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Population genomics of the Viking world

Ashot Margaryan, Daniel Lawson, Martin Sikora, Fernando Racimo, Simon Rasmussen, Ida Moltke, Lara Cassidy, Emil Jørsboe, Andrés Ingason, Mikkel Pedersen, Thorfinn Korneliussen, Helene Wilhelmson, Magdalena Buś, Peter de Barros Damgaard, Rui Martiniano, Gabriel Renaud, Claude Bhérer, J. Víctor Moreno-Mayar, Anna Fotakis, Marie Allen, Martyna Molak, Enrico Cappellini, Gabriele Scorrano, Alexandra Buzhilova, Allison Fox, Anders Albrechtsen, Berit Schütz, Birgitte Skar, Caroline Arcini, Ceri Falys, Charlotte Hedenstierna Jonson, Dariusz Błaszczyk, Denis Pezhemsky, Gordon Turner-Walker, Hildur Gestsdóttir, Inge Lundstrøm, Ingrid Gustin, Ingrid Mainland, Inna Potekhina, Italo Muntoni, Jade Cheng, Jesper Stenderup, Jilong Ma, Julie Gibson, Jüri Peets, Jörgen Gustafsson, Katrine Iversen, Linzi Simpson, Lisa Strand, Louise Loe, Maeve Sikora, Marek Florek, Maria Vretemark, Mark Redknap, Monika Bajka, Tamara Pushkina, Morten Søvsø, Natalia Grigoreva, Tom Christensen, Ole Kastholm, Otto Uldum, Pasquale Favia, Per Holck, Raili Allmäe, Sabine Sten, Símun Arge, Sturla Ellingvåg, Vayacheslav Moiseyev, Wiesław Bogdanowicz, Yvonne Magnusson, Ludovic Orlando, Daniel Bradley, Marie Louise Jørkov, Jette Arneborg, Niels Lynnerup, Neil Price, M. Thomas Gilbert, Morten Allentoft, Jan Bill, Søren Sindbæk, Lotte Hedeager, Kristian Kristiansen, Rasmus Nielsen, Thomas Werge, Eske Willerslev
doi: https://doi.org/10.1101/703405
Ashot Margaryan
1Lundbeck Foundation GeoGenetics Centre, GLOBE Institute, University of Copenhagen, Øster Voldgade 5-7, 1350 Copenhagen K, Denmark
2Institute of Molecular Biology, National Academy of Sciences, 7, Hasratian St., 0014, Yerevan, Armenia
3Section for Evolutionary Genomics, GLOBE Institute, University of Copenhagen, Øster Voldgade 5-7, 1350 Copenhagen K, Denmark
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Daniel Lawson
4MRC Integrative Epidemiology Unit, University of Bristol, Bristol, UK
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Martin Sikora
1Lundbeck Foundation GeoGenetics Centre, GLOBE Institute, University of Copenhagen, Øster Voldgade 5-7, 1350 Copenhagen K, Denmark
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Fernando Racimo
1Lundbeck Foundation GeoGenetics Centre, GLOBE Institute, University of Copenhagen, Øster Voldgade 5-7, 1350 Copenhagen K, Denmark
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Simon Rasmussen
5Novo Nordisk Foundation Center for Protein Research, Faculty of Health and Medical Sciences, University of Copenhagen, Blegdamsvej 3B, 2200 Copenhagen, Denmark
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Ida Moltke
6Department of Biology, The Bioinformatics Centre, University of Copenhagen, 2200 Copenhagen N, Denmark
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Lara Cassidy
7Smurfit Institute of Genetics, Trinity College Dublin, Dublin
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Emil Jørsboe
6Department of Biology, The Bioinformatics Centre, University of Copenhagen, 2200 Copenhagen N, Denmark
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Andrés Ingason
1Lundbeck Foundation GeoGenetics Centre, GLOBE Institute, University of Copenhagen, Øster Voldgade 5-7, 1350 Copenhagen K, Denmark
58Institute of Biological Psychiatry, Mental Health Services Copenhagen, Copenhagen, Denmark
59The Lundbeck Foundation Initiative for Integrative Psychiatric Research, iPSYCH, Denmark
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Mikkel Pedersen
1Lundbeck Foundation GeoGenetics Centre, GLOBE Institute, University of Copenhagen, Øster Voldgade 5-7, 1350 Copenhagen K, Denmark
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Thorfinn Korneliussen
1Lundbeck Foundation GeoGenetics Centre, GLOBE Institute, University of Copenhagen, Øster Voldgade 5-7, 1350 Copenhagen K, Denmark
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Helene Wilhelmson
8Historical archaeology, Department of Archaeology and Ancient history, Lund University, PB 192, SE 22100 Lund, Sweden
9Sydsvensk arkeologi AB, PB 134, SE 29122 Kristianstad, Sweden
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Magdalena Buś
10Department of Immunology, Genetics and Pathology, Science for Life Laboratory, Uppsala University, 751 08 Uppsala, Sweden
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Peter de Barros Damgaard
1Lundbeck Foundation GeoGenetics Centre, GLOBE Institute, University of Copenhagen, Øster Voldgade 5-7, 1350 Copenhagen K, Denmark
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Rui Martiniano
11Department of Genetics, University of Cambridge, Downing Street, Cambridge CB2 3EH, UK
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Gabriel Renaud
1Lundbeck Foundation GeoGenetics Centre, GLOBE Institute, University of Copenhagen, Øster Voldgade 5-7, 1350 Copenhagen K, Denmark
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Claude Bhérer
12New York Genome Center, 101 Avenue of the Americas, New York, NY, USA, 10013
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J. Víctor Moreno-Mayar
1Lundbeck Foundation GeoGenetics Centre, GLOBE Institute, University of Copenhagen, Øster Voldgade 5-7, 1350 Copenhagen K, Denmark
13National Institute of Genomic Medicine (INMEGEN), Periférico Sur 4809, 14610 Mexico City, Mexico
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Anna Fotakis
3Section for Evolutionary Genomics, GLOBE Institute, University of Copenhagen, Øster Voldgade 5-7, 1350 Copenhagen K, Denmark
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Marie Allen
10Department of Immunology, Genetics and Pathology, Science for Life Laboratory, Uppsala University, 751 08 Uppsala, Sweden
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Martyna Molak
14Museum and Institute of Zoology, Polish Academy of Sciences, Wilcza 64, 00-679 Warsaw, Poland
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Enrico Cappellini
3Section for Evolutionary Genomics, GLOBE Institute, University of Copenhagen, Øster Voldgade 5-7, 1350 Copenhagen K, Denmark
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Gabriele Scorrano
3Section for Evolutionary Genomics, GLOBE Institute, University of Copenhagen, Øster Voldgade 5-7, 1350 Copenhagen K, Denmark
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Alexandra Buzhilova
15Anuchin Research Institute and Museum of Anthropology, Moscow State University
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Allison Fox
16Manx National Heritage, Kingswood Grove, Douglas, Isle of Man, British Isles IM1 3LY
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6Department of Biology, The Bioinformatics Centre, University of Copenhagen, 2200 Copenhagen N, Denmark
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Berit Schütz
17Upplandsmuseet, Drottninggatan 7, 753 10 Uppsala, Sweden
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Birgitte Skar
18NTNU University Museum, Department of Archaeology and Cultural History Norway
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Caroline Arcini
19Arkeologerna
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Ceri Falys
20Thames Valley Archaeological Services (TVAS), Reading, UK
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Charlotte Hedenstierna Jonson
21Department of Archaeology and Ancient History, Uppsala University, Box 626, 751 26 Uppsala, Sweden
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22Institute of Archaeology, University of Warsaw, ul. Krakowskie Przedmieście 26/28, 00-927 Warsaw, Poland
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15Anuchin Research Institute and Museum of Anthropology, Moscow State University
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23Department of Cultural Heritage Conservation, National Yunlin University of Science and Technology, Douliou, Taiwan
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24Institute of Archaeology, Iceland. Bárugata 3, 101 Reykjavík, Iceland
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3Section for Evolutionary Genomics, GLOBE Institute, University of Copenhagen, Øster Voldgade 5-7, 1350 Copenhagen K, Denmark
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Ingrid Gustin
8Historical archaeology, Department of Archaeology and Ancient history, Lund University, PB 192, SE 22100 Lund, Sweden
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25UHI Archaeology Institute, University of the Highlands and Islands, Orkney College, Kirkwall, Orkney, KW15 1LX
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27Soprintendenza Archeologia, Belle Arti e Paesaggio per le Province di Barletta - Andria - Trani e Foggia, Via Alberto Alvarez Valentini, 8 - 71121 Foggia, Italy
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Jade Cheng
1Lundbeck Foundation GeoGenetics Centre, GLOBE Institute, University of Copenhagen, Øster Voldgade 5-7, 1350 Copenhagen K, Denmark
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Jesper Stenderup
1Lundbeck Foundation GeoGenetics Centre, GLOBE Institute, University of Copenhagen, Øster Voldgade 5-7, 1350 Copenhagen K, Denmark
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Jilong Ma
1Lundbeck Foundation GeoGenetics Centre, GLOBE Institute, University of Copenhagen, Øster Voldgade 5-7, 1350 Copenhagen K, Denmark
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Julie Gibson
25UHI Archaeology Institute, University of the Highlands and Islands, Orkney College, Kirkwall, Orkney, KW15 1LX
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Jüri Peets
28Archaeological Research Collection, Tallinn University, Rüütli 10, Tallinn 10130, Estonia
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Jörgen Gustafsson
29Jönköping county museum, Jönköping, Sweden
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Katrine Iversen
5Novo Nordisk Foundation Center for Protein Research, Faculty of Health and Medical Sciences, University of Copenhagen, Blegdamsvej 3B, 2200 Copenhagen, Denmark
64Department of Health Technology, Section for Bioinformatics, Technical University of Denmark, DTU, 2800 Kgs. Lyngby, Denmark
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30Trinity College Dublin
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18NTNU University Museum, Department of Archaeology and Cultural History Norway
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Louise Loe
31Oxford Archaeology, Janus House, Osney Mead, Oxford OX2 0ES, UK
32Heritage Burial Services, Oxford Archaeology, Janus House, Osney Mead, Oxford OX2 0ES, UK
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Maeve Sikora
33National Museum of Ireland, Kildare Street, Dublin 2, Ireland
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Marek Florek
34Institute of Archaeology, Maria Curie-Sklodowska University in Lublin, Pl. M. Curie-Sklodowska 4, 20-035 Lublin, Poland
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35Västergötlands museum, Box 253, 532 23 Skara Sweden
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36National Museum Cardiff
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37“Trzy Epoki” Archaeological Service, Poland
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15Anuchin Research Institute and Museum of Anthropology, Moscow State University
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38Museum of Southwest Jutland
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Natalia Grigoreva
39Institute for the history of material culture, Russian Academy of Sciences, Dvotsovaya Emb., 18, Saint-Petersburg, Russia, 191186
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40National Museum of Denmark, Frederiksholms Kanal 12, DK-1220 Copenhagen, Denmark
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41Roskilde Museum, Museum Organization ROMU, Sankt Ols Stræde 3, DK-4000 Roskilde, Denmark
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Otto Uldum
42Langelands Museum, Jens Winthersvej 12. 5900 Rudkøbing, Langeland, Denmark
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Pasquale Favia
43Department of Humanities, University of Foggia, Via Arpi, 176, 71121 Foggia, Italy
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Per Holck
44Department of Molecular Medicine, Faculty of Medicine, University of Oslo
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28Archaeological Research Collection, Tallinn University, Rüütli 10, Tallinn 10130, Estonia
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45Department of Archaeology and Ancient History, Uppsala University Campus Gotland
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46Tjóðsavnið - Faroe Islands National Museum. Kúrdalsvegur 15. Postboks 1155. FO-110 Tórshavn
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Sturla Ellingvåg
1Lundbeck Foundation GeoGenetics Centre, GLOBE Institute, University of Copenhagen, Øster Voldgade 5-7, 1350 Copenhagen K, Denmark
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Vayacheslav Moiseyev
47Peter the Great Museum of Anthropology and Ethnography (Kunstkamera), Russian Academy of Science, University Emb, 3, SPb, Russia, 199034
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Wiesław Bogdanowicz
14Museum and Institute of Zoology, Polish Academy of Sciences, Wilcza 64, 00-679 Warsaw, Poland
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Yvonne Magnusson
48Malmö Museum, Box 406, 201 24 Malmö, Sweden
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Ludovic Orlando
49Laboratoire d’Anthropobiologie Moléculaire et d’Imagerie de Synthèse, CNRS UMR 5288, Université de Toulouse, Université Paul Sabatier, 31000 Toulouse, France
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Daniel Bradley
7Smurfit Institute of Genetics, Trinity College Dublin, Dublin
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Marie Louise Jørkov
50Department of Forensic Medicine, University of Copenhagen, Frederik V’s vej 11, 2100 Copenhagen
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Jette Arneborg
40National Museum of Denmark, Frederiksholms Kanal 12, DK-1220 Copenhagen, Denmark
63School of GeoSciences, University of Edinburgh
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Niels Lynnerup
50Department of Forensic Medicine, University of Copenhagen, Frederik V’s vej 11, 2100 Copenhagen
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Neil Price
21Department of Archaeology and Ancient History, Uppsala University, Box 626, 751 26 Uppsala, Sweden
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M. Thomas Gilbert
3Section for Evolutionary Genomics, GLOBE Institute, University of Copenhagen, Øster Voldgade 5-7, 1350 Copenhagen K, Denmark
51Department of Natural History, NTNU
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Morten Allentoft
1Lundbeck Foundation GeoGenetics Centre, GLOBE Institute, University of Copenhagen, Øster Voldgade 5-7, 1350 Copenhagen K, Denmark
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Jan Bill
52Museum of Cultural History, University of Oslo, P.O. Box 6762 St. Olavs plass, 0160 Oslo, Norway
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Søren Sindbæk
53Centre for Urban Network Evolutions (UrbNet), Aarhus University, School of Culture and Society, Moesgård Allé 20, building 4215, DK-8270 Højbjerg, Denmark
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Lotte Hedeager
54Institute of Archaeology, Conservation and History, Pb. 1019 Blindern, 0315 Oslo, Norway
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Kristian Kristiansen
55Department of Historical Studies, University of Gothenburg
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Rasmus Nielsen
1Lundbeck Foundation GeoGenetics Centre, GLOBE Institute, University of Copenhagen, Øster Voldgade 5-7, 1350 Copenhagen K, Denmark
56Departments of Integrative Biology and Statistics, UC Berkeley, Berkeley, CA 94720, USA
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Thomas Werge
1Lundbeck Foundation GeoGenetics Centre, GLOBE Institute, University of Copenhagen, Øster Voldgade 5-7, 1350 Copenhagen K, Denmark
57Department of Clinical Medicine, University of Copenhagen, Copenhagen, Denmark
58Institute of Biological Psychiatry, Mental Health Services Copenhagen, Copenhagen, Denmark
59The Lundbeck Foundation Initiative for Integrative Psychiatric Research, iPSYCH, Denmark
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Eske Willerslev
1Lundbeck Foundation GeoGenetics Centre, GLOBE Institute, University of Copenhagen, Øster Voldgade 5-7, 1350 Copenhagen K, Denmark
60Department of Zoology, University of Cambridge, UK
61The Danish Institute for Advanced Study, University of Southern Denmark
62The Wellcome Trust Sanger Institute, Cambridge, UK
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  • For correspondence: ewillerslev@snm.ku.dk
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Abstract

The Viking maritime expansion from Scandinavia (Denmark, Norway, and Sweden) marks one of the swiftest and most far-flung cultural transformations in global history. During this time (c. 750 to 1050 CE), the Vikings reached most of western Eurasia, Greenland, and North America, and left a cultural legacy that persists till today. To understand the genetic structure and influence of the Viking expansion, we sequenced the genomes of 442 ancient humans from across Europe and Greenland ranging from the Bronze Age (c. 2400 BC) to the early Modern period (c. 1600 CE), with particular emphasis on the Viking Age. We find that the period preceding the Viking Age was accompanied by foreign gene flow into Scandinavia from the south and east: spreading from Denmark and eastern Sweden to the rest of Scandinavia. Despite the close linguistic similarities of modern Scandinavian languages, we observe genetic structure within Scandinavia, suggesting that regional population differences were already present 1,000 years ago. We find evidence for a majority of Danish Viking presence in England, Swedish Viking presence in the Baltic, and Norwegian Viking presence in Ireland, Iceland, and Greenland. Additionally, we see substantial foreign European ancestry entering Scandinavia during the Viking Age. We also find that several of the members of the only archaeologically well-attested Viking expedition were close family members. By comparing Viking Scandinavian genomes with present-day Scandinavian genomes, we find that pigmentation-associated loci have undergone strong population differentiation during the last millennia. Finally, we are able to trace the allele frequency dynamics of positively selected loci with unprecedented detail, including the lactase persistence allele and various alleles associated with the immune response. We conclude that the Viking diaspora was characterized by substantial foreign engagement: distinct Viking populations influenced the genomic makeup of different regions of Europe, while Scandinavia also experienced increased contact with the rest of the continent.

Footnotes

  • ↵† e-mail: ewillerslev{at}snm.ku.dk; Thomas.Werge{at}regionh.dk; rasmus_nielsen{at}berkeley.edu

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The copyright holder for this preprint is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under a CC-BY-NC-ND 4.0 International license.
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Population genomics of the Viking world
Ashot Margaryan, Daniel Lawson, Martin Sikora, Fernando Racimo, Simon Rasmussen, Ida Moltke, Lara Cassidy, Emil Jørsboe, Andrés Ingason, Mikkel Pedersen, Thorfinn Korneliussen, Helene Wilhelmson, Magdalena Buś, Peter de Barros Damgaard, Rui Martiniano, Gabriel Renaud, Claude Bhérer, J. Víctor Moreno-Mayar, Anna Fotakis, Marie Allen, Martyna Molak, Enrico Cappellini, Gabriele Scorrano, Alexandra Buzhilova, Allison Fox, Anders Albrechtsen, Berit Schütz, Birgitte Skar, Caroline Arcini, Ceri Falys, Charlotte Hedenstierna Jonson, Dariusz Błaszczyk, Denis Pezhemsky, Gordon Turner-Walker, Hildur Gestsdóttir, Inge Lundstrøm, Ingrid Gustin, Ingrid Mainland, Inna Potekhina, Italo Muntoni, Jade Cheng, Jesper Stenderup, Jilong Ma, Julie Gibson, Jüri Peets, Jörgen Gustafsson, Katrine Iversen, Linzi Simpson, Lisa Strand, Louise Loe, Maeve Sikora, Marek Florek, Maria Vretemark, Mark Redknap, Monika Bajka, Tamara Pushkina, Morten Søvsø, Natalia Grigoreva, Tom Christensen, Ole Kastholm, Otto Uldum, Pasquale Favia, Per Holck, Raili Allmäe, Sabine Sten, Símun Arge, Sturla Ellingvåg, Vayacheslav Moiseyev, Wiesław Bogdanowicz, Yvonne Magnusson, Ludovic Orlando, Daniel Bradley, Marie Louise Jørkov, Jette Arneborg, Niels Lynnerup, Neil Price, M. Thomas Gilbert, Morten Allentoft, Jan Bill, Søren Sindbæk, Lotte Hedeager, Kristian Kristiansen, Rasmus Nielsen, Thomas Werge, Eske Willerslev
bioRxiv 703405; doi: https://doi.org/10.1101/703405
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Population genomics of the Viking world
Ashot Margaryan, Daniel Lawson, Martin Sikora, Fernando Racimo, Simon Rasmussen, Ida Moltke, Lara Cassidy, Emil Jørsboe, Andrés Ingason, Mikkel Pedersen, Thorfinn Korneliussen, Helene Wilhelmson, Magdalena Buś, Peter de Barros Damgaard, Rui Martiniano, Gabriel Renaud, Claude Bhérer, J. Víctor Moreno-Mayar, Anna Fotakis, Marie Allen, Martyna Molak, Enrico Cappellini, Gabriele Scorrano, Alexandra Buzhilova, Allison Fox, Anders Albrechtsen, Berit Schütz, Birgitte Skar, Caroline Arcini, Ceri Falys, Charlotte Hedenstierna Jonson, Dariusz Błaszczyk, Denis Pezhemsky, Gordon Turner-Walker, Hildur Gestsdóttir, Inge Lundstrøm, Ingrid Gustin, Ingrid Mainland, Inna Potekhina, Italo Muntoni, Jade Cheng, Jesper Stenderup, Jilong Ma, Julie Gibson, Jüri Peets, Jörgen Gustafsson, Katrine Iversen, Linzi Simpson, Lisa Strand, Louise Loe, Maeve Sikora, Marek Florek, Maria Vretemark, Mark Redknap, Monika Bajka, Tamara Pushkina, Morten Søvsø, Natalia Grigoreva, Tom Christensen, Ole Kastholm, Otto Uldum, Pasquale Favia, Per Holck, Raili Allmäe, Sabine Sten, Símun Arge, Sturla Ellingvåg, Vayacheslav Moiseyev, Wiesław Bogdanowicz, Yvonne Magnusson, Ludovic Orlando, Daniel Bradley, Marie Louise Jørkov, Jette Arneborg, Niels Lynnerup, Neil Price, M. Thomas Gilbert, Morten Allentoft, Jan Bill, Søren Sindbæk, Lotte Hedeager, Kristian Kristiansen, Rasmus Nielsen, Thomas Werge, Eske Willerslev
bioRxiv 703405; doi: https://doi.org/10.1101/703405

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