The diverse genetic origins of a Classical period Greek army

Author:

Reitsema Laurie J.1ORCID,Mittnik Alissa23456ORCID,Kyle Britney7,Catalano Giulio8ORCID,Fabbri Pier Francesco9ORCID,Kazmi Adam C. S.1,Reinberger Katherine L.1ORCID,Sineo Luca8ORCID,Vassallo Stefano10,Bernardos Rebecca3ORCID,Broomandkhoshbacht Nasreen311ORCID,Callan Kim310,Candilio Francesca12ORCID,Cheronet Olivia13ORCID,Curtis Elizabeth311,Fernandes Daniel1314ORCID,Lari Martina15ORCID,Lawson Ann Marie311ORCID,Mah Matthew31116,Mallick Swapan31116,Mandl Kirsten13ORCID,Micco Adam31116,Modi Alessandra15,Oppenheimer Jonas311,Özdogan Kadir Toykan13,Rohland Nadin3,Stewardson Kristin311,Vai Stefania15ORCID,Vergata Chiara15ORCID,Workman J. Noah3ORCID,Zalzala Fatma311ORCID,Zaro Valentina15ORCID,Achilli Alessandro17ORCID,Anagnostopoulos Achilles18ORCID,Capelli Cristian1920,Constantinou Varnavas18ORCID,Lancioni Hovirag21ORCID,Olivieri Anna17,Papadopoulou Anastasia18ORCID,Psatha Nikoleta18,Semino Ornella17ORCID,Stamatoyannopoulos John2223,Valliannou Ioanna18,Yannaki Evangelia18,Lazaridis Iosif23ORCID,Patterson Nick216,Ringbauer Harald2456,Caramelli David15ORCID,Pinhasi Ron1524ORCID,Reich David234511ORCID

Affiliation:

1. Department of Anthropology, University of Georgia, Athens, GA 30602

2. Department of Human Evolutionary Biology, Harvard University, Cambridge, MA 02138

3. Department of Genetics, Harvard Medical School, Boston, MA 02115

4. Max Planck−Harvard Research Center for the Archaeoscience of the Ancient Mediterranean, 04103 Leipzig, Germany

5. Max Planck−Harvard Research Center for the Archaeoscience of the Ancient Mediterranean, Cambridge, MA 02138

6. Department of Archaeogenetics, Max Planck Institute for Evolutionary Anthropology, 04103 Leipzig, Germany

7. Department of Anthropology, University of Northern Colorado, Greeley, CO 80639

8. Dipartimento di Scienze e Tecnologie Biologiche Chimiche e Farmaceutiche, Università degli Studi di Palermo, Palermo, 90123 Italy

9. Department of Cultural Heritage, Università del Salento, 73100 Lecce, Italy

10. Soprintendenza per i Beni Culturali e Ambientali di Palermo, 90143 Palermo, Italy

11. Howard Hughes Medical Institute (HHMI), Harvard Medical School, Boston, MA 02115

12. Servizio di Bioarcheologia, Museo delle Civiltà, 00144 Rome, Italy

13. Department of Evolutionary Anthropology, Universität Wien, 1090 Vienna, Austria

14. CIAS, Department of Life Sciences, Universidade de Coimbra, 3000-456 Coimbra, Portugal

15. Dipartimento di Biologia, Università degli Studi di Firenze, Florence, 50122 Italy

16. Broad Institute of MIT and Harvard, Cambridge, MA 02142

17. Department of Biology and Biotechnology “L. Spallanzani,” Università di Pavia, Pavia 27100, Italy

18. Gene and Cell Therapy Center, Hematology-HCT Unit, General Hospital of Thessaloniki “George Papanikolaou”, 55710 Thessaloniki, Greece

19. Department of Zoology, University of Oxford, Oxford OX1 3SZ, United Kingdom

20. Department of Chemistry, Life Sciences and Environmental Sustainability, Università di Parma, 43124 Parma, Italy

21. Department of Chemistry, Biology and Biotechnology, Università degli Studi di Perugia, 06123 Perugia, Italy

22. Department of Medicine, University of Washington, Seattle, WA 98195

23. Department of Genome Sciences, University of Washington, Seattle, WA 98195

24. Human Evolution and Archaeological Sciences, Universität Wien, A-1030 Vienna, Austria

Abstract

Trade and colonization caused an unprecedented increase in Mediterranean human mobility in the first millennium BCE. Often seen as a dividing force, warfare is in fact another catalyst of culture contact. We provide insight into the demographic dynamics of ancient warfare by reporting genome-wide data from fifth-century soldiers who fought for the army of the Greek Sicilian colony of Himera, along with representatives of the civilian population, nearby indigenous settlements, and 96 present-day individuals from Italy and Greece. Unlike the rest of the sample, many soldiers had ancestral origins in northern Europe, the Steppe, and the Caucasus. Integrating genetic, archaeological, isotopic, and historical data, these results illustrate the significant role mercenaries played in ancient Greek armies and highlight how participation in war contributed to continental-scale human mobility in the Classical world.

Funder

Howard Hughes Medical Institute

John Templeton Foundation

Allen Foundation

Fondazione Cariplo

Dipartimenti di Eccellenza

HHS | NIH | National Institute of General Medical Sciences

National Science Foundation

Publisher

Proceedings of the National Academy of Sciences

Subject

Multidisciplinary

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