title: Population genomics of Bronze Age Eurasia.
abstract: The Bronze Age of Eurasia (around 3000-1000 BC) was a period of major cultural changes. However, there is debate about whether these changes resulted from the circulation of ideas or from human migrations, potentially also facilitating the spread of languages and certain phenotypic traits. We investigated this by using new, improved methods to sequence low-coverage genomes from 101 ancient humans from across Eurasia. We show that the Bronze Age was a highly dynamic period involving large-scale population migrations and replacements, responsible for shaping major parts of present-day demographic structure in both Europe and Asia. Our findings are consistent with the hypothesized spread of Indo-European languages during the Early Bronze Age. We also demonstrate that light skin pigmentation in Europeans was already present at high frequency in the Bronze Age, but not lactose tolerance, indicating a more recent onset of positive selection on lactose tolerance than previously thought.
authors.affiliation: Centre for GeoGenetics, Natural History Museum, University of Copenhagen, Øster Voldgade 5-7, 1350 Copenhagen K, Denmark.
authors.forename: Morten E
authors.initials: ME
authors.lastname: Allentoft
authors.affiliation: Centre for GeoGenetics, Natural History Museum, University of Copenhagen, Øster Voldgade 5-7, 1350 Copenhagen K, Denmark.
authors.forename: Martin
authors.initials: M
authors.lastname: Sikora
authors.affiliation: Department of Historical Studies, University of Gothenburg, 405 30 Gothenburg, Sweden.
authors.forename: Karl-Göran
authors.initials: KG
authors.lastname: Sjögren
authors.affiliation: Center for Biological Sequence Analysis, Department of Systems Biology, Technical University of Denmark, 2800 Kgs Lyngby, Denmark.
authors.forename: Simon
authors.initials: S
authors.lastname: Rasmussen
authors.affiliation: Centre for GeoGenetics, Natural History Museum, University of Copenhagen, Øster Voldgade 5-7, 1350 Copenhagen K, Denmark.
authors.forename: Morten
authors.initials: M
authors.lastname: Rasmussen
authors.affiliation: Centre for GeoGenetics, Natural History Museum, University of Copenhagen, Øster Voldgade 5-7, 1350 Copenhagen K, Denmark.
authors.forename: Jesper
authors.initials: J
authors.lastname: Stenderup
authors.affiliation: Centre for GeoGenetics, Natural History Museum, University of Copenhagen, Øster Voldgade 5-7, 1350 Copenhagen K, Denmark.
authors.forename: Peter B
authors.initials: PB
authors.lastname: Damgaard
authors.affiliation: 1] Centre for GeoGenetics, Natural History Museum, University of Copenhagen, Øster Voldgade 5-7, 1350 Copenhagen K, Denmark [2] Faculty of Archaeology, Leiden University, 2300 Leiden, The Netherlands.
authors.forename: Hannes
authors.initials: H
authors.lastname: Schroeder
authors.affiliation: Department of Archaeology and Ancient History, Lund University, 221 00 Lund, Sweden.
authors.forename: Torbjörn
authors.initials: T
authors.lastname: Ahlström
authors.affiliation: Centre for GeoGenetics, Natural History Museum, University of Copenhagen, Øster Voldgade 5-7, 1350 Copenhagen K, Denmark.
authors.forename: Lasse
authors.initials: L
authors.lastname: Vinner
authors.affiliation: Centre for GeoGenetics, Natural History Museum, University of Copenhagen, Øster Voldgade 5-7, 1350 Copenhagen K, Denmark.
authors.forename: Anna-Sapfo
authors.initials: AS
authors.lastname: Malaspinas
authors.affiliation: Centre for GeoGenetics, Natural History Museum, University of Copenhagen, Øster Voldgade 5-7, 1350 Copenhagen K, Denmark.
authors.forename: Ashot
authors.initials: A
authors.lastname: Margaryan
authors.affiliation: Oxford Radiocarbon Accelerator Unit, University of Oxford, Oxford OX1 3QY, UK.
authors.forename: Tom
authors.initials: T
authors.lastname: Higham
authors.affiliation: Oxford Radiocarbon Accelerator Unit, University of Oxford, Oxford OX1 3QY, UK.
authors.forename: David
authors.initials: D
authors.lastname: Chivall
authors.affiliation: Unit of Forensic Anthropology, Department of Forensic Medicine, University of Copenhagen, 2100 Copenhagen, Denmark.
authors.forename: Niels
authors.initials: N
authors.lastname: Lynnerup
authors.affiliation: Unit of Forensic Anthropology, Department of Forensic Medicine, University of Copenhagen, 2100 Copenhagen, Denmark.
authors.forename: Lise
authors.initials: L
authors.lastname: Harvig
authors.affiliation: Institute of Archaeology, University of Wrocław, 50-139 Wrocław, Poland.
authors.forename: Justyna
authors.initials: J
authors.lastname: Baron
authors.affiliation: Archaeological Institute, University of Zurich, CH-8006, Zurich, Switzerland.
authors.forename: Philippe
authors.initials: P
authors.lastname: Della Casa
authors.affiliation: Department of Anatomy, Wrocław Medical University, 50-368 Wrocław, Poland.
authors.forename: Paweł
authors.initials: P
authors.lastname: Dąbrowski
authors.affiliation: Department of Anthropology, University of Toronto, Toronto ONM5S 2S2, Canada.
authors.forename: Paul R
authors.initials: PR
authors.lastname: Duffy
authors.affiliation: Department of Archeology and General History, Gorno-Altaisk State University, 649000 Gorno-Altaisk, Russia.
authors.forename: Alexander V
authors.initials: AV
authors.lastname: Ebel
authors.affiliation: Institute of History and Archaeology RAS (South Ural Department), South Ural State University, 454080 Chelyabinsk, Russia.
authors.forename: Andrey
authors.initials: A
authors.lastname: Epimakhov
authors.affiliation: Environmental Research and Material Science and Centre for Textile Research, The National Museum of Denmark, 1471 Copenhagen K, Denmark.
authors.forename: Karin
authors.initials: K
authors.lastname: Frei
authors.affiliation: Institute of Archaeology, University of Wrocław, 50-139 Wrocław, Poland.
authors.forename: Mirosław
authors.initials: M
authors.lastname: Furmanek
authors.affiliation: Institute of Archaeology, University of Wrocław, 50-139 Wrocław, Poland.
authors.forename: Tomasz
authors.initials: T
authors.lastname: Gralak
authors.affiliation: Peter the Great Museum of Anthropology and Ethnography (Kunstkamera) RAS, 199034 St Petersburg, Russia.
authors.forename: Andrey
authors.initials: A
authors.lastname: Gromov
authors.affiliation: Department of Anthropology, Polish Academy of Sciences, 50-449 Wrocław, Poland.
authors.forename: Stanisław
authors.initials: S
authors.lastname: Gronkiewicz
authors.affiliation: Biocentre of the Ludwig-Maximilian-University München, 82152 Munich, Germany.
authors.forename: Gisela
authors.initials: G
authors.lastname: Grupe
authors.affiliation: 1] Department of Biological Anthropology, Institute of Biology, Eötvös Loránd University, H-1117 Budapest, Hungary [2] Department of Anthropology, Hungarian Natural History Museum, H-1083 Budapest, Hungary.
authors.forename: Tamás
authors.initials: T
authors.lastname: Hajdu
authors.affiliation: The Archaeological Museum of Wrocław, 50-077 Wrocław, Poland.
authors.forename: Radosław
authors.initials: R
authors.lastname: Jarysz
authors.affiliation: Peter the Great Museum of Anthropology and Ethnography (Kunstkamera) RAS, 199034 St Petersburg, Russia.
authors.forename: Valeri
authors.initials: V
authors.lastname: Khartanovich
authors.affiliation: Samara State Academy of Social Science and Humanities, 443099 Samara, Russia.
authors.forename: Alexandr
authors.initials: A
authors.lastname: Khokhlov
authors.affiliation: Institute of Archaeology of the Hungarian Academy of Sciences, Research Center for the Humanities, H-1250 Budapest, Hungary.
authors.forename: Viktória
authors.initials: V
authors.lastname: Kiss
authors.affiliation: 1] Institute of Archaeology and Museology, Faculty of Arts, Masaryk University, CZ-602 00 Brno, Czech Republic [2] Department of Vegetation Ecology, Institute of Botany of the Czech Academy of Sciences, CZ-602 00 Brno, Czech Republic.
authors.forename: Jan
authors.initials: J
authors.lastname: Kolář
authors.affiliation: Department of Archaeology, University of Tartu, 51003 Tartu, Estonia.
authors.forename: Aivar
authors.initials: A
authors.lastname: Kriiska
authors.affiliation: Institute of Archaeology, University of Wrocław, 50-139 Wrocław, Poland.
authors.forename: Irena
authors.initials: I
authors.lastname: Lasak
authors.affiliation: Archaeological Superintendence of Lombardy, 20123 Milano, Italy.
authors.forename: Cristina
authors.initials: C
authors.lastname: Longhi
authors.affiliation: Biocentre of the Ludwig-Maximilian-University München, 82152 Munich, Germany.
authors.forename: George
authors.initials: G
authors.lastname: McGlynn
authors.affiliation: Department of Archaeology, University of Vilnius, LT-01513 Vilnius, Lithuania.
authors.forename: Algimantas
authors.initials: A
authors.lastname: Merkevicius
authors.affiliation: The SAXO Institute, University of Copenhagen, 2300 Copenhagen S, Denmark.
authors.forename: Inga
authors.initials: I
authors.lastname: Merkyte
authors.affiliation: Department of Evolutionary Biology, Estonian Biocentre and University of Tartu, 51010 Tartu, Estonia.
authors.forename: Mait
authors.initials: M
authors.lastname: Metspalu
authors.affiliation: Department of History, Yerevan State University, 0025 Yerevan, Armenia.
authors.forename: Ruzan
authors.initials: R
authors.lastname: Mkrtchyan
authors.affiliation: Peter the Great Museum of Anthropology and Ethnography (Kunstkamera) RAS, 199034 St Petersburg, Russia.
authors.forename: Vyacheslav
authors.initials: V
authors.lastname: Moiseyev
authors.affiliation: 1] Hungarian National Museum, H-1083 Budapest, Hungary [2] Department of Biological Anthropology, University of Szeged, H-6726 Szeged, Hungary.
authors.forename: László
authors.initials: L
authors.lastname: Paja
authors.affiliation: Department of Biological Anthropology, University of Szeged, H-6726 Szeged, Hungary.
authors.forename: György
authors.initials: G
authors.lastname: Pálfi
authors.affiliation: Department of Historical Studies, University of Gothenburg, 405 30 Gothenburg, Sweden.
authors.forename: Dalia
authors.initials: D
authors.lastname: Pokutta
authors.affiliation: Institute of Archaeology and Ethnology of the Polish Academy of Sciences, 61-612 Poznań, Poland.
authors.forename: Łukasz
authors.initials: Ł
authors.lastname: Pospieszny
authors.affiliation: Laboratory for Archaeological Chemistry, University of Wisconsin-Madison, Madison, Wisconsin 53706, USA.
authors.forename: T Douglas
authors.initials: TD
authors.lastname: Price
authors.affiliation: Department of Evolutionary Biology, Estonian Biocentre and University of Tartu, 51010 Tartu, Estonia.
authors.forename: Lehti
authors.initials: L
authors.lastname: Saag
authors.affiliation: Zoological Institute of the Russian Academy of Sciences, 199034 St Petersburg, Russia.
authors.forename: Mikhail
authors.initials: M
authors.lastname: Sablin
authors.affiliation: Department of Archaeology, State Historical Museum, 109012 Moscow, Russia.
authors.forename: Natalia
authors.initials: N
authors.lastname: Shishlina
authors.affiliation: Institute for History of Medicine and Foreign Languages of the First Faculty of Medicine, Charles University, 121 08 Prague, Czech Republic.
authors.forename: Václav
authors.initials: V
authors.lastname: Smrčka
authors.affiliation: Research Center for the History and Culture of the Turkic Peoples, Gorno-Altaisk State University, 649000 Gorno-Altaisk, Russia.
authors.forename: Vasilii I
authors.initials: VI
authors.lastname: Soenov
authors.affiliation: Institute of Archaeology of the Hungarian Academy of Sciences, Research Center for the Humanities, H-1250 Budapest, Hungary.
authors.forename: Vajk
authors.initials: V
authors.lastname: Szeverényi
authors.affiliation: Department of Pre- and Early History, Institute of Archaeological Sciences, Faculty of Humanities, Eötvös Loránd University, H-1088 Budapest, Hungary.
authors.forename: Gusztáv
authors.initials: G
authors.lastname: Tóth
authors.affiliation: Research Center for the History and Culture of the Turkic Peoples, Gorno-Altaisk State University, 649000 Gorno-Altaisk, Russia.
authors.forename: Synaru V
authors.initials: SV
authors.lastname: Trifanova
authors.affiliation: Department of Archaeology, University of Tartu, 51003 Tartu, Estonia.
authors.forename: Liivi
authors.initials: L
authors.lastname: Varul
authors.affiliation: Matrica Museum, 2440 Százhalombatta, Hungary.
authors.forename: Magdolna
authors.initials: M
authors.lastname: Vicze
authors.affiliation: Laboratory of Ethnogenomics, Institute of Molecular Biology, National Academy of Sciences, 0014 Yerevan, Armenia.
authors.forename: Levon
authors.initials: L
authors.lastname: Yepiskoposyan
authors.affiliation: Department of Archaeology, Faculty of History, Moscow State University, 119991 Moscow, Russia.
authors.forename: Vladislav
authors.initials: V
authors.lastname: Zhitenev
authors.affiliation: Centre for GeoGenetics, Natural History Museum, University of Copenhagen, Øster Voldgade 5-7, 1350 Copenhagen K, Denmark.
authors.forename: Ludovic
authors.initials: L
authors.lastname: Orlando
authors.affiliation: Center for Biological Sequence Analysis, Department of Systems Biology, Technical University of Denmark, 2800 Kgs Lyngby, Denmark.
authors.forename: Thomas
authors.initials: T
authors.lastname: Sicheritz-Pontén
authors.affiliation: 1] Center for Biological Sequence Analysis, Department of Systems Biology, Technical University of Denmark, 2800 Kgs Lyngby, Denmark [2] Novo Nordisk Foundation Center for Protein Research, University of Copenhagen, 2200 Copenhagen, Denmark.
authors.forename: Søren
authors.initials: S
authors.lastname: Brunak
authors.affiliation: Center for Theoretical Evolutionary Genetics, University of California, Berkeley, California 94720-3140, USA.
authors.forename: Rasmus
authors.initials: R
authors.lastname: Nielsen
authors.affiliation: Department of Historical Studies, University of Gothenburg, 405 30 Gothenburg, Sweden.
authors.forename: Kristian
authors.initials: K
authors.lastname: Kristiansen
authors.affiliation: Centre for GeoGenetics, Natural History Museum, University of Copenhagen, Øster Voldgade 5-7, 1350 Copenhagen K, Denmark.
authors.forename: Eske
authors.initials: E
authors.lastname: Willerslev
mesh_terms: D001106:Archaeology / Q000379:methods; D001208:Asia / Q000208:ethnology; D044466:Asian People / Q000235:genetics*; D003468:Cultural Evolution / Q000266:history*; D004247:DNA / Q000235:genetics / Q000302:isolation & purification; D005060:Europe / Q000208:ethnology; D005580:Fossils; D005787:Gene Frequency / Q000235:genetics; D005828:Genetics, Population; D015894:Genome, Human / Q000235:genetics*; D023281:Genomics; D049690:History, Ancient; D063426:Human Migration / Q000266:history; D006801:Humans; D007787:Lactose Intolerance / Q000235:genetics; D007802:Language / Q000266:history*; D020641:Polymorphism, Single Nucleotide / Q000235:genetics; D012880:Skin Pigmentation / Q000235:genetics; D044465:White People / Q000235:genetics*
chemical_list: D004247:DNA