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There is a general consensus that these languages are genealogically related and descend from a common ancestral language, conventionally called ‘Proto-Tungusic’. However, the exact geographical location where the ancestral speakers of Proto-Tungusic originated from is subject to debate. Here we take an unprecedented approach to this problem, by integrating linguistic, archaeological and genetic evidence in a single study. Our analysis of ancient DNA suggests genetic continuity between an ancient Amur genetic lineage and the contemporary speakers of the Tungusic languages. Adding an archaeolinguistic perspective, we infer that the most plausible homeland for the speakers of Proto-Tungusic is the region around Lake Khanka in the Russian Far East. Our study pushes the field forward in answering the tantalizing question about the location of the Tungusic homeland and in illustrating how these three disciplines can converge into a holistic approach to the human past."> <meta name="citation_doi" content="10.1017/ehs.2020.8"> <link rel="alternate" href="/core/journals/evolutionary-human-sciences/article/homeland-of-prototungusic-inferred-from-contemporary-words-and-ancient-genomes/950020A1799B4D7B71CCC57D04743B3E" hreflang="en" /> <link rel="icon" href="/core/cambridge-core/public/images/favicon.ico" type="image/x-icon"/> <link rel="shortcut icon" href="/core/cambridge-core/public/images/favicon.ico" type="image/x-icon"/> <link href='//fonts.googleapis.com/css?family=Noto+Sans:400,700,400italic,700italic' rel='stylesheet' type='text/css'>  <link rel="stylesheet" href="/core/cambridge-core/public/css/app.css?version=v7.337.1-hotfix"/> <link rel="dns-prefetch" href="https://usage.prod.aop.cambridge.org/v1/events"> <link 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data-v-3692cf84><span data-v-63dfaf6e>Volume 2</span> </a></li><li class="page-breadcrumbs__item" data-v-3692cf84><span aria-hidden="true" class="breadcrumbs-wrapper__arrow" data-v-3692cf84>></span><span data-v-3692cf84>The homeland of Proto-Tungusic inferred from contemporary...</span></li></ul></div></div> <div class="language" data-v-3692cf84><ul class="language-switch" data-v-6b1118dd data-v-3692cf84><li aria-label="English" data-v-6b1118dd><span class="language-option current divider" data-v-6b1118dd>English</span></li><li aria-label="Français" data-v-6b1118dd><span role="button" tabindex="0" href="#" lang="fr" class="language-option" data-v-6b1118dd> Français </span></li></ul></div></div></div></div> <div class="container container__modified" data-v-01274b1d>  <div class="row" data-v-01274b1d><div role="complementary" aria-label="table of content" class="column__left" data-v-01274b1d><div class="col journal-container row" data-v-146270e8 data-v-01274b1d><img src="https://static.cambridge.org/covers/EHS_0_0_0/evolutionary-human-sciences.jpg" alt="" class="journal__image" data-v-146270e8> <a href="/core/journals/evolutionary-human-sciences" class="app-link journal__title app-link__text app-link--underlined" data-v-63dfaf6e data-v-146270e8><span class="text" data-v-63dfaf6e>Evolutionary Human Sciences </span> </a> <hr aria-hidden="true" class="separator default" data-v-7036083a data-v-146270e8></div>  <div id="toc" class="table-of-content" data-v-01274b1d><h2>Article contents</h2> <div id="toc-list-wrapper" class="table-of-content__wrapper"><ul id="toc-list" class="list"><li class="list__item"><a href="#sec0" class="list__item__link"><span class="toc-title">Abstract</span></a></li> <li class="list__item"><a href="#sec1" class="list__item__link"> <span><div class="toc-title">Introduction</div></span></a></li><li class="list__item"><a href="#sec2" class="list__item__link"> <span><div class="toc-title">Methods</div></span></a></li><li class="list__item"><a href="#sec3" class="list__item__link"> <span><div class="toc-title">Results</div></span></a></li><li class="list__item"><a href="#sec4" class="list__item__link"> <span><div class="toc-title">Discussion</div></span></a></li>  <li class="list__item"><a href="#references-list" class="list__item__link"><span class="toc-title">References</span></a></li></ul></div></div></div> <div class="column__main" data-v-01274b1d><div class="row" data-v-01274b1d><div class="column__main__left" data-v-01274b1d><div id="maincontent" class="col" data-v-862424e6 data-v-01274b1d> <hgroup data-v-862424e6><h1 data-v-862424e6>The homeland of Proto-Tungusic inferred from contemporary words and ancient genomes</h1> </hgroup>     <div class="row published-date" data-v-862424e6><p data-v-862424e6> Published online by Cambridge University Press: <strong data-v-862424e6>22 April 2020</strong></p></div>   <div class="contributors-details" data-v-99f6eb26 data-v-862424e6><div class="row contributors" data-v-99f6eb26><div class="col" data-v-99f6eb26><div class="row contributor-type" data-v-792406ce data-v-99f6eb26> <div class="contributor-type__contributor" data-v-792406ce><a href="/core/search?filters%5BauthorTerms%5D=Chuan-Chao%20Wang&eventCode=SE-AU" class="app-link app-link__text app-link--accent" data-v-63dfaf6e data-v-792406ce><span data-v-63dfaf6e>Chuan-Chao Wang</span> </a>  <span data-v-792406ce> and</span></div><div class="contributor-type__contributor" data-v-792406ce><a href="/core/search?filters%5BauthorTerms%5D=Martine%20Robbeets&eventCode=SE-AU" class="app-link app-link__text app-link--accent" data-v-63dfaf6e data-v-792406ce><span data-v-63dfaf6e>Martine Robbeets</span> </a> <a target="_blank" href="https://orcid.org/0000-0002-2860-0230" data-test-orcid="Martine Robbeets" class="app-link contributor-type__contributor__orcid app-link__icon app-link--" data-v-63dfaf6e data-v-792406ce><img src="data:image/svg+xml;base64,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" alt="Open the ORCID record for Martine Robbeets" class="app-icon icon orcid" data-v-d2c09870 data-v-63dfaf6e> <span class="sr-only" data-v-63dfaf6e>[Opens in a new window]</span></a> <span data-v-792406ce></span></div> </div> </div> <div class="col-2 collapse-link" data-v-99f6eb26><a href="#authors-details" data-toggle="collapse" aria-expanded="false" aria-controls="authors-details" class="app-link collapsed app-link__text-icon app-link--secondary reverse" data-v-63dfaf6e data-v-99f6eb26><img src="data:image/svg+xml;base64,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" alt="" class="app-icon icon arrow-down" data-v-d2c09870 data-v-63dfaf6e><span class="text" data-v-63dfaf6e>Show author details </span> </a></div></div> <hr aria-hidden="true" class="separator default" data-v-7036083a data-v-99f6eb26> <dl id="authors-details" class="authors-details collapse" data-v-2edb8da6 data-v-99f6eb26><div data-test-author="Chuan-Chao Wang" class="row author" data-v-2edb8da6><dt class="col-12 col-sm-2 title" data-v-2edb8da6>Chuan-Chao Wang</dt> <dd class="col content d-inline d-sm-flex" data-v-2edb8da6><span class="content__title" data-v-2edb8da6>Affiliation:</span> <div class="d-sm-flex flex-column flex-sm-1 d-inline" data-v-2edb8da6><span data-v-2edb8da6><span data-v-2edb8da6>Department of Anthropology and Ethnology, Institute of Anthropology, National Institute for Data Science in Health and Medicine, and School of Life Sciences, Xiamen University, Xiamen361005, China</span> </span></div></dd></div><div data-test-author="Martine Robbeets" class="row author" data-v-2edb8da6><dt class="col-12 col-sm-2 title" data-v-2edb8da6>Martine Robbeets*</dt> <dd class="col content d-inline d-sm-flex" data-v-2edb8da6><span class="content__title" data-v-2edb8da6>Affiliation:</span> <div class="d-sm-flex flex-column flex-sm-1 d-inline" data-v-2edb8da6><span data-v-2edb8da6><span data-v-2edb8da6>Eurasia3angle Research Group, Max Planck Institute for the Science of Human History, Jena, Germany</span> </span></div></dd></div> <div class="row" data-v-2edb8da6><dt class="col-sm-2 col-12 title" data-v-2edb8da6> * </dt> <dd class="col content" data-v-2edb8da6><div class="row" data-v-2edb8da6><div class="d-sm-flex d-inline flex-sm-1 flex-sm-wrap" data-v-2edb8da6><div class="d-inline" data-v-2edb8da6><span data-v-2edb8da6><div class="corresp"><span class="label">*</span>Corresponding author. E-mail: <a href="mailto:robbeets@shh.mpg.de">robbeets@shh.mpg.de</a></div></span></div></div></div></dd></div> <hr aria-hidden="true" class="separator default" data-v-7036083a data-v-2edb8da6></dl></div></div> <div id="app-tabs" class="tabs" data-v-1d90c6ce data-v-01274b1d><div id="app-tabs-wrapper" class="tabs__wrapper" data-v-1d90c6ce><div role="navigation" aria-label="tab navigation" class="container" data-v-1d90c6ce><a data-toggle="collapse" href="#appTabs" role="button" aria-expanded="false" aria-controls="appTabs" class="tabs__collapse collapsed d-sm-none d-print-none" data-v-1d90c6ce><span data-v-1d90c6ce></span> <span class="tabs-arrow-up" data-v-1d90c6ce></span> <span class="tabs-arrow-down" data-v-1d90c6ce></span></a> <ul id="appTabs" role="tablist" class="nav nav-tabs tabs__list collapse show" data-v-1d90c6ce><li role="none" class="tabs__tab" data-v-1d90c6ce><a aria-selected="true" href="#article-tab" role="tab" aria-controls="article-tab" tabindex="-1" class="active" 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class="toc-arrow-up"></span> <span class="toc-arrow-down"></span></a> <ul id="toc-list-mobile" class="list collapse"><li class="list__item"><a href="#sec0" class="list__item__link"><span class="toc-title">Abstract</span></a></li> <li class="list__item"><a href="#sec1" class="list__item__link"> <span>Introduction</span></a></li><li class="list__item"><a href="#sec2" class="list__item__link"> <span>Methods</span></a></li><li class="list__item"><a href="#sec3" class="list__item__link"> <span>Results</span></a></li><li class="list__item"><a href="#sec4" class="list__item__link"> <span>Discussion</span></a></li>  <li class="list__item"><a href="#references-list" class="list__item__link"><span class="toc-title">References</span></a></li></ul></div> <div class="action-bar" data-v-43a4d572><div class="row items"> <div class="app-dropdown d-print-none" data-v-fab090b8 data-v-113567da data-v-43a4d572><button aria-expanded="false" data-test-id="buttonSavePDFOptions" id="save-pdf-dropdown" 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<hr aria-hidden="true" class="separator default" data-v-7036083a></div> <div class="share-modal-overlay" style="display:none;" data-v-43a4d572></div> <div data-spy="scroll" data-target="#toc" class="scrollspy-content" data-v-43a4d572> <div id="sec0" class="col article-abstract sec" data-v-2fa8b348 data-v-43a4d572>  <div class="abstract-graphical" data-v-2fa8b348><h2>Abstract</h2> <div class="abstract-graphical__content"><div class="abstract" data-abstract-type="graphical"><p><img class="graphic" orientation mimesubtype="png" mimetype="image" position="float" type href="S2513843X20000080_figAb.png" src="https://static.cambridge.org/content/id/urn%3Acambridge.org%3Aid%3Aarticle%3AS2513843X20000080/resource/name/S2513843X20000080_figAb.png?pub-status=live" /></p></div></div> <div class="abstract-graphical__text"><div class="abstract" data-abstract-type="normal"><p>The Tungusic languages form a language family spoken in Xinjiang, Siberia, Manchuria and the Russian Far East. There is a general consensus that these languages are genealogically related and descend from a common ancestral language, conventionally called ‘Proto-Tungusic’. However, the exact geographical location where the ancestral speakers of Proto-Tungusic originated from is subject to debate. Here we take an unprecedented approach to this problem, by integrating linguistic, archaeological and genetic evidence in a single study. Our analysis of ancient DNA suggests genetic continuity between an ancient Amur genetic lineage and the contemporary speakers of the Tungusic languages. Adding an archaeolinguistic perspective, we infer that the most plausible homeland for the speakers of Proto-Tungusic is the region around Lake Khanka in the Russian Far East. Our study pushes the field forward in answering the tantalizing question about the location of the Tungusic homeland and in illustrating how these three disciplines can converge into a holistic approach to the human past.</p></div></div> <hr aria-hidden="true" class="separator default" data-v-7036083a></div> </div>  <div class="keywords" data-v-86c27100 data-v-43a4d572><h2 data-v-86c27100>Keywords</h2> <div class="row keywords__pills" data-v-86c27100><a href="/core/search?filters[keywords]=archaeolinguistics" data-v-f0b31360 data-v-86c27100><span data-v-f0b31360 data-v-86c27100>archaeolinguistics</span></a><a href="/core/search?filters[keywords]=ancient DNA" data-v-f0b31360 data-v-86c27100><span data-v-f0b31360 data-v-86c27100>ancient DNA</span></a><a href="/core/search?filters[keywords]=Tungusic language family" data-v-f0b31360 data-v-86c27100><span data-v-f0b31360 data-v-86c27100>Tungusic language family</span></a><a href="/core/search?filters[keywords]=homeland" data-v-f0b31360 data-v-86c27100><span data-v-f0b31360 data-v-86c27100>homeland</span></a><a href="/core/search?filters[keywords]=Amur genome" data-v-f0b31360 data-v-86c27100><span data-v-f0b31360 data-v-86c27100>Amur genome</span></a></div> <hr aria-hidden="true" class="separator default" data-v-7036083a data-v-86c27100></div>  <dl class="article-details" data-v-6e32a161 data-v-43a4d572><div class="row" data-v-6e32a161><dt class="col-12 col-sm-3 col-md-2_5 title" data-v-6e32a161> Type </dt> <dd class="col content" data-v-6e32a161>Research Article</dd></div> <div class="row" data-v-6e32a161><dt class="col-12 col-sm-3 col-md-2_5 title" data-v-6e32a161> Information </dt> <dd class="col content" data-v-6e32a161><div class="content__journal" data-v-6e32a161><a href="/core/journals/evolutionary-human-sciences" class="app-link app-link__text app-link--underlined" data-v-63dfaf6e data-v-6e32a161><span class="text" data-v-63dfaf6e>Evolutionary Human Sciences </span> </a> <span data-v-6e32a161> , <a href="/core/journals/evolutionary-human-sciences/volume/F2345807349E632AB7170A2CDEAF9DF5" class="app-link app-link__text app-link--underlined" data-v-63dfaf6e data-v-6e32a161><span class="text" data-v-63dfaf6e>Volume 2 </span> </a></span>  <span data-v-6e32a161>, 2020</span>   <span data-v-6e32a161>, e8</span></div> <div class="doi-data" data-v-6e32a161><div data-v-6e32a161>DOI: <a target="_blank" href="https://doi.org/10.1017/ehs.2020.8" class="app-link app-link__text app-link--accent" data-v-63dfaf6e data-v-6e32a161><span class="text" data-v-63dfaf6e>https://doi.org/10.1017/ehs.2020.8 </span> <span class="sr-only" data-v-63dfaf6e>[Opens in a new window]</span></a></div> </div>  </dd></div> <div class="row creative-commons" data-v-6e32a161><dt class="col-12 col-sm-3 col-md-2_5 creative-commons__title"> Creative Commons </dt> <dd class="col"><div class="creative-commons__content"><div class="creative-commons__icons"><img src="/core/page-component/img/license-cc-icon.e3a74ed.svg" alt="Creative Common License - CC" class="app-icon creative-commons__icon license-cc" data-v-d2c09870><img src="/core/page-component/img/license-by-icon.33e212c.svg" alt="Creative Common License - BY" class="app-icon creative-commons__icon license-by" data-v-d2c09870></div> <div class="creative-commons__description">This is an Open Access article, distributed under the terms of the Creative Commons Attribution licence (<a href="http://creativecommons.org/licenses/by/4.0/" target="_blank">http://creativecommons.org/licenses/by/4.0/</a>), which permits unrestricted re-use, distribution, and reproduction in any medium, provided the original work is properly cited.</div></div></dd></div>  <div class="row" data-v-6e32a161><dt class="col-12 col-sm-3 col-md-2_5 title" data-v-6e32a161> Copyright </dt> <dd class="col content" data-v-6e32a161><div data-v-6e32a161> Copyright © The Author(s), 2020 </div></dd></div></dl>  <div id="content-container" class="content-container" data-v-43a4d572><div class="content-box"><div class="article research-article NLM"><div class="body"><p class="p"> <strong>Media summary:</strong> The Tungusic languages form a language family spoken in Xinjiang, Siberia, Manchuria and the Russian Far East. There is a general consensus that these languages descend from a common ancestral language, conventionally called ‘Proto-Tungusic’. However, the exact geographical location where the ancestral speakers of Proto-Tungusic originated from is subject to debate. Integrating linguistic, archaeological and genetic evidence in a single study, we here show that the most plausible homeland for the speakers of Proto-Tungusic is the region around Lake Khanka in the Russian Far East. Our study pushes the field forward in answering the tantalizing question about the location of the Tungusic homeland and in illustrating how these three disciplines can converge into a holistic approach on the human past.</p><div class="sec intro" data-magellan-destination="sec1" id="sec1"><h2 class="A"> Introduction</h2><p class="p"> Even if they are rapidly losing ground owing to Chinese and Russian influence, the speakers of Tungusic languages are widely distributed all over Northeast Asia, including Xinjiang, Siberia, Manchuria and the Russian Far East. Their wide distribution and the considerable internal variation of populations have made the exact geographical location where the ancestral speakers of Proto-Tungusic originated from subject to debate. Traditionally, this issue was approached through the combined investigation of languages and material remains, but our recently acquired ability to sequence DNA directly from ancient human remains has considerably expanded the available scientific tool kit. Therefore, here we intend to complement linguistic and archaeological observations with genetic evidence to infer the most plausible homeland for the speakers of Proto-Tungusic.</p><p class="p"> The contemporary and historically attested Tungusic languages are spread from the Okhotsk Sea in the east to the Yenisei Basin in the west, and from the Bohai Sea in the south to the Arctic Ocean in the north. <a class="xref fig" href="#fig01">Figure 1</a> shows the distribution of 12 Tungusic languages, Oroch, Udihe, Hezhe, Nanai, Orok, Olcha, Xibo, Even, Solon, Evenki, Negidal and Oroqen, along with dialectal varieties such as Kamnigan Evenki, Momsky Even, Olsky Even, Najkhin Nanai, Kur-Urmi Nanai and Bikin Nanai as well as two historical varieties, Jurchen and Manchu. Since written materials in Jurchen, the now-extinct language of the Jin dynasty (AD 1115–1234) are only partially deciphered, the earliest well-documented stage is Manchu, the official language of the Qing dynasty (AD 1644–1911). </p><section><div class="fig fig" data-magellan-destination="fig01" id="fig01"><div class="figure-thumb"><img src="data:image/gif;base64,R0lGODlhAQABAIAAAMLCwgAAACH5BAAAAAAALAAAAAABAAEAAAICRAEAOw==" data-src="https://static.cambridge.org/binary/version/id/urn:cambridge.org:id:binary:20200421175645641-0667:S2513843X20000080:S2513843X20000080_fig1.png?pub-status=live" class="aop-lazy-load-image" data-original-image="/binary/version/id/urn:cambridge.org:id:binary:20200421175645641-0667:S2513843X20000080:S2513843X20000080_fig1.png" data-zoomable="true"></div><div class="caption"><p class="p"> </p><p class="p"><span class="label">Figure 1.</span> (a) The distribution of the Tungusic languages; (b) proposed locations for the homeland of the ancestral speakers of Proto-Tungusic.</p></div></div></section><p class="p"> Although there is a general consensus that these languages are genealogically related and descend from a common ancestral language, conventionally called ‘Proto-Tungusic’, there are various competing hypotheses concerning the possible homeland of this language. Among the proposals, we find (a) the Baikal region (Vasilevich <a class="xref bibr" href="#ref36"><span class="show-for-sr">Reference Vasilevich</span>1960</a>; Menges <a class="xref bibr" href="#ref20"><span class="show-for-sr">Reference Menges</span>1968</a>; Derevyanko <a class="xref bibr" href="#ref4"><span class="show-for-sr">Reference Derevyanko</span>1976</a>; Helimski <a class="xref bibr" href="#ref8"><span class="show-for-sr">Reference Helimski and Ubriatova</span>1985</a>), (b) the Mid Amur and the lower part of the Upper Amur region (Janhunen <a class="xref bibr" href="#ref10"><span class="show-for-sr">Reference Janhunen</span>1996</a>: 169; Korovina <a class="xref bibr" href="#ref15"><span class="show-for-sr">Reference Korovina</span>2011</a>; Pevnov <a class="xref bibr" href="#ref25"><span class="show-for-sr">Reference Pevnov, Malchukov and Whaley</span>2012</a>; Wichmann, personal communication, 3 October 2019), (c) the Ussuri region around Lake Khanka, a tributary of the Amur (Robbeets <em class="italic">et al</em>. <a class="xref bibr" href="#ref30"><span class="show-for-sr">Reference Robbeets, Janhunen, Savelyev, Korovina, Robbeets and Savelyev</span>2020</a>) and (d) the Yalu River region on the border between present-day Liaoning and Northern Korea (Janhunen <a class="xref bibr" href="#ref11"><span class="show-for-sr">Reference Janhunen, Malchukov and Whaley</span>2012</a>). These locations are situated on the map in <a class="xref fig" href="#fig01">Figure 1</a>.</p><p class="p"> After assessing these four different hypotheses from an archaeolinguistic perspective, we will open a genetic window into this question. By supplementing our archaeolinguistic argumentation with genetic evidence for continuity between an ancient Amur genetic profile and contemporary speakers of Tungusic, we will conclude that the region around Lake Khanka constitutes the most plausible homeland for the speakers of Proto-Tungusic. Through this holistic approach, we hope to answer the tantalizing question about the Tungusic homeland, posed by numerous specialists in the past.</p></div><div class="sec methods" data-magellan-destination="sec2" id="sec2"><h2 class="A"> Methods</h2><div class="sec other" data-magellan-destination="sec2-1" id="sec2-1"><h3 class="B"> Archaeolinguistic methods</h3><p class="p"> Traditionally, there are two principal techniques for attempting to determine the location of the original homeland of the speakers of a certain proto-language: the diversity hotspot principle and cultural reconstruction. The ‘diversity hotspot principle’ is a loose principle that was originally introduced by Edward Sapir (<a class="xref bibr" href="#ref31"><span class="show-for-sr">Reference Sapir</span>1916</a>: 87) as the ‘center of gravity principle’, but it is also known as the ‘focus of diversity’ principle (Heggarty <a class="xref bibr" href="#ref7"><span class="show-for-sr">Reference Heggarty, Bowern and Evans</span>2015</a>: 612–613). Assuming that the deepest splits within a family reflect the greatest age, the location of these splits on the map is thought to point to the area where the proto-language began to diversify. The principle is thus based on the assumption that the homeland is closest to where one finds the greatest diversity with regard to the deepest subgroups of the language family. Attempts have been made to formalize this principle in a computer-automated approach, computing a diversity measure for each language and identifying the homeland with the location of the language with the highest diversity measure (Wichmann <em class="italic">et al</em>. <a class="xref bibr" href="#ref40"><span class="show-for-sr">Reference Wichmann, Müller and Velupillai</span>2010</a>). A schoolbook example is the Austronesian family, that extends across a huge geographical range, all the way from Madagascar to Easter Island, but the deepest subgroups are found on just one Island, Taiwan, which is therefore identified as the homeland.</p><p class="p"> Although the diversity hotspot principle can provide some clues about the homeland of a language family, it must also contend with certain limitations. First, the identification of the homeland depends on the location of the deepest subgroups and therefore on how robustly the internal structure of a given family has been established. A second limitation of this principle is that the contemporary hotspot of linguistic diversity may diverge from the earlier one. Looking at the present map of Indo-European with the Balkan Peninsula hosting the highest diversity of deep subgroups, we might conclude that the homeland is there, instead of the Pontic Steppe or Anatolia. This example makes it clear that the application of the diversity hotspot principle at more profound time-depths may be speculative because earlier diversity may have been erased through time. Third, the diversity hotspot principle might also be upset when a migration was suddenly directed over a long distance rather than representing slow, gradual and random movement into adjacent areas. Finally, linguistic diversity is a function not only of time but also of other factors such as environmental change and disease. These may have made the original homeland unsuitable for human habitation at a certain point in time. In this way, original linguistic diversity may have been erased and it may no longer be possible to pinpoint the homeland using the diversity hotspot principle. However, even if the principle is not foolproof, it offers valuable clues for the location of a homeland at less remote time-depths. In the case of Tungusic, there is only mild disagreement on the primary branching structure, while the time depth is agreed to be relatively shallow, i.e. between 600 BCE and AD 600. Except for the historical transfer of the Xibo tribe speaking Manchu in 1763 from Manchuria to Xinjiang, there are no known cases of sudden long-distance movement. Therefore, we can gain more from applying the diversity hotspot principle to Tungusic than we can lose from ignoring it.</p><p class="p"> The second technique, cultural reconstruction, looks at the cultural vocabulary revealed in the reconstructed vocabulary of a proto-language. The method was first introduced under the label ‘linguistic paleontology’ by Adolphe Pictet (<a class="xref bibr" href="#ref26"><span class="show-for-sr">Reference Pictet</span>1859</a>), but it became widespread under the banner of the ‘Wörter und Sachen’ movement, which used reconstructed proto-words as a way of determining proto-culture (Shuchardt <a class="xref bibr" href="#ref33"><span class="show-for-sr">Reference Shuchardt</span>1912</a>). Cultural reconstruction is a vocabulary-based approach that enables us to study human prehistory by reconstructing lexical items that are ecologically or culturally diagnostic for a particular region at a particular time. Clues for fauna and flora can help to delimit a homeland if several reconstructed plant and animal names converge in a particular region to the exclusion of other regions. Reconstruction of cultural items such as cultivars and feedstock can help to identify an archaeological culture yielding the relevant material remains. The method can be combined with diachronic contact linguistics, the study of the ways in which ancestral languages influenced each other when their speakers interacted. As shown by the assumed borrowing of the term ‘barley’ below, this approach can be useful to shed light on prehistoric ethnic interaction and help us determine the chronology of our data.</p><p class="p"> Among the limitations of cultural reconstruction are the potential lack of accuracy in semantic reconstruction, the occurrence of lexical recycling, the deception of a single item not backed up by a semantic domain and the shakiness of inferences made on the basis of absence. Since semantic reconstruction is less precise than phonological reconstruction, we should first be cautious that the meaning assigned to a reconstructed form can be no more specific than the meaning shared by all the cognate forms.</p><p class="p"> Second, we should be aware of the possibility of ‘lexical recycling’, a process whereby words with a general, non-cultural meaning become repurposed as words with a specific, cultural meaning after the importation or invention of the corresponding innovation. The reconstruction for ‘foxtail millet’ in Supplementary Table S1 (3), for instance, could also indicate the wild variety because the crop is native to the Russian Far East.</p><p class="p"> Third, we should avoid reconstructing a single cultural item that is not backed up by other members of the semantic domain. Paying attention to the clustering of different cultural reconstructions in a specific semantic domain, such as agriculture, can contribute to a fuller picture of prehistory than the study of individual cultural reconstructions.</p><p class="p"> And fourth, ‘absence of evidence is not evidence of absence’. The observation that an agricultural lexicon cannot be reconstructed for a certain proto-language may be explained by the fact that the proto-speakers simply were not familiar with farming, but it could also be due to the lack of exhaustive research or to the attrition of agricultural cognates over time. Therefore, inferences made on the basis of absence are not necessarily wrong, but they should not be taken as absolute proof for an argument.</p><p class="p"> Nevertheless, when combined with the necessary checks and balances, cultural reconstruction is reasonably robust as a way of determining proto-ecology and proto-culture through proto-language. Therefore, we will apply this method in order to make predictions about the cultural and natural environment available to the speakers of Proto-Tungusic.</p></div><div class="sec other" data-magellan-destination="sec2-2" id="sec2-2"><h3 class="B"> Genetic methods</h3><p class="p"> We reanalyzed the genome-wide data of ancient and present-day East Eurasian populations related to Tungusic-speaking populations from the following literature: Siska <em class="italic">et al</em>. (<a class="xref bibr" href="#ref34"><span class="show-for-sr">Reference Siska, Jones, Jeon, Bhak, Kim, Cho, Kim, Lee, Veselovskaya, Balueva, Gallego-Llorente, Hofreiter, Bradley, Eriksson, Pinhasi, Bhak and Manica</span>2017</a>), de Barros Damgaard <em class="italic">et al</em>. (<a class="xref bibr" href="#ref24"><span class="show-for-sr">Reference Damgaard, Marchi, Rasmussen, Peyrot, Renaud, Korneliussen, Moreno-Mayar, Pedersen, Goldberg, Usmanova, Baimukhanov, Loman, Hedeager, Pedersen, Nielsen, Afanasiev, Akmatov, Aldashev, Alpaslan, Baimbetov, Bazaliiskii, Beisenov, Boldbaatar, Boldgiv, Dorzhu, Ellingvag, Erdenebaatar, Dajani, Dmitriev, Evdokimov, Frei, Gromov, Goryachev, Hakonarson, Hegay, Khachatryan, Khaskhanov, Kitov, Kolbina, Kubatbek, Kukushkin, Kukushkin, Lau, Margaryan, Merkyte, Mertz, Mertz, Mijiddorj, Moiyesev, Mukhtarova, Nurmukhanbetov, Orozbekova, Panyushkina, Pieta, Smrčka, Shevnina, Logvin, Sjogren, Štolcova, Taravella, Tashbaeva, Tkachev, Tulegenov, Voyakin, Yepiskoposyan, Undrakhbold, Varfolomeev, Weber, Wilson Sayres, Kradin, Allentoft, Orlando, Nielsen, Sikora, Heyer, Kristiansen and Willerslev</span>2018</a>), Kanzawa-Kiriyama <em class="italic">et al</em>. (<a class="xref bibr" href="#ref13"><span class="show-for-sr">Reference Kanzawa-Kiriyama, Kryukov, Jinam, Hosomichi, Saso, Suwa, Ueda, Yoneda, Tajima, Shinoda, Inoue and Saitou</span>2016</a>), Patterson <em class="italic">et al</em>. (<a class="xref bibr" href="#ref23"><span class="show-for-sr">Reference Patterson, Moorjani, Luo, Mallick, Rohland, Zhan, Genschoreck, Webster and Reich</span>2012</a>), Jeong <em class="italic">et al</em>. (<a class="xref bibr" href="#ref12"><span class="show-for-sr">Reference Jeong, Balanovsky, Lukianova, Kahbatkyzy, Flegontov, Zaporozhchenko, Immel, Wang, Ixan, Khussainova, Bekmanov, Zaibert, Lavryashina, Pocheshkhova, Yusupov, Agdzhoyan, Koshel, Bukin, Nymadawa, Turdikulova, Dalimova, Churnosov, Skhalyakho, Daragan, Bogunov, Bogunova, Shtrunov, Dubova, Zhabagin, Yepiskoposyan, Churakov, Pislegin, Damba, Saroyants, Dibirova, Atramentova, Utevska, Idrisov, Kamenshchikova, Evseeva, Metspalu, Outram, Robbeets, Djansugurova, Balanovska, Schiffels, Haak, Reich and Krause</span>2019</a>) and Lazaridis <em class="italic">et al</em>. (<a class="xref bibr" href="#ref16"><span class="show-for-sr">Reference Lazaridis, Patterson, Mittnik, Renaud, Mallick, Kirsanow, Sudmant, Schraiber, Castellano, Lipson, Berger, Economou, Bollongino, Fu, Bos, Nordenfelt, Li, de Filippo, Prüfer, Sawyer, Posth, Haak, Hallgren, Fornander, Rohland, Delsate, Francken, Guinet, Wahl, Ayodo, Babiker, Bailliet, Balanovska, Balanovsky, Barrantes, Bedoya, Ben-Ami, Bene, Berrada, Bravi, Brisighelli, Busby, Cali, Churnosov, Cole, Corach, Damba, van Driem, Dryomov, Dugoujon, Fedorova, Gallego Romero, Gubina, Hammer, Henn, Hervig, Hodoglugil, Jha, Karachanak-Yankova, Khusainova, Khusnutdinova, Kittles, Kivisild, Klitz, Kučinskas, Kushniarevich, Laredj, Litvinov, Loukidis, Mahley, Melegh, Metspalu, Molina, Mountain, Näkkäläjärvi, Nesheva, Nyambo, Osipova, Parik, Platonov, Posukh, Romano, Rothhammer, Rudan, Ruizbakiev, Sahakyan, Sajantila, Salas, Starikovskaya, Tarekegn, Toncheva, Turdikulova, Uktveryte, Utevska, Vasquez, Villena, Voevoda, Winkler, Yepiskoposyan, Zalloua, Zemunik, Cooper, Capelli, Thomas, Ruiz-Linares, Tishkoff, Singh, Thangaraj, Villems, Comas, Sukernik, Metspalu, Meyer, Eichler, Burger, Slatkin, Pääbo, Kelso, Reich and Krause</span>2014</a>).</p><p class="p"> We first carried out principal components analysis in the smartpca program of EIGENSOFT (Patterson <em class="italic">et al</em>. <a class="xref bibr" href="#ref22"><span class="show-for-sr">Reference Patterson, Price and Reich</span>2006</a>), using default parameters and the lsqproject: YES and numoutlieriter: 0 options. We projected ancient Devil's Gate Ustlda and Jomon samples onto the variation of present-day East Eurasians from the published Human Origin Dataset over 591,642 SNPs. We carried out ADMIXTURE analysis after pruning for linkage disequilibrium in PLINK with parameters --indep-pairwise 200 25 0.4 Chang <em class="italic">et al.</em> (<a class="xref bibr" href="#ref2a"><span class="show-for-sr">Reference Chang, Chow, Tellier, Vattikuti, Purcell and Lee</span>2015</a>) which retained 332,959 SNPs. We ran ADMIXTURE (Alexander <em class="italic">et al</em>. <a class="xref bibr" href="#ref1"><span class="show-for-sr">Reference Alexander, Novembre and Lange</span>2009</a>) with default 5-fold cross-validation (--cv = 5), varying the number of ancestral populations between <em class="italic">K</em> = 2 and <em class="italic">K</em> = 12 in 100 bootstraps with different random seeds. We observed the lowest CV error at <em class="italic">K</em> = 8. The samples used in this analysis are present-day East Eurasians, and individuals from worldwide representative populations. We used outgroup <em class="italic">f</em> <sub>3</sub>-statistics of the form <em class="italic">f</em> <sub>3</sub> (Mbuti; X, Y) to test the genetic drift sharing between ancient Devil's Gate and present-day East Eurasians. Analysis of <em class="italic">f</em> <sub>3</sub>-statistics was carried out using ADMIXTOOLS with standard errors computed with a block jackknife (Patterson <em class="italic">et al</em>. <a class="xref bibr" href="#ref23"><span class="show-for-sr">Reference Patterson, Moorjani, Luo, Mallick, Rohland, Zhan, Genschoreck, Webster and Reich</span>2012</a>). We used <em class="italic">qpAdm</em> as implemented in ADMIXTOOLS to estimate West Eurasian mixture proportions in Even and Evenk with a set of outgroup populations. Weighted linkage disequilibrium (LD) decay was calculated using ALDER to infer admixture parameters including dates and mixture proportions (Loh <em class="italic">et al</em>. <a class="xref bibr" href="#ref19"><span class="show-for-sr">Reference Loh, Lipson, Patterson, Moorjani, Pickrell, Reich and Berger</span>2013</a>). We added Table S2 representing West Eurasian-related admixture proportions estimated using <em class="italic">qpAdm</em> and Table S3 listing the admixture time with West Eurasian-related groups estimated by ALDER in the Supplementary Information.</p></div></div><div class="sec results" data-magellan-destination="sec3" id="sec3"><h2 class="A"> Results</h2><div class="sec other" data-magellan-destination="sec3-1" id="sec3-1"><h3 class="B"> Archaeolinguistic results</h3><p class="p"> The principle that the geographical area of maximal primary diversity within a language family corresponds to the original homeland implies that the primary splits in the family are determinant for the location of the homeland. There is some disagreement on the internal structure of the Tungusic language family, mainly with respect to the separation of the Manchuric branch. As shown in <a class="xref fig" href="#fig02">Figure 2</a>, some authors proposed a north–south classification, in which the separation of Manchuric from the other Tungusic languages does not constitute the earliest split in the family (Cincius <a class="xref bibr" href="#ref3"><span class="show-for-sr">Reference Cincius</span>1949</a>; Benzing <a class="xref bibr" href="#ref2"><span class="show-for-sr">Reference Benzing</span>1955</a>; Kormušin <a class="xref bibr" href="#ref14"><span class="show-for-sr">Reference Kormušin</span>1998</a>; Georg <a class="xref bibr" href="#ref6"><span class="show-for-sr">Reference Georg, Naeher, Stary and Weiers</span>2004</a>; Janhunen <a class="xref bibr" href="#ref11"><span class="show-for-sr">Reference Janhunen, Malchukov and Whaley</span>2012</a>), while others proposed an early breakup between Manchuric and the rest of Tungusic (Sunik <a class="xref bibr" href="#ref35"><span class="show-for-sr">Reference Sunik</span>1959</a>; Vasilevich <a class="xref bibr" href="#ref36"><span class="show-for-sr">Reference Vasilevich</span>1960</a>; Doerfer <a class="xref bibr" href="#ref5"><span class="show-for-sr">Reference Doerfer and Weiers</span>1978</a>; Vovin <a class="xref bibr" href="#ref37"><span class="show-for-sr">Reference Vovin</span>1993</a>; Robbeets <a class="xref bibr" href="#ref27"><span class="show-for-sr">Reference Robbeets</span>2015</a>; Whaley and Oskolskaya <a class="xref bibr" href="#ref39"><span class="show-for-sr">Reference Whaley, Oskolskaya, Robbeets and Savelyev</span>2020</a>). Depending on which of both classifications of Tungusic we favor, the center of primary diversity – and thus also the inferred location of the homeland – will move on the map, as illustrated in <a class="xref fig" href="#fig02">Figure 2</a>. Whereas the north–south classification pushes the homeland to the north into the Mid Amur region (<a class="xref fig" href="#fig02">Figure 2a</a>), the Manchu–Tungusic classification pulls it more southwards, into the area around Lake Khanka (<a class="xref fig" href="#fig02">Figure 2b</a>). </p><section><div class="fig fig" data-magellan-destination="fig02" id="fig02"><div class="figure-thumb"><img src="data:image/gif;base64,R0lGODlhAQABAIAAAMLCwgAAACH5BAAAAAAALAAAAAABAAEAAAICRAEAOw==" data-src="https://static.cambridge.org/binary/version/id/urn:cambridge.org:id:binary:20200421175645641-0667:S2513843X20000080:S2513843X20000080_fig2.png?pub-status=live" class="aop-lazy-load-image" data-original-image="/binary/version/id/urn:cambridge.org:id:binary:20200421175645641-0667:S2513843X20000080:S2513843X20000080_fig2.png" data-zoomable="true"></div><div class="caption"><p class="p"> </p><p class="p"><span class="label">Figure 2.</span> Diversity hotspot of the Tungusic languages under the north–south classification in (a) as compared with the hotspot under the Manchu–Tungusic classification in (b).</p></div></div></section><p class="p"> Restricting himself to 11 Tungusic languages, notably Even, Evenki, Negidal, Oroqen, Naykhin Nanai, Oroch, Orok, Olcha, Udehe, Manchu and Xibo, Wichmann (personal communication) situates the Tungusic homeland around the location of Naykhin Nanai at latitude 49.28 and longitude 136.47. As more southern varieties such as Solon on the river Nonni in Inner Mongolia and Bikin Nanai in the southern part of the Primorye province as well as Hezhe in Heilongjiang and Jilin are lacking from this analysis, we may expect the real diversity hotspot to be more to the south than the inferred one. In sum, the diversity hotspot principle leaves room for both the Mid Amur region and the area around Lake Khanka but excludes the Baikal and the Yalu river region as plausible homelands.</p><p class="p"> The breakup of proto-Tungusic took place relatively recently. Different linguistic dating methods, such as lexicostatistics (Korovina <a class="xref bibr" href="#ref15"><span class="show-for-sr">Reference Korovina</span>2011</a>: 600 BCE), dating of ethnonym shift (Robbeets <a class="xref bibr" href="#ref27"><span class="show-for-sr">Reference Robbeets</span>2015</a>: 206 BC to AD 220), measurement of mutual intelligibility (Pevnov <a class="xref bibr" href="#ref25"><span class="show-for-sr">Reference Pevnov, Malchukov and Whaley</span>2012</a>: AD 1), Bayesian phylolinguistics (Oskolskaya <em class="italic">et al</em>. (<a class="xref bibr" href="#ref21"><span class="show-for-sr">Reference Oskolskaya, Robbeets and Koile</span>under review</a>): AD 200), cultural reconstruction (Janhunen <a class="xref bibr" href="#ref11"><span class="show-for-sr">Reference Janhunen, Malchukov and Whaley</span>2012</a>: 500 BC to AD 500) and the automated similarity judgment program (Holman <em class="italic">et al</em>. <a class="xref bibr" href="#ref9"><span class="show-for-sr">Reference Holman, Brown, Wichmann, Müller, Velupillai, Hammarström, Sauppe, Jung, Bakker, Brown, Belyaev, Urban, Mailhammer, List and Egorov</span>2011</a>: AD 681) yield different dates, but all dates range between 600 BC and AD 700, which supports a relatively shallow time-depth for the family.</p><p class="p"> Applying cultural reconstruction to the Tungusic languages, we find that Proto-Tungusic culture can be associated with the reconstruction of PTg *<em class="italic">sele</em> ‘iron’, reflected in corresponding forms in the contemporary languages all sharing the meaning ‘iron’; see (1) in Supplementary Table S1. The first uncontested findings of iron in the Russian Far East go back to the Krounovska culture (600 BC to AD 200), which marks the beginning of the Early Iron Age in the region. From there, iron spread northwards, to the northern Primorye and the Priamurye during the ensuing period of the Pol'tse and Ol'ga cultures (200 BC to AD 500), but there is no evidence for iron as north as the Upper Amur region around that time. Iron reached Liaoning and southwestern Jilin no later than the third century BCE. Broadly speaking iron begins to appear early first millennium BCE in the Baikal region. Thus, the reconstruction of iron is compatible with all locations, except the Mid and Upper Amur region.</p><p class="p"> As shown in the etymologies (2)–(6) in Supplementary Table S1, we can reconstruct agricultural vocabulary to Proto-Tungusic, such as *<em class="italic">pisi-ke</em> ‘broomcorn millet (<em class="italic">Panicum miliaceum</em>)’, *<em class="italic">jiya</em> ‘foxtail millet (<em class="italic">Setaria italica</em>)’, *<em class="italic">üse</em>- ~ <em class="italic">üsi</em>- ‘to plant’, *<em class="italic">üse</em> ~ <em class="italic">üsi</em> ‘seed, seedling’, *<em class="italic">üsin</em> ‘field for cultivation’ and *<em class="italic">tari</em>- ‘to sow, plant, cultivate’. The use of these words by the speakers of Proto-Tungusic implies that they were familiar with agriculture. In addition to this familiarity with agricultural concepts, there are a number of other reasons to assume that the speakers of Proto-Tungusic were cultivating plants. First, the derivation of the Proto-Tungusic word *<em class="italic">üse</em> ~ <em class="italic">üsi</em> ‘seed, seedling’ as a deverbal noun from the verb *<em class="italic">üse</em>- ~ <em class="italic">üsi</em>- ‘to plant’ suggests that seeds were not just collected for consumption but that they were planted as part of a cultivation process.</p><p class="p"> Second, for some crop names such as ‘barley’ and ‘broomcorn millet’, we can argue that the word refers to the domesticated crop rather than to the wild variety of the plant because both crops are not native to the region and have been imported as domesticated crops. It is commonly assumed that broomcorn millet has been imported from the West Liao River region by the people who introduced the Zaisanovska culture (3200–1300 BCE) to the area around Lake Khanka (Sergusheva and Vostretsov <a class="xref bibr" href="#ref32"><span class="show-for-sr">Reference Sergusheva, Vostretsov, Fairbairn and Weiss</span>2009</a>; Leipe <em class="italic">et al</em>. <a class="xref bibr" href="#ref17"><span class="show-for-sr">Reference Leipe, Long, Sergusheva, Wagner and Tarasov</span>2019</a>; Li <em class="italic">et al</em>. <a class="xref bibr" href="#ref18"><span class="show-for-sr">Reference Li, Ning, Zhushchikhovskaya, Hudson and Robbeets</span>2020</a>). That the origins of the agricultural vocabulary in Proto-Tungusic go back as early as the fourth millennium BC in the southern part of the Primorye is further supported by the external etymologies of some words. Since some of the terms such as *<em class="italic">tari</em>- ‘to sow, to plant’ have cognates in Turkic and Mongolic; these agricultural terms must have been present in proto-Tungusic since its separation from proto-Altaic, estimated at 3300 BC (Robbeets and Bouckaert <a class="xref bibr" href="#ref29"><span class="show-for-sr">Reference Robbeets and Bouckaert</span>2018</a>; Robbeets <em class="italic">et al</em>. <a class="xref bibr" href="#ref30"><span class="show-for-sr">Reference Robbeets, Janhunen, Savelyev, Korovina, Robbeets and Savelyev</span>2020</a>). Only in the Zaisanovska culture around Lake Khanka and in the Wangjiacun site in Dalian on the North Korean border does agriculture go back that deep in time. Thus, the presence of agricultural vocabulary excludes the Baikal region and the Mid and Upper Amur region as plausible homelands.</p><p class="p"> The rather large time difference between the separation of Proto-Tungusic from Proto-Altaic around 3300 BC and the actual dispersal of Tungusic languages after 600 BC does not imply that proto-Tungusic continued as a single language without any diversification during three millennia. On the contrary, it is safe to assume that the contemporary Tungusic languages represent but a tiny proportion of the original diversity and that in the first millennia BCE various sister languages of Proto-Tungusic were spoken in the area. The adoption of agriculture involved a previously unparallelled demographic growth in certain production areas in North East China. With its relatively mild climate, the protected region of Lake Khanka constituted such an area, close to the northern edge of the climate range for East Asian Neolithic agriculture. Demographically, this implies a large population, while linguistically, it means that speech communities which happened to be in the center of agricultural innovation underwent rapid growth and were able to absorb neighboring speech communities by way of language shift. In this way, certain Proto-Tungusic speech communities in the region became highly expansive at the expense of others. This expansion must have affected a large number of individual languages belonging to various other language families, such as the Amuric family that was reduced to only one surviving language, Nivkh, but it may also have affected now-extinct sister branches of Proto-Tungusic. From the number of surviving contemporary languages, it is clear that the Transeurasian languages in general have a relatively high extinction rate. By way of comparison, Austronesian has 1500 surviving languages, whereas hardly 50 Transeurasian languages have survived to the present. This supports the assumption that only a happy few expansive languages survived in Northeast Asia, at the expense of numerous others.</p><p class="p"> A location around Lake Khanka for the Proto-Tungusic speech community is further corroborated by contact linguistics. The term for ‘barley and similar crops’, PTg *<em class="italic">murgi</em>, is probably borrowed from an Old Chinese donor word, 來*<em class="italic">mə.rˤək</em> > *<em class="italic">mə.rˤə</em>, ‘a kind of wheat’ (Robbeets <a class="xref bibr" href="#ref28"><span class="show-for-sr">Reference Robbeets, Robbeets and Savelyev</span>2017</a>: 28–29). The linguistic reconstruction can be correlated to the archaeological evidence for barley being first imported through Chinese contact at the time of the Krounovska culture situated in the Southern Primorye south of Lake Khanka (Sergusheva and Vostretsov <a class="xref bibr" href="#ref32"><span class="show-for-sr">Reference Sergusheva, Vostretsov, Fairbairn and Weiss</span>2009</a>: 214–215; Leipe <em class="italic">et al</em>. <a class="xref bibr" href="#ref17"><span class="show-for-sr">Reference Leipe, Long, Sergusheva, Wagner and Tarasov</span>2019</a>: 12).</p><p class="p"> Proceeding to the reconstruction of ecological vocabulary, we can reconstruct tree names to Proto-Tungusic, such as *<em class="italic">inʹŋamu-kta</em> ‘cloudberry (<em class="italic">Rubus chamaemorus</em>)’, *<em class="italic">xiba-gda</em> ‘ash tree (<em class="italic">Fráxinus mandshurica</em>)’, *<em class="italic">xami-gda</em> ‘poplar (<em class="italic">Populus maximowiczii</em>)’, *<em class="italic">kilden</em> ‘linden (<em class="italic">Tilia amurensis</em>)’ and *<em class="italic">molon</em> ‘maple (<em class="italic">Acer ukurunduense</em>)’, as shown in Supplementary Table S1 (7)–(11). Except for ‘cloudberry’, none of these species appears in the Baikal region, while maple does not appear in the Mid and Upper Amur region. Among the Proto-Tungusic tree names, the terms for ‘pine tree’ in Supplementary Table S1 (12)–(14) are particularly telling. In contrast to the wide distribution of the ‘creeping pine’, that of the ‘Korean pine (<em class="italic">Pínus koraiénsis</em>)’ is limited to the area around Lake Khanka. In addition, it is the only plant for which we have archaeobotanical evidence. The nutshells of this pine were found in pit-dwelling deposits in the Southern Primorye, from 1300 BC onwards (Sergusheva and Vostretsov <a class="xref bibr" href="#ref32"><span class="show-for-sr">Reference Sergusheva, Vostretsov, Fairbairn and Weiss</span>2009</a>: 213). Therefore the reconstruction of the word for Korean pine strongly supports a homeland around Lake Khanka.</p><p class="p"> <a class="xref table" href="#tab01">Table 1</a> assesses the different homeland proposals from an archaeolinguistic viewpoint. The hypothesis of Lake Khanka is best supported by the linguistic data, while the Baikal hypothesis is the least compatible. Situating the Proto-Tungusic homeland in the Lake Khanka area is further supported by the continuity of political formations in the region such as the Mohe (AD 500–1000) in the area around Lake Khanka, the Bohai state (AD 698–925) southwards from lake Khanka into present-day North Korea and the Jin Empire of the Jurchen (AD 1115–1234) to the west of Lake Khanka in the Sungari, Nonni and Middle Amur region. Since Neolithic times, the cultural edge in the Russian Far East was in the south, not in the north. Therefore, the area around Lake Khanka is the most plausible homeland for the ancestral Tungusic speech community. </p><div class="table-wrap" data-magellan-destination="tab01" id="tab01"><div class="caption"><p class="p"><span class="label">Table 1.</span> Evaluating the four homeland proposals for Proto-Tungusic using different archaeolinguistic techniques: 0 implausible, 1 plausible</p></div><span><div class="figure-thumb"><img src="data:image/gif;base64,R0lGODlhAQABAIAAAMLCwgAAACH5BAAAAAAALAAAAAABAAEAAAICRAEAOw==" data-src="https://static.cambridge.org/binary/version/id/urn:cambridge.org:id:binary:20200421175645641-0667:S2513843X20000080:S2513843X20000080_tab1.png?pub-status=live" class="aop-lazy-load-image" data-original-image="/binary/version/id/urn:cambridge.org:id:binary:20200421175645641-0667:S2513843X20000080:S2513843X20000080_tab1.png" data-zoomable="false"></div></span></div></div><div class="sec other" data-magellan-destination="sec3-2" id="sec3-2"><h3 class="B"> Genetic results</h3><p class="p"> From a paternal perspective, Tungusic speakers are associated with the Y chromosomal haplogroup C3-M217. The haplogroup C3c-M48 is prevalent in Evenki and Even, as well as in other Tungusic-speaking populations in the Amur River Basin, including Oroqen, Olcha, Negidal, Udehe and Nanai with frequencies ranging from about 20% to even 100%. The C3-M217 type also reaches high frequencies in populations surrounding the Amur River Basin, such as Nivkh (38%) and Ainu (12.5–25%) (Wei <a class="xref bibr" href="#ref38"><span class="show-for-sr">Reference Wei</span>2011</a>). The current geographic and genetic diversity distribution pattern suggests that the dispersal of Y chromosomal haplogroup C3-M217 was most likely associated with Tungusic-related population expansion from the Amur River.</p><p class="p"> From a genome-wide perspective, we reanalyzed the data of ancient samples from Devil's Gate (Chertovy Vorota) Cave dating to ~7.7 kya in the Primorye Region of the Russian Far East (Siska <em class="italic">et al</em>. <a class="xref bibr" href="#ref34"><span class="show-for-sr">Reference Siska, Jones, Jeon, Bhak, Kim, Cho, Kim, Lee, Veselovskaya, Balueva, Gallego-Llorente, Hofreiter, Bradley, Eriksson, Pinhasi, Bhak and Manica</span>2017</a>; de Barros Damgaard <em class="italic">et al</em>. <a class="xref bibr" href="#ref24"><span class="show-for-sr">Reference Damgaard, Marchi, Rasmussen, Peyrot, Renaud, Korneliussen, Moreno-Mayar, Pedersen, Goldberg, Usmanova, Baimukhanov, Loman, Hedeager, Pedersen, Nielsen, Afanasiev, Akmatov, Aldashev, Alpaslan, Baimbetov, Bazaliiskii, Beisenov, Boldbaatar, Boldgiv, Dorzhu, Ellingvag, Erdenebaatar, Dajani, Dmitriev, Evdokimov, Frei, Gromov, Goryachev, Hakonarson, Hegay, Khachatryan, Khaskhanov, Kitov, Kolbina, Kubatbek, Kukushkin, Kukushkin, Lau, Margaryan, Merkyte, Mertz, Mertz, Mijiddorj, Moiyesev, Mukhtarova, Nurmukhanbetov, Orozbekova, Panyushkina, Pieta, Smrčka, Shevnina, Logvin, Sjogren, Štolcova, Taravella, Tashbaeva, Tkachev, Tulegenov, Voyakin, Yepiskoposyan, Undrakhbold, Varfolomeev, Weber, Wilson Sayres, Kradin, Allentoft, Orlando, Nielsen, Sikora, Heyer, Kristiansen and Willerslev</span>2018</a>). We observe that ancient Devil's Gate samples cluster together with present-day Tungusic-speaking populations and Ulchi in the outgroup <em class="italic">f</em> <sub>3</sub>-statistics, as shown in <a class="xref fig" href="#fig03">Figure 3</a>. We find a genetic substructure within Tungusic-speaking populations. The North Tungusic Evenki and Even cluster with surrounding Siberian populations other than with Amur Tungusic groups. The substructure has been caused by the fact that North Tungusic Evenki and Even have received gene flow from West Eurasian-related populations. </p><section><div class="fig fig" data-magellan-destination="fig03" id="fig03"><div class="figure-thumb"><img src="data:image/gif;base64,R0lGODlhAQABAIAAAMLCwgAAACH5BAAAAAAALAAAAAABAAEAAAICRAEAOw==" data-src="https://static.cambridge.org/binary/version/id/urn:cambridge.org:id:binary:20200421175645641-0667:S2513843X20000080:S2513843X20000080_fig3.png?pub-status=live" class="aop-lazy-load-image" data-original-image="/binary/version/id/urn:cambridge.org:id:binary:20200421175645641-0667:S2513843X20000080:S2513843X20000080_fig3.png" data-zoomable="true"></div><div class="caption"><p class="p"> </p><p class="p"><span class="label">Figure 3.</span> Outgroup <em class="italic">f</em> <sub>3</sub>-statistics of the form <em class="italic">f</em> <sub>3</sub> (Mbuti; X, Y) to test the genetic drift sharing between ancient Devil's Gate and present-day East Eurasians.</p></div></div></section><p class="p"> Our principal component analysis in <a class="xref fig" href="#fig04">Figure 4</a> shows that the population structure in East Asia is strongly correlated with geographical and linguistic categories. The ancient Devil's Gate genomes are shown to be genetically most similar to contemporary Tungusic populations, such as Ulchi, Negidal and Nanai at the lower Amur River basin, as well as Nivkh people on the nearby Sakhalin island, documenting a continuous presence of this type of ancestry in the Amur River Basin stretching back at least 8000 years. The principal component analysis plot in <a class="xref fig" href="#fig04">Figure 4</a> shows that Evenks and Evens are shifted towards West Eurasians and they have about 14–35% West Eurasian-related ancestry estimated by <em class="italic">qpAdm</em> method (Supplementary Table S2). The admixture time estimated by LD decay suggests a very recent time of around four to six generations ago (Supplementary Table S3). Late Neolithic and Early Bronze Age ancient samples from Ustlda site near the Lake Baikal region show a similar genetic profile with Even people deriving a large amount of Devil's Gate-related ancestry and also having admixture from West Eurasians. </p><section><div class="fig fig" data-magellan-destination="fig04" id="fig04"><div class="figure-thumb"><img src="data:image/gif;base64,R0lGODlhAQABAIAAAMLCwgAAACH5BAAAAAAALAAAAAABAAEAAAICRAEAOw==" data-src="https://static.cambridge.org/binary/version/id/urn:cambridge.org:id:binary:20200421175645641-0667:S2513843X20000080:S2513843X20000080_fig4.png?pub-status=live" class="aop-lazy-load-image" data-original-image="/binary/version/id/urn:cambridge.org:id:binary:20200421175645641-0667:S2513843X20000080:S2513843X20000080_fig4.png" data-zoomable="true"></div><div class="caption"><p class="p"> </p><p class="p"><span class="label">Figure 4.</span> Principal component analysis projecting ancient Devil's Gate, Ustlda and Jomon samples onto the variation of present-day East Eurasians.</p></div></div></section><p class="p"> The ADMIXTURE analysis in <a class="xref fig" href="#fig05">Figure 5</a> also confirms the above observation. We observed a large scale of shared components from ancient Devil's Gate samples to present-day Tungusic-speaking populations. We have not found evidence of any West Eurasian-related admixture in ancient Devil's Gate samples. In contrast, the Baikal Ustlda hunter-gatherers (~4700 years ago) were reported to largely have a Devil's Gate-related component, but also have West Eurasian-related ancestry as is shown in the ADMIXTURE plot. </p><section><div class="fig fig" data-magellan-destination="fig05" id="fig05"><div class="figure-thumb"><img src="data:image/gif;base64,R0lGODlhAQABAIAAAMLCwgAAACH5BAAAAAAALAAAAAABAAEAAAICRAEAOw==" data-src="https://static.cambridge.org/binary/version/id/urn:cambridge.org:id:binary:20200421175645641-0667:S2513843X20000080:S2513843X20000080_fig5.png?pub-status=live" class="aop-lazy-load-image" data-original-image="/binary/version/id/urn:cambridge.org:id:binary:20200421175645641-0667:S2513843X20000080:S2513843X20000080_fig5.png" data-zoomable="true"></div><div class="caption"><p class="p"> </p><p class="p"><span class="label">Figure 5.</span> Admixture plot from <em class="italic">K</em> = 2 to <em class="italic">K</em> = 12.</p></div></div></section><p class="p"> Therefore, our genetic findings suggest that the Tungusic-related Amur ancestry distributed in a vast region from the Russian Far East to Lake Baikal. The ancient Devil's Gate population seems to be an unadmixed form of this type of ancestry, which tends to support the larger Amur River region as the ancestral homeland for present-day Tungusic-speaking populations.</p></div></div><div class="sec discussion" data-magellan-destination="sec4" id="sec4"><h2 class="A"> Discussion</h2><p class="p"> Aligning the evidence, we find that our linguistic and genetic results converge in outlining the broader Amur River region as the ancestral homeland of the speakers of Proto-Tungusic, but our archaeolinguistic results more specifically point to the area around Lake Khanka in the southern part of the Primorye. The Amur-like genetic profile displays a deep history, going back at least 8000 years and covering an extensive region from Baikal and Liao River in the west to the Sea of Japan in the East. Our archaeolinguistic results suggest a dispersal route for agriculture and language from the West Liao River Region to the Southern Primorye around 3300 BCE. As the proposed farming/language dispersal falls within the geographical and chronological reach of the Amur-like genetic profile, we would not expect to observe traces of genetic admixture even if population movement was involved, because the admixing of two similar Amur-like genomes would not be perceivable. This explains why in the southern Primorye both Tungusic-speaking populations and non-Tungusic speaking populations such as the Nivkh are genetically similar and continuous with the ancient Devil's Gate genome in the region. Bringing the archaeolinguistic and genetic evidence together thus makes a case for imperceptible genetic admixture and indicates that the area around Lake Khanka in the southern part of the Primorye is the most plausible homeland for the ancestral speakers of Tungusic.</p></div></div><div class="back"><div class="notes supplementary-material" data-magellan-destination="nts1" id="nts1"><h2 class="A">Supplementary material</h2><p class="p"> The supplementary material for this article can be found at <a class="uri" href="https://doi.org/10.1017/ehs.2020.8">https://doi.org/10.1017/ehs.2020.8</a>.</p></div> <div class="notes other" data-magellan-destination="nts2" id="nts2"><h2 class="A">Author contributions</h2><p class="p"> MR and CCW conceived and designed the study. MR and CCW conducted data gathering. MR performed archaeolinguistic and CCW genetic analyses. MR and CCW wrote the article.</p></div> <div class="notes other" data-magellan-destination="nts3" id="nts3"><h2 class="A">Financial support</h2><p class="p"> The research leading to these results has received funding from the European Research Council (ERC) under the European Union's Horizon 2020 research and innovation program (grant agreement no. 646612) granted to Martine Robbeets. Chuan-Chao Wang was supported by the National Natural Science Foundation of China (31801040), Nanqiang Outstanding Young Talents Program of Xiamen University (X2123302) and Fundamental Research Funds for the Central Universities (ZK1144).</p></div> <div class="notes other" data-magellan-destination="nts4" id="nts4"><h2 class="A">Data availibility statement</h2><p class="p"> The linguistic data underlying this article are made available in the Supplementary Material (1). 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<div class="circle-list__item__grouped"><div id="reference-18-content" class="circle-list__item__grouped__content"><span class="string-name"><span class="surname">Lazaridis</span>, <span class="given-names">I</span></span>, <span class="string-name"><span class="surname">Patterson</span>, <span class="given-names">N</span></span>, <span class="string-name"><span class="surname">Mittnik</span>, <span class="given-names">A</span></span>, <span class="string-name"><span class="surname">Renaud</span>, <span class="given-names">G</span></span>, <span class="string-name"><span class="surname">Mallick</span>, <span class="given-names">S</span></span>, <span class="string-name"><span class="surname">Kirsanow</span>, <span class="given-names">K</span></span>, <span class="string-name"><span class="surname">Sudmant</span>, <span class="given-names">PH</span></span>, <span class="string-name"><span class="surname">Schraiber</span>, <span class="given-names">JG</span></span>, <span class="string-name"><span class="surname">Castellano</span>, <span class="given-names">S</span></span>, <span class="string-name"><span class="surname">Lipson</span>, <span class="given-names">M</span></span>, <span class="string-name"><span class="surname">Berger</span>, <span class="given-names">B</span></span>, <span class="string-name"><span class="surname">Economou</span>, <span class="given-names">C</span></span>, <span class="string-name"><span class="surname">Bollongino</span>, <span class="given-names">R</span></span>, <span class="string-name"><span class="surname">Fu</span>, <span class="given-names">Q</span></span>, <span class="string-name"><span class="surname">Bos</span>, <span class="given-names">KI</span></span>, <span class="string-name"><span class="surname">Nordenfelt</span>, <span class="given-names">S</span></span>, <span class="string-name"><span class="surname">Li</span>, <span class="given-names">H</span></span>, <span class="string-name"><span class="surname">de Filippo</span>, <span class="given-names">C</span></span>, <span class="string-name"><span class="surname">Prüfer</span>, <span class="given-names">K</span></span>, <span class="string-name"><span class="surname">Sawyer</span>, <span class="given-names">S</span></span>, <span class="string-name"><span class="surname">Posth</span>, <span class="given-names">C</span></span>, <span class="string-name"><span class="surname">Haak</span>, <span class="given-names">W</span></span>, <span class="string-name"><span class="surname">Hallgren</span>, <span class="given-names">F</span></span>, <span class="string-name"><span class="surname">Fornander</span>, <span class="given-names">E</span></span>, <span class="string-name"><span class="surname">Rohland</span>, <span class="given-names">N</span></span>, <span class="string-name"><span class="surname">Delsate</span>, <span class="given-names">D</span></span>, <span class="string-name"><span class="surname">Francken</span>, <span class="given-names">M</span></span>, <span class="string-name"><span class="surname">Guinet</span>, <span class="given-names">JM</span></span>, <span class="string-name"><span class="surname">Wahl</span>, <span class="given-names">J</span></span>, <span class="string-name"><span class="surname">Ayodo</span>, <span class="given-names">G</span></span>, <span class="string-name"><span class="surname">Babiker</span>, <span class="given-names">HA</span></span>, <span class="string-name"><span class="surname">Bailliet</span>, <span class="given-names">G</span></span>, <span class="string-name"><span class="surname">Balanovska</span>, <span class="given-names">E</span></span>, <span class="string-name"><span class="surname">Balanovsky</span>, <span class="given-names">O</span></span>, <span class="string-name"><span class="surname">Barrantes</span>, <span class="given-names">R</span></span>, <span class="string-name"><span class="surname">Bedoya</span>, <span class="given-names">G</span></span>, <span class="string-name"><span class="surname">Ben-Ami</span>, <span class="given-names">H</span></span>, <span class="string-name"><span class="surname">Bene</span>, <span class="given-names">J</span></span>, <span class="string-name"><span class="surname">Berrada</span>, <span class="given-names">F</span></span>, <span class="string-name"><span class="surname">Bravi</span>, <span class="given-names">CM</span></span>, <span class="string-name"><span class="surname">Brisighelli</span>, <span class="given-names">F</span></span>, <span class="string-name"><span class="surname">Busby</span>, <span class="given-names">GB</span></span>, <span class="string-name"><span class="surname">Cali</span>, <span class="given-names">F</span></span>, <span class="string-name"><span class="surname">Churnosov</span>, <span class="given-names">M</span></span>, <span class="string-name"><span class="surname">Cole</span>, <span class="given-names">DE</span></span>, <span class="string-name"><span class="surname">Corach</span>, <span class="given-names">D</span></span>, <span class="string-name"><span class="surname">Damba</span>, <span class="given-names">L</span></span>, <span class="string-name"><span class="surname">van Driem</span>, <span class="given-names">G</span></span>, <span class="string-name"><span class="surname">Dryomov</span>, <span class="given-names">S</span></span>, <span class="string-name"><span class="surname">Dugoujon</span>, <span class="given-names">JM</span></span>, <span class="string-name"><span class="surname">Fedorova</span>, <span class="given-names">SA</span></span>, <span class="string-name"><span class="surname">Gallego Romero</span>, <span class="given-names">I</span></span>, <span class="string-name"><span class="surname">Gubina</span>, <span class="given-names">M</span></span>, <span class="string-name"><span class="surname">Hammer</span>, <span class="given-names">M</span></span>, <span class="string-name"><span class="surname">Henn</span>, <span class="given-names">BM</span></span>, <span class="string-name"><span class="surname">Hervig</span>, <span class="given-names">T</span></span>, <span class="string-name"><span class="surname">Hodoglugil</span>, <span class="given-names">U</span></span>, <span class="string-name"><span class="surname">Jha</span>, <span class="given-names">AR</span></span>, <span class="string-name"><span class="surname">Karachanak-Yankova</span>, <span class="given-names">S</span></span>, <span class="string-name"><span class="surname">Khusainova</span>, <span class="given-names">R</span></span>, <span class="string-name"><span class="surname">Khusnutdinova</span>, <span class="given-names">E</span></span>, <span class="string-name"><span class="surname">Kittles</span>, <span class="given-names">R</span></span>, <span class="string-name"><span class="surname">Kivisild</span>, <span class="given-names">T</span></span>, <span class="string-name"><span class="surname">Klitz</span>, <span class="given-names">W</span></span>, <span class="string-name"><span class="surname">Kučinskas</span>, <span class="given-names">V</span></span>, <span class="string-name"><span class="surname">Kushniarevich</span>, <span class="given-names">A</span></span>, <span class="string-name"><span class="surname">Laredj</span>, <span class="given-names">L</span></span>, <span class="string-name"><span class="surname">Litvinov</span>, <span class="given-names">S</span></span>, <span class="string-name"><span class="surname">Loukidis</span>, <span class="given-names">T</span></span>, <span class="string-name"><span class="surname">Mahley</span>, <span class="given-names">RW</span></span>, <span class="string-name"><span class="surname">Melegh</span>, <span class="given-names">B</span></span>, <span class="string-name"><span class="surname">Metspalu</span>, <span class="given-names">E</span></span>, <span class="string-name"><span class="surname">Molina</span>, <span class="given-names">J</span></span>, <span class="string-name"><span class="surname">Mountain</span>, <span class="given-names">J</span></span>, <span class="string-name"><span class="surname">Näkkäläjärvi</span>, <span class="given-names">K</span></span>, <span class="string-name"><span class="surname">Nesheva</span>, <span class="given-names">D</span></span>, <span class="string-name"><span class="surname">Nyambo</span>, <span class="given-names">T</span></span>, <span class="string-name"><span class="surname">Osipova</span>, <span class="given-names">L</span></span>, <span class="string-name"><span class="surname">Parik</span>, <span class="given-names">J</span></span>, <span class="string-name"><span class="surname">Platonov</span>, <span class="given-names">F</span></span>, <span class="string-name"><span class="surname">Posukh</span>, <span class="given-names">O</span></span>, <span class="string-name"><span class="surname">Romano</span>, <span class="given-names">V</span></span>, <span class="string-name"><span class="surname">Rothhammer</span>, <span class="given-names">F</span></span>, <span class="string-name"><span class="surname">Rudan</span>, <span class="given-names">I</span></span>, <span class="string-name"><span class="surname">Ruizbakiev</span>, <span class="given-names">R</span></span>, <span class="string-name"><span class="surname">Sahakyan</span>, <span class="given-names">H</span></span>, <span class="string-name"><span class="surname">Sajantila</span>, <span class="given-names">A</span></span>, <span class="string-name"><span class="surname">Salas</span>, <span class="given-names">A</span></span>, <span class="string-name"><span class="surname">Starikovskaya</span>, <span class="given-names">EB</span></span>, <span class="string-name"><span class="surname">Tarekegn</span>, <span class="given-names">A</span></span>, <span class="string-name"><span class="surname">Toncheva</span>, <span 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A quantitative approach</span>. <span class="source">Diachronica</span> <span class="volume">27</span>(<span class="issue">2</span>), <span class="fpage">247</span>–<span class="lpage">276</span>. doi <span class="pub-id doi">10.1075/dia.27.2.05wic</span><a class='ref-link' target='_blank' aria-label='Google Scholar link for Homelands of the world's language families. A quantitative approach' href=https://scholar.google.com/scholar_lookup?title=Homelands+of+the+world's+language+families.+A+quantitative+approach&author=Wichmann+S&author=M%C3%BCller+A&author=Velupillai+V&publication+year=2010&journal=Diachronica&volume=27&pages=247-276>Google Scholar</a></div></div></div></div></div> </div></div>  <div id="figures-tab" class="figures tab-pane" data-v-241a4b23><div data-v-241a4b23><div class="figures__item" data-v-241a4b23><div class="figures__item__image-box" data-v-241a4b23><button class="figures__ref" data-v-241a4b23> View in content </button> <img src="data:image/gif;base64,R0lGODlhAQABAIAAAMLCwgAAACH5BAAAAAAALAAAAAABAAEAAAICRAEAOw==" alt="Figure 0" data-zoomable="true" data-src="https://static.cambridge.org/binary/version/id/urn:cambridge.org:id:binary:20200421175645641-0667:S2513843X20000080:S2513843X20000080_fig1.png" data-enlarged-image="https://static.cambridge.org/binary/version/id/urn:cambridge.org:id:binary:20200421175645641-0667:S2513843X20000080:S2513843X20000080_fig1.png" class="graphic" data-v-241a4b23></div> <div data-v-241a4b23><div class="caption" data-v-241a4b23><p data-v-241a4b23><span class="label" data-v-241a4b23>Figure 1.</span> <span data-v-241a4b23><span class="p">(a) The distribution of the Tungusic languages; (b) proposed locations for the homeland of the ancestral speakers of Proto-Tungusic.</span></span></p></div> </div></div> <hr aria-hidden="true" class="separator dashed" data-v-7036083a data-v-241a4b23></div><div data-v-241a4b23><div class="figures__item" data-v-241a4b23><div class="figures__item__image-box" data-v-241a4b23><button class="figures__ref" data-v-241a4b23> View in content </button> <img src="data:image/gif;base64,R0lGODlhAQABAIAAAMLCwgAAACH5BAAAAAAALAAAAAABAAEAAAICRAEAOw==" alt="Figure 1" data-zoomable="true" data-src="https://static.cambridge.org/binary/version/id/urn:cambridge.org:id:binary:20200421175645641-0667:S2513843X20000080:S2513843X20000080_fig2.png" data-enlarged-image="https://static.cambridge.org/binary/version/id/urn:cambridge.org:id:binary:20200421175645641-0667:S2513843X20000080:S2513843X20000080_fig2.png" class="graphic" data-v-241a4b23></div> <div data-v-241a4b23><div class="caption" data-v-241a4b23><p data-v-241a4b23><span class="label" data-v-241a4b23>Figure 2.</span> <span data-v-241a4b23><span class="p">Diversity hotspot of the Tungusic languages under the north–south classification in (a) as compared with the hotspot under the Manchu–Tungusic classification in (b).</span></span></p></div> </div></div> <hr aria-hidden="true" class="separator dashed" data-v-7036083a data-v-241a4b23></div><div data-v-241a4b23><div class="figures__item" data-v-241a4b23><div class="figures__item__image-box" data-v-241a4b23><button class="figures__ref" data-v-241a4b23> View in content </button> <img src="data:image/gif;base64,R0lGODlhAQABAIAAAMLCwgAAACH5BAAAAAAALAAAAAABAAEAAAICRAEAOw==" alt="Figure 2" data-zoomable="true" data-src="https://static.cambridge.org/binary/version/id/urn:cambridge.org:id:binary:20200421175645641-0667:S2513843X20000080:S2513843X20000080_tab1.png" data-enlarged-image="https://static.cambridge.org/binary/version/id/urn:cambridge.org:id:binary:20200421175645641-0667:S2513843X20000080:S2513843X20000080_tab1.png" class="graphic" data-v-241a4b23></div> <div data-v-241a4b23><div class="caption" data-v-241a4b23><p data-v-241a4b23><span class="label" data-v-241a4b23>Table 1.</span> <span data-v-241a4b23><span class="p">Evaluating the four homeland proposals for Proto-Tungusic using different archaeolinguistic techniques: 0 implausible, 1 plausible</span></span></p></div> </div></div> <hr aria-hidden="true" class="separator dashed" data-v-7036083a data-v-241a4b23></div><div data-v-241a4b23><div class="figures__item" data-v-241a4b23><div class="figures__item__image-box" data-v-241a4b23><button class="figures__ref" data-v-241a4b23> View in content </button> <img src="data:image/gif;base64,R0lGODlhAQABAIAAAMLCwgAAACH5BAAAAAAALAAAAAABAAEAAAICRAEAOw==" alt="Figure 3" data-zoomable="true" data-src="https://static.cambridge.org/binary/version/id/urn:cambridge.org:id:binary:20200421175645641-0667:S2513843X20000080:S2513843X20000080_fig3.png" data-enlarged-image="https://static.cambridge.org/binary/version/id/urn:cambridge.org:id:binary:20200421175645641-0667:S2513843X20000080:S2513843X20000080_fig3.png" class="graphic" data-v-241a4b23></div> <div data-v-241a4b23><div class="caption" data-v-241a4b23><p data-v-241a4b23><span class="label" data-v-241a4b23>Figure 3.</span> <span data-v-241a4b23><span class="p">Outgroup <span class="italic">f</span><span class="sub">3</span>-statistics of the form <span class="italic">f</span><span class="sub">3</span> (Mbuti; X, Y) to test the genetic drift sharing between ancient Devil's Gate and present-day East Eurasians.</span></span></p></div> </div></div> <hr aria-hidden="true" class="separator dashed" data-v-7036083a data-v-241a4b23></div><div data-v-241a4b23><div class="figures__item" data-v-241a4b23><div class="figures__item__image-box" data-v-241a4b23><button class="figures__ref" data-v-241a4b23> View in content </button> <img src="data:image/gif;base64,R0lGODlhAQABAIAAAMLCwgAAACH5BAAAAAAALAAAAAABAAEAAAICRAEAOw==" alt="Figure 4" data-zoomable="true" data-src="https://static.cambridge.org/binary/version/id/urn:cambridge.org:id:binary:20200421175645641-0667:S2513843X20000080:S2513843X20000080_fig4.png" data-enlarged-image="https://static.cambridge.org/binary/version/id/urn:cambridge.org:id:binary:20200421175645641-0667:S2513843X20000080:S2513843X20000080_fig4.png" class="graphic" data-v-241a4b23></div> <div data-v-241a4b23><div class="caption" data-v-241a4b23><p data-v-241a4b23><span class="label" data-v-241a4b23>Figure 4.</span> <span data-v-241a4b23><span class="p">Principal component analysis projecting ancient Devil's Gate, Ustlda and Jomon samples onto the variation of present-day East Eurasians.</span></span></p></div> </div></div> <hr aria-hidden="true" class="separator dashed" data-v-7036083a data-v-241a4b23></div><div data-v-241a4b23><div class="figures__item" data-v-241a4b23><div class="figures__item__image-box" data-v-241a4b23><button class="figures__ref" data-v-241a4b23> View in content </button> <img src="data:image/gif;base64,R0lGODlhAQABAIAAAMLCwgAAACH5BAAAAAAALAAAAAABAAEAAAICRAEAOw==" alt="Figure 5" data-zoomable="true" data-src="https://static.cambridge.org/binary/version/id/urn:cambridge.org:id:binary:20200421175645641-0667:S2513843X20000080:S2513843X20000080_fig5.png" data-enlarged-image="https://static.cambridge.org/binary/version/id/urn:cambridge.org:id:binary:20200421175645641-0667:S2513843X20000080:S2513843X20000080_fig5.png" class="graphic" data-v-241a4b23></div> <div data-v-241a4b23><div class="caption" data-v-241a4b23><p data-v-241a4b23><span class="label" data-v-241a4b23>Figure 5.</span> <span data-v-241a4b23><span class="p">Admixture plot from <span class="italic">K</span> = 2 to <span class="italic">K</span> = 12.</span></span></p></div> </div></div> </div></div> <div id="supplementary-materials-tab" class="supplementary-materials tab-pane"><div class="materials-table"><div class="materials-table__item"><div class="materials-table__row"><img 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Wang",searchUrl:"\u002Fcore\u002Fsearch?filters%5BauthorTerms%5D=Chuan-Chao%20Wang&eventCode=SE-AU",orcidUrl:f},{givenNames:"Martine",surname:"Robbeets",nameStyle:Q,affiliations:[{text:"Eurasia3angle Research Group, Max Planck Institute for the Science of Human History, Jena, Germany"}],isCorresponding:c,notes:"\u003Cdiv class=\"corresp\"\u003E\u003Cspan class=\"label\"\u003E*\u003C\u002Fspan\u003ECorresponding author. E-mail: \u003Ca href=\"mailto:robbeets@shh.mpg.de\"\u003Erobbeets@shh.mpg.de\u003C\u002Fa\u003E\u003C\u002Fdiv\u003E",isAnonymous:b,fullName:"Martine Robbeets",searchUrl:"\u002Fcore\u002Fsearch?filters%5BauthorTerms%5D=Martine%20Robbeets&eventCode=SE-AU",orcidUrl:"https:\u002F\u002Forcid.org\u002F0000-0002-2860-0230"}]},translators:{contributors:[],label:"Translated by"}},collections:[],publishedDate:"22 April 2020",keywords:[{url:"\u002Fcore\u002Fsearch?filters[keywords]=archaeolinguistics",name:"archaeolinguistics"},{url:"\u002Fcore\u002Fsearch?filters[keywords]=ancient DNA",name:"ancient DNA"},{url:"\u002Fcore\u002Fsearch?filters[keywords]=Tungusic language family",name:"Tungusic language family"},{url:"\u002Fcore\u002Fsearch?filters[keywords]=homeland",name:"homeland"},{url:"\u002Fcore\u002Fsearch?filters[keywords]=Amur genome",name:"Amur genome"}],openPracticeBadges:[],doi:{url:"https:\u002F\u002Fdoi.org\u002F10.1017\u002Fehs.2020.8",value:"10.1017\u002Fehs.2020.8"},copyright:{statement:["Copyright © The Author(s), 2020"],holder:["The Author(s)"],year:[2020]},creativeCommons:{descriptions:["This is an Open Access article, distributed under the terms of the Creative Commons Attribution licence (\u003Ca href=\"http:\u002F\u002Fcreativecommons.org\u002Flicenses\u002Fby\u002F4.0\u002F\" target=\"_blank\"\u003Ehttp:\u002F\u002Fcreativecommons.org\u002Flicenses\u002Fby\u002F4.0\u002F\u003C\u002Fa\u003E), which permits unrestricted re-use, distribution, and reproduction in any medium, provided the original work is properly cited."],oaLicenseType:"CC-BY",licenses:["CC","BY"]},acceptedManuscript:b,type:"research-article",typeDescription:"Research Article",eNumber:"e8",commentsCount:R,topicsAndSubtopics:f},journal:{id:S,title:m,titleSlug:T,mnemonic:"EHS",titleHistory:[],isFirstView:b,journalSlug:T,isCompanion:b,parentCompanionJournalName:m,associatedParentCollection:f,url:U,firstViewUrl:"\u002Fcore\u002Fjournals\u002Fevolutionary-human-sciences\u002Ffirstview",coverUrl:"https:\u002F\u002Fstatic.cambridge.org\u002Fcovers\u002FEHS_0_0_0\u002Fevolutionary-human-sciences.jpg",submitMaterialsUrl:"\u002Fcore\u002Fjournals\u002Fevolutionary-human-sciences\u002Finformation\u002Fauthor-instructions\u002Fsubmitting-your-materials",hasHistory:b,latestTitle:m,latestId:S,hasPastTitle:b,volume:{id:"F2345807349E632AB7170A2CDEAF9DF5",number:t,title:V,publishedDate:n,printPublishTimestamp:1577836800000,url:W}},abstract:{graphicalAbstract:{title:"Abstract",graphicalContent:"\u003Cdiv class=\"abstract\" data-abstract-type=\"graphical\"\u003E\u003Cp\u003E\u003Cimg class=\"graphic\" orientation mimesubtype=\"png\" mimetype=\"image\" position=\"float\" type href=\"S2513843X20000080_figAb.png\" src=\"https:\u002F\u002Fstatic.cambridge.org\u002Fcontent\u002Fid\u002Furn%3Acambridge.org%3Aid%3Aarticle%3AS2513843X20000080\u002Fresource\u002Fname\u002FS2513843X20000080_figAb.png?pub-status=live\" \u002F\u003E\u003C\u002Fp\u003E\u003C\u002Fdiv\u003E",textContent:"\u003Cdiv class=\"abstract\" data-abstract-type=\"normal\"\u003E\u003Cp\u003EThe Tungusic languages form a language family spoken in Xinjiang, Siberia, Manchuria and the Russian Far East. There is a general consensus that these languages are genealogically related and descend from a common ancestral language, conventionally called ‘Proto-Tungusic’. However, the exact geographical location where the ancestral speakers of Proto-Tungusic originated from is subject to debate. Here we take an unprecedented approach to this problem, by integrating linguistic, archaeological and genetic evidence in a single study. Our analysis of ancient DNA suggests genetic continuity between an ancient Amur genetic lineage and the contemporary speakers of the Tungusic languages. Adding an archaeolinguistic perspective, we infer that the most plausible homeland for the speakers of Proto-Tungusic is the region around Lake Khanka in the Russian Far East. Our study pushes the field forward in answering the tantalizing question about the location of the Tungusic homeland and in illustrating how these three disciplines can converge into a holistic approach to the human past.\u003C\u002Fp\u003E\u003C\u002Fdiv\u003E"}},content:{html:"\u003Cdiv class=\"article research-article NLM\"\u003E\u003Cdiv class=\"body\"\u003E\u003Cp class=\"p\"\u003E \u003Cstrong\u003EMedia summary:\u003C\u002Fstrong\u003E The Tungusic languages form a language family spoken in Xinjiang, Siberia, Manchuria and the Russian Far East. There is a general consensus that these languages descend from a common ancestral language, conventionally called ‘Proto-Tungusic’. However, the exact geographical location where the ancestral speakers of Proto-Tungusic originated from is subject to debate. Integrating linguistic, archaeological and genetic evidence in a single study, we here show that the most plausible homeland for the speakers of Proto-Tungusic is the region around Lake Khanka in the Russian Far East. Our study pushes the field forward in answering the tantalizing question about the location of the Tungusic homeland and in illustrating how these three disciplines can converge into a holistic approach on the human past.\u003C\u002Fp\u003E\u003Cdiv class=\"sec intro\" data-magellan-destination=\"sec1\" id=\"sec1\"\u003E\u003Ch2 class=\"A\"\u003E Introduction\u003C\u002Fh2\u003E\u003Cp class=\"p\"\u003E Even if they are rapidly losing ground owing to Chinese and Russian influence, the speakers of Tungusic languages are widely distributed all over Northeast Asia, including Xinjiang, Siberia, Manchuria and the Russian Far East. Their wide distribution and the considerable internal variation of populations have made the exact geographical location where the ancestral speakers of Proto-Tungusic originated from subject to debate. Traditionally, this issue was approached through the combined investigation of languages and material remains, but our recently acquired ability to sequence DNA directly from ancient human remains has considerably expanded the available scientific tool kit. Therefore, here we intend to complement linguistic and archaeological observations with genetic evidence to infer the most plausible homeland for the speakers of Proto-Tungusic.\u003C\u002Fp\u003E\u003Cp class=\"p\"\u003E The contemporary and historically attested Tungusic languages are spread from the Okhotsk Sea in the east to the Yenisei Basin in the west, and from the Bohai Sea in the south to the Arctic Ocean in the north. \u003Ca class=\"xref fig\" href=\"#fig01\"\u003EFigure 1\u003C\u002Fa\u003E shows the distribution of 12 Tungusic languages, Oroch, Udihe, Hezhe, Nanai, Orok, Olcha, Xibo, Even, Solon, Evenki, Negidal and Oroqen, along with dialectal varieties such as Kamnigan Evenki, Momsky Even, Olsky Even, Najkhin Nanai, Kur-Urmi Nanai and Bikin Nanai as well as two historical varieties, Jurchen and Manchu. Since written materials in Jurchen, the now-extinct language of the Jin dynasty (AD 1115–1234) are only partially deciphered, the earliest well-documented stage is Manchu, the official language of the Qing dynasty (AD 1644–1911).\n\u003C\u002Fp\u003E\u003Csection\u003E\u003Cdiv class=\"fig fig\" data-magellan-destination=\"fig01\" id=\"fig01\"\u003E\u003Cdiv class=\"figure-thumb\"\u003E\u003Cimg src=\"data:image\u002Fgif;base64,R0lGODlhAQABAIAAAMLCwgAAACH5BAAAAAAALAAAAAABAAEAAAICRAEAOw==\" data-src=\"https:\u002F\u002Fstatic.cambridge.org\u002Fbinary\u002Fversion\u002Fid\u002Furn:cambridge.org:id:binary:20200421175645641-0667:S2513843X20000080:S2513843X20000080_fig1.png?pub-status=live\" class=\"aop-lazy-load-image\" data-original-image=\"\u002Fbinary\u002Fversion\u002Fid\u002Furn:cambridge.org:id:binary:20200421175645641-0667:S2513843X20000080:S2513843X20000080_fig1.png\" data-zoomable=\"true\"\u003E\u003C\u002Fdiv\u003E\u003Cdiv class=\"caption\"\u003E\u003Cp class=\"p\"\u003E \u003C\u002Fp\u003E\u003Cp class=\"p\"\u003E\u003Cspan class=\"label\"\u003EFigure 1.\u003C\u002Fspan\u003E (a) The distribution of the Tungusic languages; (b) proposed locations for the homeland of the ancestral speakers of Proto-Tungusic.\u003C\u002Fp\u003E\u003C\u002Fdiv\u003E\u003C\u002Fdiv\u003E\u003C\u002Fsection\u003E\u003Cp class=\"p\"\u003E Although there is a general consensus that these languages are genealogically related and descend from a common ancestral language, conventionally called ‘Proto-Tungusic’, there are various competing hypotheses concerning the possible homeland of this language. Among the proposals, we find (a) the Baikal region (Vasilevich \u003Ca class=\"xref bibr\" href=\"#ref36\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Vasilevich\u003C\u002Fspan\u003E1960\u003C\u002Fa\u003E; Menges \u003Ca class=\"xref bibr\" href=\"#ref20\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Menges\u003C\u002Fspan\u003E1968\u003C\u002Fa\u003E; Derevyanko \u003Ca class=\"xref bibr\" href=\"#ref4\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Derevyanko\u003C\u002Fspan\u003E1976\u003C\u002Fa\u003E; Helimski \u003Ca class=\"xref bibr\" href=\"#ref8\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Helimski and Ubriatova\u003C\u002Fspan\u003E1985\u003C\u002Fa\u003E), (b) the Mid Amur and the lower part of the Upper Amur region (Janhunen \u003Ca class=\"xref bibr\" href=\"#ref10\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Janhunen\u003C\u002Fspan\u003E1996\u003C\u002Fa\u003E: 169; Korovina \u003Ca class=\"xref bibr\" href=\"#ref15\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Korovina\u003C\u002Fspan\u003E2011\u003C\u002Fa\u003E; Pevnov \u003Ca class=\"xref bibr\" href=\"#ref25\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Pevnov, Malchukov and Whaley\u003C\u002Fspan\u003E2012\u003C\u002Fa\u003E; Wichmann, personal communication, 3 October 2019), (c) the Ussuri region around Lake Khanka, a tributary of the Amur (Robbeets \u003Cem class=\"italic\"\u003Eet al\u003C\u002Fem\u003E. \u003Ca class=\"xref bibr\" href=\"#ref30\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Robbeets, Janhunen, Savelyev, Korovina, Robbeets and Savelyev\u003C\u002Fspan\u003E2020\u003C\u002Fa\u003E) and (d) the Yalu River region on the border between present-day Liaoning and Northern Korea (Janhunen \u003Ca class=\"xref bibr\" href=\"#ref11\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Janhunen, Malchukov and Whaley\u003C\u002Fspan\u003E2012\u003C\u002Fa\u003E). These locations are situated on the map in \u003Ca class=\"xref fig\" href=\"#fig01\"\u003EFigure 1\u003C\u002Fa\u003E.\u003C\u002Fp\u003E\u003Cp class=\"p\"\u003E After assessing these four different hypotheses from an archaeolinguistic perspective, we will open a genetic window into this question. By supplementing our archaeolinguistic argumentation with genetic evidence for continuity between an ancient Amur genetic profile and contemporary speakers of Tungusic, we will conclude that the region around Lake Khanka constitutes the most plausible homeland for the speakers of Proto-Tungusic. Through this holistic approach, we hope to answer the tantalizing question about the Tungusic homeland, posed by numerous specialists in the past.\u003C\u002Fp\u003E\u003C\u002Fdiv\u003E\u003Cdiv class=\"sec methods\" data-magellan-destination=\"sec2\" id=\"sec2\"\u003E\u003Ch2 class=\"A\"\u003E Methods\u003C\u002Fh2\u003E\u003Cdiv class=\"sec other\" data-magellan-destination=\"sec2-1\" id=\"sec2-1\"\u003E\u003Ch3 class=\"B\"\u003E Archaeolinguistic methods\u003C\u002Fh3\u003E\u003Cp class=\"p\"\u003E Traditionally, there are two principal techniques for attempting to determine the location of the original homeland of the speakers of a certain proto-language: the diversity hotspot principle and cultural reconstruction. The ‘diversity hotspot principle’ is a loose principle that was originally introduced by Edward Sapir (\u003Ca class=\"xref bibr\" href=\"#ref31\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Sapir\u003C\u002Fspan\u003E1916\u003C\u002Fa\u003E: 87) as the ‘center of gravity principle’, but it is also known as the ‘focus of diversity’ principle (Heggarty \u003Ca class=\"xref bibr\" href=\"#ref7\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Heggarty, Bowern and Evans\u003C\u002Fspan\u003E2015\u003C\u002Fa\u003E: 612–613). Assuming that the deepest splits within a family reflect the greatest age, the location of these splits on the map is thought to point to the area where the proto-language began to diversify. The principle is thus based on the assumption that the homeland is closest to where one finds the greatest diversity with regard to the deepest subgroups of the language family. Attempts have been made to formalize this principle in a computer-automated approach, computing a diversity measure for each language and identifying the homeland with the location of the language with the highest diversity measure (Wichmann \u003Cem class=\"italic\"\u003Eet al\u003C\u002Fem\u003E. \u003Ca class=\"xref bibr\" href=\"#ref40\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Wichmann, Müller and Velupillai\u003C\u002Fspan\u003E2010\u003C\u002Fa\u003E). A schoolbook example is the Austronesian family, that extends across a huge geographical range, all the way from Madagascar to Easter Island, but the deepest subgroups are found on just one Island, Taiwan, which is therefore identified as the homeland.\u003C\u002Fp\u003E\u003Cp class=\"p\"\u003E Although the diversity hotspot principle can provide some clues about the homeland of a language family, it must also contend with certain limitations. First, the identification of the homeland depends on the location of the deepest subgroups and therefore on how robustly the internal structure of a given family has been established. A second limitation of this principle is that the contemporary hotspot of linguistic diversity may diverge from the earlier one. Looking at the present map of Indo-European with the Balkan Peninsula hosting the highest diversity of deep subgroups, we might conclude that the homeland is there, instead of the Pontic Steppe or Anatolia. This example makes it clear that the application of the diversity hotspot principle at more profound time-depths may be speculative because earlier diversity may have been erased through time. Third, the diversity hotspot principle might also be upset when a migration was suddenly directed over a long distance rather than representing slow, gradual and random movement into adjacent areas. Finally, linguistic diversity is a function not only of time but also of other factors such as environmental change and disease. These may have made the original homeland unsuitable for human habitation at a certain point in time. In this way, original linguistic diversity may have been erased and it may no longer be possible to pinpoint the homeland using the diversity hotspot principle. However, even if the principle is not foolproof, it offers valuable clues for the location of a homeland at less remote time-depths. In the case of Tungusic, there is only mild disagreement on the primary branching structure, while the time depth is agreed to be relatively shallow, i.e. between 600 BCE and AD 600. Except for the historical transfer of the Xibo tribe speaking Manchu in 1763 from Manchuria to Xinjiang, there are no known cases of sudden long-distance movement. Therefore, we can gain more from applying the diversity hotspot principle to Tungusic than we can lose from ignoring it.\u003C\u002Fp\u003E\u003Cp class=\"p\"\u003E The second technique, cultural reconstruction, looks at the cultural vocabulary revealed in the reconstructed vocabulary of a proto-language. The method was first introduced under the label ‘linguistic paleontology’ by Adolphe Pictet (\u003Ca class=\"xref bibr\" href=\"#ref26\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Pictet\u003C\u002Fspan\u003E1859\u003C\u002Fa\u003E), but it became widespread under the banner of the ‘Wörter und Sachen’ movement, which used reconstructed proto-words as a way of determining proto-culture (Shuchardt \u003Ca class=\"xref bibr\" href=\"#ref33\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Shuchardt\u003C\u002Fspan\u003E1912\u003C\u002Fa\u003E). Cultural reconstruction is a vocabulary-based approach that enables us to study human prehistory by reconstructing lexical items that are ecologically or culturally diagnostic for a particular region at a particular time. Clues for fauna and flora can help to delimit a homeland if several reconstructed plant and animal names converge in a particular region to the exclusion of other regions. Reconstruction of cultural items such as cultivars and feedstock can help to identify an archaeological culture yielding the relevant material remains. The method can be combined with diachronic contact linguistics, the study of the ways in which ancestral languages influenced each other when their speakers interacted. As shown by the assumed borrowing of the term ‘barley’ below, this approach can be useful to shed light on prehistoric ethnic interaction and help us determine the chronology of our data.\u003C\u002Fp\u003E\u003Cp class=\"p\"\u003E Among the limitations of cultural reconstruction are the potential lack of accuracy in semantic reconstruction, the occurrence of lexical recycling, the deception of a single item not backed up by a semantic domain and the shakiness of inferences made on the basis of absence. Since semantic reconstruction is less precise than phonological reconstruction, we should first be cautious that the meaning assigned to a reconstructed form can be no more specific than the meaning shared by all the cognate forms.\u003C\u002Fp\u003E\u003Cp class=\"p\"\u003E Second, we should be aware of the possibility of ‘lexical recycling’, a process whereby words with a general, non-cultural meaning become repurposed as words with a specific, cultural meaning after the importation or invention of the corresponding innovation. The reconstruction for ‘foxtail millet’ in Supplementary Table S1 (3), for instance, could also indicate the wild variety because the crop is native to the Russian Far East.\u003C\u002Fp\u003E\u003Cp class=\"p\"\u003E Third, we should avoid reconstructing a single cultural item that is not backed up by other members of the semantic domain. Paying attention to the clustering of different cultural reconstructions in a specific semantic domain, such as agriculture, can contribute to a fuller picture of prehistory than the study of individual cultural reconstructions.\u003C\u002Fp\u003E\u003Cp class=\"p\"\u003E And fourth, ‘absence of evidence is not evidence of absence’. The observation that an agricultural lexicon cannot be reconstructed for a certain proto-language may be explained by the fact that the proto-speakers simply were not familiar with farming, but it could also be due to the lack of exhaustive research or to the attrition of agricultural cognates over time. Therefore, inferences made on the basis of absence are not necessarily wrong, but they should not be taken as absolute proof for an argument.\u003C\u002Fp\u003E\u003Cp class=\"p\"\u003E Nevertheless, when combined with the necessary checks and balances, cultural reconstruction is reasonably robust as a way of determining proto-ecology and proto-culture through proto-language. Therefore, we will apply this method in order to make predictions about the cultural and natural environment available to the speakers of Proto-Tungusic.\u003C\u002Fp\u003E\u003C\u002Fdiv\u003E\u003Cdiv class=\"sec other\" data-magellan-destination=\"sec2-2\" id=\"sec2-2\"\u003E\u003Ch3 class=\"B\"\u003E Genetic methods\u003C\u002Fh3\u003E\u003Cp class=\"p\"\u003E We reanalyzed the genome-wide data of ancient and present-day East Eurasian populations related to Tungusic-speaking populations from the following literature: Siska \u003Cem class=\"italic\"\u003Eet al\u003C\u002Fem\u003E. (\u003Ca class=\"xref bibr\" href=\"#ref34\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Siska, Jones, Jeon, Bhak, Kim, Cho, Kim, Lee, Veselovskaya, Balueva, Gallego-Llorente, Hofreiter, Bradley, Eriksson, Pinhasi, Bhak and Manica\u003C\u002Fspan\u003E2017\u003C\u002Fa\u003E), de Barros Damgaard \u003Cem class=\"italic\"\u003Eet al\u003C\u002Fem\u003E. (\u003Ca class=\"xref bibr\" href=\"#ref24\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Damgaard, Marchi, Rasmussen, Peyrot, Renaud, Korneliussen, Moreno-Mayar, Pedersen, Goldberg, Usmanova, Baimukhanov, Loman, Hedeager, Pedersen, Nielsen, Afanasiev, Akmatov, Aldashev, Alpaslan, Baimbetov, Bazaliiskii, Beisenov, Boldbaatar, Boldgiv, Dorzhu, Ellingvag, Erdenebaatar, Dajani, Dmitriev, Evdokimov, Frei, Gromov, Goryachev, Hakonarson, Hegay, Khachatryan, Khaskhanov, Kitov, Kolbina, Kubatbek, Kukushkin, Kukushkin, Lau, Margaryan, Merkyte, Mertz, Mertz, Mijiddorj, Moiyesev, Mukhtarova, Nurmukhanbetov, Orozbekova, Panyushkina, Pieta, Smrčka, Shevnina, Logvin, Sjogren, Štolcova, Taravella, Tashbaeva, Tkachev, Tulegenov, Voyakin, Yepiskoposyan, Undrakhbold, Varfolomeev, Weber, Wilson Sayres, Kradin, Allentoft, Orlando, Nielsen, Sikora, Heyer, Kristiansen and Willerslev\u003C\u002Fspan\u003E2018\u003C\u002Fa\u003E), Kanzawa-Kiriyama \u003Cem class=\"italic\"\u003Eet al\u003C\u002Fem\u003E. (\u003Ca class=\"xref bibr\" href=\"#ref13\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Kanzawa-Kiriyama, Kryukov, Jinam, Hosomichi, Saso, Suwa, Ueda, Yoneda, Tajima, Shinoda, Inoue and Saitou\u003C\u002Fspan\u003E2016\u003C\u002Fa\u003E), Patterson \u003Cem class=\"italic\"\u003Eet al\u003C\u002Fem\u003E. (\u003Ca class=\"xref bibr\" href=\"#ref23\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Patterson, Moorjani, Luo, Mallick, Rohland, Zhan, Genschoreck, Webster and Reich\u003C\u002Fspan\u003E2012\u003C\u002Fa\u003E), Jeong \u003Cem class=\"italic\"\u003Eet al\u003C\u002Fem\u003E. (\u003Ca class=\"xref bibr\" href=\"#ref12\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Jeong, Balanovsky, Lukianova, Kahbatkyzy, Flegontov, Zaporozhchenko, Immel, Wang, Ixan, Khussainova, Bekmanov, Zaibert, Lavryashina, Pocheshkhova, Yusupov, Agdzhoyan, Koshel, Bukin, Nymadawa, Turdikulova, Dalimova, Churnosov, Skhalyakho, Daragan, Bogunov, Bogunova, Shtrunov, Dubova, Zhabagin, Yepiskoposyan, Churakov, Pislegin, Damba, Saroyants, Dibirova, Atramentova, Utevska, Idrisov, Kamenshchikova, Evseeva, Metspalu, Outram, Robbeets, Djansugurova, Balanovska, Schiffels, Haak, Reich and Krause\u003C\u002Fspan\u003E2019\u003C\u002Fa\u003E) and Lazaridis \u003Cem class=\"italic\"\u003Eet al\u003C\u002Fem\u003E. (\u003Ca class=\"xref bibr\" href=\"#ref16\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Lazaridis, Patterson, Mittnik, Renaud, Mallick, Kirsanow, Sudmant, Schraiber, Castellano, Lipson, Berger, Economou, Bollongino, Fu, Bos, Nordenfelt, Li, de Filippo, Prüfer, Sawyer, Posth, Haak, Hallgren, Fornander, Rohland, Delsate, Francken, Guinet, Wahl, Ayodo, Babiker, Bailliet, Balanovska, Balanovsky, Barrantes, Bedoya, Ben-Ami, Bene, Berrada, Bravi, Brisighelli, Busby, Cali, Churnosov, Cole, Corach, Damba, van Driem, Dryomov, Dugoujon, Fedorova, Gallego Romero, Gubina, Hammer, Henn, Hervig, Hodoglugil, Jha, Karachanak-Yankova, Khusainova, Khusnutdinova, Kittles, Kivisild, Klitz, Kučinskas, Kushniarevich, Laredj, Litvinov, Loukidis, Mahley, Melegh, Metspalu, Molina, Mountain, Näkkäläjärvi, Nesheva, Nyambo, Osipova, Parik, Platonov, Posukh, Romano, Rothhammer, Rudan, Ruizbakiev, Sahakyan, Sajantila, Salas, Starikovskaya, Tarekegn, Toncheva, Turdikulova, Uktveryte, Utevska, Vasquez, Villena, Voevoda, Winkler, Yepiskoposyan, Zalloua, Zemunik, Cooper, Capelli, Thomas, Ruiz-Linares, Tishkoff, Singh, Thangaraj, Villems, Comas, Sukernik, Metspalu, Meyer, Eichler, Burger, Slatkin, Pääbo, Kelso, Reich and Krause\u003C\u002Fspan\u003E2014\u003C\u002Fa\u003E).\u003C\u002Fp\u003E\u003Cp class=\"p\"\u003E We first carried out principal components analysis in the smartpca program of EIGENSOFT (Patterson \u003Cem class=\"italic\"\u003Eet al\u003C\u002Fem\u003E. \u003Ca class=\"xref bibr\" href=\"#ref22\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Patterson, Price and Reich\u003C\u002Fspan\u003E2006\u003C\u002Fa\u003E), using default parameters and the lsqproject: YES and numoutlieriter: 0 options. We projected ancient Devil's Gate Ustlda and Jomon samples onto the variation of present-day East Eurasians from the published Human Origin Dataset over 591,642 SNPs. We carried out ADMIXTURE analysis after pruning for linkage disequilibrium in PLINK with parameters --indep-pairwise 200 25 0.4 Chang \u003Cem class=\"italic\"\u003Eet al.\u003C\u002Fem\u003E (\u003Ca class=\"xref bibr\" href=\"#ref2a\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Chang, Chow, Tellier, Vattikuti, Purcell and Lee\u003C\u002Fspan\u003E2015\u003C\u002Fa\u003E) which retained 332,959 SNPs. We ran ADMIXTURE (Alexander \u003Cem class=\"italic\"\u003Eet al\u003C\u002Fem\u003E. \u003Ca class=\"xref bibr\" href=\"#ref1\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Alexander, Novembre and Lange\u003C\u002Fspan\u003E2009\u003C\u002Fa\u003E) with default 5-fold cross-validation (--cv = 5), varying the number of ancestral populations between \u003Cem class=\"italic\"\u003EK\u003C\u002Fem\u003E = 2 and \u003Cem class=\"italic\"\u003EK\u003C\u002Fem\u003E = 12 in 100 bootstraps with different random seeds. We observed the lowest CV error at \u003Cem class=\"italic\"\u003EK\u003C\u002Fem\u003E = 8. The samples used in this analysis are present-day East Eurasians, and individuals from worldwide representative populations. We used outgroup \u003Cem class=\"italic\"\u003Ef\u003C\u002Fem\u003E \u003Csub\u003E3\u003C\u002Fsub\u003E-statistics of the form \u003Cem class=\"italic\"\u003Ef\u003C\u002Fem\u003E \u003Csub\u003E3\u003C\u002Fsub\u003E (Mbuti; X, Y) to test the genetic drift sharing between ancient Devil's Gate and present-day East Eurasians. Analysis of \u003Cem class=\"italic\"\u003Ef\u003C\u002Fem\u003E \u003Csub\u003E3\u003C\u002Fsub\u003E-statistics was carried out using ADMIXTOOLS with standard errors computed with a block jackknife (Patterson \u003Cem class=\"italic\"\u003Eet al\u003C\u002Fem\u003E. \u003Ca class=\"xref bibr\" href=\"#ref23\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Patterson, Moorjani, Luo, Mallick, Rohland, Zhan, Genschoreck, Webster and Reich\u003C\u002Fspan\u003E2012\u003C\u002Fa\u003E). We used \u003Cem class=\"italic\"\u003EqpAdm\u003C\u002Fem\u003E as implemented in ADMIXTOOLS to estimate West Eurasian mixture proportions in Even and Evenk with a set of outgroup populations. Weighted linkage disequilibrium (LD) decay was calculated using ALDER to infer admixture parameters including dates and mixture proportions (Loh \u003Cem class=\"italic\"\u003Eet al\u003C\u002Fem\u003E. \u003Ca class=\"xref bibr\" href=\"#ref19\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Loh, Lipson, Patterson, Moorjani, Pickrell, Reich and Berger\u003C\u002Fspan\u003E2013\u003C\u002Fa\u003E). We added Table S2 representing West Eurasian-related admixture proportions estimated using \u003Cem class=\"italic\"\u003EqpAdm\u003C\u002Fem\u003E and Table S3 listing the admixture time with West Eurasian-related groups estimated by ALDER in the Supplementary Information.\u003C\u002Fp\u003E\u003C\u002Fdiv\u003E\u003C\u002Fdiv\u003E\u003Cdiv class=\"sec results\" data-magellan-destination=\"sec3\" id=\"sec3\"\u003E\u003Ch2 class=\"A\"\u003E Results\u003C\u002Fh2\u003E\u003Cdiv class=\"sec other\" data-magellan-destination=\"sec3-1\" id=\"sec3-1\"\u003E\u003Ch3 class=\"B\"\u003E Archaeolinguistic results\u003C\u002Fh3\u003E\u003Cp class=\"p\"\u003E The principle that the geographical area of maximal primary diversity within a language family corresponds to the original homeland implies that the primary splits in the family are determinant for the location of the homeland. There is some disagreement on the internal structure of the Tungusic language family, mainly with respect to the separation of the Manchuric branch. As shown in \u003Ca class=\"xref fig\" href=\"#fig02\"\u003EFigure 2\u003C\u002Fa\u003E, some authors proposed a north–south classification, in which the separation of Manchuric from the other Tungusic languages does not constitute the earliest split in the family (Cincius \u003Ca class=\"xref bibr\" href=\"#ref3\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Cincius\u003C\u002Fspan\u003E1949\u003C\u002Fa\u003E; Benzing \u003Ca class=\"xref bibr\" href=\"#ref2\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Benzing\u003C\u002Fspan\u003E1955\u003C\u002Fa\u003E; Kormušin \u003Ca class=\"xref bibr\" href=\"#ref14\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Kormušin\u003C\u002Fspan\u003E1998\u003C\u002Fa\u003E; Georg \u003Ca class=\"xref bibr\" href=\"#ref6\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Georg, Naeher, Stary and Weiers\u003C\u002Fspan\u003E2004\u003C\u002Fa\u003E; Janhunen \u003Ca class=\"xref bibr\" href=\"#ref11\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Janhunen, Malchukov and Whaley\u003C\u002Fspan\u003E2012\u003C\u002Fa\u003E), while others proposed an early breakup between Manchuric and the rest of Tungusic (Sunik \u003Ca class=\"xref bibr\" href=\"#ref35\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Sunik\u003C\u002Fspan\u003E1959\u003C\u002Fa\u003E; Vasilevich \u003Ca class=\"xref bibr\" href=\"#ref36\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Vasilevich\u003C\u002Fspan\u003E1960\u003C\u002Fa\u003E; Doerfer \u003Ca class=\"xref bibr\" href=\"#ref5\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Doerfer and Weiers\u003C\u002Fspan\u003E1978\u003C\u002Fa\u003E; Vovin \u003Ca class=\"xref bibr\" href=\"#ref37\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Vovin\u003C\u002Fspan\u003E1993\u003C\u002Fa\u003E; Robbeets \u003Ca class=\"xref bibr\" href=\"#ref27\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Robbeets\u003C\u002Fspan\u003E2015\u003C\u002Fa\u003E; Whaley and Oskolskaya \u003Ca class=\"xref bibr\" href=\"#ref39\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Whaley, Oskolskaya, Robbeets and Savelyev\u003C\u002Fspan\u003E2020\u003C\u002Fa\u003E). Depending on which of both classifications of Tungusic we favor, the center of primary diversity – and thus also the inferred location of the homeland – will move on the map, as illustrated in \u003Ca class=\"xref fig\" href=\"#fig02\"\u003EFigure 2\u003C\u002Fa\u003E. Whereas the north–south classification pushes the homeland to the north into the Mid Amur region (\u003Ca class=\"xref fig\" href=\"#fig02\"\u003EFigure 2a\u003C\u002Fa\u003E), the Manchu–Tungusic classification pulls it more southwards, into the area around Lake Khanka (\u003Ca class=\"xref fig\" href=\"#fig02\"\u003EFigure 2b\u003C\u002Fa\u003E).\n\u003C\u002Fp\u003E\u003Csection\u003E\u003Cdiv class=\"fig fig\" data-magellan-destination=\"fig02\" id=\"fig02\"\u003E\u003Cdiv class=\"figure-thumb\"\u003E\u003Cimg src=\"data:image\u002Fgif;base64,R0lGODlhAQABAIAAAMLCwgAAACH5BAAAAAAALAAAAAABAAEAAAICRAEAOw==\" data-src=\"https:\u002F\u002Fstatic.cambridge.org\u002Fbinary\u002Fversion\u002Fid\u002Furn:cambridge.org:id:binary:20200421175645641-0667:S2513843X20000080:S2513843X20000080_fig2.png?pub-status=live\" class=\"aop-lazy-load-image\" data-original-image=\"\u002Fbinary\u002Fversion\u002Fid\u002Furn:cambridge.org:id:binary:20200421175645641-0667:S2513843X20000080:S2513843X20000080_fig2.png\" data-zoomable=\"true\"\u003E\u003C\u002Fdiv\u003E\u003Cdiv class=\"caption\"\u003E\u003Cp class=\"p\"\u003E \u003C\u002Fp\u003E\u003Cp class=\"p\"\u003E\u003Cspan class=\"label\"\u003EFigure 2.\u003C\u002Fspan\u003E Diversity hotspot of the Tungusic languages under the north–south classification in (a) as compared with the hotspot under the Manchu–Tungusic classification in (b).\u003C\u002Fp\u003E\u003C\u002Fdiv\u003E\u003C\u002Fdiv\u003E\u003C\u002Fsection\u003E\u003Cp class=\"p\"\u003E Restricting himself to 11 Tungusic languages, notably Even, Evenki, Negidal, Oroqen, Naykhin Nanai, Oroch, Orok, Olcha, Udehe, Manchu and Xibo, Wichmann (personal communication) situates the Tungusic homeland around the location of Naykhin Nanai at latitude 49.28 and longitude 136.47. As more southern varieties such as Solon on the river Nonni in Inner Mongolia and Bikin Nanai in the southern part of the Primorye province as well as Hezhe in Heilongjiang and Jilin are lacking from this analysis, we may expect the real diversity hotspot to be more to the south than the inferred one. In sum, the diversity hotspot principle leaves room for both the Mid Amur region and the area around Lake Khanka but excludes the Baikal and the Yalu river region as plausible homelands.\u003C\u002Fp\u003E\u003Cp class=\"p\"\u003E The breakup of proto-Tungusic took place relatively recently. Different linguistic dating methods, such as lexicostatistics (Korovina \u003Ca class=\"xref bibr\" href=\"#ref15\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Korovina\u003C\u002Fspan\u003E2011\u003C\u002Fa\u003E: 600 BCE), dating of ethnonym shift (Robbeets \u003Ca class=\"xref bibr\" href=\"#ref27\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Robbeets\u003C\u002Fspan\u003E2015\u003C\u002Fa\u003E: 206 BC to AD 220), measurement of mutual intelligibility (Pevnov \u003Ca class=\"xref bibr\" href=\"#ref25\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Pevnov, Malchukov and Whaley\u003C\u002Fspan\u003E2012\u003C\u002Fa\u003E: AD 1), Bayesian phylolinguistics (Oskolskaya \u003Cem class=\"italic\"\u003Eet al\u003C\u002Fem\u003E. (\u003Ca class=\"xref bibr\" href=\"#ref21\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Oskolskaya, Robbeets and Koile\u003C\u002Fspan\u003Eunder review\u003C\u002Fa\u003E): AD 200), cultural reconstruction (Janhunen \u003Ca class=\"xref bibr\" href=\"#ref11\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Janhunen, Malchukov and Whaley\u003C\u002Fspan\u003E2012\u003C\u002Fa\u003E: 500 BC to AD 500) and the automated similarity judgment program (Holman \u003Cem class=\"italic\"\u003Eet al\u003C\u002Fem\u003E. \u003Ca class=\"xref bibr\" href=\"#ref9\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Holman, Brown, Wichmann, Müller, Velupillai, Hammarström, Sauppe, Jung, Bakker, Brown, Belyaev, Urban, Mailhammer, List and Egorov\u003C\u002Fspan\u003E2011\u003C\u002Fa\u003E: AD 681) yield different dates, but all dates range between 600 BC and AD 700, which supports a relatively shallow time-depth for the family.\u003C\u002Fp\u003E\u003Cp class=\"p\"\u003E Applying cultural reconstruction to the Tungusic languages, we find that Proto-Tungusic culture can be associated with the reconstruction of PTg *\u003Cem class=\"italic\"\u003Esele\u003C\u002Fem\u003E ‘iron’, reflected in corresponding forms in the contemporary languages all sharing the meaning ‘iron’; see (1) in Supplementary Table S1. The first uncontested findings of iron in the Russian Far East go back to the Krounovska culture (600 BC to AD 200), which marks the beginning of the Early Iron Age in the region. From there, iron spread northwards, to the northern Primorye and the Priamurye during the ensuing period of the Pol'tse and Ol'ga cultures (200 BC to AD 500), but there is no evidence for iron as north as the Upper Amur region around that time. Iron reached Liaoning and southwestern Jilin no later than the third century BCE. Broadly speaking iron begins to appear early first millennium BCE in the Baikal region. Thus, the reconstruction of iron is compatible with all locations, except the Mid and Upper Amur region.\u003C\u002Fp\u003E\u003Cp class=\"p\"\u003E As shown in the etymologies (2)–(6) in Supplementary Table S1, we can reconstruct agricultural vocabulary to Proto-Tungusic, such as *\u003Cem class=\"italic\"\u003Episi-ke\u003C\u002Fem\u003E ‘broomcorn millet (\u003Cem class=\"italic\"\u003EPanicum miliaceum\u003C\u002Fem\u003E)’, *\u003Cem class=\"italic\"\u003Ejiya\u003C\u002Fem\u003E ‘foxtail millet (\u003Cem class=\"italic\"\u003ESetaria italica\u003C\u002Fem\u003E)’, *\u003Cem class=\"italic\"\u003Eüse\u003C\u002Fem\u003E- ~ \u003Cem class=\"italic\"\u003Eüsi\u003C\u002Fem\u003E- ‘to plant’, *\u003Cem class=\"italic\"\u003Eüse\u003C\u002Fem\u003E ~ \u003Cem class=\"italic\"\u003Eüsi\u003C\u002Fem\u003E ‘seed, seedling’, *\u003Cem class=\"italic\"\u003Eüsin\u003C\u002Fem\u003E ‘field for cultivation’ and *\u003Cem class=\"italic\"\u003Etari\u003C\u002Fem\u003E- ‘to sow, plant, cultivate’. The use of these words by the speakers of Proto-Tungusic implies that they were familiar with agriculture. In addition to this familiarity with agricultural concepts, there are a number of other reasons to assume that the speakers of Proto-Tungusic were cultivating plants. First, the derivation of the Proto-Tungusic word *\u003Cem class=\"italic\"\u003Eüse\u003C\u002Fem\u003E ~ \u003Cem class=\"italic\"\u003Eüsi\u003C\u002Fem\u003E ‘seed, seedling’ as a deverbal noun from the verb *\u003Cem class=\"italic\"\u003Eüse\u003C\u002Fem\u003E- ~ \u003Cem class=\"italic\"\u003Eüsi\u003C\u002Fem\u003E- ‘to plant’ suggests that seeds were not just collected for consumption but that they were planted as part of a cultivation process.\u003C\u002Fp\u003E\u003Cp class=\"p\"\u003E Second, for some crop names such as ‘barley’ and ‘broomcorn millet’, we can argue that the word refers to the domesticated crop rather than to the wild variety of the plant because both crops are not native to the region and have been imported as domesticated crops. It is commonly assumed that broomcorn millet has been imported from the West Liao River region by the people who introduced the Zaisanovska culture (3200–1300 BCE) to the area around Lake Khanka (Sergusheva and Vostretsov \u003Ca class=\"xref bibr\" href=\"#ref32\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Sergusheva, Vostretsov, Fairbairn and Weiss\u003C\u002Fspan\u003E2009\u003C\u002Fa\u003E; Leipe \u003Cem class=\"italic\"\u003Eet al\u003C\u002Fem\u003E. \u003Ca class=\"xref bibr\" href=\"#ref17\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Leipe, Long, Sergusheva, Wagner and Tarasov\u003C\u002Fspan\u003E2019\u003C\u002Fa\u003E; Li \u003Cem class=\"italic\"\u003Eet al\u003C\u002Fem\u003E. \u003Ca class=\"xref bibr\" href=\"#ref18\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Li, Ning, Zhushchikhovskaya, Hudson and Robbeets\u003C\u002Fspan\u003E2020\u003C\u002Fa\u003E). That the origins of the agricultural vocabulary in Proto-Tungusic go back as early as the fourth millennium BC in the southern part of the Primorye is further supported by the external etymologies of some words. Since some of the terms such as *\u003Cem class=\"italic\"\u003Etari\u003C\u002Fem\u003E- ‘to sow, to plant’ have cognates in Turkic and Mongolic; these agricultural terms must have been present in proto-Tungusic since its separation from proto-Altaic, estimated at 3300 BC (Robbeets and Bouckaert \u003Ca class=\"xref bibr\" href=\"#ref29\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Robbeets and Bouckaert\u003C\u002Fspan\u003E2018\u003C\u002Fa\u003E; Robbeets \u003Cem class=\"italic\"\u003Eet al\u003C\u002Fem\u003E. \u003Ca class=\"xref bibr\" href=\"#ref30\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Robbeets, Janhunen, Savelyev, Korovina, Robbeets and Savelyev\u003C\u002Fspan\u003E2020\u003C\u002Fa\u003E). Only in the Zaisanovska culture around Lake Khanka and in the Wangjiacun site in Dalian on the North Korean border does agriculture go back that deep in time. Thus, the presence of agricultural vocabulary excludes the Baikal region and the Mid and Upper Amur region as plausible homelands.\u003C\u002Fp\u003E\u003Cp class=\"p\"\u003E The rather large time difference between the separation of Proto-Tungusic from Proto-Altaic around 3300 BC and the actual dispersal of Tungusic languages after 600 BC does not imply that proto-Tungusic continued as a single language without any diversification during three millennia. On the contrary, it is safe to assume that the contemporary Tungusic languages represent but a tiny proportion of the original diversity and that in the first millennia BCE various sister languages of Proto-Tungusic were spoken in the area. The adoption of agriculture involved a previously unparallelled demographic growth in certain production areas in North East China. With its relatively mild climate, the protected region of Lake Khanka constituted such an area, close to the northern edge of the climate range for East Asian Neolithic agriculture. Demographically, this implies a large population, while linguistically, it means that speech communities which happened to be in the center of agricultural innovation underwent rapid growth and were able to absorb neighboring speech communities by way of language shift. In this way, certain Proto-Tungusic speech communities in the region became highly expansive at the expense of others. This expansion must have affected a large number of individual languages belonging to various other language families, such as the Amuric family that was reduced to only one surviving language, Nivkh, but it may also have affected now-extinct sister branches of Proto-Tungusic. From the number of surviving contemporary languages, it is clear that the Transeurasian languages in general have a relatively high extinction rate. By way of comparison, Austronesian has 1500 surviving languages, whereas hardly 50 Transeurasian languages have survived to the present. This supports the assumption that only a happy few expansive languages survived in Northeast Asia, at the expense of numerous others.\u003C\u002Fp\u003E\u003Cp class=\"p\"\u003E A location around Lake Khanka for the Proto-Tungusic speech community is further corroborated by contact linguistics. The term for ‘barley and similar crops’, PTg *\u003Cem class=\"italic\"\u003Emurgi\u003C\u002Fem\u003E, is probably borrowed from an Old Chinese donor word, 來*\u003Cem class=\"italic\"\u003Emə.rˤək\u003C\u002Fem\u003E > *\u003Cem class=\"italic\"\u003Emə.rˤə\u003C\u002Fem\u003E, ‘a kind of wheat’ (Robbeets \u003Ca class=\"xref bibr\" href=\"#ref28\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Robbeets, Robbeets and Savelyev\u003C\u002Fspan\u003E2017\u003C\u002Fa\u003E: 28–29). The linguistic reconstruction can be correlated to the archaeological evidence for barley being first imported through Chinese contact at the time of the Krounovska culture situated in the Southern Primorye south of Lake Khanka (Sergusheva and Vostretsov \u003Ca class=\"xref bibr\" href=\"#ref32\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Sergusheva, Vostretsov, Fairbairn and Weiss\u003C\u002Fspan\u003E2009\u003C\u002Fa\u003E: 214–215; Leipe \u003Cem class=\"italic\"\u003Eet al\u003C\u002Fem\u003E. \u003Ca class=\"xref bibr\" href=\"#ref17\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Leipe, Long, Sergusheva, Wagner and Tarasov\u003C\u002Fspan\u003E2019\u003C\u002Fa\u003E: 12).\u003C\u002Fp\u003E\u003Cp class=\"p\"\u003E Proceeding to the reconstruction of ecological vocabulary, we can reconstruct tree names to Proto-Tungusic, such as *\u003Cem class=\"italic\"\u003Einʹŋamu-kta\u003C\u002Fem\u003E ‘cloudberry (\u003Cem class=\"italic\"\u003ERubus chamaemorus\u003C\u002Fem\u003E)’, *\u003Cem class=\"italic\"\u003Exiba-gda\u003C\u002Fem\u003E ‘ash tree (\u003Cem class=\"italic\"\u003EFráxinus mandshurica\u003C\u002Fem\u003E)’, *\u003Cem class=\"italic\"\u003Exami-gda\u003C\u002Fem\u003E ‘poplar (\u003Cem class=\"italic\"\u003EPopulus maximowiczii\u003C\u002Fem\u003E)’, *\u003Cem class=\"italic\"\u003Ekilden\u003C\u002Fem\u003E ‘linden (\u003Cem class=\"italic\"\u003ETilia amurensis\u003C\u002Fem\u003E)’ and *\u003Cem class=\"italic\"\u003Emolon\u003C\u002Fem\u003E ‘maple (\u003Cem class=\"italic\"\u003EAcer ukurunduense\u003C\u002Fem\u003E)’, as shown in Supplementary Table S1 (7)–(11). Except for ‘cloudberry’, none of these species appears in the Baikal region, while maple does not appear in the Mid and Upper Amur region. Among the Proto-Tungusic tree names, the terms for ‘pine tree’ in Supplementary Table S1 (12)–(14) are particularly telling. In contrast to the wide distribution of the ‘creeping pine’, that of the ‘Korean pine (\u003Cem class=\"italic\"\u003EPínus koraiénsis\u003C\u002Fem\u003E)’ is limited to the area around Lake Khanka. In addition, it is the only plant for which we have archaeobotanical evidence. The nutshells of this pine were found in pit-dwelling deposits in the Southern Primorye, from 1300 BC onwards (Sergusheva and Vostretsov \u003Ca class=\"xref bibr\" href=\"#ref32\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Sergusheva, Vostretsov, Fairbairn and Weiss\u003C\u002Fspan\u003E2009\u003C\u002Fa\u003E: 213). Therefore the reconstruction of the word for Korean pine strongly supports a homeland around Lake Khanka.\u003C\u002Fp\u003E\u003Cp class=\"p\"\u003E \u003Ca class=\"xref table\" href=\"#tab01\"\u003ETable 1\u003C\u002Fa\u003E assesses the different homeland proposals from an archaeolinguistic viewpoint. The hypothesis of Lake Khanka is best supported by the linguistic data, while the Baikal hypothesis is the least compatible. Situating the Proto-Tungusic homeland in the Lake Khanka area is further supported by the continuity of political formations in the region such as the Mohe (AD 500–1000) in the area around Lake Khanka, the Bohai state (AD 698–925) southwards from lake Khanka into present-day North Korea and the Jin Empire of the Jurchen (AD 1115–1234) to the west of Lake Khanka in the Sungari, Nonni and Middle Amur region. Since Neolithic times, the cultural edge in the Russian Far East was in the south, not in the north. Therefore, the area around Lake Khanka is the most plausible homeland for the ancestral Tungusic speech community.\n\u003C\u002Fp\u003E\u003Cdiv class=\"table-wrap\" data-magellan-destination=\"tab01\" id=\"tab01\"\u003E\u003Cdiv class=\"caption\"\u003E\u003Cp class=\"p\"\u003E\u003Cspan class=\"label\"\u003ETable 1.\u003C\u002Fspan\u003E Evaluating the four homeland proposals for Proto-Tungusic using different archaeolinguistic techniques: 0 implausible, 1 plausible\u003C\u002Fp\u003E\u003C\u002Fdiv\u003E\u003Cspan\u003E\u003Cdiv class=\"figure-thumb\"\u003E\u003Cimg src=\"data:image\u002Fgif;base64,R0lGODlhAQABAIAAAMLCwgAAACH5BAAAAAAALAAAAAABAAEAAAICRAEAOw==\" data-src=\"https:\u002F\u002Fstatic.cambridge.org\u002Fbinary\u002Fversion\u002Fid\u002Furn:cambridge.org:id:binary:20200421175645641-0667:S2513843X20000080:S2513843X20000080_tab1.png?pub-status=live\" class=\"aop-lazy-load-image\" data-original-image=\"\u002Fbinary\u002Fversion\u002Fid\u002Furn:cambridge.org:id:binary:20200421175645641-0667:S2513843X20000080:S2513843X20000080_tab1.png\" data-zoomable=\"false\"\u003E\u003C\u002Fdiv\u003E\u003C\u002Fspan\u003E\u003C\u002Fdiv\u003E\u003C\u002Fdiv\u003E\u003Cdiv class=\"sec other\" data-magellan-destination=\"sec3-2\" id=\"sec3-2\"\u003E\u003Ch3 class=\"B\"\u003E Genetic results\u003C\u002Fh3\u003E\u003Cp class=\"p\"\u003E From a paternal perspective, Tungusic speakers are associated with the Y chromosomal haplogroup C3-M217. The haplogroup C3c-M48 is prevalent in Evenki and Even, as well as in other Tungusic-speaking populations in the Amur River Basin, including Oroqen, Olcha, Negidal, Udehe and Nanai with frequencies ranging from about 20% to even 100%. The C3-M217 type also reaches high frequencies in populations surrounding the Amur River Basin, such as Nivkh (38%) and Ainu (12.5–25%) (Wei \u003Ca class=\"xref bibr\" href=\"#ref38\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Wei\u003C\u002Fspan\u003E2011\u003C\u002Fa\u003E). The current geographic and genetic diversity distribution pattern suggests that the dispersal of Y chromosomal haplogroup C3-M217 was most likely associated with Tungusic-related population expansion from the Amur River.\u003C\u002Fp\u003E\u003Cp class=\"p\"\u003E From a genome-wide perspective, we reanalyzed the data of ancient samples from Devil's Gate (Chertovy Vorota) Cave dating to ~7.7 kya in the Primorye Region of the Russian Far East (Siska \u003Cem class=\"italic\"\u003Eet al\u003C\u002Fem\u003E. \u003Ca class=\"xref bibr\" href=\"#ref34\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Siska, Jones, Jeon, Bhak, Kim, Cho, Kim, Lee, Veselovskaya, Balueva, Gallego-Llorente, Hofreiter, Bradley, Eriksson, Pinhasi, Bhak and Manica\u003C\u002Fspan\u003E2017\u003C\u002Fa\u003E; de Barros Damgaard \u003Cem class=\"italic\"\u003Eet al\u003C\u002Fem\u003E. \u003Ca class=\"xref bibr\" href=\"#ref24\"\u003E\u003Cspan class=\"show-for-sr\"\u003EReference Damgaard, Marchi, Rasmussen, Peyrot, Renaud, Korneliussen, Moreno-Mayar, Pedersen, Goldberg, Usmanova, Baimukhanov, Loman, Hedeager, Pedersen, Nielsen, Afanasiev, Akmatov, Aldashev, Alpaslan, Baimbetov, Bazaliiskii, Beisenov, Boldbaatar, Boldgiv, Dorzhu, Ellingvag, Erdenebaatar, Dajani, Dmitriev, Evdokimov, Frei, Gromov, Goryachev, Hakonarson, Hegay, Khachatryan, Khaskhanov, Kitov, Kolbina, Kubatbek, Kukushkin, Kukushkin, Lau, Margaryan, Merkyte, Mertz, Mertz, Mijiddorj, Moiyesev, Mukhtarova, Nurmukhanbetov, Orozbekova, Panyushkina, Pieta, Smrčka, Shevnina, Logvin, Sjogren, Štolcova, Taravella, Tashbaeva, Tkachev, Tulegenov, Voyakin, Yepiskoposyan, Undrakhbold, Varfolomeev, Weber, Wilson Sayres, Kradin, Allentoft, Orlando, Nielsen, Sikora, Heyer, Kristiansen and Willerslev\u003C\u002Fspan\u003E2018\u003C\u002Fa\u003E). We observe that ancient Devil's Gate samples cluster together with present-day Tungusic-speaking populations and Ulchi in the outgroup \u003Cem class=\"italic\"\u003Ef\u003C\u002Fem\u003E \u003Csub\u003E3\u003C\u002Fsub\u003E-statistics, as shown in \u003Ca class=\"xref fig\" href=\"#fig03\"\u003EFigure 3\u003C\u002Fa\u003E. We find a genetic substructure within Tungusic-speaking populations. The North Tungusic Evenki and Even cluster with surrounding Siberian populations other than with Amur Tungusic groups. The substructure has been caused by the fact that North Tungusic Evenki and Even have received gene flow from West Eurasian-related populations.\n\u003C\u002Fp\u003E\u003Csection\u003E\u003Cdiv class=\"fig fig\" data-magellan-destination=\"fig03\" id=\"fig03\"\u003E\u003Cdiv class=\"figure-thumb\"\u003E\u003Cimg src=\"data:image\u002Fgif;base64,R0lGODlhAQABAIAAAMLCwgAAACH5BAAAAAAALAAAAAABAAEAAAICRAEAOw==\" data-src=\"https:\u002F\u002Fstatic.cambridge.org\u002Fbinary\u002Fversion\u002Fid\u002Furn:cambridge.org:id:binary:20200421175645641-0667:S2513843X20000080:S2513843X20000080_fig3.png?pub-status=live\" class=\"aop-lazy-load-image\" data-original-image=\"\u002Fbinary\u002Fversion\u002Fid\u002Furn:cambridge.org:id:binary:20200421175645641-0667:S2513843X20000080:S2513843X20000080_fig3.png\" data-zoomable=\"true\"\u003E\u003C\u002Fdiv\u003E\u003Cdiv class=\"caption\"\u003E\u003Cp class=\"p\"\u003E \u003C\u002Fp\u003E\u003Cp class=\"p\"\u003E\u003Cspan class=\"label\"\u003EFigure 3.\u003C\u002Fspan\u003E Outgroup \u003Cem class=\"italic\"\u003Ef\u003C\u002Fem\u003E \u003Csub\u003E3\u003C\u002Fsub\u003E-statistics of the form \u003Cem class=\"italic\"\u003Ef\u003C\u002Fem\u003E \u003Csub\u003E3\u003C\u002Fsub\u003E (Mbuti; X, Y) to test the genetic drift sharing between ancient Devil's Gate and present-day East Eurasians.\u003C\u002Fp\u003E\u003C\u002Fdiv\u003E\u003C\u002Fdiv\u003E\u003C\u002Fsection\u003E\u003Cp class=\"p\"\u003E Our principal component analysis in \u003Ca class=\"xref fig\" href=\"#fig04\"\u003EFigure 4\u003C\u002Fa\u003E shows that the population structure in East Asia is strongly correlated with geographical and linguistic categories. The ancient Devil's Gate genomes are shown to be genetically most similar to contemporary Tungusic populations, such as Ulchi, Negidal and Nanai at the lower Amur River basin, as well as Nivkh people on the nearby Sakhalin island, documenting a continuous presence of this type of ancestry in the Amur River Basin stretching back at least 8000 years. The principal component analysis plot in \u003Ca class=\"xref fig\" href=\"#fig04\"\u003EFigure 4\u003C\u002Fa\u003E shows that Evenks and Evens are shifted towards West Eurasians and they have about 14–35% West Eurasian-related ancestry estimated by \u003Cem class=\"italic\"\u003EqpAdm\u003C\u002Fem\u003E method (Supplementary Table S2). The admixture time estimated by LD decay suggests a very recent time of around four to six generations ago (Supplementary Table S3). Late Neolithic and Early Bronze Age ancient samples from Ustlda site near the Lake Baikal region show a similar genetic profile with Even people deriving a large amount of Devil's Gate-related ancestry and also having admixture from West Eurasians.\n\u003C\u002Fp\u003E\u003Csection\u003E\u003Cdiv class=\"fig fig\" data-magellan-destination=\"fig04\" id=\"fig04\"\u003E\u003Cdiv class=\"figure-thumb\"\u003E\u003Cimg src=\"data:image\u002Fgif;base64,R0lGODlhAQABAIAAAMLCwgAAACH5BAAAAAAALAAAAAABAAEAAAICRAEAOw==\" data-src=\"https:\u002F\u002Fstatic.cambridge.org\u002Fbinary\u002Fversion\u002Fid\u002Furn:cambridge.org:id:binary:20200421175645641-0667:S2513843X20000080:S2513843X20000080_fig4.png?pub-status=live\" class=\"aop-lazy-load-image\" data-original-image=\"\u002Fbinary\u002Fversion\u002Fid\u002Furn:cambridge.org:id:binary:20200421175645641-0667:S2513843X20000080:S2513843X20000080_fig4.png\" data-zoomable=\"true\"\u003E\u003C\u002Fdiv\u003E\u003Cdiv class=\"caption\"\u003E\u003Cp class=\"p\"\u003E \u003C\u002Fp\u003E\u003Cp class=\"p\"\u003E\u003Cspan class=\"label\"\u003EFigure 4.\u003C\u002Fspan\u003E Principal component analysis projecting ancient Devil's Gate, Ustlda and Jomon samples onto the variation of present-day East Eurasians.\u003C\u002Fp\u003E\u003C\u002Fdiv\u003E\u003C\u002Fdiv\u003E\u003C\u002Fsection\u003E\u003Cp class=\"p\"\u003E The ADMIXTURE analysis in \u003Ca class=\"xref fig\" href=\"#fig05\"\u003EFigure 5\u003C\u002Fa\u003E also confirms the above observation. We observed a large scale of shared components from ancient Devil's Gate samples to present-day Tungusic-speaking populations. We have not found evidence of any West Eurasian-related admixture in ancient Devil's Gate samples. In contrast, the Baikal Ustlda hunter-gatherers (~4700 years ago) were reported to largely have a Devil's Gate-related component, but also have West Eurasian-related ancestry as is shown in the ADMIXTURE plot.\n\u003C\u002Fp\u003E\u003Csection\u003E\u003Cdiv class=\"fig fig\" data-magellan-destination=\"fig05\" id=\"fig05\"\u003E\u003Cdiv class=\"figure-thumb\"\u003E\u003Cimg src=\"data:image\u002Fgif;base64,R0lGODlhAQABAIAAAMLCwgAAACH5BAAAAAAALAAAAAABAAEAAAICRAEAOw==\" data-src=\"https:\u002F\u002Fstatic.cambridge.org\u002Fbinary\u002Fversion\u002Fid\u002Furn:cambridge.org:id:binary:20200421175645641-0667:S2513843X20000080:S2513843X20000080_fig5.png?pub-status=live\" class=\"aop-lazy-load-image\" data-original-image=\"\u002Fbinary\u002Fversion\u002Fid\u002Furn:cambridge.org:id:binary:20200421175645641-0667:S2513843X20000080:S2513843X20000080_fig5.png\" data-zoomable=\"true\"\u003E\u003C\u002Fdiv\u003E\u003Cdiv class=\"caption\"\u003E\u003Cp class=\"p\"\u003E \u003C\u002Fp\u003E\u003Cp class=\"p\"\u003E\u003Cspan class=\"label\"\u003EFigure 5.\u003C\u002Fspan\u003E Admixture plot from \u003Cem class=\"italic\"\u003EK\u003C\u002Fem\u003E = 2 to \u003Cem class=\"italic\"\u003EK\u003C\u002Fem\u003E = 12.\u003C\u002Fp\u003E\u003C\u002Fdiv\u003E\u003C\u002Fdiv\u003E\u003C\u002Fsection\u003E\u003Cp class=\"p\"\u003E Therefore, our genetic findings suggest that the Tungusic-related Amur ancestry distributed in a vast region from the Russian Far East to Lake Baikal. The ancient Devil's Gate population seems to be an unadmixed form of this type of ancestry, which tends to support the larger Amur River region as the ancestral homeland for present-day Tungusic-speaking populations.\u003C\u002Fp\u003E\u003C\u002Fdiv\u003E\u003C\u002Fdiv\u003E\u003Cdiv class=\"sec discussion\" data-magellan-destination=\"sec4\" id=\"sec4\"\u003E\u003Ch2 class=\"A\"\u003E Discussion\u003C\u002Fh2\u003E\u003Cp class=\"p\"\u003E Aligning the evidence, we find that our linguistic and genetic results converge in outlining the broader Amur River region as the ancestral homeland of the speakers of Proto-Tungusic, but our archaeolinguistic results more specifically point to the area around Lake Khanka in the southern part of the Primorye. The Amur-like genetic profile displays a deep history, going back at least 8000 years and covering an extensive region from Baikal and Liao River in the west to the Sea of Japan in the East. Our archaeolinguistic results suggest a dispersal route for agriculture and language from the West Liao River Region to the Southern Primorye around 3300 BCE. As the proposed farming\u002Flanguage dispersal falls within the geographical and chronological reach of the Amur-like genetic profile, we would not expect to observe traces of genetic admixture even if population movement was involved, because the admixing of two similar Amur-like genomes would not be perceivable. This explains why in the southern Primorye both Tungusic-speaking populations and non-Tungusic speaking populations such as the Nivkh are genetically similar and continuous with the ancient Devil's Gate genome in the region. Bringing the archaeolinguistic and genetic evidence together thus makes a case for imperceptible genetic admixture and indicates that the area around Lake Khanka in the southern part of the Primorye is the most plausible homeland for the ancestral speakers of Tungusic.\u003C\u002Fp\u003E\u003C\u002Fdiv\u003E\u003C\u002Fdiv\u003E\u003Cdiv class=\"back\"\u003E\u003Cdiv class=\"notes supplementary-material\" data-magellan-destination=\"nts1\" id=\"nts1\"\u003E\u003Ch2 class=\"A\"\u003ESupplementary material\u003C\u002Fh2\u003E\u003Cp class=\"p\"\u003E The supplementary material for this article can be found at \u003Ca class=\"uri\" href=\"https:\u002F\u002Fdoi.org\u002F10.1017\u002Fehs.2020.8\"\u003Ehttps:\u002F\u002Fdoi.org\u002F10.1017\u002Fehs.2020.8\u003C\u002Fa\u003E.\u003C\u002Fp\u003E\u003C\u002Fdiv\u003E \u003Cdiv class=\"notes other\" data-magellan-destination=\"nts2\" id=\"nts2\"\u003E\u003Ch2 class=\"A\"\u003EAuthor contributions\u003C\u002Fh2\u003E\u003Cp class=\"p\"\u003E MR and CCW conceived and designed the study. MR and CCW conducted data gathering. MR performed archaeolinguistic and CCW genetic analyses. MR and CCW wrote the article.\u003C\u002Fp\u003E\u003C\u002Fdiv\u003E \u003Cdiv class=\"notes other\" data-magellan-destination=\"nts3\" id=\"nts3\"\u003E\u003Ch2 class=\"A\"\u003EFinancial support\u003C\u002Fh2\u003E\u003Cp class=\"p\"\u003E The research leading to these results has received funding from the European Research Council (ERC) under the European Union's Horizon 2020 research and innovation program (grant agreement no. 646612) granted to Martine Robbeets. Chuan-Chao Wang was supported by the National Natural Science Foundation of China (31801040), Nanqiang Outstanding Young Talents Program of Xiamen University (X2123302) and Fundamental Research Funds for the Central Universities (ZK1144).\u003C\u002Fp\u003E\u003C\u002Fdiv\u003E \u003Cdiv class=\"notes other\" data-magellan-destination=\"nts4\" id=\"nts4\"\u003E\u003Ch2 class=\"A\"\u003EData availibility statement\u003C\u002Fh2\u003E\u003Cp class=\"p\"\u003E The linguistic data underlying this article are made available in the Supplementary Material (1). The genetic data are accessible through the following URL: \u003Ca class=\"uri\" href=\"https:\u002F\u002Freich.hms.harvard.edu\u002Fdownloadable-genotypes-present-day-and-ancient-dna-data-compiled-published-papers\"\u003Ehttps:\u002F\u002Freich.hms.harvard.edu\u002Fdownloadable-genotypes-present-day-and-ancient-dna-data-compiled-published-papers\u003C\u002Fa\u003E\u003C\u002Fp\u003E\u003C\u002Fdiv\u003E \u003C\u002Fdiv\u003E\u003C\u002Fdiv\u003E",tableOfContent:[{level:o,current:b,title:"\u003Cdiv class=\"toc-title\"\u003EIntroduction\u003C\u002Fdiv\u003E",url:"sec1"},{level:o,current:b,title:"\u003Cdiv class=\"toc-title\"\u003EMethods\u003C\u002Fdiv\u003E",url:"sec2"},{level:o,current:b,title:"\u003Cdiv class=\"toc-title\"\u003EResults\u003C\u002Fdiv\u003E",url:"sec3"},{level:o,current:b,title:"\u003Cdiv class=\"toc-title\"\u003EDiscussion\u003C\u002Fdiv\u003E",url:"sec4"}],footnotes:[],fulltextNotes:[],references:[{id:"ref1",displayNumber:a,existInContent:c,content:"\u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EAlexander\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EDH\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ENovembre\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EJ\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E and \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ELange\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EK\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E (\u003Cspan class=\"year\"\u003E2009\u003C\u002Fspan\u003E) \u003Cspan class=\"article-title\"\u003EFast model-based estimation of ancestry in unrelated individuals\u003C\u002Fspan\u003E. \u003Cspan class=\"source\"\u003EGenome Research\u003C\u002Fspan\u003E \u003Cspan class=\"volume\"\u003E19\u003C\u002Fspan\u003E(\u003Cspan class=\"issue\"\u003E9\u003C\u002Fspan\u003E), \u003Cspan class=\"fpage\"\u003E1655\u003C\u002Fspan\u003E–\u003Cspan class=\"lpage\"\u003E1664\u003C\u002Fspan\u003E.\u003Ca class='ref-link' target='_blank' aria-label='CrossRef link for Fast model-based estimation of ancestry in unrelated individuals' href=https:\u002F\u002Fdx.doi.org\u002F10.1101\u002Fgr.094052.109\u003ECrossRef\u003C\u002Fa\u003E\u003Ca class='ref-link' target='_blank' aria-label='Google Scholar link for Fast model-based estimation of ancestry in unrelated individuals' href=https:\u002F\u002Fscholar.google.com\u002Fscholar_lookup?title=Fast+model-based+estimation+of+ancestry+in+unrelated+individuals&author=Alexander+DH&author=Novembre+J&author=Lange+K&publication+year=2009&journal=Genome+Research&volume=19&doi=10.1101%2Fgr.094052.109&pages=1655-1664\u003EGoogle Scholar\u003C\u002Fa\u003E\u003Ca class='ref-link' target='_blank' aria-label='PubMed link for Fast model-based estimation of ancestry in unrelated individuals' href=https:\u002F\u002Fwww.ncbi.nlm.nih.gov\u002Fpubmed\u002F19648217\u003EPubMed\u003C\u002Fa\u003E",item:[{googleScholarLink:"https:\u002F\u002Fscholar.google.com\u002Fscholar_lookup?title=Fast+model-based+estimation+of+ancestry+in+unrelated+individuals&author=Alexander+DH&author=Novembre+J&author=Lange+K&publication+year=2009&journal=Genome+Research&volume=19&doi=10.1101%2Fgr.094052.109&pages=1655-1664",openUrlParams:{genre:g,atitle:X,jtitle:Y,title:Y,volume:"19",artnum:"5e9fc31112658c0a2bac0e79",spage:"1655",epage:"1664",date:Z,sid:e,aulast:a,aufirst:a,doi:_,au:a},innerRefId:"cite1",title:X,pubMedId:"19648217",doi:_,crossRefLink:"https:\u002F\u002Fdx.doi.org\u002F10.1101\u002Fgr.094052.109",pubMedLink:"https:\u002F\u002Fwww.ncbi.nlm.nih.gov\u002Fpubmed\u002F19648217"}]},{id:"ref2",displayNumber:a,existInContent:c,content:"\u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EBenzing\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EJ\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E (\u003Cspan class=\"year\"\u003E1955\u003C\u002Fspan\u003E) \u003Cspan class=\"chapter-title\"\u003EDie tungusischen Sprachen: Versuch einer vergleichenden Grammatik\u003C\u002Fspan\u003E. 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class=\"given-names\"\u003EVI\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E (\u003Cspan class=\"year\"\u003E1949\u003C\u002Fspan\u003E) \u003Cspan class=\"source\"\u003ESravnitel'naja fonetika tunguso-man'chzhurskikh jazykov\u003C\u002Fspan\u003E. [Comparative phonetics of the Tungus-Manchu languages]. \u003Cspan class=\"publisher-loc\"\u003ELeningrad\u003C\u002Fspan\u003E: \u003Cspan class=\"publisher-name\"\u003EUchpedgiz\u003C\u002Fspan\u003E.\u003Ca class='ref-link' target='_blank' aria-label='Google Scholar link for Sravnitel'naja fonetika tunguso-man'chzhurskikh jazykov' href=https:\u002F\u002Fscholar.google.com\u002Fscholar_lookup?title=Sravnitel'naja+fonetika+tunguso-man'chzhurskikh+jazykov&author=Cincius+VI&publication+year=1949\u003EGoogle Scholar\u003C\u002Fa\u003E",item:[{googleScholarLink:"https:\u002F\u002Fscholar.google.com\u002Fscholar_lookup?title=Sravnitel'naja+fonetika+tunguso-man'chzhurskikh+jazykov&author=Cincius+VI&publication+year=1949",openUrlParams:{genre:i,btitle:w,title:w,aulast:a,aufirst:a,au:a,pub:"Uchpedgiz",date:"1949",tpages:f,doi:a,sid:e},innerRefId:"cite3",title:w,doi:a,crossRefLink:a,pubMedLink:a}]},{id:"ref24",displayNumber:a,existInContent:c,content:"\u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EDamgaard\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EPB\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EMarchi\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EN\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ERasmussen\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ES\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EPeyrot\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EM\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ERenaud\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EG\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EKorneliussen\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ET\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EMoreno-Mayar\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EJV\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EPedersen\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EMW\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EGoldberg\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EA\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EUsmanova\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EE\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EBaimukhanov\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EN\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ELoman\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EV\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EHedeager\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EL\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EPedersen\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EAG\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ENielsen\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EK\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EAfanasiev\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EG\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EAkmatov\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EK\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EAldashev\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EA\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EAlpaslan\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EA\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EBaimbetov\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EG\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EBazaliiskii\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EVI\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EBeisenov\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EA\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EBoldbaatar\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EB\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EBoldgiv\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EB\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EDorzhu\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EC\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EEllingvag\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ES\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EErdenebaatar\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ED\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EDajani\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ER\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EDmitriev\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EE\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EEvdokimov\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EV\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EFrei\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EKM\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EGromov\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EA\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EGoryachev\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EA\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EHakonarson\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EH\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EHegay\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ET\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EKhachatryan\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EZ\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EKhaskhanov\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ER\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EKitov\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EE\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EKolbina\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EA\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EKubatbek\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ET\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EKukushkin\u003C\u002Fspan\u003E, \u003Cspan 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class=\"given-names\"\u003EB\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EOrozbekova\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EZ\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EPanyushkina\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EI\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EPieta\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EK\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ESmrčka\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EV\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EShevnina\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EI\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ELogvin\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EA\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ESjogren\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EKG\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EŠtolcova\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ET\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ETaravella\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EAM\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ETashbaeva\u003C\u002Fspan\u003E, \u003Cspan 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class=\"given-names\"\u003ES\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EVarfolomeev\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EV\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EWeber\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EA\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EWilson Sayres\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EMA\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EKradin\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EN\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EAllentoft\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EME\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EOrlando\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EL\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ENielsen\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ER\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ESikora\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EM\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EHeyer\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EE\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EKristiansen\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EK\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E and \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EWillerslev\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EE\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E (\u003Cspan class=\"year\"\u003E2018\u003C\u002Fspan\u003E) \u003Cspan class=\"article-title\"\u003E137 ancient human genomes from across the Eurasian steppes\u003C\u002Fspan\u003E. \u003Cspan class=\"source\"\u003ENature\u003C\u002Fspan\u003E \u003Cspan class=\"volume\"\u003E557\u003C\u002Fspan\u003E(\u003Cspan class=\"issue\"\u003E7705\u003C\u002Fspan\u003E), \u003Cspan class=\"fpage\"\u003E369\u003C\u002Fspan\u003E–\u003Cspan class=\"lpage\"\u003E374\u003C\u002Fspan\u003E.\u003Ca class='ref-link' target='_blank' aria-label='CrossRef link for 137 ancient human genomes from across the Eurasian steppes' href=https:\u002F\u002Fdx.doi.org\u002F10.1038\u002Fs41586-018-0094-2\u003ECrossRef\u003C\u002Fa\u003E\u003Ca class='ref-link' target='_blank' aria-label='Google Scholar link for 137 ancient human genomes from across the Eurasian steppes' href=https:\u002F\u002Fscholar.google.com\u002Fscholar_lookup?title=137+ancient+human+genomes+from+across+the+Eurasian+steppes&author=Damgaard+PB&author=Marchi+N&author=Rasmussen+S&author=Peyrot+M&author=Renaud+G&author=Korneliussen+T&author=Moreno-Mayar+JV&author=Pedersen+MW&author=Goldberg+A&author=Usmanova+E&author=Baimukhanov+N&author=Loman+V&author=Hedeager+L&author=Pedersen+AG&author=Nielsen+K&author=Afanasiev+G&author=Akmatov+K&author=Aldashev+A&author=Alpaslan+A&author=Baimbetov+G&author=Bazaliiskii+VI&author=Beisenov+A&author=Boldbaatar+B&author=Boldgiv+B&author=Dorzhu+C&author=Ellingvag+S&author=Erdenebaatar+D&author=Dajani+R&author=Dmitriev+E&author=Evdokimov+V&author=Frei+KM&author=Gromov+A&author=Goryachev+A&author=Hakonarson+H&author=Hegay+T&author=Khachatryan+Z&author=Khaskhanov+R&author=Kitov+E&author=Kolbina+A&author=Kubatbek+T&author=Kukushkin+A&author=Kukushkin+I&author=Lau+N&author=Margaryan+A&author=Merkyte+I&author=Mertz+IV&author=Mertz+VK&author=Mijiddorj+E&author=Moiyesev+V&author=Mukhtarova+G&author=Nurmukhanbetov+B&author=Orozbekova+Z&author=Panyushkina+I&author=Pieta+K&author=Smr%C4%8Dka+V&author=Shevnina+I&author=Logvin+A&author=Sjogren+KG&author=%C5%A0tolcova+T&author=Taravella+AM&author=Tashbaeva+K&author=Tkachev+A&author=Tulegenov+T&author=Voyakin+D&author=Yepiskoposyan+L&author=Undrakhbold+S&author=Varfolomeev+V&author=Weber+A&author=Wilson+Sayres+MA&author=Kradin+N&author=Allentoft+ME&author=Orlando+L&author=Nielsen+R&author=Sikora+M&author=Heyer+E&author=Kristiansen+K&author=Willerslev+E&publication+year=2018&journal=Nature&volume=557&doi=10.1038%2Fs41586-018-0094-2&pages=369-374\u003EGoogle Scholar\u003C\u002Fa\u003E\u003Ca class='ref-link' target='_blank' aria-label='PubMed link for 137 ancient human genomes from across the Eurasian steppes' href=https:\u002F\u002Fwww.ncbi.nlm.nih.gov\u002Fpubmed\u002F29743675\u003EPubMed\u003C\u002Fa\u003E",item:[{googleScholarLink:"https:\u002F\u002Fscholar.google.com\u002Fscholar_lookup?title=137+ancient+human+genomes+from+across+the+Eurasian+steppes&author=Damgaard+PB&author=Marchi+N&author=Rasmussen+S&author=Peyrot+M&author=Renaud+G&author=Korneliussen+T&author=Moreno-Mayar+JV&author=Pedersen+MW&author=Goldberg+A&author=Usmanova+E&author=Baimukhanov+N&author=Loman+V&author=Hedeager+L&author=Pedersen+AG&author=Nielsen+K&author=Afanasiev+G&author=Akmatov+K&author=Aldashev+A&author=Alpaslan+A&author=Baimbetov+G&author=Bazaliiskii+VI&author=Beisenov+A&author=Boldbaatar+B&author=Boldgiv+B&author=Dorzhu+C&author=Ellingvag+S&author=Erdenebaatar+D&author=Dajani+R&author=Dmitriev+E&author=Evdokimov+V&author=Frei+KM&author=Gromov+A&author=Goryachev+A&author=Hakonarson+H&author=Hegay+T&author=Khachatryan+Z&author=Khaskhanov+R&author=Kitov+E&author=Kolbina+A&author=Kubatbek+T&author=Kukushkin+A&author=Kukushkin+I&author=Lau+N&author=Margaryan+A&author=Merkyte+I&author=Mertz+IV&author=Mertz+VK&author=Mijiddorj+E&author=Moiyesev+V&author=Mukhtarova+G&author=Nurmukhanbetov+B&author=Orozbekova+Z&author=Panyushkina+I&author=Pieta+K&author=Smr%C4%8Dka+V&author=Shevnina+I&author=Logvin+A&author=Sjogren+KG&author=%C5%A0tolcova+T&author=Taravella+AM&author=Tashbaeva+K&author=Tkachev+A&author=Tulegenov+T&author=Voyakin+D&author=Yepiskoposyan+L&author=Undrakhbold+S&author=Varfolomeev+V&author=Weber+A&author=Wilson+Sayres+MA&author=Kradin+N&author=Allentoft+ME&author=Orlando+L&author=Nielsen+R&author=Sikora+M&author=Heyer+E&author=Kristiansen+K&author=Willerslev+E&publication+year=2018&journal=Nature&volume=557&doi=10.1038%2Fs41586-018-0094-2&pages=369-374",openUrlParams:{genre:g,atitle:ad,jtitle:p,title:p,volume:"557",artnum:"5e9fc31112658c0a2bac0e7d",spage:"369",epage:"374",date:ae,sid:e,aulast:a,aufirst:a,doi:af,au:a},innerRefId:"cite24",title:ad,pubMedId:"29743675",doi:af,crossRefLink:"https:\u002F\u002Fdx.doi.org\u002F10.1038\u002Fs41586-018-0094-2",pubMedLink:"https:\u002F\u002Fwww.ncbi.nlm.nih.gov\u002Fpubmed\u002F29743675"}]},{id:"ref4",displayNumber:a,existInContent:c,content:"\u003Cspan 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[Priamur'e in antiquity. 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class=\"surname\"\u003EBalanovsky\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EO\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ELukianova\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EE\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EKahbatkyzy\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EN\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EFlegontov\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EP\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EZaporozhchenko\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EV\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan 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class=\"given-names\"\u003EO\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EIdrisov\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EE\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EKamenshchikova\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EE\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EEvseeva\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EI\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EMetspalu\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EM\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EOutram\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EAK\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ERobbeets\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EM\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EDjansugurova\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EL\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EBalanovska\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EE\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ESchiffels\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ES\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EHaak\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EW\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EReich\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ED\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E and \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EKrause\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EJ\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E (\u003Cspan class=\"year\"\u003E2019\u003C\u002Fspan\u003E) \u003Cspan class=\"article-title\"\u003EThe genetic history of admixture across inner Eurasia\u003C\u002Fspan\u003E. \u003Cspan class=\"source\"\u003ENature Ecology and Evolution\u003C\u002Fspan\u003E \u003Cspan class=\"volume\"\u003E3\u003C\u002Fspan\u003E(\u003Cspan class=\"issue\"\u003E6\u003C\u002Fspan\u003E), \u003Cspan class=\"fpage\"\u003E966\u003C\u002Fspan\u003E–\u003Cspan class=\"lpage\"\u003E976\u003C\u002Fspan\u003E.\u003Ca class='ref-link' target='_blank' 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href=https:\u002F\u002Fwww.ncbi.nlm.nih.gov\u002Fpubmed\u002F31036896\u003EPubMed\u003C\u002Fa\u003E",item:[{googleScholarLink:"https:\u002F\u002Fscholar.google.com\u002Fscholar_lookup?title=The+genetic+history+of+admixture+across+inner+Eurasia&author=Jeong+C&author=Balanovsky+O&author=Lukianova+E&author=Kahbatkyzy+N&author=Flegontov+P&author=Zaporozhchenko+V&author=Immel+A&author=Wang+CC&author=Ixan+O&author=Khussainova+E&author=Bekmanov+B&author=Zaibert+V&author=Lavryashina+M&author=Pocheshkhova+E&author=Yusupov+Y&author=Agdzhoyan+A&author=Koshel+S&author=Bukin+A&author=Nymadawa+P&author=Turdikulova+S&author=Dalimova+D&author=Churnosov+M&author=Skhalyakho+R&author=Daragan+D&author=Bogunov+Y&author=Bogunova+A&author=Shtrunov+A&author=Dubova+N&author=Zhabagin+M&author=Yepiskoposyan+L&author=Churakov+V&author=Pislegin+N&author=Damba+L&author=Saroyants+L&author=Dibirova+K&author=Atramentova+L&author=Utevska+O&author=Idrisov+E&author=Kamenshchikova+E&author=Evseeva+I&author=Metspalu+M&author=Outram+AK&author=Robbeets+M&author=Djansugurova+L&author=Balanovska+E&author=Schiffels+S&author=Haak+W&author=Reich+D&author=Krause+J&publication+year=2019&journal=Nature+Ecology+and+Evolution&volume=3&doi=10.1038%2Fs41559-019-0878-2&pages=966-976",openUrlParams:{genre:g,atitle:ak,jtitle:al,title:al,volume:H,artnum:"5e9fc31112658c0a2bac0e86",spage:"966",epage:"976",date:am,sid:e,aulast:a,aufirst:a,doi:an,au:a},innerRefId:"cite12",title:ak,pubMedId:"31036896",doi:an,crossRefLink:"https:\u002F\u002Fdx.doi.org\u002F10.1038\u002Fs41559-019-0878-2",pubMedLink:"https:\u002F\u002Fwww.ncbi.nlm.nih.gov\u002Fpubmed\u002F31036896"}]},{id:"ref13",displayNumber:a,existInContent:c,content:"\u003Cspan 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class=\"surname\"\u003ESuwa\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EG\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EUeda\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ES\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EYoneda\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EM\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ETajima\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EA\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EShinoda\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EKI\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EInoue\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EI\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ESaitou\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EN\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E (\u003Cspan class=\"year\"\u003E2016\u003C\u002Fspan\u003E) \u003Cspan class=\"article-title\"\u003EA partial nuclear genome of the Jomons who lived 3000 years ago in Fukushima, Japan\u003C\u002Fspan\u003E. \u003Cspan class=\"source\"\u003EJournal of Human Genetics\u003C\u002Fspan\u003E \u003Cspan class=\"volume\"\u003E62\u003C\u002Fspan\u003E(\u003Cspan class=\"issue\"\u003E2\u003C\u002Fspan\u003E), \u003Cspan class=\"fpage\"\u003E213\u003C\u002Fspan\u003E–\u003Cspan class=\"lpage\"\u003E221\u003C\u002Fspan\u003E.\u003Ca class='ref-link' target='_blank' aria-label='CrossRef link for A partial nuclear genome of the Jomons who lived 3000 years ago in Fukushima, Japan' href=https:\u002F\u002Fdx.doi.org\u002F10.1038\u002Fjhg.2016.110\u003ECrossRef\u003C\u002Fa\u003E\u003Ca class='ref-link' target='_blank' aria-label='Google Scholar link for A partial nuclear genome of the Jomons who lived 3000 years ago in Fukushima, Japan' href=https:\u002F\u002Fscholar.google.com\u002Fscholar_lookup?title=A+partial+nuclear+genome+of+the+Jomons+who+lived+3000+years+ago+in+Fukushima%2C+Japan&author=Kanzawa-Kiriyama+H&author=Kryukov+K&author=Jinam+TA&author=Hosomichi+K&author=Saso+A&author=Suwa+G&author=Ueda+S&author=Yoneda+M&author=Tajima+A&author=Shinoda+KI&author=Inoue+I&author=Saitou+N&publication+year=2016&journal=Journal+of+Human+Genetics&volume=62&doi=10.1038%2Fjhg.2016.110&pages=213-221\u003EGoogle Scholar\u003C\u002Fa\u003E\u003Ca class='ref-link' target='_blank' aria-label='PubMed link for A partial nuclear genome of the Jomons who lived 3000 years ago in Fukushima, Japan' href=https:\u002F\u002Fwww.ncbi.nlm.nih.gov\u002Fpubmed\u002F27581845\u003EPubMed\u003C\u002Fa\u003E",item:[{googleScholarLink:"https:\u002F\u002Fscholar.google.com\u002Fscholar_lookup?title=A+partial+nuclear+genome+of+the+Jomons+who+lived+3000+years+ago+in+Fukushima%2C+Japan&author=Kanzawa-Kiriyama+H&author=Kryukov+K&author=Jinam+TA&author=Hosomichi+K&author=Saso+A&author=Suwa+G&author=Ueda+S&author=Yoneda+M&author=Tajima+A&author=Shinoda+KI&author=Inoue+I&author=Saitou+N&publication+year=2016&journal=Journal+of+Human+Genetics&volume=62&doi=10.1038%2Fjhg.2016.110&pages=213-221",openUrlParams:{genre:g,atitle:ao,jtitle:ap,title:ap,volume:"62",artnum:"5e9fc31112658c0a2bac0e87",spage:ag,epage:"221",date:"2016",sid:e,aulast:a,aufirst:a,doi:aq,au:a},innerRefId:"cite13",title:ao,pubMedId:"27581845",doi:aq,crossRefLink:"https:\u002F\u002Fdx.doi.org\u002F10.1038\u002Fjhg.2016.110",pubMedLink:"https:\u002F\u002Fwww.ncbi.nlm.nih.gov\u002Fpubmed\u002F27581845"}]},{id:"ref14",displayNumber:a,existInContent:c,content:"\u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EKormušin\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EIV\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E (\u003Cspan class=\"year\"\u003E1998\u003C\u002Fspan\u003E) \u003Cspan class=\"source\"\u003EUdyhejskij (udegejskij) jazyk\u003C\u002Fspan\u003E. [The Udihe language.] \u003Cspan class=\"publisher-loc\"\u003EMoscow\u003C\u002Fspan\u003E: \u003Cspan class=\"publisher-name\"\u003ENauka\u003C\u002Fspan\u003E.\u003Ca class='ref-link' target='_blank' aria-label='Google Scholar link for Udyhejskij (udegejskij) jazyk' href=https:\u002F\u002Fscholar.google.com\u002Fscholar_lookup?title=Udyhejskij+(udegejskij)+jazyk&author=Kormu%C5%A1in+IV&publication+year=1998\u003EGoogle Scholar\u003C\u002Fa\u003E",item:[{googleScholarLink:"https:\u002F\u002Fscholar.google.com\u002Fscholar_lookup?title=Udyhejskij+(udegejskij)+jazyk&author=Kormu%C5%A1in+IV&publication+year=1998",openUrlParams:{genre:i,btitle:I,title:I,aulast:a,aufirst:a,au:a,pub:y,date:"1998",tpages:f,doi:a,sid:e},innerRefId:"cite14",title:I,doi:a,crossRefLink:a,pubMedLink:a}]},{id:"ref15",displayNumber:a,existInContent:c,content:"\u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EKorovina\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EE\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E (\u003Cspan class=\"year\"\u003E2011\u003C\u002Fspan\u003E) Leksika prirodnogo i kul'turnogo okruženiya v Tunguso-Man’čžurskich jazykach (v istoriko-tipologičeskom osveščenii). [Tungus-Manchu vocabulary related to natural environment and cultural activities (in historical and typological perspective).] Moscow: Russian State University of Humanities, BA dissertation.\u003Ca class='ref-link' target='_blank' aria-label='Google Scholar link for Korovina, E (2011) Leksika prirodnogo i kul'turnogo okruženiya v Tunguso-Man’čžurskich jazykach (v istoriko-tipologičeskom osveščenii). [Tungus-Manchu vocabulary related to natural environment and cultural activities (in historical and typological perspective).] Moscow: Russian State University of Humanities, BA dissertation.' href=https:\u002F\u002Fscholar.google.com\u002Fscholar?q=Korovina,+E+(2011)+Leksika+prirodnogo+i+kul'turnogo+okruženiya+v+Tunguso-Man’čžurskich+jazykach+(v+istoriko-tipologičeskom+osveščenii).+[Tungus-Manchu+vocabulary+related+to+natural+environment+and+cultural+activities+(in+historical+and+typological+perspective).]+Moscow:+Russian+State+University+of+Humanities,+BA+dissertation.\u003EGoogle Scholar\u003C\u002Fa\u003E",item:[{googleScholarLink:"https:\u002F\u002Fscholar.google.com\u002Fscholar?q=Korovina,+E+(2011)+Leksika+prirodnogo+i+kul'turnogo+okruženiya+v+Tunguso-Man’čžurskich+jazykach+(v+istoriko-tipologičeskom+osveščenii).+[Tungus-Manchu+vocabulary+related+to+natural+environment+and+cultural+activities+(in+historical+and+typological+perspective).]+Moscow:+Russian+State+University+of+Humanities,+BA+dissertation.",openUrlParams:{genre:ar,date:D,sid:e,title:f},innerRefId:"cite15",title:"Korovina, E (2011) Leksika prirodnogo i kul'turnogo okruženiya v Tunguso-Man’čžurskich jazykach (v istoriko-tipologičeskom osveščenii). [Tungus-Manchu vocabulary related to natural environment and cultural activities (in historical and typological perspective).] Moscow: Russian State University of Humanities, BA dissertation.",doi:a,crossRefLink:a,pubMedLink:a}]},{id:"ref16",displayNumber:a,existInContent:c,content:"\u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ELazaridis\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EI\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EPatterson\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EN\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EMittnik\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EA\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ERenaud\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EG\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EMallick\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ES\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EKirsanow\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EK\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ESudmant\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EPH\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ESchraiber\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EJG\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ECastellano\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ES\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ELipson\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EM\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EBerger\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EB\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EEconomou\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EC\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EBollongino\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ER\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EFu\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EQ\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EBos\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EKI\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ENordenfelt\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ES\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ELi\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EH\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003Ede Filippo\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EC\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EPrüfer\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EK\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ESawyer\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ES\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EPosth\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EC\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EHaak\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EW\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EHallgren\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EF\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EFornander\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EE\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ERohland\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EN\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EDelsate\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ED\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EFrancken\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EM\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EGuinet\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EJM\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EWahl\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EJ\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EAyodo\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EG\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EBabiker\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EHA\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EBailliet\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EG\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EBalanovska\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EE\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EBalanovsky\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EO\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EBarrantes\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ER\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EBedoya\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EG\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EBen-Ami\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EH\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EBene\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EJ\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EBerrada\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EF\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EBravi\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ECM\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EBrisighelli\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EF\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EBusby\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EGB\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ECali\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EF\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EChurnosov\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EM\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ECole\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EDE\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ECorach\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ED\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EDamba\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EL\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003Evan Driem\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EG\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EDryomov\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ES\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EDugoujon\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EJM\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EFedorova\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ESA\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EGallego Romero\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EI\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EGubina\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EM\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EHammer\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EM\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EHenn\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EBM\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EHervig\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ET\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EHodoglugil\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EU\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EJha\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EAR\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EKarachanak-Yankova\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ES\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EKhusainova\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ER\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EKhusnutdinova\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EE\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EKittles\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ER\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EKivisild\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ET\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EKlitz\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EW\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EKučinskas\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EV\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EKushniarevich\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EA\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ELaredj\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EL\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ELitvinov\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ES\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ELoukidis\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ET\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EMahley\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ERW\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EMelegh\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EB\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EMetspalu\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EE\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EMolina\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EJ\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EMountain\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EJ\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ENäkkäläjärvi\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EK\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ENesheva\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ED\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ENyambo\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ET\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EOsipova\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EL\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EParik\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EJ\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EPlatonov\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EF\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EPosukh\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EO\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ERomano\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EV\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ERothhammer\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EF\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ERudan\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EI\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ERuizbakiev\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ER\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ESahakyan\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EH\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ESajantila\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EA\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ESalas\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EA\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EStarikovskaya\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EEB\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ETarekegn\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EA\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EToncheva\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ED\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ETurdikulova\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ES\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EUktveryte\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EI\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EUtevska\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EO\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EVasquez\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ER\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EVillena\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EM\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EVoevoda\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EM\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EWinkler\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ECA\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EYepiskoposyan\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EL\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EZalloua\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EP\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EZemunik\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ET\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ECooper\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EA\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ECapelli\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EC\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EThomas\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EMG\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ERuiz-Linares\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EA\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ETishkoff\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ESA\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ESingh\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EL\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EThangaraj\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EK\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EVillems\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ER\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EComas\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ED\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ESukernik\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ER\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EMetspalu\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EM\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EMeyer\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EM\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EEichler\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EEE\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EBurger\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EJ\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003ESlatkin\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EM\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EPääbo\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ES\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EKelso\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EJ\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E, \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EReich\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003ED\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E and \u003Cspan class=\"string-name\"\u003E\u003Cspan class=\"surname\"\u003EKrause\u003C\u002Fspan\u003E, \u003Cspan class=\"given-names\"\u003EJ\u003C\u002Fspan\u003E\u003C\u002Fspan\u003E (\u003Cspan class=\"year\"\u003E2014\u003C\u002Fspan\u003E) \u003Cspan class=\"article-title\"\u003EAncient human genomes suggest three ancestral populations for present-day Europeans\u003C\u002Fspan\u003E. \u003Cspan class=\"source\"\u003ENature\u003C\u002Fspan\u003E \u003Cspan class=\"volume\"\u003E513\u003C\u002Fspan\u003E(\u003Cspan class=\"issue\"\u003E7518\u003C\u002Fspan\u003E), \u003Cspan class=\"fpage\"\u003E409\u003C\u002Fspan\u003E–\u003Cspan class=\"lpage\"\u003E413\u003C\u002Fspan\u003E.\u003Ca class='ref-link' target='_blank' aria-label='CrossRef link for Ancient human genomes suggest three ancestral populations for present-day Europeans' href=https:\u002F\u002Fdx.doi.org\u002F10.1038\u002Fnature13673\u003ECrossRef\u003C\u002Fa\u003E\u003Ca class='ref-link' target='_blank' aria-label='Google Scholar link for Ancient human genomes suggest three ancestral populations for present-day Europeans' 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locations for the homeland of the ancestral speakers of Proto-Tungusic.\u003C\u002Fspan\u003E",thumbnailSrc:a_,enlargedSrc:a_,attrib:[]},{contentId:"fig02",label:"Figure 2.",description:"\u003Cspan class=\"p\"\u003EDiversity hotspot of the Tungusic languages under the north–south classification in (a) as compared with the hotspot under the Manchu–Tungusic classification in (b).\u003C\u002Fspan\u003E",thumbnailSrc:a$,enlargedSrc:a$,attrib:[]},{contentId:"tab01",label:"Table 1.",description:"\u003Cspan class=\"p\"\u003EEvaluating the four homeland proposals for Proto-Tungusic using different archaeolinguistic techniques: 0 implausible, 1 plausible\u003C\u002Fspan\u003E",thumbnailSrc:ba,enlargedSrc:ba,attrib:[]},{contentId:"fig03",label:"Figure 3.",description:"\u003Cspan class=\"p\"\u003EOutgroup \u003Cspan class=\"italic\"\u003Ef\u003C\u002Fspan\u003E\u003Cspan class=\"sub\"\u003E3\u003C\u002Fspan\u003E-statistics of the form \u003Cspan class=\"italic\"\u003Ef\u003C\u002Fspan\u003E\u003Cspan class=\"sub\"\u003E3\u003C\u002Fspan\u003E (Mbuti; X, Y) to test the genetic drift sharing between ancient Devil's Gate and present-day East Eurasians.\u003C\u002Fspan\u003E",thumbnailSrc:bb,enlargedSrc:bb,attrib:[]},{contentId:"fig04",label:"Figure 4.",description:"\u003Cspan class=\"p\"\u003EPrincipal component analysis projecting ancient Devil's Gate, Ustlda and Jomon samples onto the variation of present-day East Eurasians.\u003C\u002Fspan\u003E",thumbnailSrc:bc,enlargedSrc:bc,attrib:[]},{contentId:"fig05",label:"Figure 5.",description:"\u003Cspan class=\"p\"\u003EAdmixture plot from \u003Cspan class=\"italic\"\u003EK\u003C\u002Fspan\u003E = 2 to \u003Cspan class=\"italic\"\u003EK\u003C\u002Fspan\u003E = 12.\u003C\u002Fspan\u003E",thumbnailSrc:bd,enlargedSrc:bd,attrib:[]}]},pdf:{standardResolution:{fileUrl:"\u002Fcore\u002Fservices\u002Faop-cambridge-core\u002Fcontent\u002Fview\u002F950020A1799B4D7B71CCC57D04743B3E\u002FS2513843X20000080a.pdf\u002Fthe-homeland-of-proto-tungusic-inferred-from-contemporary-words-and-ancient-genomes.pdf",fileSizeInMb:.56,articleTitle:l,slugTitle:be},highResolution:{fileUrl:"\u002Fcore\u002Fservices\u002Faop-cambridge-core\u002Fcontent\u002Fview\u002F950020A1799B4D7B71CCC57D04743B3E\u002FS2513843X20000080a_hi.pdf\u002Fthe-homeland-of-proto-tungusic-inferred-from-contemporary-words-and-ancient-genomes.pdf",fileSizeInMb:.82,articleTitle:l,slugTitle:be},media:f},classification:[],supplementaryMaterials:[{title:"Wang and Robbeets supplementary material",description:"\u003Cp\u003ETables 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Advances in Tungusic Linguistics","The Oxford Guide to the Transeurasian Languages","The homeland of Proto-Tungusic inferred from contemporary words and ancient genomes","Evolutionary Human Sciences","2020",1,"Nature","Otto Harrassowitz","Science Advances","Genetics","2","Abhandlungen der geistes- und sozialwissenschaftlichen Klasse","2015","Sravnitel'naja fonetika tunguso-man'chzhurskikh jazykov","Priamur'ye v drevnosti (do nachala nashej ery)","Nauka","Beiträge zur nordasiatischen Kulturgeschichte","Proceedings of the First International Conference on Manchu-Tungus Studies. 2: Trends in Tungusic and Siberian Linguistics","The Routledge Handbook of Historical Linguistics","Uralo-Altaistika","2011","Manchuria: an ethnic history","2012","5","3","Udyhejskij (udegejskij) jazyk","Tungusologie. Handbuch der Orientalistik","Les origines indo-européennes ou les aryas primitifs: essai de paléontologie linguistique (première partie, seconde partie)","Diachrony of verb morphology. Japanese and the Transeurasian languages","Language Dispersal Beyond Farming","Time Perspective in Aboriginal American Culture: A Study in Method","From Foragers to Farmers: Papers in Honour of Gordon C. Hillman","Mlado-pis'mennye jazyki narodov SSSR","western",0,"F9A99C4602D4F4A5277A9D3A04AE7353","evolutionary-human-sciences","\u002Fcore\u002Fjournals\u002Fevolutionary-human-sciences","Volume 2","\u002Fcore\u002Fjournals\u002Fevolutionary-human-sciences\u002Fvolume\u002FF2345807349E632AB7170A2CDEAF9DF5","Fast model-based estimation of ancestry in unrelated individuals","Genome Research","2009","10.1101\u002Fgr.094052.109","Second-generation PLINK: rising to the challenge of larger and richer datasets","Gigascience","7","10.1186\u002Fs13742-015-0047-8","137 ancient human genomes from across the Eurasian steppes","2018","10.1038\u002Fs41586-018-0094-2","213","Automated dating of the world's language families based on lexical similarity","Current Anthropology","10.1086\u002F662127","The genetic history of admixture across inner Eurasia","Nature Ecology and Evolution","2019","10.1038\u002Fs41559-019-0878-2","A partial nuclear genome of the Jomons who lived 3000 years ago in Fukushima, Japan","Journal of Human Genetics","10.1038\u002Fjhg.2016.110","unknown","Ancient human genomes suggest three ancestral populations for present-day Europeans","10.1038\u002Fnature13673","Discontinuous spread of millet agriculture in eastern Asia and prehistoric population dynamics","10.1126\u002Fsciadv.aax6225","Millet agriculture dispersed from Northeast China to the Russian Far East: integrating archaeology, genetics, and linguistics","Archaeological Research in Asia","10.1016\u002Fj.ara.2020.100177","Inferring admixture histories of human populations using linkage disequilibrium","10.1534\u002Fgenetics.112.147330","Population structure and eigenanalysis","PLoS Genetics","10.1371\u002Fjournal.pgen.0020190","Ancient admixture in human history","10.1534\u002Fgenetics.112.145037","10.1515\u002F9783110399943","2017","10.1075\u002Fz.215.05rob","Bayesian phylolinguistics reveals the internal structure of the Transeurasian family","Journal of Linguistic Evolution","Oxford University Press","10.4095\u002F103486","Sachen und Wörter","Revue Internationale d'Ethnologie et de Linguistique","Genome-wide data from two early Neolithic East Asian individuals dating to 7700 years ago","10.1126\u002Fsciadv.1601877","K voprosu o klassifikacii tunguso-man’čžurskih jazykov. [About the classification of Tungus–Manchu languages.]","Voprosy jazykoznanija","Towards a new classification of Tungusic languages","Eurasian Studies Yearbook","Genetic evidences are against a common origin of the Altaic populations","Communication on Contemporary Anthropology","10.4236\u002Fcoca.2011.51038","Homelands of the world's language families. 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