were performed at the ¹⁴CHRONO Centre laboratory at Queen's Uni-
versity, Belfast, following published sample pretreatment and labo-
ratory protocols¹¹². Additional samples were analysed by the Oxford
Radiocarbon Accelerator Unit (ORAU) laboratory (n = 24) and by the
Keck-CCAMS Group (n = 6) (see previous reports¹¹³,¹¹⁴ for laboratory
procedures). Only datings with a C/N ratio of 2.9–3.6 were accepted;
both δ¹³C and δ¹⁵N collagen measurements were also performed, and
were used in estimates of marine and freshwater reservoir effects (MRE
and FRE, respectively) (see Supplementary Note 4 and Supplementary
Data 4). Published values of MRE and FRE were used where available, but
for some regions, such as sites in western Russia, a standard FRE value
of 500 years was applied. A diet-weighted reservoir offset was then
applied to the ¹⁴C central value before calibration. Calibrations were
made in Oxcal 4.4 using the Intcal20 calibration curve¹¹⁵. For display
and calculation purposes a midpoint of the reservoir-corrected and
calibrated 95% interval was calculated. Full details of the reservoir cor-
rection and calibration procedure are given in Supplementary Note 4
and the calculations are in Supplementary Table 4.¹

**Reporting summary**

Further information on research design is available in the Nature
Portfolio Reporting Summary linked to this article.

Data availability

All adapter-trimmed sequence data (fastq) for the samples sequenced in this study are publicly available on the European Nucleotide Archive under accession PRJEB64656, together with sequence alignment map files, aligned using human build GRCh37. The full analysis dataset including both imputed and pseudo-haploid genotypes for all ancient individuals used in this study is available at https://doi.org/10.17894/ucph.d71a6a5a-8107-4fd9-9440-bdafdf61455. Aggregated IBD-sharing data as well as high-resolution versions of supplementary figures are available at Zenodo (https://doi.org/10.5281/zenodo.8196989). Previously published ancient genomic data used in this study are detailed in Supplementary Data 7, and are all already publicly available. Bioarchaeological data (including accelerator mass spectrometry results) are included in the online supplementary materials of this submission. Map figures were created using Natural Earth Data (in Figs. 1–3 and 6 and Extended Data Figs. 1, 3, 4 and 8–11.).

Code availability

All analyses relied on available software, which has been fully referenced
in the manuscript and is detailed in the relevant supplementary notes.
A collection of R functions for IBD-based mixture model inference is
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**Acknowledgements** We acknowledge P. Bennike, who was involved in initiating this project, for her substantial contributions to its conception and to prehistoric research more broadly; she passed away in 2017. We thank L. Olsen and P. Selmer Olsen for administrative and technical assistance, respectively; the UK Biobank for access to the UK Biobank genomic resource; Illumina for collaboration; and S. Ellingvåg for assistance with sample access. E.W. thanks St John's College, Cambridge, for providing a stimulating environment of discussion and learning. The Lundbeck Foundation GeoGenetics Centre is supported by grants from the Lundbeck Foundation (R302-2018-2155 and R155-2013-16338), the Novo Nordisk Foundation (NNF18SA0035006), the Wellcome Trust (214300), the Carlsberg Foundation (CF18-0024), the Danish National Research Foundation (DNRF94, DNRF174), the University of Copenhagen (KU2016 programme) and Ferring Pharmaceuticals A/S to E.W. This research has been conducted using the UK Biobank Resource and the iPSYCH Initiative, funded by the Lundbeck Foundation (R102-A9118 and R155-2014-1724). This work was further supported by the Swedish Foundation for Humanities and Social Sciences grant (Riksbankens Jubileumsfond M16-0455:1) to K.K. M.E.A. was supported by Marie Skłodowska-Curie Actions of the EU (grant no. 300554), The Villum Foundation (grant no. 10120) and Independent Research Fund Denmark (grant no. 7027-00147B). W.B. is supported by the Hanne and Torkel Weis-Fogh Fund (Department of Zoology, University of Cambridge). A.P. is funded by the Wellcome grant WT214300; B.S.d.M and O.D. by the Swiss National Science Foundation (SFNS P00P3_176977) and the European Research Council (ERC 679330); R. Macleod by an SSHRC doctoral studentship grant (G101449: 'Individual Life Histories in Long-Term Cultural Change'); G.R. by a Novo Nordisk Foundation Fellowship (gNNF200C0062491); N.N.J. by Aarhus University Research Foundation; B.S.P. by an ERC-Starter Grant 'NEOSEA' (grant no. 949424); H.S. by a Carlsberg Foundation
Fellowship (CF19-0601); G.S. by Marie Skłodowska-Curie Individual Fellowship 'PALAEO-ENO' (grant agreement number 751349); A. J. Schork by a Lundbeckfonden Fellowship (R335-2019-2318) and the National Institute on Aging (NIH award numbers U19AG023122, U24AG051129 and UH2AG064706); A.L. and I.S. by the Science Committee, Ministry of Education and Science of the Republic of Kazakhstan (AP08856317); B.G.-R. and M.G.-M. by the Spanish Ministry of Science and Innovation (project HAR2016-75605-R); C.M.-L. and O.R. by the Italian Ministry for the Universities (grants 2010-11 prot.2010EL8TXP_001, 'Biological and cultural heritage of the central-southern Italian population through 30 thousand years' and 2008 prot. 2008B4J2HS_001, 'Origin and diffusion of farming in central-southern Italy: a molecular approach'); and D.C.-S. and I.G.-Z. by the Spanish Ministry of Science and Innovation (project HAR2017-86262-P). D.C.S.-G. acknowledges funding from the Generalitat Valenciana (CIDEGENT/2019/061) and the Spanish Government (EUR2020-112213). D.B. was supported by the NOMIS Foundation and Marie Skłodowska-Curie Global Fellowship 'CUSP' (grant no. 846856); E.R.U. by the Science Committee, Ministry of Education and Science of the Republic of Kazakhstan (AP09261083: 'Transcultural Communications in the Late Bronze Age (Western Siberia-Kazakhstan-Central Asia)'); E.C. by Villum Fonden (17649); J.E.A.T. by the Spanish Ministry of Economy and Competitiveness (HAR2013-46861-R) and Generalitat Valenciana (Aico/ 2018/125 and Aico 2020/97); and P.K. by the Russian Ministry of Science and Higher Education (Ural Federal University Program of Development within the Priority-2030 Program). P.K. also acknowledges the Museum of the Institute of Plant and Animal Ecology (UB RAS, Ekaterinburg). L.Y. acknowledges funding by the Science Committee of the Armenian Ministry of Education and Science (project 21AG-1F025); L.O. by the ERC Consolidator Grant 'PEGASUS' (agreement no. 681605); M. Sablin by the Russian Ministry of Science and Higher Education (075-15-2021-1069); N.C. by Historic Environment Scotland; S.V. and E.V.V. by the Russian Ministry of Science and Higher Education (075-15-2022-328); and V.M. by the Science Committee, Ministry of Education and Science of the Republic of Kazakhstan (AR08856925). V.A. is supported by a Lundbeckfonden Fellowship (R335-2019-2318); P.H.S. by the National Institute of General Medical Sciences (R35GM142916); S.R. by the Novo Nordisk Foundation (NNF14CC0001); T.S.K. is funded by Carlsberg grant CF19-0712; R.D. by the Wellcome Trust (WT214300); R.N. by the National Institute of General Medical Sciences (NIH grant R01GM138634); and F. Racimo by a Villum Fonden Young Investigator Grant (no. 00025300); by a Novo Nordisk Fonden Data Science Ascending Investigator Award (NNF22OC0076816) and by the European Research Council (ERC) under the European Union's Horizon Europe programme (grant agreement No. 101077592). T.W. and V.A. are supported by the Lundbeck Foundation iPSYCH initiative (R248-2017-2003).

**Author contributions** E.W. initiated the study. M.E.A., M.S., T.S.K., R.D., R.N., O.D., T.W., F. Racimo, K.K. and E.W. led the study. M.E.A., M.S., A.F., M.M., C.L.-F., R.N., E.C., T.W., K.K. and E.W. conceptualized the study. M.E.A., M.S., H.S., L.O., T.S.K., R.D., R.N., O.D., T.W., F. Racimo, K.K. and E.W. supervised the research. M.E.A., L.O., R.D., R.N., T.W., K.H.K., K.K. and E.W. acquired

funding for research. M.E.A., A.F., J.S., K.-G.S., M.L.S.J., M.U.H., A.A.T., A.C., A.Z., A.M.S., A.J.H., A.G., A.L., B.H.N., B.G.-R., C.B., C.L., C.M.-L., D.V.P., D.C.-S., D.O.L., D.E., D.C.S.-G., D.B., E.B.P., E.K., E.V.V., E.R.U., E. Kannegaard, F. Radina, H.D., I.G.-Z., I.P., I.S., J.G., J.H., J.E.A.T., J.Z., J.V., K.B.P., K.T., L.N., L.L., L.N.M., L.Y., L.P., L. Sarti, L. Slimak, L.K., M.G.-M., M. Silvestrini, M.D., M.V., M.S.N., M.R., M.S., M.P., M.C., M. Sablin, N.C., N.S., O.P., O.R., O.V.L., P.A., P.K., P.C., P. Ríos, P. Lotz, P. Lysdahl, P.M., P.P., P.B., P.d.B.D., P.V.P., P.P.M., P.W., R.V.S., R. Maring, R. Menduiña, R.B., R.T., S.V., S.W., S.B., S.S., S.A.S., S.H.A., T.D.P., T.Z.T.J., Y.B.S., V.I.M., V.S., V.M., Y.M., I.M., O.G. and N.L. were involved in sample collection. M.E.A., M.S., A.R.-M., E.K.I.-P., W.B., A.I., J.S., A.P., B.S.d.M., M.I., M.M., L.V., A. Stern, C.G., F.E.Y, D.J.L., T.S.K., R.D., R.N., O.D., F. Racimo, K.H.K. and E.W. were involved in developing and applying methodology. M.E.A., J.S., C.G. and L.V. led the DNA laboratory work research component. K.-G.S., A.F. and M.E.A. led bioarchaeological data curation. M.E.A., M.S., A.R.-M., E.K.I.-P., W.B., A.I., A.P., B.S.d.M., B.S.P., A.H., R.A.H., T.V., H.M., A.M., A.V., A.B.N., P. Rasmussen, G.R., A. D. Ramsøe, A. Skorobogatov, A. J. Schork, A. Rosengren, C.J.M., I.A., L.Z., R. Maring, V.S., V.A., P.H.S., S.R., T.S.K., O.D. and F. Racimo undertook formal analyses of data. M.E.A., M.S., A.R.-M., E.K.I.-P., A.F., W.B., A.I., K.-G.S., R. Macleod, D.J.L., P.H.S., T.S.K., F. Racimo and E.W. drafted the main text (M.E.A. and M.S. led this). M.E.A., M.S., A.R.-M., E.K.I.-P., A.F., W.B., A.I., K.-G.S., A.P., B.S.d.M., B.S.P., A.H., R. Macleod, R.A.H., T.V., M.F.M., A.B.N., M.U.H., P. Rasmussen, A. J. Stern, N.N.J., H.S., G.S., A. Ramsøe, A. Skorobogatov, A. Rosengren, A.O., A.B., A.C., A.G., A.L., A.B.G., C.J.M., D.C.S.-G., E.B.P., E. Kostyleva, E.R.U., E. Kannegaard, I.G.-Z., I.P., I.S., J.G., J.H., J.E.A.T., K.H.K., L.Z, L.Y., L.P., L.K., M.B., M.G.-M., M.V., M.R., M.J., N.B., O.V.L., O.C.U., P.K., P. Lysdahl, P.B., P.W., R.V.S., R. Maring, R.B., R.I., S.V., S.W., S.B., S.H.A., T.Z.T.J., V.S., D.J.L., P.H.S., S.R., T.S.K., O.D. and F. Racimo drafted supplementary notes and materials. M.E.A., M.S., A.R.-M., E.K.I.-P., A.F., W.B., A.I., G.S., A.H., M.L.S.J., F.D., R. Macleod, L. Sørensen, P.O.N., R.A.H., T.V., H.M., A.M., N.N.J., H.S., A. Ramsøe, A. Skorobogatov, A. J. Schork, A. Ruter, A.O., B.H.N., B.G.-R., D.C.-S., D.C.S.-G., I.G.-Z., I.P., J.G., J.E.A.T., L.Z., L.O., L.K., M.G.-M., P.d.B.D., R.I., S.A.S., D.J.L., I.M., O.G., P.H.S., T.S.K., R.D., R.N., O.D., T.W., F. Racimo, K.K. and E.W. were involved in reviewing drafts and editing (M.E.A., M.S., A.F., K.-G.S., R. Macleod and E.W. led this, and subsequent finalization of the study). All co-authors read, commented on and agreed on the submitted manuscript.

**Competing interests** The authors declare no competing interests.

**Additional information**

**Supplementary information** The online version contains supplementary material available at https://doi.org/10.1038/s41586-023-06865-0.

**Correspondence and requests for materials** should be addressed to Morten E. Allentoft, Martin Sikora or Eske Willerslev.

**Peer review information** Nature thanks Benjamin Peter and the other, anonymous, reviewer(s) for their contribution to the peer review of this work.

**Reprints and permissions information** is available at http://www.nature.com/reprints.
Fellowship (CF19-0601); G.S. by Marie Skłodowska-Curie Individual Fellowship 'PALAEO-ENO' (grant agreement number 751349); A. J. Schork by a Lundbeckfonden Fellowship (R335-2019-2318) and the National Institute on Aging (NIH award numbers U19AG023122, U24AG051129 and UH2AG064706); A.L. and I.S. by the Science Committee, Ministry of Education and Science of the Republic of Kazakhstan (AP08856317); B.G.-R. and M.G.-M. by the Spanish Ministry of Science and Innovation (project HAR2016-75605-R); C.M.-L. and O.R. by the Italian Ministry for the Universities (grants 2010-11 prot.2010EL8TXP_001, 'Biological and cultural heritage of the central-southern Italian population through 30 thousand years' and 2008 prot. 2008B4J2HS_001, 'Origin and diffusion of farming in central-southern Italy: a molecular approach'); and D.C.-S. and I.G.-Z. by the Spanish Ministry of Science and Innovation (project HAR2017-86262-P). D.C.S.-G. acknowledges funding from the Generalitat Valenciana (CIDEGENT/2019/061) and the Spanish Government (EUR2020-112213). D.B. was supported by the NOMIS Foundation and Marie Skłodowska-Curie Global Fellowship 'CUSP' (grant no. 846856); E.R.U. by the Science Committee, Ministry of Education and Science of the Republic of Kazakhstan (AP09261083: 'Transcultural Communications in the Late Bronze Age (Western Siberia-Kazakhstan-Central Asia)'); E.C. by Villum Fonden (17649); J.E.A.T. by the Spanish Ministry of Economy and Competitiveness (HAR2013-46861-R) and Generalitat Valenciana (Aico/ 2018/125 and Aico 2020/97); and P.K. by the Russian Ministry of Science and Higher Education (Ural Federal University Program of Development within the Priority-2030 Program). P.K. also acknowledges the Museum of the Institute of Plant and Animal Ecology (UB RAS, Ekaterinburg). L.Y. acknowledges funding by the Science Committee of the Armenian Ministry of Education and Science (project 21AG-1F025); L.O. by the ERC Consolidator Grant 'PEGASUS' (agreement no. 681605); M. Sablin by the Russian Ministry of Science and Higher Education (075-15-2021-1069); N.C. by Historic Environment Scotland; S.V. and E.V.V. by the Russian Ministry of Science and Higher Education (075-15-2022-328); and V.M. by the Science Committee, Ministry of Education and Science of the Republic of Kazakhstan (AR08856925). V.A. is supported by a Lundbeckfonden Fellowship (R335-2019-2318); P.H.S. by the National Institute of General Medical Sciences (R35GM142916); S.R. by the Novo Nordisk Foundation (NNF14CC0001); T.S.K. is funded by Carlsberg grant CF19-0712; R.D. by the Wellcome Trust (WT214300); R.N. by the National Institute of General Medical Sciences (NIH grant R01GM138634); and F. Racimo by a Villum Fonden Young Investigator Grant (no. 00025300); by a Novo Nordisk Fonden Data Science Ascending Investigator Award (NNF22OC0076816) and by the European Research Council (ERC) under the European Union's Horizon Europe programme (grant agreement No. 101077592). T.W. and V.A. are supported by the Lundbeck Foundation iPSYCH initiative (R248-2017-2003).

**Author contributions** E.W. initiated the study. M.E.A., M.S., T.S.K., R.D., R.N., O.D., T.W., F. Racimo, K.K. and E.W. led the study. M.E.A., M.S., A.F., M.M., C.L.-F., R.N., E.C., T.W., K.K. and E.W. conceptualized the study. M.E.A., M.S., H.S., L.O., T.S.K., R.D., R.N., O.D., T.W., F. Racimo, K.K. and E.W. supervised the research. M.E.A., L.O., R.D., R.N., T.W., K.H.K., K.K. and E.W. acquired

funding for research. M.E.A., A.F., J.S., K.-G.S., M.L.S.J., M.U.H., A.A.T., A.C., A.Z., A.M.S., A.J.H., A.G., A.L., B.H.N., B.G.-R., C.B., C.L., C.M.-L., D.V.P., D.C.-S., D.O.L., D.E., D.C.S.-G., D.B., E.B.P., E.K., E.V.V., E.R.U., E. Kannegaard, F. Radina, H.D., I.G.-Z., I.P., I.S., J.G., J.H., J.E.A.T., J.Z., J.V., K.B.P., K.T., L.N., L.L., L.N.M., L.Y., L.P., L. Sarti, L. Slimak, L.K., M.G.-M., M. Silvestrini, M.D., M.V., M.S.N., M.R., M.S., M.P., M.C., M. Sablin, N.C., N.S., O.P., O.R., O.V.L., P.A., P.K., P.C., P. Rios, P. Lotz, P. Lysdahl, P.M., P.P., P.B., P.d.B.D., P.V.P., P.P.M., P.W., R.V.S., R. Maring, R. Menduiña, R.B., R.T., S.V., S.W., S.B., S.S., S.A.S., S.H.A., T.D.P., T.Z.T.J., Y.B.S., V.I.M., V.S., V.M., Y.M., I.M., O.G. and N.L. were involved in sample collection. M.E.A., M.S., A.R.-M., E.K.I.-P., W.B., A.I., J.S., A.P., B.S.d.M., M.I., M.M., L.V., A. Stern, C.G., F.E.Y, D.J.L., T.S.K., R.D., R.N., O.D., F. Racimo, K.H.K. and E.W. were involved in developing and applying methodology. M.E.A., J.S., C.G. and L.V. led the DNA laboratory work research component. K.-G.S., A.F. and M.E.A. led bioarchaeological data curation. M.E.A., M.S., A.R.-M., E.K.I.-P., W.B., A.I., A.P., B.S.d.M., B.S.P., A.H., R.A.H., T.V., H.M., A.M., A.V., A.B.N., P. Rasmussen, G.R., A. D. Ramsøe, A. Skorobogatov, A. J. Schork, A. Rosengren, C.J.M., I.A., L.Z., R. Maring, V.S., V.A., P.H.S., S.R., T.S.K., O.D. and F. Racimo undertook formal analyses of data. M.E.A., M.S., A.R.-M., E.K.I.-P., A.F., W.B., A.I., K.-G.S., R. Macleod, D.J.L., P.H.S., T.S.K., F. Racimo and E.W. drafted the main text (M.E.A. and M.S. led this). M.E.A., M.S., A.R.-M., E.K.I.-P., A.F., W.B., A.I., K.-G.S., A.P., B.S.d.M., B.S.P., A.H., R. Macleod, R.A.H., T.V., M.F.M., A.B.N., M.U.H., P. Rasmussen, A. J. Stern, N.N.J., H.S., G.S., A. Ramsøe, A. Skorobogatov, A. Rosengren, A.O., A.B., A.C., A.G., A.L., A.B.G., C.J.M., D.C.S.-G., E.B.P., E. Kostyleva, E.R.U., E. Kannegaard, I.G.-Z., I.P., I.S., J.G., J.H., J.E.A.T., K.H.K., L.Z, L.Y., L.P., L.K., M.B., M.G.-M., M.V., M.R., M.J., N.B., O.V.L., O.C.U., P.K., P. Lysdahl, P.B., P.W., R.V.S., R. Maring, R.B., R.I., S.V., S.W., S.B., S.H.A., T.Z.T.J., V.S., D.J.L., P.H.S., S.R., T.S.K., O.D. and F. Racimo drafted supplementary notes and materials. M.E.A., M.S., A.R.-M., E.K.I.-P., A.F., W.B., A.I., G.S., A.H., M.L.S.J., F.D., R. Macleod, L. Sørensen, P.O.N., R.A.H., T.V., H.M., A.M., N.N.J., H.S., A. Ramsøe, A. Skorobogatov, A. J. Schork, A. Ruter, A.O., B.H.N., B.G.-R., D.C.-S., D.C.S.-G., I.G.-Z., I.P., J.G., J.E.A.T., L.Z., L.O., L.K., M.G.-M., P.d.B.D., R.I., S.A.S., D.J.L., I.M., O.G., P.H.S., T.S.K., R.D., R.N., O.D., T.W., F. Racimo, K.K. and E.W. were involved in reviewing drafts and editing (M.E.A., M.S., A.F., K.-G.S., R. Macleod and E.W. led this, and subsequent finalization of the study). All co-authors read, commented on and agreed on the submitted manuscript.

**Competing interests** The authors declare no competing interests.

**Additional information**

**Supplementary information** The online version contains supplementary material available at https://doi.org/10.1038/s41586-023-06865-0.

**Correspondence and requests for materials** should be addressed to Morten E. Allentoft, Martin Sikora or Eske Willerslev.

**Peer review information** Nature thanks Benjamin Peter and the other, anonymous, reviewer(s) for their contribution to the peer review of this work.

**Reprints and permissions information** is available at http://www.nature.com/reprints.
<figure><img src="image_13.png" /></figure>

e

<figure><img src="image_14.png" /><figcaption>**Extended Data Fig. 1** | Genetic structure of the 317 herein-reported ancient genomes. **a–d**, PCA of 3,316 modern and ancient individuals from Eurasia, Oceania and the Americas (**a, b**), as well as restricted to 2,126 individuals from western Eurasia (west of the Urals) (**c, d**). Shown are analyses with principal components inferred either using both modern and imputed ancient genomes passing all filters, and projecting low coverage ancient genomes (**a, c**); or only modern genomes and projecting all ancient genomes (**b, d**). Ancient genomes sequenced in this study are indicated either with black circles</figcaption></figure>

(imputed genomes) or grey diamonds (projected genomes). **e**, Model-based clustering results using ADMIXTURE for 284 newly reported genomes (excluding close relatives and individuals flagged for possible contamination). Results shown are based on ADMIXTURE runs from K = 2 to K = 15 on 1,593 ancient individuals, corresponding to the full set of 1,492 imputed genomes passing filters as well as 101 low coverage genomes represented by pseudo-haploid genotypes (flags "lowcov" or "lowGpAvg", Supplementary Data 7; indicated with alpha transparency in plot).
<figure><img src="image_15.png" /></figure>

e

<figure><img src="image_16.png" /></figure>

**Extended Data Fig. 1** | Genetic structure of the 317 herein-reported ancient genomes. **a–d**, PCA of 3,316 modern and ancient individuals from Eurasia, Oceania and the Americas (**a, b**), as well as restricted to 2,126 individuals from western Eurasia (west of the Urals) (**c, d**). Shown are analyses with principal components inferred either using both modern and imputed ancient genomes passing all filters, and projecting low coverage ancient genomes (**a, c**); or only modern genomes and projecting all ancient genomes (**b, d**). Ancient genomes sequenced in this study are indicated either with black circles

(imputed genomes) or grey diamonds (projected genomes). **e**, Model-based clustering results using ADMIXTURE for 284 newly reported genomes (excluding close relatives and individuals flagged for possible contamination). Results shown are based on ADMIXTURE runs from K = 2 to K = 15 on 1,593 ancient individuals, corresponding to the full set of 1,492 imputed genomes passing filters as well as 101 low coverage genomes represented by pseudo-haploid genotypes (flags "lowcov" or "lowGpAvg", Supplementary Data 7; indicated with alpha transparency in plot).
<figure><img src="image_15.png" /><figcaption>a. Depth of Coverage</figcaption></figure>

<figure><img src="image_16.png" /><figcaption>b. Prehistoric Europe 1x</figcaption></figure>

<figure><img src="image_17.png" /><figcaption>c. Viking Age Europe 1x</figcaption></figure>

<figure><img src="image_18.png" /><figcaption>d. Hungary 1x</figcaption></figure>

**Extended Data Fig. 2** | Imputation accuracy of ancient DNA. a, Imputation accuracy across 42 high-coverage ancient genomes when downsampled to lower depth of coverage values (see Supplementary Note 2 and Supplementary Table 2.1). b, Imputation accuracy for 1× depth of coverage across 9 prehistoric

European genomes; c, across 5 Viking age genomes; and d, across 7 ancient genomes from Early Medieval Hungary. In all panels, imputation accuracy is shown as the squared Pearson correlation between imputed and true genotype dosages as a function of MAF of the target variant sites.
<figure><img src="image_17.png" /></figure>

<figure><img src="image_18.png" /></figure>

<figure><img src="image_19.png" /></figure>

<figure><img src="image_20.png" /></figure>

**Extended Data Fig. 2** | Imputation accuracy of ancient DNA. a, Imputation accuracy across 42 high-coverage ancient genomes when downsampled to lower depth of coverage values (see Supplementary Note 2 and Supplementary Table 2.1). b, Imputation accuracy for 1× depth of coverage across 9 prehistoric

European genomes; c, across 5 Viking age genomes; and d, across 7 ancient genomes from Early Medieval Hungary. In all panels, imputation accuracy is shown as the squared Pearson correlation between imputed and true genotype dosages as a function of MAF of the target variant sites.
a

<figure><img src="image_19.png" /></figure>

b

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c

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f

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d

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e

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g

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<figure><img src="image_21.png" /></figure>

<figure><img src="image_22.png" /></figure>

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<figure><img src="image_25.png" /></figure>

<figure><img src="image_26.png" /></figure>

<figure><img src="image_27.png" /></figure>
**Extended Data Fig. 3 | Genetic clustering of ancient individuals.**

Characterization of genetic clusters for 1,401 imputed ancient individuals from Eurasia (that is, excluding 91 individuals from Africa and Americas), inferred from pairwise IBD sharing (indicated using coloured symbols throughout), a, Temporal distribution of clustered individuals, grouped by broad ancestry cluster. b,c, Geographical distribution of clustered individuals, shown for individuals predating 3,000 BP (b) and after 3,000 BP (c). d, Network graph of pairwise IBD sharing between 596 ancient Eurasians predating 3,000 BP, highlighting within- and between-cluster relationships. Each node represents

an individual, and the width of edges connecting nodes indicates the fraction
of the genome shared IBD between the respective pair of individuals. Network
edges were restricted to the 10 highest sharing connections for each individual,
and the layout was computed using the force-directed Fruchterman-Reingold
algorithm. e, Neighbour-joining tree showing relationships between genetic
clusters, inferred using total variation distance (TVD) of IBD painting palettes.
f,g, PCA of 3,119 Eurasian (f) or 2,126 west Eurasian (g) ancient and modern
individuals ("HO" dataset).
**Extended Data Fig. 3 | Genetic clustering of ancient individuals.**

Characterization of genetic clusters for 1,401 imputed ancient individuals from Eurasia (that is, excluding 91 individuals from Africa and Americas), inferred from pairwise IBD sharing (indicated using coloured symbols throughout), a, Temporal distribution of clustered individuals, grouped by broad ancestry cluster. b,c, Geographical distribution of clustered individuals, shown for individuals predating 3,000 BP (b) and after 3,000 BP (c). d, Network graph of pairwise IBD sharing between 596 ancient Eurasians predating 3,000 BP, highlighting within- and between-cluster relationships. Each node represents

an individual, and the width of edges connecting nodes indicates the fraction
of the genome shared IBD between the respective pair of individuals. Network
edges were restricted to the 10 highest sharing connections for each individual,
and the layout was computed using the force-directed Fruchterman-Reingold
algorithm. e, Neighbour-joining tree showing relationships between genetic
clusters, inferred using total variation distance (TVD) of IBD painting palettes.
f,g, PCA of 3,119 Eurasian (f) or 2,126 west Eurasian (g) ancient and modern
individuals ("HO" dataset).
a

<figure><img src="image_26.png" /></figure>

<figure><img src="image_27.png" /></figure>

<figure><img src="image_28.png" /></figure>

b

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c

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d

<figure><img src="image_34.png" /></figure>

<figure><img src="image_35.png" /></figure>
a

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b

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c

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d

<figure><img src="image_36.png" /></figure>

<figure><img src="image_37.png" /></figure>
**Extended Data Fig. 4 | Genetic structure of European HGs after the LGM.**
a, Supervised ancestry modelling using non-negative least squares on IBD sharing profiles. Panels show estimated ancestry proportions for target individuals from genetic clusters representing European HGs, using different sets of increasingly proximal source groups. Individuals used as sources in a particular set are indicated with black crosses and coloured bars with 100% ancestry proportion. Black lines indicate 1 standard error for the respective ancestry component. b, Residuals for model fit of target individuals from selected genetic clusters across different source sets. c, Moon charts showing

spatial distribution of ancestry proportions in European HGs deriving from four European source groups (set "hgEur2"; source origins shown with coloured symbol). Estimated ancestry proportions are indicated by both size and amount of fill of moon symbols. Note that 'Italy_15000BP_9000BP' and 'RussiaNW_11000BP_8000BP' correspond to 'WHG' and 'EHG' labels used in previous studies. d, Maps showing networks of highest between-cluster IBD sharing (top 10 highest sharing per individual) for individuals from two genetic clusters representing Scandinavian HGs. See Supplementary Data 1 and 7 for details of individual sample IDs presented here.
**Extended Data Fig. 4 | Genetic structure of European HGs after the LGM.**
a, Supervised ancestry modelling using non-negative least squares on IBD sharing profiles. Panels show estimated ancestry proportions for target individuals from genetic clusters representing European HGs, using different sets of increasingly proximal source groups. Individuals used as sources in a particular set are indicated with black crosses and coloured bars with 100% ancestry proportion. Black lines indicate 1 standard error for the respective ancestry component. b, Residuals for model fit of target individuals from selected genetic clusters across different source sets. c, Moon charts showing

spatial distribution of ancestry proportions in European HGs deriving from four European source groups (set "hgEur2"; source origins shown with coloured symbol). Estimated ancestry proportions are indicated by both size and amount of fill of moon symbols. Note that 'Italy_15000BP_9000BP' and 'RussiaNW_11000BP_8000BP' correspond to 'WHG' and 'EHG' labels used in previous studies. d, Maps showing networks of highest between-cluster IBD sharing (top 10 highest sharing per individual) for individuals from two genetic clusters representing Scandinavian HGs. See Supplementary Data 1 and 7 for details of individual sample IDs presented here.
<figure><img src="image_36.png" /></figure>

**Extended Data Fig. 5** | Ancestry modelling for HG and Neolithic farmer-associated genetic clusters. Supervised ancestry modelling using non-negative least squares on IBD sharing profiles. Panels show estimated ancestry proportions of two global Eurasian clusters, corresponding to European HGs before 4,000 BP and individuals from Europe and western Asia from around 10,000 BP until historical times, including Anatolian-associated (Neolithic) farmers, Caucasus HGs and recent individuals with genetic affinity to the Levant. Columns show results of modelling target individuals using three

panels of increasingly distal source groups: “postBA”: Bronze Age and Neolithic source groups; “postNeol”, Bronze Age and later targets using Late Neolithic/ early Bronze Age and earlier source groups; “deep”, Mesolithic and later targets using deep ancestry source groups. Individuals used as sources in a particular set are indicated with black crosses and coloured bars with 100% ancestry proportion. Black lines indicate 1 standard error for the respective ancestry component.
<figure><img src="image_38.png" /></figure>

**Extended Data Fig. 5** | Ancestry modelling for HG and Neolithic farmer-associated genetic clusters. Supervised ancestry modelling using non-negative least squares on IBD sharing profiles. Panels show estimated ancestry proportions of two global Eurasian clusters, corresponding to European HGs before 4,000 BP and individuals from Europe and western Asia from around 10,000 BP until historical times, including Anatolian-associated (Neolithic) farmers, Caucasus HGs and recent individuals with genetic affinity to the Levant. Columns show results of modelling target individuals using three

panels of increasingly distal source groups: “postBA”: Bronze Age and Neolithic source groups; “postNeol”, Bronze Age and later targets using Late Neolithic/ early Bronze Age and earlier source groups; “deep”, Mesolithic and later targets using deep ancestry source groups. Individuals used as sources in a particular set are indicated with black crosses and coloured bars with 100% ancestry proportion. Black lines indicate 1 standard error for the respective ancestry component.
<figure><img src="image_37.png" /><figcaption>**Extended Data Fig. 6** | Ancestry modelling for post-Neolithic genetic clusters. Supervised ancestry modelling using non-negative least squares on IBD sharing profiles. Panels show estimated ancestry proportions of a global Eurasian cluster corresponding to European individuals after 5,000 BP, as well as pastoralist groups from the Eurasian steppe. Columns show results of modelling target individuals using three panels of increasingly distal source</figcaption></figure>

groups: "postBA": Bronze Age and Neolithic source groups; "postNeol", Bronze Age and later targets using Late Neolithic/early Bronze Age and earlier source groups; "deep", Mesolithic and later targets using deep ancestry source groups. Individuals used as sources in a particular set are indicated with black crosses and coloured bars with 100% ancestry proportion. Black lines indicate 1 standard error for the respective ancestry component.
<figure><img src="image_39.png" /><figcaption>**Extended Data Fig. 6** | Ancestry modelling for post-Neolithic genetic clusters. Supervised ancestry modelling using non-negative least squares on IBD sharing profiles. Panels show estimated ancestry proportions of a global Eurasian cluster corresponding to European individuals after 5,000 BP, as well as pastoralist groups from the Eurasian steppe. Columns show results of modelling target individuals using three panels of increasingly distal source</figcaption></figure>

groups: "postBA": Bronze Age and Neolithic source groups; "postNeol", Bronze Age and later targets using Late Neolithic/early Bronze Age and earlier source groups; "deep", Mesolithic and later targets using deep ancestry source groups. Individuals used as sources in a particular set are indicated with black crosses and coloured bars with 100% ancestry proportion. Black lines indicate 1 standard error for the respective ancestry component.
<figure><img src="image_38.png" /><figcaption>**Extended Data Fig. 7 | Ancestry modelling for genetic clusters east of the Urals.** Supervised ancestry modelling using non-negative least squares on IBDaring profiles. Panels show estimated ancestry proportions of a global Eurasian cluster corresponding to central, east and north Asian individuals with east Eurasian genetic affinities. Columns show results of modelling target individuals using three panels of increasingly distal source groups: “postBA”:</figcaption></figure>

Bronze Age and Neolithic source groups; “postNeol”, Bronze Age and later targets using Late Neolithic/early Bronze Age and earlier source groups; “deep”, Mesolithic and later targets using deep ancestry source groups. Individuals used as sources in a particular set are indicated with black crosses and coloured bars with 100% ancestry proportion. Black lines indicate 1 standard error for the respective ancestry component.
<figure><img src="image_40.png" /><figcaption>**Extended Data Fig. 7 | Ancestry modelling for genetic clusters east of the Urals.** Supervised ancestry modelling using non-negative least squares on IBDaring profiles. Panels show estimated ancestry proportions of a global Eurasian cluster corresponding to central, east and north Asian individuals with east Eurasian genetic affinities. Columns show results of modelling target individuals using three panels of increasingly distal source groups: "postBA":</figcaption></figure>

Bronze Age and Neolithic source groups; "postNeol", Bronze Age and later targets using Late Neolithic/early Bronze Age and earlier source groups; "deep", Mesolithic and later targets using deep ancestry source groups. Individuals used as sources in a particular set are indicated with black crosses and coloured bars with 100% ancestry proportion. Black lines indicate 1 standard error for the respective ancestry component.
a

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b

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c

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d

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e

<figure><img src="image_43.png" /><figcaption>**Extended Data Fig. 8** | See next page for caption.</figcaption></figure>
a

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b

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c

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d

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e

<figure><img src="image_45.png" /><figcaption>Extended Data Fig. 8 | See next page for caption.</figcaption></figure>
**Extended Data Fig. 8 | Dynamics of the Neolithic transition in Europe.**

**a**, Supervised ancestry modelling using non-negative least squares on IBD sharing profiles. Panels show estimated ancestry proportions for target individuals from genetic clusters representing European Neolithic farmer individuals ("fEur" source set). Individuals used as sources in a particular set are indicated with black crosses and coloured bars with 100% ancestry proportion. Black lines indicate 1 standard error for the respective ancestry component. **b**, Composition of HG ancestry proportions from different source groups in individuals with Neolithic farmer ancestry, shown as bar plots. Grey bars represent contributions from a source with ancestry related to local HGs. **c**, Moon charts showing spatial distribution of estimated ancestry proportions related to local HGs across Europe. Estimated ancestry proportions are

indicated by size and amount of fill of moon symbols. Coloured areas indicate the geographical extent of individuals included as local sources in the respective regions. d, Estimated time of admixture between local HG groups and Neolithic farmers. Black diamonds and error bars represent point estimate and standard errors of admixture time, coloured bars show temporal range of included target individuals. The time to admixture was adjusted backwards by the average age of individuals for each region. e, Moon charts showing spatial distribution of estimated ancestry proportions derived from genetic clusters of early Neolithic European farmers (locations indicated with coloured symbols). Estimated ancestry proportions are indicated by size and amount of fill of moon symbols. Red symbols indicate individuals where standard errors exceed the point estimates for the respective ancestry source.
**Extended Data Fig. 8 | Dynamics of the Neolithic transition in Europe.**

**a**, Supervised ancestry modelling using non-negative least squares on IBD sharing profiles. Panels show estimated ancestry proportions for target individuals from genetic clusters representing European Neolithic farmer individuals ("fEur" source set). Individuals used as sources in a particular set are indicated with black crosses and coloured bars with 100% ancestry proportion. Black lines indicate 1 standard error for the respective ancestry component. **b**, Composition of HG ancestry proportions from different source groups in individuals with Neolithic farmer ancestry, shown as bar plots. Grey bars represent contributions from a source with ancestry related to local HGs. **c**, Moon charts showing spatial distribution of estimated ancestry proportions related to local HGs across Europe. Estimated ancestry proportions are

indicated by size and amount of fill of moon symbols. Coloured areas indicate the geographical extent of individuals included as local sources in the respective regions. d, Estimated time of admixture between local HG groups and Neolithic farmers. Black diamonds and error bars represent point estimate and standard errors of admixture time, coloured bars show temporal range of included target individuals. The time to admixture was adjusted backwards by the average age of individuals for each region. e, Moon charts showing spatial distribution of estimated ancestry proportions derived from genetic clusters of early Neolithic European farmers (locations indicated with coloured symbols). Estimated ancestry proportions are indicated by size and amount of fill of moon symbols. Red symbols indicate individuals where standard errors exceed the point estimates for the respective ancestry source.
a

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b

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c

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d

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**Extended Data Fig. 9 | Dynamics of the steppe transition in Europe.**

a, Estimated time of admixture between local HG groups and Neolithic farmers. Black diamonds and error bars represent point estimate and standard errors of admixture time, coloured bars show temporal range of included target individuals. The time to admixture was adjusted backwards by the average age of individuals for each region. b, Moon charts showing spatial distribution of estimated ancestry proportions related to local Neolithic farmers across Europe. Estimated ancestry proportions are indicated by size and amount of fill

of moon symbols. Coloured areas indicate the geographical extent of individuals included as local sources in the respective regions. c, Map showing networks of highest between-cluster IBD sharing (top 10 highest sharing per individual) for individuals from genetic cluster "Steppe_5000BP_4300BP" representing the major steppe ancestry source for Europeans. d, Distributions of difference in estimated steppe-related ancestry proportions, using individuals from the genetic cluster "Steppe_5000BP_4300BP", associated with either Yamnaya or Afanasievo cultural contexts as separate sources.
a

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b

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c

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d

<figure><img src="image_49.png" /></figure>

**Extended Data Fig. 9 | Dynamics of the steppe transition in Europe.**
a, Estimated time of admixture between local HG groups and Neolithic farmers. Black diamonds and error bars represent point estimate and standard errors of admixture time, coloured bars show temporal range of included target individuals. The time to admixture was adjusted backwards by the average age of individuals for each region. b, Moon charts showing spatial distribution of estimated ancestry proportions related to local Neolithic farmers across Europe. Estimated ancestry proportions are indicated by size and amount of fill

of moon symbols. Coloured areas indicate the geographical extent of individuals included as local sources in the respective regions. c, Map showing networks of highest between-cluster IBD sharing (top 10 highest sharing per individual) for individuals from genetic cluster "Steppe_5000BP_4300BP" representing the major steppe ancestry source for Europeans. d, Distributions of difference in estimated steppe-related ancestry proportions, using individuals from the genetic cluster "Steppe_5000BP_4300BP", associated with either Yamnaya or Afanasievo cultural contexts as separate sources.
a

<figure><img src="image_48.png" /></figure>

<figure><img src="image_49.png" /></figure>

<figure><img src="image_50.png" /></figure>

c

<figure><img src="image_51.png" /></figure>

d

<figure><img src="image_52.png" /></figure>

b

<figure><img src="image_53.png" /></figure>

<figure><img src="image_54.png" /></figure>

**Extended Data Fig. 10** | Genetic transformations across the Eurasian steppe. **a**, Moon charts showing spatial distribution of estimated ancestry proportions of Siberian HGs from the “deep” Siberian ancestry sources (names and locations indicated with coloured symbols). Estimated ancestry proportions are indicated by size and amount of fill of moon symbols. **b**, Timelines of ancestry proportions from “postNeol” sources in central and north Asian ancient individuals after 5,000 BP. Symbol shape and colour indicate the genetic cluster

of each individual. Black lines indicate 1 standard error. c,d, Difference in estimated steppe-related ancestry proportions, using individuals from genetic cluster “Steppe_5000BP_4300BP” associated with either Yamnaya or Afanasievo cultural contexts as separate sources, as a function of time (c) or total estimated steppe-ancestry proportion (d). Individuals from genetic clusters of individuals associated with Okunevo (blue stars) or Sintashta/Andronovo (green diamonds) contexts are indicated with coloured symbols.
a

<figure><img src="image_50.png" /></figure>

<figure><img src="image_51.png" /></figure>

<figure><img src="image_52.png" /></figure>

c

<figure><img src="image_53.png" /></figure>

d

<figure><img src="image_54.png" /></figure>

b

<figure><img src="image_55.png" /></figure>

<figure><img src="image_56.png" /></figure>

**Extended Data Fig. 10** | Genetic transformations across the Eurasian steppe.
a, Moon charts showing spatial distribution of estimated ancestry proportions of Siberian HGs from the “deep” Siberian ancestry sources (names and locations indicated with coloured symbols). Estimated ancestry proportions are indicated by size and amount of fill of moon symbols. b, Timelines of ancestry proportions from “postNeol” sources in central and north Asian ancient individuals after 5,000 BP. Symbol shape and colour indicate the genetic cluster

of each individual. Black lines indicate 1 standard error. c,d, Difference in estimated steppe-related ancestry proportions, using individuals from genetic cluster “Steppe_5000BP_4300BP” associated with either Yamnaya or Afanasievo cultural contexts as separate sources, as a function of time (c) or total estimated steppe-ancestry proportion (d). Individuals from genetic clusters of individuals associated with Okunevo (blue stars) or Sintashta/Andronovo (green diamonds) contexts are indicated with coloured symbols.
<figure><img src="image_55.png" /><figcaption>**Extended Data Fig. 11 | Patterns of co-ancestry.** a, Panels show within-cluster genetic relatedness over time, measured as the total length of genomic segments shared IBD between individuals. Results for both measures are shown separately for individuals from western versus eastern Eurasia. Small grey dots indicate estimates for individual pairs, with larger coloured symbols indicating median values within genetic clusters. Ranges of median values for</figcaption></figure>

major ancestry groups are indicated with labelled convex hulls. b, Distribution of ROH lengths for 29 individuals with evidence for recent parental relatedness (>50 cM total in ROHs > 20 cM). c, Karyogram showing genomic distribution of ROH in individual tem003, an ancient case of uniparental disomy for chromosome 2. Regions within ROH are indicated with blue colour.
<figure><img src="image_57.png" /><figcaption>**Extended Data Fig. 11 | Patterns of co-ancestry.** a, Panels show within-cluster genetic relatedness over time, measured as the total length of genomic segments shared IBD between individuals. Results for both measures are shown separately for individuals from western versus eastern Eurasia. Small grey dots indicate estimates for individual pairs, with larger coloured symbols indicating median values within genetic clusters. Ranges of median values for</figcaption></figure>

major ancestry groups are indicated with labelled convex hulls. b, Distribution of ROH lengths for 29 individuals with evidence for recent parental relatedness (>50 cM total in ROHs > 20 cM). c, Karyogram showing genomic distribution of ROH in individual tem003, an ancient case of uniparental disomy for chromosome 2. Regions within ROH are indicated with blue colour.
# Reporting Summary

Nature Portfolio wishes to improve the reproducibility of the work that we publish. This form provides structure for consistency and transparency in reporting. For further information on Nature Portfolio policies, see our [Editorial Policies](#) and the [Editorial Policy Checklist](#).

## Statistics

For all statistical analyses, confirm that the following items are present in the figure legend, table legend, main text, or Methods section.

n/a | Confirmed

☐ ☑ The exact sample size (n) for each experimental group/condition, given as a discrete number and unit of measurement

☐ ☑ A statement on whether measurements were taken from distinct samples or whether the same sample was measured repeatedly

☐ ☑ The statistical test(s) used AND whether they are one- or two-sided
*Only common tests should be described solely by name; describe more complex techniques in the Methods section.*

☐ ☑ A description of all covariates tested

☐ ☑ A description of any assumptions or corrections, such as tests of normality and adjustment for multiple comparisons

☐ ☑ A full description of the statistical parameters including central tendency (e.g. means) or other basic estimates (e.g. regression coefficient) AND variation (e.g. standard deviation) or associated estimates of uncertainty (e.g. confidence intervals)

☐ ☑ For null hypothesis testing, the test statistic (e.g. *F*, *t*, *r*) with confidence intervals, effect sizes, degrees of freedom and *P* value noted
*Give *P* values as exact values whenever suitable.*

☐ ☑ For Bayesian analysis, information on the choice of priors and Markov chain Monte Carlo settings

☐ ☑ For hierarchical and complex designs, identification of the appropriate level for tests and full reporting of outcomes

☐ ☑ Estimates of effect sizes (e.g. Cohen's *d*, Pearson's *r*), indicating how they were calculated

*Our web collection on statistics for biologists contains articles on many of the points above.*

## Software and code

Policy information about [availability of computer code](#)

**Data collection**

Sequencing data and metadata pertaining to the sequencing of ancient genomes is managed on secure servers at the Globe Institute, University of Copenhagen, and via the Illumina Inc. BaseSpace platform. Accelerator Mass Spectrometry data and associated metadata is managed at the Department of Historical Studies, University of Gothenberg.

**Data analysis**

Custom scripts used to apply chromopainter from large-scale phased data are available at https://github.com/will-camb/Nero/tree/master/scripts/cp; panel scripts. All other analyses relied upon available software which has been fully referenced in the manuscript and detailed in the relevant supplementary notes. These comprise:

CASAVA (v.1.8.2)

AdapterRemoval (v.2.1.3)

Picard (v.1.127)

GATK (v.3.3.0)

Samtools (1.9)

Samtools calmd (v.1.10)

pysam (https://github.com/pysam-developers/pysam)

BEDtools (v.2.23.0)

mapDamage2.0 (v2.2.1)

BWA (0.7-17)

Schmutzi (VERSION?)

ContamMix (VERSION?)

ANGSD (0.938)

GLIMPSE (v1.0.1)
## Reporting Summary

Nature Portfolio wishes to improve the reproducibility of the work that we publish. This form provides structure for consistency and transparency in reporting. For further information on Nature Portfolio policies, see our [Editorial Policies](#) and the [Editorial Policy Checklist](#).

## Statistics

For all statistical analyses, confirm that the following items are present in the figure legend, table legend, main text, or Methods section.

n/a | Confirmed

☐ ☑ The exact sample size (n) for each experimental group/condition, given as a discrete number and unit of measurement

☐ ☑ A statement on whether measurements were taken from distinct samples or whether the same sample was measured repeatedly

☐ ☑ The statistical test(s) used AND whether they are one- or two-sided
*Only common tests should be described solely by name; describe more complex techniques in the Methods section.*

☐ ☑ A description of all covariates tested

☐ ☑ A description of any assumptions or corrections, such as tests of normality and adjustment for multiple comparisons

☐ ☑ A full description of the statistical parameters including central tendency (e.g. means) or other basic estimates (e.g. regression coefficient) AND variation (e.g. standard deviation) or associated estimates of uncertainty (e.g. confidence intervals)

☐ ☑ For null hypothesis testing, the test statistic (e.g. *F*, *t*, *r*) with confidence intervals, effect sizes, degrees of freedom and *P* value noted
*Give *P* values as exact values whenever suitable.*

☐ ☑ For Bayesian analysis, information on the choice of priors and Markov chain Monte Carlo settings

☐ ☑ For hierarchical and complex designs, identification of the appropriate level for tests and full reporting of outcomes

☐ ☑ Estimates of effect sizes (e.g. Cohen's *d*, Pearson's *r*), indicating how they were calculated

*Our web collection on statistics for biologists contains articles on many of the points above.*

## Software and code

Policy information about [availability of computer code](#)

**Data collection** Sequencing data and metadata pertaining to the sequencing of ancient genomes is managed on secure servers at the Globe Institute, University of Copenhagen, and via the Illumina Inc. BaseSpace platform. Accelerator Mass Spectrometry data and associated metadata is managed at the Department of Historical Studies, University of Gothenberg.

**Data analysis**

Custom scripts used to apply chromopainter from large-scale phased data are available at https://github.com/will-camb/Nero/tree/master/scripts/cp; scripts. All other analyses relied upon available software which has been fully referenced in the manuscript and detailed in the relevant supplementary notes. These comprise:
* CASAVA (v.1.8.2)
* AdapterRemoval (v.2.1.3)
* Picard (v.1.127)
* GATK (v.3.3.0)
* Samtools (1.9)
* Samtools calmd (v.1.10)
* pysam (https://github.com/pysam-developers/pysam)
* BEDtools (v.2.23.0)
* mapDamage2.0 (v2.2.1)
* BWA (0.7-17)
* Schmutzi (VERSION?)
* ContamMix (VERSION?)
* ANGSD (0.938)
* GLIMPSE (v1.0.1)
Beagle (v4.1)
BCFtools 1.10
MAFFT (v7.490)
RAxML-ng (v1.1.0)
rynettle (v0.4)
EPA-ng (0.3.8)
NgsRelate (v2)
ADMIXTURE (1.3.0)
smartpca
GCTA
R version 4.0
ADMIXTOOLS2
gstat (2.0-9)
IBDseq (version r1206)
GenomicRanges (3.15)
leidenAlg (v1.01, https://github.com/kharchenkolab/leidenAlg)
igraph (0.9.9)
limSolve (1.5.6)
Scikit-learn (0.21.2)
Chromopainter (0.0.4)
fineSTRUCTURE (0.0.5)
QCTOOL v2 (https://www.well.ox.ac.uk/~gav/qctool_v2/)
ArcGIS Online (www.arcgis.com)
OxCal v4.4
LRA v0.1.0 (https://github.com/petrkunes/LRA)

For manuscripts utilizing custom algorithms or software that are central to the research but not yet described in published literature, software must be made available to editors and reviewers. We strongly encourage code deposition in a community repository (e.g. GitHub). See the Nature Portfolio guidelines for submitting code & software for further information.

## Data

Policy information about availability of data

All manuscripts must include a data availability statement. This statement should provide the following information, where applicable:

- Accession codes, unique identifiers, or web links for publicly available datasets

- A description of any restrictions on data availability

- For clinical datasets or third party data, please ensure that the statement adheres to our policy

All collapsed and paired-end sequence data for novel samples sequenced in this study will be made publicly available on the European Nucleotide Archive, together with trimmed sequence alignment map files, aligned using human build GRCh37. Previously published ancient genomic data used in this study is detailed in Supplementary Table VII, and are all already publicly available. Bioarchaeological data (including stable isotopes and radiocarbon dating results) are included in the online supplementary materials of this submission.

## Human research participants

Policy information about studies involving human research participants and Sex and Gender in Research.

Reporting on sex and gender

No living or recently deceased human research participants were affected by this study

Population characteristics

Describe the covariate-relevant population characteristics of the human research participants (e.g. age, genotypic information, past and current diagnosis and treatment categories). If you filled out the behavioural & social sciences study design questions and have nothing to add here, write "See above."

Recruitment

Describe how participants were recruited. Outline any potential self-selection bias or other biases that may be present and how these are likely to impact results.

Ethics oversight

Identify the organization(s) that approved the study protocol.

Note that full information on the approval of the study protocol must also be provided in the manuscript.

## Field-specific reporting

Please select the one below that is the best fit for your research. If you are not sure, read the appropriate sections before making your selection.

<figure><img src="image_56.png" /></figure>

Life sciences

<figure><img src="image_57.png" /></figure>

Behavioural & social sciences

<figure><img src="image_58.png" /></figure>

Ecological, evolutionary & environmental sciences

For a reference copy of the document with all sections, see [nature.com/documents/nr-reporting-summary-flat.pdf](https://nature.com/documents/nr-reporting-summary-flat.pdf)
Beagle (v4.1)
BCFtools 1.10
MAFFT (v7.490)
RAxML-ng (v1.1.0)
rynettle (v0.4)
EPA-ng (0.3.8)
NgsRelate (v2)
ADMIXTURE (1.3.0)
smartpca
GCTA
R version 4.0
ADMIXTOOLS2
gstat (2.0-9)
IBDseq (version r1206)
GenomicRanges (3.15)
leidenAlg (v1.01, https://github.com/kharchenkolab/leidenAlg)
igraph (0.9.9)
limSolve (1.5.6)
Scikit-learn (0.21.2)
Chromopainter (0.0.4)
fineSTRUCTURE (0.0.5)
QCTOOL v2 (https://www.well.ox.ac.uk/~gav/qctool_v2/)
ArcGIS Online (www.arcgis.com)
OxCal v4.4
LRA v0.1.0 (https://github.com/petrkunes/LRA)

For manuscripts utilizing custom algorithms or software that are central to the research but not yet described in published literature, software must be made available to editors and reviewers. We strongly encourage code deposition in a community repository (e.g. GitHub). See the Nature Portfolio guidelines for submitting code & software for further information.

## Data

Policy information about availability of data

All manuscripts must include a data availability statement. This statement should provide the following information, where applicable:

- Accession codes, unique identifiers, or web links for publicly available datasets

- A description of any restrictions on data availability

- For clinical datasets or third party data, please ensure that the statement adheres to our policy

All collapsed and paired-end sequence data for novel samples sequenced in this study will be made publicly available on the European Nucleotide Archive, together with trimmed sequence alignment map files, aligned using human build GRCh37. Previously published ancient genomic data used in this study is detailed in Supplementary Table VII, and are all already publicly available. Bioarchaeological data (including stable isotopes and radiocarbon dating results) are included in the online supplementary materials of this submission.

## Human research participants

Policy information about studies involving human research participants and Sex and Gender in Research.

Reporting on sex and gender

No living or recently deceased human research participants were affected by this study

Population characteristics

Describe the covariate-relevant population characteristics of the human research participants (e.g. age, genotypic information, past and current diagnosis and treatment categories). If you filled out the behavioural & social sciences study design questions and have nothing to add here, write "See above."

Recruitment

Describe how participants were recruited. Outline any potential self-selection bias or other biases that may be present and how these are likely to impact results.

Ethics oversight

Identify the organization(s) that approved the study protocol.

Note that full information on the approval of the study protocol must also be provided in the manuscript.

## Field-specific reporting

Please select the one below that is the best fit for your research. If you are not sure, read the appropriate sections before making your selection.

<figure><img src="image_58.png" /></figure>

Life sciences

<figure><img src="image_59.png" /></figure>

Behavioural & social sciences

<figure><img src="image_60.png" /></figure>

Ecological, evolutionary & environmental sciences

For a reference copy of the document with all sections, see [nature.com/documents/nr-reporting-summary-flat.pdf](https://nature.com/documents/nr-reporting-summary-flat.pdf)
# Ecological, evolutionary & environmental sciences study design

All studies must disclose on these points even when the disclosure is negative.

**Study description**
We sequenced 317 novel ancient human genomes from primarily Mesolithic and Neolithic individuals across Eurasia, combined with new radiocarbon dates, stable isotope data. We additionally test and apply GLIMPSE to impute 1492 previously published ancient genomes, and incorporate this data in our analyses. We undertake a suite of analyses of this data to generate new insights into population structure through time, natural selection and health-related adaptation.

**Research sample**
Ancient human remains (bone and dental cementum) were collected, sampled, extracted and screened by shallow shotgun sequencing (see Supplementary Note 1). This resulted in 317 samples found to be suitable for deep sequencing (of 962 initial specimens). These represent a considerable geographic range across Eurasia (from Lake Baikal to the Atlantic coast). A high sample density for Denmark (100) was undertaken to provide high resolution insights at this location. A particular focus upon individuals related to Mesolithic or Neolithic culture was made, with 300 of the individuals from the time span 11,000 cal. BP to 3,000 cal. BP.

**Sampling strategy**
Sampling was dependent upon the availability of ancient human remains, though a considerable transect of the Mesolithic and Neolithic periods in Eurasia was represented, together with a detailed continuous sequence of human occupation of Denmark specifically. Our assemblage is well-represented with individuals of such key archaeological complexes as the Maglemose, Ertebølle and Funnel Beaker cultures in Scandinavia, the Cardial in the Mediterranean, the Körös and Linear Pottery complexes in SE and Central Europe, and many archaeological cultures in Ukraine, western Russia, and the trans-Ural (e.g. Veretye, Lyalovo, Volosovo, Kitoi).

**Data collection**
Sampled remains consisted of teeth (sampled for dental cementum, N=211), petrous bones (N=91), and other bone types (N=15; long bones, ribs and cranial bones). Sampling and ancient DNA laboratory work was undertaken in dedicated ancient DNA clean lab facilities at the GeoGenetics Centre. Additionally, 272 novel accelerator mass spectrometry results were generated at the 14CHRONO laboratory, Queen's University Belfast (242 samples), at the Oxford Radiocarbon Accelerator Unit (ORAU) laboratory (24 samples) and at the Keck-CCAMS Group, Irvine, California, USA (6 samples).

**Timing and spatial scale**
Sample chronology was generated by radiocarbon dating, with dates corrected for marine and freshwater reservoir effects. These showed samples ranging from the Upper Palaeolithic (c. 25,700 cal. BP) to the mediaeval period (c. 1200 cal. BP). Most individuals (97%, N=309) span 11,000 cal. BP to 3,000 cal. BP, the period broadly associated with the Mesolithic and Neolithic in Eurasia. Our research area can broadly be divided into three large regions: 1) central, western and northern Europe, 2) eastern Europe including western Russia and Ukraine, and 3) the Urals and western Siberia.

**Data exclusions**
Samples that were screened by shallow sequencing and found to have endogenous DNA of >1% were not sequenced further and not included in analyses. This amounted to 645 specimens.

**Reproducibility**
Data quality and uncertainty (e.g. contamination) was accounted for in computational analyses to assess robustness of inferences, and all methods and data are made available for future replication.

**Randomization**
Sample groups were defined to reflect archaeological populations, based on phylogenetic inferences, temporal and geographic provenance, and cultural interpretations evidenced by archaeological contexts.

**Blinding**
Blinding was not applicable to this study.

Did the study involve field work?

<figure><img src="image_59.png" /></figure>

Yes

<figure><img src="image_60.png" /></figure>

No

# Reporting for specific materials, systems and methods

We require information from authors about some types of materials, experimental systems and methods used in many studies. Here, indicate whether each material, system or method listed is relevant to your study. If you are not sure if a list item applies to your research, read the appropriate section before selecting a response.

## Materials & experimental systems

<table><thead><tr><th>n/a</th><th>Involved in the study</th></tr></thead><tbody><tr><td>☑</td><td>Antibodies</td></tr><tr><td>☑</td><td>Eukaryotic cell lines</td></tr><tr><td>☐</td><td>Palaeontology and archaeology</td></tr><tr><td>☑</td><td>Animals and other organisms</td></tr><tr><td>☑</td><td>Clinical data</td></tr><tr><td>☑</td><td>Dual use research of concern</td></tr></tbody></table>

## Methods

<table><thead><tr><th>n/a</th><th>Involved in the study</th></tr></thead><tbody><tr><td>☑</td><td>ChIP-seq</td></tr><tr><td>☑</td><td>Flow cytometry</td></tr><tr><td>☑</td><td>MRI-based neuroimaging</td></tr></tbody></table>
# Ecological, evolutionary & environmental sciences study design

All studies must disclose on these points even when the disclosure is negative.

**Study description**
We sequenced 317 novel ancient human genomes from primarily Mesolithic and Neolithic individuals across Eurasia, combined with new radiocarbon dates, stable isotope data. We additionally test and apply GLIMPSE to impute 1492 previously published ancient genomes, and incorporate this data in our analyses. We undertake a suite of analyses of this data to generate new insights into population structure through time, natural selection and health-related adaptation.

**Research sample**
Ancient human remains (bone and dental cementum) were collected, sampled, extracted and screened by shallow shotgun sequencing (see Supplementary Note 1). This resulted in 317 samples found to be suitable for deep sequencing (of 962 initial specimens). These represent a considerable geographic range across Eurasia (from Lake Baikal to the Atlantic coast). A high sample density for Denmark (100) was undertaken to provide high resolution insights at this location. A particular focus upon individuals related to Mesolithic or Neolithic culture was made, with 300 of the individuals from the time span 11,000 cal. BP to 3,000 cal. BP.

**Sampling strategy**
Sampling was dependent upon the availability of ancient human remains, though a considerable transect of the Mesolithic and Neolithic periods in Eurasia was represented, together with a detailed continuous sequence of human occupation of Denmark specifically. Our assemblage is well-represented with individuals of such key archaeological complexes as the Maglemose, Ertebølle and Funnel Beaker cultures in Scandinavia, the Cardial in the Mediterranean, the Körös and Linear Pottery complexes in SE and Central Europe, and many archaeological cultures in Ukraine, western Russia, and the trans-Ural (e.g. Veretye, Lyalovo, Volosovo, Kitoi).

**Data collection**
Sampled remains consisted of teeth (sampled for dental cementum, N=211), petrous bones (N=91), and other bone types (N=15; long bones, ribs and cranial bones). Sampling and ancient DNA laboratory work was undertaken in dedicated ancient DNA clean lab facilities at the GeoGenetics Centre. Additionally, 272 novel accelerator mass spectrometry results were generated at the 14CHRONO laboratory, Queen's University Belfast (242 samples), at the Oxford Radiocarbon Accelerator Unit (ORAU) laboratory (24 samples) and at the Keck-CCAMS Group, Irvine, California, USA (6 samples).

**Timing and spatial scale**
Sample chronology was generated by radiocarbon dating, with dates corrected for marine and freshwater reservoir effects. These showed samples ranging from the Upper Palaeolithic (c. 25,700 cal. BP) to the mediaeval period (c. 1200 cal. BP). Most individuals (97%, N=309) span 11,000 cal. BP to 3,000 cal. BP, the period broadly associated with the Mesolithic and Neolithic in Eurasia. Our research area can broadly be divided into three large regions: 1) central, western and northern Europe, 2) eastern Europe including western Russia and Ukraine, and 3) the Urals and western Siberia.

**Data exclusions**
Samples that were screened by shallow sequencing and found to have endogenous DNA of >1% were not sequenced further and not included in analyses. This amounted to 645 specimens.

**Reproducibility**
Data quality and uncertainty (e.g. contamination) was accounted for in computational analyses to assess robustness of inferences, and all methods and data are made available for future replication.

**Randomization**
Sample groups were defined to reflect archaeological populations, based on phylogenetic inferences, temporal and geographic provenance, and cultural interpretations evidenced by archaeological contexts.

**Blinding**
Blinding was not applicable to this study.

Did the study involve field work?

<figure><img src="image_61.png" /></figure>

Yes

<figure><img src="image_62.png" /></figure>

No

# Reporting for specific materials, systems and methods

We require information from authors about some types of materials, experimental systems and methods used in many studies. Here, indicate whether each material, system or method listed is relevant to your study. If you are not sure if a list item applies to your research, read the appropriate section before selecting a response.

## Materials & experimental systems

<table><thead><tr><th>n/a</th><th>Involved in the study</th></tr></thead><tbody><tr><td>☑</td><td>Antibodies</td></tr><tr><td>☑</td><td>Eukaryotic cell lines</td></tr><tr><td>☐</td><td>Palaeontology and archaeology</td></tr><tr><td>☑</td><td>Animals and other organisms</td></tr><tr><td>☑</td><td>Clinical data</td></tr><tr><td>☑</td><td>Dual use research of concern</td></tr></tbody></table>

## Methods

<table><thead><tr><th>n/a</th><th>Involved in the study</th></tr></thead><tbody><tr><td>☑</td><td>ChIP-seq</td></tr><tr><td>☑</td><td>Flow cytometry</td></tr><tr><td>☑</td><td>MRI-based neuroimaging</td></tr></tbody></table>
<table>
  <thead>
    <tr>
      <th>Specimen provenance</th>
      <th>Description</th>
    </tr>
  </thead>
  <tbody>
    <tr>
      <td></td>
      <td>Samples analysed in this study represent a considerable transect of Eurasian Mesolithic and Neolithic archaeological sites. Institution: 176 in total. Given the highly collaborative, international scale of the sampling effort, key contact persons for each site helped facilitate the access to material and coordinate legal authorisation for accessing sample material. Samples were collected with permission from organisations holding specimens and documented via permissions or agreements letters held by the Lundbeck Foundation GeoGenetics Centre. Full details and documentation for these is maintained in a dedicated database by F.B., K.G.S and Pernille Olsen of the GeoGenetics Centre. The following list provides the site name, country, sample ID, contact and institution. Further details of each site and excavation history are provided in Supplementary Notes 5-6.</td>
    </tr>
    <tr>
      <td></td>
      <td>Afontova Gora, Russia. Samples: NEO102. Key contact: Mikhail Sablin. Institution: Zoological Institute of the Russian Academy of Sciences.</td>
    </tr>
    <tr>
      <td></td>
      <td>Aknashen, Armenia. Samples: NEO110. Key contact: Levon Yeppiskoposyan. Institution: National Academy of Sciences of Armenia, Laboratory of Ethnogenomics, Yerevan, Armenia.</td>
    </tr>
    <tr>
      <td></td>
      <td>Ångdala, Sweden. Samples: NEO046. Key contact: Catharina Ödman / Yvonne Magnusson. Institution: Malmö museum.</td>
    </tr>
    <tr>
      <td></td>
      <td>Avlebjerg, Denmark. Samples: NEO961. Key contact: Anders Fischer. Institution: Sealand Archaeology/University of Copenhagen.</td>
    </tr>
    <tr>
      <td></td>
      <td>Baile herculane, Romania. Samples: NEO674. Key contact: Dušan Borić. Institution: Columbia university.</td>
    </tr>
    <tr>
      <td></td>
      <td>Banks Tomb, UK. Samples: NEO624, NEO625, NEO626, NEO627, NEO630, NEO717. Key contact: Nick Card. Institution: Orkney Research Centre for Archaeology. Institution: ORCA.</td>
    </tr>
    <tr>
      <td></td>
      <td>Bazaiha, Russia. Samples: NEO070. Key contact: Mikhail Sablin. Institution: Zoological Institute of the Russian Academy of Sciences.</td>
    </tr>
    <tr>
      <td></td>
      <td>Biestamak, Kazakhstan. Samples: NEO057. Key contact: Emma Usmanova, Irina Shevnina, Andrey Logvin. Institution: Karaganda State University.</td>
    </tr>
    <tr>
      <td></td>
      <td>Bjørnsholm, Denmark. Samples: NEO751. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/ University of Copenhagen.</td>
    </tr>
    <tr>
      <td></td>
      <td>Bodal K, Denmark. Samples: NEO814. Key contact: Lasse Sørensen, Peter Vang, Anders Fischer. Institution: National Museum of Denmark</td>
    </tr>
    <tr>
      <td></td>
      <td>Bol'shoy Oleni Ostrov, Russia. Samples: NEO060, NEO061, NEO062. Key contact: Vyacheslav Moiseyev. Institution: Peter the Great Museum of Anthropology and ethnography (Kunstkamera).</td>
    </tr>
    <tr>
      <td></td>
      <td>Borovjanka XVII, Russia. Samples: NEO080, NEO081, NEO083. Key contact: Vyacheslav Moiseyev. Institution: Peter the Great Museum of Anthropology and Ethnography (Kunstkamera).</td>
    </tr>
    <tr>
      <td></td>
      <td>Borreby, Denmark. Samples: NEO735, NEO737. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.</td>
    </tr>
    <tr>
      <td></td>
      <td>Bredgården, Sweden. Samples: NEO027. Key contact: Maria Vretemark. Institution: Västergötlands museum.</td>
    </tr>
    <tr>
      <td></td>
      <td>Bybjerg, Denmark. Samples: NEO563. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/ University of Copenhagen.</td>
    </tr>
    <tr>
      <td></td>
      <td>Bygholm, Denmark. Samples: NEO564. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/ University of Copenhagen.</td>
    </tr>
    <tr>
      <td></td>
      <td>Camino de las Yeseras, Spain. Samples: NEO721. Key contact: Corina Liesau, Pilar Prieto. Institution: Universidad Autónoma de Madrid.</td>
    </tr>
    <tr>
      <td></td>
      <td>Dalmosegaard, Denmark. Samples: NEO886. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.</td>
    </tr>
    <tr>
      <td></td>
      <td>Dezsk, Hungary. Samples: NEO130. Key contact: Vaclav Smrcka. Institution: Institute for History of Medicine and Foreign Languages of the First Faculty of Medicine, Charles University.</td>
    </tr>
    <tr>
      <td></td>
      <td>Døjringe, Denmark. Samples: NEO566. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/ University of Copenhagen.</td>
    </tr>
    <tr>
      <td></td>
      <td>Dolgoe Ozero, Russia. Samples: NEO292. Key contact: Sergey Vasilyev. Institution: Inst. of Ethnography and Anthropology, Moscow.</td>
    </tr>
    <tr>
      <td></td>
      <td>Dösemarken, Sweden. Samples: NEO044. Key contact: Catharina Ödman / Yvonne Magnusson. Institution: Malmö museum.</td>
    </tr>
    <tr>
      <td></td>
      <td>Dragsholm, Denmark. Samples: NEO822. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Danish National Museum.</td>
    </tr>
    <tr>
      <td></td>
      <td>Dragsholm, Denmark. Samples: NEO732, NEO733. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.</td>
    </tr>
    <tr>
      <td></td>
      <td>El Mazo, Spain. Samples: NEO646, NEO938. Key contact: Igor Gutiérrez Zugasti. Institution: Universidad de Cantabria.</td>
    </tr>
    <tr>
      <td></td>
      <td>Elkenøre, Denmark. Samples: NEO888. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/ University of Copenhagen.</td>
    </tr>
    <tr>
      <td></td>
      <td>Ertebølle, Denmark. Samples: NEO568, NEO569. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.</td>
    </tr>
    <tr>
      <td></td>
      <td>Evensås, Sweden. Samples: NEO260. Key contact: Karl-Göran Sjögren. Institution: Gothenburg university.</td>
    </tr>
    <tr>
      <td></td>
      <td>Falköping 5, Sweden. Samples: NEO220, NEO221, NEO223, NEO224, NEO225, NEO226, NEO227, NEO228. Key contact: Malou Blank. Institution: Gothenburg university.</td>
    </tr>
    <tr>
      <td></td>
      <td>Fannerup D, Denmark. Samples: NEO855. Key contact: Lutz Klassen. Institution: Museum Østjylland.</td>
    </tr>
    <tr>
      <td></td>
      <td>Fannerup E, Denmark. Samples: NEO570. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/ University of Copenhagen.</td>
    </tr>
    <tr>
      <td></td>
      <td>Fannerup F, Denmark. Samples: NEO930. Key contact: Lutz Klassen / Rikke Maring. Institution: Museum Østjylland.</td>
    </tr>
    <tr>
      <td></td>
      <td>Femhøve Vig, Denmark. Samples: NEO744. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.</td>
    </tr>
    <tr>
      <td></td>
      <td>Fofonovo, Russia. Samples: NEO199, NEO200, NEO201. Key contact: Sergey Vasilyev. Institution: Inst. of Ethnography and Anthropology, Moscow.</td>
    </tr>
    <tr>
      <td></td>
      <td>Fontenoce, Italy. Samples: NEO830. Key contact: Gabriele Scorrano. Institution: University of Copenhagen.</td>
    </tr>
    <tr>
      <td></td>
      <td>Frälsegården, Sweden. Samples: NEO259. Key contact: Karl-Göran Sjögren. Institution: Gothenburg university.</td>
    </tr>
    <tr>
      <td></td>
      <td>Gaudo, Italy. Samples: NEO828. Key contact: Gabriele Scorrano. Institution: University of Copenhagen.</td>
    </tr>
    <tr>
      <td></td>
      <td>Gazel Cave, France. Samples: NEO812, NEO813. Key contact: Patrice Cortaud. Institution: Universite de Bordeaux.</td>
    </tr>
    <tr>
      <td></td>
      <td>Golubaya Krinitsa, Russia. Samples: NEO204, NEO207, NEO209, NEO210, NEO212. Key contact: Ruslan Turin. Institution: Karaganda</td>
    </tr>
  </tbody>
</table>
<table>
  <tr>
    <td>Specimen provenance</td>
    <td>Samples analysed in this study represent a considerable transect of Eurasian Mesolithic and Neolithic archaeological sites. Institution: 176 in total. Given the highly collaborative, international scale of the sampling effort, key contact persons for each site helped facilitate the access to material and coordinate legal authorisation for accessing sample material. Samples were collected with permission from organisations holding specimens and documented via permissions or agreements letters held by the Lundbeck Foundation GeoGenetics Centre. Full details and documentation for these is maintained in a dedicated database by F.B., K.G.S and Pernille Olsen of the GeoGenetics Centre. The following list provides the site name, country, sample ID, contact and institution. Further details of each site and excavation history are provided in Supplementary Notes 5-6.</td>
  </tr>
  <tr>
    <td>Afontova Gora, Russia.</td>
    <td>Samples: NEO102. Key contact: Mikhail Sablin. Institution: Zoological Institute of the Russian Academy of Sciences.</td>
  </tr>
  <tr>
    <td>Aknashen, Armenia.</td>
    <td>Samples: NEO110. Key contact: Levon Yeppiskoposyan. Institution: National Academy of Sciences of Armenia, Laboratory of Ethnogenomics, Yerevan, Armenia.</td>
  </tr>
  <tr>
    <td>Ångdala, Sweden.</td>
    <td>Samples: NEO046. Key contact: Catharina Ödman / Yvonne Magnusson. Institution: Malmö museum.</td>
  </tr>
  <tr>
    <td>Avlebjerg, Denmark.</td>
    <td>Samples: NEO961. Key contact: Anders Fischer. Institution: Sealand Archaeology/University of Copenhagen.</td>
  </tr>
  <tr>
    <td>Baile herculane, Romania.</td>
    <td>Samples: NEO674. Key contact: Dušan Borić. Institution: Columbia university.</td>
  </tr>
  <tr>
    <td>Banks Tomb, UK.</td>
    <td>Samples: NEO624, NEO625, NEO626, NEO627, NEO630, NEO717. Key contact: Nick Card. Institution: Orkney Research Centre for Archaeology. Institution: ORCA.</td>
  </tr>
  <tr>
    <td>Bazaiha, Russia.</td>
    <td>Samples: NEO070. Key contact: Mikhail Sablin. Institution: Zoological Institute of the Russian Academy of Sciences.</td>
  </tr>
  <tr>
    <td>Biestamak, Kazakhstan.</td>
    <td>Samples: NEO057. Key contact: Emma Usmanova, Irina Shevnina, Andrey Logvin. Institution: Karaganda State University.</td>
  </tr>
  <tr>
    <td>Bjørnsholm, Denmark.</td>
    <td>Samples: NEO751. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/ University of Copenhagen.</td>
  </tr>
  <tr>
    <td>Bodal K, Denmark.</td>
    <td>Samples: NEO814. Key contact: Lasse Sørensen, Peter Vang, Anders Fischer. Institution: National Museum of Denmark</td>
  </tr>
  <tr>
    <td>Bol'shoy Oleni Ostrov, Russia.</td>
    <td>Samples: NEO060, NEO061, NEO062. Key contact: Vyacheslav Moiseyev. Institution: Peter the Great Museum of Anthropology and ethnography (Kunstkamera).</td>
  </tr>
  <tr>
    <td>Borovjanka XVII, Russia.</td>
    <td>Samples: NEO080, NEO081, NEO083. Key contact: Vyacheslav Moiseyev. Institution: Peter the Great Museum of Anthropology and Ethnography (Kunstkamera).</td>
  </tr>
  <tr>
    <td>Borreby, Denmark.</td>
    <td>Samples: NEO735, NEO737. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.</td>
  </tr>
  <tr>
    <td>Bredgården, Sweden.</td>
    <td>Samples: NEO027. Key contact: Maria Vretemark. Institution: Västergötlands museum.</td>
  </tr>
  <tr>
    <td>Bybjerg, Denmark.</td>
    <td>Samples: NEO563. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/ University of Copenhagen.</td>
  </tr>
  <tr>
    <td>Bygholm, Denmark.</td>
    <td>Samples: NEO564. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/ University of Copenhagen.</td>
  </tr>
  <tr>
    <td>Camino de las Yeseras, Spain.</td>
    <td>Samples: NEO721. Key contact: Corina Liesau, Pilar Prieto. Institution: Universidad Autónoma de Madrid.</td>
  </tr>
  <tr>
    <td>Dalmosegaard, Denmark.</td>
    <td>Samples: NEO886. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.</td>
  </tr>
  <tr>
    <td>Dezsk, Hungary.</td>
    <td>Samples: NEO130. Key contact: Vaclav Smrcka. Institution: Institute for History of Medicine and Foreign Languages of the First Faculty of Medicine, Charles University.</td>
  </tr>
  <tr>
    <td>Døjringe, Denmark.</td>
    <td>Samples: NEO566. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/ University of Copenhagen.</td>
  </tr>
  <tr>
    <td>Dolgoe Ozero, Russia.</td>
    <td>Samples: NEO292. Key contact: Sergey Vasilyev. Institution: Inst. of Ethnography and Anthropology, Moscow.</td>
  </tr>
  <tr>
    <td>Dösemarken, Sweden.</td>
    <td>Samples: NEO044. Key contact: Catharina Ödman / Yvonne Magnusson. Institution: Malmö museum.</td>
  </tr>
  <tr>
    <td>Dragsholm, Denmark.</td>
    <td>Samples: NEO822. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Danish National Museum.</td>
  </tr>
  <tr>
    <td>Dragsholm, Denmark.</td>
    <td>Samples: NEO732, NEO733. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.</td>
  </tr>
  <tr>
    <td>El Mazo, Spain.</td>
    <td>Samples: NEO646, NEO938. Key contact: Igor Gutiérrez Zugasti. Institution: Universidad de Cantabria.</td>
  </tr>
  <tr>
    <td>Elkenøre, Denmark.</td>
    <td>Samples: NEO888. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/ University of Copenhagen.</td>
  </tr>
  <tr>
    <td>Ertebølle, Denmark.</td>
    <td>Samples: NEO568, NEO569. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.</td>
  </tr>
  <tr>
    <td>Evensås, Sweden.</td>
    <td>Samples: NEO260. Key contact: Karl-Göran Sjögren. Institution: Gothenburg university.</td>
  </tr>
  <tr>
    <td>Falköping 5, Sweden.</td>
    <td>Samples: NEO220, NEO221, NEO223, NEO224, NEO225, NEO226, NEO227, NEO228. Key contact: Malou Blank. Institution: Gothenburg university.</td>
  </tr>
  <tr>
    <td>Fannerup D, Denmark.</td>
    <td>Samples: NEO855. Key contact: Lutz Klassen. Institution: Museum Østjylland.</td>
  </tr>
  <tr>
    <td>Fannerup E, Denmark.</td>
    <td>Samples: NEO570. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/ University of Copenhagen.</td>
  </tr>
  <tr>
    <td>Fannerup F, Denmark.</td>
    <td>Samples: NEO930. Key contact: Lutz Klassen / Rikke Maring. Institution: Museum Østjylland.</td>
  </tr>
  <tr>
    <td>Femhøve Vig, Denmark.</td>
    <td>Samples: NEO744. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.</td>
  </tr>
  <tr>
    <td>Fofonovo, Russia.</td>
    <td>Samples: NEO199, NEO200, NEO201. Key contact: Sergey Vasilyev. Institution: Inst. of Ethnography and Anthropology, Moscow.</td>
  </tr>
  <tr>
    <td>Fontenoce, Italy.</td>
    <td>Samples: NEO830. Key contact: Gabriele Scorrano. Institution: University of Copenhagen.</td>
  </tr>
  <tr>
    <td>Frälsegården, Sweden.</td>
    <td>Samples: NEO259. Key contact: Karl-Göran Sjögren. Institution: Gothenburg university.</td>
  </tr>
  <tr>
    <td>Gaudo, Italy.</td>
    <td>Samples: NEO828. Key contact: Gabriele Scorrano. Institution: University of Copenhagen.</td>
  </tr>
  <tr>
    <td>Gazel Cave, France.</td>
    <td>Samples: NEO812, NEO813. Key contact: Patrice Cortaud. Institution: Universite de Bordeaux.</td>
  </tr>
  <tr>
    <td>Golubaya Krinitsa, Russia.</td>
    <td>Samples: NEO204, NEO207, NEO209, NEO210, NEO212. Key contact: Ruslan Turin. Institution: Karaganda</td>
  </tr>
</table>
State University.

Golubaya Krinitsa, Russia. Samples: NEO113. Key contact: Sergey Vasilyev. Institution: Inst. of Ethnography and Anthropology, Moscow.

Gorzsa Cukormajor, Hungary. Samples: NEO140, NEO142, NEO143, NEO145, NEO147, NEO148, NEO149. Key contact: Vaclav Smrcka.

Institution: Institute for History of Medicine and Foreign Languages of the First Faculty of Medicine, Charles University.

Gregorievka 1, Kazakhstan. Samples: NEO899. Key contact: Victor Merc/Emma Usmanova. Institution: Karaganda State University.

Grøfte, Denmark. Samples: NEO571. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/ University of Copenhagen.

Grotta Delle Mura, Italy. Samples: NEO806. Key contact: Gabriele Scorrano. Institution: University of Copenhagen.

Grotta Nisco, Italy. Samples: NEO823. Key contact: Gabriele Scorrano. Institution: University of Copenhagen.

Gruta do Caldeirão, Portugal. Samples: NEO631, NEO632. Key contact: Joao Zilhao. Institution: Universitat de Barcelona.

Hanaskede, Sweden. Samples: NEO018. Key contact: Maria Vretemark. Institution: Västergötlands museum.

Havnø, Denmark. Samples: NEO941. Key contact: Anders Fischer/Niels Lynnerup. Institution: Sealand Archaeology/University of Copenhagen.

Hedegaard. Institution: Bislev), Denmark. Samples: NEO013. Key contact: Bjarne Nielsen.

Henriksholm-Bøgebakken. Institution: Vedbæk), Denmark. Samples: NEO745, NEO746, NEO747, NEO748, NEO749. Key contact: Niels

Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Hetty Pegler's Tump, UK. Samples: NEO847. Key contact: Alan Outram. Institution: University of Exeter.

Hindby Mosse, Sweden. Samples: NEO036, NEO038, NEO039. Key contact: Catharina Ödman / Yvonne Magnusson. Institution: Malmö museum.

Hödmezöväsähely Kotac, Hungary. Samples: NEO137. Key contact: Vaclav Smrcka. Institution: Institute for History of Medicine and Foreign Languages of the First Faculty of Medicine, Charles University.

Holmegård-Djursland, Denmark. Samples: NEO001. Key contact: Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/ University of Copenhagen.

Hove Å, Denmark. Samples: NEO946. Key contact: Anders Fischer/Niels Lynnerup. Institution: Sealand Archaeology/University of Copenhagen.

Hummervik, Norway. Samples: NEO017. Key contact: Per Åke Persson. Institution: UiO: Kulturhistorisk museum.

Igren, Ukraine. Samples: NEO509, NEO516, NEO518, NEO521. Key contact: Alexandra Buzhilova. Institution: Lomonosov Moscow State University.

Itkul', Russia. Samples: NEO063, NEO064, NEO065. Key contact: Vyacheslav Moiseyev. Institution: Peter the Great Museum of Anthropology and Ethnography (Kunstkamera).

Jørløse Mose, Denmark. Samples: NEO023. Key contact: Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Jørlundegaard, Denmark. Samples: NEO702. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Kainsbakke, Denmark. Samples: NEO025. Key contact: Lutz Klassen. Institution: Museum Østjylland.

Karavaikha, Russia. Samples: NEO555, NEO556, NEO557, NEO558, NEO559, NEO560, NEO561. Key contact: Alexandra Buzhilova.

Institution: Lomonosov Moscow State University.

Kastanjegården, Sweden. Samples: NEO051. Key contact: Catharina Ödman / Yvonne Magnusson. Institution: Malmö museum.

Klastrup Holm, Denmark. Samples: NEO951. Key contact: Anders Fischer/Niels Lynnerup. Institution: Sealand Archaeology/University of Copenhagen.

Kleshnya III 1998, Ukraine. Samples: NEO278. Key contact: Inna Potekhina. Institution: Institute of Archaeology of Ukrainian National Academy of Sciences.

Klokkehøj, Denmark. Samples: NEO580. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/ University of Copenhagen.

Koed I, Denmark. Samples: NEO583. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/ University of Copenhagen.

Koed IV, Denmark. Samples: NEO586. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/ University of Copenhagen.

Koelbjerg, Denmark. Samples: NEO254. Key contact: Jesper Hansen. Institution: Odense Bys Museer.

Køge Sønakke, Denmark. Samples: NEO759. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Kolind, Denmark. Samples: NEO738, NEO739. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Kongemose, Denmark. Samples: NEO587. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/ University of Copenhagen.

Korsør Glasværk, Denmark. Samples: NEO589. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Korsør Nor, Denmark. Samples: NEO791. Key contact: Lasse Sørensen, Poul Otto Nielsen. Institution: National Museum of Denmark.

Kotias Klde Cave, Georgia. Samples: NEO281, NEO283. Key contact: David O. Lordkipanidze. Institution: Georgian National Museum.

Ksizovo 6, Russia. Samples: NEO172, NEO173, NEO174, NEO175. Key contact: Sergey Vasilyev. Institution: Inst. of Ethnography and Anthropology, Moscow.

Kumyshanskaya Cave, Russia. Samples: NEO687. Key contact: Andrej Evteev. Institution: Moscow State University.

Kyndeløse, Denmark. Samples: NEO878. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/ University of Copenhagen.

Læsten Mose, Denmark. Samples: NEO945. Key contact: Anders Fischer/Niels Lynnerup. Institution: Sealand Archaeology/University of Copenhagen.

Langø Skaldynge, Denmark. Samples: NEO853. Key contact: Anders Fischer/ Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Lendemark, Denmark. Samples: NEO896. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/ University of Copenhagen.

Lepenski Vir, Serbia. Samples: NEO669. Key contact: Dušan Borić. Institution: Columbia university.

Lohals Nord, Denmark. Samples: NEO029. Key contact: Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Lollikehuse, Denmark. Samples: NEO857. Key contact: Kristian Gregersen. Institution: Statens Naturhistoriske Museum.

Lundby-Falster, Denmark. Samples: NEO865, NEO866. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand
State University.

Golubaya Krinitsa, Russia. Samples: NEO113. Key contact: Sergey Vasilyev. Institution: Inst. of Ethnography and Anthropology, Moscow.

Gorzsa Cukormajor, Hungary. Samples: NEO140, NEO142, NEO143, NEO145, NEO147, NEO148, NEO149. Key contact: Vaclav Smrcka.

Institution: Institute for History of Medicine and Foreign Languages of the First Faculty of Medicine, Charles University.

Gregorievka 1, Kazakhstan. Samples: NEO899. Key contact: Victor Merc/Emma Usmanova. Institution: Karaganda State University.

Grøfte, Denmark. Samples: NEO571. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/ University of Copenhagen.

Grotta Delle Mura, Italy. Samples: NEO806. Key contact: Gabriele Scorrano. Institution: University of Copenhagen.

Grotta Nisco, Italy. Samples: NEO823. Key contact: Gabriele Scorrano. Institution: University of Copenhagen.

Gruta do Caldeirão, Portugal. Samples: NEO631, NEO632. Key contact: Joao Zilhao. Institution: Universitat de Barcelona.

Hanaskede, Sweden. Samples: NEO018. Key contact: Maria Vretemark. Institution: Västergötlands museum.

Havnø, Denmark. Samples: NEO941. Key contact: Anders Fischer/Niels Lynnerup. Institution: Sealand Archaeology/University of Copenhagen.

Hedegaard. Institution: Bislev), Denmark. Samples: NEO013. Key contact: Bjarne Nielsen.

Henriksholm-Bøgebakken. Institution: Vedbæk), Denmark. Samples: NEO745, NEO746, NEO747, NEO748, NEO749. Key contact: Niels

Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Hetty Pegler's Tump, UK. Samples: NEO847. Key contact: Alan Outram. Institution: University of Exeter.

Hindby Mosse, Sweden. Samples: NEO036, NEO038, NEO039. Key contact: Catharina Ödman / Yvonne Magnusson. Institution: Malmö museum.

Hödmezöväsähely Kotac, Hungary. Samples: NEO137. Key contact: Vaclav Smrcka. Institution: Institute for History of Medicine and Foreign Languages of the First Faculty of Medicine, Charles University.

Holmegård-Djursland, Denmark. Samples: NEO001. Key contact: Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/ University of Copenhagen.

Hove Å, Denmark. Samples: NEO946. Key contact: Anders Fischer/Niels Lynnerup. Institution: Sealand Archaeology/University of Copenhagen.

Hummervik, Norway. Samples: NEO017. Key contact: Per Åke Persson. Institution: UiO: Kulturhistorisk museum.

Igren, Ukraine. Samples: NEO509, NEO516, NEO518, NEO521. Key contact: Alexandra Buzhilova. Institution: Lomonosov Moscow State University.

Itkul', Russia. Samples: NEO063, NEO064, NEO065. Key contact: Vyacheslav Moiseyev. Institution: Peter the Great Museum of Anthropology and Ethnography (Kunstkamera).

Jørløse Mose, Denmark. Samples: NEO023. Key contact: Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Jørlundegaard, Denmark. Samples: NEO702. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Kainsbakke, Denmark. Samples: NEO025. Key contact: Lutz Klassen. Institution: Museum Østjylland.

Karavaikha, Russia. Samples: NEO555, NEO556, NEO557, NEO558, NEO559, NEO560, NEO561. Key contact: Alexandra Buzhilova.

Institution: Lomonosov Moscow State University.

Kastanjegården, Sweden. Samples: NEO051. Key contact: Catharina Ödman / Yvonne Magnusson. Institution: Malmö museum.

Klastrup Holm, Denmark. Samples: NEO951. Key contact: Anders Fischer/Niels Lynnerup. Institution: Sealand Archaeology/University of Copenhagen.

Kleshnya III 1998, Ukraine. Samples: NEO278. Key contact: Inna Potekhina. Institution: Institute of Archaeology of Ukrainian National Academy of Sciences.

Klokkehøj, Denmark. Samples: NEO580. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/ University of Copenhagen.

Koed I, Denmark. Samples: NEO583. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/ University of Copenhagen.

Koed IV, Denmark. Samples: NEO586. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/ University of Copenhagen.

Koelbjerg, Denmark. Samples: NEO254. Key contact: Jesper Hansen. Institution: Odense Bys Museer.

Køge Sønakke, Denmark. Samples: NEO759. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Kolind, Denmark. Samples: NEO738, NEO739. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Kongemose, Denmark. Samples: NEO587. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/ University of Copenhagen.

Korsør Glasværk, Denmark. Samples: NEO589. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Korsør Nor, Denmark. Samples: NEO791. Key contact: Lasse Sørensen, Poul Otto Nielsen. Institution: National Museum of Denmark.

Kotias Klde Cave, Georgia. Samples: NEO281, NEO283. Key contact: David O. Lordkipanidze. Institution: Georgian National Museum.

Ksizovo 6, Russia. Samples: NEO172, NEO173, NEO174, NEO175. Key contact: Sergey Vasilyev. Institution: Inst. of Ethnography and Anthropology, Moscow.

Kumyshanskaya Cave, Russia. Samples: NEO687. Key contact: Andrej Evteev. Institution: Moscow State University.

Kyndeløse, Denmark. Samples: NEO878. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/ University of Copenhagen.

Læsten Mose, Denmark. Samples: NEO945. Key contact: Anders Fischer/Niels Lynnerup. Institution: Sealand Archaeology/University of Copenhagen.

Langø Skaldynge, Denmark. Samples: NEO853. Key contact: Anders Fischer/ Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Lendemark, Denmark. Samples: NEO896. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/ University of Copenhagen.

Lepenski Vir, Serbia. Samples: NEO669. Key contact: Dušan Borić. Institution: Columbia University.

Lohals Nord, Denmark. Samples: NEO029. Key contact: Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Lollikehuse, Denmark. Samples: NEO857. Key contact: Kristian Gregersen. Institution: Statens Naturhistoriske Museum.

Lundby-Falster, Denmark. Samples: NEO865, NEO866. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand
Archaeology/University of Copenhagen.

Lysa Gora, Ukraine. Samples: NEO262, NEO265. Key contact: Inna Potekhina. Institution: Institute of Archaeology of Ukrainian National Academy of Sciences.

Maddalena, Italy. Samples: NEO695. Key contact: Alfredo Coppa. Institution: Universita di Roma Sapienza.

Madesø, Denmark. Samples: NEO752. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Magleø, Denmark. Samples: NEO590. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Mamaj Gora, Ukraine. Samples: NEO268, NEO270. Key contact: Inna Potekhina. Institution: Institute of Archaeology of Ukrainian National Academy of Sciences.

Mandrin Cave, France. Samples: NEO119, NEO120, NEO121. Key contact: Ludovic Slimak. Institution: Université Toulouse.

Mandzuli-Depe, Turkmenistan. Samples: NEO310. Key contact: Sergey Vasilyev. Institution: Inst. of Ethnography and Anthropology, Moscow.

Mergen' 6, Russia. Samples: NEO072, NEO073. Key contact: Vyacheslav Moiseyev. Institution: Peter the Great Museum of Anthropology and ethnography (Kunstkamera).

Minino, Russia. Samples: NEO536, NEO537, NEO538, NEO539. Key contact: Alexandra Buzhilova. Institution: Lomonosov Moscow State University.

Mora Cavorso, Italy. Samples: NEO834. Key contact: Gabriele Scorrano. Institution: University of Copenhagen.

Mosede Mose. Institution: Karlslunde Mose), Denmark. Samples: NEO860. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Mosede Mose III, Denmark. Samples: NEO861. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Myrebjerg Mose, Denmark. Samples: NEO925. Key contact: Otto Christian Uldum. Institution: Langelands Museum.

Næs, Denmark. Samples: NEO792. Key contact: Lasse Sørensen, Poul Otto Nielsen. Institution: National Museum of Denmark.

Nederst, Denmark. Samples: NEO856. Key contact: Lutz Klassen. Institution: Museum Østjylland.

Neverkær Mose I, Denmark. Samples: NEO594. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Norsminde, Denmark. Samples: NEO852. Key contact: Anders Fischer/ Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Okunevo 5, Russia. Samples: NEO068, NEO077. Key contact: Vyacheslav Moiseyev. Institution: Peter the Great Museum of Anthropology and ethnography (Kunstkamera).

Okunevo 7, Russia. Samples: NEO079. Key contact: Vyacheslav Moiseyev. Institution: Peter the Great Museum of Anthropology and ethnography (Kunstkamera).

Omskaya Stoyanka II, Russia. Samples: NEO075, NEO078. Key contact: Vyacheslav Moiseyev. Institution: Peter the Great Museum of Anthropology and ethnography (Kunstkamera).

Orehoved Sejlrende, Denmark. Samples: NEO122, NEO123. Key contact: Anders Fischer, Pia Bennike, Morten Johansen. Institution: Sealand Archaeology/University of Copenhagen.

Ostrov 2, Russia. Samples: NEO076. Key contact: Vyacheslav Moiseyev. Institution: Peter the Great Museum of Anthropology and ethnography (Kunstkamera).

Pad Tokui, Russia. Samples: NEO116. Key contact: Sergey Vasilyev. Institution: Inst. of Ethnography and Anthropology, Moscow.

Pandebjerg, Denmark. Samples: NEO595. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Peschanitsa, Russia. Samples: NEO202. Key contact: Sergey Vasilyev. Institution: Inst. of Ethnography and Anthropology, Moscow.

Pogostishche, Russia. Samples: NEO554. Key contact: Alexandra Buzhilova. Institution: Lomonosov Moscow State University.

Porsmose, Denmark. Samples: NEO0795. Key contact: Lasse Sørensen, Poul Otto Nielsen. Institution: National Museum of Denmark.

Protoka, Russia. Samples: NEO309. Key contact: Sergey Vasilyev. Institution: Inst. of Ethnography and Anthropology, Moscow.

Ravnsbjerggård, Denmark. Samples: NEO960. Key contact: Anders Fischer. Institution: Sealand Archaeology/University of Copenhagen.

Rødhals (Sejerø), Denmark. Samples: NEO645. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Røntesten, Denmark. Samples: NEO019. Key contact: Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Roskilde Fjord, Denmark. Samples: NEO891. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Rude, Denmark. Samples: NEO041, NEO043. Key contact: Niels Lynnerup. Institution: Department of Forensic Medicine, Copenhagen university.

Sakhtish 8, Russia. Samples: NEO184. Key contact: Sergey Vasilyev. Institution: Inst. of Ethnography and Anthropology, Moscow.

Sakhtish II, Russia. Samples: NEO158, NEO178, NEO179, NEO192. Key contact: Sergey Vasilyev. Institution: Inst. of Ethnography and Anthropology, Moscow.

Sakhtish IIa, Russia. Samples: NEO180, NEO181, NEO182, NEO183, NEO185, NEO186, NEO187, NEO188, NEO189, NEO193, NEO194, NEO195, NEO197. Key contact: Sergey Vasilyev. Institution: Inst. of Ethnography and Anthropology, Moscow.

Salpetermosen, Denmark. Samples: NEO028. Key contact: Niels Lynnerup/Thomas Jørgensen.

Santa Maira, Spain. Samples: NEO694. Key contact: Carles Lalueza-Fox. Institution: Pompeu Fabra.

Sao Paulo II, Portugal. Samples: NEO603, NEO609. Key contact: Ana Maria Silva. Institution: University of Coimbra.

Schela Cladovei, Romania. Samples: NEO671, NEO672. Key contact: Dušan Borić. Institution: Columbia university.

Sejerby (Sejerø), Denmark. Samples: NEO757. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Sigersdal, Denmark. Samples: NEO007. Key contact: Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Sigersdal Mose, Denmark. Samples: NEO753. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Sillvik, Sweden. Samples: NEO261. Key contact: Karl-Göran Sjögren. Institution: Gothenburg university.

Sjauke, Kazakhstan. Samples: NEO900. Key contact: Victor Merc/Emma Usmanova. Institution: Karaganda State University.

Sjauke 1, Kazakhstan. Samples: NEO902. Key contact: Victor Merc/Emma Usmanova. Institution: Karaganda State University.

Sjauke settlement, Kazakhstan. Samples: NEO901. Key contact: Victor Merc/Emma Usmanova. Institution: Karaganda State University.
Archaeology/University of Copenhagen.

Lysa Gora, Ukraine. Samples: NEO262, NEO265. Key contact: Inna Potekhina. Institution: Institute of Archaeology of Ukrainian National Academy of Sciences.

Maddalena, Italy. Samples: NEO695. Key contact: Alfredo Coppa. Institution: Universita di Roma Sapienza.

Madesø, Denmark. Samples: NEO752. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Magleø, Denmark. Samples: NEO590. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Mamaj Gora, Ukraine. Samples: NEO268, NEO270. Key contact: Inna Potekhina. Institution: Institute of Archaeology of Ukrainian National Academy of Sciences.

Mandrin Cave, France. Samples: NEO119, NEO120, NEO121. Key contact: Ludovic Slimak. Institution: Université Toulouse.

Mandzuli-Depe, Turkmenistan. Samples: NEO310. Key contact: Sergey Vasilyev. Institution: Inst. of Ethnography and Anthropology, Moscow.

Mergen' 6, Russia. Samples: NEO072, NEO073. Key contact: Vyacheslav Moiseyev. Institution: Peter the Great Museum of Anthropology and ethnography (Kunstkamera).

Minino, Russia. Samples: NEO536, NEO537, NEO538, NEO539. Key contact: Alexandra Buzhilova. Institution: Lomonosov Moscow State University.

Mora Cavorso, Italy. Samples: NEO834. Key contact: Gabriele Scorrano. Institution: University of Copenhagen.

Mosede Mose. Institution: Karlslunde Mose), Denmark. Samples: NEO860. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Mosede Mose III, Denmark. Samples: NEO861. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Myrebjerg Mose, Denmark. Samples: NEO925. Key contact: Otto Christian Uldum. Institution: Langelands Museum.

Næs, Denmark. Samples: NEO792. Key contact: Lasse Sørensen, Poul Otto Nielsen. Institution: National Museum of Denmark.

Nederst, Denmark. Samples: NEO856. Key contact: Lutz Klassen. Institution: Museum Østjylland.

Neverkær Mose I, Denmark. Samples: NEO594. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Norsminde, Denmark. Samples: NEO852. Key contact: Anders Fischer/ Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Okunevo 5, Russia. Samples: NEO068, NEO077. Key contact: Vyacheslav Moiseyev. Institution: Peter the Great Museum of Anthropology and ethnography (Kunstkamera).

Okunevo 7, Russia. Samples: NEO079. Key contact: Vyacheslav Moiseyev. Institution: Peter the Great Museum of Anthropology and ethnography (Kunstkamera).

Omskaya Stoyanka II, Russia. Samples: NEO075, NEO078. Key contact: Vyacheslav Moiseyev. Institution: Peter the Great Museum of Anthropology and ethnography (Kunstkamera).

Orehoved Sejlrende, Denmark. Samples: NEO122, NEO123. Key contact: Anders Fischer, Pia Bennike, Morten Johansen. Institution: Sealand Archaeology/University of Copenhagen.

Ostrov 2, Russia. Samples: NEO076. Key contact: Vyacheslav Moiseyev. Institution: Peter the Great Museum of Anthropology and ethnography (Kunstkamera).

Pad Tokui, Russia. Samples: NEO116. Key contact: Sergey Vasilyev. Institution: Inst. of Ethnography and Anthropology, Moscow.

Pandebjerg, Denmark. Samples: NEO595. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Peschanitsa, Russia. Samples: NEO202. Key contact: Sergey Vasilyev. Institution: Inst. of Ethnography and Anthropology, Moscow.

Pogostishche, Russia. Samples: NEO554. Key contact: Alexandra Buzhilova. Institution: Lomonosov Moscow State University.

Porsmose, Denmark. Samples: NEO0795. Key contact: Lasse Sørensen, Poul Otto Nielsen. Institution: National Museum of Denmark.

Protoka, Russia. Samples: NEO309. Key contact: Sergey Vasilyev. Institution: Inst. of Ethnography and Anthropology, Moscow.

Ravnsbjerggård, Denmark. Samples: NEO960. Key contact: Anders Fischer. Institution: Sealand Archaeology/University of Copenhagen.

Rødhals (Sejerø), Denmark. Samples: NEO645. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Røntesten, Denmark. Samples: NEO019. Key contact: Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Roskilde Fjord, Denmark. Samples: NEO891. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Rude, Denmark. Samples: NEO041, NEO043. Key contact: Niels Lynnerup. Institution: Department of Forensic Medicine, Copenhagen university.

Sakhtish 8, Russia. Samples: NEO184. Key contact: Sergey Vasilyev. Institution: Inst. of Ethnography and Anthropology, Moscow.

Sakhtish II, Russia. Samples: NEO158, NEO178, NEO179, NEO192. Key contact: Sergey Vasilyev. Institution: Inst. of Ethnography and Anthropology, Moscow.

Sakhtish IIa, Russia. Samples: NEO180, NEO181, NEO182, NEO183, NEO185, NEO186, NEO187, NEO188, NEO189, NEO193, NEO194, NEO195, NEO197. Key contact: Sergey Vasilyev. Institution: Inst. of Ethnography and Anthropology, Moscow.

Salpetermosen, Denmark. Samples: NEO028. Key contact: Niels Lynnerup/Thomas Jørgensen.

Santa Maira, Spain. Samples: NEO694. Key contact: Carles Lalueza-Fox. Institution: Pompeu Fabra.

Sao Paulo II, Portugal. Samples: NEO603, NEO609. Key contact: Ana Maria Silva. Institution: University of Coimbra.

Schela Cladovei, Romania. Samples: NEO671, NEO672. Key contact: Dušan Borić. Institution: Columbia university.

Sejerby (Sejerø), Denmark. Samples: NEO757. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Sigersdal, Denmark. Samples: NEO007. Key contact: Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Sigersdal Mose, Denmark. Samples: NEO753. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Sillvik, Sweden. Samples: NEO261. Key contact: Karl-Göran Sjögren. Institution: Gothenburg university.

Sjauke, Kazakhstan. Samples: NEO900. Key contact: Victor Merc/Emma Usmanova. Institution: Karaganda State University.

Sjauke 1, Kazakhstan. Samples: NEO902. Key contact: Victor Merc/Emma Usmanova. Institution: Karaganda State University.

Sjauke settlement, Kazakhstan. Samples: NEO901. Key contact: Victor Merc/Emma Usmanova. Institution: Karaganda State University.
Sjیدری 10, Kazakhstan. Samples: NEO904. Key contact: Victor Merc/Emma Usmanova. Institution: Karaganda State University.

Skateholm I, Sweden. Samples: NEO679. Key contact: Douglas Price / Lars Larsson, Institution: Lund university.

Słonowice, Poland. Samples: NEO640, NEO641. Key contact: Piotr Wlodarczak. Institution: Polish Academy of Sciences. Institution: Instytut Archeologii i Etnologii.

Sludegård Sømose, Denmark. Samples: NEO933. Key contact: Anders Fischer / Jesper Hansen. Institution: Odense Bys museer.

Sølager, Denmark. Samples: NEO598. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/ University of Copenhagen.

Sope-2, Estonia. Samples: NEO306. Key contact: Sergey Vasilyev. Institution: Inst. of Ethnography and Anthropology, Moscow.

Sosnovyiy Myis, Russia. Samples: NEO841, NEO843. Key contact: Vyacheslav Moiseyev. Institution: Peter the Great Museum of Anthropology and ethnography (Kunstkamera).

Stenderup Hage, Denmark. Samples: NEO943. Key contact: Anders Fischer/Niels Lynnerup.

Storelyng. Institution: Øgårde boat III), Denmark. Samples: NEO597. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike.

Institution: Sealand Archaeology/University of Copenhagen.

Storelyng (Østrup Homo II), Denmark. Samples: NEO602. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution:

Sealand Archaeology/University of Copenhagen.

Strøby Egede, Denmark. Samples: NEO092. Key contact: Niels Lynnerup. Institution: Department of Forensic Medicine, Copenhagen university.

Strøby Grøftemark, Denmark. Samples: NEO091. Key contact: Niels Lynnerup, Anders Fischer. Institution: Department of Forensic Medicine, Copenhagen university.

Strøby Ladeplads, Denmark. Samples: NEO093. Key contact: Niels Lynnerup. Institution: Department of Forensic Medicine, Copenhagen university.

Svinninge Vejle, Denmark. Samples: NEO898. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Tepe Guran, Iran. Samples: NEO816, NEO817, NEO819. Key contact: Peder Mortensen, Pernille Bangsgaard. Institution: Statens Naturhistoriske Museum.

Tingbjerggård Vest, Denmark. Samples: NEO957. Key contact: Anders Fischer/Niels Lynnerup. Institution: Sealand Archaeology/ University of Copenhagen.

Tissøe, Denmark. Samples: NEO942. Key contact: Anders Fischer/Niels Lynnerup. Institution: Sealand Archaeology/University of Copenhagen.

Toftum, Skanderborg, Denmark. Samples: NEO870, NEO872, NEO875, NEO876. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Toral III, Spain. Samples: NEO648, NEO649, NEO650, NEO653. Key contact: Igor Gutiérrez Zugasti. Institution: Universidad de Cantabria.

Troldebjerg, Denmark. Samples: NEO934. Key contact: Anders Fischer/Otto Uldum. Institution: Langelands Museum, Rudkøbing.

Tudse Hage, Denmark. Samples: NEO932. Key contact: Anders Fischer. Institution: Museum Vestsjælland, Museum Vestsjælland.

Tybrind Vig, Denmark. Samples: NEO683. Key contact: Anders Fischer. Institution: Sealand Archaeology/University of Copenhagen.

Tysmosen II, Denmark. Samples: NEO790. Key contact: Lasse Sørensen, Anders Fischer, Poul Otto Nielsen. Institution: National Museum of Denmark.

Ural River Beach, Russia. Samples: NEO100. Key contact: Mikhail Sablin. Institution: Zoological Institute of the Russian Academy of Sciences.

Ust'-Isha, Russia. Samples: NEO067. Key contact: Vyacheslav Moiseyev. Institution: Peter the Great Museum of Anthropology and ethnography (Kunstkamera).

Vængesø II, Denmark. Samples: NEO003. Key contact: Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Vanløse Mose II, Denmark. Samples: NEO599. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Vasagård, Denmark. Samples: NEO815. Key contact: Lasse Sørensen. Institution: National Museum of Denmark.

Vasilevka-I, Ukraine. Samples: NEO492, NEO494, NEO496, NEO497, NEO501, NEO545, NEO549, NEO550. Key contact: Alexandra Buzhilova. Institution: Lomonosov Moscow State University.

Vasilevsky, Ukraine. Samples: NEO305. Key contact: Sergey Vasilyev. Institution: Inst. of Ethnography and Anthropology, Moscow.

Vasilyevskiy kordon 17, Russia. Samples: NEO160, NEO162, NEO163, NEO164, NEO166, NEO167, NEO168, NEO170, NEO171. Key contact: Sergey Vasilyev. Institution: Inst. of Ethnography and Anthropology, Moscow.

Vattenledningen, Sweden. Samples: NEO052. Key contact: Catharina Ödman / Yvonne Magnusson. Institution: Malmö museum.

Vedbæk Boldbaner, Denmark. Samples: NEO600. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Vedrovice, Czech Republic. Samples: NEO128. Key contact: Vaclav Smrcka. Institution: Institute for History of Medicine and Foreign Languages of the First Faculty of Medicine, Charles University.

Vengerovo-2, Russia. Samples: NEO907, NEO910, NEO911, NEO912, NEO915, NEO916, NEO917, NEO918, NEO921, NEO922, NEO923. Key contact: V. I. Molodin. Institution: Institute of Archaeology and Ethnography, Russian Academy of Science, Siberian Branch.

Vibygårds Mose, Denmark. Samples: NEO935. Key contact: Anders Fischer / Per Lotz. Institution: Værløse Museum.

Viksø Mose, Denmark. Samples: NEO601. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/ University of Copenhagen.

Vittrup Mose, Denmark. Samples: NEO033. Key contact: Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Vlasac, Serbia. Samples: NEO655, NEO657, NEO658, NEO677. Key contact: Dušan Borić. Institution: Columbia university.

Volnensky, Ukraine. Samples: NEO300, NEO302, NEO304. Key contact: Sergey Vasilyev. Institution: Inst. of Ethnography and Anthropology, Moscow.

Voloshskoe, Ukraine. Samples: NEO522, NEO527. Key contact: Alexandra Buzhilova. Institution: Lomonosov Moscow State University.

Vovnigi, Ukraine. Samples: NEO498, NEO502, NEO503, NEO507, NEO508, NEO514, NEO524, NEO528, NEO529, NEO551, NEO552, NEO553. Key contact: Alexandra Buzhilova. Institution: Lomonosov Moscow State University.

Zamostje 2, Russia. Samples: NEO087, NEO088. Key contact: Alisa Zubova/Vyacheslav Moiseyev. Institution: Peter the Great Museum of Anthropology and ethnography (Kunstkamera).

Zhindo, Russia. Samples: NEO115, NEO117. Key contact: Sergey Vasilyev. Institution: Inst. of Ethnography and Anthropology, Moscow.
Sjیدری 10, Kazakhstan. Samples: NEO904. Key contact: Victor Merc/Emma Usmanova. Institution: Karaganda State University.

Skateholm I, Sweden. Samples: NEO679. Key contact: Douglas Price / Lars Larsson, Institution: Lund university.

Słonowice, Poland. Samples: NEO640, NEO641. Key contact: Piotr Wlodarczak. Institution: Polish Academy of Sciences. Institution: Instytut Archeologii i Etnologii.

Sludegård Sømose, Denmark. Samples: NEO933. Key contact: Anders Fischer / Jesper Hansen. Institution: Odense Bys museer.

Sølager, Denmark. Samples: NEO598. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/ University of Copenhagen.

Sope-2, Estonia. Samples: NEO306. Key contact: Sergey Vasilyev. Institution: Inst. of Ethnography and Anthropology, Moscow.

Sosnovyiy Myis, Russia. Samples: NEO841, NEO843. Key contact: Vyacheslav Moiseyev. Institution: Peter the Great Museum of Anthropology and ethnography (Kunstkamera).

Stenderup Hage, Denmark. Samples: NEO943. Key contact: Anders Fischer/Niels Lynnerup.

Storelyng. Institution: Øgårde boat III), Denmark. Samples: NEO597. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike.

Institution: Sealand Archaeology/University of Copenhagen.

Storelyng (Østrup Homo II), Denmark. Samples: NEO602. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution:

Sealand Archaeology/University of Copenhagen.

Strøby Egede, Denmark. Samples: NEO092. Key contact: Niels Lynnerup. Institution: Department of Forensic Medicine, Copenhagen university.

Strøby Grøftemark, Denmark. Samples: NEO091. Key contact: Niels Lynnerup, Anders Fischer. Institution: Department of Forensic Medicine, Copenhagen university.

Strøby Ladeplads, Denmark. Samples: NEO093. Key contact: Niels Lynnerup. Institution: Department of Forensic Medicine, Copenhagen university.

Svinninge Vejle, Denmark. Samples: NEO898. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Tepe Guran, Iran. Samples: NEO816, NEO817, NEO819. Key contact: Peder Mortensen, Pernille Bangsgaard. Institution: Statens Naturhistoriske Museum.

Tingbjerggård Vest, Denmark. Samples: NEO957. Key contact: Anders Fischer/Niels Lynnerup. Institution: Sealand Archaeology/ University of Copenhagen.

Tissøe, Denmark. Samples: NEO942. Key contact: Anders Fischer/Niels Lynnerup. Institution: Sealand Archaeology/University of Copenhagen.

Toftum, Skanderborg, Denmark. Samples: NEO870, NEO872, NEO875, NEO876. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Toral III, Spain. Samples: NEO648, NEO649, NEO650, NEO653. Key contact: Igor Gutiérrez Zugasti. Institution: Universidad de Cantabria.

Troldebjerg, Denmark. Samples: NEO934. Key contact: Anders Fischer/Otto Uldum. Institution: Langelands Museum, Rudkøbing.

Tudse Hage, Denmark. Samples: NEO932. Key contact: Anders Fischer. Institution: Museum Vestsjælland, Museum Vestsjælland.

Tybrind Vig, Denmark. Samples: NEO683. Key contact: Anders Fischer. Institution: Sealand Archaeology/University of Copenhagen.

Tysmosen II, Denmark. Samples: NEO790. Key contact: Lasse Sørensen, Anders Fischer, Poul Otto Nielsen. Institution: National Museum of Denmark.

Ural River Beach, Russia. Samples: NEO100. Key contact: Mikhail Sablin. Institution: Zoological Institute of the Russian Academy of Sciences.

Ust'-Isha, Russia. Samples: NEO067. Key contact: Vyacheslav Moiseyev. Institution: Peter the Great Museum of Anthropology and ethnography (Kunstkamera).

Vængesø II, Denmark. Samples: NEO003. Key contact: Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Vanløse Mose II, Denmark. Samples: NEO599. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Vasagård, Denmark. Samples: NEO815. Key contact: Lasse Sørensen. Institution: National Museum of Denmark.

Vasilevka-I, Ukraine. Samples: NEO492, NEO494, NEO496, NEO497, NEO501, NEO545, NEO549, NEO550. Key contact: Alexandra Buzhilova. Institution: Lomonosov Moscow State University.

Vasilevsky, Ukraine. Samples: NEO305. Key contact: Sergey Vasilyev. Institution: Inst. of Ethnography and Anthropology, Moscow.

Vasilyevskiy kordon 17, Russia. Samples: NEO160, NEO162, NEO163, NEO164, NEO166, NEO167, NEO168, NEO170, NEO171. Key contact: Sergey Vasilyev. Institution: Inst. of Ethnography and Anthropology, Moscow.

Vattenledningen, Sweden. Samples: NEO052. Key contact: Catharina Ödman / Yvonne Magnusson. Institution: Malmö museum.

Vedbæk Boldbaner, Denmark. Samples: NEO600. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Vedrovice, Czech Republic. Samples: NEO128. Key contact: Vaclav Smrcka. Institution: Institute for History of Medicine and Foreign Languages of the First Faculty of Medicine, Charles University.

Vengerovo-2, Russia. Samples: NEO907, NEO910, NEO911, NEO912, NEO915, NEO916, NEO917, NEO918, NEO921, NEO922, NEO923. Key contact: V. I. Molodin. Institution: Institute of Archaeology and Ethnography, Russian Academy of Science, Siberian Branch.

Vibygårds Mose, Denmark. Samples: NEO935. Key contact: Anders Fischer / Per Lotz. Institution: Værløse Museum.

Viksø Mose, Denmark. Samples: NEO601. Key contact: Niels Lynnerup, Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/ University of Copenhagen.

Vittrup Mose, Denmark. Samples: NEO033. Key contact: Anders Fischer, Pia Bennike. Institution: Sealand Archaeology/University of Copenhagen.

Vlasac, Serbia. Samples: NEO655, NEO657, NEO658, NEO677. Key contact: Dušan Borić. Institution: Columbia university.

Volnensky, Ukraine. Samples: NEO300, NEO302, NEO304. Key contact: Sergey Vasilyev. Institution: Inst. of Ethnography and Anthropology, Moscow.

Voloshskoe, Ukraine. Samples: NEO522, NEO527. Key contact: Alexandra Buzhilova. Institution: Lomonosov Moscow State University.

Vovnigi, Ukraine. Samples: NEO498, NEO502, NEO503, NEO507, NEO508, NEO514, NEO524, NEO528, NEO529, NEO551, NEO552, NEO553. Key contact: Alexandra Buzhilova. Institution: Lomonosov Moscow State University.

Zamostje 2, Russia. Samples: NEO087, NEO088. Key contact: Alisa Zubova/Vyacheslav Moiseyev. Institution: Peter the Great Museum of Anthropology and ethnography (Kunstkamera).

Zhindo, Russia. Samples: NEO115, NEO117. Key contact: Sergey Vasilyev. Institution: Inst. of Ethnography and Anthropology, Moscow.
Zvejnieki, Latvia. Samples: NEO307. Key contact: Sergey Vasilyev. Institution: Inst. of Ethnography and Anthropology, Moscow.

**Specimen deposition** All specimens studied are available upon direct contact/request to the archaeologists, curators or officials responsible for their curation at the organisation where they are held.

**Dating methods** 272 novel radiocarbon dates were generated at the 14CHRONO laboratory, Queen's University Belfast (242 samples), at the Oxford Radiocarbon Accelerator Unit (ORAU) laboratory (24 samples) and at the Keck-CCAMS Group, Irvine, California, USA (6 samples).

<figure><img src="image_61.png" /></figure>

Tick this box to confirm that the raw and calibrated dates are available in the paper or in Supplementary Information.

**Ethics oversight** No ethical oversight was required - all specimens were sampled following protocols designed to minimise invasive impact of sampling, with the full agreement of curators or archaeologists responsible for collections where specimens are held. Our use of shotgun whole genome data also reduces the need to resequence samples in the future (in contrast to targeting specific loci).

Note that full information on the approval of the study protocol must also be provided in the manuscript.
Zvejnieki, Latvia. Samples: NEO307. Key contact: Sergey Vasilyev. Institution: Inst. of Ethnography and Anthropology, Moscow.

**Specimen deposition** All specimens studied are available upon direct contact/request to the archaeologists, curators or officials responsible for their curation at the organisation where they are held.

**Dating methods** 272 novel radiocarbon dates were generated at the 14CHRONO laboratory, Queen's University Belfast (242 samples), at the Oxford Radiocarbon Accelerator Unit (ORAU) laboratory (24 samples) and at the Keck-CCAMS Group, Irvine, California, USA (6 samples).

<figure><img src="image_63.png" /></figure>

Tick this box to confirm that the raw and calibrated dates are available in the paper or in Supplementary Information.

**Ethics oversight** No ethical oversight was required - all specimens were sampled following protocols designed to minimise invasive impact of sampling, with the full agreement of curators or archaeologists responsible for collections where specimens are held. Our use of shotgun whole genome data also reduces the need to resequence samples in the future (in contrast to targeting specific loci).

Note that full information on the approval of the study protocol must also be provided in the manuscript.