Abstract
During the Early to Middle Pleistocene, Java was inhabited by hominid taxa of great diversity. However, their seasonal dietary strategies have never been explored. We undertook geochemical analyses of orangutan (Pongo sp.), Homo erectus and other mammalian Pleistocene teeth from Sangiran. We reconstructed past dietary strategies at subweekly resolution and inferred seasonal ecological patterns. Histologically controlled spatially resolved elemental analyses by laser-based plasma mass spectrometry confirmed the preservation of authentic biogenic signals despite the effect of spatially restricted diagenetic overprint. The Sr/Ca record of faunal remains is in line with expected trophic positions, contextualizing fossil hominid diet. Pongo sp. displays marked seasonal cycles with ~3 month-long strongly elevated Sr/Ca peaks, reflecting contrasting plant food consumption presumably during the monsoon season, while lower Sr/Ca ratios suggest different food availability during the dry season. In contrast, omnivorous H. erectus shows low and less accentuated intra-annual Sr/Ca variability compared to Pongo sp., with δ13C data of one individual indicating a dietary shift from C4 to a mix of C3 and C4 plants. Our data suggest that H. erectus on Java was maximizing the resources available in more open mosaic habitats and was less dependent on variations in seasonal resource availability. While still influenced by seasonal food availability, we infer that H. erectus was affected to a lesser degree than Pongo sp., which inhabited monsoonal rain forests on Java. We suggest that H. erectus maintained a greater degree of nutritional independence by exploiting the regional diversity of food resources across the seasons.
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Data availability
Data of elemental analyses and corresponding contextual information obtained in this study are available as Supplementary Data.
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Acknowledgements
We express our gratitude to the Werner Reimers Foundation in Bad Homburg (Germany), which provides the Gustav Heinrich Ralph von Koenigswald collection as a permanent loan for scientific research to the Senckenberg Research Institute and Natural History Museum Frankfurt. FIERCE, where all LA-ICPMS analyses were performed, is financially supported by the Wilhelm and Else Heraeus Foundation and by the Deutsche Forschungsgemeinschaft (DFG, INST 161/921-1 FUGG and INST 161/923-1 FUGG), which are gratefully acknowledged. We thank L. Marko and A. Gerdes for help with analytical work. This is FIERCE contribution no. 113. We thank R. Brocke and G. Riedel for assistance with microscopic imaging. C.Z. acknowledges the support of the French CNRS (Centre National de la Recherche Scientifique). J.K. received funding from the Erasmus+ Traineeship programme (2019). A.N. received funding from the European Union’s Horizon 2020 research and innovation programme under the Marie Skłodowska-Curie grant agreement (no. 842812). T.L. received funding from the DFG (Emmy Noether Fellowship LU 2199/2-1).
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The study was initiated by W.M., F.S. and J.K. and forms part of J.K.’s MSc research project completed under the supervision of W.M., L.B. and A.N. J.K., W.M., A.N., L.B., F.S. and O.K. designed research. J.K., W.M., A.N., L.B., B.P., T.L. and R.A. performed research. J.K., W.M., A.N., T.L., P.M. and L.B. analysed data. J.K., W.M., A.N., L.B., F.S., O.K., C.Z., T.L. and C.H. wrote the manuscript, with discussions on data interpretation and contributions to the manuscript text from M.C.D., N.B., A.-M.B. and F.D.
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Extended data
Extended Data Fig. 1 Scatter plots of [Sr] or [Ba] vs. [U], respectively, for representative examples of each faunal group.
Scatter plots of [Sr] or [Ba] vs. [U], respectively, for representative examples of each faunal group, to illustrate the diagenesis assessment of the fossil assemblage. While both [Sr] and [Ba] broadly positively correlate with [U], [Ba] increases proportionally much more than [Sr], namely 1.4 to 1.7-fold for Sr and 3 to 7-fold for Ba. For simplicity, data are shown here as concentrations, whereas elsewhere they are displayed as Element/Ca to facilitate comparison throughout. See Fig. 1 for equivalent plots relative to [Mn].
Extended Data Fig. 2 Time-resolved compositional profiles for H. erectus SMF-8865 molar.
a) Sr/Ca, Ba/Ca, [U] and [Mn] along the buccal EDJ plotted against relative days. b) Sr/Ca, Ba/Ca, [U] and [Mn] along the lingual EDJ plotted against relative days. c) Comparative Sr/Ca ratio profiles for the mesiolingual and mesiobuccal cusps plotted against relative days. Based on the overall lower [U]-signal, the lingual Sr/Ca data are considered slightly more reliable.
Extended Data Fig. 3 Time-resolved compositional profiles for H. erectus S7-13 molar.
a) Sr/Ca, Ba/Ca, [U] and [Mn] along the buccal EDJ plotted against relative days. b) Sr/Ca, Ba/Ca, [U] and [Mn] along the lingual EDJ plotted against relative days. c) Comparative Sr/Ca ratio profiles for the mesiolingual and mesiobuccal cusps plotted against relative days. Based on the overall lower [U] and [Mn] signals, the lingual Sr/Ca data are considered more reliable, which is corroborated by the more pronounced diagenetic overprint for buccal Ba/Ca.
Extended Data Fig. 4 Time-resolved compositional profiles for Pongo sp. SMF-8864 molar.
a) Sr/Ca, U and Mn signals along the EDJ and corresponding prism orientations plotted against relative days. b) Ba/Ca, U and Mn signals along the EDJ and corresponding prism orientations plotted against relative days. c) Micrograph of the molar section with laser ablation paths along the EDJ and prism orientations highlighted.
Extended Data Fig. 5 Time-resolved compositional profiles for H. erectus SMF-8865 with emphasis on comparative EDJ vs. prism orientations.
a) Sr/Ca and U as well as b) Ba/Ca and U signals, all along the EDJ vs. corresponding prism orientations (that is across enamel) plotted against relative days. c) Micrograph of the molar section with laser ablation paths along the EDJ and prism orientations highlighted. Relatively low [U] betray limited diagenetic overprint; all prism profiles show substantially lower Sr/Ca and Ba/Ca values towards outer enamel due to maturation overprint26, which confirms that EDJ profiles preserve initial dietary information more faithfully.
Extended Data Fig. 6 Time-resolved compositional profiles for H. erectus S7-13 with emphasis on comparative EDJ vs. prism orientations.
a) Sr/Ca and U as well as b) Ba/Ca and U signals, all along the EDJ vs. corresponding prism orientations (that is across enamel) plotted against relative days. c) Micrograph of the molar section with laser ablation paths along the EDJ and prism orientations highlighted. Relatively low [U] along EDJ indicate limited diagenetic overprint of inner enamel, which contrasts all prism profiles with their increasing U-profiles, with [U] up to 10 ppm. Sr/Ca and Ba/Ca along the prisms increase accordingly by ~60% and ~300%, respectively, parallel to U, which confirms the greater susceptibility of Ba/Ca to diagenetic overprint.
Extended Data Fig. 7 Sr/Ca and Ba/Ca ratios boxplots comparing H. erectus and Pongo sp. specimens to the other taxa with known trophic levels.
a) Sr/Ca ratios of all data along the EDJ profiles. b) Sr/Ca ratios of the same dataset after diagenesis filtering, including points with [U] < 1 ppm and [Mn]<400 ppm (µg/g). c) Ba/Ca ratios of all data along the EDJ profiles. d) Ba/Ca ratios of the same dataset after diagenesis filtering as in b). Colour dots outside the whiskers represent outliers, lower whisker is equal to minimum value (excluding outliers), lower hinge equals to first quartile, thick line represents the median value, upper hinge equals to third quartile and upper whisker to maximum value (excluding outliers). H. erectus S7 − 13 N points=1032; H. erectus S7 − 37 N points=1012; H. erectus SMF − 8865 N points=606; Pongo SMF − 8864 N points=708; Felidae N points=3232, N sample=2; Rhinoceratidae N points=631, N sample=2; Suidae N points=189, N sample=2; Cervidae N points=920, N sample=2; Hippopotamidae N points=1263, N sample=2.
Extended Data Fig. 8 Representative micrographs of Pongo sp. SMF-8664.
a) The whole crown before laser ablation (5x objective). b) Lingual aspect after laser ablation (10x objective). Purple lines highlight the laser ablation paths and green lines highlight accentuated lines. The zig-zag path used for the chronology of the crown development follows Retzius lines (red lines) and enamel prisms (yellow lines).
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Data of elemental analyses.
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Kubat, J., Nava, A., Bondioli, L. et al. Dietary strategies of Pleistocene Pongo sp. and Homo erectus on Java (Indonesia). Nat Ecol Evol 7, 279–289 (2023). https://doi.org/10.1038/s41559-022-01947-0
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DOI: https://doi.org/10.1038/s41559-022-01947-0