Skip to main content

Thank you for visiting You are using a browser version with limited support for CSS. To obtain the best experience, we recommend you use a more up to date browser (or turn off compatibility mode in Internet Explorer). In the meantime, to ensure continued support, we are displaying the site without styles and JavaScript.

  • Article
  • Published:

Paths and timings of the peopling of Polynesia inferred from genomic networks


Polynesia was settled in a series of extraordinary voyages across an ocean spanning one third of the Earth1, but the sequences of islands settled remain unknown and their timings disputed. Currently, several centuries separate the dates suggested by different archaeological surveys2,3,4. Here, using genome-wide data from merely 430 modern individuals from 21 key Pacific island populations and novel ancestry-specific computational analyses, we unravel the detailed genetic history of this vast, dispersed island network. Our reconstruction of the branching Polynesian migration sequence reveals a serial founder expansion, characterized by directional loss of variants, that originated in Samoa and spread first through the Cook Islands (Rarotonga), then to the Society (Tōtaiete mā) Islands (11th century), the western Austral (Tuha’a Pae) Islands and Tuāmotu Archipelago (12th century), and finally to the widely separated, but genetically connected, megalithic statue-building cultures of the Marquesas (Te Henua ‘Enana) Islands in the north, Raivavae in the south, and Easter Island (Rapa Nui), the easternmost of the Polynesian islands, settled in approximately ad 1200 via Mangareva.

This is a preview of subscription content, access via your institution

Access options

Buy this article

Prices may be subject to local taxes which are calculated during checkout

Fig. 1: Dimensionality reduction of genetic variation in Pacific Islanders.
Fig. 2: Serial bottlenecks and relatedness define the settlement sequence and timings for the Polynesian Islands.

Similar content being viewed by others

Data availability

The samples for this project were collected by the University of Oxford, Stanford University and the University of Chile as part of different studies. SNP data for all newly genotyped individuals are available through a data access agreement to respect the privacy of the participants for the transfer of genetic data from the European Genome-Phenome Archive under accession number EGAS00001005362.

Code availability

All new techniques described in Methods are available from and all existing software packages and versions used are noted in Methods.


  1. Low, S. Hawaiki Rising: Hōkūle‘a, Nainoa Thompson, and the Hawaiian Renaissance (Univ. of Hawaii Press, 2019).

  2. Kirch, P. V. On the Road of the Winds (Univ. of California Press, 2017).

  3. Mulrooney, M. A., Bickler, S. H., Allen, M. S. & Ladefoged, T. N. High-precision dating of colonization and settlement in East Polynesia. Proc. Natl Acad. Sci. USA 108, E192–E194 (2011).

    Article  ADS  CAS  PubMed  PubMed Central  Google Scholar 

  4. Schmid, M. M. E. et al. How 14C dates on wood charcoal increase precision when dating colonization: the examples of Iceland and Polynesia. Quat. Geochronol. 48, 64–71 (2018).

    Article  Google Scholar 

  5. Kahō‘āli‘i Keauokalani, K. Kepelino’s traditions of Hawaii. Bernice P. Bishop Museum Bulletin 206 (1932).

  6. Cook, J. The Journals of Captain James Cook on his Voyages of Discovery (Cambridge Univ. Press, 1955).

  7. Kirch, P. V. & Green, R. C. Hawaiki, Ancestral Polynesia (Cambridge Univ. Press, 2001).

  8. Minster, R. L. et al. A thrifty variant in CREBRF strongly influences body mass index in Samoans. Nat. Genet. 48, 1049–1054 (2016).

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  9. Gray, R. D., Drummond, A. J. & Greenhill, S. J. Language phylogenies reveal expansion pulses and pauses in Pacific settlement. Science 323, 479–483 (2009).

    Article  ADS  CAS  PubMed  Google Scholar 

  10. Walworth, M. Eastern Polynesian: the linguistic evidence revisited. Ocean. Linguist. 53, 256–272 (2014).

    Article  Google Scholar 

  11. Martinsson-Wallin, H., Wallin, P. & Anderson, A. Chronogeographic variation in initial East Polynesian construction of monumental ceremonial sites. J. Island Coastal Archaeol. 8, 405–421 (2013).

    Article  Google Scholar 

  12. Wilmshurst, J. M., Hunt, T. L., Lipo, C. P. & Anderson, A. J. High-precision radiocarbon dating shows recent and rapid initial human colonization of East Polynesia. Proc. Natl Acad. Sci. USA 108, 1815–1820 (2011).

    Article  ADS  CAS  PubMed  Google Scholar 

  13. Spriggs, M. & Anderson, A. Late colonization of east Polynesia. Antiquity 67, 200–217 (1993).

    Article  Google Scholar 

  14. Hill, A. V. S. et al. Polynesian origins and affinities: globin gene variants in eastern Polynesia. Am. J. Hum. Genet. 40, 453–463 (1987).

    CAS  PubMed  PubMed Central  Google Scholar 

  15. Wollstein, A. et al. Demographic history of Oceania inferred from genome-wide data. Curr. Biol. 20, 1983–1992 (2010).

    Article  CAS  PubMed  Google Scholar 

  16. Hudjashov, G. et al. Investigating the origins of eastern Polynesians using genome-wide data from the Leeward Society Isles. Sci. Rep. 8, 1823 (2018).

    Article  ADS  PubMed  PubMed Central  CAS  Google Scholar 

  17. Skoglund, P. et al. Genomic insights into the peopling of the Southwest Pacific. Nature 538, 510–513 (2016).

    Article  ADS  PubMed  PubMed Central  CAS  Google Scholar 

  18. Posth, C. et al. Language continuity despite population replacement in Remote Oceania. Nat. Ecol. Evol. 2, 731–740 (2018).

    Article  PubMed  PubMed Central  Google Scholar 

  19. McColl, H. et al. The prehistoric peopling of Southeast Asia. Science 361, 88–92 (2018).

    Article  ADS  CAS  PubMed  Google Scholar 

  20. Emory, K. P. The Tuamotuan creation charts by Paiore. J. Polynesian Soc. 48, 1–29 (1939).

    Google Scholar 

  21. Hunt, T. & Lipo, C. The Statues that Walked (Free Press, 2011).

  22. Whyte, A. L. H., Marshall, S. J. & Chambers, G. K. Human evolution in Polynesia. Hum. Biol. 77, 157–177 (2005).

    Article  PubMed  Google Scholar 

  23. Duncan, R. P., Boyer, A. G. & Blackburn, T. M. Magnitude and variation of prehistoric bird extinctions in the Pacific. Proc. Natl Acad. Sci. USA 110, 6436–6441 (2013).

    Article  ADS  CAS  PubMed  PubMed Central  Google Scholar 

  24. Steadman, D. W. Extinction and Biogeography of Tropical Pacific Birds (Univ. of Chicago Press, 2006).

  25. Kirch, P. V. et al. Human ecodynamics in the Mangareva Islands: a stratified sequence from Nenega-Iti Rock Shelter (site AGA-3, Agakauitai Island). Archaeol. Oceania 50, 23–42 (2015).

    Article  Google Scholar 

  26. Rolett, B. V. Voyaging and interaction in ancient East Polynesia. Asian Perspect. 41, 182–194 (2002).

    Article  Google Scholar 

  27. Handy, E. S. C. The Native Culture in the Marquesas (The Bishop Museum, 1923).

  28. Weisler, M. I. et al. Cook Island artifact geochemistry demonstrates spatial and temporal extent of pre-European interarchipelago voyaging in East Polynesia. Proc. Natl Acad. Sci. USA 113, 8150–8155 (2016).

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  29. Collerson, K. D. & Weisler, M. I. Stone adze compositions and the extent of ancient Polynesian voyaging and trade. Science 317, 1907–1911 (2007).

    Article  ADS  CAS  PubMed  Google Scholar 

  30. Slatkin, M. & Excoffier, L. Serial founder effects during range expansion: a spatial analog of genetic drift. Genetics 191, 171–181 (2012).

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  31. Stephens, M. & Novembre, J. Interpreting principal component analyses of spatial population genetic variation. Nat. Genet. 40, 646–649 (2008).

    Article  PubMed  PubMed Central  CAS  Google Scholar 

  32. Wang, C. et al. Comparing spatial maps of human population-genetic variation using procrustes analysis. Stat. Appl. Genet. Mol. Biol. 9, 13 (2010).

  33. Ioannidis, A. G. et al. Native American gene flow into Polynesia predating Easter Island settlement. Nature 583, 572–577 (2020).

    Article  ADS  CAS  PubMed  PubMed Central  Google Scholar 

  34. Novembre, J. et al. Genes mirror geography within Europe. Nature 456, 98–101 (2008).

    Article  ADS  CAS  PubMed  PubMed Central  Google Scholar 

  35. Nei, M. & Kumar, S. Molecular Evolution and Phylogenetics (Oxford Univ. Press, 2000).

  36. Patterson, N. et al. Ancient admixture in human history. Genetics 192, 1065–1093 (2012).

    Article  PubMed  PubMed Central  Google Scholar 

  37. Peter, B. M. & Slatkin, M. Detecting range expansions from genetic data. Evolution 67, 3274–3289 (2013).

    Article  PubMed  PubMed Central  Google Scholar 

  38. Zhan, S. et al. The genetics of monarch butterfly migration and warning colouration. Nature 514, 317–321 (2014).

    Article  ADS  CAS  PubMed  PubMed Central  Google Scholar 

  39. Lipson, M. et al. Efficient moment-based inference of admixture parameters and sources of gene flow. Mol. Biol. Evol. 30, 1788–1802 (2013).

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  40. Pickrell, J. K. & Pritchard, J. K. Inference of population splits and mixtures from genome-wide allele frequency data. PLoS Genet. 8, e1002967 (2012).

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  41. Leppälä, K., Nielsen, S. V. & Mailund, T. admixturegraph: an R package for admixture graph manipulation and fitting. Bioinformatics 33, 1738–1740 (2017).

    Article  PubMed  PubMed Central  CAS  Google Scholar 

  42. Anderson, A. J., Conte, E., Smith, I. & Szabo, K. New excavations at Fa’ahia (Huahine, Society Islands) and chronologies of central East Polynesian colonization. J. Pac. Arch. 10, 1–14 (2019).

    Google Scholar 

  43. Hunt, T. L. & Lipo, C. P. Evidence for a shorter chronology on Rapa Nui (Easter Island). J. Island Coast. Archaeol. (2008).

  44. Mulrooney, M. A. An island-wide assessment of the chronology of settlement and land use on Rapa Nui (Easter Island) based on radiocarbon data. J. Archaeol. Sci. 40, 4377–4399 (2013).

    Article  Google Scholar 

  45. Pirazzoli, P. A. & Montaggioni, L. F. Late Holocene sea-level changes in the northwest Tuamotu islands, French Polynesia. Quat. Res. 25, 350–368 (1986).

    Article  Google Scholar 

  46. Di Piazza, A., Di Piazza, P. & Pearthree, E. Sailing virtual canoes across Oceania: revisiting island accessibility. J. Archaeol. Sci. 34, 1219–1225 (2007).

    Article  Google Scholar 

  47. Walworth, M. The Language of Rapa Iti (Univ. Hawaii, 2015).

  48. Dickinson, W. Pacific atoll living: how long already and until when. Geol. Soc. Am. Today 19, 4–10 (2009).

    Google Scholar 

  49. Fischer, S. R. Mangarevan doublets: preliminary evidence for proto-southeastern Polynesian. Ocean. Linguist. 40, 112–124 (2001).

    Article  Google Scholar 

  50. Flenley, J. & Bahn, P. The Enigmas of Easter Island (Oxford Univ. Press, 2003).

  51. Buck Te Rangi Hīroa, P. H. Vikings of the Sunrise (J. B. Lippincott, 1938).

  52. Belbin, G. M. et al. Toward a fine-scale population health monitoring system. Cell 184, 2068–2083.e11 (2021).

    Article  CAS  PubMed  Google Scholar 

  53. Claw, K. G. et al. A framework for enhancing ethical genomic research with Indigenous communities. Nat. Commun. 9, 2957 (2018).

    Article  ADS  PubMed  PubMed Central  CAS  Google Scholar 

  54. Chang, C. C. et al. Second-generation PLINK: rising to the challenge of larger and richer datasets. GigaScience 4, 7 (2015).

    Article  PubMed  PubMed Central  CAS  Google Scholar 

  55. Tyner, C. et al. The UCSC Genome Browser database: 2017 update. Nucleic Acids Res. 45, D626–D634 (2017).

    CAS  PubMed  Google Scholar 

  56. Thornton, T. et al. Estimating kinship in admixed populations. Am. J. Hum. Genet. 91, 122–138 (2012).

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  57. The 1000 Genomes Project Consortium. A map of human genome variation from population-scale sequencing. Nature 467, 1061–1073 (2010).

    Article  PubMed Central  CAS  Google Scholar 

  58. Patterson, N., Price, A. L. & Reich, D. Population structure and eigenanalysis. PLoS Genet. 2, e190 (2006).

    Article  PubMed  PubMed Central  CAS  Google Scholar 

  59. Wickham, H. ggplot2 (Springer, 2016).

  60. R Core Team. R: a Language and Environment for Statistical Computing (2017).

  61. Alexander, D. H. & Lange, K. Enhancements to the ADMIXTURE algorithm for individual ancestry estimation. BMC Bioinf. 12, 246 (2011).

    Article  Google Scholar 

  62. Holmes, S. & Huber, W. Modern Statistics for Modern Biology (Cambridge Univ. Press, 2019).

  63. Maples, B. K., Gravel, S., Kenny, E. E. & Bustamante, C. D. RFMix: A discriminative modeling approach for rapid and robust local-ancestry inference. Am. J. Hum. Genet. 93, 278–288 (2013).

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  64. O’Connell, J. et al. A general approach for haplotype phasing across the full spectrum of relatedness. PLoS Genet. 10, e1004234 (2014).

    Article  PubMed  PubMed Central  CAS  Google Scholar 

  65. D’Arcy, P. The Chinese Pacifics: a brief historical review. J. Pacific Hist. 49, 396–420 (2014).

    Article  Google Scholar 

  66. Browning, S. R. et al. Ancestry-specific recent effective population size in the Americas. PLoS Genet. 14, e1007385 (2018).

    Article  PubMed  PubMed Central  CAS  Google Scholar 

  67. Schroeder, H. et al. Origins and genetic legacies of the Caribbean Taino. Proc. Natl Acad. Sci. USA 115, 2341–2346 (2018).

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  68. Moreno-Estrada, A. et al. The genetics of Mexico recapitulates Native American substructure and affects biomedical traits. Science 344, 1280–1285 (2014).

    Article  ADS  CAS  PubMed  PubMed Central  Google Scholar 

  69. Mazumder, R., Hastie, T. & Tibshirani, R. Spectral regularization algorithms for learning large incomplete matrices. J. Mach. Learn. Res. 11, 2287–2322 (2010).

    MathSciNet  PubMed  PubMed Central  MATH  Google Scholar 

  70. Reich, D. et al. Reconstructing Native American population history. Nature 488, 370–374 (2012).

    Article  ADS  CAS  PubMed  PubMed Central  Google Scholar 

  71. Skoglund, P. et al. Origins and genetic legacy of Neolithic farmers and hunter-gatherers in Europe. Science 336, 466–469 (2012).

    Article  ADS  CAS  PubMed  Google Scholar 

  72. Moreno-Estrada, A. et al. Reconstructing the population genetic history of the Caribbean. PLoS Genet. 9, e1003925 (2013).

    Article  PubMed  PubMed Central  CAS  Google Scholar 

  73. Nguyen, L. H. & Holmes, S. Ten quick tips for effective dimensionality reduction. PLoS Comp. Biol. 15, e1006907 (2019).

    Article  ADS  CAS  Google Scholar 

  74. Maaten, L. V. D. & Hinton, G. Visualizing data using t-SNE. J. Mach. Learn. Res. 9, 2579–2605 (2008).

    MATH  Google Scholar 

  75. Van Der Maaten, L. Accelerating t-SNE using tree-based algorithms. J. Mach. Learn. Res. 15, 3221–3245 (2014).

  76. McInnes, L., Healy, J. & Melville, J. UMAP: Uniform manifold approximation and projection for dimension reduction. Preprint at (2018).

  77. Diaz-Papkovich, A., Anderson-Trocmé, L., Ben-Eghan, C. & Gravel, S. UMAP reveals cryptic population structure and phenotype heterogeneity in large genomic cohorts. PLoS Genet. 15, e1008432 (2019).

    Article  PubMed  PubMed Central  CAS  Google Scholar 

  78. Hastie, T., Tibshirani, R. & Friedman, J. The Elements of Statistical Learning (Springer, 2009).

  79. Wittek, P., Gao, S. C., Lim, I. S. & Zhao, L. Somoclu: an efficient parallel library for self-organizing maps. J. Stat. Softw. 78, (2017).

  80. Nguyen, L. H. & Holmes, S. Bayesian unidimensional scaling for visualizing uncertainty in high dimensional datasets with latent ordering of observations. BMC Bioinf. 18, 65–79 (2017).

    Article  Google Scholar 

  81. Peter, B. M. & Slatkin, M. The effective founder effect in a spatially expanding population. Evolution 69, 721–734 (2015).

    Article  PubMed  PubMed Central  Google Scholar 

  82. Pugach, I. et al. The complex admixture history and recent southern origins of Siberian populations. Mol. Biol. Evol. 33, 1777–1795 (2016).

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  83. Takahata, N. & Nei, M. Gene genealogy and variance of interpopulational nucleotide differences. Genetics 110, 325–344 (1985).

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  84. Peter, B. M. Admixture, population structure, and F-statistics. Genetics 202, 1485–1501 (2016).

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  85. Nei, M. Molecular Evolutionary Genetics (Columbia Univ. Press, 1987).

  86. Patterson, N. et al. Reconstructing Indian population history. Nature 461, 489–494 (2009).

    Article  ADS  PubMed  PubMed Central  CAS  Google Scholar 

  87. Bhatia, G., Patterson, N., Sankararaman, S. & Price, A. L. Estimating and interpreting FST: the impact of rare variants. Genome Res. 23, 1514–1521 (2013).

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  88. Nei, M. & Roychoudhury, A. K. Sampling variances of heterozygosity and genetic distance. Genetics 76, 379–390 (1974).

    Article  MathSciNet  CAS  PubMed  PubMed Central  Google Scholar 

  89. Davison, A. C. & Hinkley, D. V. Bootstrap Methods and Their Application (Cambridge Univ. Press, 1997).

  90. Chu, Y. J. & Lui, T. H. On the shortest arborescence of a directed graph. Science Sinica 14, 1396–1400 (1965).

  91. Edmonds, J. Optimum branchings. J. Res. Natl. Bur. Stand. 71B, 233–240 (1967).

    Article  MathSciNet  MATH  Google Scholar 

  92. Ceballos, F. C., Joshi, P. K., Clark, D. W., Ramsay, M. & Wilson, J. F. Runs of homozygosity: windows into population history and trait architecture. Nat. Rev. Genet. 19, 220–234 (2018).

    Article  CAS  PubMed  Google Scholar 

  93. Huff, C. D. et al. Maximum-likelihood estimation of recent shared ancestry (ERSA). Genome Res. 21, 768–774 (2011).

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  94. Baharian, S. et al. The Great Migration and African-American genomic diversity. PLoS Genet. 12, e1006059 (2016).

    Article  PubMed  PubMed Central  CAS  Google Scholar 

  95. Gusev, A. et al. Whole population, genome-wide mapping of hidden relatedness. Genome Res. 19, 318–326 (2009).

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  96. Efron, B. Bootstrap methods: another look at the jackknife. Ann. Stat. 7, 1–26 (1979).

    Article  MathSciNet  MATH  Google Scholar 

  97. Botigué, L. R. et al. Gene flow from North Africa contributes to differential human genetic diversity in southern Europe. Proc. Natl Acad. Sci. USA 110, 11791–11796 (2013).

    Article  ADS  PubMed  PubMed Central  Google Scholar 

  98. Atzmon, G. et al. Abraham’s children in the genome era: major Jewish diaspora populations comprise distinct genetic clusters with shared Middle Eastern ancestry. Am. J. Hum. Genet. 86, 850–859 (2010).

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  99. Jobling, M., Hurles, M. & Tyler-Smith, C. Human Evolutionary Genetics (Garland Science, 2013).

  100. Liang, M. & Nielsen, R. The lengths of admixture tracts. Genetics 197, 953–967 (2014).

    Article  PubMed  PubMed Central  Google Scholar 

  101. Palamara, P. F., Lencz, T., Darvasi, A. & Pe'er, I. Length distributions of identity by descent reveal fine-scale demographic history. Am. J. Hum. Genet. 91, 809–822 (2012).

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  102. Ralph, P. & Coop, G. The geography of recent genetic ancestry across Europe. PLoS Biol. 11, e1001555 (2013).

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  103. Deemer, W. L. Jr & Votaw, D. F. Jr Estimation of parameters of truncated or censored exponential distributions. Ann. Math. Stat. 26, 498–504 (1955).

  104. Hill, W. G. & White, I. M. S. Identification of pedigree relationship from genome sharing. G3 Genes Genom. Genet. 3, 1553–1571 (2013).

    Google Scholar 

  105. McVean, G. A. T. et al. The fine-scale structure of recombination rate variation in the human genome. Science 304, 581–584 (2004).

    Article  ADS  CAS  PubMed  Google Scholar 

  106. Makarenkov, V. & Lapointe, F.-J. A weighted least-squares approach for inferring phylogenies from incomplete distance matrices. Bioinformatics 20, 2113–2121 (2004).

    Article  CAS  PubMed  Google Scholar 

  107. Fehren-Schmitz, L. et al. Genetic ancestry of Rapanui before and after European contact. Curr. Biol. 27, 3209–3215 (2017).

    Article  CAS  PubMed  Google Scholar 

  108. Crowe, A. Pathway of the Birds (Univ. Hawai'i Press, 2018).

  109. Marck, J. C. Topics in Polynesian Language and Culture History (The Australian National Univ., 2000).

  110. Niespolo, E. M., Sharp, W. D. & Kirch, P. V. 230Th dating of coral abraders from stratified deposits at Tangatatau Rockshelter, Mangaia, Cook Islands: implications for building precise chronologies in Polynesia. J. Archaeol. Sci. 101, 21–33 (2019).

    Article  CAS  Google Scholar 

  111. Kirch, P. V. Tangatatau Rockshelter: The Evolution of an Eastern Polynesian Socio-ecosystem (Cotsen Institute of Archaeology Press, 2017).

  112. Sear, D. A. et al. Human settlement of East Polynesia earlier, incremental, and coincident with prolonged South Pacific drought. Proc. Natl Acad. Sci. USA 117, 8813–8819 (2020).

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  113. Kahn, J. G. & Sinoto, Y. Refining the Society Islands cultural sequence: colonisation phase and developmental phase coastal occupation on Mo'orea Island. J. Polynesian Soc. 126, 33 (2017).

    Article  Google Scholar 

  114. Conte, E. & Molle, G. Reinvestigating a key site for Polynesian prehistory: new results from the Hane dune site, Ua Huka (Marquesas). Archaeol. Oceania 49, 121–136 (2014).

    Article  Google Scholar 

  115. Allen, M. S. Marquesan colonisation chronologies and postcolonisation interaction: implications for Hawaiian origins and the ‘Marquesan homeland’ hypothesis. J. Pac. Arch. 5, 1–17 (2014).

    Google Scholar 

  116. Prebble, M. & Wilmshurst, J. M. Detecting the initial impact of humans and introduced species on island environments in Remote Oceania using palaeoecology. Biol Invasions 11, 1529–1556 (2009).

    Article  Google Scholar 

  117. Anderson, A., Kennett, D. J., Culleton, B. J. & Southon, J. in Taking the High Ground (eds. Anderson, A. & Kennett, D. J.) 288 (ANU Press, 2012).

  118. Kirch, P. V., Conte, E., Sharp, W. & Nickelsen, C. The Onemea Site (Taravai Island, Mangareva) and the human colonization of Southeastern Polynesia. Archaeol. Oceania 45, 66–79 (2010).

    Article  Google Scholar 

  119. Allen, M. S. & Steadman, D. W. Excavations at the Ureia site, Aitutaki, Cook Islands: preliminary results. Archaeol. Oceania 25, 24–37 (1990).

    Article  Google Scholar 

  120. Matisoo-Smith, E. et al. Patterns of prehistoric human mobility in Polynesia indicated by mtDNA from the Pacific rat. Proc. Natl Acad. Sci. USA 95, 15145–15150 (1998).

    Article  ADS  CAS  PubMed  PubMed Central  Google Scholar 

Download references


We thank the participants and volunteers who donated DNA samples for this study and the local community leaders and authorities who provided approval and logistics support in the various sampling sites, including T. Teariki (Cook Islands), S. Foliaki (Tonga), N. Tuuau (Samoa), H.-W. Peng (Taiwan), J. Roux (French Polynesia), E. Paoa (Rapa Nui), as well as J. Martinson, D. Weatherall and J. Clegg for their pioneering fieldwork in the region that ultimately led to this work. We thank H. Kane and S. Kahanamoku for critiques and suggestions on the text, as well as M. Spriggs for his comments on the manuscript concerning archaeological dates in Polynesia. We thank the Core Staff at the UCSF Institute for Human Genetics for contributing genotyping capacity, and the Stanford Center for Computational, Evolutionary and Human Genomics (CEHG) for supporting the initial stages of this project. We thank J. Cervantes for IT support and M. Ortega and G. Mireles for technical support at LANGEBIO’s Genomics Core Facility at CINVESTAV, Mexico. This work was supported by the George Rosenkranz Prize for Health Care Research in Developing Countries, Mexico’s CONACYT Basic Research Program (grant number CB-2015-01-251380), and the International Center for Genetic Engineering and Biotechnology (ICGEB, Italy) Grant CRP/MEX15-04_EC (each awarded to A.M.-E.); the Chilean funding programs FONDEF, FONDECYT and CONICYT (grants D10I1007, 1130303 and USA2013-0015, respectively); the National Institute for Health Research (NIHR) Oxford Biomedical Research Centre (BRC) and a Wellcome Trust Fellowship with reference 106289/Z/14/Z (A.J.M.); together with National Library of Medicine (NLM) training grant T15LM007033 and an American Society of Engineering Education NDSEG Fellowship (each awarded to A.G.I.). Views expressed are those of the author(s) and not necessarily those of the NIHR, the NHS, or the Department of Health.

Author information

Authors and Affiliations



A.M.-E. and A.G.I. conceived the study. A.G.I., A.J.M., C.D.B. and A.M.-E. provided overall project supervision and management. A.M.-E., A.G.I., E.H., K.S., A.J.M. and C.R.G. contributed to study design. A.S., C.E., S.H., E.G.B., C.D.B. and A.M.-E. carried out genotyping experiments and quality control. A.G.I., J.B.-P., C. B.-J., J.E.R.-R., C.D.Q.-C., J.R.H. and A.M.-E. analysed the data. A.G.I. developed the analytical methods. A.G.I., J.B.-P. and A.M.-E. interpreted the results. A.M.-E., K.S., E.H., A.V.S.H., J.F.M.-P., K.A., T.P., K.R., M.C.A.-A., A.S., A.S.M.S., C.E., S.H., E.G.B., R.A.V., M.M., A.J.M. and C.D.B. contributed to acquisition of the data. A.G.I. wrote the manuscript, and A.M.-E., E.H., K.F. and S.H.-C. provided feedback on the manuscript.

Corresponding authors

Correspondence to Alexander G. Ioannidis or Andrés Moreno-Estrada.

Ethics declarations

Competing interests

C.D.B. is a member of the scientific advisory boards for Liberty Biosecurity, Personalis, 23andMe Roots into the Future,, IdentifyGenomics, Genomelink and Etalon, and is a founder of CDB Consulting. C.R.G. owns stock in 23andMe and is member of the scientific advisory board for Encompass Bioscience.

Additional information

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

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

Extended data figures and tables

Extended Data Fig. 1 Comparison of genetic and geographic coordinates for European vs. Polynesian samples.

a, A principal component analysis of samples from Europe (15 from each nation) is shown to closely fit the geography of Europe. (See Extended Data Fig. 2 for a quantitative comparison.) b, c, A principal component analysis (b) of samples from Polynesia (with non-Polynesian ancestry masked) is shown not to match the vast geography of the Pacific (c), and instead splits out island groups one at a time, reflecting the founder effects that dominate the variance of these populations.

Extended Data Fig. 2 Permutation test for fit between genetic and geographic coordinates.

100,000 random permutations of the population labels were created for the European populations’ genetic data (blue, left) versus the Polynesian populations (orange, right). For the European populations, out of 100,000 random permutations of the population labels on the genetic PCA, none better fits the geography of Europe (after fitting using a Procrustes analysis32), than the correct labels, showing that the genetic coordinates of Europeans fit the geographic coordinates of Europe better than chance every time. However, for the Polynesian data 5% of the random permutations of the labels on the genetic PCA fit the geographic coordinates of the Pacific islands better (after fitting using Procrustes), showing that the genetic data in Polynesia does not fit Polynesia’s geography better than random chance. In the box and whiskers plots the mean and upper and lower quartiles of the rms error of the fits of the random permutations of population labels are indicated by horizontal lines. The fits of the actual population labels are indicated by asterisks.

Extended Data Fig. 3 Continuity between ancient and modern Polynesian island populations.

F3 statistics were computed between ancient Rapanui samples and the Polynesian component from modern samples from each island in our dataset (top)107. Indigenous Austronesian language speakers from Taiwan (the Atayal) were used as an outgroup. The ancient Rapanui were found to be the most similar genetically to the modern Rapanui, indicating genetic continuity. A similar comparison was performed between the only other ancient samples from an island in our study, Tonga (bottom)18. Again, the modern Tongans appear most similar genetically; however, all islands downstream from Tonga in our inferred settlement path also share the same amount of genetic drift with the ancient Tongan samples (to within one standard error), as they should, since they are all descendants of these ancient Tongan sample according to our settlement reconstruction.

Extended Data Fig. 4 Statistics used for settlement path inference.

All statistics are based on the Polynesian-specific aggregate SNP frequency vectors computed for each island from all sampled individuals. The number (n) of individuals used are given for each island in Supplementary Table 1. a, Directionality index (ψ), used to define sets of potential parent islands, plotted for each island relative to Samoa (equivalent to the top row of the matrix in Fig. 2b). b, Average number of pairwise differences (π), measuring genetic distance and used to select the closest of potential parents, plotted for each island relative to Rapa Nui. c, F3 statistic, used to find additional shared genetic drift, plotted for each island relative to Rapa Nui, with Taiwan as an outgroup. Standard errors in ac were determined by a block bootstrap analysis. d, Exponential decay constant (λ) for the Polynesian-specific IBD fragment length distributions between all pairs of individuals from Rapa Nui and each plotted island. The λ values can be used to calculate the number of generations elapsed since each pair of island populations were joined. Error bars show 95% confidence intervals of the maximum likelihood estimates determined analytically from the Fisher Information.

Extended Data Fig. 5 Settlement map with candidate intermediate islands added.

A reproduction of the map of Fig. 2a showing intermediate islands that are in the settlement path but not in our dataset that are possible candidates for explaining the additional shared drift observed in the corresponding colored settlement branches, that is, genetic drift shared between the child islands but not shared with the parent island. The additional shared drift of the Austral islands (Rimatara and Tubuai) with the Society islands (Tahiti) and Tuamotus (Palliser) beyond what they each share with their parental island (Rarotonga in the Cooks) could indicate that there exists a shared intermediate island in their settlement path that we do not have in our dataset, for instance Mangaia108. Geological analyses of ancient tools found on Mangaia (green) have shown that it served as a connection between the Cook islands and remote eastern Polynesia28, now uninhabited Nororotu (Maria Atoll) is also believed to have played a role as an intermediary island108. Traditional histories give Raiatea (pink) and its surrounding islands a role in the settling of remote eastern Polynesia108. Finally, linguistic studies have found connections between Marquesic languages (Marquesas and Mangareva) and the central Tuamotus (orange)109. North Marquesas, South Marquesas, and Mangareva share drift with one another beyond what they share with Palliser, the westernmost island group in the Tuamotus, which could indicate that these three populations shared a common settlement path eastward through some of the Tuamotu Archipelago before diverging. Another possible explanation for additional shared drift is the settlement of each child island from a common subpopulation within the parental island, such as from the same clan or village.

Extended Data Fig. 6 Effect of phasing errors on IBD dates.

IBD segments on the island of Rapa Nui were identified between all male X chromosomes. The log of the number of IBD segments (y axis) of a given genetic length (x axis) is plotted (orange; bottom left). The expected exponential decay of IBD segment lengths (linear semilog plot) is seen. The slope of this line (−0.161) is the exponential (decay) constant lambda. Since the X chromosome is perfectly phased in men, because it is haploid, the identification of these IBD segments is unaffected by errors introduced through phasing algorithms. To quantify the effect of such errors, synthetic-female individuals were constructed by combining two male X chromosomes to make a diploid pair and to erase the phase information by recording only the genotype. The unphased diploid genotypes so constructed were phased and IBD segments were again identified and plotted (green; bottom right). The difference between the exponential decay constant (−0.166) of these statistically phased genotypes and the previous one is seen to be minor (top panel), amounting to three per cent (3.01%), which corresponds to a difference of around 25 years for dates approximately eight hundred years ago (as in Polynesia). Uncertainty in the slope of the lines (equivalent to the uncertainty in the estimate exponential decay constant) is shaded.

Extended Data Fig. 7 Polynesian ancestry-specific shared drift ordination plot with principal curve.

A principal coordinate analysis (PCoA) projection of the pairwise shared drift distances (the Polynesian ancestry-specific outgroup-F3) between each Pacific island population using Taiwan as an outgroup (Supplementary Fig. 12). This PCoA projection uses only the pairwise distance matrix and is fully unsupervised; that is, it does not presuppose that Rapa Nui is a terminal island along some settlement path. Nevertheless, it shows the same ordering as in Supplementary Fig. 9, confirming that Rapa Nui is indeed the terminal island in our dataset along the longest drift path, and confirming the drift ordering along that path. For further confirmation, a principal curve was also fit to the full dimensional space (Supplementary Fig. 12) and then projected into the two-dimensional PCoA space for visualization. The orthogonal projections of each island onto the principal curve are shown as thinner grey lines. This fully unsupervised principal curve confirms the visually apparent path from Island Southeast Asia (Sumatra, far right) through Samoa, Fiji, Tonga and ending in Raivavae, Mangareva, and Rapa Nui (far left) in that order (cf. migration map in Fig. 2a). This projection of the high dimensional principal curve does not double back on itself, showing that the apparent ordering in this projection is consistent with the original high dimensional ordering. Note that this principal curve is able to fit only one settlement path (the principal one, that is, the longest drift path), which ends in Rapa Nui. Other settlement paths that branch away from this principal (longest) path appear simply as clusters projected onto the principal curve, since islands on those paths share no further drift with the principal path. That is, islands settled along secondary branching paths appear as clusters lying very close to one another along the principal curve. For example, Rapa Iti, which branches off from Rarotonga separately from the main settlement path (Fig. 2a), appears here as coincident with Rarotonga along the principal curve. The eigenvalue for PC1 over the sum of eigenvalues is .997 and for PC 2 is .002 (all eigenvalues are non-negative).

Extended Data Table 1 Archaeological and genetic inferred dates for first settlement

Supplementary information

Supplementary Information

This file contains supplementary text, Figs. 1–33, Tables 1–4, discussion and references.

Reporting Summary

Peer Review File

Rights and permissions

Reprints and permissions

About this article

Check for updates. Verify currency and authenticity via CrossMark

Cite this article

Ioannidis, A.G., Blanco-Portillo, J., Sandoval, K. et al. Paths and timings of the peopling of Polynesia inferred from genomic networks. Nature 597, 522–526 (2021).

Download citation

  • Received:

  • Accepted:

  • Published:

  • Issue Date:

  • DOI:

This article is cited by


By submitting a comment you agree to abide by our Terms and Community Guidelines. If you find something abusive or that does not comply with our terms or guidelines please flag it as inappropriate.


Quick links

Nature Briefing: Translational Research

Sign up for the Nature Briefing: Translational Research newsletter — top stories in biotechnology, drug discovery and pharma.

Get what matters in translational research, free to your inbox weekly. Sign up for Nature Briefing: Translational Research