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The spread of steppe and Iranian-related ancestry in the islands of the western Mediterranean

An Author Correction to this article was published on 15 April 2020

This article has been updated


Steppe-pastoralist-related ancestry reached Central Europe by at least 2500 bc, whereas Iranian farmer-related ancestry was present in Aegean Europe by at least 1900 bc. However, the spread of these ancestries into the western Mediterranean, where they have contributed to many populations that live today, remains poorly understood. Here, we generated genome-wide ancient-DNA data from the Balearic Islands, Sicily and Sardinia, increasing the number of individuals with reported data from 5 to 66. The oldest individual from the Balearic Islands (~2400 bc) carried ancestry from steppe pastoralists that probably derived from west-to-east migration from Iberia, although two later Balearic individuals had less ancestry from steppe pastoralists. In Sicily, steppe pastoralist ancestry arrived by ~2200 bc, in part from Iberia; Iranian-related ancestry arrived by the mid-second millennium bc, contemporary to its previously documented spread to the Aegean; and there was large-scale population replacement after the Bronze Age. In Sardinia, nearly all ancestry derived from the island’s early farmers until the first millennium bc, with the exception of an outlier from the third millennium bc, who had primarily North African ancestry and who—along with an approximately contemporary Iberian—documents widespread Africa-to-Europe gene flow in the Chalcolithic. Major immigration into Sardinia began in the first millennium bc and, at present, no more than 56–62% of Sardinian ancestry is from its first farmers. This value is lower than previous estimates, highlighting that Sardinia, similar to every other region in Europe, has been a stage for major movement and mixtures of people.

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Fig. 1: Geographical origins and temporal distribution of newly reported data.
Fig. 2: Overview of the genetic structure.
Fig. 3: Pairwise qpWave testing to group individuals.
Fig. 4: Distal modelling of ancestry proportions using qpAdm.

Data availability

All raw data are available at the European Nucleotide Archive under accession number PRJEB35980 and at

Code availability

All custom code used in this study is provided at

Change history

  • 15 April 2020

    An amendment to this paper has been published and can be accessed via a link at the top of the paper.


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This manuscript is dedicated to the memory of S. Tusa of the Soprintendenza del Mare in Palermo, who would have been an author of this study had he not tragically died in the crash of Ethiopia Airlines flight 302 on 10 March 2019. We thank Z. Zhang for database support; the Soprintendenza BBCCAA Palermo and R. Schicchi (director of Museum of Castelbuono) for facilitating access to important skeletal materials. D.M.F. was supported by an Irish Research Council grant GOIPG/2013/36. Radiocarbon work was supported in part by the NSF Archaeometry program BCS-1460369 (to D.J.K. and B.J.C). C.L.-F. was supported by Obra Social La Caixa and by FEDER-MINECO (BFU2015-64699-P and PGC2018-095931-B-100). D.C. was supported by grant 20177PJ9XF MIUR PRIN 2017. D.Reich is an Investigator of the Howard Hughes Medical Institute, and his ancient-DNA laboratory work was supported by National Science Foundation HOMINID grant BCS-1032255, a National Institutes of Health grant GM100233, an Allen Discovery Center grant, and grant no. 61220 from the John Templeton Foundation.

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D.M.F., D.Reich and R.P. conceived the study. D.M.F., E.C., C.C., G.C., M.C., V.F., M.Lozano, E.M., M.Michel, R.M.M., D.Ramis, M.R.P., V.S., P.S., L.T., M.T.-N., C.L.-F., L.S., D.C., A.C., M.Lucci, G.G., F.C., G.S. and R.P. excavated, assembled and/or studied the osteological material. D.M.F., O.C., N.R., N.B., M.F., B.G., M.Lari, M.Micheletti, A.Modi, M.N., F.C., J.O., K.A.S., K.S., K.M., C.S., K.T.Ö. and S.V. performed laboratory work under the supervision of N.R., D.C. and R.P. J.C. provided computing resources. B.J.C. performed radiocarbon analysis under the supervision of D.J.K. D.M.F., I.O., R.B., S.M. and M.Mah performed bioinformatics and population genetics analysis with input from A.Mittnik, I.L., N.P. and D.Reich.

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Correspondence to Daniel M. Fernandes or David Caramelli or Ron Pinhasi or David Reich.

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Supplementary Figs. 1–4, Tables 1–27, notes and references, and legends for Supplementary Data 1–6.

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Supplementary Data 1–6: six supplementary tables in Excel (a single file with six tabs).

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Fernandes, D.M., Mittnik, A., Olalde, I. et al. The spread of steppe and Iranian-related ancestry in the islands of the western Mediterranean. Nat Ecol Evol 4, 334–345 (2020).

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