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Genetic origins of the Minoans and Mycenaeans

Abstract

The origins of the Bronze Age Minoan and Mycenaean cultures have puzzled archaeologists for more than a century. We have assembled genome-wide data from 19 ancient individuals, including Minoans from Crete, Mycenaeans from mainland Greece, and their eastern neighbours from southwestern Anatolia. Here we show that Minoans and Mycenaeans were genetically similar, having at least three-quarters of their ancestry from the first Neolithic farmers of western Anatolia and the Aegean1,2, and most of the remainder from ancient populations related to those of the Caucasus3 and Iran4,5. However, the Mycenaeans differed from Minoans in deriving additional ancestry from an ultimate source related to the hunter–gatherers of eastern Europe and Siberia6,7,8, introduced via a proximal source related to the inhabitants of either the Eurasian steppe1,6,9 or Armenia4,9. Modern Greeks resemble the Mycenaeans, but with some additional dilution of the Early Neolithic ancestry. Our results support the idea of continuity but not isolation in the history of populations of the Aegean, before and after the time of its earliest civilizations.

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Figure 1: Samples and PCA.
Figure 2: Genetic differentiation of Bronze Age populations to present-day populations.

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European Nucleotide Archive

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Acknowledgements

We thank M. McCormick for comments and critiques, F. Göhringer, I. Kucukkalipci, and G. Brandt for wet laboratory support, and S. Pääbo for providing access to the clean room facilities at the MPI-EVA, Leipzig. We thank the Hellenic Ministry of Culture, the Hellenic Archaeological Service, and the Turkish Ministry of Culture and Tourism for approval of our studies, and the personnel of the Hagios Nikolaos, Herakleion, Pireas, Olympia, Chora (Trifylia), and Isparta Museums for facilitating sample collection. All maps were plotted in R using the worldHiRes map of the ‘mapdata’ package (using data in the public domain from the CIA World Data Bank II). Research on Hagios Charalambos cave by P.J.P.McG. was supported by the Royal Society and the Institute for Aegean Prehistory (INSTAP). D.M.F. was supported by an Irish Research Council grant (GOIPG/2013/36). J.K. and A.M. were funded by Deutsche Forschungsgemeinschaft grant KR 4015/1-1 and the Max Planck Society. D.R. was supported by National Institutes of Health grant GM100233, by National Science Foundation HOMINID BCS-1032255, and is a Howard Hughes Medical Institute investigator. The study of the ancient Minoans and Mycenaeans was supported by the Lucille P. Markey Charitable Trust to G.S.

Author information

Affiliations

Authors

Contributions

G.S. conceived the study. D.R. and J.K. co-supervised the ancient DNA work, sequencing, and data analysis. I.L. performed population genetics analysis and wrote the manuscript with input from other authors. P.J.P.McG., E.K.-Y., G.K., H.M., M.M., M.Ö., N.Ö., A.Pa., M.R., S.A.R., Y.T., A.V., R.A., P.S., R.P., J.K., and G.S. assembled, studied, or described archaeological and osteological material. A.M., S.P., N.R., A.F., C.P., D.M.F., J.R.H., D.M.L., Y.M., J.A.S., K.St., R.P., G.S., D.R., P.A.N., and J.K. performed wet laboratory work. A.M., N.P., S.M., and A.Pe., performed bioinformatics analyses.

Corresponding authors

Correspondence to Iosif Lazaridis or David Reich or Johannes Krause or George Stamatoyannopoulos.

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Competing interests

The authors declare no competing financial interests.

Additional information

Reviewer Information Nature thanks R. Nielsen, C. Renfrew, B. Shapiro and the other anonymous reviewer(s) for their contribution to the peer review of this work.

Publisher's note: Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.

Extended data figures and tables

Extended Data Figure 1 ADMIXTURE analysis.

ADMIXTURE analysis (Methods) with K = 2 to K = 17 is shown. Three hundred and fifty-one ancient and 2,616 present-day individuals were used in this analysis; ancient samples and present-day Greeks are displayed. To avoid visual clutter of labels, individuals in populations with sample size ≤ 5 are shown with thicker lines.

Extended Data Figure 2 Symmetry testing of Aegean Bronze Age populations.

The statistic f4(X, Y; Test, Chimp) is shown with ±3 standard errors. Each panel is titled with the pair X, Y. Populations are ordered according to the value of the statistic. Positive values indicate that Test shares more alleles with X than Y, and negative values that it shares more with Y than X. a, ‘Northern’ and ‘eastern’ populations share more alleles with Minoans than with Neolithic Greece. b, ‘Northern’ and ‘eastern’ populations share more alleles with Mycenaeans than with Neolithic Greece. c, Minoans from Lasithi and Moni Odigitria are symmetrically related to diverse populations. d, Neolithic populations from Anatolia, Europe, Greece, and the Levant share fewer alleles with Mycenaeans than with Minoans.

Extended Data Figure 3 Symmetry testing of Anatolian Bronze Age populations.

The statistic f4(X, Y; Test, Chimp) is shown with ±3 standard errors. Each panel is titled with the pair X, Y. Populations are ordered according to the value of the statistic. Positive values indicate that Test shares more alleles with X than Y, and negative values that it shares more with Y than X. a, European, Siberian, and Caucasus hunter–gatherers share fewer alleles with Bronze Age Anatolians from Harmanören Göndürle than with a Chalcolithic Anatolian from Barcın. b, Bronze Age Anatolians differ from Neolithic ones in sharing more alleles with populations of Iran, the Caucasus, and the Steppe than with those of Europe. c, Bronze Age Anatolians differ from Minoans in sharing more alleles with populations from Neolithic Iran than Neolithic Anatolia and Europe. d, Bronze Age Anatolians differ from Mycenaeans in sharing more alleles with Neolithic and Bronze Age populations of the Levant.

Extended Data Figure 4 The f3-statistics of Mycenaeans as a target with different pairs of reference populations.

The value of the statistic f3(Ref1, Ref2; Mycenaean) with ±3 standard errors; only the population pairs (Ref1, Ref2) for which the Z-score of the statistic is less than −2 are shown. Negative values indicate that the Mycenaean population is admixed from sources related to the two reference populations.

Extended Data Figure 5 Correspondence of qpAdm estimates with PCA.

As a way of validating qpAdm models of admixture for Mycenaeans from three ancestral populations (Anatolia_N or Minoan_Lasithi), (Armenia_ChL or Armenia_MLBA), (Steppe_EMBA, Steppe_MLBA, Europe_LNBA), representing substratum, ‘eastern’, and ‘northern’ ancestry, respectively (Supplementary Information section 2), we plot the qpAdm-predicted position in the PCA space of Fig. 1 versus the actual position of the Mycenaean population.

Extended Data Figure 6 Comparison of Mycenaeans and simulated admixed populations.

We simulate admixed individuals with known ancestry from three ancestral populations (Anatolia_N or Minoan_Lasithi), (Armenia_ChL or Armenia_MLBA), (Steppe_EMBA, Steppe_MLBA, Europe_LNBA), representing substratum, ‘eastern’, and ‘northern’ ancestry, respectively (Methods and Supplementary Information section 2). The maximum |Z|-score of statistics f4(Mycenaean, Simulated; Outgroup1, Outgroup2) is plotted with circles of varying size (proportional to log(|Z|)) for each assignment of ancestry proportions. The best estimate (red) corresponds to the proportions that minimize |Z|, and they are compared against the qpAdm estimate for the same ancestral sources (blue).

Extended Data Figure 7 FST between Bronze Age and present-day West Eurasian populations.

a, The population of Early Bronze Age Armenia4 shows an affinity to present-day populations from Armenia, Anatolia, the Caucasus, and Iran, as does (b) Middle/Late Bronze Age Armenia4,9. c, The Bronze Age Levant4 has an affinity to Levantine and Arabian populations. d, Late Neolithic/Bronze Age Europeans1,6,9,43 most resemble present-day northern/central Europeans, as do (e) Early/Middle Bronze Age steppe populations1,6,9, who also resemble populations of the northeast Caucasus, while (f) Middle/Late Bronze Age steppe populations resemble central/northern Europeans1,9. Jewish populations are plotted with a square to distinguish them from non-Jewish populations from the same geographical area. The plots for the newly reported populations of Mycenaeans, Minoans, and Bronze Age Anatolians are shown in Fig. 2.

Extended Data Figure 8 Symmetry testing of Mycenaeans with Modern Greek populations.

The statistic f4(Mycenaean, Modern Greek; Test, Chimp) is shown with ±3 standard errors. Modern Greeks share fewer alleles with Levantine/Anatolian/European Neolithic populations and with Minoans than Mycenaeans, suggesting a dilution of Early Neolithic ancestry since the Bronze Age. Human Origins genotype data: a, Greeks from the Coriell repository10; b, Greeks from Thessaloniki10; c, Cypriots10. Whole-genome data: d, Cretans40. Illumina genotype data: e, Greeks from Thessaly41; f, Greeks from central Greece41; g, Greeks from the study in ref. 27.

Extended Data Table 1 Information on ancient samples reported in this study
Extended Data Table 2 Phenotypic inference of ancient individuals

Supplementary information

Supplementary Information

This file contains four sections with accompanying Supplementary Figures and Tables. Section 1: Archaeological and osteological context of ancient samples; Section 2: Admixture modeling of ancient populations; Section 3: Y-chromosome haplogroup determination; Section 4: Phenotypic inference.

Supplementary Table 1

Overview of processing steps for ancient samples

Supplementary Table 2

Screening results

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Lazaridis, I., Mittnik, A., Patterson, N. et al. Genetic origins of the Minoans and Mycenaeans. Nature 548, 214–218 (2017). https://doi.org/10.1038/nature23310

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