Krause, J. et al. The complete mitochondrial DNA genome of an unknown hominin from southern Siberia. Nature 464, 894–897 (2010).
Reich, D. et al. Genetic history of an archaic hominin group from Denisova Cave in Siberia. Nature 468, 1053–1060 (2010).
Meyer, M. et al. A high-coverage genome sequence from an archaic Denisovan individual. Science 338, 222–226 (2012).
Chen, F. et al. A late Middle Pleistocene Denisovan mandible from the Tibetan Plateau. Nature 569, 409–412 (2019).
Zhang, D. et al. Denisovan DNA in late Pleistocene sediments from Baishiya Karst Cave on the Tibetan Plateau. Science 370, 584–587 (2020).
Demeter, F. et al. A Middle Pleistocene Denisovan molar from the Annamite Chain of northern Laos. Nat. Commun. 13, 2557 (2022).
Tsutaya, T. et al. A male Denisovan mandible from Pleistocene Taiwan. Science 388, 176–180 (2025).
Fu, Q. et al. The proteome of the late Middle Pleistocene Harbin individual. Science 389, 704–707 (2025).
Fu, Q. et al. Denisovan mitochondrial DNA from dental calculus of the >146,000-year-old Harbin cranium. Cell 188, 3919–3926 (2025).
Rao, H. et al. Ancient proteins identify various Denisovan remains from Southwest China. Nature https://doi.org/10.1038/s41586-026-10976-9 (2026).
Slon, V. et al. A fourth Denisovan individual. Sci. Adv. 3, e1700186 (2017).
Xia, H. et al. Middle and Late Pleistocene Denisovan subsistence at Baishiya Karst Cave. Nature 632, 108–113 (2024).
Larena, M. et al. Philippine Ayta possess the highest level of Denisovan ancestry in the world. Curr. Biol. 31, 4219–4230 (2021).
Reich, D. et al. Denisova admixture and the first modern human dispersals into Southeast Asia and Oceania. Am. J. Hum. Genet. 89, 516–528 (2011).
Sawyer, S. et al. Nuclear and mitochondrial DNA sequences from two Denisovan individuals. Proc. Natl Acad. Sci. USA 112, 15696–15700 (2015).
Slon, V. et al. The genome of the offspring of a Neanderthal mother and a Denisovan father. Nature 561, 113–116 (2018).
Massilani, D. et al. Denisovan ancestry and population history of early East Asians. Science 370, 579–583 (2020).
Ongaro, L. & Huerta-Sanchez, E. A history of multiple Denisovan introgression events in modern humans. Nat. Genet. 56, 2612–2622 (2024).
Peyrégne, S., Slon, V. & Kelso, J. More than a decade of genetic research on the Denisovans. Nat. Rev. Genet. 25, 83–103 (2024).
Brown, S. et al. The earliest Denisovans and their cultural adaptation. Nat. Ecol. Evol. 6, 28–35 (2022).
Chang, C.-H. et al. The first archaic Homo from Taiwan. Nat. Commun. 6, 6037 (2015).
Jacobs, Z. et al. Pleistocene chronology and history of hominins and fauna at Denisova Cave. Nat. Commun. 16, 4738 (2025).
Douka, K. et al. Age estimates for hominin fossils and the onset of the Upper Palaeolithic at Denisova Cave. Nature 565, 640–644 (2019).
Shunkov, M. V., Kozlikin, M. B. & Derevianko, A. P. Dynamics of the Altai Paleolithic industries in the archaeological record of Denisova Cave. Quat. Int. 559, 34–46 (2020).
Zavala, E. I. et al. Pleistocene sediment DNA reveals hominin and faunal turnovers at Denisova Cave. Nature 595, 399–403 (2021).
Zilhão, J., d’Errico, F., Banks, W. E. & Teyssandier, N. A data-driven paradigm shift for the Middle-to-Upper Palaeolithic transition and the Neandertal debate. Quat. Environ. Hum. 2, 100037 (2024).
Browning, S. R., Browning, B. L., Zhou, Y., Tucci, S. & Akey, J. M. Analysis of human sequence data reveals two pulses of archaic Denisovan admixture. Cell 173, 53–61 (2018).
Jacobs, G. S. et al. Multiple deeply divergent Denisovan ancestries in Papuans. Cell 177, 1010–1021 (2019).
Choin, J. et al. Genomic insights into population history and biological adaptation in Oceania. Nature 592, 583–589 (2021).
Grün, R. Direct dating of human fossils. Am. J. Phys. Anthropol. 131, 2–48 (2006).
Lisiecki, L. E. & Raymo, M. E. A Pliocene-Pleistocene stack of 57 globally distributed benthic δ18O records. Paleoceanogr. Paleoclimatol. 20, PA1003 (2005).
Cheng, H. et al. The Asian monsoon over the past 640,000 years and ice age termination. Nature 534, 640–646 (2016).
AlQahtani, S. J., Hector, M. P. & Liversidge, H. M. Brief communication: the London atlas of human tooth development and eruption. Am. J. Phys. Anthropol. 142, 481–490 (2010).
Kaifu, Y. et al. Taxonomic affinities and evolutionary history of the early Pleistocene hominids of Java: dentognathic evidence. Am. J. Phys. Anthropol. 128, 709–726 (2005).
Xing, S., Martinón-Torres, M. & Bermúdez de Castro, J. M. The fossil teeth of the Peking Man. Sci Rep. 8, 2066 (2018).
Martinón-Torres, M. et al. Hominin lower second premolar morphology: evolutionary inferences through geometric morphometric analysis. J. Hum. Evol. 50, 523–533 (2006).
Martinón-Torres, M., Bermúdez de Castro, J. M., Gómez-Robles, A., Prado-Simón, L. & Arsuaga, J. L. Morphological description and comparison of the dental remains from Atapuerca-Sima de los Huesos site (Spain). J. Hum. Evol. 62, 7–58 (2012).
Wu, X.-J. et al. Archaic human remains from Hualongdong, China, and Middle Pleistocene human continuity and variation. Proc. Natl Acad. Sci. USA 116, 9820–9824 (2019).
Bailey, S. E. A closer look at Neanderthal postcanine dental morphology: the mandibular dentition. Anat. Rec. 269, 148–156 (2002).
Weidenreich, F. The skull of Sinanthropus pekinensis: a comparative study on a primitive hominid skull. Paleontol. Sin. Ser. D 10, 43–52 (1943).
Wu, R., Wu, X. & Zhang, S. Early Humankind in China (Science Press, 1989).
Wu, X. Middle Pleistocene Human Skull from Dali, China (Science Press, 2020).
Bae, C. J. & Wu, X. Making sense of eastern Asian Late Quaternary hominin variability. Nat. Commun. 15, 9479 (2024).
Trinkaus, E. & Churchill, S. E. Neandertal radial tuberosity orientation. Am. J. Phys. Anthropol. 75, 15–21 (1988).
Carretero, J. M., Lorenzo, C. & Arsuaga, J. L. Axial and appendicular skeleton of Homo antecessor. J. Hum. Evol. 37, 459–499 (1999).
Rodríguez, L., García-González, R., Arsuaga, J. L. & Carretero, J.-M. Uncovering the adult morphology of the forearm bones from the Sima de los Huesos Site in Atapuerca (Spain), with comments on biomechanical features. Anat. Rec. 307, 2550–2574 (2024).
Kuhn, S. L. A geometric index of reduction for unifacial stone tools. J. Archaeol. Sci. 17, 583–593 (1990).
Xiao, X., Shen, J., Wang, S., Xiao, H. & Tong, G. Palynological evidence for vegetational and climatic changes from the HQ deep drilling core in Yunnan Province, China. Sci. China Ser. D Earth Sci. 50, 1189–1201 (2007).
An, Z. et al. Glacial-interglacial Indian summer monsoon dynamics. Science 333, 719–723 (2011).
Bradfield, J. & Langley, M. C. Bone tool diversity during the stone age: more insights into the human story. J. Archaeol. Res. 34, 151–205 (2026).
Richards, D. A. & Dorale, J. A. Uranium-series chronology and environmental applications of speleothems. Rev Mineral. Geochem. 52, 407–460 (2003).
Cheng, H. et al. Improvements in 230Th dating, 230Th and 234U half-life values, and U–Th isotopic measurements by multi-collector inductively coupled plasma mass spectrometry. Earth Planet. Sci. Lett. 371–372, 82–91 (2013).
Douville, E. et al. Rapid and accurate U–Th dating of ancient carbonates using inductively coupled plasma-quadrupole mass spectrometry. Chem. Geol. 272, 1–11 (2010).
Jin, M. et al. High-precision MC-ICP-MS static measurements of uranium isotopes using Faraday cups. Chin. Sci. Bull. 67, 2651–2661 (2022).
Shao, Q.-F. et al. Interactive programs of MC-ICPMS data processing for 230Th/U geochronology. Quat. Geochronol. 51, 43–52 (2019).
Holden, N. E. Total half-lives for selected nuclides. Pure Appl. Chem. 62, 941–958 (1990).
Goldstein, S. J., Murrell, M. T. & Janecky, D. R. Th and U isotopic systematics of basalts from the Juan de Fuca and Gorda Ridges by mass spectrometry. Earth Planet. Sci. Lett. 96, 134–146 (1989).
Huntley, D. J., Godfrey-Smith, D. I. & Thewalt, M. L. W. Optical dating of sediments. Nature 313, 105–107 (1985).
Wintle, A. G. Luminescence dating: laboratory procedures and protocols. Radiat. Meas. 27, 769–817 (1997).
Aitken, M. J. Thermoluminescence Dating (Academic Press, 1985).
Liritzis, I., Stamoulis, K., Papachristodoulou, C. & Ioannides, K. A re-evaluation of radiation dose-rate conversion factors. Mediterr. Archaeol. Archaeom. 13, 1–15 (2013).
Brennan, B. J. Beta doses to spherical grains. Radiat. Meas. 37, 299–303 (2003).
Bøtter-Jensen, L., Bulur, E., Duller, G. A. T. & Murray, A. S. Advances in luminescence instrument systems. Radiat. Meas. 32, 523–528 (2000).
Bøtter-Jensen, L., Andersen, C. E., Duller, G. A. T. & Murray, A. S. Developments in radiation, stimulation and observation facilities in luminescence measurements. Radiat. Meas. 37, 535–541 (2003).
Roberts, R. G., Galbraith, R. F., Olley, J. M., Yoshida, H. & Laslett, G. M. Optical dating of single and multiple grains of quartz from Jinmium rock shelter, northern Australia: part II, results and implications. Archaeometry 41, 365–395 (1999).
Murray, A. S. & Wintle, A. G. Luminescence dating of quartz using an improved single-aliquot regenerative-dose protocol. Radiat. Meas. 32, 57–73 (2000).
Li, B., Jacobs, Z. L., Roberts, R. G., Galbraith, R. F. & Peng, J. Variability in quartz OSL signals caused by measurement uncertainties: problems and solutions. Quat. Geochronol. 41, 11–25 (2017).
Li, B., Jacobs, Z. & Roberts, R. G. A Bayesian hierarchical age model for optical dating of single grains of quartz. Quat. Geochronol. 77, 101455 (2023).
Faegri, K. & Iversen, J. Textbook of Pollen Analysis (Wiley, 1989).
Grimm, E. C. CONISS: a FORTRAN 77 program for stratigraphically constrained cluster analysis by the method of incremental sum of squares. Comput. Geosci. 13, 13–35 (1987).
Prentice, C., Guiot, J., Huntley, B., Jolly, D. & Cheddadi, R. Reconstructing biomes from palaeoecological data: a general method and its application to European pollen data at 0 and 6 ka. Clim. Dyn. 12, 185–194 (1996).
Turner, C. G., Nichol, C. R. & Scott, G. R. in Advances in Dental Anthropology (eds Kelley, M. A. & Larsen, C. S.) 13–31 (Wiley-Liss, 1991).
Bône, A. Louis, M., Martin, B. & Durrleman, S. Deformetrica 4: an open-source software for statistical shape analysis. In Proc. International Workshop on Shape in Medical Imaging vol. 11167 (eds Reuter, M. et al.) 3–13 (Springer, 2018).
Jean, D. R Tools for Deformetrica: R script for Deformetrica ouput. R package version 0.1. gitlab.com https://gitlab.com/jeandumoncel/tools-for-deformetrica (2020).
Dray, S. & Dufour, A.-B. The ade4 package: implementing the duality diagram for ecologists. J. Stat. Softw. 22, 1–20 https://doi.org/10.18637/JSS.V022.I04 (2007).
Schlager, S. in Statistical Shape and Deformation Analysis (eds Zheng, G. et al.) 217–256 (Academic Press, 2017).
R Core Team. R: a language and environment for statistical computing. http://www.R-project.org/ (R Foundation for Statistical Computing, 2021).
Warnes, G., Bolker, B. & Lumley, T. gtools: various R programming tools. R package v.3.9.2. GitHub https://github.com/r-gregmisc/gtools (2021).
Schäfer, T. free surfer formats: Read and Write ‘Free Surfer’ Neuroimaging File Formats. R package v.0.1.15. GitHub https://github.com/dfsp-spirit/freesurferformats (2021).
Clay, R. B. Typological classification, attribute analysis, and lithic variability. J Field Archaeol. 3, 303–311 (1976).
Andrefsky, W. Jr. Lithics: Macroscopic Approaches to Analysis (Cambridge Univ. Press, 1998).
Tafelmaier, Y., Bataille, G., Schmid, V., Taller, A. & Will, M. in Methods for the Analysis of Stone Artefacts: An Overview (eds Tafelmaier, Y. et al.) 5-13 (Springer, 2023).
Sellet, F. Chaîne opératoire; the concept and its applications. Lithic Technol. 18, 106–112 (1993).
Debénath, A. & Dibble, H. L. Handbook of Paleolithic Typology: Lower and Middle Paleolithic of Europe (Univ. of Pennsylvania Museum of Archaeology, 1994).
Hiscock, P. & Attenbrow, V. J. Early Australian implement variation: a reduction model. J. Archaeol. Sci. 30, 239–249 (2003).
Anderson, M. J. A new method for non-parametric multivariate analysis of variance. Austral. Ecol. 26, 32–46 (2001).
Anderson, M. J. Distance-based tests for homogeneity of multivariate dispersions. Biometrics 62, 245–253 (2006).
R Core Team. R: a language and environment for statistical computing. https://www.R-project.org (R Foundation for Statistical Computing, 2024).
R Studio Team. RStudio: integrated development environment for R. RStudio http://www.rstudio.com (PBC, 2021).
Oksanen, J. et al. vegan: Community Ecology Package. R package v.2.6–4. https://doi.org/10.32614/CRAN.package.vegan (2024).
Pales, L., Lambert, C. & Garcia, M. A. Atlas Ostéologique Pour Servir à l’Identification des Mammifères du Quaternaire (CNRS, 1971).
O’Connor, T. The Archaeology of Animal Bones (Texas A&M Univ. Press, 2000).
Reitz, E. J. & Wing, E. S. Zooarchaeology 2nd edn (Cambridge Univ. Press, 2008).
France, D. L. Comparative Bone Identification: Human Subadult to Nonhuman (CRC Press, 2017).
Morin, E., Beauval, C., Boileau, A., Ready, E. & Laroulandie, V. The number of distinct elements: extending a landmark-based counting unit to other taxa. J. Archaeol. Sci. Rep. 24, 773–784 (2019).
Brain, C. K. The Hunters or the Hunted: An Introduction to African Cave Taphonomy (Univ. Chicago Press, 1983).
Discamps, E. & Costamagno, S. Improving mortality profile analysis in zooarchaeology: a revised zoning for ternary diagrams. J. Archaeol. Sci. 58, 62–76 (2015).
Villa, P. & Mahieu, E. Breakage patterns of human long bones. J. Hum. Evol. 21, 27–48 (1991).
Vettese, D. et al. A way to break bones? The weight of intuitiveness. PLoS ONE 16, e0259136 (2021).
Behrensmeyer, A. K. Taphonomic and ecologic information from bone weathering. Paleobiology 4, 150–162 (1978).
Fernández-Jalvo, Y. & Andrews, P. Atlas of Taphonomic Identifications, Vertebrate Paleobiology and Paleoanthropology. (Springer, 2016).
Fisher, J. W. Jr. Bone surface modifications in zooarchaeology. J. Archaeol. Method Theory 2, 7–68 (1995).
Cáceres, I. Tafonomía de Yacimientos Antrópicos en Karst. Complejo Galería (Sierra de Atapuerca, Burgos), Vanguard Cave (Gibraltar) y Abric Romani (Capellades, Barcelona). PhD dissertation, Univ. Rovira i Virgili, Tarragona (2002).
Lyman, R. L. Vertebrate Taphonomy (Cambridge Univ. Press, 1994).
Abrams, G. et al. Earliest evidence of Neanderthal multifunctional bone tool production from cave lion (Panthera spelaea) remains. Sci. Rep. 15, 24010 (2025).
Patou-Mathis, M. (ed.) Retouchoirs, Compresseurs, Percuteurs … Os à Impressions et Éraillures. In Fiches de la Commission de Nomenclature sur l’Industrie de l’Os Préhistorique. Cahier X (Éditions de la Société Préhistorique Française, 2002).
Selvaggio, M. M. & Wilder, J. Identifying the involvement of multiple carnivore taxa with archaeological bone assemblages. J. Archaeol. Sci. 28, 465–470 (2001).
Pobiner, B. Paleoecological information in predator tooth marks. J. Taphonomy 6, 373–397 (2008).
Andrés, M., Gidna, A. O., Yravedra, J. & Domínguez-Rodrigo, M. A study of dimensional differences of tooth marks (pits and scores) on bones modified by small and large carnivores. Archaeol. Anthropol. Sci. 4, 209–219 (2012).
Pei, W. The Upper Cave fauna of Choukoutien. Palaeontol. Sin. Ser. C 10, 1–101 (1940).
Soulier, M.-C. & Costamagno, S. Let the cutmarks speak! Experimental butchery to reconstruct carcass processing. J. Archaeol. Sci. Rep. 11, 782–802 (2017).
Costamagno, S., Soulier, M.-C., Val, A. & Chong, S. Le référentiel de stries de boucherie. Palethnologie 10, 195–291 (2019).