Genetic evidence for anatomically modern humans (AMH) out of Africa before 75 thousand years ago (ka)1 and in island southeast Asia (ISEA) before 60 ka (93–61 ka)2 predates accepted archaeological records of occupation in the region3. Claims that AMH arrived in ISEA before 60 ka (ref. 4) have been supported only by equivocal5 or non-skeletal evidence6. AMH evidence from this period is rare and lacks robust chronologies owing to a lack of direct dating applications7, poor preservation and/or excavation strategies8 and questionable taxonomic identifications9. Lida Ajer is a Sumatran Pleistocene cave with a rich rainforest fauna associated with fossil human teeth7,10. The importance of the site is unclear owing to unsupported taxonomic identification of these fossils and uncertainties regarding the age of the deposit, therefore it is rarely considered in models of human dispersal. Here we reinvestigate Lida Ajer to identify the teeth confidently and establish a robust chronology using an integrated dating approach. Using enamel–dentine junction morphology, enamel thickness and comparative morphology, we show that the teeth are unequivocally AMH. Luminescence and uranium-series techniques applied to bone-bearing sediments and speleothems, and coupled uranium-series and electron spin resonance dating of mammalian teeth, place modern humans in Sumatra between 73 and 63 ka. This age is consistent with biostratigraphic estimations7, palaeoclimate and sea-level reconstructions, and genetic evidence for a pre-60 ka arrival of AMH into ISEA2. Lida Ajer represents, to our knowledge, the earliest evidence of rainforest occupation by AMH, and underscores the importance of reassessing the timing and environmental context of the dispersal of modern humans out of Africa.

Access optionsAccess options

Rent or Buy article

Get time limited or full article access on ReadCube.

from $8.99

All prices are NET prices.


  1. 1.

    et al. Genomic analyses inform on migration events during the peopling of Eurasia. Nature 538, 238–242 (2016)

  2. 2.

    et al. A revised timescale for human evolution based on ancient mitochondrial genomes. Curr. Biol. 23, 553–559 (2013)

  3. 3.

    The great arc of dispersal of modern humans: Africa to Australia. Quat. Int. 202, 2–13 (2009)

  4. 4.

    & The dispersal of Homo sapiens across southern Asia: how early, how often, how complex? Quat. Sci. Rev. 47, 15–22 (2012)

  5. 5.

    et al. The earliest unequivocally modern humans in southern China. Nature 526, 696–699 (2015)

  6. 6.

    et al. Climate, people and faunal succession on Java, Indonesia: evidence from Song Gupuh. J. Archaeol. Sci. 35, 1776–1789 (2008)

  7. 7.

    in The Encyclopedia of Quaternary Science (ed. ) 3232–3249 (Elsevier, 2007)

  8. 8.

    et al. Preface: research at Liang Bua, Flores, Indonesia. J. Hum. Evol. 57, 437–449 (2009)

  9. 9.

    , , & A review of the Pleistocene hominoid fauna of the Socialist Republic of Vietnam (excluding Hylobatidae). Anthropol. Pap. Am. Mus. Nat. Hist. 76, 1–24 (1995)

  10. 10.

    Voorlopig bericht omtrent het onderzoek naar de Pleistocene en Tertiaire vertebraten-fauna van Sumatra en Java, gedurende het jaar 1890. Nat. Tijdschr. Ned. Indië 51, 93–100 (1891)

  11. 11.

    The Pongo faunas from Java and Sumatra and their significance for biostratigraphical and paleo-ecological interpretations. Proc. Koninklijke Nederlandse Akademie Wetenschappen 86, 417–425 (1983)

  12. 12.

    Prehistoric teeth of man and of the orang-utan from central Sumatra, with notes on the fossil orang-utan from Java and Southern China. Zool. Meded. 29, 175–301 (1948)

  13. 13.

    The Systematics and Paleodemography of Fossil Orangutans. PhD Thesis, Univ. California (Davis, 1994)

  14. 14.

    & Palaeoecology of southeast Asian megafauna-bearing sites from the Pleistocene and a review of environmental changes in the region. J. Biogeogr. 37, 1432–1449 (2010)

  15. 15.

    et al. Age and biostratigraphic significance of the Punung rainforest fauna, East Java, Indonesia; implications for Pongo and Homo. J. Hum. Evol. 53, 709–717 (2007)

  16. 16.

    et al. The age of the 20 meter Solo River terrace, Java, Indonesia and the survival of Homo erectus in Asia. PLoS ONE 6, e21562 (2011)

  17. 17.

    et al. U-series and radiocarbon analyses of human and faunal remains from Wajak, Indonesia. J. Hum. Evol. 64, 356–365 (2013)

  18. 18.

    et al. Preliminary U-series and thermoluminescence dating of deposits in Liang Bua sub-chamber, Flores, Indonesia. J. Archaeol. Sci. 40, 148–155 (2013)

  19. 19.

    et al. Geochronology of cave deposits in Liang Bua and of adjacent river terraces in the Wae Racang valley, western Flores, Indonesi: a synthesis of age estimates for the type locality of Homo floresiensis. J. Hum. Evol. 57, 484–502 (2009)

  20. 20.

    & Environment, preferred habitats and potential refugia for Pleistocene Homo in Southeast Asia. C. R. Palevol 11, 203–211 (2012)

  21. 21.

    & (eds) Archaeological Investigations in the Niah Caves, Sarawak, 1954–2004 Monographs 2 (McDonald Institute for Archaeological Research, 2016)

  22. 22.

    et al. Additional evidence for early modern human morphological diversity in Southeast Asia at Tam Pa Ling, Laos. Quat. Int. (in the press) (2017)

  23. 23.

    A single southern exit of modern humans from Africa: before or after Toba? Quat. Int. 258, 88–99 (2012)

  24. 24.

    , , , & The Toba volcanic super-eruption, environmental change, and hominin occupation history in India over the last 140,000 years. Quat. Int. 258, 119–134 (2012)

  25. 25.

    et al. Rethinking the dispersal of Homo sapiens out of Africa. Evol. Anthropol. 24, 149–164 (2015)

  26. 26.

    et al. The influence of the ~73 ka Toba super-eruption on the ecosystems of northern Sumatra as recorded in marine core BAR94-25. Quat. Int. 258, 45–53 (2012)

  27. 27.

    & Late Quaternary vegetational history of the Central Highlands of Sumatra. II. Palaeopalynology and vegetational history. J. Biogeogr. 15, 555–578 (1988)

  28. 28.

    & Coasting out of Africa: the potential of mangrove forests and marine habitats to facilitate human coastal expansion via the Southern Dispersal Route. Quat. Int. 382, 31–41 (2015)

  29. 29.

    & Pleistocene rainforests: barriers or attractive environments for early human foragers? World Archaeol. 47, 718–739 (2015)

  30. 30.

    The dentition of Sinanthropus pekinensis: a comparative odontography of the hominids (China Geological survey Palaeontologia sinica, new series D, 1937)

  31. 31.

    The Aleut Dentition (Harvard Univ. Press, 1957)

  32. 32.

    (National Institutes of Health, USA, 1997–2008)

  33. 33.

    ., . & In Advances in Dental Anthropology (eds . & ) 13–31 (Wiley-Liss, 1991)

  34. 34.

    , , , & Modern human molar enamel thickness and enamel–dentine junction shape. Arch. Oral Biol. 51, 974–995 (2006)

  35. 35.

    et al. Dental tissue proportions in fossil orangutans from mainland Asia and Indonesia. Hum. Origins Res. 1, e1 (2011)

  36. 36.

    , , , & Enamel thickness in Bornean and Sumatran orangutan dentitions. Am. J. Phys. Anthropol. 147, 417–426 (2012)

  37. 37.

    Relationships of the later Miocene Hominoidea. PhD thesis, Univ. College London (1983)

  38. 38.

    Significance of enamel thickness in hominoid evolution. Nature 314, 260–263 (1985)

  39. 39.

    , , & Enamel–dentine junction (EDJ) morphology distinguishes the lower molars of Australopithecus africanus and Paranthropus robustus. J. Hum. Evol. 55, 979–988 (2008)

  40. 40.

    , , , & Discrimination of extant Pan species and subspecies using the enamel–dentine junction morphology of lower molars. Am. J. Phys. Anthropol. 140, 234–243 (2009)

  41. 41.

    , & Optical dating of sediments. Nature 313, 105–107 (1985)

  42. 42.

    & A dual-aliquot regenerative-dose protocol (DAP) for thermoluminescence (TL) dating of quartz sediments using the light-sensitive and isothermally stimulated red emissions. Quat. Sci. Rev. 25, 2513–2528 (2006)

  43. 43.

    & Effect of the heating rate on the red TL of quartz. Radiat. Meas. 35, 59–66 (2002)

  44. 44.

    & Comparison of regenerative-dose single-aliquot and multiple-aliquot (SARA) protocols using heated quartz from archaeological sites. Quat. Sci. Rev. 18, 223–229 (1999)

  45. 45.

    , & Isothermal thermoluminescence signals from quartz. Radiat. Meas. 41, 796–802 (2006)

  46. 46.

    & Luminescence dating of quartz using an improved single-aliquot regenerative-dose protocol. Radiat. Meas. 32, 57–73 (2000)

  47. 47.

    et al. Archaeology and age of a new hominin from Flores in eastern Indonesia. Nature 431, 1087–1091 (2004)

  48. 48.

    , , & Laboratory fading rates of various luminescence signals from feldspar-rich sediment extracts. Radiat. Meas. 43, 1474–1486 (2008)

  49. 49.

    , , & The effect of preheating on the IRSL signal from feldspar. Radiat. Meas. 44, 554–559 (2009)

  50. 50.

    et al. Luminescence dating of the Stratzing loess profile (Austria)—testing the potential of an elevated temperature post-IR IRSL protocol. Quat. Int. 234, 23–31 (2011)

  51. 51.

    , & Stability of IRSL signals from sedimentary K-feldspar samples. Geochronometria 38, 1–13 (2011)

  52. 52.

    , , & Testing the potential of an elevated temperature IRSL signal from K-feldspar. Radiat. Meas. 44, 560–565 (2009)

  53. 53.

    et al. Alternative chronologies for Late Quaternary (Last Interglacial–Holocene) deep sea sediments via optical dating of silt-sized quartz. Quat. Sci. Rev. 22, 925–941 (2003)

  54. 54.

    Thermoluminescence dating: beta-dose attenuation in quartz grains. Archaeometry 21, 61–72 (1979)

  55. 55.

    & Luminescence dating at Katanda—a reassessment. Quat. Sci. Rev. 20, 961–966 (2001)

  56. 56.

    & The K content of the K-feldspars being measured in optical dating or in thermoluminescence dating. Anc. TL 15, 11–13 (1997)

  57. 57.

    & The Rb contents of the K-feldspars being measured in optical dating. Anc. TL 19, 43–46 (2001)

  58. 58.

    & Cosmic-ray contributions to dose rates for luminescence and ESR dating: large depths and long-term time variations. Radiat. Meas. 23, 497–500 (1994)

  59. 59.

    , , & Thermal ionization mass spectrometry U-series dating of a hominid site near Nanjing, China. Geology 29, 27–30 (2001)

  60. 60.

    , , , & Speleothem-derived Asian summer monsoon variations in Central China during 54–46 ka. J. Quat. Sci. 26, 781–790 (2011)

  61. 61.

    et al. Discerning the timing and cause of historical mortality events in modern Porites from the Great Barrier Reef. Geochim. Cosmochim. Acta 138, 57–80 (2014)

  62. 62.

    User’s Manual for Isoplot 3.75. A Geochronological Toolkit for Microsoft Excel (Berkeley Geochronology Center Special Publication No. 5, 2012)

  63. 63.

    et al. The half-lives of uranium-234 and thorium-230. Chem. Geol. 169, 17–33 (2000)

  64. 64.

    et al. In situ U-series dating by laser-ablation multi-collector ICPMS: new prospects for Quaternary geochronology. Quat. Sci. Rev. 24, 2523–2538 (2005)

  65. 65.

    , , , & Laser ablation U-series analysis of fossil bones and teeth. Palaeogeogr. Palaeoclimatol. Palaeoecol. 416, 120–167 (2014)

  66. 66.

    et al. ESR and U-series analyses of teeth from the palaeoanthropological site of Hexian, Anhui Province, China. J. Hum. Evol. 34, 555–564 (1998)

  67. 67.

    User’s Manual for Isoplot 3.00 (Berkeley Geochronology Center, 2003)

  68. 68.

    Detailed protocol for an accurate non-destructive direct dating of tooth enamel fragment using electron spin resonance. Geochronometria 40, 322–333 (2013)

  69. 69.

    & Are published ESR dose assessments on fossil tooth enamel reliable? Quat. Geochronol. 31, 19–27 (2016)

  70. 70.

    , , , & A new U-uptake model for combined ESR/U-series dating of tooth enamel. Quat. Geochronol. 10, 406–411 (2012)

  71. 71.

    , & Dose–rate conversion factors: update. Anc. TL 29, 5–8 (2011)

  72. 72.

    Radiocarbon calibration and analysis of stratigraphy: the OxCal program. Radiocarbon 37, 425–430 (1995)

  73. 73.

    , , , & Hominin teeth from the early Late Pleistocene site of Xujiayao, Northern China. Am. J. Phys. Anthropol. 156, 224–240 (2015)

Download references


This research was funded by Australian Research Council Discovery grants (DP1093049, DP140100919, and DP120101752 to K.E.W., R.J.-B., and G.J.P. et al., respectively) and a Leaky Foundation grant and Research School of Asia and the Pacific Grant Development Support grant to J.L. C.S.’s research is supported by the Human Origins Research Fund and the Calleva Foundation. We acknowledge the Max Planck Society for funding micro-CT scanning of the teeth, A. Olejnicak and J. P. Zermeno for assistance with section preparation, and support provided by the Centre from Archaeology in Padang Sumatra and ARKENAS in Jakarta and for allowing access to the site and four fossil faunal teeth for dating. We thank the Department of Geology, Naturalis Biodiversity Center in Leiden, The Netherlands for providing access to Dubois’s fieldnote book, excavation details, the two modern human teeth for scanning and the Pongo tooth for dating, and we thank C. Bronk Ramsey for assistance with age modelling.

Author information

Author notes

    • R. Due Awe
    •  & M. J. Morwood



  1. Department of Environmental Sciences, Faculty of Science and Engineering, Macquarie University, Sydney, New South Wales 2109, Australia

    • K. E. Westaway
  2. School of Culture, History, and Languages, ANU College of Asia and the Pacific, Australian National University, Canberra, Australia

    • J. Louys
  3. Indonesian Centre for Archaeology, Jl. Raya Condet Pejaten No. 4, Jakarta 12001, Indonesia

    • R. Due Awe
    • , E. W. Saptomo
    •  & B. Sulistyanto
  4. Centre for Archaeological Sciences, School of Earth and Environmental Sciences, University of Wollongong, Wollongong, New South Wales 2522, Australia

    • M. J. Morwood
    •  & G. D. van den Bergh
  5. School of Earth and Environmental Sciences, University of Queensland, Brisbane, Queensland 4072, Australia

    • G. J. Price
    •  & J.-x. Zhao
  6. Place, Evolution and Rock Art Heritage Unit (PERAHU), Griffith University, Gold Coast, Queensland 4222, Australia

    • M. Aubert
  7. Southern Cross GeoScience, Southern Cross University, Military Road, Lismore, New South Wales 2480, Australia

    • R. Joannes-Boyau
  8. Australian Research Centre for Human Evolution, Environmental Futures Research Institute, Griffith University, 170 Kessels Road, Nathan, Queensland 4111, Australia

    • T. M. Smith
  9. Department of Human Evolutionary Biology, Harvard University, 11 Divinity Avenue, Cambridge, Massachusetts 02138, USA

    • T. M. Smith
  10. School of Anthropology and Conservation, University of Kent, Canterbury CT2 7NR, UK

    • M. M. Skinner
  11. Department of Human Evolution, Max Planck Institute for Evolutionary Anthropology, Deutscher Platz 6, Leipzig 04103, Germany

    • M. M. Skinner
  12. Department of Earth Sciences, Natural History Museum, Cromwell Road, London SW7 5BD, UK

    • T. Compton
    •  & C. Stringer
  13. School of Geography and the Environment, University of Oxford, Oxford OX1 2JD, UK

    • R. M. Bailey
  14. Department of Geology, Naturalis Biodiversity Center, Leiden, The Netherlands

    • J. de Vos
  15. Department of Archaeology, University of Southampton, Highfield Road, Southampton SO17 1BF, UK

    • A. W. G. Pike
  16. Geology Study Program, Institut Teknologi Bandung, Java, Indonesia

    • Y. Rizal
    • , J. Zaim
    • , W. D. Santoso
    •  & A. Trihascaryo
  17. Research School of Earth Sciences, Australian National University, Canberra, Australia

    • L. Kinsley


  1. Search for K. E. Westaway in:

  2. Search for J. Louys in:

  3. Search for R. Due Awe in:

  4. Search for M. J. Morwood in:

  5. Search for G. J. Price in:

  6. Search for J.-x. Zhao in:

  7. Search for M. Aubert in:

  8. Search for R. Joannes-Boyau in:

  9. Search for T. M. Smith in:

  10. Search for M. M. Skinner in:

  11. Search for T. Compton in:

  12. Search for R. M. Bailey in:

  13. Search for G. D. van den Bergh in:

  14. Search for J. de Vos in:

  15. Search for A. W. G. Pike in:

  16. Search for C. Stringer in:

  17. Search for E. W. Saptomo in:

  18. Search for Y. Rizal in:

  19. Search for J. Zaim in:

  20. Search for W. D. Santoso in:

  21. Search for A. Trihascaryo in:

  22. Search for L. Kinsley in:

  23. Search for B. Sulistyanto in:


K.E.W., R.D.A., J.L., G.J.P., W.D.S. mapped and excavated the site and collected faunal and dating samples, K.E.W. conducted the red thermoluminescence and pIR-IRSL dating, J.-x.Z. and G.J.P. conducted the U-series measurements on the speleothem, while M.A. and L.K. conducted U-series profiling on the fossil teeth and R.J.-B. conducted the U-series/ESR dating. G.D.v.d.B. and R.D.A. analysed the fauna and M.J.M., G.D.v.d.B., J.d.V., Y.R., J.Z., W.D.S. and A.T. helped to find and organize access to the site. T.M.S. and J.d.V. conducted the micro-CT scanning of the teeth, and T.M.S. measured enamel thickness. T.C. and C.S. described the teeth and M.M.S. analysed the enamel–dentine junction. R.M.B. aided in the design of the dating approach and conducted the Bayesian modelling, while A.W.G.P. conducted the modelling of the U-series age estimates. Finally, E.W.S. and B.S. helped with the dating of the fauna and K.E.W., J.L., G.J.P., J.-x.Z., R.J.-B., G.D.v.d.B., M.A., T.M.S., T.C., M.M.S., C.S. and J.d.V. wrote the paper, with early contributions from M.J.M. and R.D.A.

Competing interests

The authors declare no competing financial interests.

Corresponding author

Correspondence to K. E. Westaway.

Reviewer Information Nature thanks G. Barker, R. Dennell and the other anonymous reviewer(s) for their contribution to the peer review of this work.

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

Extended data

Supplementary information

PDF files

  1. 1.

    Supplementary Information

    This file contains SI sections 1-8.

About this article

Publication history







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.