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Denisovans from southwestern China and their subsistence strategies

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Data availability

The raw data used for analysis are available in the Supplementary Information and can be accessed online at OSF (https://doi.org/10.17605/OSF.IO/PFWNR). All archaeological materials studied in this paper are stored in the Yunnan Provincial Institute of Cultural Relics and Archaeology, China. Access to the materials can be arranged by contacting our first author Q.J.R.

Code availability

Sample code to analyse the available data to reproduce part of the above results is available at OSF (https://doi.org/10.17605/OSF.IO/PFWNR) under an open licence.

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Acknowledgements

We thank C. C. Yang, S. Yang, J. W. Zhou, X. M. Zhang, L. G. Wu, X. R. Luo, J. Y. Li, Z. J. Song, J. Sun and R. B. Ye for assistance with field excavation and materials curation; G. Y. Bai for help on pollen analysis; R. Grün for suggestions on the U-series dating; and X. J. Wu for providing Hualongdong specimens for comparisons.

Funding

This research was eusoft-did-translate-comment-enfunded by the National Key R&D Program of China (2023YFF0905700 to S.X.), the NSFC Excellent Research Group Program for Tibetan Plateau Earth System (42588201 to F.H.C.), the National Natural Science Foundation of China (U2574202 and 42471180 to H.L. and 42471185 to K.L.Z.), the Archaeological Talent Promotion Program of China (no. 2025-198 to H.L. and no. 2024-278 to Q.M.F. and Q.J.R.), the Open Research Fund of TPESER (no. TPESER202210 to Q.J.R.), the Association for Trans-Eurasia Exchange and Silk Road Civilization Development (ATES) (to H.L., L.D., F.D. and F.H.C.), the University of Ferrara (FAR2023 to M.P.), the European Research Council (ERC) Starting ExOsTech (grant no. 101161065 to L.D.), the French government in the framework of the University of Bordeaux’s IdEx ‘Investments for the Future’ program/GPR ‘Human Past’ (to L.D. and F.D.), the Australian Research Council (project DP210100717 to B.L.), the ERC Synergy grant QUANTA (grant no. 951388 to F.D.), and the Research Council of Norway through its Centres of Excellence funding scheme (SFF Centre for Early Sapiens Behaviour, SapienCE-project no. 262618 to F.D.).

Author information

Author notes

  1. These authors contributed equally: Qijun Ruan, Hao Li, Song Xing, Keliang Zhao

Authors and Affiliations

  1. State Key Laboratory of Tibetan Plateau Earth System, Environment and Resources, Institute of Tibetan Plateau Research, Chinese Academy of Sciences, Beijing, China

    Qijun Ruan, Hao Li, Peiyuan Xiao, Yuqing Wang, Zhenxiu Jia & Fahu Chen

  2. Yunnan Provincial Institute of Cultural Relics and Archaeology, Kunming, China

    Qijun Ruan & Jianhui Liu

  3. Center for Excellence in Archaeological Science and Cultural Heritage, MOE Key Laboratory of Western China’s Environmental System, Lanzhou University, Lanzhou, China

    Hao Li & Fahu Chen

  4. Key Scientific Research Base on Paleolithic Human Evolution and Paleogenetics, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing, China

    Song Xing, Keliang Zhao, Qiaomei Fu, Qiyahui Sun & Sungui Lin

  5. Centro Nacional de Investigación sobre la Evolución Humana (CENIEH), Burgos, Spain

    Song Xing

  6. Univ. Bordeaux, CNRS, MCC, PACEA, UMR 5199, Pessac, France

    Clément Zanolli & Francesco d’Errico

  7. GIZAPRE Consolidated Research Group (IT1852-26), Department of Geography, Prehistory and Archaeology, Euskal Herriko Unibertsitatea (EHU), Vitoria-Gasteiz, Spain

    Luc Doyon

  8. Basque Foundation for Science IKERBASQUE, Bilbao, 48009, Spain

    Luc Doyon

  9. Department of Human Studies, Prehistoric and Anthropological Science Unit, University of Ferrara, Ferrara, Italy

    Davide Delpiano & Marco Peresani

  10. CNR-Institute of Environmental Geology and Geoengineering, Laboratory of Palynology and Palaeoecology, Research Group on Vegetation, Climate and Human Stratigraphy, Milan, Italy

    Marco Peresani

  11. SFF Centre for Early Sapiens Behaviour (SapienCE), University of Bergen, Bergen, Norway

    Francesco d’Errico

  12. GeoGenetics Centre, Globe Institute, University of Copenhagen, Copenhagen, Denmark

    Fabrice Demeter

  13. Eco-anthropologie (EA), Muséum national d’Histoire naturelle, CNRS, Université de Paris, Paris, France

    Fabrice Demeter

  14. Collège de France-CNRS/UMR, Paris, France

    Jean-Jacques Hublin

  15. University of Chinese Academy of Sciences, Beijing, China

    Peiyuan Xiao, Yuqing Wang, Qiyahui Sun & Sungui Lin

  16. College of Resources and Environment, Chengdu University of Information Technology, Chengdu, China

    Qingjiang Yang

  17. School of Archaeology and Museology, Peking University, Beijing, China

    Yuhao Zhao & Youping Wang

  18. Department for the History of Science and Scientific Archaeology, University of Science and Technology of China, Hefei, China

    Anchuan Fan

  19. School of Geography, Nanjing Normal University, Nanjing, China

    Qingfeng Shao

  20. Center for Excellence in Archaeological Science and Cultural Heritage, Lanzhou University, Lanzhou, China

    Qingfeng Shao

  21. School of Science, University of Wollongong, Wollongong, New South Wales, Australia

    Bo Li

  22. Environmental Futures Research Centre, University of Wollongong, Wollongong, New South Wales, Australia

    Bo Li

Authors

  1. Qijun Ruan
  2. Hao Li
  3. Song Xing
  4. Keliang Zhao
  5. Jianhui Liu
  6. Clément Zanolli
  7. Luc Doyon
  8. Davide Delpiano
  9. Marco Peresani
  10. Francesco d’Errico
  11. Fabrice Demeter
  12. Jean-Jacques Hublin
  13. Peiyuan Xiao
  14. Yuqing Wang
  15. Zhenxiu Jia
  16. Qingjiang Yang
  17. Qiaomei Fu
  18. Qiyahui Sun
  19. Yuhao Zhao
  20. Sungui Lin
  21. Anchuan Fan
  22. Youping Wang
  23. Qingfeng Shao
  24. Bo Li
  25. Fahu Chen

Contributions

Q.R., H.L., S.X., B.L., Youping Wang and F.C. designed the study; Q.R. and J.L. led field excavation and site sampling; B.L., A.F. and Q. Shao performed luminescence and U-series dating analyses; S.X., C.Z., F.D., J.-J.H., Q.F., Q. Sun, Y.Z. and S.L. performed analysis of hominin fossils; Q.R., H.L., P.X., D.D., M.P. and Z.J. performed analysis of stone artefacts; J.L., Yuqing Wang, L.D. and F.d.E. performed zooarchaeological analysis; L.D. and Yuqing Wang performed analysis of bone tools; K.Z. and Q.Y. performed pollen analysis; Q.R., H.L., S.X., K.Z., L.D., Q. Shao, B.L., F.C. and F.d.E. drafted and substantively revised the paper, with additional input from all other authors.

Corresponding authors

Correspondence to Hao Li, Qingfeng Shao, Bo Li or Fahu Chen.

Ethics declarations

Competing interests

The authors declare no competing interests.

Peer review

Peer review information

Nature thanks Xuefeng Sun, Bence Viola, Wei Wang and the other, anonymous, reviewer(s) for their contribution to the peer review of this work. Peer reviewer reports are available.

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Extended data figures and tables

Extended Data Fig. 1 Excavation area and locations of optical dating samples at the site.

a, Aerial photo showing the geographical context of the site; b, Plan view of the excavated trenches; c-d, Locations of optical dating samples from Trench 2.

Extended Data Fig. 2 The percentages diagram of main pollen taxa, and pollen-based biome reconstruction results from Bianfu Cave.

(STEP: Cool-temperate steppe; TEFS: Cool-temperate forest steppe; WTFO: Middle subtropical broadleaf evergreen forest; TEDE: Warm-temperate mixed forest; WAMF: North subtropical mixed forest; COMX: Cool-temperate mixed forest; CLEC: Cold-temperate evergreen conifer forest).

Extended Data Fig. 3 The hominin molars unearthed from the Bianfu Cave.

a, From left to right, the occlusal, buccal, mesial, lingual, and distal views of the fossil (top row), the virtual reconstruction of the outer enamel surface (middle row), and enamel-dentine junction (bottom row) of Bianfu Cave right M2 (YHB2890) were shown; b, From left to right, the occlusal, buccal, mesial, lingual, and distal views of the fossil (top row), the virtual reconstruction of the outer enamel surface (middle row), and enamel-dentine junction (bottom row) of Bianfu Cave right M2 (YHB3075) were shown.

Extended Data Fig. 4 The hominin incisor and premolar unearthed from the Bianfu Cave.

a, From left to right, the occlusal, labial, mesial, lingual, and distal views of the fossil (top row), virtual reconstruction of the outer enamel surface (middle row), and enamel-dentine junction (bottom row) of Bianfu Cave I2 (YHB3513) were shown; b, From left to right, the occlusal, buccal, and distal views of the fossil (top row), occlusal, buccal, mesial, lingual, and distal view of the virtual reconstruction of the outer enamel surface (middle row) and enamel-dentine junction (bottom row) of Bianfu Cave P4 (YHB3518) were shown.

Extended Data Fig. 7 Bi-variable plots of the crown metrics of fossil hominins from Bianfu Cave and comparative samples.

Elliptical contours represent the 95% confidence intervals of the distribution. MD: mesiodistal dimension; BL: buccolingual dimension.

Extended Data Fig. 8 Stone artefacts from Bianfu Cave.

a-g, Cores; h-o, Flakes; p, Hammerstone; q-r, Retouched tools.

Extended Data Fig. 10 Bone tools from Bianfu Cave.

a-c, Bone retouchers from layer 7; d, Intermediate tool from layer 3; e, Potential expedient tool from layer 4; f, Bone retoucher from layer 3.

Supplementary informations

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Ruan, Q., Li, H., Xing, S. et al. Denisovans from southwestern China and their subsistence strategies. Nature (2026). https://doi.org/10.1038/s41586-026-10997-4

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