The Beaker phenomenon and the genomic transformation of northwest Europe (original) (raw)

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• 21 March 2018

Please see accompanying Erratum (https://doi.org/10.1038/nature26164). The surname of author Alessandra Modi was incorrectly listed as ‘Mod’. The original Article has been corrected online.

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Acknowledgements

We thank D. Anthony, J. Koch, I. Mathieson and C. Renfrew for comments; A. Cooper for support from the Australian Centre for Ancient DNA; the Bristol Radiocarbon Accelerator Mass Spectrometry Facility (BRAMS); A. C. Sousa, A. M. Cólliga, L. Loe, C. Roth, E. Carmona Ballesteros, M. Kunst, S.-A. Coupar, M. Giesen, T. Lord, M. Green, A. Chamberlain and G. Drinkall for assistance with samples; E. Willerslev for supporting several co-authors at the Centre for GeoGenetics; the Museo Arqueológico Regional de la Comunidad de Madrid, the Hunterian Museum, University of Glasgow, the Orkney Museum, the Museu Municipal de Torres Vedras, the Great North Museum: Hancock, the Society of Antiquaries of Newcastle upon Tyne, the Sunderland Museum, the National Museum of Wales, the Duckworth Laboratory, the Wiltshire Museum, the Wells Museum, the Brighton Museum, the Somerset Heritage Museum and the Museum of London for facilitating sample collection. Support for this project was provided by Czech Academy of Sciences grant RVO:67985912; by the Momentum Mobility Research Group of the Hungarian Academy of Sciences; by the Wellcome Trust (100713/Z/12/Z); by Irish Research Council grant GOIPG/2013/36 to D.F.; by the Heidelberg Academy of Sciences (WIN project ‘Times of Upheaval’) to P.W.S., J.K. and A.Mi.; by the Swedish Foundation for Humanities and Social Sciences grant M16-0455:1 to K.Kr.; by the National Science Centre, Poland grant DEC-2013/10/E/HS3/00141 to M.Fu.; by Obra Social La Caixa and by a Spanish MINECO grant BFU2015-64699-P to C.L.-F.; by a Spanish MINECO grant HAR2016-77600-P to C.L., P.R. and C.Bl.; by the NSF Archaeometry program BCS-1460369 to D.J.K.; by the NFS Archaeology program BCS-1725067 to D.J.K. and T.Ha.; and by an Allen Discovery Center grant from the Paul Allen Foundation, US National Science Foundation HOMINID grant BCS-1032255, US National Institutes of Health grant GM100233, and the Howard Hughes Medical Institute to D.R.

Author information

Author notes

1. Ian Armit, Kristian Kristiansen, Ron Pinhasi, Wolfgang Haak, Ian Barnes, Carles Lalueza-Fox and David Reich: These authors jointly supervised this work.

Authors and Affiliations

1. Department of Genetics, Harvard Medical School, Boston, 02115, Massachusetts, USA
   Iñigo Olalde, Nadin Rohland, Swapan Mallick, Mark Lipson, Iosif Lazaridis, Nasreen Broomandkhoshbacht, Matthew Ferry, Eadaoin Harney, Megan Michel, Jonas Oppenheimer, Kristin Stewardson & David Reich
2. Department of Earth Sciences, Natural History Museum, London, SW7 5BD, UK
   Selina Brace, Thomas Booth & Ian Barnes
3. Centre for GeoGenetics, Natural History Museum, University of Copenhagen, Copenhagen, 1350, Denmark
   Morten E. Allentoft
4. School of Archaeological and Forensic Sciences, University of Bradford, Bradford, BD7 1DP, UK
   Ian Armit, Lindsey Büster & Laura Castells Navarro
5. University of Gothenburg, Gothenburg, 405 30, Sweden
   Kristian Kristiansen & Karl-Göran Sjögren
6. Broad Institute of MIT and Harvard, Cambridge, 02142, Massachusetts, USA
   Swapan Mallick, Iosif Lazaridis, Nick Patterson & David Reich
7. Howard Hughes Medical Institute, Harvard Medical School, Boston, 02115, Massachusetts, USA
   Swapan Mallick, Nasreen Broomandkhoshbacht, Matthew Ferry, Megan Michel, Jonas Oppenheimer, Kristin Stewardson & David Reich
8. Laboratory of Archaeogenetics, Institute of Archaeology, Research Centre for the Humanities, Hungarian Academy of Sciences, Budapest, 1097, Hungary
   Anna Szécsényi-Nagy & Balázs Gusztáv Mende
9. Institute for Archaeological Sciences, Archaeo- and Palaeogenetics, University of Tübingen, Tübingen, 72070, Germany
   Alissa Mittnik & Saskia Pfrengle
10. Department of Archaeogenetics, Max Planck Institute for the Science of Human History, Jena, 07745, Germany
    Alissa Mittnik
11. Department of Human Genetics, Leiden University Medical Center, Leiden, 2333 ZC, The Netherlands
    Eveline Altena & Peter de Knijff
12. Department of Anthropology, The Pennsylvania State University, University Park, Pennsylvania, 16802, USA
    Thomas K. Harper
13. Research Department of Genetics, Evolution and Environment, University College London, London, WC1E 6BT, UK
    Yoan Diekmann, Zuzana Faltyskova & Mark G. Thomas
14. Earth Institute, University College Dublin, Dublin 4, Ireland
    Daniel Fernandes, Olivia Cheronet & Ron Pinhasi
15. Department of Anthropology, University of Vienna, Vienna, 1090, Austria
    Daniel Fernandes, Olivia Cheronet & Ron Pinhasi
16. Department of Life Science, Research Center for Anthropology and Health, University of Coimbra, Coimbra, 3000-456, Portugal
    Daniel Fernandes
17. Wessex Archaeology, Salisbury, SP4 6EB, UK
    Alistair Barclay, Kirsten Egging Dinwiddy & Jacqueline I. McKinley
18. Center of Natural and Cultural History of Man, Danube Private University, Krems, 3500, Austria
    Kurt Werner Alt
19. Department of Biomedical Engineering, Basel University, Basel, 4123, Switzerland
    Kurt Werner Alt
20. Integrative Prehistory and Archaeological Science, Basel University, Basel, Switzerland
    Kurt Werner Alt
21. Departamento de Prehistoria y Arqueología, Universidad Autónoma de Madrid, Madrid, 28049, Spain
    Corina Liesau, Patricia Ríos, Concepción Blasco & Rafael Garrido Pena
22. ARGEA S.L., Madrid, 28011, Spain
    Jorge Vega Miguel & Roberto Menduiña García
23. Área de Prehistoria, Universidad de Murcia, Murcia, 30001, Spain
    Azucena Avilés Fernández, María Haber-Uriarte & Joaquín Lomba Maurandi
24. Institute of Archaeology, Research Centre for the Humanities, Hungarian Academy of Sciences, Budapest, 1097, Hungary
    Eszter Bánffy, Gabriella Kulcsár, Viktória Kiss & Kitti Köhler
25. Romano-Germanic Commission, German Archaeological Institute, Frankfurt am Main, 60325, Germany
    Eszter Bánffy
26. Museo Archeologico Nazionale di Parma, Parma, 43100, Italy
    Maria Bernabò-Brea
27. INRAP, Institut National de Recherches Archéologiques Préventives, Buffard, 25440, France
    David Billoin
28. School of History, Classics and Archaeology, University of Edinburgh, Edinburgh, EH8 9AG, UK
    Clive Bonsall & Kathleen McSweeney
29. 10 Merchiston Gardens, Edinburgh, EH10 5DD, UK
    Laura Bonsall
30. Oxford Archaeology, Oxford, OX2 0ES, UK
    Tim Allen, Gill Hey & Fraser Brown
31. Department of Archaeology and Anthropology, University of Bristol, Bristol, BS8 1UU, UK
    Sophie Carver & Volker Heyd
32. Department of Archaeology, BioArCh, University of York, York, YO10 5DD, UK
    Oliver E. Craig
33. Scottish Universities Environmental Research Centre, East Kilbride, G75 0QF, UK
    Gordon T. Cook
34. Institute of Archaeology, University of Oxford, Oxford, OX1 2PG, UK
    Barry Cunliffe
35. University of Burgundy, Dijon, 21000, France
    Anthony Denaire
36. Oxford Archaeology East, Cambridge, CB23 8SQ, UK
    Natasha Dodwell & Richard Mortimer
37. Institute of Archaeology, Czech Academy of Sciences, Prague, 118 01, Czech Republic
    Michal Ernée & Miroslav Dobeš
38. Department of Archaeology, Cambridge Archaeological Unit, University of Cambridge, Cambridge, CB3 0DT, UK
    Christopher Evans, Benjamin Neil & Marcus Brittain
39. Labrys o.p.s., Prague, 198 00, Czech Republic
    Milan Kuchařík
40. Museu i Poblat Ibèric de Ca n'Oliver, Cerdanyola del Vallès, 08290, Spain
    Joan Francès Farré
41. School of History, Classics & Archaeology, Newcastle University, Newcastle upon Tyne, NE1 7RU, UK
    Chris Fowler
42. INRAP, Institut National de Recherches Archéologiques Préventives, Nice, 06300, France
    Michiel Gazenbeek
43. Institute of Zoology and Biomedical Research, Jagiellonian University, Kraków, 31-007, Poland
    Elżbieta Haduch
44. Great Orme Mines, Great Orme, Llandudno, LL30 2XG, UK
    Nick Jowett
45. Bristol Radiocarbon Accelerator Mass Spectrometry Facility, University of Bristol, Bristol, BS8 1UU, UK
    Timothy Knowles
46. Institut für Vor- und Frühgeschichtliche Archäologie und Provinzialrömische Archäologie, Ludwig-Maximilians-Universität München, Munich, 80539, Germany
    Ken Massy & Philipp W. Stockhammer
47. INRAP, Institut National de Recherches Archéologiques Préventives, Strasbourg, 67100, France
    Philippe Lefranc
48. Université Paul-Valéry - Montpellier 3, UMR 5140 ASM, Montpellier, 34199, France
    Olivier Lemercier
49. INRAP, Institut National de Recherches Archéologiques Préventives, Metz, 57063, France
    Arnaud Lefebvre
50. UMR 5199, Pacea, équipe A3P, Université de Bordeaux, Talence, 33400, France
    Arnaud Lefebvre
51. TRÉBEDE, Patrimonio y Cultura SL, Torres de la Alameda, 28813, Spain
    César Heras Martínez & Ana Bastida Ramírez
52. Departamento de Ciencias de la Vida, Universidad de Alcalá, Alcalá de Henares, 28801, Spain
    César Heras Martínez & Virginia Galera Olmo
53. Instituto Universitario de Investigación en Ciencias Policiales (IUICP), Alcalá de Henares, 28801, Spain
    César Heras Martínez & Virginia Galera Olmo
54. Archaeom, Departament de Prehistòria, Universitat Autònoma de Barcelona, Cerdanyola del Vallès, 08193, Spain
    Tona Majó
55. Department of Biology, University of Florence, Florence, 50121, Italy
    Alessandra Modi & David Caramelli
56. Salisbury Ltd, Budaörs, 2040, Hungary
    András Czene
57. Ferenczy Museum Center, Szentendre, 2100, Hungary
    Róbert Patay
58. Budapest History Museum, Budapest, 1014, Hungary
    Anna Endrődi
59. Department of Biological Anthropology, Eötvös Loránd University, Budapest, 1117, Hungary
    Tamás Hajdu & Tamás Szeniczey
60. Hungarian Natural History Museum, Budapest, 1083, Hungary
    Tamás Hajdu & Zsolt Bernert
61. Déri Museum, Debrecen, 4026, Hungary
    János Dani
62. Historic Environment Scotland, Edinburgh, EH9 1SH, UK
    Maya Hoole
63. Humanities Institute, University College Dublin, Dublin 4, Ireland
    Denise Keating
64. Department of Anthropology, National Museum, Prague, 115 79, Czech Republic
    Petr Velemínský
65. Soprintendenza Archeologia belle arti e paesaggio per la città metropolitana di Cagliari e per le province di Oristano e Sud Sardegna, Cagliari, 9124, Italy
    Francesca Candilio
66. Physical Anthropology Section, University of Philadelphia Museum of Archaeology and Anthropology, Philadelphia, 19104, Pennsylvania, USA
    Francesca Candilio
67. Department of Environmental Biology, Sapienza University of Rome, Rome, 00185, Italy
    Francesca Candilio
68. 46 Cuidad Real Street, Parla, 28982, Spain
    Raúl Flores Fernández
69. Departamento de Prehistoria, Universidad Complutense de Madrid, Madrid, 28040, Spain
    Ana-Mercedes Herrero-Corral
70. Soprintendenza del Mare, Palermo, 90133, Italy
    Sebastiano Tusa
71. Facoltà di Lettere e Filosofia, Università di Palermo, Palermo, 90133, Italy
    Emiliano Carnieri
72. Soprintendenza per i beni culturali e ambientali di Trapani, Trapani, 91100, Italy
    Luigi Lentini
73. Prima Archeologia del Mediterraneo, Partanna, 91028, Italy
    Antonella Valenti
74. Università degli Studi di Palermo, Agrigento, 92100, Italy
    Alessandro Zanini
75. Archaeological Research Services Ltd, Bakewell, DE45 1HB, UK
    Clive Waddington
76. Departamento de Prehistoria, Facultad de Filosofía y Letras, Universidad de Valladolid, Valladolid, 47011, Spain
    Germán Delibes & Elisa Guerra-Doce
77. Albion Archaeology, Bedford, MK42 0AS, UK
    Mike Luke
78. Department F.-A. Forel for Environmental and Aquatic Sciences, Laboratory of Prehistoric Archaeology and Anthropology, University of Geneva, Geneva 4, Switzerland
    Jocelyne Desideri & Marie Besse
79. General Department of Cultural Heritage Rhineland Palatinate, Department of Archaeology, Mainz, 55116, Germany
    Günter Brücken
80. Institute of Archaeology, University of Wroclaw, Wrocław, 50-137, Poland
    Mirosław Furmanek, Agata Hałuszko & Maksym Mackiewicz
81. Institute of Archaeology, Silesian University in Opava, Opava, 746 01, Czech Republic
    Artur Rapiński
82. Department of Archaeology, University of Exeter, Exeter, EX4 4QE, UK
    Stephany Leach
83. Departament de Prehistòria, Universitat Autònoma de Barcelona, Cerdanyola del Vallès, 08193, Spain
    Ignacio Soriano & Roberto Risch
84. Department of Anthropology, University of Iowa, Iowa City, 52240, Iowa, USA
    Katina T. Lillios
85. Centro de Arqueologia, Universidade de Lisboa, Lisboa, 1600-214, Portugal
    João Luís Cardoso & João Zilhão
86. Universidade Aberta, Lisboa, 1269-001, Portugal
    João Luís Cardoso
87. Institute of Archaeology, University College London, London, WC1H 0PY, UK
    Michael Parker Pearson
88. Institute of Archaeology and Ethnology, Polish Academy of Sciences, Kraków, 31-016, Poland
    Piotr Włodarczak
89. Laboratory for Archaeological Chemistry, University of Wisconsin-Madison, Madison, 53706, Wisconsin, USA
    T. Douglas Price
90. University of Santiago de Compostela, Santiago de Compostela, 15782, Spain
    Pilar Prieto
91. UMR 5204 Laboratoire Edytem, Université Savoie Mont Blanc, Chambéry, 73376, France
    Pierre-Jérôme Rey
92. Department of Prehistory and Archaeology, Faculty of Philosophy and Letters, Valladolid University, Valladolid, 47011, Spain
    Manuel A. Rojo Guerra
93. UMR 7268 ADES, CNRS, Aix-Marseille Univ, EFS, Faculté de médecine Nord, Marseille, 13015, France
    Aurore Schmitt
94. Service archéologique, Conseil Général de la Haute-Savoie, Annecy, 74000, France
    Joël Serralongue
95. Department of Life Science, Laboratory of Prehistory, Research Center for Anthropology and Health, University of Coimbra, Coimbra, 3000-456, Portugal
    Ana Maria Silva
96. Institute for History of Medicine and Foreign Languages, First Faculty of Medicine, Charles University, Prague, 121 08, Czech Republic
    Václav Smrčka
97. ANTEA Bureau d'étude en Archéologie, Habsheim, 68440, France
    Luc Vergnaud
98. Institució Catalana de Recerca i Estudis Avançats, Barcelona, 08010, Spain
    João Zilhão
99. Departament d'Història i Arqueologia, Universitat de Barcelona, Barcelona, 08001, Spain
    João Zilhão
100. Oxford Radiocarbon Accelerator Unit, RLAHA, University of Oxford, Oxford, OX1 3QY, UK
     Thomas Higham
101. Department of Anthropology & Institute for Energy and the Environment, The Pennsylvania State University, University Park, Pennsylvania, 16802, USA
     Douglas J. Kennett
102. Faculty of Archaeology, Leiden University, Leiden, 2333 CC, The Netherlands
     Harry Fokkens
103. Department of Philosophy, History, Culture and Art Studies, Section of Archaeology, University of Helsinki, Helsinki, 00014, Finland
     Volker Heyd
104. National Museums Scotland, Edinburgh, EH1 1JF, UK
     Alison Sheridan
105. Max Planck Institute for the Science of Human History, Jena, 07745, Germany
     Philipp W. Stockhammer, Johannes Krause & Wolfgang Haak
106. Australian Centre for Ancient DNA, School of Biological Sciences, University of Adelaide, Adelaide, 5005, South Australia, Australia
     Wolfgang Haak
107. Institute of Evolutionary Biology, CSIC-Universitat Pompeu Fabra, Barcelona, 08003, Spain
     Carles Lalueza-Fox

Authors

1. Iñigo Olalde
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2. Selina Brace
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3. Morten E. Allentoft
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4. Ian Armit
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5. Kristian Kristiansen
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6. Thomas Booth
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Contributions

S.B., M.E.A., N.R., A.S.-N., A.Mi., N.B., M.Fe., E.Har., M.Mi., J.O., K.S., O.C., D.K., F.C., R.Pi., J.K., W.H., I.B. and D.R. performed or supervised laboratory work. G.T.C. and D.J.K. undertook the radiocarbon dating of a large fraction of samples. I.A., K.Kr., A.B., K.W.A., A.A.F., E.B., M.B.-B., D.B., C.Bl., J.V.M., R.M.G., C.Bo., L.Bo., T.A., L.Bü., S.C., L.C.N., O.E.C., G.T.C., B.C., A.D., K.E.D., N.D., M.E., C.E., M.K., J.F.F., H.F., C.F., M.G., R.G.P., M.H.-U., E.Had., G.H., N.J., T.K., K.Ma., S.P., P.L., O.L., A.L., C.H.M., V.G.O., A.B.R., J.L.M., T.M., J.I.M., K.Mc., B.G.M., A.Mo., G.K., V.K., A.C., R.Pa., A.E., K.Kö., T.Ha., T.S., J.Da., Z.B., M.H., P.V., M.D., F.B., R.F.F., A.-M.H.-C., S.T., E.C., L.L., A.V., A.Z., C.W., G.D., E.G.-D., B.N., M.Br., M.Lu., R.M., J.De., M.Be., G.B., M.Fu., A.H., M.Ma., A.R., S.L., I.S., K.T.L., J.L.C., C.L., M.P.P., P.W., T.D.P., P.P., P.-J.R., P.R., R.R., M.A.R.G., A.Sc., J.S., A.M.S., V.S., L.V., J.Z., D.C., T.Hi., V.H., A.Sh., K.-G.S., P.W.S., R.Pi., J.K., W.H., I.B., C.L.-F. and D.R. assembled archaeological material. I.O., S.M., T.B., A.Mi., E.A., M.Li., I.L., N.P., Y.D., Z.F., D.F., D.J.K., P.d.K., T.K.H., M.G.T. and D.R. analysed data. I.O., C.L.-F. and D.R. wrote the manuscript with input from all co-authors.

Corresponding authors

Correspondence toIñigo Olalde or David Reich.

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Additional information

Reviewer Information Nature thanks C. Renfrew, E. Rhodes, M. Richards and the other anonymous reviewer(s) for their contribution to the peer review of this work.

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Extended data figures and tables

Extended Data Figure 1 Beaker-complex artefacts.

a, ‘All-Over-Cord’ Beaker from Bathgate, West Lothian, Scotland. Photograph: © National Museums Scotland. b, Beaker-complex grave goods from La Sima III barrow, Soria, Spain61. The set includes Beaker pots of the so-called ‘Maritime style’. Photograph: Junta de Castilla y León, Archivo Museo Numantino, Alejandro Plaza.

Extended Data Figure 2 Ancient individuals with previously published genome-wide data used in this study.

a, Sampling locations. b, Time ranges. WHG, western hunter-gatherers; EHG, eastern hunter-gatherers; SHG, Scandinavian hunter-gatherers; CHG, Caucasus hunter-gatherers; E, Early; M, Middle; L, Late; N, Neolithic; CA, Copper Age; and BA, Bronze Age. Map data from the R package ‘maps’.

Extended Data Figure 3 Population structure.

a, Principal component analysis of 990 present-day west Eurasian individuals (grey dots), with previously published (pale yellow) and new ancient samples projected onto the first two principal components. b, ADMIXTURE clustering analysis with K = 8 showing ancient individuals. WHG, western hunter-gatherers; EHG, eastern hunter-gatherers; SHG, Scandinavian hunter-gatherers; CHG, Caucasus hunter-gatherers; E, Early; M, Middle; L, Late; N, Neolithic; CA, Copper Age; and BA, Bronze Age.

Extended Data Figure 4 Hunter-gatherer affinities in Neolithic and Copper Age Europe.

Differential affinity to hunter-gatherer individuals (La Braña156 from Spain and KO162 from Hungary) in European populations before the emergence of the Beaker complex. See Supplementary Information section 8 for mixture proportions and standard errors computed with qpAdm2. E, Early; M, Middle; L, Late; N, Neolithic; CA, Copper Age; BA, Bronze Age; N_Iberia, northern Iberia; and C_Iberia, central Iberia.

Extended Data Figure 5 Modelling the relationships between Neolithic populations.

a, Admixture graph fitting a test population as a mixture of sources related to both Iberia_EN and Hungary_EN. b, Likelihood distribution for models with different proportions of the source related to Iberia_EN (green admixture edge in a) when the test population is England_N, Scotland_N or France_MLN. E, Early; M, Middle; L, Late; and N, Neolithic.

Extended Data Figure 6 Genetic affinity between Beaker-complex-associated individuals from southern England and the Netherlands.

a, _f_-statistics of the form _f_4(Mbuti, test; BK_Netherlands_Tui, BK_England_SOU). Negative values indicate that test population is closer to BK_Netherlands_Tui than to BK_England_SOU; positive values indicate that the test population is closer to BK_England_SOU than to BK_Netherlands_Tui. Error bars represent ± 3 standard errors. b, Outgroup _f_3-statistics of the form _f_3(Mbuti; BK_England_SOU, test) measuring shared genetic drift between BK_England_SOU and other Beaker-complex-associated groups. Error bars represent ± 1 standard errors. Number of individuals for each group is given in parentheses. BK_Netherlands_Tui, Beaker-complex-associated individuals from De Tuithoorn (Oostwoud, the Netherlands); BK_England_SOU, Beaker-complex-associated individuals from southern England. See Supplementary Table 1 for individuals associated with each population label.

Extended Data Figure 7 Derived allele frequencies at three SNPs of functional importance.

Error bars represent 1.9-log-likelihood support interval. The red dashed lines show allele frequencies in the 1000 Genomes Project (http://www.internationalgenome.org/) ‘GBR’ population (present-day people from Great Britain). Sample sizes are 50, 98 and 117 for Britain Neolithic, Britain Copper Age and Bronze Age, and central European Beaker-complex-associated individuals, respectively. BC, Beaker complex; CA, Copper Age; and BA, Bronze Age.

Extended Data Table 1 Sites from outside Britain with new genome-wide data reported in this study

Full size table

Extended Data Table 2 Sites from Britain with new genome-wide data reported in this study

Full size table

Extended Data Table 3 111 newly reported radiocarbon dates

Full size table

Supplementary information

Life Sciences Reporting Summary (PDF 72 kb)

Supplementary Information

This file contains Supplementary Text and Data, Supplementary Figures 1-5, Supplementary Tables S1-S12 and Supplementary References. (PDF 6895 kb)

Supplementary Data

This file contains Supplementary Tables 1-5. Supplementary Table 1 shows the ancient individuals included in this study. Supplementary Table 2 contains mitochondrial haplogroup calls for individuals with newly reported data. Supplementary Table 3 contains mitochondrial haplogroup frequencies for relevant ancient populations. Supplementary Table 4 contains Y-chromosome calls for males with newly reported data and Supplementary Table 5 contains the radiocarbon database. (XLSX 282 kb)

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Olalde, I., Brace, S., Allentoft, M. et al. The Beaker phenomenon and the genomic transformation of northwest Europe.Nature 555, 190–196 (2018). https://doi.org/10.1038/nature25738

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• Received: 08 May 2017
• Accepted: 04 January 2018
• Published: 21 February 2018
• Issue Date: 08 March 2018
• DOI: https://doi.org/10.1038/nature25738