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<!DOCTYPE html> <html lang="en" dir="ltr" xmlns="http://www.w3.org/1999/xhtml" xmlns:mml="http://www.w3.org/1998/Math/MathML"> <head prefix="og: http://ogp.me/ns# article: http://ogp.me/ns/article# book: http://ogp.me/ns/book#" >  <link rel="dns-prefetch" href="//d33xdlntwy0kbs.cloudfront.net" /> <link rel="dns-prefetch" href="//www.google.com" /> <link rel="dns-prefetch" href="//scholar.google.com" /> <link rel="dns-prefetch" href="//www.googletagmanager.com" /> <meta http-equiv="Content-Type" content="text/html; charset=utf-8" /> <link rel="shortcut icon" href="https://www.biorxiv.org/sites/default/files/images/favicon.ico" type="image/vnd.microsoft.icon" /> <meta name="viewport" content="width=device-width, initial-scale=1" /> <link rel="alternate" type="application/pdf" title="Full Text (PDF)" href="/content/10.1101/2024.03.13.584607v1.full.pdf" /> <link rel="alternate" type="text/plain" title="Full Text (Plain)" href="/content/10.1101/2024.03.13.584607v1.full.txt" /> <meta name="type" content="article" /> <meta name="category" content="article" /> <meta name="HW.identifier" content="/biorxiv/early/2024/03/14/2024.03.13.584607.atom" /> <meta name="HW.pisa" content="biorxiv;2024.03.13.584607v1" /> <meta name="DC.Format" content="text/html" /> <meta name="DC.Language" content="en" /> <meta name="DC.Title" content="Steppe Ancestry in western Eurasia and the spread of the Germanic Languages" /> <meta name="DC.Identifier" content="10.1101/2024.03.13.584607" /> <meta name="DC.Date" content="2024-03-14" /> <meta name="DC.Publisher" content="Cold Spring Harbor Laboratory" /> <meta name="DC.Rights" content="© 2024, Posted by Cold Spring Harbor Laboratory. This pre-print is available under a Creative Commons License (Attribution-NonCommercial-NoDerivs 4.0 International), CC BY-NC-ND 4.0, as described at http://creativecommons.org/licenses/by-nc-nd/4.0/" /> <meta name="DC.AccessRights" content="restricted" /> <meta name="DC.Description" content="Germanic-speaking populations historically form an integral component of the North and Northwest European cultural configuration. According to linguistic consensus, the common ancestor of the Germanic languages, which include German, English, Frisian, Dutch as well as the Nordic languages, was spoken in Northern Europe during the Pre-Roman Iron Age. However, important questions remain concerning the earlier Bronze Age distribution of this Indo-European language branch in Scandinavia as well as the driving factors behind its Late Iron Age diversification and expansion across the European continent. A key difficulty in addressing these questions are the existence of striking differences in the interpretation of the archaeological record, leading to various hypotheses of correlations with linguistic dispersals and changes in material culture. Moreover, these interpretations have been difficult to assess using genomics due to limited ancient genomes and the difficulty in differentiating closely related populations. Here we integrate multidisciplinary evidence from population genomics, historical sources, archaeology and linguistics to offer a fully revised model for the origins and spread of Germanic languages and for the formation of the genomic ancestry of Germanic-speaking northern European populations, while acknowledging that coordinating archaeology, linguistics and genetics is complex and potentially controversial. We sequenced 710 ancient human genomes from western Eurasia and analysed them together with 3,940 published genomes suitable for imputing diploid genotypes. We find evidence of a previously unknown, large-scale Bronze Age migration within Scandinavia, originating in the east and becoming widespread to the west and south, thus providing a new potential driving factor for the expansion of the Germanic speech community. This East Scandinavian genetic cluster is first seen 800 years after the arrival of the Corded Ware Culture, the first Steppe-related population to emerge in Northern Europe, opening a new scenario implying a Late rather than an Middle Neolithic arrival of the Germanic language group in Scandinavia. Moreover, the non-local Hunter-Gatherer ancestry of this East Scandinavian cluster is indicative of a cross-Baltic maritime rather than a southern Scandinavian land-based entry. Later in the Iron Age around 1700 BP, we find a southward push of admixed Eastern and Southern Scandinavians into areas including Germany and the Netherlands, previously associated with Celtic speakers, mixing with local populations from the Eastern North Sea coast. During the Migration Period (1575-1200 BP), we find evidence of this structured, admixed Southern Scandinavian population representing the Western Germanic Anglo-Saxon migrations into Britain and Langobards into southern Europe. During the Migration Period, we detect a previously unknown northward migration back into Southern Scandinavia, partly replacing earlier inhabitants and forming the North Germanic-speaking Viking-Age populations of Denmark and southern Sweden, corresponding with historically attested Danes. However, the origin and character of these major changes in Scandinavia before the Viking Age remain contested. In contrast to these Western and Northern Germanic-speaking populations, we find the Wielbark population from Poland to be primarily of Eastern Scandinavian ancestry, supporting a Swedish origin for East Germanic groups. In contrast, the later cultural descendants, the Ostrogoths and Visigoths are predominantly of Southern European ancestry implying the adoption of Gothic culture. Together, these results highlight the use of archaeology, linguistics and genetics as distinct but complementary lines of evidence. ### Competing Interest Statement The authors have declared no competing interest." /> <meta name="DC.Contributor" content="Hugh McColl" /> <meta name="DC.Contributor" content="Guus Kroonen" /> <meta name="DC.Contributor" content="J. Víctor Moreno-Mayar" /> <meta name="DC.Contributor" content="Frederik Valeur Seersholm" /> <meta name="DC.Contributor" content="Gabriele Scorrano" /> <meta name="DC.Contributor" content="Thomaz Pinotti" /> <meta name="DC.Contributor" content="Tharsika Vimala" /> <meta name="DC.Contributor" content="Søren M. Sindbæk" /> <meta name="DC.Contributor" content="Per Ethelberg" /> <meta name="DC.Contributor" content="Ralph Fyfe" /> <meta name="DC.Contributor" content="Marie-José Gaillard" /> <meta name="DC.Contributor" content="Hanne M. Ellegård Larsen" /> <meta name="DC.Contributor" content="Morten Fischer Mortensen" /> <meta name="DC.Contributor" content="Fabrice Demeter" /> <meta name="DC.Contributor" content="Marie Louise S. Jørkov" /> <meta name="DC.Contributor" content="Sophie Bergerbrant" /> <meta name="DC.Contributor" content="Peter de Barros Damgaard" /> <meta name="DC.Contributor" content="Morten E. Allentoft" /> <meta name="DC.Contributor" content="Lasse Vinner" /> <meta name="DC.Contributor" content="Charleen Gaunitz" /> <meta name="DC.Contributor" content="Abigail Ramsøe" /> <meta name="DC.Contributor" content="Isin Altinkaya" /> <meta name="DC.Contributor" content="Rasmus Amund Henriksen" /> <meta name="DC.Contributor" content="Evan K. Irving-Pease" /> <meta name="DC.Contributor" content="Serena Sabatini" /> <meta name="DC.Contributor" content="Anders Fischer" /> <meta name="DC.Contributor" content="William Barrie" /> <meta name="DC.Contributor" content="Andrés Ingason" /> <meta name="DC.Contributor" content="Anders Rosengren" /> <meta name="DC.Contributor" content="Andrew Vaughn" /> <meta name="DC.Contributor" content="Jialu Cao" /> <meta name="DC.Contributor" content="Jacqueline Staring" /> <meta name="DC.Contributor" content="Jesper Stenderup" /> <meta name="DC.Contributor" content="Fulya Eylem Yediay" /> <meta name="DC.Contributor" content="Torbjörn Ahlström" /> <meta name="DC.Contributor" content="Sofie Laurine Albris" /> <meta name="DC.Contributor" content="Biyaslan Atabiev" /> <meta name="DC.Contributor" content="Pernille Bangsgaard" /> <meta name="DC.Contributor" content="Maria Giovanna Belcastro" /> <meta name="DC.Contributor" content="Nick Card" /> <meta name="DC.Contributor" content="Philippe Charlier" /> <meta name="DC.Contributor" content="Elizaveta Chernykh" /> <meta name="DC.Contributor" content="Torben Trier Christiansen" /> <meta name="DC.Contributor" content="Alfredo Coppa" /> <meta name="DC.Contributor" content="Maura De Coster" /> <meta name="DC.Contributor" content="Sean Dexter Denham" /> <meta name="DC.Contributor" content="Sophie Desenne" /> <meta name="DC.Contributor" content="Jane Downes" /> <meta name="DC.Contributor" content="Karin Margarita Frei" /> <meta name="DC.Contributor" content="Olivér Gábor" /> <meta name="DC.Contributor" content="Johan Zakarias Gårdsvoll" /> <meta name="DC.Contributor" content="Zanette Tsigaridas Glørstad" /> <meta name="DC.Contributor" content="Jesper Hansen" /> <meta name="DC.Contributor" content="Stijn Heeren" /> <meta name="DC.Contributor" content="Merete Henriksen" /> <meta name="DC.Contributor" content="Volker Heyd" /> <meta name="DC.Contributor" content="Mette Høj" /> <meta name="DC.Contributor" content="Mads Kähler Holst" /> <meta name="DC.Contributor" content="Rimantas Jankauskas" /> <meta name="DC.Contributor" content="Henrik Janson" /> <meta name="DC.Contributor" content="Mads Dengsø Jessen" /> <meta name="DC.Contributor" content="Jens Winther Johannsen" /> <meta name="DC.Contributor" content="Torkel Johansen" /> <meta name="DC.Contributor" content="Ole Thirup Kastholm" /> <meta name="DC.Contributor" content="Anton Kern" /> <meta name="DC.Contributor" content="Ruslan Khaskhanov" /> <meta name="DC.Contributor" content="Katrine Kjær" /> <meta name="DC.Contributor" content="Vladimir Kolosov" /> <meta name="DC.Contributor" content="Lisette M. Kootker" /> <meta name="DC.Contributor" content="Anne Christine Larsen" /> <meta name="DC.Contributor" content="Thierry Lejars" /> <meta name="DC.Contributor" content="Mette Løvschal" /> <meta name="DC.Contributor" content="Niels Lynnerup" /> <meta name="DC.Contributor" content="Yvonne Magnusson" /> <meta name="DC.Contributor" content="Kristiina Mannermaa" /> <meta name="DC.Contributor" content="Vyacheslav Masyakin" /> <meta name="DC.Contributor" content="Anne Lene Melheim" /> <meta name="DC.Contributor" content="Inga Merkyte" /> <meta name="DC.Contributor" content="Vyacheslav Moiseyev" /> <meta name="DC.Contributor" content="Stig Bergmann Møller" /> <meta name="DC.Contributor" content="Erika Molnár" /> <meta name="DC.Contributor" content="Nadja Mortensen" /> <meta name="DC.Contributor" content="Eileen Murphy" /> <meta name="DC.Contributor" content="Bjarne Henning Nielsen" /> <meta name="DC.Contributor" content="Doris Pany-Kucera" /> <meta name="DC.Contributor" content="Bettina Schulz Paulsson" /> <meta name="DC.Contributor" content="Marcia S Ponce de León" /> <meta name="DC.Contributor" content="Håkon Reiersen" /> <meta name="DC.Contributor" content="Walter Reinhard" /> <meta name="DC.Contributor" content="Antti Sajantila" /> <meta name="DC.Contributor" content="Birgitte Skar" /> <meta name="DC.Contributor" content="Vladimir Slavchev" /> <meta name="DC.Contributor" content="Václav Smrčka" /> <meta name="DC.Contributor" content="Lasse Sørensen" /> <meta name="DC.Contributor" content="Georg Tiefengraber" /> <meta name="DC.Contributor" content="Otto Christian Uldum" /> <meta name="DC.Contributor" content="Jorge Vega" /> <meta name="DC.Contributor" content="Daniele Vitali" /> <meta name="DC.Contributor" content="Alexey Voloshinov" /> <meta name="DC.Contributor" content="Sidsel Wåhlin" /> <meta name="DC.Contributor" content="Holger Wendling" /> <meta name="DC.Contributor" content="Anna Wessman" /> <meta name="DC.Contributor" content="Helene Wilhelmson" /> <meta name="DC.Contributor" content="Karin Wiltschke" /> <meta name="DC.Contributor" content="João Zilhao" /> <meta name="DC.Contributor" content="Christoph PE Zollikofer" /> <meta name="DC.Contributor" content="Thorfinn Sand Korneliussen" /> <meta name="DC.Contributor" content="Bruno Chaume" /> <meta name="DC.Contributor" content="Jean-Paul Demoule" /> <meta name="DC.Contributor" content="Thomas Werge" /> <meta name="DC.Contributor" content="Line Olsen" /> <meta name="DC.Contributor" content="Rasmus Nielsen" /> <meta name="DC.Contributor" content="Lotte Hedeager" /> <meta name="DC.Contributor" content="Kristian Kristiansen" /> <meta name="DC.Contributor" content="Martin Sikora" /> <meta name="DC.Contributor" content="Eske Willerslev" /> <meta name="article:published_time" content="2024-03-14" /> <meta name="article:section" content="New Results" /> <meta name="citation_title" content="Steppe Ancestry in western Eurasia and the spread of the Germanic Languages" /> <meta name="citation_abstract" lang="en" content="<h3>Summary</h3> <p>Germanic-speaking populations historically form an integral component of the North and Northwest European cultural configuration. According to linguistic consensus, the common ancestor of the Germanic languages, which include German, English, Frisian, Dutch as well as the Nordic languages, was spoken in Northern Europe during the Pre-Roman Iron Age. However, important questions remain concerning the earlier Bronze Age distribution of this Indo-European language branch in Scandinavia as well as the driving factors behind its Late Iron Age diversification and expansion across the European continent. A key difficulty in addressing these questions are the existence of striking differences in the interpretation of the archaeological record, leading to various hypotheses of correlations with linguistic dispersals and changes in material culture. Moreover, these interpretations have been difficult to assess using genomics due to limited ancient genomes and the difficulty in differentiating closely related populations. Here we integrate multidisciplinary evidence from population genomics, historical sources, archaeology and linguistics to offer a fully revised model for the origins and spread of Germanic languages and for the formation of the genomic ancestry of Germanic-speaking northern European populations, while acknowledging that coordinating archaeology, linguistics and genetics is complex and potentially controversial. We sequenced 710 ancient human genomes from western Eurasia and analysed them together with 3,940 published genomes suitable for imputing diploid genotypes. We find evidence of a previously unknown, large-scale Bronze Age migration within Scandinavia, originating in the east and becoming widespread to the west and south, thus providing a new potential driving factor for the expansion of the Germanic speech community. This East Scandinavian genetic cluster is first seen 800 years after the arrival of the Corded Ware Culture, the first Steppe-related population to emerge in Northern Europe, opening a new scenario implying a Late rather than an Middle Neolithic arrival of the Germanic language group in Scandinavia. Moreover, the non-local Hunter-Gatherer ancestry of this East Scandinavian cluster is indicative of a cross-Baltic maritime rather than a southern Scandinavian land-based entry. Later in the Iron Age around 1700 BP, we find a southward push of admixed Eastern and Southern Scandinavians into areas including Germany and the Netherlands, previously associated with Celtic speakers, mixing with local populations from the Eastern North Sea coast. During the Migration Period (1575-1200 BP), we find evidence of this structured, admixed Southern Scandinavian population representing the Western Germanic Anglo-Saxon migrations into Britain and Langobards into southern Europe. During the Migration Period, we detect a previously unknown northward migration back into Southern Scandinavia, partly replacing earlier inhabitants and forming the North Germanic-speaking Viking-Age populations of Denmark and southern Sweden, corresponding with historically attested Danes. However, the origin and character of these major changes in Scandinavia before the Viking Age remain contested. In contrast to these Western and Northern Germanic-speaking populations, we find the Wielbark population from Poland to be primarily of Eastern Scandinavian ancestry, supporting a Swedish origin for East Germanic groups. In contrast, the later cultural descendants, the Ostrogoths and Visigoths are predominantly of Southern European ancestry implying the adoption of Gothic culture. Together, these results highlight the use of archaeology, linguistics and genetics as distinct but complementary lines of evidence.</p>" /> <meta name="citation_journal_title" content="bioRxiv" /> <meta name="citation_publisher" content="Cold Spring Harbor Laboratory" /> <meta name="citation_publication_date" content="2024/01/01" /> <meta name="citation_mjid" content="biorxiv;2024.03.13.584607v1" /> <meta name="citation_id" content="2024.03.13.584607v1" /> <meta name="citation_public_url" content="https://www.biorxiv.org/content/10.1101/2024.03.13.584607v1" /> <meta name="citation_abstract_html_url" content="https://www.biorxiv.org/content/10.1101/2024.03.13.584607v1.abstract" /> <meta name="citation_full_html_url" content="https://www.biorxiv.org/content/10.1101/2024.03.13.584607v1.full" /> <meta name="citation_pdf_url" content="https://www.biorxiv.org/content/biorxiv/early/2024/03/14/2024.03.13.584607.full.pdf" /> <meta name="citation_doi" content="10.1101/2024.03.13.584607" /> <meta name="citation_num_pages" content="50" /> <meta name="citation_article_type" content="Article" /> <meta name="citation_section" content="New Results" /> <meta name="citation_firstpage" content="2024.03.13.584607" /> <meta name="citation_author" content="Hugh McColl" /> <meta name="citation_author_institution" content="Lundbeck Foundation GeoGenetics Center, Globe Institute, University of Copenhagen" /> <meta name="citation_author_orcid" content="http://orcid.org/0000-0002-7568-4270" /> <meta name="citation_author" content="Guus Kroonen" /> <meta name="citation_author_institution" content="Leiden University Centre for Linguistics, Leiden University" /> <meta name="citation_author_institution" content="Department of Nordic Studies and Linguistics, University of Copenhagen" /> <meta name="citation_author_orcid" content="http://orcid.org/0000-0002-3708-0476" /> <meta name="citation_author" content="J. Víctor Moreno-Mayar" /> <meta name="citation_author_institution" content="Lundbeck Foundation GeoGenetics Center, Globe Institute, University of Copenhagen" /> <meta name="citation_author" content="Frederik Valeur Seersholm" /> <meta name="citation_author_institution" content="Lundbeck Foundation GeoGenetics Center, Globe Institute, University of Copenhagen" /> <meta name="citation_author" content="Gabriele Scorrano" /> <meta name="citation_author_institution" content="Lundbeck Foundation GeoGenetics Center, Globe Institute, University of Copenhagen" /> <meta name="citation_author_orcid" content="http://orcid.org/0000-0002-0887-4023" /> <meta name="citation_author" content="Thomaz Pinotti" /> <meta name="citation_author_institution" content="Lundbeck Foundation GeoGenetics Center, Globe Institute, University of Copenhagen" /> <meta name="citation_author_institution" content="Laboratório de Biodiversidade e Evolução Molecular (LBEM), Universidade Federal de Minas Gerais" /> <meta name="citation_author" content="Tharsika Vimala" /> <meta name="citation_author_institution" content="Lundbeck Foundation GeoGenetics Center, Globe Institute, University of Copenhagen" /> <meta name="citation_author_orcid" content="http://orcid.org/0000-0003-2709-5815" /> <meta name="citation_author" content="Søren M. Sindbæk" /> <meta name="citation_author_institution" content="Centre for Urban Network Evolutions (UrbNet), School of Culture and Society, Aarhus University" /> <meta name="citation_author_orcid" content="http://orcid.org/0000-0001-9737-1243" /> <meta name="citation_author" content="Per Ethelberg" /> <meta name="citation_author_institution" content="Museum Sønderjylland" /> <meta name="citation_author" content="Ralph Fyfe" /> <meta name="citation_author_institution" content="School of Geography, Earth and Environmental Sciences, University of Plymouth UK," /> <meta name="citation_author_orcid" content="http://orcid.org/0000-0002-5676-008X" /> <meta name="citation_author" content="Marie-José Gaillard" /> <meta name="citation_author_institution" content="Department of Biology and Environmental Sciences, Linnaeus University" /> <meta name="citation_author_orcid" content="http://orcid.org/0000-0002-2025-410X" /> <meta name="citation_author" content="Hanne M. Ellegård Larsen" /> <meta name="citation_author_institution" content="National Museum of Denmark" /> <meta name="citation_author" content="Morten Fischer Mortensen" /> <meta name="citation_author_institution" content="National Museum of Denmark. Environmental Archaeology and Materials Science" /> <meta name="citation_author_orcid" content="http://orcid.org/0000-0002-4167-8227" /> <meta name="citation_author" content="Fabrice Demeter" /> <meta name="citation_author_institution" content="Lundbeck Foundation GeoGenetics Center, Globe Institute, University of Copenhagen" /> <meta name="citation_author_institution" content="Eco-anthropologie (EA), Dpt ABBA, Muséum national d’Histoire naturelle, CNRS, Université Paris Cité, Musée de l’Homme 17 place du Trocadéro" /> <meta name="citation_author" content="Marie Louise S. 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A global analysis of matches and mismatches between human genetic and linguistic histories. Proc. Natl. Acad. Sci. 119, e2122084119 (2022)." /> <meta name="twitter:title" content="Steppe Ancestry in western Eurasia and the spread of the Germanic Languages" /> <meta name="twitter:site" content="@biorxivpreprint" /> <meta name="twitter:card" content="summary" /> <meta name="twitter:image" content="https://www.biorxiv.org/sites/default/files/images/biorxiv_logo_homepage7-5-small.png" /> <meta name="twitter:description" content="Germanic-speaking populations historically form an integral component of the North and Northwest European cultural configuration. According to linguistic consensus, the common ancestor of the Germanic languages, which include German, English, Frisian, Dutch as well as the Nordic languages, was spoken in Northern Europe during the Pre-Roman Iron Age. However, important questions remain concerning the earlier Bronze Age distribution of this Indo-European language branch in Scandinavia as well as the driving factors behind its Late Iron Age diversification and expansion across the European continent. A key difficulty in addressing these questions are the existence of striking differences in the interpretation of the archaeological record, leading to various hypotheses of correlations with linguistic dispersals and changes in material culture. Moreover, these interpretations have been difficult to assess using genomics due to limited ancient genomes and the difficulty in differentiating closely related populations. Here we integrate multidisciplinary evidence from population genomics, historical sources, archaeology and linguistics to offer a fully revised model for the origins and spread of Germanic languages and for the formation of the genomic ancestry of Germanic-speaking northern European populations, while acknowledging that coordinating archaeology, linguistics and genetics is complex and potentially controversial. We sequenced 710 ancient human genomes from western Eurasia and analysed them together with 3,940 published genomes suitable for imputing diploid genotypes. We find evidence of a previously unknown, large-scale Bronze Age migration within Scandinavia, originating in the east and becoming widespread to the west and south, thus providing a new potential driving factor for the expansion of the Germanic speech community. This East Scandinavian genetic cluster is first seen 800 years after the arrival of the Corded Ware Culture, the first Steppe-related population to emerge in Northern Europe, opening a new scenario implying a Late rather than an Middle Neolithic arrival of the Germanic language group in Scandinavia. Moreover, the non-local Hunter-Gatherer ancestry of this East Scandinavian cluster is indicative of a cross-Baltic maritime rather than a southern Scandinavian land-based entry. Later in the Iron Age around 1700 BP, we find a southward push of admixed Eastern and Southern Scandinavians into areas including Germany and the Netherlands, previously associated with Celtic speakers, mixing with local populations from the Eastern North Sea coast. During the Migration Period (1575-1200 BP), we find evidence of this structured, admixed Southern Scandinavian population representing the Western Germanic Anglo-Saxon migrations into Britain and Langobards into southern Europe. During the Migration Period, we detect a previously unknown northward migration back into Southern Scandinavia, partly replacing earlier inhabitants and forming the North Germanic-speaking Viking-Age populations of Denmark and southern Sweden, corresponding with historically attested Danes. However, the origin and character of these major changes in Scandinavia before the Viking Age remain contested. In contrast to these Western and Northern Germanic-speaking populations, we find the Wielbark population from Poland to be primarily of Eastern Scandinavian ancestry, supporting a Swedish origin for East Germanic groups. In contrast, the later cultural descendants, the Ostrogoths and Visigoths are predominantly of Southern European ancestry implying the adoption of Gothic culture. Together, these results highlight the use of archaeology, linguistics and genetics as distinct but complementary lines of evidence. ### Competing Interest Statement The authors have declared no competing interest." /> <meta name="og-title" property="og:title" content="Steppe Ancestry in western Eurasia and the spread of the Germanic Languages" /> <meta name="og-url" property="og:url" content="https://www.biorxiv.org/content/10.1101/2024.03.13.584607v1" /> <meta name="og-site-name" property="og:site_name" content="bioRxiv" /> <meta name="og-description" property="og:description" content="Germanic-speaking populations historically form an integral component of the North and Northwest European cultural configuration. According to linguistic consensus, the common ancestor of the Germanic languages, which include German, English, Frisian, Dutch as well as the Nordic languages, was spoken in Northern Europe during the Pre-Roman Iron Age. However, important questions remain concerning the earlier Bronze Age distribution of this Indo-European language branch in Scandinavia as well as the driving factors behind its Late Iron Age diversification and expansion across the European continent. A key difficulty in addressing these questions are the existence of striking differences in the interpretation of the archaeological record, leading to various hypotheses of correlations with linguistic dispersals and changes in material culture. Moreover, these interpretations have been difficult to assess using genomics due to limited ancient genomes and the difficulty in differentiating closely related populations. Here we integrate multidisciplinary evidence from population genomics, historical sources, archaeology and linguistics to offer a fully revised model for the origins and spread of Germanic languages and for the formation of the genomic ancestry of Germanic-speaking northern European populations, while acknowledging that coordinating archaeology, linguistics and genetics is complex and potentially controversial. We sequenced 710 ancient human genomes from western Eurasia and analysed them together with 3,940 published genomes suitable for imputing diploid genotypes. We find evidence of a previously unknown, large-scale Bronze Age migration within Scandinavia, originating in the east and becoming widespread to the west and south, thus providing a new potential driving factor for the expansion of the Germanic speech community. This East Scandinavian genetic cluster is first seen 800 years after the arrival of the Corded Ware Culture, the first Steppe-related population to emerge in Northern Europe, opening a new scenario implying a Late rather than an Middle Neolithic arrival of the Germanic language group in Scandinavia. Moreover, the non-local Hunter-Gatherer ancestry of this East Scandinavian cluster is indicative of a cross-Baltic maritime rather than a southern Scandinavian land-based entry. Later in the Iron Age around 1700 BP, we find a southward push of admixed Eastern and Southern Scandinavians into areas including Germany and the Netherlands, previously associated with Celtic speakers, mixing with local populations from the Eastern North Sea coast. During the Migration Period (1575-1200 BP), we find evidence of this structured, admixed Southern Scandinavian population representing the Western Germanic Anglo-Saxon migrations into Britain and Langobards into southern Europe. During the Migration Period, we detect a previously unknown northward migration back into Southern Scandinavia, partly replacing earlier inhabitants and forming the North Germanic-speaking Viking-Age populations of Denmark and southern Sweden, corresponding with historically attested Danes. However, the origin and character of these major changes in Scandinavia before the Viking Age remain contested. In contrast to these Western and Northern Germanic-speaking populations, we find the Wielbark population from Poland to be primarily of Eastern Scandinavian ancestry, supporting a Swedish origin for East Germanic groups. In contrast, the later cultural descendants, the Ostrogoths and Visigoths are predominantly of Southern European ancestry implying the adoption of Gothic culture. Together, these results highlight the use of archaeology, linguistics and genetics as distinct but complementary lines of evidence. ### Competing Interest Statement The authors have declared no competing interest." /> <meta name="og-type" property="og:type" content="article" /> <meta name="og-image" property="og:image" content="https://www.biorxiv.org/sites/default/files/images/biorxiv_logo_homepage7-5-small.png" /> <meta name="citation_date" content="2024-03-14" /> <link rel="alternate" type="application/vnd.ms-powerpoint" title="Powerpoint" href="/content/10.1101/2024.03.13.584607v1.ppt" /> <meta name="description" content="bioRxiv - the preprint server for biology, operated by Cold Spring Harbor Laboratory, a research and educational institution" /> <meta name="generator" content="Drupal 7 (http://drupal.org)" /> <link rel="canonical" href="https://www.biorxiv.org/content/10.1101/2024.03.13.584607v1" /> <link rel="shortlink" href="https://www.biorxiv.org/node/3713187" /> 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class="highwire-article-citation highwire-citation-type-highwire-article node3713187" data-node-nid="3713187" id="node-3713187--22118244168" data-pisa="biorxiv;2024.03.13.584607v1" data-pisa-master="biorxiv;2024.03.13.584607" data-apath="/biorxiv/early/2024/03/14/2024.03.13.584607.atom" data-hw-author-tooltip-instance="highwire_author_tooltip"><div class="highwire-cite highwire-cite-highwire-article highwire-citation-biorxiv-article-top clearfix has-author-tooltip" > <span class="biorxiv-article-type"> New Results </span> <h1 class="highwire-cite-title" id="page-title">Steppe Ancestry in western Eurasia and the spread of the Germanic Languages</h1> <div class="highwire-cite-authors" ><span class="highwire-citation-authors"><span class="highwire-citation-author first hw-author-orcid-logo-wrapper" data-delta="0"><a href="http://orcid.org/0000-0002-7568-4270" target="_blank" class="hw-author-orcid-logo link-icon-only link-icon"><span class="hw-icon-orcid hw-icon-color-orcid"></span> <span class="title element-invisible">View ORCID Profile</span></a><span class="nlm-given-names">Hugh</span> <span class="nlm-surname">McColl</span></span>, <span class="highwire-citation-author hw-author-orcid-logo-wrapper" data-delta="1"><a href="http://orcid.org/0000-0002-3708-0476" target="_blank" class="hw-author-orcid-logo link-icon-only link-icon"><span class="hw-icon-orcid hw-icon-color-orcid"></span> <span class="title element-invisible">View ORCID Profile</span></a><span class="nlm-given-names">Guus</span> <span class="nlm-surname">Kroonen</span></span>, <span class="highwire-citation-author" data-delta="2"><span class="nlm-given-names">J. Víctor</span> <span class="nlm-surname">Moreno-Mayar</span></span>, <span class="highwire-citation-author" data-delta="3"><span class="nlm-given-names">Frederik</span> <span class="nlm-surname">Valeur Seersholm</span></span>, <span class="highwire-citation-author hw-author-orcid-logo-wrapper" data-delta="4"><a href="http://orcid.org/0000-0002-0887-4023" target="_blank" class="hw-author-orcid-logo link-icon-only link-icon"><span class="hw-icon-orcid hw-icon-color-orcid"></span> <span class="title element-invisible">View ORCID Profile</span></a><span class="nlm-given-names">Gabriele</span> <span class="nlm-surname">Scorrano</span></span>, <span class="highwire-citation-author" data-delta="5"><span class="nlm-given-names">Thomaz</span> <span class="nlm-surname">Pinotti</span></span>, <span class="highwire-citation-author hw-author-orcid-logo-wrapper" data-delta="6"><a href="http://orcid.org/0000-0003-2709-5815" target="_blank" class="hw-author-orcid-logo link-icon-only link-icon"><span class="hw-icon-orcid hw-icon-color-orcid"></span> <span class="title element-invisible">View ORCID Profile</span></a><span class="nlm-given-names">Tharsika</span> <span class="nlm-surname">Vimala</span></span>, <span class="highwire-citation-author hw-author-orcid-logo-wrapper" data-delta="7"><a href="http://orcid.org/0000-0001-9737-1243" target="_blank" class="hw-author-orcid-logo link-icon-only link-icon"><span class="hw-icon-orcid hw-icon-color-orcid"></span> <span class="title element-invisible">View ORCID Profile</span></a><span class="nlm-given-names">Søren M.</span> <span class="nlm-surname">Sindbæk</span></span>, <span class="highwire-citation-author" data-delta="8"><span class="nlm-given-names">Per</span> <span class="nlm-surname">Ethelberg</span></span>, <span class="highwire-citation-author hw-author-orcid-logo-wrapper" data-delta="9"><a href="http://orcid.org/0000-0002-5676-008X" target="_blank" class="hw-author-orcid-logo link-icon-only link-icon"><span class="hw-icon-orcid hw-icon-color-orcid"></span> <span class="title element-invisible">View ORCID Profile</span></a><span class="nlm-given-names">Ralph</span> <span class="nlm-surname">Fyfe</span></span>, <span class="highwire-citation-author hw-author-orcid-logo-wrapper" data-delta="10"><a href="http://orcid.org/0000-0002-2025-410X" target="_blank" class="hw-author-orcid-logo link-icon-only link-icon"><span class="hw-icon-orcid hw-icon-color-orcid"></span> <span class="title element-invisible">View ORCID Profile</span></a><span class="nlm-given-names">Marie-José</span> <span class="nlm-surname">Gaillard</span></span>, <span class="highwire-citation-author" data-delta="11"><span class="nlm-given-names">Hanne M. Ellegård</span> <span class="nlm-surname">Larsen</span></span>, <span class="highwire-citation-author hw-author-orcid-logo-wrapper" data-delta="12"><a href="http://orcid.org/0000-0002-4167-8227" target="_blank" class="hw-author-orcid-logo link-icon-only link-icon"><span class="hw-icon-orcid hw-icon-color-orcid"></span> <span class="title element-invisible">View ORCID Profile</span></a><span class="nlm-given-names">Morten Fischer</span> <span class="nlm-surname">Mortensen</span></span>, <span class="highwire-citation-author" data-delta="13"><span class="nlm-given-names">Fabrice</span> <span class="nlm-surname">Demeter</span></span>, <span class="highwire-citation-author hw-author-orcid-logo-wrapper" data-delta="14"><a href="http://orcid.org/0000-0002-5283-4328" target="_blank" class="hw-author-orcid-logo link-icon-only link-icon"><span class="hw-icon-orcid hw-icon-color-orcid"></span> <span class="title element-invisible">View ORCID Profile</span></a><span class="nlm-given-names">Marie Louise S.</span> <span class="nlm-surname">Jørkov</span></span>, <span class="highwire-citation-author hw-author-orcid-logo-wrapper" data-delta="15"><a href="http://orcid.org/0000-0002-9614-0264" target="_blank" class="hw-author-orcid-logo link-icon-only link-icon"><span class="hw-icon-orcid hw-icon-color-orcid"></span> <span class="title element-invisible">View ORCID Profile</span></a><span class="nlm-given-names">Sophie</span> <span class="nlm-surname">Bergerbrant</span></span>, <span class="highwire-citation-author" data-delta="16"><span class="nlm-given-names">Peter de Barros</span> <span class="nlm-surname">Damgaard</span></span>, <span class="highwire-citation-author" data-delta="17"><span class="nlm-given-names">Morten E.</span> <span class="nlm-surname">Allentoft</span></span>, <span class="highwire-citation-author hw-author-orcid-logo-wrapper" data-delta="18"><a href="http://orcid.org/0000-0002-3081-3702" target="_blank" class="hw-author-orcid-logo link-icon-only 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class="hw-icon-orcid hw-icon-color-orcid"></span> <span class="title element-invisible">View ORCID Profile</span></a><span class="nlm-given-names">Isin</span> <span class="nlm-surname">Altinkaya</span></span>, <span class="highwire-citation-author hw-author-orcid-logo-wrapper" data-delta="22"><a href="http://orcid.org/0000-0003-3657-1983" target="_blank" class="hw-author-orcid-logo link-icon-only link-icon"><span class="hw-icon-orcid hw-icon-color-orcid"></span> <span class="title element-invisible">View ORCID Profile</span></a><span class="nlm-given-names">Rasmus</span> <span class="nlm-surname">Amund Henriksen</span></span>, <span class="highwire-citation-author hw-author-orcid-logo-wrapper" data-delta="23"><a href="http://orcid.org/0000-0003-1940-2192" target="_blank" class="hw-author-orcid-logo link-icon-only link-icon"><span class="hw-icon-orcid hw-icon-color-orcid"></span> <span class="title element-invisible">View ORCID Profile</span></a><span class="nlm-given-names">Evan K.</span> <span class="nlm-surname">Irving-Pease</span></span>, <span class="highwire-citation-author hw-author-orcid-logo-wrapper" data-delta="24"><a href="http://orcid.org/0000-0001-6404-1280" target="_blank" class="hw-author-orcid-logo link-icon-only link-icon"><span class="hw-icon-orcid hw-icon-color-orcid"></span> <span class="title element-invisible">View ORCID Profile</span></a><span class="nlm-given-names">Serena</span> <span class="nlm-surname">Sabatini</span></span>, <span class="highwire-citation-author hw-author-orcid-logo-wrapper" data-delta="25"><a href="http://orcid.org/0000-0003-0852-5904" target="_blank" class="hw-author-orcid-logo link-icon-only link-icon"><span class="hw-icon-orcid hw-icon-color-orcid"></span> <span class="title element-invisible">View ORCID Profile</span></a><span class="nlm-given-names">Anders</span> <span class="nlm-surname">Fischer</span></span>, <span class="highwire-citation-author hw-author-orcid-logo-wrapper" data-delta="26"><a href="http://orcid.org/0000-0003-0698-4955" target="_blank" class="hw-author-orcid-logo link-icon-only link-icon"><span class="hw-icon-orcid hw-icon-color-orcid"></span> <span class="title element-invisible">View ORCID Profile</span></a><span class="nlm-given-names">William</span> <span class="nlm-surname">Barrie</span></span>, <span class="highwire-citation-author" data-delta="27"><span class="nlm-given-names">Andrés</span> <span class="nlm-surname">Ingason</span></span>, <span class="highwire-citation-author hw-author-orcid-logo-wrapper" data-delta="28"><a href="http://orcid.org/0000-0002-6682-1288" target="_blank" class="hw-author-orcid-logo link-icon-only link-icon"><span class="hw-icon-orcid hw-icon-color-orcid"></span> <span class="title element-invisible">View ORCID Profile</span></a><span class="nlm-given-names">Anders</span> <span class="nlm-surname">Rosengren</span></span>, <span class="highwire-citation-author hw-author-orcid-logo-wrapper" data-delta="29"><a href="http://orcid.org/0000-0003-3113-2981" target="_blank" class="hw-author-orcid-logo link-icon-only link-icon"><span class="hw-icon-orcid hw-icon-color-orcid"></span> <span class="title element-invisible">View ORCID Profile</span></a><span class="nlm-given-names">Andrew</span> <span class="nlm-surname">Vaughn</span></span>, <span class="highwire-citation-author" data-delta="30"><span class="nlm-given-names">Jialu</span> <span class="nlm-surname">Cao</span></span>, <span class="highwire-citation-author" data-delta="31"><span class="nlm-given-names">Jacqueline</span> <span class="nlm-surname">Staring</span></span>, <span class="highwire-citation-author" data-delta="32"><span class="nlm-given-names">Jesper</span> <span class="nlm-surname">Stenderup</span></span>, <span class="highwire-citation-author" data-delta="33"><span class="nlm-given-names">Fulya Eylem</span> <span class="nlm-surname">Yediay</span></span>, <span class="highwire-citation-author hw-author-orcid-logo-wrapper" data-delta="34"><a href="http://orcid.org/0000-0001-7468-5979" target="_blank" class="hw-author-orcid-logo link-icon-only link-icon"><span class="hw-icon-orcid hw-icon-color-orcid"></span> <span class="title element-invisible">View ORCID Profile</span></a><span class="nlm-given-names">Torbjörn</span> <span class="nlm-surname">Ahlström</span></span>, <span class="highwire-citation-author" data-delta="35"><span class="nlm-given-names">Sofie Laurine</span> <span class="nlm-surname">Albris</span></span>, <span class="highwire-citation-author hw-author-orcid-logo-wrapper" data-delta="36"><a href="http://orcid.org/0000-0002-9195-2482" target="_blank" class="hw-author-orcid-logo link-icon-only link-icon"><span class="hw-icon-orcid hw-icon-color-orcid"></span> <span class="title element-invisible">View ORCID Profile</span></a><span class="nlm-given-names">Biyaslan</span> <span class="nlm-surname">Atabiev</span></span>, <span class="highwire-citation-author hw-author-orcid-logo-wrapper" 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<span class="highwire-citation-author hw-author-orcid-logo-wrapper" data-delta="53"><a href="http://orcid.org/0000-0001-9265-3882" target="_blank" class="hw-author-orcid-logo link-icon-only link-icon"><span class="hw-icon-orcid hw-icon-color-orcid"></span> <span class="title element-invisible">View ORCID Profile</span></a><span class="nlm-given-names">Stijn</span> <span class="nlm-surname">Heeren</span></span>, <span class="highwire-citation-author" data-delta="54"><span class="nlm-given-names">Merete</span> <span class="nlm-surname">Henriksen</span></span>, <span class="highwire-citation-author hw-author-orcid-logo-wrapper" data-delta="55"><a href="http://orcid.org/0000-0002-0889-6513" target="_blank" class="hw-author-orcid-logo link-icon-only link-icon"><span class="hw-icon-orcid hw-icon-color-orcid"></span> <span class="title element-invisible">View ORCID Profile</span></a><span class="nlm-given-names">Volker</span> <span class="nlm-surname">Heyd</span></span>, <span class="highwire-citation-author" data-delta="56"><span class="nlm-given-names">Mette</span> <span class="nlm-surname">Høj</span></span>, <span class="highwire-citation-author" data-delta="57"><span class="nlm-given-names">Mads Kähler</span> <span class="nlm-surname">Holst</span></span>, <span class="highwire-citation-author" data-delta="58"><span class="nlm-given-names">Rimantas</span> <span class="nlm-surname">Jankauskas</span></span>, <span class="highwire-citation-author" data-delta="59"><span class="nlm-given-names">Henrik</span> <span class="nlm-surname">Janson</span></span>, <span class="highwire-citation-author" data-delta="60"><span class="nlm-given-names">Mads Dengsø</span> <span class="nlm-surname">Jessen</span></span>, <span class="highwire-citation-author" data-delta="61"><span class="nlm-given-names">Jens Winther</span> <span class="nlm-surname">Johannsen</span></span>, <span class="highwire-citation-author" data-delta="62"><span class="nlm-given-names">Torkel</span> <span class="nlm-surname">Johansen</span></span>, <span class="highwire-citation-author hw-author-orcid-logo-wrapper" data-delta="63"><a href="http://orcid.org/0000-0003-4793-5607" target="_blank" class="hw-author-orcid-logo link-icon-only link-icon"><span class="hw-icon-orcid hw-icon-color-orcid"></span> <span class="title element-invisible">View ORCID Profile</span></a><span class="nlm-given-names">Ole Thirup</span> <span class="nlm-surname">Kastholm</span></span>, <span class="highwire-citation-author" data-delta="64"><span class="nlm-given-names">Anton</span> <span class="nlm-surname">Kern</span></span>, <span class="highwire-citation-author" data-delta="65"><span class="nlm-given-names">Ruslan</span> <span class="nlm-surname">Khaskhanov</span></span>, <span class="highwire-citation-author" data-delta="66"><span class="nlm-given-names">Katrine</span> <span class="nlm-surname">Kjær</span></span>, <span class="highwire-citation-author" data-delta="67"><span 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hw-author-orcid-logo-wrapper" data-delta="72"><a href="http://orcid.org/0000-0002-7771-5376" target="_blank" class="hw-author-orcid-logo link-icon-only link-icon"><span class="hw-icon-orcid hw-icon-color-orcid"></span> <span class="title element-invisible">View ORCID Profile</span></a><span class="nlm-given-names">Niels</span> <span class="nlm-surname">Lynnerup</span></span>, <span class="highwire-citation-author hw-author-orcid-logo-wrapper" data-delta="73"><a href="http://orcid.org/0000-0002-7076-2583" target="_blank" class="hw-author-orcid-logo link-icon-only link-icon"><span class="hw-icon-orcid hw-icon-color-orcid"></span> <span class="title element-invisible">View ORCID Profile</span></a><span class="nlm-given-names">Yvonne</span> <span class="nlm-surname">Magnusson</span></span>, <span class="highwire-citation-author hw-author-orcid-logo-wrapper" data-delta="74"><a href="http://orcid.org/0000-0002-8510-1120" target="_blank" class="hw-author-orcid-logo link-icon-only 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Profile</span></a><span class="nlm-given-names">Vyacheslav</span> <span class="nlm-surname">Moiseyev</span></span>, <span class="highwire-citation-author" data-delta="79"><span class="nlm-given-names">Stig Bergmann</span> <span class="nlm-surname">Møller</span></span>, <span class="highwire-citation-author hw-author-orcid-logo-wrapper" data-delta="80"><a href="http://orcid.org/0000-0001-6660-9239" target="_blank" class="hw-author-orcid-logo link-icon-only link-icon"><span class="hw-icon-orcid hw-icon-color-orcid"></span> <span class="title element-invisible">View ORCID Profile</span></a><span class="nlm-given-names">Erika</span> <span class="nlm-surname">Molnár</span></span>, <span class="highwire-citation-author" data-delta="81"><span class="nlm-given-names">Nadja</span> <span class="nlm-surname">Mortensen</span></span>, <span class="highwire-citation-author" data-delta="82"><span class="nlm-given-names">Eileen</span> <span class="nlm-surname">Murphy</span></span>, <span class="highwire-citation-author" data-delta="83"><span class="nlm-given-names">Bjarne Henning</span> <span class="nlm-surname">Nielsen</span></span>, <span class="highwire-citation-author hw-author-orcid-logo-wrapper" data-delta="84"><a href="http://orcid.org/0000-0003-4140-3220" target="_blank" class="hw-author-orcid-logo link-icon-only link-icon"><span class="hw-icon-orcid hw-icon-color-orcid"></span> <span class="title element-invisible">View ORCID Profile</span></a><span class="nlm-given-names">Doris</span> <span class="nlm-surname">Pany-Kucera</span></span>, <span class="highwire-citation-author" data-delta="85"><span class="nlm-given-names">Bettina Schulz</span> <span class="nlm-surname">Paulsson</span></span>, <span class="highwire-citation-author" data-delta="86"><span class="nlm-given-names">Marcia S</span> <span class="nlm-surname">Ponce de León</span></span>, <span class="highwire-citation-author hw-author-orcid-logo-wrapper" data-delta="87"><a href="http://orcid.org/0000-0001-7429-2112" target="_blank" class="hw-author-orcid-logo link-icon-only link-icon"><span class="hw-icon-orcid hw-icon-color-orcid"></span> <span class="title element-invisible">View ORCID Profile</span></a><span class="nlm-given-names">Håkon</span> <span class="nlm-surname">Reiersen</span></span>, <span class="highwire-citation-author" data-delta="88"><span class="nlm-given-names">Walter</span> <span class="nlm-surname">Reinhard</span></span>, <span class="highwire-citation-author hw-author-orcid-logo-wrapper" data-delta="89"><a href="http://orcid.org/0000-0003-4117-5248" target="_blank" class="hw-author-orcid-logo link-icon-only link-icon"><span class="hw-icon-orcid hw-icon-color-orcid"></span> <span class="title element-invisible">View ORCID Profile</span></a><span class="nlm-given-names">Antti</span> <span class="nlm-surname">Sajantila</span></span>, <span class="highwire-citation-author hw-author-orcid-logo-wrapper" data-delta="90"><a href="http://orcid.org/0000-0002-4784-2948" target="_blank" class="hw-author-orcid-logo link-icon-only link-icon"><span class="hw-icon-orcid hw-icon-color-orcid"></span> <span class="title element-invisible">View ORCID Profile</span></a><span class="nlm-given-names">Birgitte</span> <span class="nlm-surname">Skar</span></span>, <span class="highwire-citation-author hw-author-orcid-logo-wrapper" data-delta="91"><a href="http://orcid.org/0000-0002-6369-4527" target="_blank" class="hw-author-orcid-logo link-icon-only link-icon"><span class="hw-icon-orcid hw-icon-color-orcid"></span> <span class="title element-invisible">View ORCID Profile</span></a><span class="nlm-given-names">Vladimir</span> <span class="nlm-surname">Slavchev</span></span>, <span class="highwire-citation-author" data-delta="92"><span class="nlm-given-names">Václav</span> <span class="nlm-surname">Smrčka</span></span>, <span class="highwire-citation-author hw-author-orcid-logo-wrapper" data-delta="93"><a href="http://orcid.org/0000-0001-5687-5357" target="_blank" class="hw-author-orcid-logo link-icon-only link-icon"><span class="hw-icon-orcid hw-icon-color-orcid"></span> <span class="title element-invisible">View ORCID Profile</span></a><span class="nlm-given-names">Lasse</span> <span class="nlm-surname">Sørensen</span></span>, <span class="highwire-citation-author hw-author-orcid-logo-wrapper" data-delta="94"><a href="http://orcid.org/0000-0002-1800-6684" target="_blank" class="hw-author-orcid-logo link-icon-only link-icon"><span class="hw-icon-orcid hw-icon-color-orcid"></span> <span class="title element-invisible">View ORCID Profile</span></a><span class="nlm-given-names">Georg</span> <span class="nlm-surname">Tiefengraber</span></span>, <span class="highwire-citation-author" data-delta="95"><span class="nlm-given-names">Otto Christian</span> <span class="nlm-surname">Uldum</span></span>, <span class="highwire-citation-author" data-delta="96"><span class="nlm-given-names">Jorge</span> <span class="nlm-surname">Vega</span></span>, <span class="highwire-citation-author" data-delta="97"><span class="nlm-given-names">Daniele</span> <span class="nlm-surname">Vitali</span></span>, <span class="highwire-citation-author" data-delta="98"><span class="nlm-given-names">Alexey</span> <span class="nlm-surname">Voloshinov</span></span>, <span class="highwire-citation-author" data-delta="99"><span class="nlm-given-names">Sidsel</span> <span class="nlm-surname">Wåhlin</span></span>, <span class="highwire-citation-author hw-author-orcid-logo-wrapper" data-delta="100"><a href="http://orcid.org/0000-0003-0295-0248" target="_blank" class="hw-author-orcid-logo link-icon-only link-icon"><span class="hw-icon-orcid hw-icon-color-orcid"></span> <span class="title element-invisible">View ORCID Profile</span></a><span class="nlm-given-names">Holger</span> <span class="nlm-surname">Wendling</span></span>, <span class="highwire-citation-author hw-author-orcid-logo-wrapper" data-delta="101"><a href="http://orcid.org/0000-0001-6886-5455" target="_blank" class="hw-author-orcid-logo link-icon-only link-icon"><span class="hw-icon-orcid hw-icon-color-orcid"></span> <span class="title element-invisible">View ORCID Profile</span></a><span class="nlm-given-names">Anna</span> <span class="nlm-surname">Wessman</span></span>, <span class="highwire-citation-author" data-delta="102"><span class="nlm-given-names">Helene</span> <span class="nlm-surname">Wilhelmson</span></span>, <span class="highwire-citation-author hw-author-orcid-logo-wrapper" data-delta="103"><a href="http://orcid.org/0000-0002-8321-5095" target="_blank" class="hw-author-orcid-logo link-icon-only link-icon"><span class="hw-icon-orcid hw-icon-color-orcid"></span> <span class="title element-invisible">View ORCID Profile</span></a><span class="nlm-given-names">Karin</span> <span class="nlm-surname">Wiltschke</span></span>, <span class="highwire-citation-author hw-author-orcid-logo-wrapper" data-delta="104"><a href="http://orcid.org/0000-0001-5937-3061" target="_blank" class="hw-author-orcid-logo link-icon-only link-icon"><span class="hw-icon-orcid hw-icon-color-orcid"></span> <span class="title element-invisible">View ORCID Profile</span></a><span class="nlm-given-names">João</span> <span class="nlm-surname">Zilhao</span></span>, <span class="highwire-citation-author" data-delta="105"><span class="nlm-given-names">Christoph PE</span> <span class="nlm-surname">Zollikofer</span></span>, <span class="highwire-citation-author hw-author-orcid-logo-wrapper" data-delta="106"><a href="http://orcid.org/0000-0001-7576-5380" target="_blank" class="hw-author-orcid-logo link-icon-only link-icon"><span class="hw-icon-orcid hw-icon-color-orcid"></span> <span class="title element-invisible">View ORCID Profile</span></a><span class="nlm-given-names">Thorfinn</span> <span class="nlm-surname">Sand Korneliussen</span></span>, <span class="highwire-citation-author" data-delta="107"><span class="nlm-given-names">Bruno</span> <span class="nlm-surname">Chaume</span></span>, <span class="highwire-citation-author hw-author-orcid-logo-wrapper" data-delta="108"><a href="http://orcid.org/0000-0003-4712-5110" target="_blank" class="hw-author-orcid-logo link-icon-only link-icon"><span class="hw-icon-orcid hw-icon-color-orcid"></span> <span class="title element-invisible">View ORCID Profile</span></a><span class="nlm-given-names">Jean-Paul</span> <span class="nlm-surname">Demoule</span></span>, <span class="highwire-citation-author" data-delta="109"><span class="nlm-given-names">Thomas</span> <span class="nlm-surname">Werge</span></span>, <span class="highwire-citation-author hw-author-orcid-logo-wrapper" data-delta="110"><a href="http://orcid.org/0000-0001-6785-4981" target="_blank" class="hw-author-orcid-logo link-icon-only link-icon"><span class="hw-icon-orcid hw-icon-color-orcid"></span> <span class="title element-invisible">View ORCID Profile</span></a><span class="nlm-given-names">Line</span> <span class="nlm-surname">Olsen</span></span>, <span class="highwire-citation-author" data-delta="111"><span class="nlm-given-names">Rasmus</span> <span class="nlm-surname">Nielsen</span></span>, <span class="highwire-citation-author" data-delta="112"><span class="nlm-given-names">Lotte</span> <span class="nlm-surname">Hedeager</span></span>, <span class="highwire-citation-author hw-author-orcid-logo-wrapper" data-delta="113"><a href="http://orcid.org/0000-0003-2423-308X" target="_blank" class="hw-author-orcid-logo link-icon-only link-icon"><span class="hw-icon-orcid hw-icon-color-orcid"></span> <span class="title element-invisible">View ORCID Profile</span></a><span class="nlm-given-names">Kristian</span> <span class="nlm-surname">Kristiansen</span></span>, <span class="highwire-citation-author hw-author-orcid-logo-wrapper" data-delta="114"><a href="http://orcid.org/0000-0003-2818-8319" target="_blank" class="hw-author-orcid-logo link-icon-only link-icon"><span class="hw-icon-orcid hw-icon-color-orcid"></span> <span class="title element-invisible">View ORCID Profile</span></a><span class="nlm-given-names">Martin</span> <span class="nlm-surname">Sikora</span></span>, <span class="highwire-citation-author hw-author-orcid-logo-wrapper" data-delta="115"><a href="http://orcid.org/0000-0002-7081-6748" target="_blank" class="hw-author-orcid-logo link-icon-only link-icon"><span class="hw-icon-orcid hw-icon-color-orcid"></span> <span class="title element-invisible">View ORCID Profile</span></a><span class="nlm-given-names">Eske</span> <span class="nlm-surname">Willerslev</span></span></span></div> <div class="highwire-cite-metadata" ><span class="highwire-cite-metadata-doi highwire-cite-metadata"><span class="label">doi:</span> https://doi.org/10.1101/2024.03.13.584607 </span></div> </div> <div id="hw-article-author-popups-node-3713187--22118244168" style="display: none;"><div class="author-tooltip-0"><div class="author-tooltip-name">Hugh McColl </div><div class="author-tooltip-affiliation"><span class="author-tooltip-text"><div class='author-affiliation'><span class='nlm-sup'>1</span><span class='nlm-institution'>Lundbeck Foundation GeoGenetics Center, Globe Institute, University of Copenhagen</span>, Copenhagen, <span class='nlm-country'>Denmark</span>.,</div></span></div><ul class="author-tooltip-find-more"><li class="author-tooltip-gs-link first"><a href="/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=Hugh%2BMcColl%2B" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on Google Scholar</a></li><li class="author-tooltip-pubmed-link"><a href="/lookup/external-ref?access_num=McColl%20H&link_type=AUTHORSEARCH" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on PubMed</a></li><li class="author-site-search-link"><a href="/search/author1%3AHugh%2BMcColl%2B" rel="nofollow" class="" data-icon-position="" data-hide-link-title="0">Search for this author on this site</a></li><li class="author-orcid-link last"><a href="http://orcid.org/0000-0002-7568-4270" target="_blank" class="" data-icon-position="" data-hide-link-title="0">ORCID record for Hugh McColl</a></li></ul></div><div class="author-tooltip-1"><div class="author-tooltip-name">Guus Kroonen </div><div class="author-tooltip-affiliation"><span class="author-tooltip-text"><div class='author-affiliation'><span class='nlm-sup'>2</span><span class='nlm-institution'>Leiden University Centre for Linguistics, Leiden University</span>, Leiden, <span class='nlm-country'>The Netherlands</span>,</div><div class='author-affiliation'><span class='nlm-sup'>3</span><span class='nlm-institution'>Department of Nordic Studies and Linguistics, University of Copenhagen</span>, Copenhagen, <span class='nlm-country'>Denmark</span>,</div></span></div><ul class="author-tooltip-find-more"><li class="author-tooltip-gs-link first"><a href="/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=Guus%2BKroonen%2B" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on Google Scholar</a></li><li class="author-tooltip-pubmed-link"><a href="/lookup/external-ref?access_num=Kroonen%20G&link_type=AUTHORSEARCH" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on PubMed</a></li><li class="author-site-search-link"><a href="/search/author1%3AGuus%2BKroonen%2B" rel="nofollow" class="" data-icon-position="" data-hide-link-title="0">Search for this author on this site</a></li><li class="author-orcid-link last"><a href="http://orcid.org/0000-0002-3708-0476" target="_blank" class="" data-icon-position="" data-hide-link-title="0">ORCID record for Guus Kroonen</a></li></ul></div><div class="author-tooltip-2"><div class="author-tooltip-name">J. Víctor Moreno-Mayar </div><div class="author-tooltip-affiliation"><span class="author-tooltip-text"><div class='author-affiliation'><span class='nlm-sup'>1</span><span class='nlm-institution'>Lundbeck Foundation GeoGenetics Center, Globe Institute, University of Copenhagen</span>, Copenhagen, <span class='nlm-country'>Denmark</span>.,</div></span></div><ul class="author-tooltip-find-more"><li class="author-tooltip-gs-link first"><a href="/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=J.%2BV%C3%ADctor%2BMoreno-Mayar%2B" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on Google Scholar</a></li><li class="author-tooltip-pubmed-link"><a href="/lookup/external-ref?access_num=Moreno-Mayar%20JV&link_type=AUTHORSEARCH" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on PubMed</a></li><li class="author-site-search-link last"><a 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class="author-tooltip-53"><div class="author-tooltip-name">Stijn Heeren </div><div class="author-tooltip-affiliation"><span class="author-tooltip-text"><div class='author-affiliation'><span class='nlm-sup'>39</span><span class='nlm-institution'>Vrije Universiteit Amsterdam</span>, <span class='nlm-country'>the Netherlands</span>,</div></span></div><ul class="author-tooltip-find-more"><li class="author-tooltip-gs-link first"><a href="/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=Stijn%2BHeeren%2B" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on Google Scholar</a></li><li class="author-tooltip-pubmed-link"><a href="/lookup/external-ref?access_num=Heeren%20S&link_type=AUTHORSEARCH" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on PubMed</a></li><li class="author-site-search-link"><a href="/search/author1%3AStijn%2BHeeren%2B" rel="nofollow" class="" data-icon-position="" 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class='author-affiliation'><span class='nlm-sup'>1</span><span class='nlm-institution'>Lundbeck Foundation GeoGenetics Center, Globe Institute, University of Copenhagen</span>, Copenhagen, <span class='nlm-country'>Denmark</span>.,</div><div class='author-affiliation'><span class='nlm-sup'>16</span><span class='nlm-institution'>Department of Historical Studies, University of Gothenburg</span>, <span class='nlm-country'>Sweden</span>,</div></span></div><ul class="author-tooltip-find-more"><li class="author-tooltip-gs-link first"><a href="/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=Kristian%2BKristiansen%2B" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on Google Scholar</a></li><li class="author-tooltip-pubmed-link"><a href="/lookup/external-ref?access_num=Kristiansen%20K&link_type=AUTHORSEARCH" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on PubMed</a></li><li 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href="/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=Martin%2BSikora%2B" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on Google Scholar</a></li><li class="author-tooltip-pubmed-link"><a href="/lookup/external-ref?access_num=Sikora%20M&link_type=AUTHORSEARCH" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on PubMed</a></li><li class="author-site-search-link"><a href="/search/author1%3AMartin%2BSikora%2B" rel="nofollow" class="" data-icon-position="" data-hide-link-title="0">Search for this author on this site</a></li><li class="author-orcid-link last"><a href="http://orcid.org/0000-0003-2818-8319" target="_blank" class="" data-icon-position="" data-hide-link-title="0">ORCID record for Martin Sikora</a></li></ul></div><div class="author-tooltip-115"><div class="author-tooltip-name">Eske Willerslev </div><div class="author-tooltip-affiliation"><span class="author-tooltip-text"><div class='author-affiliation'><span class='nlm-sup'>1</span><span class='nlm-institution'>Lundbeck Foundation GeoGenetics Center, Globe Institute, University of Copenhagen</span>, Copenhagen, <span class='nlm-country'>Denmark</span>.,</div><div class='author-affiliation'><span class='nlm-sup'>90</span><span class='nlm-institution'>GeoGenetics Group, Department of Zoology, University of Cambridge</span>, Cambridge, <span class='nlm-country'>UK</span></div></span></div><ul class="author-tooltip-find-more"><li class="author-tooltip-gs-link first"><a href="/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=Eske%2BWillerslev%2B" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on Google Scholar</a></li><li class="author-tooltip-pubmed-link"><a href="/lookup/external-ref?access_num=Willerslev%20E&link_type=AUTHORSEARCH" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find 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pane-highwire-markup" > <div class="pane-content"> <div class="highwire-markup"><div xmlns="http://www.w3.org/1999/xhtml" data-highwire-cite-ref-tooltip-instance="highwire_reflinks_tooltip" class="content-block-markup" xmlns:xhtml="http://www.w3.org/1999/xhtml"><div class="article abstract-view "><span class="highwire-journal-article-marker-start"></span><div class="section abstract" id="abstract-1"><h2 class="">Summary</h2><p id="p-2">Germanic-speaking populations historically form an integral component of the North and Northwest European cultural configuration. According to linguistic consensus, the common ancestor of the Germanic languages, which include German, English, Frisian, Dutch as well as the Nordic languages, was spoken in Northern Europe during the Pre-Roman Iron Age. However, important questions remain concerning the earlier Bronze Age distribution of this Indo-European language branch in Scandinavia as well as the driving factors behind its Late Iron Age diversification and expansion across the European continent. A key difficulty in addressing these questions are the existence of striking differences in the interpretation of the archaeological record, leading to various hypotheses of correlations with linguistic dispersals and changes in material culture. Moreover, these interpretations have been difficult to assess using genomics due to limited ancient genomes and the difficulty in differentiating closely related populations. Here we integrate multidisciplinary evidence from population genomics, historical sources, archaeology and linguistics to offer a fully revised model for the origins and spread of Germanic languages and for the formation of the genomic ancestry of Germanic-speaking northern European populations, while acknowledging that coordinating archaeology, linguistics and genetics is complex and potentially controversial. We sequenced 710 ancient human genomes from western Eurasia and analysed them together with 3,940 published genomes suitable for imputing diploid genotypes. We find evidence of a previously unknown, large-scale Bronze Age migration within Scandinavia, originating in the east and becoming widespread to the west and south, thus providing a new potential driving factor for the expansion of the Germanic speech community. This East Scandinavian genetic cluster is first seen 800 years after the arrival of the Corded Ware Culture, the first Steppe-related population to emerge in Northern Europe, opening a new scenario implying a Late rather than an Middle Neolithic arrival of the Germanic language group in Scandinavia. Moreover, the non-local Hunter-Gatherer ancestry of this East Scandinavian cluster is indicative of a cross-Baltic maritime rather than a southern Scandinavian land-based entry. Later in the Iron Age around 1700 BP, we find a southward push of admixed Eastern and Southern Scandinavians into areas including Germany and the Netherlands, previously associated with Celtic speakers, mixing with local populations from the Eastern North Sea coast. During the Migration Period (1575-1200 BP), we find evidence of this structured, admixed Southern Scandinavian population representing the Western Germanic Anglo-Saxon migrations into Britain and Langobards into southern Europe. During the Migration Period, we detect a previously unknown northward migration back into Southern Scandinavia, partly replacing earlier inhabitants and forming the North Germanic-speaking Viking-Age populations of Denmark and southern Sweden, corresponding with historically attested Danes. However, the origin and character of these major changes in Scandinavia before the Viking Age remain contested. In contrast to these Western and Northern Germanic-speaking populations, we find the Wielbark population from Poland to be primarily of Eastern Scandinavian ancestry, supporting a Swedish origin for East Germanic groups. In contrast, the later cultural descendants, the Ostrogoths and Visigoths are predominantly of Southern European ancestry implying the adoption of Gothic culture. Together, these results highlight the use of archaeology, linguistics and genetics as distinct but complementary lines of evidence.</p></div><h3>Competing Interest Statement</h3><p id="p-3">The authors have declared no competing interest.</p><span class="highwire-journal-article-marker-end"></span></div><span class="related-urls"></span></div></div> </div> </div> <div class="panel-separator"></div><div class="panel-pane pane-biorxiv-copyright" > <div class="pane-content"> <div class="field field-name-field-highwire-copyright field-type-text field-label-inline clearfix"><div class="field-label">Copyright </div><div class="field-items"><div class="field-item even">The copyright holder for this preprint is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity.<span class="license-type"> It is made available under a <a href="http://creativecommons.org/licenses/by-nc-nd/4.0/" class="" data-icon-position="" data-hide-link-title="0">CC-BY-NC-ND 4.0 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<div class="inside"><div class="panel-pane pane-highwire-article-citation" > <div class="pane-content"> <div class="highwire-article-citation highwire-citation-type-highwire-article node3713187--3" data-node-nid="3713187" id="node-3713187--4621701837" data-pisa="biorxiv;2024.03.13.584607v1" data-pisa-master="biorxiv;2024.03.13.584607" data-seqnum="3713187" data-apath="/biorxiv/early/2024/03/14/2024.03.13.584607.atom"><div class="highwire-cite highwire-cite-highwire-article highwire-citation-biorxiv-article-pap-list clearfix" > <div class="highwire-cite-title" > <div class="highwire-cite-title">Steppe Ancestry in western Eurasia and the spread of the Germanic Languages</div> </div> <div class="highwire-cite-authors" ><span class="highwire-citation-authors"><span class="highwire-citation-author first" data-delta="0"><span class="nlm-given-names">Hugh</span> <span class="nlm-surname">McColl</span></span>, <span class="highwire-citation-author" data-delta="1"><span class="nlm-given-names">Guus</span> <span class="nlm-surname">Kroonen</span></span>, <span class="highwire-citation-author" data-delta="2"><span class="nlm-given-names">J. Víctor</span> <span class="nlm-surname">Moreno-Mayar</span></span>, <span class="highwire-citation-author" data-delta="3"><span class="nlm-given-names">Frederik</span> <span class="nlm-surname">Valeur Seersholm</span></span>, <span class="highwire-citation-author" data-delta="4"><span class="nlm-given-names">Gabriele</span> <span class="nlm-surname">Scorrano</span></span>, <span class="highwire-citation-author" data-delta="5"><span class="nlm-given-names">Thomaz</span> <span class="nlm-surname">Pinotti</span></span>, <span class="highwire-citation-author" data-delta="6"><span class="nlm-given-names">Tharsika</span> <span class="nlm-surname">Vimala</span></span>, <span class="highwire-citation-author" data-delta="7"><span class="nlm-given-names">Søren M.</span> <span class="nlm-surname">Sindbæk</span></span>, <span class="highwire-citation-author" data-delta="8"><span class="nlm-given-names">Per</span> <span class="nlm-surname">Ethelberg</span></span>, <span class="highwire-citation-author" data-delta="9"><span class="nlm-given-names">Ralph</span> <span class="nlm-surname">Fyfe</span></span>, <span class="highwire-citation-author" data-delta="10"><span class="nlm-given-names">Marie-José</span> <span class="nlm-surname">Gaillard</span></span>, <span class="highwire-citation-author" data-delta="11"><span class="nlm-given-names">Hanne M. Ellegård</span> <span class="nlm-surname">Larsen</span></span>, <span class="highwire-citation-author" data-delta="12"><span class="nlm-given-names">Morten Fischer</span> <span class="nlm-surname">Mortensen</span></span>, <span class="highwire-citation-author" data-delta="13"><span class="nlm-given-names">Fabrice</span> <span class="nlm-surname">Demeter</span></span>, <span class="highwire-citation-author" data-delta="14"><span class="nlm-given-names">Marie Louise S.</span> <span class="nlm-surname">Jørkov</span></span>, <span class="highwire-citation-author" data-delta="15"><span class="nlm-given-names">Sophie</span> <span class="nlm-surname">Bergerbrant</span></span>, <span class="highwire-citation-author" data-delta="16"><span class="nlm-given-names">Peter de Barros</span> <span class="nlm-surname">Damgaard</span></span>, <span class="highwire-citation-author" data-delta="17"><span class="nlm-given-names">Morten E.</span> <span class="nlm-surname">Allentoft</span></span>, <span class="highwire-citation-author" data-delta="18"><span class="nlm-given-names">Lasse</span> <span class="nlm-surname">Vinner</span></span>, <span class="highwire-citation-author" data-delta="19"><span class="nlm-given-names">Charleen</span> <span class="nlm-surname">Gaunitz</span></span>, <span class="highwire-citation-author" data-delta="20"><span class="nlm-given-names">Abigail</span> <span class="nlm-surname">Ramsøe</span></span>, <span class="highwire-citation-author" data-delta="21"><span class="nlm-given-names">Isin</span> <span class="nlm-surname">Altinkaya</span></span>, <span class="highwire-citation-author" data-delta="22"><span class="nlm-given-names">Rasmus</span> <span class="nlm-surname">Amund Henriksen</span></span>, <span class="highwire-citation-author" data-delta="23"><span class="nlm-given-names">Evan K.</span> <span class="nlm-surname">Irving-Pease</span></span>, <span class="highwire-citation-author" data-delta="24"><span class="nlm-given-names">Serena</span> <span class="nlm-surname">Sabatini</span></span>, <span class="highwire-citation-author" data-delta="25"><span class="nlm-given-names">Anders</span> <span class="nlm-surname">Fischer</span></span>, <span class="highwire-citation-author" data-delta="26"><span class="nlm-given-names">William</span> <span class="nlm-surname">Barrie</span></span>, <span class="highwire-citation-author" data-delta="27"><span class="nlm-given-names">Andrés</span> <span class="nlm-surname">Ingason</span></span>, <span class="highwire-citation-author" data-delta="28"><span class="nlm-given-names">Anders</span> <span class="nlm-surname">Rosengren</span></span>, <span class="highwire-citation-author" data-delta="29"><span class="nlm-given-names">Andrew</span> <span class="nlm-surname">Vaughn</span></span>, <span class="highwire-citation-author" data-delta="30"><span class="nlm-given-names">Jialu</span> <span class="nlm-surname">Cao</span></span>, <span class="highwire-citation-author" data-delta="31"><span class="nlm-given-names">Jacqueline</span> <span class="nlm-surname">Staring</span></span>, <span class="highwire-citation-author" data-delta="32"><span class="nlm-given-names">Jesper</span> <span class="nlm-surname">Stenderup</span></span>, <span class="highwire-citation-author" data-delta="33"><span class="nlm-given-names">Fulya Eylem</span> <span class="nlm-surname">Yediay</span></span>, <span class="highwire-citation-author" data-delta="34"><span class="nlm-given-names">Torbjörn</span> <span class="nlm-surname">Ahlström</span></span>, <span class="highwire-citation-author" data-delta="35"><span class="nlm-given-names">Sofie Laurine</span> <span class="nlm-surname">Albris</span></span>, <span class="highwire-citation-author" data-delta="36"><span class="nlm-given-names">Biyaslan</span> <span class="nlm-surname">Atabiev</span></span>, <span class="highwire-citation-author" data-delta="37"><span class="nlm-given-names">Pernille</span> <span class="nlm-surname">Bangsgaard</span></span>, <span class="highwire-citation-author" data-delta="38"><span class="nlm-given-names">Maria Giovanna</span> <span class="nlm-surname">Belcastro</span></span>, <span class="highwire-citation-author" data-delta="39"><span class="nlm-given-names">Nick</span> <span class="nlm-surname">Card</span></span>, <span class="highwire-citation-author" data-delta="40"><span class="nlm-given-names">Philippe</span> <span class="nlm-surname">Charlier</span></span>, <span class="highwire-citation-author" data-delta="41"><span class="nlm-given-names">Elizaveta</span> <span class="nlm-surname">Chernykh</span></span>, <span class="highwire-citation-author" data-delta="42"><span class="nlm-given-names">Torben Trier</span> <span class="nlm-surname">Christiansen</span></span>, <span class="highwire-citation-author" data-delta="43"><span class="nlm-given-names">Alfredo</span> <span class="nlm-surname">Coppa</span></span>, <span class="highwire-citation-author" data-delta="44"><span class="nlm-given-names">Maura</span> <span class="nlm-surname">De Coster</span></span>, <span class="highwire-citation-author" data-delta="45"><span class="nlm-given-names">Sean Dexter</span> <span class="nlm-surname">Denham</span></span>, <span class="highwire-citation-author" data-delta="46"><span class="nlm-given-names">Sophie</span> <span class="nlm-surname">Desenne</span></span>, <span class="highwire-citation-author" data-delta="47"><span class="nlm-given-names">Jane</span> <span class="nlm-surname">Downes</span></span>, <span class="highwire-citation-author" data-delta="48"><span class="nlm-given-names">Karin Margarita</span> <span class="nlm-surname">Frei</span></span>, <span class="highwire-citation-author" data-delta="49"><span class="nlm-given-names">Olivér</span> <span class="nlm-surname">Gábor</span></span>, <span class="highwire-citation-author" data-delta="50"><span class="nlm-given-names">Johan Zakarias</span> <span class="nlm-surname">Gårdsvoll</span></span>, <span class="highwire-citation-author" data-delta="51"><span class="nlm-given-names">Zanette Tsigaridas</span> <span class="nlm-surname">Glørstad</span></span>, <span class="highwire-citation-author" data-delta="52"><span class="nlm-given-names">Jesper</span> <span class="nlm-surname">Hansen</span></span>, <span class="highwire-citation-author" data-delta="53"><span class="nlm-given-names">Stijn</span> <span class="nlm-surname">Heeren</span></span>, <span class="highwire-citation-author" data-delta="54"><span class="nlm-given-names">Merete</span> <span class="nlm-surname">Henriksen</span></span>, <span class="highwire-citation-author" data-delta="55"><span class="nlm-given-names">Volker</span> <span class="nlm-surname">Heyd</span></span>, <span class="highwire-citation-author" data-delta="56"><span class="nlm-given-names">Mette</span> <span class="nlm-surname">Høj</span></span>, <span class="highwire-citation-author" data-delta="57"><span class="nlm-given-names">Mads Kähler</span> <span class="nlm-surname">Holst</span></span>, <span class="highwire-citation-author" data-delta="58"><span class="nlm-given-names">Rimantas</span> <span class="nlm-surname">Jankauskas</span></span>, <span class="highwire-citation-author" data-delta="59"><span class="nlm-given-names">Henrik</span> <span class="nlm-surname">Janson</span></span>, <span class="highwire-citation-author" data-delta="60"><span class="nlm-given-names">Mads Dengsø</span> <span class="nlm-surname">Jessen</span></span>, <span class="highwire-citation-author" data-delta="61"><span class="nlm-given-names">Jens Winther</span> <span class="nlm-surname">Johannsen</span></span>, <span class="highwire-citation-author" data-delta="62"><span class="nlm-given-names">Torkel</span> <span class="nlm-surname">Johansen</span></span>, <span class="highwire-citation-author" data-delta="63"><span class="nlm-given-names">Ole Thirup</span> <span class="nlm-surname">Kastholm</span></span>, <span class="highwire-citation-author" data-delta="64"><span class="nlm-given-names">Anton</span> <span class="nlm-surname">Kern</span></span>, <span class="highwire-citation-author" data-delta="65"><span class="nlm-given-names">Ruslan</span> <span class="nlm-surname">Khaskhanov</span></span>, <span class="highwire-citation-author" data-delta="66"><span class="nlm-given-names">Katrine</span> <span class="nlm-surname">Kjær</span></span>, <span class="highwire-citation-author" data-delta="67"><span class="nlm-given-names">Vladimir</span> <span class="nlm-surname">Kolosov</span></span>, <span class="highwire-citation-author" data-delta="68"><span class="nlm-given-names">Lisette M.</span> <span class="nlm-surname">Kootker</span></span>, <span class="highwire-citation-author" data-delta="69"><span class="nlm-given-names">Anne Christine</span> <span class="nlm-surname">Larsen</span></span>, <span class="highwire-citation-author" data-delta="70"><span class="nlm-given-names">Thierry</span> <span class="nlm-surname">Lejars</span></span>, <span class="highwire-citation-author" data-delta="71"><span class="nlm-given-names">Mette</span> <span class="nlm-surname">Løvschal</span></span>, <span class="highwire-citation-author" data-delta="72"><span class="nlm-given-names">Niels</span> <span class="nlm-surname">Lynnerup</span></span>, <span class="highwire-citation-author" data-delta="73"><span class="nlm-given-names">Yvonne</span> <span class="nlm-surname">Magnusson</span></span>, <span class="highwire-citation-author" data-delta="74"><span class="nlm-given-names">Kristiina</span> <span class="nlm-surname">Mannermaa</span></span>, <span class="highwire-citation-author" data-delta="75"><span class="nlm-given-names">Vyacheslav</span> <span class="nlm-surname">Masyakin</span></span>, <span class="highwire-citation-author" data-delta="76"><span class="nlm-given-names">Anne Lene</span> <span class="nlm-surname">Melheim</span></span>, <span class="highwire-citation-author" data-delta="77"><span class="nlm-given-names">Inga</span> <span class="nlm-surname">Merkyte</span></span>, <span class="highwire-citation-author" data-delta="78"><span class="nlm-given-names">Vyacheslav</span> <span class="nlm-surname">Moiseyev</span></span>, <span class="highwire-citation-author" data-delta="79"><span class="nlm-given-names">Stig Bergmann</span> <span class="nlm-surname">Møller</span></span>, <span class="highwire-citation-author" data-delta="80"><span class="nlm-given-names">Erika</span> <span class="nlm-surname">Molnár</span></span>, <span class="highwire-citation-author" data-delta="81"><span class="nlm-given-names">Nadja</span> <span class="nlm-surname">Mortensen</span></span>, <span class="highwire-citation-author" data-delta="82"><span class="nlm-given-names">Eileen</span> <span class="nlm-surname">Murphy</span></span>, <span class="highwire-citation-author" data-delta="83"><span class="nlm-given-names">Bjarne Henning</span> <span class="nlm-surname">Nielsen</span></span>, <span class="highwire-citation-author" data-delta="84"><span class="nlm-given-names">Doris</span> <span class="nlm-surname">Pany-Kucera</span></span>, <span class="highwire-citation-author" data-delta="85"><span class="nlm-given-names">Bettina Schulz</span> <span class="nlm-surname">Paulsson</span></span>, <span class="highwire-citation-author" data-delta="86"><span class="nlm-given-names">Marcia S</span> <span class="nlm-surname">Ponce de León</span></span>, <span class="highwire-citation-author" data-delta="87"><span class="nlm-given-names">Håkon</span> <span class="nlm-surname">Reiersen</span></span>, <span class="highwire-citation-author" data-delta="88"><span class="nlm-given-names">Walter</span> <span class="nlm-surname">Reinhard</span></span>, <span class="highwire-citation-author" data-delta="89"><span class="nlm-given-names">Antti</span> <span class="nlm-surname">Sajantila</span></span>, <span class="highwire-citation-author" data-delta="90"><span class="nlm-given-names">Birgitte</span> <span class="nlm-surname">Skar</span></span>, <span class="highwire-citation-author" data-delta="91"><span class="nlm-given-names">Vladimir</span> <span class="nlm-surname">Slavchev</span></span>, <span class="highwire-citation-author" data-delta="92"><span class="nlm-given-names">Václav</span> <span class="nlm-surname">Smrčka</span></span>, <span class="highwire-citation-author" data-delta="93"><span class="nlm-given-names">Lasse</span> <span class="nlm-surname">Sørensen</span></span>, <span class="highwire-citation-author" data-delta="94"><span class="nlm-given-names">Georg</span> <span class="nlm-surname">Tiefengraber</span></span>, <span class="highwire-citation-author" data-delta="95"><span class="nlm-given-names">Otto Christian</span> <span class="nlm-surname">Uldum</span></span>, <span class="highwire-citation-author" data-delta="96"><span class="nlm-given-names">Jorge</span> <span class="nlm-surname">Vega</span></span>, <span class="highwire-citation-author" data-delta="97"><span class="nlm-given-names">Daniele</span> <span class="nlm-surname">Vitali</span></span>, <span class="highwire-citation-author" data-delta="98"><span class="nlm-given-names">Alexey</span> <span class="nlm-surname">Voloshinov</span></span>, <span class="highwire-citation-author" data-delta="99"><span class="nlm-given-names">Sidsel</span> <span class="nlm-surname">Wåhlin</span></span>, <span class="highwire-citation-author" data-delta="100"><span class="nlm-given-names">Holger</span> <span class="nlm-surname">Wendling</span></span>, <span class="highwire-citation-author" data-delta="101"><span class="nlm-given-names">Anna</span> <span class="nlm-surname">Wessman</span></span>, <span class="highwire-citation-author" data-delta="102"><span class="nlm-given-names">Helene</span> <span class="nlm-surname">Wilhelmson</span></span>, <span class="highwire-citation-author" data-delta="103"><span class="nlm-given-names">Karin</span> <span class="nlm-surname">Wiltschke</span></span>, <span class="highwire-citation-author" data-delta="104"><span class="nlm-given-names">João</span> <span class="nlm-surname">Zilhao</span></span>, <span class="highwire-citation-author" data-delta="105"><span class="nlm-given-names">Christoph PE</span> <span class="nlm-surname">Zollikofer</span></span>, <span class="highwire-citation-author" data-delta="106"><span class="nlm-given-names">Thorfinn</span> <span class="nlm-surname">Sand Korneliussen</span></span>, <span class="highwire-citation-author" data-delta="107"><span class="nlm-given-names">Bruno</span> <span class="nlm-surname">Chaume</span></span>, <span class="highwire-citation-author" data-delta="108"><span class="nlm-given-names">Jean-Paul</span> <span class="nlm-surname">Demoule</span></span>, <span class="highwire-citation-author" data-delta="109"><span class="nlm-given-names">Thomas</span> <span class="nlm-surname">Werge</span></span>, <span class="highwire-citation-author" data-delta="110"><span class="nlm-given-names">Line</span> <span class="nlm-surname">Olsen</span></span>, <span class="highwire-citation-author" data-delta="111"><span class="nlm-given-names">Rasmus</span> <span class="nlm-surname">Nielsen</span></span>, <span class="highwire-citation-author" data-delta="112"><span class="nlm-given-names">Lotte</span> <span class="nlm-surname">Hedeager</span></span>, <span class="highwire-citation-author" data-delta="113"><span class="nlm-given-names">Kristian</span> <span class="nlm-surname">Kristiansen</span></span>, <span class="highwire-citation-author" data-delta="114"><span class="nlm-given-names">Martin</span> <span class="nlm-surname">Sikora</span></span>, <span class="highwire-citation-author" data-delta="115"><span class="nlm-given-names">Eske</span> <span class="nlm-surname">Willerslev</span></span></span></div> <div class="highwire-cite-metadata" ><span class="highwire-cite-metadata-journal highwire-cite-metadata">bioRxiv </span><span class="highwire-cite-metadata-pages highwire-cite-metadata">2024.03.13.584607; </span><span class="highwire-cite-metadata-doi highwire-cite-metadata"><span class="doi_label">doi:</span> https://doi.org/10.1101/2024.03.13.584607 </span></div> </div> </div> </div> </div> </div> </div> <div class="panel-panel grid-12 panel-region-content"> <div class="inside"><div class="panel-pane pane-highwire-article-clipboard-copy" > <div class="pane-content"> <div class = "clipboard-copy"> <span class="label-url"> <label for="dynamic">Share This Article:</label> </span> <span class="input-text-url"> <input type="text" id="dynamic" value="https://www.biorxiv.org/content/10.1101/2024.03.13.584607v1" size="50"/> </span> <span class="copy-button button"> <button id="copy-dynamic" class="clipboardjs-button" data-clipboard-target="#dynamic" data-clipboard-alert-style="tooltip" data-clipboard-alert-text="Copied!">Copy</button> </span> </div> </div> </div> </div> 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highwire-citation-type-highwire-article cite-tool-node3713187--5" data-node-nid="3713187" id="citation-node-3713187--61656010386" data-pisa="biorxiv;2024.03.13.584607v1" data-pisa-master="biorxiv;2024.03.13.584607" data-seqnum="3713187" data-apath="/biorxiv/early/2024/03/14/2024.03.13.584607.atom"><div class="highwire-cite highwire-cite-highwire-article highwire-citation-biorxiv-article-pap-list clearfix" > <div class="highwire-cite-title" > <div class="highwire-cite-title">Steppe Ancestry in western Eurasia and the spread of the Germanic Languages</div> </div> <div class="highwire-cite-authors" ><span class="highwire-citation-authors"><span class="highwire-citation-author first" data-delta="0"><span class="nlm-given-names">Hugh</span> <span class="nlm-surname">McColl</span></span>, <span class="highwire-citation-author" data-delta="1"><span class="nlm-given-names">Guus</span> <span class="nlm-surname">Kroonen</span></span>, <span class="highwire-citation-author" data-delta="2"><span class="nlm-given-names">J. Víctor</span> <span class="nlm-surname">Moreno-Mayar</span></span>, <span class="highwire-citation-author" data-delta="3"><span class="nlm-given-names">Frederik</span> <span class="nlm-surname">Valeur Seersholm</span></span>, <span class="highwire-citation-author" data-delta="4"><span class="nlm-given-names">Gabriele</span> <span class="nlm-surname">Scorrano</span></span>, <span class="highwire-citation-author" data-delta="5"><span class="nlm-given-names">Thomaz</span> <span class="nlm-surname">Pinotti</span></span>, <span class="highwire-citation-author" data-delta="6"><span class="nlm-given-names">Tharsika</span> <span class="nlm-surname">Vimala</span></span>, <span class="highwire-citation-author" data-delta="7"><span class="nlm-given-names">Søren M.</span> <span class="nlm-surname">Sindbæk</span></span>, <span class="highwire-citation-author" data-delta="8"><span class="nlm-given-names">Per</span> <span class="nlm-surname">Ethelberg</span></span>, <span class="highwire-citation-author" data-delta="9"><span class="nlm-given-names">Ralph</span> <span class="nlm-surname">Fyfe</span></span>, <span class="highwire-citation-author" data-delta="10"><span class="nlm-given-names">Marie-José</span> <span class="nlm-surname">Gaillard</span></span>, <span class="highwire-citation-author" data-delta="11"><span class="nlm-given-names">Hanne M. Ellegård</span> <span class="nlm-surname">Larsen</span></span>, <span class="highwire-citation-author" data-delta="12"><span class="nlm-given-names">Morten Fischer</span> <span class="nlm-surname">Mortensen</span></span>, <span class="highwire-citation-author" data-delta="13"><span class="nlm-given-names">Fabrice</span> <span class="nlm-surname">Demeter</span></span>, <span class="highwire-citation-author" data-delta="14"><span class="nlm-given-names">Marie Louise S.</span> <span class="nlm-surname">Jørkov</span></span>, <span class="highwire-citation-author" data-delta="15"><span class="nlm-given-names">Sophie</span> <span class="nlm-surname">Bergerbrant</span></span>, <span class="highwire-citation-author" data-delta="16"><span class="nlm-given-names">Peter de Barros</span> <span class="nlm-surname">Damgaard</span></span>, <span class="highwire-citation-author" data-delta="17"><span class="nlm-given-names">Morten E.</span> <span class="nlm-surname">Allentoft</span></span>, <span class="highwire-citation-author" data-delta="18"><span class="nlm-given-names">Lasse</span> <span class="nlm-surname">Vinner</span></span>, <span class="highwire-citation-author" data-delta="19"><span class="nlm-given-names">Charleen</span> <span class="nlm-surname">Gaunitz</span></span>, <span class="highwire-citation-author" data-delta="20"><span class="nlm-given-names">Abigail</span> <span class="nlm-surname">Ramsøe</span></span>, <span class="highwire-citation-author" data-delta="21"><span class="nlm-given-names">Isin</span> <span class="nlm-surname">Altinkaya</span></span>, <span class="highwire-citation-author" data-delta="22"><span class="nlm-given-names">Rasmus</span> <span class="nlm-surname">Amund Henriksen</span></span>, <span class="highwire-citation-author" data-delta="23"><span class="nlm-given-names">Evan K.</span> <span class="nlm-surname">Irving-Pease</span></span>, <span class="highwire-citation-author" data-delta="24"><span class="nlm-given-names">Serena</span> <span class="nlm-surname">Sabatini</span></span>, <span class="highwire-citation-author" data-delta="25"><span class="nlm-given-names">Anders</span> <span class="nlm-surname">Fischer</span></span>, <span class="highwire-citation-author" data-delta="26"><span class="nlm-given-names">William</span> <span class="nlm-surname">Barrie</span></span>, <span class="highwire-citation-author" data-delta="27"><span class="nlm-given-names">Andrés</span> <span class="nlm-surname">Ingason</span></span>, <span class="highwire-citation-author" data-delta="28"><span class="nlm-given-names">Anders</span> <span class="nlm-surname">Rosengren</span></span>, <span class="highwire-citation-author" data-delta="29"><span class="nlm-given-names">Andrew</span> <span class="nlm-surname">Vaughn</span></span>, <span class="highwire-citation-author" data-delta="30"><span class="nlm-given-names">Jialu</span> <span class="nlm-surname">Cao</span></span>, <span class="highwire-citation-author" data-delta="31"><span 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data-delta="77"><span class="nlm-given-names">Inga</span> <span class="nlm-surname">Merkyte</span></span>, <span class="highwire-citation-author" data-delta="78"><span class="nlm-given-names">Vyacheslav</span> <span class="nlm-surname">Moiseyev</span></span>, <span class="highwire-citation-author" data-delta="79"><span class="nlm-given-names">Stig Bergmann</span> <span class="nlm-surname">Møller</span></span>, <span class="highwire-citation-author" data-delta="80"><span class="nlm-given-names">Erika</span> <span class="nlm-surname">Molnár</span></span>, <span class="highwire-citation-author" data-delta="81"><span class="nlm-given-names">Nadja</span> <span class="nlm-surname">Mortensen</span></span>, <span class="highwire-citation-author" data-delta="82"><span class="nlm-given-names">Eileen</span> <span class="nlm-surname">Murphy</span></span>, <span class="highwire-citation-author" data-delta="83"><span class="nlm-given-names">Bjarne Henning</span> <span class="nlm-surname">Nielsen</span></span>, <span class="highwire-citation-author" data-delta="84"><span class="nlm-given-names">Doris</span> <span class="nlm-surname">Pany-Kucera</span></span>, <span class="highwire-citation-author" data-delta="85"><span class="nlm-given-names">Bettina Schulz</span> <span class="nlm-surname">Paulsson</span></span>, <span class="highwire-citation-author" data-delta="86"><span class="nlm-given-names">Marcia S</span> <span class="nlm-surname">Ponce de León</span></span>, <span class="highwire-citation-author" data-delta="87"><span class="nlm-given-names">Håkon</span> <span class="nlm-surname">Reiersen</span></span>, <span class="highwire-citation-author" data-delta="88"><span class="nlm-given-names">Walter</span> <span class="nlm-surname">Reinhard</span></span>, <span class="highwire-citation-author" data-delta="89"><span class="nlm-given-names">Antti</span> <span class="nlm-surname">Sajantila</span></span>, <span class="highwire-citation-author" 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class="nlm-surname">Vega</span></span>, <span class="highwire-citation-author" data-delta="97"><span class="nlm-given-names">Daniele</span> <span class="nlm-surname">Vitali</span></span>, <span class="highwire-citation-author" data-delta="98"><span class="nlm-given-names">Alexey</span> <span class="nlm-surname">Voloshinov</span></span>, <span class="highwire-citation-author" data-delta="99"><span class="nlm-given-names">Sidsel</span> <span class="nlm-surname">Wåhlin</span></span>, <span class="highwire-citation-author" data-delta="100"><span class="nlm-given-names">Holger</span> <span class="nlm-surname">Wendling</span></span>, <span class="highwire-citation-author" data-delta="101"><span class="nlm-given-names">Anna</span> <span class="nlm-surname">Wessman</span></span>, <span class="highwire-citation-author" data-delta="102"><span class="nlm-given-names">Helene</span> <span class="nlm-surname">Wilhelmson</span></span>, <span class="highwire-citation-author" data-delta="103"><span class="nlm-given-names">Karin</span> <span class="nlm-surname">Wiltschke</span></span>, <span class="highwire-citation-author" data-delta="104"><span class="nlm-given-names">João</span> <span class="nlm-surname">Zilhao</span></span>, <span class="highwire-citation-author" data-delta="105"><span class="nlm-given-names">Christoph PE</span> <span class="nlm-surname">Zollikofer</span></span>, <span class="highwire-citation-author" data-delta="106"><span class="nlm-given-names">Thorfinn</span> <span class="nlm-surname">Sand Korneliussen</span></span>, <span class="highwire-citation-author" data-delta="107"><span class="nlm-given-names">Bruno</span> <span class="nlm-surname">Chaume</span></span>, <span class="highwire-citation-author" data-delta="108"><span class="nlm-given-names">Jean-Paul</span> <span class="nlm-surname">Demoule</span></span>, <span class="highwire-citation-author" data-delta="109"><span class="nlm-given-names">Thomas</span> <span class="nlm-surname">Werge</span></span>, <span class="highwire-citation-author" data-delta="110"><span class="nlm-given-names">Line</span> <span class="nlm-surname">Olsen</span></span>, <span class="highwire-citation-author" data-delta="111"><span class="nlm-given-names">Rasmus</span> <span class="nlm-surname">Nielsen</span></span>, <span class="highwire-citation-author" data-delta="112"><span class="nlm-given-names">Lotte</span> <span class="nlm-surname">Hedeager</span></span>, <span class="highwire-citation-author" data-delta="113"><span class="nlm-given-names">Kristian</span> <span class="nlm-surname">Kristiansen</span></span>, <span class="highwire-citation-author" data-delta="114"><span class="nlm-given-names">Martin</span> <span class="nlm-surname">Sikora</span></span>, <span class="highwire-citation-author" data-delta="115"><span class="nlm-given-names">Eske</span> <span class="nlm-surname">Willerslev</span></span></span></div> <div class="highwire-cite-metadata" ><span class="highwire-cite-metadata-journal highwire-cite-metadata">bioRxiv </span><span class="highwire-cite-metadata-pages highwire-cite-metadata">2024.03.13.584607; </span><span class="highwire-cite-metadata-doi highwire-cite-metadata"><span class="doi_label">doi:</span> https://doi.org/10.1101/2024.03.13.584607 </span></div> </div> </div> </div> <div class="highwire-citation-formats"> <h2>Citation Manager Formats</h2> <div class="highwire-citation-formats-links"> <span class="Z3988" 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