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.
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Investigating the palaeoenvironmental context of Late Pleistocene human dispersals into Southeast Asia: a review of stable isotope applications
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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.
The authors declare no competing financial interests.
Reviewer Information Nature thanks G. Barker, R. Dennell and the other anonymous reviewer(s) for their contribution to the peer review of this work.
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Extended data figures and tables
a, Corridor of dispersal of fauna into southeast Asia during periods of connection (redrawn with permission from ref. 7). b, The main fossil faunal sites in southeast Asia. In southern China: 1, Luijiang; 2, Liucheng; 3, Hoshantung; 4, Hei-Tu’ung; 5, Changyang; 6, Hsing-an. In Vietnam: 7, Lang Trang; 8, Tham Khuyen; 9, Thung Lang; 10, Hang Hum; 11, Ma U ‘Oi; 12, Tham Om; 13, Keo Leng. In Laos: 14, Tham Hang; 15, Tham P’a Loi. In Thailand: 16, Thum Wiman Nakin; 17, Thum Phra Khai Phet. In Cambodia: 18, Phnom Loang. In Borneo; 19, Niah Cave. In Indonesia: 20, Lida Ajer; 21, Sibrambang; 22, Punung (redrawn with permission from ref. 7). c, Dubois’ field sketches of Lida Ajer cave location copied directly from his field notebook—now housed in Leiden (with permission from the Naturalis, the Netherlands). His rough sketch of the cave location close to Payakumbuh village has had annotations added to make the features clearer. d, Our map of the cave location for comparison, note the similar relationship between Mount Sago, River Agam and Lida Ajer. e, Dubois’ plan of the cave, annotations have been added to identify the chambers discussed in the text. f, Our plan of the cave for comparison, with the only differences being the absence of the sinkhole passage on our plan (unmapped).
a, Cervid sp. b, Cervid sp. c, Pongo sp., upper premolar. d, Rusa sp. e, Pongo sp., molar. f, Pongo sp., molar mesial view from c. g, Siamang gibbon, molar. h, Pongo sp., molar mesial view from e. i, Hystrix sp. j, Soda straw stalactite samples LA08-29 (own scale on photograph). k, Photograph of areas 3 and 4 in the cave where the majority of fossil fauna were discovered.
a, b, The incisor (a) and molar (b) mesio:distal ratio versus bucco:lingual ratio metrics plotted against data from 37 and 353 fossil Pongo teeth15, respectively. c, d, The incisor (c) and molar (d) data are plotted against the full range of Homo teeth from African early Homo to recent modern (data from ref. 73). In all four plots, the Lida Ajer teeth are denoted by a red star, with the key for symbols in c, d, representing the different human teeth is indicated on the right. e, f, The incisor (e) and molar (f) from Lida Ajer.
a, Virtual sections of the Lida Ajer teeth. The labio-lingual section of the incisor is shown on the left, the bucco-lingual section through the mesial molar cusps is shown on the right. Scale bar, 5 mm. b, EDJ anatomical landmarks. Landmark protocol for geometric morphometric analysis of EDJ shape. Numbers in brackets represent the number of equidistantly spaced landmarks between main landmarks (red spheres) and around the cervix.
Top, CT-based volume renderings of the external surface (left) and surface models of the EDJ (right) of the Lida Ajer molar in six anatomical views. Bottom, initial landmark placement (yellow spheres) capturing the main dentine horns, EDJ ridge and cervix (left) and noting the presence of an accessory dentine horn mesial to the metacone (right).
Extended Data Figure 6 Principal component analysis of the EDJ shape of the comparative sample and the Lida Ajer molar.
a, b, A comparison of the red thermoluminescence signal characteristics using glow curves derived from a Liang Bua sample WR1 (a) and from the Lida Ajer sample (b). The glow curves demonstrate that after 500 Gy dosing the low temperature peaks disappear with the introduction of the 260 °C preheat, and the presence of a light-sensitive shoulder (260–305 °C) that is removed by 1 h of bleaching. The Lida Ajer sample shows similarities with the Liang Bua sample, but has a more defined bleachable shoulder and a more intense signal. c, Isothermal decay of the red thermoluminescence signal from sample LA-1. d, Dose–response curve for the unbleachable signal derived from aliquot A providing a De of 132 ± 13 Gy (see ref. 42 for further methodological details). e, pIR-IRSL intensity and shine down from red-diode stimulation for 250 s at 270 °C, displaying the natural curve and a regenerative dose for comparison. f, pIR-IRSL sensitivity corrected dose–response providing a De of 103 ± 9 Gy. g, The De values of the 22 aliquots of feldspars plotted on a radial plot. Each aliquot was corrected for minor fading and residual dose and was plotted producing an overdispersion of 17.6%. Prior to running the minimum age model a value of 10% was added to the errors as an estimation of inherent overdispersion within the grains. This was determined by estimating the distribution of De values of 12 aliquots after a 4 h bleaching period in a solar simulator. The minimum age model produced a De of 105 ± 3 Gy as depicted by a solid black line, which lies within ±10% of the Central age (shaded box), owing to the low overdispersion. This produces an age estimate of 62 ± 5 kyr. h, Fading tests for the Lida Ajer feldspars comparing the IR50 measurement with a g value of 17.67 with the pIR-IRSL270 measurement, which has reduced the g value to 1.74.
a, 7/LA/5/08, a molar of siamang gibbon sp., sampled during our excavations. b, Sample 12/LA/5/08, a premolar of Pongo sp., sampled during our excavations. c, 13/LA/5/08, a molar of Pongo sp., sampled during our excavations. d, Dubois 9967A, a Pongo sp. molar from Dubois’s original excavation—borrowed from the Naturalis Museum in the Netherlands. e–h, U-series profiling tracks on the 4 fossil teeth (7-, 12-, 13-, and 21/LA/5/08). i, Example of the best fit D–A (diffusion–absorption model) date profile for sample 13/LA/5/08 (with 4/8 = 1.066, t′ = 1.0) demonstrating that the age estimate fits the model at around 55 kyr. The possibility of delayed uptake of uranium and the absence of evidence for uranium leaching means that this should be treated as a minimum age. The U-series profiles from other teeth did not fit well with the predictions of the D–A model owing to complex U-uptake (and potentially U-loss) processes in the sampled teeth.
Extended Data Figure 9 ESR dating of two Fossil Pongo teeth 12/LA/5/08 (orange) and 13/LA/5/08 (blue).
a, ESR dose equivalent (De) calculation. Top, Markov Chain Monte-Carlo fitted dose–reponse curve for each of the samples, using McDoseE 2.0 with a single saturating exponential function and 100,000 iterations. Bottom, ESR dose equivalent distribution of the Markov Chain Monte-Carlo model with McDoseE 2.0. b, Uranium uptake model in the different tissues used for the U-series/ESR age calculation. c, Table summarizing the U-series values (averaged) obtained by LA-MC-ICPMS on the ESR fragment and dentine directly in contact (EDJ) and used in the coupled U-series/ESR age model. No ages were calculated for U concentration <1 p.p.m. or U/Th ratio <500. d, Sample 13/LA/5/08. e, Sample 12/LA/5/08.
a, Photograph of the fossil chamber, showing the location and structure of the breccia and flowstone units. b, Annotated photograph of the fossil chamber with the sampling locations and dating results found in Supplementary Tables 7, 8, 11. c, Bayesian analysis of the red thermoluminescence, U-series and coupled U-series/ESR dating results to construct the new modelled chronology for Lida Ajer. The photograph on the left (taken from the dashed box in a) depicts the boundaries between the underlying flowstone, the breccia deposit and overlying flowstone units. Note: the red thermoluminescence and ESR error on the age estimates are presented at 1σ, while the U-series errors have been presented at 2σ. The main figure uses all the available data, while inset A uses only the breccia data (from the red thermoluminescence and pIR-IRSL dating of the breccia matrix and U-series dating of the flowstones and soda straw) and inset B uses only the fossil tooth data (from U-series age depth modelling and coupled U-series/ESR dating of the teeth directly).
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Westaway, K., Louys, J., Awe, R. et al. An early modern human presence in Sumatra 73,000–63,000 years ago. Nature 548, 322–325 (2017). https://doi.org/10.1038/nature23452
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