Ancient genomics support deep divergence between Eastern and Western Mediterranean Indo-European languages

bioRxiv (Cold Spring Harbor Laboratory)

Abstract

Summary The Indo-European languages are among the most widely spoken in the world, yet their early diversification remains contentious 1–5 . It is widely accepted that the spread of this language family across Europe from the 5th millennium BP correlates with the expansion and diversification of steppe-related genetic ancestry from the onset of the Bronze Age 6,7 . However, multiple steppe-derived populations co-existed in Europe during this period, and it remains unclear how these populations diverged and which provided the demographic channels for the ancestral forms of the Italic, Celtic, Greek, and Armenian languages 8,9 . To investigate the ancestral histories of Indo-European-speaking groups in Southern Europe, we sequenced genomes from 314 ancient individuals from the Mediterranean and surrounding regions, spanning from 5,200 BP to 2,100 BP, and co-analysed these with published genome data. We additionally conducted strontium isotope analyses on 224 of these individuals. We find a deep east-west divide of steppe ancestry in Southern Europe during the Bronze Age. Specifically, we show that the arrival of steppe ancestry in Spain, France, and Italy was mediated by Bell Beaker (BB) populations of Western Europe, likely contributing to the emergence of the Italic and Celtic languages. In contrast, Armenian and Greek populations acquired steppe ancestry directly from Yamnaya groups of Eastern Europe. These results are consistent with the linguistic Italo-Celtic 10,11 and Graeco-Armenian 1,12,13 hypotheses accounting for the origins of most Mediterranean Indo-European languages of Classical Antiquity. Our findings thus align with specific linguistic divergence models for the Indo-European language family while contradicting others. This underlines the power of ancient DNA in uncovering prehistoric diversifications of human populations and language communities.

Authors 90

1. Fulya Eylem Yediay corresponding

   University of Copenhagen · Lundbeck Foundation · University of Gothenburg

   Affiliation as printed

   Department of Historical Studies, University of Gothenburg, Gothenburg, Sweden

   Lundbeck Foundation GeoGenetics Center, Globe Institute, University of Copenhagen, Copenhagen, Denmark

2. Guus Kroonen corresponding Aachen

   Leiden University · University of Copenhagen

   Affiliation as printed

   Department of Nordic Studies and Linguistics, University of Copenhagen, Copenhagen, Denmark

   Leiden University Centre for Linguistics, Leiden University, Leiden, The Netherlands

3. Serena Sabatini

   University of Gothenburg

   Affiliation as printed

   Department of Historical Studies, University of Gothenburg, Gothenburg, Sweden

4. Karin Margarita Frei

   National Museum of Denmark

   Affiliation as printed

   Department of Environmental Archaeology and Materials Science, National Museum of Denmark, Kongens Lyngby, Denmark

5. Anja B. Frank

   Universität Hamburg · National Museum of Denmark

   Affiliation as printed

   Department of Research, Collections and Conservation, Environmental Archaeology and Materials Science, National Museum of Denmark, Kongens Lyngby, Denmark

   Institute for Geology, University of Hamburg, Hamburg, Germany

6. Thomaz Pinotti

   Universidade Federal de Minas Gerais · University of Copenhagen · Lundbeck Foundation

   Affiliation as printed

   Laboratório de Biodiversidade e Evolução Molecular (LBEM), Universidade Federal de Minas Gerais, Belo Horizonte, Brazil

   Lundbeck Foundation GeoGenetics Center, Globe Institute, University of Copenhagen, Copenhagen, Denmark

7. Andrew Wigman Aachen

   Leiden University

   Affiliation as printed

   Leiden University Centre for Linguistics, Leiden University, Leiden, The Netherlands

8. Rasmus Thorsø Aachen

   Leiden University

   Affiliation as printed

   Leiden University Centre for Linguistics, Leiden University, Leiden, The Netherlands

9. Tharsika Vimala

   University of Copenhagen · Lundbeck Foundation

   Affiliation as printed

   Lundbeck Foundation GeoGenetics Center, Globe Institute, University of Copenhagen, Copenhagen, Denmark

10. Hugh McColl

    University of Copenhagen · Lundbeck Foundation · University of Gothenburg

    Affiliation as printed

    Department of Historical Studies, University of Gothenburg, Gothenburg, Sweden

    Lundbeck Foundation GeoGenetics Center, Globe Institute, University of Copenhagen, Copenhagen, Denmark

11. Ioanna Moutafi

    American School of Classical Studies at Athens · Ludwig-Maximilians-Universität München · American School of Classical Studies at Athens

    Affiliation as printed

    Institute for Pre- and Protohistoric Archaeology and Archaeology of the Roman Provinces, Ludwig-Maximilians-Universität München, Germany

    The M.H. Wiener Laboratory for Archaeological Science, American School of Classical Studies at Athens

12. Isin Altinkaya

    University of Copenhagen · Lundbeck Foundation

    Affiliation as printed

    Lundbeck Foundation GeoGenetics Center, Globe Institute, University of Copenhagen, Copenhagen, Denmark

13. Abigail Daisy Ramsøe

    University of Copenhagen · Lundbeck Foundation

    Affiliation as printed

    Lundbeck Foundation GeoGenetics Center, Globe Institute, University of Copenhagen, Copenhagen, Denmark

14. Charleen Gaunitz

    University of Copenhagen · Lundbeck Foundation

    Affiliation as printed

    Lundbeck Foundation GeoGenetics Center, Globe Institute, University of Copenhagen, Copenhagen, Denmark

15. Gabriel Renaud

    Technical University of Denmark

    Affiliation as printed

    Department of Health Technology, Technical University of Denmark, Copenhagen, Denmark

16. Alfredo Mederos Martín

    Universidad Autónoma de Madrid

    Affiliation as printed

    Department of Biology, Autonomous University of Madrid, Spain

    Universidad Autonoma de Madrid, Spain

17. Fabrice Demeter

    University of Copenhagen · Centre National de la Recherche Scientifique · Université Paris Cité · Muséum national d'Histoire naturelle · Lundbeck Foundation · Éco-Anthropologie · Musée de l'Homme

    Affiliation as printed

    Eco-anthropologie (EA), Dpt ABBA, Muséum national d'Histoire naturelle, CNRS, Université Paris Cité, Musée de l'Homme 17 place du Trocadéro, 75016 Paris, France

    Lundbeck Foundation GeoGenetics Center, Globe Institute, University of Copenhagen, Copenhagen, Denmark

18. Gabriele Scorrano

    University of Rome Tor Vergata · University of Copenhagen · Lundbeck Foundation

    Affiliation as printed

    Center for Molecular Anthropology for the study of ancient DNA, Department of Biology, University of Rome Tor Vergata, Rome Italy

    Lundbeck Foundation GeoGenetics Center, Globe Institute, University of Copenhagen, Copenhagen, Denmark

19. Alessandro Canci

    University of Udine

    Affiliation as printed

    University of Udine, Italy

20. İzzet Duyar

    Istanbul University

    Affiliation as printed

    Istanbul University

21. Claude Serhal

    British Museum · University College London

    Affiliation as printed

    British Museum, London (UK) and University College London (UK)

22. Alexander Varzari

    Phthisiopneumology Institute "Chiril Draganiuc" · National Museum of History of Moldova

    Affiliation as printed

    Laboratory of Human Genetics, Chiril Draganiuc Institute of Phthisiopneumology, Chișinău, Republic of Moldova

    The National Museum of History of Moldova, Chișinău, Republic of Moldova

23. Murat Türkteki

    Bilecik Şeyh Edebali Üniversitesi

    Affiliation as printed

    Bilecik Şeyh Edebali University, Turkey

24. John M. O’Shea

    University of Michigan

    Affiliation as printed

    Museum of Anthropological Archaeology, University of Michigan, USA

25. Lorenz Rahmstorf

    University of Göttingen

    Affiliation as printed

    University of Göttingen, Germany

26. Gürcan Polat

    Ege University

    Affiliation as printed

    Ege University, Turkey

27. Derya Atamtürk

    Istanbul University

    Affiliation as printed

    Istanbul University, Turkey

28. Lasse Vinner

    University of Copenhagen · Lundbeck Foundation

    Affiliation as printed

    Lundbeck Foundation GeoGenetics Center, Globe Institute, University of Copenhagen, Copenhagen, Denmark

29. Sachihiro Omura
    Affiliation as printed

    Japanese Institute of Anatolian Archaeology, Turkey

30. Kimiyoshi Matsumura
    Affiliation as printed

    Japanese Institute of Anatolian Archaeology, Turkey

31. Jialu Cao

    University of Copenhagen · Lundbeck Foundation

    Affiliation as printed

    Lundbeck Foundation GeoGenetics Center, Globe Institute, University of Copenhagen, Copenhagen, Denmark

32. Frederik Valeur Seersholm

    University of Copenhagen · Lundbeck Foundation

    Affiliation as printed

    Lundbeck Foundation GeoGenetics Center, Globe Institute, University of Copenhagen, Copenhagen, Denmark

33. José Miguel Morillo León

    Christian-Albrechts-Universität zu Kiel

    Affiliation as printed

    Universidad Christian Albrecht de Kiel, Germany

34. Sofia Voutsaki

    University of Groningen

    Affiliation as printed

    Groningen Institute of Archaeology, University of Groningen, The Netherlands

35. Raphaël Orgeolet

    Centre National de la Recherche Scientifique · Aix-Marseille Université · French School at Athens · Centre Camille Jullian

    Affiliation as printed

    Aix Marseille Univ, CNRS, CCJ, Aix-en-Provence, France

    École française d'Athènes

36. Brendan Burke

    University of Victoria

    Affiliation as printed

    University of Victoria, Canada

37. Nicholas P. Herrmann

    Texas State University

    Affiliation as printed

    Texas State University, USA

38. Giulia Recchia

    Sapienza University of Rome

    Affiliation as printed

    Rome University La Sapienza, Italy

39. Susi Corazza

    University of Udine

    Affiliation as printed

    University of Udine, Italy

40. Elisabetta Borgna

    University of Udine

    Affiliation as printed

    University of Udine, Italy

41. Mirella Cipolloni Sampò

    National Research Council

    Affiliation as printed

    National Research Council, Italy

42. Flavia Trucco
    Affiliation as printed

    Italian Heritage board, Italy

43. Ana Pajuelo Pando

    Universidad de Sevilla

    Affiliation as printed

    Grupo de Investigación TELLUS. Departamento de Prehistoria y Arqueología. Universidad de Sevilla. España

44. Marie Louise Schjellerup Jørkov

    University of Copenhagen

    Affiliation as printed

    Laboratory of Biological Anthropology, ArchaeoScience, Globe Institute, University of Copenhagen, Denmark

45. Patrice Courtaud

    Université de Bordeaux

    Affiliation as printed

    University of Bordeaux, France

46. Rebecca Peake

    Université de Bourgogne · Institut National de Recherches Archéologiques Préventives · Archéologie, Terre, Histoire, Sociétés

    Affiliation as printed

    Institut national de recherches archéologiques préventives (Inrap), France

    UMR 6298, Université de Bourgogne, France

47. Juan Francisco Gibaja

    Consejo Superior de Investigaciones Científicas

    Affiliation as printed

    Spanish National Research Council, Spain

48. Györgyi M. Parditka

    University of Michigan

    Affiliation as printed

    Museum of Anthropological Archaeology, University of Michigan, USA

49. Jesper T. Stenderup

    University of Copenhagen · Lundbeck Foundation

    Affiliation as printed

    Lundbeck Foundation GeoGenetics Center, Globe Institute, University of Copenhagen, Copenhagen, Denmark

50. Karl-Göran Sjögren

    University of Gothenburg

    Affiliation as printed

    Department of Historical Studies, University of Gothenburg, Gothenburg, Sweden

51. Jacqueline Staring

    University of Copenhagen · Lundbeck Foundation

    Affiliation as printed

    Lundbeck Foundation GeoGenetics Center, Globe Institute, University of Copenhagen, Copenhagen, Denmark

52. Line Olsen

    University of Copenhagen · Lundbeck Foundation

    Affiliation as printed

    Lundbeck Foundation GeoGenetics Center, Globe Institute, University of Copenhagen, Copenhagen, Denmark

53. Igor V. Deyneko

    Phthisiopneumology Institute "Chiril Draganiuc"

    Affiliation as printed

    Laboratory of Human Genetics, Chiril Draganiuc Institute of Phthisiopneumology, Chișinău, Republic of Moldova

54. György Pálfi

    University of Szeged

    Affiliation as printed

    Department of Biological Anthropology, University of Szeged, Szeged, Hungary

55. Pedro Manuel López Aldana

    Universidad de Sevilla

    Affiliation as printed

    Grupo de Investigación TELLUS. Departamento de Prehistoria y Arqueología. Universidad de Sevilla. España

56. Bryan Burns

    Wellesley College

    Affiliation as printed

    Classical Studies, Wellesley College, Boston, USA

57. László Paja

    University of Szeged

    Affiliation as printed

    Department of Biological Anthropology, University of Szeged, Szeged, Hungary

58. Christian Mühlenbock

    Lund University

    Affiliation as printed

    Lund University, Sweden

59. Claudio Cavazzuti

    University of Bologna

    Affiliation as printed

    University of Bologna, Italy

60. Alberto Cazzella

    Sapienza University of Rome

    Affiliation as printed

    Rome University La Sapienza, Italy

61. Αnna Lagia

    Ghent University

    Affiliation as printed

    Ghent University, Department of Archaeology, The Netherlands

62. Vassilis Lambrinoudakis

    National and Kapodistrian University of Athens

    Affiliation as printed

    National and Kapodistrian University of Athens

63. Lazaros Kolonas

    Hellenic Ministry of Culture and Sports

    Affiliation as printed

    Ministry of Culture (Greece)

64. Jörg Rambach

    The Archaeological Society at Athens · Deutsches Archäologisches Institut, Abteilung Athen

    Affiliation as printed

    German Archaeological Institute, Athens, Greece

    Greek Archaeological Society

65. Eugen Sava

    National Museum of History of Moldova

    Affiliation as printed

    The National Museum of History of Moldova, Chișinău, Republic of Moldova

66. Sergey M. Agulnikov

    National Agency for Archaeology

    Affiliation as printed

    National Agency for Archaeology, Chișinău, Republic of Moldova

67. Vicente Castañeda Fernández

    Universidad de Cádiz

    Affiliation as printed

    Universidad de Cádiz, Spain

68. Mia Broné

    National Museums of World Culture

    Affiliation as printed

    Världskulturmuseerna, Sweden

69. Victoria Peña Romo

    Universidad Complutense de Madrid

    Affiliation as printed

    Complutense University, Madrid, Spain

70. Fernando Ricardo Molina González

    Universidad de Granada

    Affiliation as printed

    Department of Prehistory and Archaeology, University of Granada, Spain

71. Juan Antonio Cámara Serrano

    Universidad de Granada

    Affiliation as printed

    Department of Prehistory and Archaeology, University of Granada, Spain

72. Sylvia Alejandra Jiménez Brobeil

    Universidad de Granada

    Affiliation as printed

    Department of Legal Medicine, Toxicology and Anthropology University of Granada, Spain

73. Trinidad Nájera Molino

    Universidad de Granada

    Affiliation as printed

    Department of Prehistory and Archaeology, University of Granada, Spain

74. Maria Oliva Rodríguez‐Ariza

    Universidad de Jaén

    Affiliation as printed

    Branch of Prehistory, Department of Historical Heritage, University of Jaen, Spain

75. Catalina Galán y Saulnier

    Universidad Autónoma de Madrid

    Affiliation as printed

    Department of Prehistory and Archaeology, Autonomous University of Madrid, Spain

76. Armando González Martín

    Universidad Autónoma de Madrid

    Affiliation as printed

    Department of Biology, Autonomous University of Madrid, Spain

    Universidad Autonoma de Madrid, Spain

77. Nicolas Cauwe

    Royal Museums of Art and History

    Affiliation as printed

    Royal Museums of Art and History Brussels, Belgium

78. Claude Mordant

    Université de Bourgogne · Archéologie, Terre, Histoire, Sociétés

    Affiliation as printed

    UMR 6298, Université de Bourgogne, France

79. Mafalda Roscio

    Université de Lille · Histoire, Archéologie et Littérature des Mondes Anciens

    Affiliation as printed

    UMR 8164, Université de Lille, France

80. Luc Staniaszek

    Université de Bourgogne · Institut National de Recherches Archéologiques Préventives · Archéologie, Terre, Histoire, Sociétés

    Affiliation as printed

    Institut national de recherches archéologiques préventives (Inrap), France

    UMR 6298, Université de Bourgogne, France

81. Mary Anne Tafuri

    Sapienza University of Rome

    Affiliation as printed

    Rome University La Sapienza, Italy

82. Tayfun Yıldırım

    Ankara University

    Affiliation as printed

    Department of Archaeology, Ankara University, Ankara, Turkey

83. Luciano Salzani
    Affiliation as printed

    Archaeological Superintendence of Verona, Italy

84. Thorfinn Sand Korneliussen

    University of Copenhagen · Lundbeck Foundation

    Affiliation as printed

    Lundbeck Foundation GeoGenetics Center, Globe Institute, University of Copenhagen, Copenhagen, Denmark

85. J. Víctor Moreno-Mayar

    University of Copenhagen · Lundbeck Foundation

    Affiliation as printed

    Lundbeck Foundation GeoGenetics Center, Globe Institute, University of Copenhagen, Copenhagen, Denmark

86. Morten E. Allentoft

    University of Copenhagen · Curtin University · Lundbeck Foundation

    Affiliation as printed

    Lundbeck Foundation GeoGenetics Center, Globe Institute, University of Copenhagen, Copenhagen, Denmark

    Trace and Environmental DNA (TrEnD) Laboratory, School of Molecular and Life Sciences, Curtin University, Perth, Australia

87. Martin Sikora

    University of Copenhagen · Lundbeck Foundation

    Affiliation as printed

    Lundbeck Foundation GeoGenetics Center, Globe Institute, University of Copenhagen, Copenhagen, Denmark

88. Rasmus Nielsen

    University of Copenhagen · Lundbeck Foundation · University of California, Berkeley

    Affiliation as printed

    Departments of Integrative Biology and Statistics, UC Berkeley, USA

    Lundbeck Foundation GeoGenetics Center, Globe Institute, University of Copenhagen, Copenhagen, Denmark

89. Kristian Kristiansen corresponding

    University of Copenhagen · Lundbeck Foundation · University of Gothenburg

    Affiliation as printed

    Department of Historical Studies, University of Gothenburg, Gothenburg, Sweden

    Lundbeck Foundation GeoGenetics Center, Globe Institute, University of Copenhagen, Copenhagen, Denmark

90. Eske Willerslev corresponding

    University of Copenhagen · University of Cambridge · Lundbeck Foundation

    Affiliation as printed

    GeoGenetics Group, Department of Zoology, University of Cambridge, Cambridge, UK

    Lundbeck Foundation GeoGenetics Center, Globe Institute, University of Copenhagen, Copenhagen, Denmark