comal_15-0" class="reference"><a href="#cite_note-pruf13comal-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> The question now became whether this admixture had taken place in Europe, or rather in the Levant, prior to AMH migration into Europe. </p><p>There has also been speculation about the inheritance of specific genes from Neanderthals. For example, one <a href="/wiki/Tau_protein" title="Tau protein">MAPT</a> locus <a href="/wiki/Chromosome_17_(human)" class="mw-redirect" title="Chromosome 17 (human)">17q</a>21.3 which is split into deep genetic lineages H1 and H2. Since the H2 lineage seems restricted to European populations, several authors had argued for inheritance from Neanderthals beginning in 2005.<sup id="cite_ref-16" class="reference"><a href="#cite_note-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-17" class="reference"><a href="#cite_note-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-18" class="reference"><a href="#cite_note-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-19" class="reference"><a href="#cite_note-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-20" class="reference"><a href="#cite_note-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> However the preliminary results from the sequencing of the full Neanderthal Genome at that time (2009), failed to uncover evidence of interbreeding between Neanderthals and modern humans.<sup id="cite_ref-21" class="reference"><a href="#cite_note-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-22" class="reference"><a href="#cite_note-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> By 2010, findings by <a href="/wiki/Svante_P%C3%A4%C3%A4bo" title="Svante Pääbo">Svante Pääbo</a> (Max Planck Institute for Evolutionary Anthropology at Leipzig, Germany), Richard E. Green (University of California, Santa Cruz), and <a href="/wiki/David_Reich_(geneticist)" title="David Reich (geneticist)">David Reich</a> (Harvard Medical School), comparing the genetic material from the bones of three Neanderthals with that from five modern humans, did show a relationship between Neanderthals and modern people outside Africa. </p> <div class="mw-heading mw-heading4"><h4 id="Upper_Paleolithic">Upper Paleolithic</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Genetic_history_of_Europe&action=edit&section=3" title="Edit section: Upper Paleolithic"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main articles: <a href="/wiki/Upper_Paleolithic" title="Upper Paleolithic">Upper Paleolithic</a> and <a href="/wiki/Early_European_modern_humans" class="mw-redirect" title="Early European modern humans">Early European modern humans</a></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Expansion_of_early_modern_humans_from_Africa.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/16/Expansion_of_early_modern_humans_from_Africa.jpg/330px-Expansion_of_early_modern_humans_from_Africa.jpg" decoding="async" width="330" height="155" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/16/Expansion_of_early_modern_humans_from_Africa.jpg/495px-Expansion_of_early_modern_humans_from_Africa.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/16/Expansion_of_early_modern_humans_from_Africa.jpg/660px-Expansion_of_early_modern_humans_from_Africa.jpg 2x" data-file-width="1693" data-file-height="795" /></a><figcaption>Replacement of Neanderthals by early modern humans</figcaption></figure> <p>It is thought that modern humans began to inhabit Europe during the Upper Paleolithic about 40,000 years ago. Some evidence shows the spread of the <a href="/wiki/Aurignacian" title="Aurignacian">Aurignacian</a> culture.<sup id="cite_ref-Milisauskas_2002_23-0" class="reference"><a href="#cite_note-Milisauskas_2002-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 59">: 59 </span></sup> </p><p>From a purely patrilineal, <a href="/wiki/Y-DNA_haplogroup" class="mw-redirect" title="Y-DNA haplogroup">Y-chromosome</a> perspective, it appears that <a href="/wiki/Haplogroup_C1a2_(Y-DNA)" class="mw-redirect" title="Haplogroup C1a2 (Y-DNA)">Haplogroup C1a2</a>, <a href="/wiki/Haplogroup_F-M89" title="Haplogroup F-M89">F</a> and <a href="/wiki/Haplogroup_K2a_(Y-DNA)" title="Haplogroup K2a (Y-DNA)">K2a</a> may be those with the oldest presence in Europe. They have been found in some of the oldest human remains sequenced from the <a href="/wiki/Paleolithic_era" class="mw-redirect" title="Paleolithic era">paleolithic era</a>. However, other haplogroups are far more common among modern European males, because of later demographic changes. </p><p>Currently the oldest sample of <a href="/wiki/Haplogroup_I-M170" title="Haplogroup I-M170">Haplogroup I</a> (M170), which is now relatively common and widespread within Europe, has been found to be Krems WA3 from Lower Austria dating back to about 30–31,000 ybp.<sup id="cite_ref-24" class="reference"><a href="#cite_note-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> At about this time, an Upper Palaeolithic culture also appeared, known as the <a href="/wiki/Gravettian" title="Gravettian">Gravettian</a>.<sup id="cite_ref-Semino_2000_25-0" class="reference"><a href="#cite_note-Semino_2000-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> </p><p>Earlier research into Y-DNA had instead focused on <a href="/wiki/Haplogroup_R1" title="Haplogroup R1">haplogroup R1</a> (M173): the most populous lineage among living European males; R1 was also believed to have emerged ~ 40,000 BP in <a href="/wiki/Central_Asia" title="Central Asia">Central Asia</a>.<sup id="cite_ref-Semino_2000_25-1" class="reference"><a href="#cite_note-Semino_2000-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Wells_2001_26-0" class="reference"><a href="#cite_note-Wells_2001-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> However, it is now estimated that R1 emerged substantially more recently: a 2008 study dated the most recent common ancestor of haplogroup IJ to 38,500 and haplogroup R1 to 18,000 BP. This suggested that haplogroup IJ colonists formed the first wave and haplogroup R1 arrived much later.<sup id="cite_ref-27" class="reference"><a href="#cite_note-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> </p><p>Thus the genetic data suggests that, at least from the perspective of patrilineal ancestry, separate groups of modern humans took two routes into Europe: from the Middle East via the Balkans and another from Central Asia via the <a href="/wiki/Eurasian_Steppe" title="Eurasian Steppe">Eurasian Steppe</a>, to the north of the <a href="/wiki/Black_Sea" title="Black Sea">Black Sea</a>. </p><p>Martin Richards <i>et al.</i> found that 15–40% of extant mtDNA lineages trace back to the Palaeolithic migrations (depending on whether one allows for multiple founder events).<sup id="cite_ref-Richards_2000_28-0" class="reference"><a href="#cite_note-Richards_2000-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> MtDNA haplogroup U5, dated to be ~ 40–50 kYa, arrived during the first early upper Palaeolithic colonisation. Individually, it accounts for 5–15% of total mtDNA lineages. Middle U.P. movements are marked by the haplogroups HV, I and U4. HV split into Pre-V (around 26,000 years old) and the larger branch H, both of which spread over Europe, possibly via Gravettian contacts.<sup id="cite_ref-Semino_2000_25-2" class="reference"><a href="#cite_note-Semino_2000-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-29" class="reference"><a href="#cite_note-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Haplogroup_H_(mtDNA)" title="Haplogroup H (mtDNA)">Haplogroup H</a> accounts for about half the gene lines in Europe, with many subgroups. The above mtDNA lineages or their precursors, are most likely to have arrived into Europe via the Middle East. This contrasts with Y <a href="/wiki/DNA_profiling" title="DNA profiling">DNA evidence</a>, whereby some 50%-plus of male lineages are characterised by the R1 superfamily, which is of possible central Asian origin.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (November 2021)">citation needed</span></a></i>]</sup> Ornella Semino postulates that these differences "may be due in part to the apparent more recent molecular age of Y chromosomes relative to other loci, suggesting more rapid replacement of previous Y chromosomes. Gender-based differential migratory demographic behaviors will also influence the observed patterns of mtDNA and Y variation"<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (November 2021)">citation needed</span></a></i>]</sup>. </p> <div class="mw-heading mw-heading4"><h4 id="Last_Glacial_Maximum">Last Glacial Maximum</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Genetic_history_of_Europe&action=edit&section=4" title="Edit section: Last Glacial Maximum"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Early_European_modern_humans" class="mw-redirect" title="Early European modern humans">Early European modern humans</a></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Last_Glacial_Maximum" title="Last Glacial Maximum">Last Glacial Maximum</a> and <a href="/wiki/Last_Glacial_Maximum_refugia" title="Last Glacial Maximum refugia">Last Glacial Maximum refugia</a></div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Europe20000ya.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/de/Europe20000ya.png/250px-Europe20000ya.png" decoding="async" width="250" height="166" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/de/Europe20000ya.png/375px-Europe20000ya.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/de/Europe20000ya.png/500px-Europe20000ya.png 2x" data-file-width="1900" data-file-height="1264" /></a><figcaption>European LGM refuges, 20 kya<br /> <style data-mw-deduplicate="TemplateStyles:r981673959">.mw-parser-output .legend{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .legend-color{display:inline-block;min-width:1.25em;height:1.25em;line-height:1.25;margin:1px 0;text-align:center;border:1px solid black;background-color:transparent;color:black}.mw-parser-output .legend-text{}</style><div class="legend"><span class="legend-color mw-no-invert" style="background-color:#c54b00; color:white;"> </span> Solutrean and Proto-Solutrean Cultures</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r981673959"><div class="legend"><span class="legend-color mw-no-invert" style="background-color:#ca00b0; color:white;"> </span> Epi-Gravettian Culture</div> </figcaption></figure> <p>The Last Glacial Maximum ("LGM") started c. 30 ka BCE, at the end of <a href="/wiki/Marine_isotope_stage" class="mw-redirect" title="Marine isotope stage">MIS 3</a>, leading to a depopulation of Northern Europe. According to the classical model, people took refuge in climatic sanctuaries (or refugia) as follows: </p> <ul><li>Northern Iberia and Southwest <a href="/wiki/France" title="France">France</a>, together making up the "Franco-Cantabrian" refugium</li> <li>The Balkans</li> <li><a href="/wiki/Ukraine" title="Ukraine">Ukraine</a> and more generally the northern coast of the Black Sea<sup id="cite_ref-Semino_2000_25-3" class="reference"><a href="#cite_note-Semino_2000-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup></li> <li>Italy.<sup id="cite_ref-30" class="reference"><a href="#cite_note-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup></li></ul> <p>This event decreased the overall genetic diversity in Europe, a "result of drift, consistent with an inferred population bottleneck during the Last Glacial Maximum".<sup id="cite_ref-Wells_2001_26-1" class="reference"><a href="#cite_note-Wells_2001-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> As the glaciers receded from about 16,000–13,000 years ago, Europe began to be slowly repopulated by people from refugia, leaving genetic signatures.<sup id="cite_ref-Semino_2000_25-4" class="reference"><a href="#cite_note-Semino_2000-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> </p><p>Some Y haplogroup I clades appear to have diverged from their parental haplogroups sometime during or shortly after the LGM.<sup id="cite_ref-rootsi_31-0" class="reference"><a href="#cite_note-rootsi-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> </p><p>Cinnioglu sees evidence for the existence of an Anatolian refuge, which also harboured Hg R1b1b2.<sup id="cite_ref-32" class="reference"><a href="#cite_note-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> Today, R1b dominates the y chromosome landscape of western Europe, including the British Isles, suggesting that there could have been large population composition changes based on migrations after the LGM. </p><p>Semino, Passarino and Pericic place the origins of haplogroup <b>R1a</b> within the Ukrainian <a href="/wiki/Ice_age" title="Ice age">ice-age</a> refuge. Its current distribution in eastern Europe and parts of Scandinavia are in part reflective of a re-peopling of Europe from the southern Russian/Ukrainian steppes after the <a href="/wiki/Late_Glacial_Maximum" class="mw-redirect" title="Late Glacial Maximum">Late Glacial Maximum</a>.<sup id="cite_ref-Peričic_2005_33-0" class="reference"><a href="#cite_note-Peričic_2005-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Passarino_2001_34-0" class="reference"><a href="#cite_note-Passarino_2001-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Semino_2000_25-5" class="reference"><a href="#cite_note-Semino_2000-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> </p><p>From an mtDNA perspective, Richards <i>et al.</i> found that the majority of mtDNA diversity in Europe is accounted for by post-glacial re-expansions during the late upper Palaeolithic/ Mesolithic. "The regional analyses lend some support to the suggestion that much of western and central Europe was repopulated largely from the southwest when the climate improved. The lineages involved include much of the most common haplogroup, H, as well as much of K, T, W, and X." The study could not determine whether there were new migrations of mtDNA lineages from the near east during this period; a significant input was deemed unlikely.<sup id="cite_ref-Richards_2000_28-1" class="reference"><a href="#cite_note-Richards_2000-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> </p><p>The alternative model of more refugees was discussed by Bilton et al.<sup id="cite_ref-35" class="reference"><a href="#cite_note-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> </p><p>From a study of 51 individuals, researchers were able to identify five separate genetic clusters of ancient Eurasians during the LGM: the <a href="/wiki/Doln%C3%AD_V%C4%9Bstonice_(archaeology)" class="mw-redirect" title="Dolní Věstonice (archaeology)">Věstonice Cluster</a> (34,000–26,000 years ago), associated with the <a href="/wiki/Gravettian_culture" class="mw-redirect" title="Gravettian culture">Gravettian culture</a>; the Mal'ta Cluster (24,000–17,000), associated with the <a href="/wiki/Mal%27ta-Buret%27_culture" class="mw-redirect" title="Mal'ta-Buret' culture">Mal'ta-Buret' culture</a>, the <a href="/wiki/El_Mir%C3%B3n_Cave" title="El Mirón Cave">El Mirón Cluster</a> (19,000–14,000 years ago), associated with the <a href="/wiki/Magdalenian_culture" class="mw-redirect" title="Magdalenian culture">Magdalenian culture</a>; the <a href="/wiki/Ripari_Villabruna" title="Ripari Villabruna">Villabruna Cluster</a> (14,000–7,000 years ago) and the <a href="/wiki/Caucasus_hunter-gatherer" title="Caucasus hunter-gatherer">Satsurblia cluster</a> (13,000 to 10,000 years ago).<sup id="cite_ref-Fu_2016_13-3" class="reference"><a href="#cite_note-Fu_2016-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> </p><p>From around 37,000 years ago, all ancient Europeans began to share some ancestry with modern Europeans.<sup id="cite_ref-Fu_2016_13-4" class="reference"><a href="#cite_note-Fu_2016-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> This founding population is represented by GoyetQ116-1, a 35,000 year old specimen from Belgium.<sup id="cite_ref-Fu_2016_13-5" class="reference"><a href="#cite_note-Fu_2016-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> This lineage disappears from the record and is not found again until 19,000 BP in Spain at El Mirón, which shows strong affinities to GoyetQ116-1.<sup id="cite_ref-Fu_2016_13-6" class="reference"><a href="#cite_note-Fu_2016-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> During this interval, the distinct Věstonice Cluster is predominant in Europe, even at <a href="/wiki/Goyet_Caves" title="Goyet Caves">Goyet</a>.<sup id="cite_ref-Fu_2016_13-7" class="reference"><a href="#cite_note-Fu_2016-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> The re-expansion of the El Mirón Cluster coincided with warming temperatures following the retreat of the glaciers during the <a href="/wiki/Last_Glacial_Maximum" title="Last Glacial Maximum">Last Glacial Maximum</a>.<sup id="cite_ref-Fu_2016_13-8" class="reference"><a href="#cite_note-Fu_2016-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> From 37,000 to 14,000 years ago, the population of Europe consisted of an isolated population descended from a founding population that didn't interbreed significantly with other populations.<sup id="cite_ref-Dutchen_2016_36-0" class="reference"><a href="#cite_note-Dutchen_2016-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Mesolithic">Mesolithic</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Genetic_history_of_Europe&action=edit&section=5" title="Edit section: Mesolithic"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Mesolithic_Europe" class="mw-redirect" title="Mesolithic Europe">Mesolithic Europe</a>, <a href="/wiki/Western_hunter-gatherer" title="Western hunter-gatherer">Western hunter-gatherer</a>, and <a href="/wiki/Caucasus_hunter-gatherer" title="Caucasus hunter-gatherer">Caucasus hunter-gatherer</a></div> <p><a href="/wiki/Mesolithic" title="Mesolithic">Mesolithic</a> (post-LGM) populations had diverged significantly due to their relative isolation over several millennia, to the harsh selection pressures during the LGM, and to the <a href="/wiki/Founder_effect" title="Founder effect">founder effects</a> caused by the rapid expansion from <a href="/wiki/LGM_refugia" class="mw-redirect" title="LGM refugia">LGM refugia</a> in the beginning Mesolithic.<sup id="cite_ref-LGMpressures_3-1" class="reference"><a href="#cite_note-LGMpressures-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> By the end of the LGM, around 19 to 11 ka, the familiar varieties of Eurasian phenotypes had emerged. However, the lineage of Mesolithic hunter-gatherers of Western Europe (WHG) does not survive as a majority contribution in any modern population. They were most likely blue eyed, and retained the dark skin pigmentation of pre-LGM EEMH.<sup id="cite_ref-Mathieson_2015_37-0" class="reference"><a href="#cite_note-Mathieson_2015-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> The <a href="/wiki/HERC2" title="HERC2">HERC2</a> and <a href="/wiki/OCA2" class="mw-redirect" title="OCA2">OCA2</a> variations for blue eyes are derived from the WHG lineage were also found in the <a href="/wiki/Yamnaya_culture" title="Yamnaya culture">Yamnaya people</a>.<sup id="cite_ref-Mathieson_2015_37-1" class="reference"><a href="#cite_note-Mathieson_2015-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup><sup class="noprint Inline-Template" style="white-space:nowrap;">[<i><a href="/wiki/Category:All_self-contradictory_articles" title="Category:All self-contradictory articles"><span title="This text contradicts material elsewhere on this page. (August 2019)">contradictory</span></a></i>]</sup> </p><p>Around 14,000 years ago, the <a href="/wiki/Ripari_Villabruna" title="Ripari Villabruna">Villabruna Cluster</a> shifted away from GoyetQ116-1 affinity and started to show more affinity with the Near East, a shift which coincided with the warming temperatures of the <a href="/wiki/B%C3%B8lling-Aller%C3%B8d" class="mw-redirect" title="Bølling-Allerød">Bølling-Allerød</a> interstadial.<sup id="cite_ref-Fu_2016_13-9" class="reference"><a href="#cite_note-Fu_2016-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> This genetic shift shows that Near East populations had probably already begun moving into Europe during the end of the Upper Paleolithic, about 6,000 years earlier than previously thought, before the introduction of farming.<sup id="cite_ref-Dutchen_2016_36-1" class="reference"><a href="#cite_note-Dutchen_2016-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> A few specimens from the Villabruna Cluster also show genetic affinities for East Asians that are derived from gene flow.<sup id="cite_ref-Fu_2016_13-10" class="reference"><a href="#cite_note-Fu_2016-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Dutchen_2016_36-2" class="reference"><a href="#cite_note-Dutchen_2016-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> The <a href="/wiki/HERC2" title="HERC2">HERC2</a> variation for blue eyes first appears around 13,000 to 14,000 years ago in Italy and the Caucasus.<sup id="cite_ref-Fu_2016_13-11" class="reference"><a href="#cite_note-Fu_2016-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> The <a href="/wiki/Light_skin" title="Light skin">light skin</a> pigmentation characteristic of modern Europeans is estimated to have spread across Europe in a "selective sweep" during the Mesolithic (19 to 11 ka). The associated <a href="/wiki/TYRP1" title="TYRP1">TYRP1</a> <a href="/wiki/SLC24A5" class="mw-redirect" title="SLC24A5">SLC24A5</a> and <a href="/wiki/SLC45A2" class="mw-redirect" title="SLC45A2">SLC45A2</a> alleles emerge around 19 ka, still during the LGM, most likely in the Caucasus.<sup id="cite_ref-LGMpressures_3-2" class="reference"><a href="#cite_note-LGMpressures-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Jones2015_4-1" class="reference"><a href="#cite_note-Jones2015-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Neolithic">Neolithic</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Genetic_history_of_Europe&action=edit&section=6" title="Edit section: Neolithic"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Neolithic_Europe" title="Neolithic Europe">Neolithic Europe</a>, <a href="/wiki/Neolithic_Revolution" title="Neolithic Revolution">Neolithic Revolution</a>, <a href="/wiki/Early_European_Farmers" title="Early European Farmers">Early European Farmers</a>, and <a href="/wiki/Holocene" title="Holocene">Holocene</a></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Simplified_model_for_the_recent_demographic_history_of_Europeans.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/6/69/Simplified_model_for_the_recent_demographic_history_of_Europeans.jpg/290px-Simplified_model_for_the_recent_demographic_history_of_Europeans.jpg" decoding="async" width="290" height="205" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/69/Simplified_model_for_the_recent_demographic_history_of_Europeans.jpg/435px-Simplified_model_for_the_recent_demographic_history_of_Europeans.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/69/Simplified_model_for_the_recent_demographic_history_of_Europeans.jpg/580px-Simplified_model_for_the_recent_demographic_history_of_Europeans.jpg 2x" data-file-width="1995" data-file-height="1412" /></a><figcaption>Simplified model for the demographic history of Europeans during the <a href="/wiki/Neolithic" title="Neolithic">Neolithic</a> period in the <a href="/wiki/Neolithic_Revolution" title="Neolithic Revolution">introduction of agriculture</a><sup id="cite_ref-38" class="reference"><a href="#cite_note-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup></figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Genetic_matrilineal_distances_between_European_Neolithic_Linear_Pottery_Culture_populations_(5,500%E2%80%934,900_calibrated_BC)_and_modern_Western_Eurasian_populations.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/19/Genetic_matrilineal_distances_between_European_Neolithic_Linear_Pottery_Culture_populations_%285%2C500%E2%80%934%2C900_calibrated_BC%29_and_modern_Western_Eurasian_populations.jpg/290px-thumbnail.jpg" decoding="async" width="290" height="291" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/19/Genetic_matrilineal_distances_between_European_Neolithic_Linear_Pottery_Culture_populations_%285%2C500%E2%80%934%2C900_calibrated_BC%29_and_modern_Western_Eurasian_populations.jpg/435px-thumbnail.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/19/Genetic_matrilineal_distances_between_European_Neolithic_Linear_Pottery_Culture_populations_%285%2C500%E2%80%934%2C900_calibrated_BC%29_and_modern_Western_Eurasian_populations.jpg/580px-thumbnail.jpg 2x" data-file-width="1028" data-file-height="1033" /></a><figcaption>Ancient European Neolithic farmers were genetically closest to modern Near-Eastern/ Anatolian populations. Genetic matrilineal distances between European Neolithic Linear Pottery Culture populations (5,500–4,900 calibrated BC) and modern Western Eurasian populations.<sup id="cite_ref-39" class="reference"><a href="#cite_note-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup></figcaption></figure> <p>A big cline in genetic variation that has long been recognised in Europe seems to show important dispersals from the direction of the Middle East. This has often been linked to the spread of farming technology during the Neolithic, which has been argued to be one of the most important periods in determining modern European genetic diversity. </p><p>The Neolithic started with the introduction of farming, beginning in SE Europe approximately 10,000–3000 BCE, and extending into NW Europe between 4500 and 1700 BCE. During this era, the <a href="/wiki/Neolithic_Revolution" title="Neolithic Revolution">Neolithic Revolution</a> led to drastic economic as well as socio-cultural changes in Europe and this is also thought to have had a big effect on Europe's genetic diversity, especially concerning genetic lineages entering Europe from the Middle East into the Balkans. There were several phases of this period: </p> <ul><li>In a late European Mesolithic prelude to the Neolithic, it appears that Near Eastern peoples from areas that already had farming, and who also had sea-faring technology, had a transient presence in Greece (for example at <a href="/wiki/Franchthi_Cave" title="Franchthi Cave">Franchthi Cave</a>).<sup id="cite_ref-40" class="reference"><a href="#cite_note-40"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-41" class="reference"><a href="#cite_note-41"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup></li> <li>There is consensus that <a href="/wiki/Agricultural_machinery" title="Agricultural machinery">agricultural technology</a> and the main breeds of animals and plants which are farmed entered Europe from somewhere in the area of the <a href="/wiki/Fertile_Crescent" title="Fertile Crescent">Fertile Crescent</a> and specifically the <a href="/wiki/Levant" title="Levant">Levant</a> region from the <a href="/wiki/Sinai_Peninsula" title="Sinai Peninsula">Sinai</a> to <a href="/wiki/Anatolia" title="Anatolia">Southern Anatolia</a>.<sup id="cite_ref-Milisauskas_2002_23-1" class="reference"><a href="#cite_note-Milisauskas_2002-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 1143, 1150">: 1143, 1150 </span></sup><sup id="cite_ref-42" class="reference"><a href="#cite_note-42"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup> (Less certainly, this <a href="/wiki/Neolithic_Revolution" title="Neolithic Revolution">agricultural revolution</a> is sometimes argued to have in turn been partly triggered by movements of people and technology coming across the Sinai from Africa.) For more see <a href="/wiki/Fertile_Crescent#Cosmopolitan_diffusion" title="Fertile Crescent">Fertile Crescent: Cosmopolitan diffusion</a>.</li> <li>A later stage of the Neolithic, the so-called <a href="/wiki/Neolithic" title="Neolithic">Pottery Neolithic</a>, saw an introduction of <a href="/wiki/Pottery" title="Pottery">pottery</a> into the Levant, Balkans and Southern Italy (it had been present in the area of modern Sudan for some time before it is found in the <a href="/wiki/Eastern_Mediterranean" title="Eastern Mediterranean">Eastern Mediterranean</a>, but it is thought to have developed independently), and this may have also been a period of cultural transfer from the Levant into the Balkans.</li></ul> <p>An important issue regarding the genetic impact of neolithic technologies in Europe is the manner by which they were transferred into Europe. Farming was introduced by a significant migration of farmers from the Near East (Cavalli-Sforza's biological <a href="/wiki/Demic_diffusion" title="Demic diffusion">demic diffusion</a> model) or a "<a href="/wiki/Trans-cultural_diffusion" class="mw-redirect" title="Trans-cultural diffusion">cultural diffusion</a>" or a combination of the two, and <a href="/wiki/Population_genetics" title="Population genetics">population geneticists</a> have tried to clarify whether any genetic signatures of Near Eastern origin correspond to the expansion routes postulated by the archaeological evidence.<sup id="cite_ref-Milisauskas_2002_23-2" class="reference"><a href="#cite_note-Milisauskas_2002-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 146">: 146 </span></sup> </p><p>Martin Richards estimated that only 11% of European mtDNA is due to immigration in this period, suggesting that farming was spread primarily due to being adopted by indigenous Mesolithic populations, rather than due to immigration from Near East. Gene flow from SE to NW Europe seems to have continued in the Neolithic, the percentage significantly declining towards the British Isles. <a href="/wiki/Classical_genetics" title="Classical genetics">Classical genetics</a> also suggested that the largest admixture to the European Paleolithic/Mesolithic stock was due to the <a href="/wiki/Neolithic_Revolution" title="Neolithic Revolution">Neolithic revolution</a> of the 7th to 5th millennia BCE.<sup id="cite_ref-43" class="reference"><a href="#cite_note-43"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup> Three main mtDNA gene groups have been identified as contributing Neolithic entrants into Europe: J, T1 and U3 (in that order of importance). With others, they amount up to around 20% of the <a href="/wiki/Gene_pool" title="Gene pool">gene pool</a>.<sup id="cite_ref-Richards_2000_28-2" class="reference"><a href="#cite_note-Richards_2000-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> </p><p>In 2000, Semino's study on Y DNA revealed the presence of haplotypes belonging to the large clade <a href="/wiki/Haplogroup_E1b1b_(Y-DNA)" class="mw-redirect" title="Haplogroup E1b1b (Y-DNA)">E1b1b1</a> (E-M35). These were predominantly found in the southern Balkans, southern Italy and parts of Iberia. Semino connected this pattern, along with J haplogroup subclades, to be the Y-DNA component of Cavalli-Sforza's Neolithic demic-diffusion of farmers from the Near East.<sup id="cite_ref-Semino_2000_25-6" class="reference"><a href="#cite_note-Semino_2000-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: Here, the clade E-M35 is referred to as "Eu 4"">: Here, the clade E-M35 is referred to as "Eu 4" </span></sup> Rosser et al. rather saw it as a (direct) 'North African component' in European genealogy, although they did not propose a timing and mechanism to account for it.<sup id="cite_ref-Rosser_2000_44-0" class="reference"><a href="#cite_note-Rosser_2000-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Underhill_2007_45-0" class="reference"><a href="#cite_note-Underhill_2007-45"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup> also described E1b1b as representing a late-<a href="/wiki/Pleistocene" title="Pleistocene">Pleistocene</a> migration from Africa to Europe over the <a href="/wiki/Sinai_Peninsula" title="Sinai Peninsula">Sinai Peninsula</a> in <a href="/wiki/Egypt" title="Egypt">Egypt</a>, evidence for which does not show up in mitochondrial DNA.<sup id="cite_ref-46" class="reference"><a href="#cite_note-46"><span class="cite-bracket">[</span>46<span class="cite-bracket">]</span></a></sup> </p><p>Concerning timing the distribution and diversity of V13 however, Battaglia<sup id="cite_ref-Battaglia_2008_47-0" class="reference"><a href="#cite_note-Battaglia_2008-47"><span class="cite-bracket">[</span>47<span class="cite-bracket">]</span></a></sup> proposed an earlier movement whereby the E-M78* lineage ancestral to all modern E-V13 men moved rapidly out of a Southern Egyptian homeland and arrived in Europe with only <a href="/wiki/Mesolithic" title="Mesolithic">Mesolithic</a> technologies. They then suggest that the E-V13 sub-clade of E-M78 only expanded subsequently as native Balkan 'foragers-cum-farmers' adopted Neolithic technologies from the Near East. They propose that the first major dispersal of E-V13 from the Balkans may have been in the direction of the <a href="/wiki/Adriatic_Sea" title="Adriatic Sea">Adriatic Sea</a> with the <a href="/wiki/Neolithic" title="Neolithic">Neolithic</a> <a href="/wiki/Cardium_pottery" title="Cardium pottery">Impressed Ware</a> culture often referred to as <i>Impressa</i> or <a href="/wiki/Cardium_pottery" title="Cardium pottery">Cardial</a>,<sup id="cite_ref-Peričic_2005_33-1" class="reference"><a href="#cite_note-Peričic_2005-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> rather propose that the main route of E-V13 spread was along the Vardar-Morava-Danube river 'highway' system. </p><p>In contrast to Battaglia, Cruciani<sup id="cite_ref-Cruciani_2007_48-0" class="reference"><a href="#cite_note-Cruciani_2007-48"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup> tentatively suggested (i) a different point where the V13 mutation happened on its way from Egypt to the Balkans via the Middle East, and (ii) a later dispersal time. The authors proposed that the V13 mutation first appeared in western Asia, where it is found in low but significant frequencies, whence it entered the Balkans sometime after 11 kYa. It later experienced a rapid dispersal which he dated to c. 5300 years ago in Europe, coinciding with the Balkan Bronze Age. Like Peričic et al. they consider that "the dispersion of the E-V13 and J-M12 haplogroups seems to have mainly followed the river waterways connecting the southern Balkans to north-central Europe". </p><p>More recently, Lacan<sup id="cite_ref-Lacan_2011_49-0" class="reference"><a href="#cite_note-Lacan_2011-49"><span class="cite-bracket">[</span>49<span class="cite-bracket">]</span></a></sup> announced that a 7000-year-old skeleton in a Neolithic context in a Spanish funeral cave, was an E-V13 man. (The other specimens tested from the same site were in <a href="/wiki/Haplogroup_G2a" class="mw-redirect" title="Haplogroup G2a">haplogroup G2a</a>, which has been found in Neolithic contexts throughout Europe.) Using 7 STR markers, this specimen was identified as being similar to modern individuals tested in <a href="/wiki/Albania" title="Albania">Albania</a>, <a href="/wiki/Bosnia" class="mw-redirect" title="Bosnia">Bosnia</a>, <a href="/wiki/Greece" title="Greece">Greece</a>, <a href="/wiki/Corsica" title="Corsica">Corsica</a>, and <a href="/wiki/Provence" title="Provence">Provence</a>. The authors therefore proposed that, whether or not the modern distribution of E-V13 of today is a result of more recent events, E-V13 was already in Europe within the Neolithic, carried by early farmers from the Eastern Mediterranean to the Western Mediterranean, much earlier than the Bronze Age. This supports the proposals of Battaglia et al. rather than Cruciani et al. at least concerning earliest European dispersals, but E-V13 may have dispersed more than once. Even more recent than the Bronze Age, it has also been proposed that modern E-V13's modern distribution in Europe is at least partly caused by Roman era movements of people.<sup id="cite_ref-Bird_2007_50-0" class="reference"><a href="#cite_note-Bird_2007-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup> (See below.) </p><p>The migration of Neolithic farmers into Europe brought along several new adaptations.<sup id="cite_ref-Mathieson_2015_37-2" class="reference"><a href="#cite_note-Mathieson_2015-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> The variation for light skin colour was introduced to Europe by the <a href="/wiki/Early_European_Farmers" title="Early European Farmers">neolithic farmers</a>.<sup id="cite_ref-Mathieson_2015_37-3" class="reference"><a href="#cite_note-Mathieson_2015-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> After the arrival of the neolithic farmers, a <a href="/wiki/SLC22A4" title="SLC22A4">SLC22A4</a> mutation was selected for, a mutation which probably arose to deal with <a href="/wiki/Ergothioneine" title="Ergothioneine">ergothioneine</a> deficiency but increases the risk of <a href="/wiki/Ulcerative_colitis" title="Ulcerative colitis">ulcerative colitis</a>, <a href="/wiki/Coeliac_disease" title="Coeliac disease">coeliac disease</a>, and <a href="/wiki/Irritable_bowel_syndrome" title="Irritable bowel syndrome">irritable bowel syndrome</a>. </p> <div class="mw-heading mw-heading3"><h3 id="Bronze_Age">Bronze Age</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Genetic_history_of_Europe&action=edit&section=7" title="Edit section: Bronze Age"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Bronze_Age_Europe" title="Bronze Age Europe">Bronze Age Europe</a></div> <p>The <a href="/wiki/Bronze_Age" title="Bronze Age">Bronze Age</a> saw the development of long-distance <a href="/wiki/Trade" title="Trade">trading networks</a>, particularly along the Atlantic Coast and in the Danube valley. There was migration from Norway to <a href="/wiki/Orkney" title="Orkney">Orkney</a> and <a href="/wiki/Shetland" title="Shetland">Shetland</a> in this period (and to a lesser extent to mainland Scotland and Ireland). There was also migration from Germany to eastern England. Martin Richards estimated that there was about 4% mtDNA immigration to Europe in the Bronze Age. </p> <figure typeof="mw:File/Thumb"><a href="/wiki/File:IE_expansion.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/5a/IE_expansion.png/260px-IE_expansion.png" decoding="async" width="260" height="159" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/5a/IE_expansion.png/390px-IE_expansion.png 1.5x, //upload.wikimedia.org/wikipedia/commons/5/5a/IE_expansion.png 2x" data-file-width="450" data-file-height="275" /></a><figcaption>Scheme of Indo-European migrations from ca. 4000 to 1000 BC according to the <a href="/wiki/Kurgan_hypothesis" title="Kurgan hypothesis">Kurgan hypothesis</a></figcaption></figure> <p>Another theory about the origin of the <a href="/wiki/Indo-European_languages" title="Indo-European languages">Indo-European language</a> centres around a hypothetical <a href="/wiki/Proto-Indo-Europeans" title="Proto-Indo-Europeans">Proto-Indo-European</a> people, who, according to the <a href="/wiki/Kurgan_hypothesis" title="Kurgan hypothesis">Kurgan hypothesis</a>, can be traced to north of the Black and Caspian Seas at about 4500 BCE.<sup id="cite_ref-Reich_51-0" class="reference"><a href="#cite_note-Reich-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup> They <a href="/wiki/Domestication_of_the_horse" title="Domestication of the horse">domesticated the horse</a> and possibly invented the wooden disk <a href="/wiki/Wheel" title="Wheel">wheel</a>, and are considered to have spread their culture and genes across Europe.<sup id="cite_ref-Callaway_52-0" class="reference"><a href="#cite_note-Callaway-52"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup> The Y <a href="/wiki/Haplogroup_R1a" title="Haplogroup R1a">haplogroup R1a</a> is a proposed marker of these "Kurgan" genes, as is the Y <a href="/wiki/Haplogroup_R1b" title="Haplogroup R1b">Haplogroup R1b</a>, although these haplogroups as a whole may be much older than the language family.<sup id="cite_ref-53" class="reference"><a href="#cite_note-53"><span class="cite-bracket">[</span>53<span class="cite-bracket">]</span></a></sup> </p><p>In the far north, carriers of the <a href="/wiki/Haplogroup_N_(Y-DNA)" class="mw-redirect" title="Haplogroup N (Y-DNA)">Y-haplogroup N</a> arrived to Europe from <a href="/wiki/Siberia" title="Siberia">Siberia</a>, eventually expanding as far as <a href="/wiki/Finland" title="Finland">Finland</a>, though the specific timing of their arrival is uncertain. The most common North European subclade N1c1 is estimated to be around 8,000 years old. There is evidence of human settlement in Finland dating back to 8500 BCE, linked with the <a href="/wiki/Kunda_culture" title="Kunda culture">Kunda culture</a> and its putative ancestor, the <a href="/wiki/Swiderian_culture" title="Swiderian culture">Swiderian culture</a>, but the latter is thought to have a European origin. The geographical spread of haplogroup N in Europe is well aligned with the <a href="/wiki/Pit%E2%80%93Comb_Ware_culture" class="mw-redirect" title="Pit–Comb Ware culture">Pit–Comb Ware culture</a>, whose emergence is commonly dated c. 4200 BCE, and with the distribution of <a href="/wiki/Uralic_languages" title="Uralic languages">Uralic languages</a>. Mitochondrial DNA studies of <a href="/wiki/Sami_people" class="mw-redirect" title="Sami people">Sami people</a>, haplogroup <a href="/wiki/Haplogroup_U_(mtDNA)" class="mw-redirect" title="Haplogroup U (mtDNA)">U5</a> are consistent with multiple migrations to <a href="/wiki/Scandinavia" title="Scandinavia">Scandinavia</a> from <a href="/wiki/Volga_River" class="mw-redirect" title="Volga River">Volga</a>-<a href="/wiki/Ural_(region)" title="Ural (region)">Ural</a> region, starting 6,000 to 7,000 years before present.<sup id="cite_ref-54" class="reference"><a href="#cite_note-54"><span class="cite-bracket">[</span>54<span class="cite-bracket">]</span></a></sup> </p><p>The relationship between roles of European and Asian colonists in the prehistory of Finland is a point of some contention, and some scholars insist that Finns are "predominantly Eastern European and made up of people who trekked north from the Ukrainian refuge during the Ice Age".<sup id="cite_ref-55" class="reference"><a href="#cite_note-55"><span class="cite-bracket">[</span>55<span class="cite-bracket">]</span></a></sup> Farther east, the issue is less contentious. Haplogroup N carriers account for a significant part of all non-Slavic ethnic groups in northern <a href="/wiki/Russia" title="Russia">Russia</a>, including 37% of <a href="/wiki/Karelians" title="Karelians">Karelians</a>, 35% of <a href="/wiki/Komi_people" class="mw-redirect" title="Komi people">Komi people</a> (65% according to another study<sup id="cite_ref-Mirabal_2010_56-0" class="reference"><a href="#cite_note-Mirabal_2010-56"><span class="cite-bracket">[</span>56<span class="cite-bracket">]</span></a></sup>), 67% of <a href="/wiki/Mari_people" title="Mari people">Mari people</a>, as many as 98% of <a href="/wiki/Nenets_people" class="mw-redirect" title="Nenets people">Nenets people</a>, 94% of <a href="/wiki/Nganasans" class="mw-redirect" title="Nganasans">Nganasans</a>, and 86% to 94% of <a href="/wiki/Yakuts" title="Yakuts">Yakuts</a>.<sup id="cite_ref-57" class="reference"><a href="#cite_note-57"><span class="cite-bracket">[</span>57<span class="cite-bracket">]</span></a></sup> </p><p>The Yamnaya component contains partial ancestry from an Ancient North Eurasian component, a Paleolithic Siberian lineage but closely related to European hunter-gatherers, first identified in <a href="/wiki/Mal%27ta-Buret%27_culture" class="mw-redirect" title="Mal'ta-Buret' culture">Mal'ta</a>.<sup id="cite_ref-Jones_2015_58-0" class="reference"><a href="#cite_note-Jones_2015-58"><span class="cite-bracket">[</span>58<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-59" class="reference"><a href="#cite_note-59"><span class="cite-bracket">[</span>59<span class="cite-bracket">]</span></a></sup> According to Iosif Lazaridis, "the Ancient North Eurasian ancestry is proportionally the smallest component everywhere in Europe, never more than 20 percent, but we find it in nearly every European group we’ve studied."<sup id="cite_ref-Dutchen_2014_60-0" class="reference"><a href="#cite_note-Dutchen_2014-60"><span class="cite-bracket">[</span>60<span class="cite-bracket">]</span></a></sup> This genetic component does not come directly from the Mal'ta lineage itself, but a related lineage that separated from the Mal'ta lineage.<sup id="cite_ref-Fu_2016_13-12" class="reference"><a href="#cite_note-Fu_2016-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> </p><p>Up to a half of the Yamnaya component may have come from a <a href="/wiki/Caucasus_hunter-gatherer" title="Caucasus hunter-gatherer">Caucasus hunter-gatherer</a> strand.<sup id="cite_ref-Jones_2015_58-1" class="reference"><a href="#cite_note-Jones_2015-58"><span class="cite-bracket">[</span>58<span class="cite-bracket">]</span></a></sup> On November 16, 2015, in a study published in the journal <i><a href="/wiki/Nature_Communications" title="Nature Communications">Nature Communications</a></i>,<sup id="cite_ref-Jones_2015_58-2" class="reference"><a href="#cite_note-Jones_2015-58"><span class="cite-bracket">[</span>58<span class="cite-bracket">]</span></a></sup> geneticists announced that they had found a new fourth ancestral "tribe" or "strand" which had contributed to the modern European gene pool. They analysed genomes from two hunter-gatherers from Georgia which were 13,300 and 9,700 years old, and found that these Caucasus hunter-gatherers were probably the source of the farmer-like DNA in the Yamnaya. According to co-author Dr Andrea Manica of the University of Cambridge: "The question of where the Yamnaya come from has been something of a mystery up to now....we can now answer that as we've found that their genetic make-up is a mix of <a href="/wiki/Eastern_European_hunter-gatherer" class="mw-redirect" title="Eastern European hunter-gatherer">Eastern European hunter-gatherers</a> and a population from this pocket of Caucasus hunter-gatherers who weathered much of the last Ice Age in apparent isolation."<sup id="cite_ref-bbcnov16_61-0" class="reference"><a href="#cite_note-bbcnov16-61"><span class="cite-bracket">[</span>61<span class="cite-bracket">]</span></a></sup> </p><p>According to Lazaridis et al. (2016), a population related to the people of the <a href="/wiki/Chalcolithic" title="Chalcolithic">Chalcolithic</a> <a href="/wiki/Iran" title="Iran">Iran</a> contributed to roughly half of the ancestry of Yamnaya populations of the Pontic–Caspian steppe. These Iranian Chalcolithic people were a mixture of "the Neolithic people of western Iran, the Levant, and Caucasus Hunter Gatherers."<sup id="cite_ref-62" class="reference"><a href="#cite_note-62"><span class="cite-bracket">[</span>62<span class="cite-bracket">]</span></a></sup> </p><p>The genetic variations for <a href="/wiki/Lactase_persistence" title="Lactase persistence">lactase persistence</a> and greater height came with the Yamnaya people.<sup id="cite_ref-Mathieson_2015_37-4" class="reference"><a href="#cite_note-Mathieson_2015-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> The derived allele of the <a href="/wiki/KITLG" class="mw-redirect" title="KITLG">KITLG</a> gene (SNP rs12821256) that is associated with – and likely causal for – <a href="/wiki/Blond_hair" class="mw-redirect" title="Blond hair">blond hair</a> in Europeans is found in populations with eastern but not <a href="/wiki/Western_hunter-gatherers" class="mw-redirect" title="Western hunter-gatherers">western hunter-gatherers</a> ancestry, suggesting that its origin is in the <a href="/wiki/Ancient_North_Eurasian" title="Ancient North Eurasian">Ancient North Eurasian</a> (ANE) population and may have been spread in Europe by individuals with <a href="/wiki/Steppe_ancestry" class="mw-redirect" title="Steppe ancestry">steppe ancestry</a>. Consistent with this, the earliest known individual with the derived allele is an ANE individual from the Late Upper Paleolithic <a href="/wiki/Afontova_Gora" title="Afontova Gora">Afontova Gora</a> archaeological complex in central Siberia.<sup id="cite_ref-pmid29466330_63-0" class="reference"><a href="#cite_note-pmid29466330-63"><span class="cite-bracket">[</span>63<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Recent_history">Recent history</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Genetic_history_of_Europe&action=edit&section=8" title="Edit section: Recent history"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/History_of_Europe" title="History of Europe">History of Europe</a></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Demography_of_the_Roman_Empire" title="Demography of the Roman Empire">Demography of the Roman Empire</a>, <a href="/wiki/Migration_period" class="mw-redirect" title="Migration period">Migration period</a>, <a href="/wiki/Viking_expansion" title="Viking expansion">Viking expansion</a>, <a href="/wiki/Slavic_expansion" class="mw-redirect" title="Slavic expansion">Slavic expansion</a>, <a href="/wiki/Magyar_migration" class="mw-redirect" title="Magyar migration">Magyar migration</a>, <a href="/wiki/Muslim_conquest_of_Spain" class="mw-redirect" title="Muslim conquest of Spain">Muslim conquest of Spain</a>, <a href="/wiki/Turkic_expansion" class="mw-redirect" title="Turkic expansion">Turkic expansion</a>, <a href="/wiki/African_admixture_in_Europe" title="African admixture in Europe">African admixture in Europe</a>, <a href="/wiki/Ottoman_wars_in_Europe" title="Ottoman wars in Europe">Ottoman wars in Europe</a>, <a href="/wiki/Ostsiedlung" title="Ostsiedlung">Ostsiedlung</a>, <a href="/wiki/World_War_II_evacuation_and_expulsion" title="World War II evacuation and expulsion">World War II evacuation and expulsion</a>, <a href="/wiki/Population_transfer_in_the_Soviet_Union" title="Population transfer in the Soviet Union">Population transfer in the Soviet Union</a>, and <a href="/wiki/Immigration_to_Europe" title="Immigration to Europe">Immigration to Europe</a></div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Europe_DNA_01.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/0d/Europe_DNA_01.jpg/320px-Europe_DNA_01.jpg" decoding="async" width="320" height="170" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/0d/Europe_DNA_01.jpg/480px-Europe_DNA_01.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/0d/Europe_DNA_01.jpg/640px-Europe_DNA_01.jpg 2x" data-file-width="1085" data-file-height="577" /></a><figcaption>Overview map of recent (1st to 17th centuries AD) admixture events in Europe<sup id="cite_ref-pmid26387712_64-0" class="reference"><a href="#cite_note-pmid26387712-64"><span class="cite-bracket">[</span>64<span class="cite-bracket">]</span></a></sup></figcaption></figure> <p>Expansions of the <a href="/wiki/Roman_Empire" title="Roman Empire">Roman Empire</a> do not appear to have left distinct genetic signatures in Europe. Indeed, Romance-speaking populations in the Balkans, like <a href="/wiki/Romanians" title="Romanians">Romanians</a>, <a href="/wiki/Aromanians" title="Aromanians">Aromanians</a>, <a href="/wiki/Moldovans" title="Moldovans">Moldovans</a>, etc. have been found to genetically resemble neighbouring Greek and South Slavic-speaking peoples rather than modern Italians.<sup id="cite_ref-65" class="reference"><a href="#cite_note-65"><span class="cite-bracket">[</span>65<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-66" class="reference"><a href="#cite_note-66"><span class="cite-bracket">[</span>66<span class="cite-bracket">]</span></a></sup> Steven Bird has speculated that E1b1b1a was spread during the <a href="/wiki/Ancient_Rome" title="Ancient Rome">Roman era</a> through <a href="/wiki/Thracians" title="Thracians">Thracian</a> and <a href="/wiki/Dacia" title="Dacia">Dacian</a> populations from the Balkans into the rest of Europe.<sup id="cite_ref-Bird_2007_50-1" class="reference"><a href="#cite_note-Bird_2007-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup> </p><p>Concerning the late Roman period of (not only) <a href="/wiki/German_people" class="mw-redirect" title="German people">Germanic</a> "<i><a href="/wiki/V%C3%B6lkerwanderung" class="mw-redirect" title="Völkerwanderung">Völkerwanderung</a></i>", some suggestions have been made, at least for Britain, with Y haplogroup I1a being associated with <a href="/wiki/Anglo-Saxon" class="mw-redirect" title="Anglo-Saxon">Anglo-Saxon</a> immigration in eastern England, and R1a being associated with Norse immigration in northern Scotland.<sup id="cite_ref-67" class="reference"><a href="#cite_note-67"><span class="cite-bracket">[</span>67<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Genetics_of_modern_European_populations">Genetics of modern European populations</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Genetic_history_of_Europe&action=edit&section=9" title="Edit section: Genetics of modern European populations"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Ethnic_groups_in_Europe" title="Ethnic groups in Europe">Ethnic groups in Europe</a></div> <div class="mw-heading mw-heading3"><h3 id="Patrilineal_studies">Patrilineal studies</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Genetic_history_of_Europe&action=edit&section=10" title="Edit section: Patrilineal studies"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>There are four main Y-chromosome DNA <a href="/wiki/Haplogroup" title="Haplogroup">haplogroups</a> that account for most of Europe's <a href="/wiki/Patrilineality" title="Patrilineality">patrilineal descent</a>.<sup id="cite_ref-Semino_2000_25-7" class="reference"><a href="#cite_note-Semino_2000-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> </p> <ul><li>Haplogroup <a href="/wiki/Haplogroup_R1b_(Y-DNA)" class="mw-redirect" title="Haplogroup R1b (Y-DNA)">R1b</a> is common in Europe, particularly in <a href="/wiki/Western_Europe" title="Western Europe">Western Europe</a>, with the <a href="/wiki/Haplogroup_R1b1a1a2" class="mw-redirect" title="Haplogroup R1b1a1a2">R1b1a1a2</a> being the most common among Western Europeans.<sup id="cite_ref-balaresque_68-0" class="reference"><a href="#cite_note-balaresque-68"><span class="cite-bracket">[</span>68<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Myres2010_69-0" class="reference"><a href="#cite_note-Myres2010-69"><span class="cite-bracket">[</span>69<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Cruciani2010_70-0" class="reference"><a href="#cite_note-Cruciani2010-70"><span class="cite-bracket">[</span>70<span class="cite-bracket">]</span></a></sup> Nearly all of this R1b in Europe is in the form of the R1b1a2 (2011 name) (R-M269) sub-clade, specifically within the R-L23 sub-sub-clade whereas R1b found in <a href="/wiki/Central_Asia" title="Central Asia">Central Asia</a>, <a href="/wiki/Western_Asia" class="mw-redirect" title="Western Asia">western Asia</a> and <a href="/wiki/Africa" title="Africa">Africa</a> tends to be in other clades. It has also been pointed out that <a href="/wiki/Outlier" title="Outlier">outlier</a> types are present in Europe and are particularly notable in some areas such as Sardinia and Armenia.<sup id="cite_ref-71" class="reference"><a href="#cite_note-71"><span class="cite-bracket">[</span>71<span class="cite-bracket">]</span></a></sup> Haplogroup R1b frequencies vary from highs in western Europe in a steadily decreasing cline with growing distance from the Atlantic: 80–90% (<a href="/wiki/Welsh_people" title="Welsh people">Welsh</a>, <a href="/wiki/Basque_people" class="mw-redirect" title="Basque people">Basque</a>, <a href="/wiki/Irish_people" title="Irish people">Irish</a>, <a href="/wiki/Scottish_people" title="Scottish people">Scots</a>, <a href="/wiki/Bretons" title="Bretons">Bretons</a>) around 70–80% in Spain, Britain and France and around 40–60% in parts of <a href="/wiki/Eastern_Germany" class="mw-redirect" title="Eastern Germany">eastern Germany</a>, and northern Italy. It drops outside this area and is around 30% or less in areas such as southern Italy, <a href="/wiki/Poland" title="Poland">Poland</a>, the Balkans and <a href="/wiki/Cyprus" title="Cyprus">Cyprus</a>. R1b remains the most common clade as one moves east to Germany, while farther east, in Poland, R1a is more common (see below).<sup id="cite_ref-Kayser_2005_72-0" class="reference"><a href="#cite_note-Kayser_2005-72"><span class="cite-bracket">[</span>72<span class="cite-bracket">]</span></a></sup> In <a href="/wiki/Southeast_Europe" title="Southeast Europe">Southeast Europe</a>, R1b drops behind R1a in the area in and around Hungary and Serbia but is more common both to the south and north of this region.<sup id="cite_ref-Peričic_2005_33-2" class="reference"><a href="#cite_note-Peričic_2005-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> R1b in Western Europe is dominated by at least two sub-clades, R-U106, which is distributed from the east side of the <a href="/wiki/Rhine" title="Rhine">Rhine</a> into northern and central Europe (with a strong presence in England) and R-P312, which is most common west of the Rhine, including the <a href="/wiki/British_Isles" title="British Isles">British Isles</a>.<sup id="cite_ref-Myres2010_69-1" class="reference"><a href="#cite_note-Myres2010-69"><span class="cite-bracket">[</span>69<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Cruciani2010_70-1" class="reference"><a href="#cite_note-Cruciani2010-70"><span class="cite-bracket">[</span>70<span class="cite-bracket">]</span></a></sup></li> <li>Haplogroup <a href="/wiki/R1a" class="mw-redirect" title="R1a">R1a</a>, almost entirely in the R1a1a sub-clade, is prevalent in much of <a href="/wiki/Eastern_Europe" title="Eastern Europe">Eastern</a> and <a href="/wiki/Central_Europe" title="Central Europe">Central Europe</a> (also in <a href="/wiki/South_Asia" title="South Asia">South</a> and <a href="/wiki/Central_Asia" title="Central Asia">Central Asia</a>). For example, there is a sharp increase in R1a1 and decrease in R1b1b2 as one goes east from Germany to Poland.<sup id="cite_ref-Kayser_2005_72-1" class="reference"><a href="#cite_note-Kayser_2005-72"><span class="cite-bracket">[</span>72<span class="cite-bracket">]</span></a></sup> It also has a substantial presence in Scandinavia (particularly Norway).<sup id="cite_ref-Bowden_2008_73-0" class="reference"><a href="#cite_note-Bowden_2008-73"><span class="cite-bracket">[</span>73<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Dupuy_2021_74-0" class="reference"><a href="#cite_note-Dupuy_2021-74"><span class="cite-bracket">[</span>74<span class="cite-bracket">]</span></a></sup> In the Baltic countries R1a frequencies decrease from Lithuania (45%) to Estonia (around 30%).<sup id="cite_ref-Kasperaviciūte_2004_75-0" class="reference"><a href="#cite_note-Kasperaviciūte_2004-75"><span class="cite-bracket">[</span>75<span class="cite-bracket">]</span></a></sup></li> <li><a href="/wiki/Haplogroup_I_(Y-DNA)" class="mw-redirect" title="Haplogroup I (Y-DNA)">Haplogroup I</a> is found in the form of various sub-clades throughout Europe and is found at highest frequencies in the <a href="/wiki/Nordic_countries" title="Nordic countries">Nordic countries</a> as I1 (<a href="/wiki/Norway" title="Norway">Norway</a>, <a href="/wiki/Denmark" title="Denmark">Denmark</a>, <a href="/wiki/Sweden" title="Sweden">Sweden</a>, <a href="/wiki/Finland" title="Finland">Finland</a>) and in the <a href="/wiki/Balkans" title="Balkans">Balkan Peninsula</a> as I2a (<a href="/wiki/Bosnia_and_Herzegovina" title="Bosnia and Herzegovina">Bosnia and Herzegovina</a> 65%,<sup id="cite_ref-Mirabal_2010_56-1" class="reference"><a href="#cite_note-Mirabal_2010-56"><span class="cite-bracket">[</span>56<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Croatia" title="Croatia">Croatia</a> and <a href="/wiki/Serbia" title="Serbia">Serbia</a>). I1 is also frequent in <a href="/wiki/Germany" title="Germany">Germany</a>, <a href="/wiki/Great_Britain" title="Great Britain">Great Britain</a> and <a href="/wiki/Netherlands" title="Netherlands">Netherlands</a>, while I2a is frequent also in <a href="/wiki/Sardinia" title="Sardinia">Sardinia</a>, <a href="/wiki/Romania" title="Romania">Romania</a>/<a href="/wiki/Moldova" title="Moldova">Moldova</a>, <a href="/wiki/Bulgaria" title="Bulgaria">Bulgaria</a> and <a href="/wiki/Ukraine" title="Ukraine">Ukraine</a>. This clade is found at its highest expression by far in Europe and may have been there since before the <a href="/wiki/Last_Glacial_Maximum" title="Last Glacial Maximum">LGM</a>.<sup id="cite_ref-rootsi_31-1" class="reference"><a href="#cite_note-rootsi-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup></li> <li>Haplogroup <a href="/wiki/E1b1b" class="mw-redirect" title="E1b1b">E1b1b</a> (formerly known as E3b) was part of a migration of Neolithic farmers from the Middle East, which carried E1b1b at low to medium frequency and was introduced into Neolithic Middle Easterners throughout genetic drift from a migration from Africa into the Middle East associated with the <a href="/wiki/Afroasiatic_languages" title="Afroasiatic languages">Afroasiatic languages</a>. It is believed to have first appeared in Northeast Africa approximately 26,000 years ago and dispersed to North Africa and the Near East during the late Paleolithic and Mesolithic periods. E1b1b lineages are closely linked to the diffusion of Afroasiatic languages. Although present throughout Europe, it peaks in the southern <a href="/wiki/Balkan_region" class="mw-redirect" title="Balkan region">Balkan region</a> amongst <a href="/wiki/Albanians" title="Albanians">Albanians</a> and their neighbors. It is also common in Italy and the Iberian peninsula at lower frequency. Haplogroup <a href="/wiki/Haplogroup_E1b1b_(Y-DNA)" class="mw-redirect" title="Haplogroup E1b1b (Y-DNA)">E1b1b1</a>, mainly in the form of its E1b1b1a2 (E-V13) sub-clade, reaches frequencies above 47% around the area of <a href="/wiki/Kosovo" title="Kosovo">Kosovo</a>.<sup id="cite_ref-Peričic_2005_33-3" class="reference"><a href="#cite_note-Peričic_2005-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> This clade is thought to have arrived in Europe from western Asia either in the later Mesolithic,<sup id="cite_ref-Battaglia_2008_47-1" class="reference"><a href="#cite_note-Battaglia_2008-47"><span class="cite-bracket">[</span>47<span class="cite-bracket">]</span></a></sup> or the Neolithic.<sup id="cite_ref-cruciani2004_76-0" class="reference"><a href="#cite_note-cruciani2004-76"><span class="cite-bracket">[</span>76<span class="cite-bracket">]</span></a></sup> North Africa subclade E-M81 is also present in Sicily and Andalusia.</li></ul> <p>Putting aside small enclaves, there are also several haplogroups apart from the above four that are less prominent or most common only in certain areas of Europe. </p> <ul><li><a href="/wiki/Haplogroup_G-M201" title="Haplogroup G-M201">Haplogroup G</a>, a common haplogroup among European Neolithic farmers, is common in most parts of Europe at a low frequency, reaching peaks above 70% around <a href="/wiki/Georgia_(country)" title="Georgia (country)">Georgia</a> and among the <a href="/wiki/Madjars" title="Madjars">Madjars</a> (although living in Asia they border the eastern perimeter of Europe), up to 10% in Sardinia, 12% in Corsica and Uppsala (Sweden), 11% in the Balkans and Portugal, 10% in Spain and 9% in European Russia. This clade is also found in the Near East.</li> <li><a href="/wiki/Haplogroup_N_(Y-DNA)" class="mw-redirect" title="Haplogroup N (Y-DNA)">Haplogroup N</a>, is common only in the northeast of Europe and in the form of its N1c1 sub-clade reaches frequencies of approximately 60% among Finns and approximately 40% among Estonians, Latvians, and Lithuanians.</li> <li><a href="/wiki/Haplogroup_J_(Y-DNA)" title="Haplogroup J (Y-DNA)">Haplogroup J2</a>, in various sub-clades (J2a, J2b), is found in levels of around 15–30% in the Balkans (particularly <a href="/wiki/Greece" title="Greece">Greece</a>) and <a href="/wiki/Italy" title="Italy">Italy</a>. Haplogroup J2 is frequent in <a href="/wiki/Western_Asia" class="mw-redirect" title="Western Asia">Western Asia</a> and the <a href="/wiki/Eastern_Mediterranean" title="Eastern Mediterranean">Eastern Mediterranean</a>.<sup id="cite_ref-Semino_2004_77-0" class="reference"><a href="#cite_note-Semino_2004-77"><span class="cite-bracket">[</span>77<span class="cite-bracket">]</span></a></sup></li></ul> <div class="mw-heading mw-heading3"><h3 id="Matrilineal_studies">Matrilineal studies</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Genetic_history_of_Europe&action=edit&section=11" title="Edit section: Matrilineal studies"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>There have been a number of studies about the <a href="/wiki/Human_mitochondrial_DNA_haplogroup" title="Human mitochondrial DNA haplogroup">mitochondrial DNA haplogroups</a> (mtDNA) in Europe. In contrast to Y DNA haplogroups, mtDNA haplogroups did not show as much geographical patterning, but were more evenly ubiquitous. Apart from the outlying Saami, all Europeans are characterised by the predominance of haplogroups H, U and T. The lack of observable geographic structuring of mtDNA may be due to socio-cultural factors, namely the phenomena of <a href="/wiki/Polygyny" title="Polygyny">polygyny</a> and <a href="/wiki/Patrilocal_residence" title="Patrilocal residence">patrilocality</a>.<sup id="cite_ref-Rosser_2000_44-1" class="reference"><a href="#cite_note-Rosser_2000-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> </p><p>Genetic studies suggest some maternal gene flow to eastern Europe from eastern Asia or southern Siberia 13,000 – 6,600 years <a href="/wiki/Before_Present" title="Before Present">BP</a>.<sup id="cite_ref-po2012dmd_78-0" class="reference"><a href="#cite_note-po2012dmd-78"><span class="cite-bracket">[</span>78<span class="cite-bracket">]</span></a></sup> Analysis of Neolithic skeletons in the <a href="/wiki/Great_Hungarian_Plain" title="Great Hungarian Plain">Great Hungarian Plain</a> found a high frequency of eastern Asian mtDNA haplogroups, some of which survive in modern eastern European populations.<sup id="cite_ref-po2012dmd_78-1" class="reference"><a href="#cite_note-po2012dmd-78"><span class="cite-bracket">[</span>78<span class="cite-bracket">]</span></a></sup> Maternal gene flow to Europe from sub-Saharan Africa began as early as 11,000 years BP, although the majority of lineages, approximately 65%, are estimated to have arrived more recently, including during the Romanization period, the Arab conquests of southern Europe, and during the Atlantic slave trade.<sup id="cite_ref-cerezo_79-0" class="reference"><a href="#cite_note-cerezo-79"><span class="cite-bracket">[</span>79<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="European_population_sub-structure">European population sub-structure</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Genetic_history_of_Europe&action=edit&section=12" title="Edit section: European population sub-structure"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Genetically, Europe is relatively homogeneous, but distinct sub-population patterns of various types of genetic markers have been found,<sup id="cite_ref-Cavalli-Sforza_1993_80-0" class="reference"><a href="#cite_note-Cavalli-Sforza_1993-80"><span class="cite-bracket">[</span>80<span class="cite-bracket">]</span></a></sup> particularly along a southeast–northwest cline.<sup id="cite_ref-Lao_2008_81-0" class="reference"><a href="#cite_note-Lao_2008-81"><span class="cite-bracket">[</span>81<span class="cite-bracket">]</span></a></sup> For example, Cavalli-Sforza's principal component analyses revealed five major clinal patterns throughout Europe, and similar patterns have continued to be found in more recent studies.<sup id="cite_ref-Cavalli-Sforza_1993_80-1" class="reference"><a href="#cite_note-Cavalli-Sforza_1993-80"><span class="cite-bracket">[</span>80<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 291–296">: 291–296 </span></sup> </p> <ol><li>A cline of genes with highest frequencies in the <a href="/wiki/Middle_East" title="Middle East">Middle East</a>, spreading to lowest levels northwest. Cavalli-Sforza originally described this as faithfully reflecting the spread of agriculture in Neolithic times. This has been the general tendency in interpretation of all genes with this pattern.</li> <li>A cline of genes with highest frequencies among <a href="/wiki/Finns" title="Finns">Finnish</a> and <a href="/wiki/Sami_people" class="mw-redirect" title="Sami people">Sami</a> in the extreme north east, and spreading to lowest frequencies in the south west.</li> <li>A cline of genes with highest frequencies in the area of the lower <a href="/wiki/Don_River_(Russia)" class="mw-redirect" title="Don River (Russia)">Don</a> and <a href="/wiki/Volga_River" class="mw-redirect" title="Volga River">Volga</a> rivers in <a href="/wiki/Southern_Federal_District" title="Southern Federal District">southern Russia</a>, and spreading to lowest frequencies in Spain, <a href="/wiki/Southern_Italy" title="Southern Italy">Southern Italy</a>, <a href="/wiki/Greece" title="Greece">Greece</a> and the areas inhabited by <a href="/wiki/Sami_languages" class="mw-redirect" title="Sami languages">Saami speakers</a> in the extreme north of <a href="/wiki/Scandinavia" title="Scandinavia">Scandinavia</a>. Cavalli-Sforza associated this with the spread of Indo-European languages, which he links in turn to a "secondary expansion" after the spread of agriculture, associated with animal grazing.</li> <li>A cline of genes with highest frequencies in the <a href="/wiki/Balkans" title="Balkans">Balkans</a> and Southern Italy, spreading to lowest levels in Britain and the Basque country. Cavalli-Sforza associates this with "the Greek expansion, which reached its peak in historical times around 1000 and 500 BCE but which certainly began earlier".</li> <li>A cline of genes with highest frequencies in the <a href="/wiki/Basque_Country_(greater_region)" title="Basque Country (greater region)">Basque country</a>, and lower levels beyond the area of Iberia and <a href="/wiki/Southern_France" title="Southern France">Southern France</a>. In perhaps the most well-known conclusion from Cavalli-Sforza, this weakest of the five patterns was described as isolated remnants of the pre-Neolithic population of Europe, "who at least partially withstood the expansion of the cultivators". It corresponds roughly to the geographical spread of <a href="/wiki/Rh_blood_group_system" title="Rh blood group system">rhesus negative</a> blood types. In particular, the conclusion that the Basques are a genetic isolate has become widely discussed, but also a controversial conclusion.</li></ol> <p>He also created a phylogenetic tree to analyse the internal relationships among Europeans. He found four major 'outliers'- <a href="/wiki/Basque_people" class="mw-redirect" title="Basque people">Basques</a>, <a href="/wiki/Sami_people" class="mw-redirect" title="Sami people">Sami</a>, <a href="/wiki/Sardinians" class="mw-redirect" title="Sardinians">Sardinians</a> and <a href="/wiki/Icelanders" title="Icelanders">Icelanders</a>;<sup id="cite_ref-82" class="reference"><a href="#cite_note-82"><span class="cite-bracket">[</span>82<span class="cite-bracket">]</span></a></sup> a result he attributed to their relative isolation (note: the Icelanders and the Sardinians speak <a href="/wiki/Indo-European_languages" title="Indo-European languages">Indo-European languages</a>, while the other two groups do not). <a href="/wiki/Greeks" title="Greeks">Greeks</a> and <a href="/wiki/Yugoslavs" title="Yugoslavs">Yugoslavs</a> represented a second group of less extreme outliers. The remaining populations clustered into several groups : "<a href="/wiki/Celtic_nations" title="Celtic nations">Celtic</a>", "<a href="/wiki/Germanic_peoples" title="Germanic peoples">Germanic</a>", "south-western Europeans", "<a href="/wiki/Scandinavia" title="Scandinavia">Scandinavians</a>" and "eastern Europeans".<sup id="cite_ref-Cavalli-Sforza_1993_80-2" class="reference"><a href="#cite_note-Cavalli-Sforza_1993-80"><span class="cite-bracket">[</span>80<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 268">: 268 </span></sup> </p><p>A study conducted in May of 2009<sup id="cite_ref-83" class="reference"><a href="#cite_note-83"><span class="cite-bracket">[</span>83<span class="cite-bracket">]</span></a></sup> researching 19 populations from Europe using 270,000 SNPs highlighted the genetic diversity of European populations corresponding to the northwest to southeast gradient and distinguished "four several distinct regions" within Europe: </p> <ul><li>Finland, showing the greatest distance to the rest of Europeans.</li> <li>the <a href="/wiki/Baltic_region" title="Baltic region">Baltic region</a> (<a href="/wiki/Estonia" title="Estonia">Estonia</a>, <a href="/wiki/Latvia" title="Latvia">Latvia</a> and <a href="/wiki/Lithuania" title="Lithuania">Lithuania</a>), western <a href="/wiki/Russia" title="Russia">Russia</a> and eastern <a href="/wiki/Poland" title="Poland">Poland</a>.</li> <li>Central and Western Europe.</li> <li><a href="/wiki/Italy" title="Italy">Italy</a>, due to the alps acting as a great genetic barrier.</li></ul> <p>In this study, barrier analysis revealed "genetic barriers" between Finland, Italy and other countries and that barriers could also be demonstrated within Finland (between Helsinki and Kuusamo) and Italy (between northern and southern part, Fst=0.0050). Fst (<a href="/wiki/Fixation_index" title="Fixation index">Fixation index</a>) was found to correlate considerably with geographic distances ranging from ≤0.0010 for neighbouring populations to 0.0200–0.0230 for Southern Italy and Finland. For comparisons, pair-wise Fst of non-European samples were as follows: Europeans – Africans (Yoruba) 0.1530; Europeans – Chinese 0.1100; Africans (Yoruba) – Chinese 0.1900.<sup id="cite_ref-84" class="reference"><a href="#cite_note-84"><span class="cite-bracket">[</span>84<span class="cite-bracket">]</span></a></sup> </p><p>A study by Chao Tian in August 2009 extended the analysis of European population genetic structure to include additional southern European groups and levantine populations (<a href="/wiki/Palestinians" title="Palestinians">Palestinians</a>, <a href="/wiki/Druzes" class="mw-redirect" title="Druzes">Druzes</a>...) from the Near-East. This study determined autosomal Fst between 18 population groups and concluded that, in general, genetic distances corresponded to geographical relationships with smaller values between population groups with origins in neighbouring countries/regions (for example, <a href="/wiki/Greeks" title="Greeks">Greeks</a>/<a href="/wiki/Tuscany" title="Tuscany">Tuscans</a>: Fst=0.0010, <a href="/wiki/Greeks" title="Greeks">Greeks</a>/<a href="/wiki/Palestinians" title="Palestinians">Palestinians</a>: Fst=0.0057) compared with those from very different regions in Europe (for example <a href="/wiki/Greeks" title="Greeks">Greeks</a>/<a href="/wiki/Swedes" title="Swedes">Swedish</a>: Fst=0.0087, <a href="/wiki/Greeks" title="Greeks">Greeks</a>/<a href="/wiki/Russians" title="Russians">Russians</a>: Fst=0.0108).<sup id="cite_ref-European_population_genetic_substru_85-0" class="reference"><a href="#cite_note-European_population_genetic_substru-85"><span class="cite-bracket">[</span>85<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Autosomal_DNA">Autosomal DNA</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Genetic_history_of_Europe&action=edit&section=13" title="Edit section: Autosomal DNA"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Seldin (2006) used over 5,000 autosomal SNPs. It showed "a consistent and reproducible distinction between ‘northern’ and ‘southern’ <a href="/wiki/Demographics_of_Europe" title="Demographics of Europe">European population</a> groups". Most individual participants with <a href="/wiki/Southern_Europe" title="Southern Europe">southern European</a> ancestry (<a href="/wiki/Italian_people" class="mw-redirect" title="Italian people">Italians</a>, <a href="/wiki/Greeks" title="Greeks">Greeks</a>, <a href="/wiki/Portuguese_people" title="Portuguese people">Portuguese</a>, <a href="/wiki/Spaniards" title="Spaniards">Spaniards</a>), and <a href="/wiki/Ashkenazi_Jews" title="Ashkenazi Jews">Ashkenazi Jews</a> have >85% membership in the southern population; and most northern, western, central, and eastern Europeans (<a href="/wiki/Swedes" title="Swedes">Swedes</a>, <a href="/wiki/English_people" title="English people">English</a>, <a href="/wiki/Irish_people" title="Irish people">Irish</a>, <a href="/wiki/Germans" title="Germans">Germans</a>, and <a href="/wiki/Ukrainians" title="Ukrainians">Ukrainians</a>) have >90% in the northern population group. Many of the participants in this study were American citizens who self-identified with different European ethnicities based on self-reported familial pedigree.<sup id="cite_ref-86" class="reference"><a href="#cite_note-86"><span class="cite-bracket">[</span>86<span class="cite-bracket">]</span></a></sup> </p><p>A similar study in 2007 using samples predominantly from Europe found that the most important genetic differentiation in Europe occurs on a line from the north to the south-east (<a href="/wiki/Northern_Europe" title="Northern Europe">northern Europe</a> to the Balkans), with another east–west axis of differentiation across Europe. Its findings were consistent with earlier results based on mtDNA and Y-chromosomal DNA that support the theory that modern Iberians (Spanish and Portuguese) hold the most ancient European genetic ancestry, as well as separating Basques and Sami from other European populations.<sup id="cite_ref-sitesled2007_87-0" class="reference"><a href="#cite_note-sitesled2007-87"><span class="cite-bracket">[</span>87<span class="cite-bracket">]</span></a></sup> </p><p>It suggested that the <a href="/wiki/English_people" title="English people">English</a> and Irish cluster with other Northern and Eastern Europeans such as <a href="/wiki/Germans" title="Germans">Germans</a> and <a href="/wiki/Polish_people" title="Polish people">Poles</a>, while some Basque and Italian individuals also clustered with Northern Europeans. Despite these stratifications, it noted that "there is low apparent diversity in Europe with the entire continent-wide samples only marginally more dispersed than single population samples elsewhere in the world".<sup id="cite_ref-sitesled2007_87-1" class="reference"><a href="#cite_note-sitesled2007-87"><span class="cite-bracket">[</span>87<span class="cite-bracket">]</span></a></sup> </p><p>In 2008, two international research teams published analyses of large-scale genotyping of large samples of Europeans, using over 300,000 autosomal SNPs. With the exception of usual isolates such as <a href="/wiki/Basques" title="Basques">Basques</a>, <a href="/wiki/Finns" title="Finns">Finns</a> and <a href="/wiki/Sardinians" class="mw-redirect" title="Sardinians">Sardinians</a>, the European population lacked sharp discontinuities (clustering) as previous studies have found (see Seldin <i>et al.</i> 2006 and Bauchet <i>et al.</i> 2007<sup id="cite_ref-sitesled2007_87-2" class="reference"><a href="#cite_note-sitesled2007-87"><span class="cite-bracket">[</span>87<span class="cite-bracket">]</span></a></sup>), although there was a discernible south to north gradient. Overall, they found only a low level of genetic differentiation between subpopulations, and differences which did exist were characterised by a strong continent-wide correlation between geographic and genetic distance. In addition, they found that diversity was greatest in southern Europe due a larger effective population size and/or <a href="/wiki/Population_growth" title="Population growth">population expansion</a> from southern to northern Europe.<sup id="cite_ref-Lao_2008_81-1" class="reference"><a href="#cite_note-Lao_2008-81"><span class="cite-bracket">[</span>81<span class="cite-bracket">]</span></a></sup> The researchers take this observation to imply that genetically, Europeans are not distributed into discrete populations.<sup id="cite_ref-88" class="reference"><a href="#cite_note-88"><span class="cite-bracket">[</span>88<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Lao_2008_81-2" class="reference"><a href="#cite_note-Lao_2008-81"><span class="cite-bracket">[</span>81<span class="cite-bracket">]</span></a></sup> </p><p>Two whole-genome studies of the two Eastern European populations in Ukraine (<a href="/wiki/Ukraine" title="Ukraine">Ukrainians</a> from <a href="/wiki/Ukraine" title="Ukraine">Ukraine</a>) and Russia (<a href="/wiki/Russia" title="Russia">Russians</a> from <a href="/wiki/Russia" title="Russia">Russia</a>) showed genomic diversity, which has not been represented in the previous genomic surveys, as studies in Europe are mostly biased towards the populations in the western part of the continent.<sup id="cite_ref-89" class="reference"><a href="#cite_note-89"><span class="cite-bracket">[</span>89<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-90" class="reference"><a href="#cite_note-90"><span class="cite-bracket">[</span>90<span class="cite-bracket">]</span></a></sup> Within <a href="/wiki/Russia" title="Russia">Russia</a>, <a href="/wiki/Komi_people" class="mw-redirect" title="Komi people">Komi people</a>, who live in the northeastern regions and are part of the <a href="/wiki/Uralic" class="mw-redirect" title="Uralic">Uralic</a> language family that also includes <a href="/wiki/Finns" title="Finns">Finns</a>, form a pole of genetic diversity that is distinct from other populations, and characterized by a higher European hunter-gatherer (WHG) and Ancient North Eurasian ancestry.<sup id="cite_ref-plosone130307_91-0" class="reference"><a href="#cite_note-plosone130307-91"><span class="cite-bracket">[</span>91<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-The_Arrival_of_Siberian_Ancestry_Co_92-0" class="reference"><a href="#cite_note-The_Arrival_of_Siberian_Ancestry_Co-92"><span class="cite-bracket">[</span>92<span class="cite-bracket">]</span></a></sup> </p><p>According to geneticist <a href="/wiki/David_Reich_(geneticist)" title="David Reich (geneticist)">David Reich</a>, based on <a href="/wiki/Ancient_DNA" title="Ancient DNA">ancient human genomes</a> that his laboratory sequenced in 2016, Europeans descend from a mixture of four West-Eurasian ancestral components, namely WHG (western hunter-gatherers), EHG, Neolithic farmers from the Levant/Anatolia as well as from Neolithic farmers from Iran (often summarized as "EEF"; early European farmers), in varying degrees.<sup id="cite_ref-93" class="reference"><a href="#cite_note-93"><span class="cite-bracket">[</span>93<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-94" class="reference"><a href="#cite_note-94"><span class="cite-bracket">[</span>94<span class="cite-bracket">]</span></a></sup> </p><p>Siberian geneflow is found among several Uralic-speaking European ethnic groups. This Siberian component is itself a composition of Ancient North Eurasian and East Asian-related ancestry from Eastern Siberia, maximized among <a href="/wiki/Evenks" class="mw-redirect" title="Evenks">Evenks</a> and <a href="/wiki/Evens" title="Evens">Evens</a> or <a href="/wiki/Nganasans" class="mw-redirect" title="Nganasans">Nganasans</a>. The spread of this ancestry is linked by some geneticists to the dispersal of <a href="/wiki/Uralic_languages" title="Uralic languages">Uralic languages</a>, others however maintain that the Uralic languages spread prior to the arrival of Siberian geneflow, which is a secondary source of diversity within Uralic-speaking populations.<sup id="cite_ref-95" class="reference"><a href="#cite_note-95"><span class="cite-bracket">[</span>95<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-96" class="reference"><a href="#cite_note-96"><span class="cite-bracket">[</span>96<span class="cite-bracket">]</span></a></sup> Genetic data points to a <a href="/wiki/Western_Siberia" title="Western Siberia">western Siberian</a> <a href="/wiki/Hunter-gatherer" title="Hunter-gatherer">hunter-gatherer</a> origin of the observed Siberian geneflow among Uralic-speaking groups. Western Siberian hunter-gatherers were characterized by high <a href="/wiki/Ancient_North_Eurasian" title="Ancient North Eurasian">Ancient North Eurasian</a> ancestry and lower amounts of Eastern Siberian admixture. Genetic data on <a href="/wiki/Volga_Tatars" title="Volga Tatars">Volga Tatars</a> or <a href="/wiki/Chuvash_people" title="Chuvash people">Chuvash</a>, found among "<i>Western Turkic speakers, like Chuvash and Volga Tatar, the East Asian component was detected only in low amounts (~ 5%)</i>".<sup id="cite_ref-97" class="reference"><a href="#cite_note-97"><span class="cite-bracket">[</span>97<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-98" class="reference"><a href="#cite_note-98"><span class="cite-bracket">[</span>98<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/East_Asians" class="mw-redirect" title="East Asians">East Asian</a> ancestry is found at low frequency among some Europeans, such as <a href="/wiki/British_people" title="British people">British</a> (2.5 ± 1%), <a href="/wiki/Orcadians" title="Orcadians">Orcadians</a> (3.8 ± 1%), <a href="/wiki/French_people" title="French people">French</a> (0.7 ± 0.8%) and <a href="/wiki/Germans" title="Germans">Germans</a> (0.7 ± 0.8%). Finns and Russians have more than 12% East Asian ancestry, deriving from historic intermarriages with Mongolian populations.<sup id="cite_ref-99" class="reference"><a href="#cite_note-99"><span class="cite-bracket">[</span>99<span class="cite-bracket">]</span></a></sup> But a 2017 study finds no evidence of Asian admixture among Russians, except for <a href="/wiki/Novosibirsk" title="Novosibirsk">Novosibirsk</a> residents and <a href="/wiki/Old_Believers" title="Old Believers">Old Believers</a> in Siberia.<sup id="cite_ref-100" class="reference"><a href="#cite_note-100"><span class="cite-bracket">[</span>100<span class="cite-bracket">]</span></a></sup>The <a href="/wiki/Lipka_Tatars" title="Lipka Tatars">Lipka Tatars</a>, a <a href="/wiki/Turkic_peoples" title="Turkic peoples">Turkic minority</a> in Belarus, have significant East Eurasian ancestry, making up one-third of their genome.<sup id="cite_ref-srep30197_101-0" class="reference"><a href="#cite_note-srep30197-101"><span class="cite-bracket">[</span>101<span class="cite-bracket">]</span></a></sup> </p><p>Like other Eurasian populations, Mesolithic, Neolithic or Bronze Age ancestries are not homogenously distributed in European populations. But WHG-related ancestries are highest in present-day individuals from the Baltic States, Belarus, Poland and Russia whilst EHG-related ancestries are highest in Finland and Estonia. Steppe-related ancestries are found in high levels in northern Europe, peaking in Ireland, Iceland, Norway and Sweden, but decrease further south, especially in southern Europe, where Neolithic Anatolian-related farmer ancestries dominate.<sup id="cite_ref-102" class="reference"><a href="#cite_note-102"><span class="cite-bracket">[</span>102<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Autosomal_genetic_distances_(Fst)_based_on_SNPs_(2009)"><span id="Autosomal_genetic_distances_.28Fst.29_based_on_SNPs_.282009.29"></span>Autosomal genetic distances (Fst) based on SNPs (2009)</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Genetic_history_of_Europe&action=edit&section=14" title="Edit section: Autosomal genetic distances (Fst) based on SNPs (2009)"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The genetic distance between populations is often measured by <a href="/wiki/Fixation_index" title="Fixation index">Fixation index</a> (Fst), based on genetic polymorphism data, such as <a href="/wiki/Single-nucleotide_polymorphism" title="Single-nucleotide polymorphism">single-nucleotide polymorphisms (SNPs)</a> or <a href="/wiki/Microsatellite_(genetics)" class="mw-redirect" title="Microsatellite (genetics)">microsatellites</a>. Fst is a special case of <a href="/wiki/F-statistics" title="F-statistics">F-statistics</a>, the concept developed in the 1920s by <a href="/wiki/Sewall_Wright" title="Sewall Wright">Sewall Wright</a>. Fst is simply the correlation of randomly chosen alleles within the same sub-population relative to that found in the entire population. It is often expressed as the proportion of genetic diversity due to allele frequency differences among populations. </p><p>The values range from 0 to 1. A zero value implies that the two populations are panmictic, that they are interbreeding freely. A value of one would imply that the two populations are completely separate. The greater the Fst value, the greater the genetic distance. Essentially, these low Fst values suggest that the majority of genetic variation is at the level of individuals within the same population group (~ 85%); whilst belonging to a different population group within same ‘race’/ continent, and even to different racial/ continental groups added a much smaller degree of variation (3–8%; 6–11%, respectively). </p> <table class="wikitable" style="text-align:center; width:80%;"> <caption>Intra-European/Mediterranean <a href="/wiki/Fixation_index" title="Fixation index">Autosomal genetic distances (Fst)</a> based on 3,500 <a href="/wiki/Single-nucleotide_polymorphism" title="Single-nucleotide polymorphism">SNPs</a> using the Weir and Cockerham algorithm<sup id="cite_ref-European_population_genetic_substru_85-1" class="reference"><a href="#cite_note-European_population_genetic_substru-85"><span class="cite-bracket">[</span>85<span class="cite-bracket">]</span></a></sup> </caption> <tbody><tr> <th scope="col"> </th> <th scope="col"><a href="/wiki/Italian_Americans" title="Italian Americans">Italian Americans</a> </th> <th scope="col"><a href="/wiki/Palestinians" title="Palestinians">Palestinians</a> </th> <th scope="col"><a href="/wiki/Swedes" title="Swedes">Swedes</a> </th> <th scope="col"><a href="/wiki/Druzes" class="mw-redirect" title="Druzes">Druzes</a> </th> <th scope="col"><a href="/wiki/Spaniards" title="Spaniards">Spaniards</a> </th> <th scope="col"><a href="/wiki/Germans" title="Germans">Germans</a> </th> <th scope="col"><a href="/wiki/Russians" title="Russians">Russians</a> </th> <th scope="col"><a href="/wiki/Irish_people" title="Irish people">Irish</a> </th> <th scope="col"><a href="/wiki/Greek_Americans" title="Greek Americans">Greek Americans</a> </th> <th scope="col"><a href="/wiki/Ashkenazi_Jews" title="Ashkenazi Jews">Ashkenazi Jews</a> </th> <th scope="col"><a href="/wiki/Circassians" title="Circassians">Circassians</a> </th></tr> <tr> <th scope="row">Italian Americans </th> <td> </td> <td>0.0064 </td> <td>0.0064 </td> <td>0.0057 </td> <td>0.0010 </td> <td>0.0029 </td> <td>0.0088 </td> <td>0.0048 </td> <td>0.0000 </td> <td>0.0040 </td> <td>0.0067 </td></tr> <tr> <th scope="row">Palestinians </th> <td>0.0064 </td> <td> </td> <td>0.0191 </td> <td>0.0064 </td> <td>0.0101 </td> <td>0.0136 </td> <td>0.0202 </td> <td>0.0170 </td> <td>0.0057 </td> <td>0.0093 </td> <td>0.0108 </td></tr> <tr> <th scope="row">Swedes </th> <td>0.0064 </td> <td>0.0191 </td> <td> </td> <td>0.0167 </td> <td>0.0040 </td> <td>0.0007 </td> <td>0.0030 </td> <td>0.0020 </td> <td>0.0084 </td> <td>0.0120 </td> <td>0.0117 </td></tr> <tr> <th scope="row">Druzes </th> <td>0.0057 </td> <td>0.0064 </td> <td>0.0167 </td> <td> </td> <td>0.0096 </td> <td>0.0121 </td> <td>0.0194 </td> <td>0.0154 </td> <td>0.0052 </td> <td>0.0088 </td> <td>0.0092 </td></tr> <tr> <th scope="row">Spaniards </th> <td>0.0010 </td> <td>0.0101 </td> <td>0.0040 </td> <td>0.0096 </td> <td> </td> <td>0.0015 </td> <td>0.0070 </td> <td>0.0037 </td> <td>0.0035 </td> <td>0.0056 </td> <td>0.0090 </td></tr> <tr> <th scope="row">Germans </th> <td>0.0029 </td> <td>0.0136 </td> <td>0.0007 </td> <td>0.0121 </td> <td>0.0015 </td> <td> </td> <td>0.0030 </td> <td>0.0010 </td> <td>0.0039 </td> <td>0.0072 </td> <td>0.0089 </td></tr> <tr> <th scope="row">Russians </th> <td>0.0088 </td> <td>0.0202 </td> <td>0.0030 </td> <td>0.0194 </td> <td>0.0070 </td> <td>0.0030 </td> <td> </td> <td>0.0038 </td> <td>0.0108 </td> <td>0.0137 </td> <td>0.0120 </td></tr> <tr> <th scope="row">Irish </th> <td>0.0048 </td> <td>0.0170 </td> <td>0.0020 </td> <td>0.0154 </td> <td>0.0037 </td> <td>0.0010 </td> <td>0.0038 </td> <td> </td> <td>0.0067 </td> <td>0.0109 </td> <td>0.0110 </td></tr> <tr> <th scope="row">Greek Americans </th> <td>0.0000 </td> <td>0.0057 </td> <td>0.0084 </td> <td>0.0052 </td> <td>0.0035 </td> <td>0.0039 </td> <td>0.0108 </td> <td>0.0067 </td> <td> </td> <td>0.0042 </td> <td>0.0054 </td></tr> <tr> <th scope="row">Ashkenazi Jews </th> <td>0.0040 </td> <td>0.0093 </td> <td>0.0120 </td> <td>0.0088 </td> <td>0.0056 </td> <td>0.0072 </td> <td>0.0137 </td> <td>0.0109 </td> <td>0.0042 </td> <td> </td> <td>0.0107 </td></tr> <tr> <th scope="row">Circassians </th> <td>0.0067 </td> <td>0.0108 </td> <td>0.0117 </td> <td>0.0092 </td> <td>0.0090 </td> <td>0.0089 </td> <td>0.0120 </td> <td>0.0110 </td> <td>0.0054 </td> <td>0.0107 </td> <td> </td></tr></tbody></table> <table class="wikitable" style="text-align:center; width:80%;"> <caption>European Population Genetic Substructure based on SNPs<sup id="cite_ref-nelis_103-0" class="reference"><a href="#cite_note-nelis-103"><span class="cite-bracket">[</span>103<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-FleischerNelis2009_1-1" class="reference"><a href="#cite_note-FleischerNelis2009-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </caption> <tbody><tr> <th scope="col"> </th> <th scope="col"><a href="/wiki/Austria" title="Austria">Austria</a> </th> <th scope="col"><a href="/wiki/Bulgaria" title="Bulgaria">Bulgaria</a> </th> <th scope="col"><a href="/wiki/Czech_Republic" title="Czech Republic">Czech Republic</a> </th> <th scope="col"><a href="/wiki/Estonia" title="Estonia">Estonia</a> </th> <th scope="col"><a href="/wiki/Finland" title="Finland">Finland</a> (<a href="/wiki/Helsinki" title="Helsinki">Helsinki</a>) </th> <th scope="col"><a href="/wiki/Finland" title="Finland">Finland</a> (<a href="/wiki/Kuusamo" title="Kuusamo">Kuusamo</a>) </th> <th scope="col"><a href="/wiki/France" title="France">France</a> </th> <th scope="col"><a href="/wiki/Northern_Germany" title="Northern Germany">Northern Germany</a> </th> <th scope="col"><a href="/wiki/Southern_Germany" title="Southern Germany">Southern Germany</a> </th> <th scope="col"><a href="/wiki/Hungary" title="Hungary">Hungary</a> </th> <th scope="col"><a href="/wiki/Northern_Italy" title="Northern Italy">Northern Italy</a> </th> <th scope="col"><a href="/wiki/Southern_Italy" title="Southern Italy">Southern Italy</a> </th> <th scope="col"><a href="/wiki/Latvia" title="Latvia">Latvia</a> </th> <th scope="col"><a href="/wiki/Lithuania" title="Lithuania">Lithuania</a> </th> <th scope="col"><a href="/wiki/Poland" title="Poland">Poland</a> </th> <th scope="col"><a href="/wiki/Russia" title="Russia">Russia</a> </th> <th scope="col"><a href="/wiki/Spain" title="Spain">Spain</a> </th> <th scope="col"><a href="/wiki/Sweden" title="Sweden">Sweden</a> </th> <th scope="col"><a href="/wiki/Switzerland" title="Switzerland">Switzerland</a> </th> <th scope="col">CEU </th></tr> <tr> <th scope="row">Austria </th> <td> </td> <td>1.14 </td> <td>1.08 </td> <td>1.58 </td> <td>2.24 </td> <td>3.30 </td> <td>1.16 </td> <td>1.10 </td> <td>1.04 </td> <td>1.04 </td> <td>1.49 </td> <td>1.79 </td> <td>1.85 </td> <td>1.70 </td> <td>1.19 </td> <td>1.47 </td> <td>1.41 </td> <td>1.21 </td> <td>1.19 </td> <td>1.12 </td> <td>Austria </td></tr> <tr> <th scope="row">Bulgaria </th> <td>1.14 </td> <td> </td> <td>1.21 </td> <td>1.70 </td> <td>2.19 </td> <td>2.91 </td> <td>1.22 </td> <td>1.32 </td> <td>1.19 </td> <td>1.10 </td> <td>1.32 </td> <td>1.38 </td> <td>1.86 </td> <td>1.73 </td> <td>1.29 </td> <td>1.53 </td> <td>1.30 </td> <td>1.47 </td> <td>1.13 </td> <td>1.29 </td> <td>Bulgaria </td></tr> <tr> <th scope="row">Czech Republic </th> <td>1.08 </td> <td>1.21 </td> <td> </td> <td>1.42 </td> <td>2.20 </td> <td>3.26 </td> <td>1.35 </td> <td>1.15 </td> <td>1.16 </td> <td>1.06 </td> <td>1.69 </td> <td>2.04 </td> <td>1.62 </td> <td>1.48 </td> <td>1.09 </td> <td>1.27 </td> <td>1.63 </td> <td>1.26 </td> <td>1.37 </td> <td>1.21 </td> <td>Czech Republic </td></tr> <tr> <th scope="row">Estonia </th> <td>1.58 </td> <td>1.70 </td> <td>1.42 </td> <td> </td> <td>1.71 </td> <td>2.80 </td> <td>2.08 </td> <td>1.53 </td> <td>1.70 </td> <td>1.41 </td> <td>2.42 </td> <td>2.93 </td> <td>1.24 </td> <td>1.28 </td> <td>1.17 </td> <td>1.21 </td> <td>2.54 </td> <td>1.49 </td> <td>2.16 </td> <td>1.59 </td> <td>Estonia </td></tr> <tr> <th scope="row">Finland (Helsinki) </th> <td>2.24 </td> <td>2.19 </td> <td>2.20 </td> <td>1.71 </td> <td> </td> <td>1.86 </td> <td>2.69 </td> <td>2.17 </td> <td>2.35 </td> <td>1.87 </td> <td>2.82 </td> <td>3.37 </td> <td>2.31 </td> <td>2.33 </td> <td>1.75 </td> <td>2.10 </td> <td>3.14 </td> <td>1.89 </td> <td>2.77 </td> <td>1.99 </td> <td>Finland (Helsinki) </td></tr> <tr> <th scope="row">Finland (Kuusamo) </th> <td>3.30 </td> <td>2.91 </td> <td>3.26 </td> <td>2.80 </td> <td>1.86 </td> <td> </td> <td>3.72 </td> <td>3.27 </td> <td>3.46 </td> <td>2.68 </td> <td>3.64 </td> <td>4.18 </td> <td>3.33 </td> <td>3.37 </td> <td>2.49 </td> <td>3.16 </td> <td>4.21 </td> <td>2.87 </td> <td>3.83 </td> <td>2.89 </td> <td>Finland (Kuusamo) </td></tr> <tr> <th scope="row">France </th> <td>1.16 </td> <td>1.22 </td> <td>1.35 </td> <td>2.08 </td> <td>2.69 </td> <td>3.72 </td> <td> </td> <td>1.25 </td> <td>1.12 </td> <td>1.16 </td> <td>1.38 </td> <td>1.68 </td> <td>2.40 </td> <td>2.20 </td> <td>1.44 </td> <td>1.94 </td> <td>1.13 </td> <td>1.38 </td> <td>1.10 </td> <td>1.13 </td> <td>France </td></tr> <tr> <th scope="row">Northern Germany </th> <td>1.10 </td> <td>1.32 </td> <td>1.15 </td> <td>1.53 </td> <td>2.17 </td> <td>3.27 </td> <td>1.25 </td> <td> </td> <td>1.08 </td> <td>1.11 </td> <td>1.72 </td> <td>2.14 </td> <td>1.84 </td> <td>1.66 </td> <td>1.18 </td> <td>1.49 </td> <td>1.62 </td> <td>1.12 </td> <td>1.36 </td> <td>1.06 </td> <td>Northern Germany </td></tr> <tr> <th scope="row">Southern Germany </th> <td>1.04 </td> <td>1.19 </td> <td>1.16 </td> <td>1.70 </td> <td>2.35 </td> <td>3.46 </td> <td>1.12 </td> <td>1.08 </td> <td> </td> <td>1.08 </td> <td>1.53 </td> <td>1.85 </td> <td>1.20 </td> <td>1.84 </td> <td>1.23 </td> <td>1.58 </td> <td>1.40 </td> <td>1.21 </td> <td>1.17 </td> <td>1.07 </td> <td>Southern Germany </td></tr> <tr> <th scope="row">Hungary </th> <td>1.04 </td> <td>1.10 </td> <td>1.06 </td> <td>1.41 </td> <td>1.87 </td> <td>2.68 </td> <td>1.16 </td> <td>1.11 </td> <td>1.08 </td> <td> </td> <td>1.42 </td> <td>1.63 </td> <td>1.58 </td> <td>1.46 </td> <td>1.14 </td> <td>1.28 </td> <td>1.32 </td> <td>1.22 </td> <td>1.16 </td> <td>1.13 </td> <td>Hungary </td></tr> <tr> <th scope="row">Northern Italy </th> <td>1.49 </td> <td>1.32 </td> <td>1.69 </td> <td>2.42 </td> <td>2.82 </td> <td>3.64 </td> <td>1.38 </td> <td>1.72 </td> <td>1.53 </td> <td>1.42 </td> <td> </td> <td>1.54 </td> <td>2.64 </td> <td>2.48 </td> <td>1.75 </td> <td>2.24 </td> <td>1.42 </td> <td>1.86 </td> <td>1.36 </td> <td>1.56 </td> <td>Northern Italy </td></tr> <tr> <th scope="row">Southern Italy </th> <td>1.79 </td> <td>1.38 </td> <td>2.04 </td> <td>2.93 </td> <td>3.37 </td> <td>4.18 </td> <td>1.68 </td> <td>2.14 </td> <td>1.85 </td> <td>1.63 </td> <td>1.54 </td> <td> </td> <td>3.14 </td> <td>2.96 </td> <td>1.99 </td> <td>2.68 </td> <td>1.67 </td> <td>2.28 </td> <td>1.54 </td> <td>1.84 </td> <td>Southern Italy </td></tr> <tr> <th scope="row">Latvia </th> <td>1.85 </td> <td>1.86 </td> <td>1.62 </td> <td>1.24 </td> <td>2.31 </td> <td>3.33 </td> <td>2.40 </td> <td>1.84 </td> <td>1.20 </td> <td>1.58 </td> <td>2.64 </td> <td>3.14 </td> <td> </td> <td>1.20 </td> <td>1.26 </td> <td>1.84 </td> <td>2.82 </td> <td>1.89 </td> <td>2.52 </td> <td>1.87 </td> <td>Latvia </td></tr> <tr> <th scope="row">Lithuania </th> <td>1.70 </td> <td>1.73 </td> <td>1.48 </td> <td>1.28 </td> <td>2.33 </td> <td>3.37 </td> <td>2.20 </td> <td>1.66 </td> <td>1.84 </td> <td>1.46 </td> <td>2.48 </td> <td>2.96 </td> <td>1.20 </td> <td> </td> <td>1.20 </td> <td>1.26 </td> <td>2.62 </td> <td>1.74 </td> <td>2.29 </td> <td>1.74 </td> <td>Lithuania </td></tr> <tr> <th scope="row">Poland </th> <td>1.19 </td> <td>1.29 </td> <td>1.09 </td> <td>1.17 </td> <td>1.75 </td> <td>2.49 </td> <td>1.44 </td> <td>1.18 </td> <td>1.23 </td> <td>1.14 </td> <td>1.75 </td> <td>1.99 </td> <td>1.26 </td> <td>1.20 </td> <td> </td> <td>1.18 </td> <td>1.66 </td> <td>1.30 </td> <td>1.46 </td> <td>1.28 </td> <td>Poland </td></tr> <tr> <th scope="row">Russia </th> <td>1.47 </td> <td>1.53 </td> <td>1.27 </td> <td>1.21 </td> <td>2.10 </td> <td>3.16 </td> <td>1.94 </td> <td>1.49 </td> <td>1.58 </td> <td>1.28 </td> <td>2.24 </td> <td>2.68 </td> <td>1.84 </td> <td>1.26 </td> <td>1.18 </td> <td> </td> <td>2.32 </td> <td>1.59 </td> <td>1.20 </td> <td>1.56 </td> <td>Russia </td></tr> <tr> <th scope="row">Spain </th> <td>1.41 </td> <td>1.30 </td> <td>1.63 </td> <td>2.54 </td> <td>3.14 </td> <td>4.21 </td> <td>1.13 </td> <td>1.62 </td> <td>1.40 </td> <td>1.32 </td> <td>1.42 </td> <td>1.67 </td> <td>2.82 </td> <td>2.62 </td> <td>1.66 </td> <td>2.32 </td> <td> </td> <td>1.73 </td> <td>1.16 </td> <td>1.34 </td> <td>Spain </td></tr> <tr> <th scope="row">Sweden </th> <td>1.21 </td> <td>1.47 </td> <td>1.26 </td> <td>1.49 </td> <td>1.89 </td> <td>2.87 </td> <td>1.38 </td> <td>1.12 </td> <td>1.21 </td> <td>1.22 </td> <td>1.86 </td> <td>2.28 </td> <td>1.89 </td> <td>1.74 </td> <td>1.30 </td> <td>1.59 </td> <td>1.73 </td> <td> </td> <td>1.50 </td> <td>1.09 </td> <td>Sweden </td></tr> <tr> <th scope="row">Switzerland </th> <td>1.19 </td> <td>1.13 </td> <td>1.37 </td> <td>2.16 </td> <td>2.77 </td> <td>3.83 </td> <td>1.10 </td> <td>1.36 </td> <td>1.17 </td> <td>1.16 </td> <td>1.36 </td> <td>1.54 </td> <td>2.52 </td> <td>2.29 </td> <td>1.46 </td> <td>1.20 </td> <td>1.16 </td> <td>1.50 </td> <td> </td> <td>1.21 </td> <td>Switzerland </td></tr> <tr> <th scope="row">CEU </th> <td>1.12 </td> <td>1.29 </td> <td>1.21 </td> <td>1.59 </td> <td>1.99 </td> <td>2.89 </td> <td>1.13 </td> <td>1.06 </td> <td>1.07 </td> <td>1.13 </td> <td>1.56 </td> <td>1.84 </td> <td>1.87 </td> <td>1.74 </td> <td>1.28 </td> <td>1.56 </td> <td>1.34 </td> <td>1.09 </td> <td>1.21 </td> <td> </td> <td>CEU </td></tr> <tr> <th scope="col"> </th> <th scope="col">Austria </th> <th scope="col">Bulgaria </th> <th scope="col">Czech Republic </th> <th scope="col">Estonia </th> <th scope="col">Finland (Helsinki) </th> <th scope="col">Finland (Kuusamo) </th> <th scope="col">France </th> <th scope="col">Northern Germany </th> <th scope="col">Southern Germany </th> <th scope="col">Hungary </th> <th scope="col">Northern Italy </th> <th scope="col">Southern Italy </th> <th scope="col">Latvia </th> <th scope="col">Lithuania </th> <th scope="col">Poland </th> <th scope="col">Russia </th> <th scope="col">Spain </th> <th scope="col">Sweden </th> <th scope="col">Switzerland </th> <th scope="col">CEU </th></tr> </tbody></table> <p>CEU – Utah residents with ancestry from Northern and Western Europe. </p> <div class="mw-heading mw-heading2"><h2 id="History_of_research">History of research</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Genetic_history_of_Europe&action=edit&section=15" title="Edit section: History of research"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Population_genetics" title="Population genetics">Population genetics</a></div> <div class="mw-heading mw-heading3"><h3 id="Classical_genetic_markers_(by_proxy)"><span id="Classical_genetic_markers_.28by_proxy.29"></span>Classical genetic markers (by proxy)</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Genetic_history_of_Europe&action=edit&section=16" title="Edit section: Classical genetic markers (by proxy)"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>One of the first scholars to perform genetic studies was <a href="/wiki/Luigi_Luca_Cavalli-Sforza" title="Luigi Luca Cavalli-Sforza">Luigi Luca Cavalli-Sforza</a>. He used classical genetic markers to analyse DNA by proxy. This method studies differences in the frequencies of particular allelic traits, namely <a href="/wiki/Polymorphism_(biology)" title="Polymorphism (biology)">polymorphisms</a> from proteins found within <a href="/wiki/Blood" title="Blood">human blood</a> (such as the <a href="/wiki/ABO_blood_group_system" title="ABO blood group system">ABO blood groups</a>, Rhesus blood antigens, <a href="/wiki/Major_histocompatibility_complex" title="Major histocompatibility complex">HLA loci</a>, <a href="/wiki/Antibody" title="Antibody">immunoglobulins</a>, <a href="/wiki/G6PD" class="mw-redirect" title="G6PD">G6PD</a> <a href="/wiki/Isoenzyme" class="mw-redirect" title="Isoenzyme">isoenzymes</a>, among others). Subsequently, his team calculated <a href="/wiki/Genetic_distance" title="Genetic distance">genetic distance</a> between populations, based on the principle that two populations that share similar frequencies of a trait are more closely related than populations that have more divergent frequencies of the trait.<sup id="cite_ref-Cavalli-Sforza_1993_80-3" class="reference"><a href="#cite_note-Cavalli-Sforza_1993-80"><span class="cite-bracket">[</span>80<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 51">: 51 </span></sup> </p><p>From this, he constructed <a href="/wiki/Phylogenetic_tree" title="Phylogenetic tree">phylogenetic trees</a> that showed genetic distances diagrammatically. His team also performed <a href="/wiki/Principal_component_analysis" title="Principal component analysis">principal component analyses</a>, which is good at analysing <a href="/wiki/Multivariate_statistics" title="Multivariate statistics">multivariate data</a> with minimal loss of information. The information that is lost can be partly restored by generating a second principal component, and so on.<sup id="cite_ref-Cavalli-Sforza_1993_80-4" class="reference"><a href="#cite_note-Cavalli-Sforza_1993-80"><span class="cite-bracket">[</span>80<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 39">: 39 </span></sup> In turn, the information from each individual principal component (<b>PC</b>) can be presented graphically in <i>synthetic maps</i>. These maps show peaks and troughs, which represent populations whose <a href="/wiki/Allele_frequency" title="Allele frequency">gene frequencies</a> take extreme values compared to others in the studied area.<sup id="cite_ref-Cavalli-Sforza_1993_80-5" class="reference"><a href="#cite_note-Cavalli-Sforza_1993-80"><span class="cite-bracket">[</span>80<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 51">: 51 </span></sup> </p><p>Peaks and troughs usually connected by smooth gradients are called <a href="/wiki/Cline_(biology)" title="Cline (biology)">clines</a>. Genetic clines can be generated by adaptation to environment (<a href="/wiki/Natural_selection" title="Natural selection">natural selection</a>), continuous <a href="/wiki/Gene_flow" title="Gene flow">gene flow</a> between two initially different populations or a demographic expansion into a scarcely populated environment, with little initial <a href="/wiki/Genetic_admixture" title="Genetic admixture">admixture</a> with existing populations.<sup id="cite_ref-Arredi_2007_104-0" class="reference"><a href="#cite_note-Arredi_2007-104"><span class="cite-bracket">[</span>104<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 390">: 390 </span></sup> Cavalli-Sforza connected these gradients with postulated pre-historical population movements, based on archaeological and linguistic theories. However, given that the time depths of such patterns are not known, "associating them with particular demographic events is usually speculative".<sup id="cite_ref-Rosser_2000_44-2" class="reference"><a href="#cite_note-Rosser_2000-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Direct_DNA_analysis">Direct DNA analysis</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Genetic_history_of_Europe&action=edit&section=17" title="Edit section: Direct DNA analysis"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Genetic_drift" title="Genetic drift">Genetic drift</a>, <a href="/wiki/Founder_effect" title="Founder effect">Founder effect</a>, and <a href="/wiki/Population_bottleneck" title="Population bottleneck">Population bottleneck</a></div> <p>Studies using direct DNA analysis are now abundant and may use <a href="/wiki/Mitochondrial_DNA" title="Mitochondrial DNA">mitochondrial DNA (mtDNA)</a>, the non-recombining portion of the Y chromosome (NRY), or even autosomal DNA. MtDNA and NRY DNA share some similar features, which have made them particularly useful in genetic anthropology. These properties include the direct, unaltered inheritance of mtDNA and NRY DNA from mother to offspring and father to son, respectively, without the 'scrambling' effects of <a href="/wiki/Genetic_recombination" title="Genetic recombination">genetic recombination</a>. We also presume that these genetic loci are not affected by natural selection and that the major process responsible for changes in <a href="/wiki/Base_pair" title="Base pair">base pairs</a> has been mutation (which can be calculated).<sup id="cite_ref-Milisauskas_2002_23-3" class="reference"><a href="#cite_note-Milisauskas_2002-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 58">: 58 </span></sup> </p><p>The smaller <a href="/wiki/Effective_population_size" title="Effective population size">effective population size</a> of the NRY and mtDNA enhances the consequences of drift and founder effect, relative to the autosomes, making NRY and mtDNA variation a potentially sensitive index of population composition.<sup id="cite_ref-Rosser_2000_44-3" class="reference"><a href="#cite_note-Rosser_2000-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Richards_2000_28-3" class="reference"><a href="#cite_note-Richards_2000-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Semino_2000_25-8" class="reference"><a href="#cite_note-Semino_2000-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> These biologically plausible assumptions are not concrete; Rosser suggests that climatic conditions may affect the fertility of certain lineages.<sup id="cite_ref-Rosser_2000_44-4" class="reference"><a href="#cite_note-Rosser_2000-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> </p><p>The underlying <a href="/wiki/Mutation_rate" title="Mutation rate">mutation rate</a> used by the geneticists is more questionable. They often use different mutation rates and studies frequently arrive at vastly different conclusions.<sup id="cite_ref-Rosser_2000_44-5" class="reference"><a href="#cite_note-Rosser_2000-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> NRY and mtDNA may be so susceptible to drift that some ancient patterns may have become obscured. Another assumption is that population genealogies are approximated by <a href="/wiki/Allele_frequency" title="Allele frequency">allele genealogies</a>. <a href="/wiki/Guido_Barbujani" title="Guido Barbujani">Guido Barbujani</a> points out that this only holds if population groups develop from a genetically monomorphic set of founders. Barbujani argues that there is no reason to believe that Europe was colonised by monomorphic populations. This would result in an overestimation of haplogroup age, thus falsely extending the demographic history of Europe into the <a href="/wiki/Paleolithic" title="Paleolithic">Late Paleolithic</a> rather than the <a href="/wiki/Neolithic" title="Neolithic">Neolithic</a> era.<sup id="cite_ref-Barbujani_2001_105-0" class="reference"><a href="#cite_note-Barbujani_2001-105"><span class="cite-bracket">[</span>105<span class="cite-bracket">]</span></a></sup> Greater certainty about chronology may be obtained from studies of ancient DNA (see below), but so far these have been comparatively few. </p><p>Whereas <a href="/wiki/Human_Y-chromosome_DNA_haplogroup" title="Human Y-chromosome DNA haplogroup">Y-DNA</a> and mtDNA haplogroups represent but a small component of a person's DNA pool, <a href="/wiki/Autosome" title="Autosome">autosomal DNA</a> has the advantage of containing hundreds of thousands of examinable genetic loci, thus giving a more complete picture of genetic composition. Descent relationships can only be determined on a statistical basis, because autosomal DNA undergoes recombination. A single chromosome can record a history for each gene. Autosomal studies are much more reliable for showing the relationships between existing populations, but do not offer the possibilities for unravelling their histories in the same way as mtDNA and NRY DNA studies promise, despite their many complications. </p><p>Genetic studies operate on numerous assumptions and suffer from methodological limitations, such as <a href="/wiki/Selection_bias" title="Selection bias">selection bias</a> and confounding phenomena like <a href="/wiki/Genetic_drift" title="Genetic drift">genetic drift</a>, foundation and bottleneck effects cause large errors, particularly in haplogroup studies. No matter how accurate the methodology, conclusions derived from such studies are compiled on the basis of how the author envisages their data fits with established archaeological or linguistic theories.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (February 2021)">citation needed</span></a></i>]</sup> </p> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Genetic_history_of_Europe&action=edit&section=18" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1259569809">.mw-parser-output .portalbox{padding:0;margin:0.5em 0;display:table;box-sizing:border-box;max-width:175px;list-style:none}.mw-parser-output .portalborder{border:1px solid var(--border-color-base,#a2a9b1);padding:0.1em;background:var(--background-color-neutral-subtle,#f8f9fa)}.mw-parser-output .portalbox-entry{display:table-row;font-size:85%;line-height:110%;height:1.9em;font-style:italic;font-weight:bold}.mw-parser-output .portalbox-image{display:table-cell;padding:0.2em;vertical-align:middle;text-align:center}.mw-parser-output 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.multicol-float-clear{clear:both}</style><div class="multicol-float" style=""> <dl><dt>General</dt></dl> <ul><li><a href="/wiki/Archaeogenetics" title="Archaeogenetics">Archaeogenetics</a></li> <li><a href="/wiki/Ethnic_groups_in_Europe" title="Ethnic groups in Europe">Ethnic groups in Europe</a></li> <li><a href="/wiki/European_studies" title="European studies">European studies</a></li> <li><a href="/wiki/Genetic_history_of_North_Africa" title="Genetic history of North Africa">Genetic history of North Africa</a></li> <li><a href="/wiki/Genetic_history_of_the_Middle_East" title="Genetic history of the Middle East">Genetic history of the Middle East</a></li> <li><a href="/wiki/Genetics_and_archaeogenetics_of_South_Asia" title="Genetics and archaeogenetics of South Asia">Genetics and archaeogenetics of South Asia</a></li> <li><a href="/wiki/History_of_Europe" title="History of Europe">History of Europe</a></li> <li><a href="/wiki/Human_genetic_variation" title="Human genetic variation">Human genetic variation</a></li> <li><a href="/wiki/Y-DNA_haplogroups_in_populations_of_Europe" title="Y-DNA haplogroups in populations of Europe">Y-DNA haplogroups in populations of Europe</a></li> <li><a href="/wiki/Romani_people#Genetic_evidence" title="Romani people">Romani people#Genetic evidence</a></li></ul> </div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1215345927"><div class="multicol-float" style=""> <dl><dt>Genetics by European group</dt></dl> <ul><li><a href="/wiki/Genetic_history_of_Italy" title="Genetic history of Italy">Genetic history of Italy</a></li> <li><a href="/wiki/Genetic_history_of_the_British_Isles" title="Genetic history of the British Isles">Genetic history of the British Isles</a></li> <li><a href="/wiki/Genetic_history_of_the_Iberian_Peninsula" title="Genetic history of the Iberian Peninsula">Genetic history of the Iberian Peninsula</a></li> <li><a href="/wiki/Genetic_studies_on_Bosniaks" title="Genetic studies on Bosniaks">Genetic studies on Bosniaks</a></li> <li><a href="/wiki/Genetic_studies_on_Bulgarians" title="Genetic studies on Bulgarians">Genetic studies on Bulgarians</a></li> <li><a href="/wiki/Genetic_studies_on_Croats" title="Genetic studies on Croats">Genetic studies on Croats</a></li> <li><a href="/wiki/Genetic_studies_on_Jews" class="mw-redirect" title="Genetic studies on Jews">Genetic studies on Jews</a></li> <li><a href="/wiki/Genetic_studies_on_Russians" title="Genetic studies on Russians">Genetic studies on Russians</a></li> <li><a href="/wiki/Genetic_studies_on_Sami" title="Genetic studies on Sami">Genetic studies on Sami</a></li> <li><a href="/wiki/Genetic_studies_on_Serbs" title="Genetic studies on Serbs">Genetic studies on Serbs</a></li> <li><a href="/wiki/Genetic_studies_on_Turkish_people" title="Genetic studies on Turkish people">Genetic studies on Turkish people</a></li></ul> </div><div class="multicol-float-clear" style=""></div> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Genetic_history_of_Europe&action=edit&section=19" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Inline_citations">Inline citations</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Genetic_history_of_Europe&action=edit&section=20" title="Edit section: Inline citations"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist"> <div class="mw-references-wrap mw-references-columns"><ol class="references"> <li id="cite_note-FleischerNelis2009-1"><span class="mw-cite-backlink">^ <a href="#cite_ref-FleischerNelis2009_1-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-FleischerNelis2009_1-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFNelisEskoMägiZimprich2009" class="citation journal cs1">Nelis M, Esko T, Mägi R, Zimprich F, Zimprich A, Toncheva D, et al. 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(identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/29466337">29466337</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nature&rft.atitle=The+Beaker+phenomenon+and+the+genomic+transformation+of+northwest+Europe&rft.volume=555&rft.issue=7695&rft.pages=190-196&rft.date=2018-03&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC5973796%23id-name%3DPMC&rft_id=info%3Apmid%2F29466337&rft_id=info%3Adoi%2F10.1038%2Fnature25738&rft_id=info%3Abibcode%2F2018Natur.555..190O&rft.aulast=Olalde&rft.aufirst=I&rft.au=Brace%2C+S&rft.au=Allentoft%2C+ME&rft.au=Armit%2C+I&rft.au=Kristiansen%2C+K&rft.au=Booth%2C+T&rft.au=Rohland%2C+N&rft.au=Mallick%2C+S&rft.au=Sz%C3%A9cs%C3%A9nyi-Nagy%2C+A&rft.au=Mittnik%2C+A&rft.au=Altena%2C+E&rft.au=Lipson%2C+M&rft.au=Lazaridis%2C+I&rft.au=Harper%2C+TK&rft.au=Patterson%2C+N&rft.au=Broomandkhoshbacht%2C+N&rft.au=Diekmann%2C+Y&rft.au=Faltyskova%2C+Z&rft.au=Fernandes%2C+D&rft.au=Ferry%2C+M&rft.au=Harney%2C+E&rft.au=de+Knijff%2C+P&rft.au=Michel%2C+M&rft.au=Oppenheimer%2C+J&rft.au=Stewardson%2C+K&rft.au=Barclay%2C+A&rft.au=Alt%2C+KW&rft.au=Liesau%2C+C&rft.au=R%C3%ADos%2C+P&rft.au=Blasco%2C+C&rft.au=Miguel%2C+JV&rft.au=Garc%C3%ADa%2C+RM&rft.au=Fern%C3%A1ndez%2C+AA&rft.au=B%C3%A1nffy%2C+E&rft.au=Bernab%C3%B2-Brea%2C+M&rft.au=Billoin%2C+D&rft.au=Bonsall%2C+C&rft.au=Bonsall%2C+L&rft.au=Allen%2C+T&rft.au=B%C3%BCster%2C+L&rft.au=Carver%2C+S&rft.au=Navarro%2C+LC&rft.au=Craig%2C+OE&rft.au=Cook%2C+GT&rft.au=Cunliffe%2C+B&rft.au=Denaire%2C+A&rft.au=Dinwiddy%2C+KE&rft.au=Dodwell%2C+N&rft.au=Ern%C3%A9e%2C+M&rft.au=Evans%2C+C&rft.au=Kucha%C5%99%C3%ADk%2C+M&rft.au=Farr%C3%A9%2C+JF&rft.au=Fowler%2C+C&rft.au=Gazenbeek%2C+M&rft.au=Pena%2C+RG&rft.au=Haber-Uriarte%2C+M&rft.au=Haduch%2C+E&rft.au=Hey%2C+G&rft.au=Jowett%2C+N&rft.au=Knowles%2C+T&rft.au=Massy%2C+K&rft.au=Pfrengle%2C+S&rft.au=Lefranc%2C+P&rft.au=Lemercier%2C+O&rft.au=Lefebvre%2C+A&rft.au=Mart%C3%ADnez%2C+CH&rft.au=Olmo%2C+VG&rft.au=Ram%C3%ADrez%2C+AB&rft.au=Maurandi%2C+JL&rft.au=Maj%C3%B3%2C+T&rft.au=McKinley%2C+JI&rft.au=McSweeney%2C+K&rft.au=Mende%2C+BG&rft.au=Modi%2C+A&rft.au=Kulcs%C3%A1r%2C+G&rft.au=Kiss%2C+V&rft.au=Czene%2C+A&rft.au=Patay%2C+R&rft.au=Endr%C5%91di%2C+A&rft.au=K%C3%B6hler%2C+K&rft.au=Hajdu%2C+T&rft.au=Szeniczey%2C+T&rft.au=Dani%2C+J&rft.au=Bernert%2C+Z&rft.au=Hoole%2C+M&rft.au=Cheronet%2C+O&rft.au=Keating%2C+D&rft.au=Velem%C3%ADnsk%C3%BD%2C+P&rft.au=Dobe%C5%A1%2C+M&rft.au=Candilio%2C+F&rft.au=Brown%2C+F&rft.au=Fern%C3%A1ndez%2C+RF&rft.au=Herrero-Corral%2C+AM&rft.au=Tusa%2C+S&rft.au=Carnieri%2C+E&rft.au=Lentini%2C+L&rft.au=Valenti%2C+A&rft.au=Zanini%2C+A&rft.au=Waddington%2C+C&rft.au=Delibes%2C+G&rft.au=Guerra-Doce%2C+E&rft.au=Neil%2C+B&rft.au=Brittain%2C+M&rft.au=Luke%2C+M&rft.au=Mortimer%2C+R&rft.au=Desideri%2C+J&rft.au=Besse%2C+M&rft.au=Br%C3%BCcken%2C+G&rft.au=Furmanek%2C+M&rft.au=Ha%C5%82uszko%2C+A&rft.au=Mackiewicz%2C+M&rft.au=Rapi%C5%84ski%2C+A&rft.au=Leach%2C+S&rft.au=Soriano%2C+I&rft.au=Lillios%2C+KT&rft.au=Cardoso%2C+JL&rft.au=Pearson%2C+MP&rft.au=W%C5%82odarczak%2C+P&rft.au=Price%2C+TD&rft.au=Prieto%2C+P&rft.au=Rey%2C+PJ&rft.au=Risch%2C+R&rft.au=Rojo+Guerra%2C+MA&rft.au=Schmitt%2C+A&rft.au=Serralongue%2C+J&rft.au=Silva%2C+AM&rft.au=Smr%C4%8Dka%2C+V&rft.au=Vergnaud%2C+L&rft.au=Zilh%C3%A3o%2C+J&rft.au=Caramelli%2C+D&rft.au=Higham%2C+T&rft.au=Thomas%2C+MG&rft.au=Kennett%2C+DJ&rft.au=Fokkens%2C+H&rft.au=Heyd%2C+V&rft.au=Sheridan%2C+A&rft.au=Sj%C3%B6gren%2C+KG&rft.au=Stockhammer%2C+PW&rft.au=Krause%2C+J&rft.au=Pinhasi%2C+R&rft.au=Haak%2C+W&rft.au=Barnes%2C+I&rft.au=Lalueza-Fox%2C+C&rft.au=Reich%2C+D&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC5973796&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGenetic+history+of+Europe" class="Z3988"></span></span> </li> <li id="cite_note-6"><span class="mw-cite-backlink"><b><a href="#cite_ref-6">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHaakLazaridisPattersonRohland2015" class="citation journal cs1">Haak, Wolfgang; Lazaridis, Iosif; Patterson, Nick; Rohland, Nadin; Mallick, Swapan; Llamas, Bastien; Brandt, Guido; Nordenfelt, Susanne; Harney, Eadaoin; Stewardson, Kristin; Fu, Qiaomei; Mittnik, Alissa; Bánffy, Eszter; Economou, Christos; Francken, Michael (2015). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5048219">"Massive migration from the steppe was a source for Indo-European languages in Europe"</a>. <i>Nature</i>. <b>522</b> (7555): 207–211. <a href="/wiki/ArXiv_(identifier)" class="mw-redirect" title="ArXiv (identifier)">arXiv</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://arxiv.org/abs/1502.02783">1502.02783</a></span>. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2015Natur.522..207H">2015Natur.522..207H</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2Fnature14317">10.1038/nature14317</a>. <a href="/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/1476-4687">1476-4687</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5048219">5048219</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/25731166">25731166</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nature&rft.atitle=Massive+migration+from+the+steppe+was+a+source+for+Indo-European+languages+in+Europe&rft.volume=522&rft.issue=7555&rft.pages=207-211&rft.date=2015&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC5048219%23id-name%3DPMC&rft_id=info%3Abibcode%2F2015Natur.522..207H&rft_id=info%3Aarxiv%2F1502.02783&rft.issn=1476-4687&rft_id=info%3Adoi%2F10.1038%2Fnature14317&rft_id=info%3Apmid%2F25731166&rft.aulast=Haak&rft.aufirst=Wolfgang&rft.au=Lazaridis%2C+Iosif&rft.au=Patterson%2C+Nick&rft.au=Rohland%2C+Nadin&rft.au=Mallick%2C+Swapan&rft.au=Llamas%2C+Bastien&rft.au=Brandt%2C+Guido&rft.au=Nordenfelt%2C+Susanne&rft.au=Harney%2C+Eadaoin&rft.au=Stewardson%2C+Kristin&rft.au=Fu%2C+Qiaomei&rft.au=Mittnik%2C+Alissa&rft.au=B%C3%A1nffy%2C+Eszter&rft.au=Economou%2C+Christos&rft.au=Francken%2C+Michael&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC5048219&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGenetic+history+of+Europe" class="Z3988"></span></span> </li> <li id="cite_note-Lazaridis2014-7"><span class="mw-cite-backlink"><b><a href="#cite_ref-Lazaridis2014_7-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLazaridisPattersonMittnikRenaud2014" class="citation journal cs1">Lazaridis I, Patterson N, Mittnik A, Renaud G, Mallick S, Kirsanow K, et al. (September 2014). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4170574">"Ancient human genomes suggest three ancestral populations for present-day Europeans"</a>. <i>Nature</i>. <b>513</b> (7518): 409–413. <a href="/wiki/ArXiv_(identifier)" class="mw-redirect" title="ArXiv (identifier)">arXiv</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://arxiv.org/abs/1312.6639">1312.6639</a></span>. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2014Natur.513..409L">2014Natur.513..409L</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2Fnature13673">10.1038/nature13673</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4170574">4170574</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/25230663">25230663</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nature&rft.atitle=Ancient+human+genomes+suggest+three+ancestral+populations+for+present-day+Europeans&rft.volume=513&rft.issue=7518&rft.pages=409-413&rft.date=2014-09&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4170574%23id-name%3DPMC&rft_id=info%3Abibcode%2F2014Natur.513..409L&rft_id=info%3Aarxiv%2F1312.6639&rft_id=info%3Apmid%2F25230663&rft_id=info%3Adoi%2F10.1038%2Fnature13673&rft.aulast=Lazaridis&rft.aufirst=I&rft.au=Patterson%2C+N&rft.au=Mittnik%2C+A&rft.au=Renaud%2C+G&rft.au=Mallick%2C+S&rft.au=Kirsanow%2C+K&rft.au=Sudmant%2C+PH&rft.au=Schraiber%2C+JG&rft.au=Castellano%2C+S&rft.au=Lipson%2C+M&rft.au=Berger%2C+B&rft.au=Economou%2C+C&rft.au=Bollongino%2C+R&rft.au=Fu%2C+Q&rft.au=Bos%2C+KI&rft.au=Nordenfelt%2C+S&rft.au=Li%2C+H&rft.au=de+Filippo%2C+C&rft.au=Pr%C3%BCfer%2C+K&rft.au=Sawyer%2C+S&rft.au=Posth%2C+C&rft.au=Haak%2C+W&rft.au=Hallgren%2C+F&rft.au=Fornander%2C+E&rft.au=Rohland%2C+N&rft.au=Delsate%2C+D&rft.au=Francken%2C+M&rft.au=Guinet%2C+JM&rft.au=Wahl%2C+J&rft.au=Ayodo%2C+G&rft.au=Babiker%2C+HA&rft.au=Bailliet%2C+G&rft.au=Balanovska%2C+E&rft.au=Balanovsky%2C+O&rft.au=Barrantes%2C+R&rft.au=Bedoya%2C+G&rft.au=Ben-Ami%2C+H&rft.au=Bene%2C+J&rft.au=Berrada%2C+F&rft.au=Bravi%2C+CM&rft.au=Brisighelli%2C+F&rft.au=Busby%2C+GB&rft.au=Cali%2C+F&rft.au=Churnosov%2C+M&rft.au=Cole%2C+DE&rft.au=Corach%2C+D&rft.au=Damba%2C+L&rft.au=van+Driem%2C+G&rft.au=Dryomov%2C+S&rft.au=Dugoujon%2C+JM&rft.au=Fedorova%2C+SA&rft.au=Gallego+Romero%2C+I&rft.au=Gubina%2C+M&rft.au=Hammer%2C+M&rft.au=Henn%2C+BM&rft.au=Hervig%2C+T&rft.au=Hodoglugil%2C+U&rft.au=Jha%2C+AR&rft.au=Karachanak-Yankova%2C+S&rft.au=Khusainova%2C+R&rft.au=Khusnutdinova%2C+E&rft.au=Kittles%2C+R&rft.au=Kivisild%2C+T&rft.au=Klitz%2C+W&rft.au=Ku%C4%8Dinskas%2C+V&rft.au=Kushniarevich%2C+A&rft.au=Laredj%2C+L&rft.au=Litvinov%2C+S&rft.au=Loukidis%2C+T&rft.au=Mahley%2C+RW&rft.au=Melegh%2C+B&rft.au=Metspalu%2C+E&rft.au=Molina%2C+J&rft.au=Mountain%2C+J&rft.au=N%C3%A4kk%C3%A4l%C3%A4j%C3%A4rvi%2C+K&rft.au=Nesheva%2C+D&rft.au=Nyambo%2C+T&rft.au=Osipova%2C+L&rft.au=Parik%2C+J&rft.au=Platonov%2C+F&rft.au=Posukh%2C+O&rft.au=Romano%2C+V&rft.au=Rothhammer%2C+F&rft.au=Rudan%2C+I&rft.au=Ruizbakiev%2C+R&rft.au=Sahakyan%2C+H&rft.au=Sajantila%2C+A&rft.au=Salas%2C+A&rft.au=Starikovskaya%2C+EB&rft.au=Tarekegn%2C+A&rft.au=Toncheva%2C+D&rft.au=Turdikulova%2C+S&rft.au=Uktveryte%2C+I&rft.au=Utevska%2C+O&rft.au=Vasquez%2C+R&rft.au=Villena%2C+M&rft.au=Voevoda%2C+M&rft.au=Winkler%2C+CA&rft.au=Yepiskoposyan%2C+L&rft.au=Zalloua%2C+P&rft.au=Zemunik%2C+T&rft.au=Cooper%2C+A&rft.au=Capelli%2C+C&rft.au=Thomas%2C+MG&rft.au=Ruiz-Linares%2C+A&rft.au=Tishkoff%2C+SA&rft.au=Singh%2C+L&rft.au=Thangaraj%2C+K&rft.au=Villems%2C+R&rft.au=Comas%2C+D&rft.au=Sukernik%2C+R&rft.au=Metspalu%2C+M&rft.au=Meyer%2C+M&rft.au=Eichler%2C+EE&rft.au=Burger%2C+J&rft.au=Slatkin%2C+M&rft.au=P%C3%A4%C3%A4bo%2C+S&rft.au=Kelso%2C+J&rft.au=Reich%2C+D&rft.au=Krause%2C+J&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4170574&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGenetic+history+of+Europe" class="Z3988"></span></span> </li> <li id="cite_note-8"><span class="mw-cite-backlink"><b><a href="#cite_ref-8">^</a></b></span> <span class="reference-text">Since Lazaridis et al. (2014), further studies have refined the picture of interbreeding between EEF and WHG. In a 2017 analysis of 180 ancient DNA datasets of the Chalcolithic and Neolithic periods from Hungary, Germany and Spain evidence was found of a prolonged period of EEF-WHG interbreeding. Admixture took place regionally, from local hunter-gatherer populations, so that populations from the three regions (Germany, Iberia and Hungary) were genetically distinguishable at all stages of the Neolithic period, with a gradually increasing ratio of WHG ancestry of farming populations over time. This suggests that after the initial expansion of early farmers, there were no further long-range migrations substantial enough to homogenize the farming population, and that farming and hunter-gatherer populations existed side by side for many centuries, with ongoing gradual admixture throughout the 5th to 4th millennia <a href="/wiki/Common_Era" title="Common Era">BCE</a> (rather than a single admixture event on initial contact). <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLipsonSzécsényi-NagyMallickPósa2017" class="citation journal cs1">Lipson M, Szécsényi-Nagy A, Mallick S, Pósa A, Stégmár B, Keerl V, et al. 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Archived from <a rel="nofollow" class="external text" href="https://www.nationalgeographic.com/culture/2019/07/first-europeans-immigrants-genetic-testing-feature/">the original</a> on July 9, 2019.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=National+Geographic&rft.atitle=Genetic+testing+reveals+that+Europe+is+a+melting+pot%2C+made+of+immigrants&rft.date=2019&rft.aulast=Curry&rft.aufirst=A&rft_id=https%3A%2F%2Fwww.nationalgeographic.com%2Fculture%2F2019%2F07%2Ffirst-europeans-immigrants-genetic-testing-feature%2F&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGenetic+history+of+Europe" class="Z3988"></span></span> </li> <li id="cite_note-11"><span class="mw-cite-backlink"><b><a href="#cite_ref-11">^</a></b></span> <span class="reference-text">Lipson et al. 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These results were deemed 'ambiguous'. Archaeological evidence points to an abrupt change from Neanderthal artefacts to those related to AMH during the Upper Palaeolithic.<link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKlein2003" class="citation journal cs1">Klein RG (March 2003). "Paleoanthropology. 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(September 2006). 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(September 2008). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2684794">"Evolutionary toggling of the MAPT 17q21.31 inversion region"</a>. <i>Nature Genetics</i>. <b>40</b> (9): 1076–1083. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2Fng.193">10.1038/ng.193</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2684794">2684794</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/19165922">19165922</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nature+Genetics&rft.atitle=Evolutionary+toggling+of+the+MAPT+17q21.31+inversion+region&rft.volume=40&rft.issue=9&rft.pages=1076-1083&rft.date=2008-09&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2684794%23id-name%3DPMC&rft_id=info%3Apmid%2F19165922&rft_id=info%3Adoi%2F10.1038%2Fng.193&rft.aulast=Zody&rft.aufirst=MC&rft.au=Jiang%2C+Z&rft.au=Fung%2C+HC&rft.au=Antonacci%2C+F&rft.au=Hillier%2C+LW&rft.au=Cardone%2C+MF&rft.au=Graves%2C+TA&rft.au=Kidd%2C+JM&rft.au=Cheng%2C+Z&rft.au=Abouelleil%2C+A&rft.au=Chen%2C+L&rft.au=Wallis%2C+J&rft.au=Glasscock%2C+J&rft.au=Wilson%2C+RK&rft.au=Reily%2C+AD&rft.au=Duckworth%2C+J&rft.au=Ventura%2C+M&rft.au=Hardy%2C+J&rft.au=Warren%2C+WC&rft.au=Eichler%2C+EE&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2684794&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGenetic+history+of+Europe" class="Z3988"></span></span> </li> <li id="cite_note-19"><span class="mw-cite-backlink"><b><a href="#cite_ref-19">^</a></b></span> <span class="reference-text"><a href="/wiki/Introgression" title="Introgression">Introgression</a> and microcephalin FAQ <a href="/wiki/John_D._Hawks" title="John D. Hawks">John Hawks</a> <a rel="nofollow" class="external autonumber" href="http://johnhawks.net/weblog/reviews/neandertals/neandertal_dna/introgression_faq_2006.html">[1]</a></span> </li> <li id="cite_note-20"><span class="mw-cite-backlink"><b><a href="#cite_ref-20">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFAlmosHorváthCzibulaRaskó2008" class="citation journal cs1">Almos PZ, Horváth S, Czibula A, Raskó I, Sipos B, Bihari P, et al. (November 2008). <a rel="nofollow" class="external text" href="https://doi.org/10.1038%2Fhdy.2008.70">"H1 tau haplotype-related genomic variation at 17q21.3 as an Asian heritage of the European Gypsy population"</a>. <i>Heredity</i>. <b>101</b> (5): 416–419. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1038%2Fhdy.2008.70">10.1038/hdy.2008.70</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/18648385">18648385</a>. <q>In this study, we examine the frequency of a 900 kb inversion at 17q21.3 in the Gypsy and Caucasian populations of Hungary, which may reflect the Asian origin of Gypsy populations. Of the two haplotypes (H1 and H2), H2 is thought to be exclusively of Caucasian origin, and its occurrence in other racial groups is likely to reflect admixture. In our sample, the H1 haplotype was significantly more frequent in the Gypsy population (89.8 vs 75.5%, P<0.001) and was in Hardy–Weinberg disequilibrium (P=0.017). 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(November 2000). "The genetic legacy of Paleolithic Homo sapiens sapiens in extant Europeans: a Y chromosome perspective". <i>Science</i>. <b>290</b> (5494): 1155–1159. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2000Sci...290.1155S">2000Sci...290.1155S</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1126%2Fscience.290.5494.1155">10.1126/science.290.5494.1155</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/11073453">11073453</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Science&rft.atitle=The+genetic+legacy+of+Paleolithic+Homo+sapiens+sapiens+in+extant+Europeans%3A+a+Y+chromosome+perspective&rft.volume=290&rft.issue=5494&rft.pages=1155-1159&rft.date=2000-11&rft_id=info%3Apmid%2F11073453&rft_id=info%3Adoi%2F10.1126%2Fscience.290.5494.1155&rft_id=info%3Abibcode%2F2000Sci...290.1155S&rft.aulast=Semino&rft.aufirst=O&rft.au=Passarino%2C+G&rft.au=Oefner%2C+PJ&rft.au=Lin%2C+AA&rft.au=Arbuzova%2C+S&rft.au=Beckman%2C+LE&rft.au=De+Benedictis%2C+G&rft.au=Francalacci%2C+P&rft.au=Kouvatsi%2C+A&rft.au=Limborska%2C+S&rft.au=Marcikiae%2C+M&rft.au=Mika%2C+A&rft.au=Mika%2C+B&rft.au=Primorac%2C+D&rft.au=Santachiara-Benerecetti%2C+AS&rft.au=Cavalli-Sforza%2C+LL&rft.au=Underhill%2C+PA&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGenetic+history+of+Europe" class="Z3988"></span> Note: Haplogroup names are different in this article. For example: <a href="/wiki/Haplogroup_I_(Y-DNA)" class="mw-redirect" title="Haplogroup I (Y-DNA)">Haplogroup I</a> is referred as M170</span> </li> <li id="cite_note-Wells_2001-26"><span class="mw-cite-backlink">^ <a href="#cite_ref-Wells_2001_26-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Wells_2001_26-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFWellsYuldashevaRuzibakievUnderhill2001" class="citation journal cs1">Wells RS, Yuldasheva N, Ruzibakiev R, Underhill PA, Evseeva I, Blue-Smith J, et al. (August 2001). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC56946">"The Eurasian heartland: a continental perspective on Y-chromosome diversity"</a>. <i>Proceedings of the National Academy of Sciences of the United States of America</i>. <b>98</b> (18): 10244–10249. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2001PNAS...9810244W">2001PNAS...9810244W</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1073%2Fpnas.171305098">10.1073/pnas.171305098</a></span>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC56946">56946</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/11526236">11526236</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Proceedings+of+the+National+Academy+of+Sciences+of+the+United+States+of+America&rft.atitle=The+Eurasian+heartland%3A+a+continental+perspective+on+Y-chromosome+diversity&rft.volume=98&rft.issue=18&rft.pages=10244-10249&rft.date=2001-08&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC56946%23id-name%3DPMC&rft_id=info%3Apmid%2F11526236&rft_id=info%3Adoi%2F10.1073%2Fpnas.171305098&rft_id=info%3Abibcode%2F2001PNAS...9810244W&rft.aulast=Wells&rft.aufirst=RS&rft.au=Yuldasheva%2C+N&rft.au=Ruzibakiev%2C+R&rft.au=Underhill%2C+PA&rft.au=Evseeva%2C+I&rft.au=Blue-Smith%2C+J&rft.au=Jin%2C+L&rft.au=Su%2C+B&rft.au=Pitchappan%2C+R&rft.au=Shanmugalakshmi%2C+S&rft.au=Balakrishnan%2C+K&rft.au=Read%2C+M&rft.au=Pearson%2C+NM&rft.au=Zerjal%2C+T&rft.au=Webster%2C+MT&rft.au=Zholoshvili%2C+I&rft.au=Jamarjashvili%2C+E&rft.au=Gambarov%2C+S&rft.au=Nikbin%2C+B&rft.au=Dostiev%2C+A&rft.au=Aknazarov%2C+O&rft.au=Zalloua%2C+P&rft.au=Tsoy%2C+I&rft.au=Kitaev%2C+M&rft.au=Mirrakhimov%2C+M&rft.au=Chariev%2C+A&rft.au=Bodmer%2C+WF&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC56946&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGenetic+history+of+Europe" 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Archived from <a rel="nofollow" class="external text" href="http://www.ajhg.org/AJHG/abstract/S0002-9297(07)63221-2">the original</a> on 2008-05-06.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=American+Journal+of+Human+Genetics&rft.atitle=Y-chromosomal+diversity+in+Europe+is+clinal+and+influenced+primarily+by+geography%2C+rather+than+by+language&rft.volume=67&rft.issue=6&rft.pages=1526-1543&rft.date=2000-12&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC1287948%23id-name%3DPMC&rft_id=info%3Apmid%2F11078479&rft_id=info%3Adoi%2F10.1086%2F316890&rft.aulast=Rosser&rft.aufirst=ZH&rft.au=Zerjal%2C+T&rft.au=Hurles%2C+ME&rft.au=Adojaan%2C+M&rft.au=Alavantic%2C+D&rft.au=Amorim%2C+A&rft.au=Amos%2C+W&rft.au=Armenteros%2C+M&rft.au=Arroyo%2C+E&rft.au=Barbujani%2C+G&rft.au=Beckman%2C+G&rft.au=Beckman%2C+L&rft.au=Bertranpetit%2C+J&rft.au=Bosch%2C+E&rft.au=Bradley%2C+DG&rft.au=Brede%2C+G&rft.au=Cooper%2C+G&rft.au=C%C3%B4rte-Real%2C+HB&rft.au=de+Knijff%2C+P&rft.au=Decorte%2C+R&rft.au=Dubrova%2C+YE&rft.au=Evgrafov%2C+O&rft.au=Gilissen%2C+A&rft.au=Glisic%2C+S&rft.au=G%C3%B6lge%2C+M&rft.au=Hill%2C+EW&rft.au=Jeziorowska%2C+A&rft.au=Kalaydjieva%2C+L&rft.au=Kayser%2C+M&rft.au=Kivisild%2C+T&rft.au=Kravchenko%2C+SA&rft.au=Krumina%2C+A&rft.au=Kucinskas%2C+V&rft.au=Lavinha%2C+J&rft.au=Livshits%2C+LA&rft.au=Malaspina%2C+P&rft.au=Maria%2C+S&rft.au=McElreavey%2C+K&rft.au=Meitinger%2C+TA&rft.au=Mikelsaar%2C+AV&rft.au=Mitchell%2C+RJ&rft.au=Nafa%2C+K&rft.au=Nicholson%2C+J&rft.au=N%C3%B8rby%2C+S&rft.au=Pandya%2C+A&rft.au=Parik%2C+J&rft.au=Patsalis%2C+PC&rft.au=Pereira%2C+L&rft.au=Peterlin%2C+B&rft.au=Pielberg%2C+G&rft.au=Prata%2C+MJ&rft.au=Previder%C3%A9%2C+C&rft.au=Roewer%2C+L&rft.au=Rootsi%2C+S&rft.au=Rubinsztein%2C+DC&rft.au=Saillard%2C+J&rft.au=Santos%2C+FR&rft.au=Stefanescu%2C+G&rft.au=Sykes%2C+BC&rft.au=Tolun%2C+A&rft.au=Villems%2C+R&rft.au=Tyler-Smith%2C+C&rft.au=Jobling%2C+MA&rft_id=http%3A%2F%2Fwww.ajhg.org%2FAJHG%2Fabstract%2FS0002-9297%2807%2963221-2&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGenetic+history+of+Europe" class="Z3988"></span></span> </li> <li id="cite_note-Underhill_2007-45"><span class="mw-cite-backlink"><b><a href="#cite_ref-Underhill_2007_45-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFUnderhillKivisild2007" class="citation journal cs1">Underhill PA, Kivisild T (2007). "Use of y chromosome and mitochondrial DNA population structure in tracing human migrations". <i>Annual Review of Genetics</i>. <b>41</b>: 539–564. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1146%2Fannurev.genet.41.110306.130407">10.1146/annurev.genet.41.110306.130407</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/18076332">18076332</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Annual+Review+of+Genetics&rft.atitle=Use+of+y+chromosome+and+mitochondrial+DNA+population+structure+in+tracing+human+migrations&rft.volume=41&rft.pages=539-564&rft.date=2007&rft_id=info%3Adoi%2F10.1146%2Fannurev.genet.41.110306.130407&rft_id=info%3Apmid%2F18076332&rft.aulast=Underhill&rft.aufirst=PA&rft.au=Kivisild%2C+T&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGenetic+history+of+Europe" class="Z3988"></span></span> </li> <li id="cite_note-46"><span class="mw-cite-backlink"><b><a href="#cite_ref-46">^</a></b></span> <span class="reference-text">Y chromosome data show a signal for a separate late-Pleistocene migration from Africa to Europe via Sinai as evidenced through the distribution of haplogroup E3b lineages, which is not manifested in mtDNA haplogroup distributions.</span> </li> <li id="cite_note-Battaglia_2008-47"><span class="mw-cite-backlink">^ <a href="#cite_ref-Battaglia_2008_47-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Battaglia_2008_47-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBattagliaFornarinoAl-ZaheryOlivieri2009" class="citation journal cs1">Battaglia V, Fornarino S, Al-Zahery N, Olivieri A, Pala M, Myres NM, et al. (June 2009). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2947100">"Y-chromosomal evidence of the cultural diffusion of agriculture in Southeast Europe"</a>. <i>European Journal of Human Genetics</i>. <b>17</b> (6): 820–830. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2Fejhg.2008.249">10.1038/ejhg.2008.249</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2947100">2947100</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/19107149">19107149</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=European+Journal+of+Human+Genetics&rft.atitle=Y-chromosomal+evidence+of+the+cultural+diffusion+of+agriculture+in+Southeast+Europe&rft.volume=17&rft.issue=6&rft.pages=820-830&rft.date=2009-06&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2947100%23id-name%3DPMC&rft_id=info%3Apmid%2F19107149&rft_id=info%3Adoi%2F10.1038%2Fejhg.2008.249&rft.aulast=Battaglia&rft.aufirst=V&rft.au=Fornarino%2C+S&rft.au=Al-Zahery%2C+N&rft.au=Olivieri%2C+A&rft.au=Pala%2C+M&rft.au=Myres%2C+NM&rft.au=King%2C+RJ&rft.au=Rootsi%2C+S&rft.au=Marjanovic%2C+D&rft.au=Primorac%2C+D&rft.au=Hadziselimovic%2C+R&rft.au=Vidovic%2C+S&rft.au=Drobnic%2C+K&rft.au=Durmishi%2C+N&rft.au=Torroni%2C+A&rft.au=Santachiara-Benerecetti%2C+AS&rft.au=Underhill%2C+PA&rft.au=Semino%2C+O&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2947100&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGenetic+history+of+Europe" class="Z3988"></span></span> </li> <li id="cite_note-Cruciani_2007-48"><span class="mw-cite-backlink"><b><a href="#cite_ref-Cruciani_2007_48-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFCrucianiLa_FrattaTrombettaSantolamazza2007" class="citation journal cs1">Cruciani F, La Fratta R, Trombetta B, Santolamazza P, Sellitto D, Colomb EB, et al. (June 2007). <a rel="nofollow" class="external text" href="https://wayback.archive-it.org/all/20171010093322/https://academic.oup.com/mbe/article/24/6/1300/984002/Tracing-Past-Human-Male-Movements-in-Northern">"Tracing past human male movements in northern/eastern Africa and western Eurasia: new clues from Y-chromosomal haplogroups E-M78 and J-M12"</a>. <i>Molecular Biology and Evolution</i>. <b>24</b> (6): 1300–1311. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1093%2Fmolbev%2Fmsm049">10.1093/molbev/msm049</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/17351267">17351267</a>. Archived from <a rel="nofollow" class="external text" href="http://mbe.oxfordjournals.org/cgi/reprint/24/6/1300">the original</a> on 2017-10-10<span class="reference-accessdate">. Retrieved <span class="nowrap">2009-07-22</span></span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Molecular+Biology+and+Evolution&rft.atitle=Tracing+past+human+male+movements+in+northern%2Feastern+Africa+and+western+Eurasia%3A+new+clues+from+Y-chromosomal+haplogroups+E-M78+and+J-M12&rft.volume=24&rft.issue=6&rft.pages=1300-1311&rft.date=2007-06&rft_id=info%3Adoi%2F10.1093%2Fmolbev%2Fmsm049&rft_id=info%3Apmid%2F17351267&rft.aulast=Cruciani&rft.aufirst=F&rft.au=La+Fratta%2C+R&rft.au=Trombetta%2C+B&rft.au=Santolamazza%2C+P&rft.au=Sellitto%2C+D&rft.au=Colomb%2C+EB&rft.au=Dugoujon%2C+JM&rft.au=Crivellaro%2C+F&rft.au=Benincasa%2C+T&rft.au=Pascone%2C+R&rft.au=Moral%2C+P&rft.au=Watson%2C+E&rft.au=Melegh%2C+B&rft.au=Barbujani%2C+G&rft.au=Fuselli%2C+S&rft.au=Vona%2C+G&rft.au=Zagradisnik%2C+B&rft.au=Assum%2C+G&rft.au=Brdicka%2C+R&rft.au=Kozlov%2C+AI&rft.au=Efremov%2C+GD&rft.au=Coppa%2C+A&rft.au=Novelletto%2C+A&rft.au=Scozzari%2C+R&rft_id=http%3A%2F%2Fmbe.oxfordjournals.org%2Fcgi%2Freprint%2F24%2F6%2F1300&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGenetic+history+of+Europe" class="Z3988"></span> Also see <a rel="nofollow" class="external text" href="https://archive.today/20121205040152/http://mbe.oxfordjournals.org/cgi/content/full/msm049/DC1?maxtoshow=&HITS=10&hits=10&RESULTFORMAT=&fulltext=cruciani&searchid=1&FIRSTINDEX=0&resourcetype=HWCIT">Supplementary Data</a>.</span> </li> <li id="cite_note-Lacan_2011-49"><span class="mw-cite-backlink"><b><a href="#cite_ref-Lacan_2011_49-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLacanKeyserRicautBrucato2011" class="citation journal cs1">Lacan M, Keyser C, Ricaut FX, Brucato N, Duranthon F, Guilaine J, Crubézy E, Ludes B (June 2011). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3116412">"Ancient DNA reveals male diffusion through the Neolithic Mediterranean route"</a>. <i>Proceedings of the National Academy of Sciences of the United States of America</i>. <b>108</b> (24): 9788–91. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2011PNAS..108.9788L">2011PNAS..108.9788L</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1073%2Fpnas.1100723108">10.1073/pnas.1100723108</a></span>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3116412">3116412</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/21628562">21628562</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Proceedings+of+the+National+Academy+of+Sciences+of+the+United+States+of+America&rft.atitle=Ancient+DNA+reveals+male+diffusion+through+the+Neolithic+Mediterranean+route&rft.volume=108&rft.issue=24&rft.pages=9788-91&rft.date=2011-06&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3116412%23id-name%3DPMC&rft_id=info%3Apmid%2F21628562&rft_id=info%3Adoi%2F10.1073%2Fpnas.1100723108&rft_id=info%3Abibcode%2F2011PNAS..108.9788L&rft.aulast=Lacan&rft.aufirst=M&rft.au=Keyser%2C+C&rft.au=Ricaut%2C+FX&rft.au=Brucato%2C+N&rft.au=Duranthon%2C+F&rft.au=Guilaine%2C+J&rft.au=Crub%C3%A9zy%2C+E&rft.au=Ludes%2C+B&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3116412&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGenetic+history+of+Europe" class="Z3988"></span></span> </li> <li id="cite_note-Bird_2007-50"><span class="mw-cite-backlink">^ <a href="#cite_ref-Bird_2007_50-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Bird_2007_50-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBird2007" class="citation journal cs1">Bird S (2007). <a rel="nofollow" class="external text" href="https://web.archive.org/web/20160422005652/http://www.jogg.info/32/bird.htm">"Haplogroup E3b1a2 as a Possible Indicator of Settlement in Roman Britain by Soldiers of Balkan Origin"</a>. <i>Journal of Genetic Genealogy</i>. <b>3</b> (2). 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Archived from <a rel="nofollow" class="external text" href="http://dirkschweitzer.net/E3b-papers/HG-05-428-Poland-Germany.pdf">the original</a> <span class="cs1-format">(PDF)</span> on 2009-03-04<span class="reference-accessdate">. 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"Y chromosome sequence variation and the history of human populations". <i>Nature Genetics</i>. <b>26</b> (3): 358–361. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2F81685">10.1038/81685</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/11062480">11062480</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:12893406">12893406</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nature+Genetics&rft.atitle=Y+chromosome+sequence+variation+and+the+history+of+human+populations&rft.volume=26&rft.issue=3&rft.pages=358-361&rft.date=2000-11&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A12893406%23id-name%3DS2CID&rft_id=info%3Apmid%2F11062480&rft_id=info%3Adoi%2F10.1038%2F81685&rft.aulast=Underhill&rft.aufirst=PA&rft.au=Shen%2C+P&rft.au=Lin%2C+AA&rft.au=Jin%2C+L&rft.au=Passarino%2C+G&rft.au=Yang%2C+WH&rft.au=Kauffman%2C+E&rft.au=Bonn%C3%A9-Tamir%2C+B&rft.au=Bertranpetit%2C+J&rft.au=Francalacci%2C+P&rft.au=Ibrahim%2C+M&rft.au=Jenkins%2C+T&rft.au=Kidd%2C+JR&rft.au=Mehdi%2C+SQ&rft.au=Seielstad%2C+MT&rft.au=Wells%2C+RS&rft.au=Piazza%2C+A&rft.au=Davis%2C+RW&rft.au=Feldman%2C+MW&rft.au=Cavalli-Sforza%2C+LL&rft.au=Oefner%2C+PJ&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGenetic+history+of+Europe" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFUnderhillPassarinoLinShen2001" class="citation journal cs1">Underhill PA, Passarino G, Lin AA, Shen P, Mirazón Lahr M, Foley RA, et al. (January 2001). <a rel="nofollow" class="external text" href="https://doi.org/10.1046%2Fj.1469-1809.2001.6510043.x">"The phylogeography of Y chromosome binary haplotypes and the origins of modern human populations"</a>. <i>Annals of Human Genetics</i>. <b>65</b> (Pt 1): 43–62. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1046%2Fj.1469-1809.2001.6510043.x">10.1046/j.1469-1809.2001.6510043.x</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/11415522">11415522</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:9441236">9441236</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Annals+of+Human+Genetics&rft.atitle=The+phylogeography+of+Y+chromosome+binary+haplotypes+and+the+origins+of+modern+human+populations&rft.volume=65&rft.issue=Pt+1&rft.pages=43-62&rft.date=2001-01&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A9441236%23id-name%3DS2CID&rft_id=info%3Apmid%2F11415522&rft_id=info%3Adoi%2F10.1046%2Fj.1469-1809.2001.6510043.x&rft.aulast=Underhill&rft.aufirst=PA&rft.au=Passarino%2C+G&rft.au=Lin%2C+AA&rft.au=Shen%2C+P&rft.au=Miraz%C3%B3n+Lahr%2C+M&rft.au=Foley%2C+RA&rft.au=Oefner%2C+PJ&rft.au=Cavalli-Sforza%2C+LL&rft_id=https%3A%2F%2Fdoi.org%2F10.1046%252Fj.1469-1809.2001.6510043.x&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGenetic+history+of+Europe" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFUnderhill2002" class="citation book cs1">Underhill PA (2002). "Inference of Neolithic Population Histories using Y-chromosome Haplotypes". In Bellwood and Renfrew (ed.). <i>Examining the farming/language dispersal hypothesis, McDonald Institute for Archaeological Research</i>. Cambridge: McDonald Institute for Archaeological Research. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-1-902937-20-5" title="Special:BookSources/978-1-902937-20-5"><bdi>978-1-902937-20-5</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=Inference+of+Neolithic+Population+Histories+using+Y-chromosome+Haplotypes&rft.btitle=Examining+the+farming%2Flanguage+dispersal+hypothesis%2C+McDonald+Institute+for+Archaeological+Research&rft.place=Cambridge&rft.pub=McDonald+Institute+for+Archaeological+Research&rft.date=2002&rft.isbn=978-1-902937-20-5&rft.aulast=Underhill&rft.aufirst=PA&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGenetic+history+of+Europe" class="Z3988"></span></li> <li>Perlès C, Monthel G ( 2001) The Early Neolithic in Greece: The First Farming Communities in Europe. <a href="/wiki/Cambridge_University_Press" title="Cambridge University Press">Cambridge University Press</a>, Cambridge.</li> <li>Runnels C (2003) The origins of the Greek Neolithic: a personal view, in Ammerman and Biagi (2003 eds).</li></ul> </div> <p><br /> </p> <div class="mw-heading mw-heading2"><h2 id="Further_reading">Further reading</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Genetic_history_of_Europe&action=edit&section=22" title="Edit section: Further reading"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li>Rodríguez-Varela, Ricardo et al. "The genetic history of Scandinavia from the Roman Iron Age to the present". In: <i>Cell</i>. Volume 186, Issue 1, 5 January 2023, Pages 32–46.e19. <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.cell.2022.11.024">10.1016/j.cell.2022.11.024</a></li> <li>Skourtanioti, E., Ringbauer, H., Gnecchi Ruscone, G.A. et al. "Ancient DNA reveals admixture history and endogamy in the prehistoric Aegean". In: <i>Nature Ecology & Evolution</i> (2023). <a rel="nofollow" class="external free" href="https://doi.org/10.1038/s41559-022-01952-3">https://doi.org/10.1038/s41559-022-01952-3</a></li></ul> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Genetic_history_of_Europe&action=edit&section=23" title="Edit section: External links"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1235681985">.mw-parser-output .side-box{margin:4px 0;box-sizing:border-box;border:1px solid #aaa;font-size:88%;line-height:1.25em;background-color:var(--background-color-interactive-subtle,#f8f9fa);display:flow-root}.mw-parser-output .side-box-abovebelow,.mw-parser-output .side-box-text{padding:0.25em 0.9em}.mw-parser-output .side-box-image{padding:2px 0 2px 0.9em;text-align:center}.mw-parser-output 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.navbar{display:none!important}}</style><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Human_genetics" title="Template:Human genetics"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Human_genetics" title="Template talk:Human genetics"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Human_genetics" title="Special:EditPage/Template:Human genetics"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Human_genetics" style="font-size:114%;margin:0 4em"><a href="/wiki/Human_genetics" title="Human genetics">Human genetics</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align: center;">Sub-topics</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Human_genome" title="Human genome">Human genome</a> <ul><li><a href="/wiki/Human_Genome_Project" title="Human Genome Project">Human Genome Project</a></li></ul></li> <li><a href="/wiki/Human_evolutionary_genetics" title="Human evolutionary genetics">Evolutionary genetics</a> <ul><li><a href="/wiki/Chimpanzee%E2%80%93human_last_common_ancestor" title="Chimpanzee–human last common ancestor">Human-chimp MRCA</a></li> <li><a href="/wiki/Neanderthal_genetics" title="Neanderthal genetics">Neanderthal genetics</a> <ul><li><a href="/wiki/Neanderthal_genome_project" title="Neanderthal genome project">Neanderthal genome project</a></li></ul></li> <li><a href="/wiki/Timeline_of_human_evolution" title="Timeline of human evolution">Timeline</a></li></ul></li> <li><a href="/wiki/Human_genetic_variation" title="Human genetic variation">Genetic variation</a> <ul><li><a href="/wiki/Blood_type_distribution_by_country" title="Blood type distribution by country">Blood type distribution by country</a></li> <li><a href="/wiki/Genealogical_DNA_test" title="Genealogical DNA test">Genealogical DNA test</a></li> <li><a href="/wiki/Genetic_genealogy" title="Genetic genealogy">Genetic genealogy</a></li> <li><a href="/wiki/Race_and_genetics" title="Race and genetics">Race and genetics</a></li> <li><a href="/wiki/Recent_human_evolution" title="Recent human evolution">Recent evolution</a></li> <li><a href="/wiki/Surname_DNA_project" title="Surname DNA project">Surname DNA project</a></li></ul></li> <li><a href="/wiki/Human_genetic_enhancement" title="Human genetic enhancement">Genetic enhancement</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align: center;"><a href="/wiki/Archaeogenetics" title="Archaeogenetics">Genetic history</a> <br />by region</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Genetic_history_of_Africa" title="Genetic history of Africa">Africa</a> <ul><li><a href="/wiki/Genetic_history_of_North_Africa" title="Genetic history of North Africa">North Africa</a></li> <li><a href="/wiki/Genetic_history_of_West_Africa" title="Genetic history of West Africa">West Africa</a></li> <li><a href="/wiki/Genetic_history_of_Central_Africa" title="Genetic history of Central Africa">Central Africa</a></li> <li><a href="/wiki/Genetic_history_of_Eastern_Africa" title="Genetic history of Eastern Africa">Eastern Africa</a></li> <li><a href="/wiki/Genetic_history_of_Southern_Africa" title="Genetic history of Southern Africa">Southern Africa</a></li> <li><a href="/wiki/Genetic_history_of_the_African_diaspora" title="Genetic history of the African diaspora">African diaspora</a></li></ul></li> <li><a href="/wiki/Genetics_and_archaeogenetics_of_South_Asia" title="Genetics and archaeogenetics of South Asia">South Asia</a> <ul><li><a href="/wiki/Peopling_of_India" title="Peopling of India">India</a></li></ul></li> <li><a href="/wiki/Genetic_history_of_the_Middle_East" title="Genetic history of the Middle East">Middle East</a> <ul><li><a href="/wiki/Early_Anatolian_farmers" class="mw-redirect" title="Early Anatolian farmers">Early Anatolian farmers</a></li></ul></li> <li><a href="/wiki/Genetic_history_of_the_Caucasus" class="mw-redirect" title="Genetic history of the Caucasus">Caucasus</a> <ul><li><a href="/wiki/Caucasus_hunter-gatherer" title="Caucasus hunter-gatherer">Caucasus hunter-gatherer</a></li></ul></li> <li><a class="mw-selflink selflink">Europe</a> <ul><li><a href="/wiki/Early_European_Farmers" title="Early European Farmers">Early European Farmers</a></li> <li><a href="/wiki/Western_hunter-gatherer" title="Western hunter-gatherer">Western hunter-gatherer</a></li> <li><a href="/wiki/Genetic_history_of_the_British_Isles" title="Genetic history of the British Isles">British Isles</a></li> <li><a href="/wiki/Genetic_history_of_the_Iberian_Peninsula" title="Genetic history of the Iberian Peninsula">Iberia</a></li> <li><a href="/wiki/Genetic_history_of_Italy" title="Genetic history of Italy">Italy</a></li> <li><a href="/wiki/Eastern_hunter-gatherer" title="Eastern hunter-gatherer">Eastern hunter-gatherer</a></li></ul></li> <li><a href="/wiki/Genetic_history_of_Central_Asia" class="mw-redirect" title="Genetic history of Central Asia">Central Asia</a> <ul><li><a href="/wiki/Ancient_North_Eurasian" title="Ancient North Eurasian">Ancient North Eurasian</a></li></ul></li> <li><a href="/wiki/Genetic_history_of_East_Asians" title="Genetic history of East Asians">East Asia</a> <ul><li><a href="/wiki/Ancient_Northeast_Asian" title="Ancient Northeast Asian">Ancient Northeast Asian</a></li> <li><a href="/wiki/Ancient_Paleo-Siberian" title="Ancient Paleo-Siberian">Ancient Paleo-Siberian</a></li> <li><a href="/wiki/Peopling_of_China" title="Peopling of China">China</a></li></ul></li> <li><a href="/wiki/Peopling_of_Southeast_Asia" title="Peopling of Southeast Asia">Southeast Asia</a> <ul><li><a href="/wiki/Peopling_of_Thailand" title="Peopling of Thailand">Thailand</a></li></ul></li> <li><a href="/wiki/Genetic_history_of_Indigenous_peoples_of_the_Americas" class="mw-redirect" title="Genetic history of Indigenous peoples of the Americas">America</a> <ul><li><a href="/wiki/Ancient_Beringian" title="Ancient Beringian">Ancient Beringian</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align: center;"><a href="/wiki/Population_genetics" title="Population genetics">Population genetics</a> <br />by group</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li>Europe <ul><li><a href="/wiki/Origin_of_the_Basques" title="Origin of the Basques">Basques</a></li> <li><a href="/wiki/Genetic_studies_on_Bosniaks" title="Genetic studies on Bosniaks">Bosniaks</a></li> <li><a href="/wiki/Genetic_studies_on_Bulgarians" title="Genetic studies on Bulgarians">Bulgarians</a></li> <li><a href="/wiki/Genetic_studies_on_Croats" title="Genetic studies on Croats">Croats</a></li> <li><a href="/wiki/Genetic_studies_on_Romanians" class="mw-redirect" title="Genetic studies on Romanians">Romanians</a></li> <li><a href="/wiki/Genetic_studies_on_Russians" title="Genetic studies on Russians">Russians</a></li> <li><a href="/wiki/Genetic_studies_on_Sami" title="Genetic studies on Sami">Sami</a></li> <li><a href="/wiki/Genetic_studies_on_Serbs" title="Genetic studies on Serbs">Serbs</a></li></ul></li> <li>MENA <ul><li><a href="/wiki/Genetic_studies_on_Arabs" title="Genetic studies on Arabs">Arabs</a></li> <li><a href="/wiki/Origin_of_the_Azerbaijanis" title="Origin of the Azerbaijanis">Azerbaijanis</a></li> <li><a href="/wiki/Genetic_history_of_Egypt" title="Genetic history of Egypt">Egyptians</a></li> <li><a href="/wiki/Genetic_studies_on_Jews" class="mw-redirect" title="Genetic studies on Jews">Jews</a></li> <li><a href="/wiki/Genetic_studies_on_Moroccans" title="Genetic studies on Moroccans">Moroccans</a></li> <li><a href="/wiki/Genetic_studies_on_Turkish_people" title="Genetic studies on Turkish people">Turks</a></li></ul></li> <li>South Asia <ul><li><a href="/wiki/Genetic_studies_on_Gujarati_people" title="Genetic studies on Gujarati people">Gujaratis</a></li> <li><a href="/wiki/Genetic_studies_on_Sinhalese" title="Genetic studies on Sinhalese">Sinhalese</a></li> <li><a href="/wiki/Genetic_studies_on_Sri_Lankan_Tamils" title="Genetic studies on Sri Lankan Tamils">Tamils (Sri Lankan)</a></li></ul></li> <li>East Asia <ul><li><a href="/wiki/Genetic_studies_on_Han_Chinese" class="mw-redirect" title="Genetic studies on Han Chinese">Han Chinese</a></li> <li><a href="/wiki/Genetic_and_anthropometric_studies_on_Japanese_people" title="Genetic and anthropometric studies on Japanese people">Japanese</a></li></ul></li> <li>Southeast Asia <ul><li><a href="/wiki/Genetic_studies_on_Filipinos" title="Genetic studies on Filipinos">Filipinos</a></li></ul></li> <li>Sub-Saharan Africa <ul><li><a href="/wiki/Genetic_studies_on_Hutu_and_Tutsi" class="mw-redirect" title="Genetic studies on Hutu and Tutsi">Hutu/Tutsi</a></li></ul></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><span class="noviewer" typeof="mw:File"><span title="Category"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/16px-Symbol_category_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/23px-Symbol_category_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/31px-Symbol_category_class.svg.png 2x" data-file-width="180" data-file-height="185" /></span></span><b><a href="/wiki/Category:Human_genetics" title="Category:Human genetics">Category</a></b></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="History_of_Europe" style="padding:3px"><table class="nowraplinks mw-collapsible mw-collapsed navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:History_of_Europe" title="Template:History of Europe"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:History_of_Europe" title="Template talk:History of Europe"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:History_of_Europe" title="Special:EditPage/Template:History of Europe"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="History_of_Europe" style="font-size:114%;margin:0 4em"><a href="/wiki/History_of_Europe" title="History of Europe">History of Europe</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Prehistoric_Europe" title="Prehistoric Europe">Prehistory</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Paleolithic_Europe" title="Paleolithic Europe">Paleolithic Europe</a></li> <li><a href="/wiki/Neolithic_Europe" title="Neolithic Europe">Neolithic Europe</a></li> <li><a href="/wiki/Bronze_Age_Europe" title="Bronze Age Europe">Bronze Age Europe</a></li> <li><a href="/wiki/Iron_Age_Europe" title="Iron Age Europe">Iron Age Europe</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Classical_antiquity" title="Classical antiquity">Classical antiquity</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Classical_Greece" title="Classical Greece">Classical Greece</a></li> <li><a href="/wiki/Roman_Republic" title="Roman Republic">Roman Republic</a></li> <li><a href="/wiki/Hellenistic_period" title="Hellenistic period">Hellenistic period</a></li> <li><a href="/wiki/Roman_Empire" title="Roman Empire">Roman Empire</a></li> <li><a href="/wiki/Early_Christianity" title="Early Christianity">Early Christianity</a> <ul><li><a href="/wiki/Christianity_in_late_antiquity" title="Christianity in late antiquity">Christianity in late antiquity</a></li></ul></li> <li><a href="/wiki/Crisis_of_the_Third_Century" title="Crisis of the Third Century">Crisis of the Third Century</a></li> <li><a href="/wiki/Fall_of_the_Western_Roman_Empire" title="Fall of the Western Roman Empire">Fall of the Western Roman Empire</a></li> <li><a href="/wiki/Late_antiquity" title="Late antiquity">Late antiquity</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Middle_Ages" title="Middle Ages">Middle Ages</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Early_Middle_Ages" title="Early Middle Ages">Early Middle Ages</a></li> <li><a href="/wiki/Migration_Period" title="Migration Period">Migration Period</a></li> <li><a href="/wiki/Christianity_in_the_Middle_Ages" title="Christianity in the Middle Ages">Christianity in the Middle Ages</a> <ul><li><a href="/wiki/Christianization" title="Christianization">Christianization</a></li></ul></li> <li><a href="/wiki/Francia" title="Francia">Francia</a></li> <li><a href="/wiki/History_of_Anglo-Saxon_England" title="History of Anglo-Saxon England">Anglo-Saxon England</a></li> <li><a href="/wiki/Byzantine_Empire" title="Byzantine Empire">Byzantine Empire</a></li> <li><a href="/wiki/Papal_States" title="Papal States">Papal States</a></li> <li>Bulgarian Empire <ul><li><a href="/wiki/First_Bulgarian_Empire" title="First Bulgarian Empire">First</a></li> <li><a href="/wiki/Second_Bulgarian_Empire" title="Second Bulgarian Empire">Second</a></li></ul></li> <li><a href="/wiki/Maritime_republics" title="Maritime republics">Maritime republics</a> <ul><li><a href="/wiki/Republic_of_Venice" title="Republic of Venice">Venice</a></li> <li><a href="/wiki/Republic_of_Genoa" title="Republic of Genoa">Genoa</a></li> <li><a href="/wiki/Republic_of_Pisa" title="Republic of Pisa">Pisa</a></li> <li><a href="/wiki/Duchy_of_Amalfi" title="Duchy of Amalfi">Amalfi</a></li></ul></li> <li><a href="/wiki/Viking_Age" title="Viking Age">Viking Age</a></li> <li><a href="/wiki/Kievan_Rus%27" title="Kievan Rus'">Kievan Rus'</a></li> <li><a href="/wiki/Crown_of_Aragon" title="Crown of Aragon">Crown of Aragon</a> (<a href="/wiki/Kingdom_of_Aragon" title="Kingdom of Aragon">Aragon</a>, <a href="/wiki/Principality_of_Catalonia" title="Principality of Catalonia">Catalonia</a>, <a href="/wiki/Kingdom_of_Valencia" title="Kingdom of Valencia">Valencia</a>, <a href="/wiki/Kingdom_of_Majorca" title="Kingdom of Majorca">Majorca</a>)</li> <li><a href="/wiki/Holy_Roman_Empire" title="Holy Roman Empire">Holy Roman Empire</a></li> <li><a href="/wiki/High_Middle_Ages" title="High Middle Ages">High Middle Ages</a></li> <li><a href="/wiki/Republic_of_Florence" title="Republic of Florence">Republic of Florence</a></li> <li><a href="/wiki/Feudalism" title="Feudalism">Feudalism</a></li> <li><a href="/wiki/Crusades" title="Crusades">Crusades</a></li> <li><a href="/wiki/Mongol_invasion_of_Europe" title="Mongol invasion of Europe">Mongol invasion</a></li> <li><a href="/wiki/Serbian_Empire" title="Serbian Empire">Serbian Empire</a></li> <li><a href="/wiki/Late_Middle_Ages" title="Late Middle Ages">Late Middle Ages</a></li> <li><a href="/wiki/Black_Death" title="Black Death">Black Death</a></li> <li><a href="/wiki/Hundred_Years%27_War" title="Hundred Years' War">Hundred Years' War</a></li> <li><a href="/wiki/Kalmar_Union" title="Kalmar Union">Kalmar Union</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Modern_period" class="mw-redirect" title="Modern period">Modern period</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Renaissance" title="Renaissance">Renaissance</a></li> <li><a href="/wiki/Early_modern_Europe" title="Early modern Europe">Early modern</a></li> <li><a href="/wiki/Christianity_in_the_modern_era" title="Christianity in the modern era">Christianity in the modern era</a> <ul><li><a href="/wiki/Reformation" title="Reformation">Reformation</a></li></ul></li> <li><a href="/wiki/Age_of_Discovery" title="Age of Discovery">Age of Discovery</a></li> <li><a href="/wiki/Baroque" title="Baroque">Baroque</a></li> <li><a href="/wiki/Grand_Duchy_of_Tuscany" title="Grand Duchy of Tuscany">Grand Duchy of Tuscany</a></li> <li><a href="/wiki/Thirty_Years%27_War" title="Thirty Years' War">Thirty Years' War</a></li> <li><a href="/wiki/Absolute_monarchy" title="Absolute monarchy">Absolute monarchy</a></li> <li><a href="/wiki/Ottoman_Empire" title="Ottoman Empire">Ottoman Empire</a></li> <li><a href="/wiki/Portuguese_Empire" title="Portuguese Empire">Portuguese Empire</a></li> <li><a href="/wiki/Spanish_Empire" title="Spanish Empire">Spanish Empire</a></li> <li><a href="/wiki/Early_modern_France" class="mw-redirect" title="Early modern France">Early modern France</a></li> <li><a href="/wiki/Polish%E2%80%93Lithuanian_Commonwealth" title="Polish–Lithuanian Commonwealth">Polish–Lithuanian Commonwealth</a></li> <li><a href="/wiki/Cossack_Hetmanate" title="Cossack Hetmanate">Cossack Hetmanate</a></li> <li><a href="/wiki/Swedish_Empire" title="Swedish Empire">Swedish Empire</a></li> <li><a href="/wiki/Dutch_Republic" title="Dutch Republic">Dutch Republic</a></li> <li><a href="/wiki/British_Empire" title="British Empire">British Empire</a></li> <li><a href="/wiki/Habsburg_monarchy" title="Habsburg monarchy">Habsburg monarchy</a></li> <li><a href="/wiki/Russian_Empire" title="Russian Empire">Russian Empire</a></li> <li><a href="/wiki/Age_of_Enlightenment" title="Age of Enlightenment">Age of Enlightenment</a></li> <li><a href="/wiki/Great_Divergence" title="Great Divergence">Great Divergence</a></li> <li><a href="/wiki/Industrial_Revolution" title="Industrial Revolution">Industrial Revolution</a></li> <li><a href="/wiki/French_Revolution" title="French Revolution">French Revolution</a></li> <li><a href="/wiki/Napoleonic_Wars" title="Napoleonic Wars">Napoleonic Wars</a></li> <li><a href="/wiki/Rise_of_nationalism_in_Europe" title="Rise of nationalism in Europe">Nationalism</a></li> <li><a href="/wiki/Revolutions_of_1848" title="Revolutions of 1848">Revolutions of 1848</a></li> <li><a href="/wiki/World_War_I" title="World War I">World War I</a></li> <li><a href="/wiki/Russian_Revolution" title="Russian Revolution">Russian Revolution</a></li> <li><a href="/wiki/Interwar_period" title="Interwar period">Interwar period</a></li> <li><a href="/wiki/World_War_II" title="World War II">World War II</a></li> <li><a href="/wiki/Cold_War" title="Cold War">Cold War</a></li> <li><a href="/wiki/European_integration" title="European integration">European integration</a></li> <li><a href="/wiki/European_debt_crisis" title="European debt crisis">European debt crisis</a></li> <li><a href="/wiki/COVID-19_pandemic_in_Europe" title="COVID-19 pandemic in Europe">COVID-19 pandemic</a></li> <li><a href="/wiki/Russian_invasion_of_Ukraine" title="Russian invasion of Ukraine">Russian invasion of Ukraine</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">See also</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Art_of_Europe" title="Art of Europe">Art of Europe</a></li> <li><a href="/wiki/Bibliography_of_European_history" title="Bibliography of European history">Bibliography of European history</a></li> <li><a class="mw-selflink selflink">Genetic history of Europe</a></li> <li><a href="/wiki/History_of_Christianity" title="History of Christianity">History of Christianity</a></li> <li><a href="/wiki/History_of_the_Mediterranean_region" title="History of the Mediterranean region">History of the Mediterranean region</a></li> <li><a href="/wiki/History_of_the_European_Union" title="History of the European Union">History of the European Union</a></li> <li><a href="/wiki/History_of_Western_civilization" title="History of Western civilization">History of Western civilization</a></li> <li><a href="/wiki/Maritime_history_of_Europe" title="Maritime history of Europe">Maritime history of Europe</a></li> <li><a href="/wiki/Military_history_of_Europe" title="Military history of Europe">Military history of Europe</a></li> <li><a href="/wiki/Crusading_movement" title="Crusading movement">Crusading movement</a></li></ul> </div></td></tr></tbody></table></div> <!-- NewPP limit report Parsed by mw‐web.codfw.main‐55db797859‐9v9rm Cached time: 20241218031716 Cache expiry: 2592000 Reduced expiry: false Complications: [vary‐revision‐sha1, show‐toc] CPU time usage: 2.081 seconds Real time usage: 2.373 seconds Preprocessor visited node count: 12449/1000000 Post‐expand include size: 504266/2097152 bytes Template argument size: 6520/2097152 bytes Highest expansion depth: 16/100 Expensive parser function count: 44/500 Unstrip recursion depth: 1/20 Unstrip 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