Generate QR code
Open Access
Subscription Access
The Dispersal of Denisovans and Neanderthals in China
https://doi.org/10.17746/1563-0110.2025.53.4.003-027
Abstract
The initial peopling of China by H. erectus occurred ~1.7 Ma BP, maybe even earlier. Despite climatic changes in the Pleistocene, ecological conditions for the permanent occupation of this territory by members of that taxon remained favorable, causing their continuous morphological and genetic evolution in situ along the line leading to H. sapiens. As a result, between 120–40 ka BP, they evolved convergently into early anatomically modern humans in this territory. A large part of the morphological variation of Late Middle to Early Upper Pleistocene hominins in China is attributable not only to the divergence of separate groups of H. erectus descendants, who convergently, under different ecological conditions, evolved into anatomically modern humans using various adaptive strategies, but also to the immigration of Denisovans and their assimilation of native groups. Contacts between the latter and Neanderthals in the second half of the Upper Pleistocene cannot be ruled out too.
About the Author
A. P. Derevianko
Institute of Archaeology and Ethnography, Siberian Branch, Russian Academy of Sciences; Altai State University
Russian Federation
Russian Federation
Derevianko A.P., Professor, Scientific Director; Laboratory Head
17, pr. Akademika Lavrentieva, Novosibirsk, 630090;
61, pr. Lenina Barnaul, 656049.
References
1. Ao Hong, Liu Chun-Ru, Roberts A.P., Zhang Peng, Xu Xinwen. 2017 An updated age for the Xujiayao hominin from the Nihewan Basin, North China: Implications for Middle Pleistocene human evolution in East Asia. Journal of Human Evolution, vol. 106: 54–65.
2. Bae Ch. 2010 The late Middle Pleistocene hominin fossil record of eastern Asia: Synthesis and review. Yearbook of Physical Anthropology, vol. 53: 75–93.
3. Bae Ch.J., Wu X.-J. 2024 Making sense of eastern Asian Late Quaternary hominin variability. Nature communication. doi:10.1038/s41467-024-53918-7
4. Bailey Sh., Glantz M., Weaver T.D., Viola B. 2008 The affi nity of the dental remains from Obi-Rakhmat grotto, Uzbekistan. Journal of Human Evolution, vol. 55: 238–248.
5. Bailey Sh.E., Liu W. 2010 A comparative dental metrical and morphological analysis of a Middle Pleistocene hominin maxilla from Chaoxian (Chaohu), China. Quaternary International, vol. 211: 14–23.
6. Bergström A., Stringer C., Hajdinjak M., Scerri E.M.L., Skoglund P. 2021 Origins of modern human ancestry. Nature, vol. 590 (7845): 229–237.
7. Bräuer G. 2001a The “Out-of-Africa” model and the question of regional continuity. In Humanity from African Naissance to Coming Millennia, P.V. Tobias et al. (eds). Johannesburg: Witwatersrand Univ. Press, pp. 183–189.
8. Bräuer G. 2001b The KNM-ER 3884 hominid and the emergence of modern anatomy in Africa. In Humanity from African Naissance to Coming Millennia, P.V. Tobias et al. (eds.). Firenze: Univ. Press, pp. 191–199.
9. Bräuer G. 2007 Origin of modern humans. In Handbook of Paleoanthropology. Vol. III: Phylogeny of Hominids, W. Henke, I. Tattersall (eds.). Berlin: Springer, pp. 1749–1779.
10. Bräuer G. 2008 The origin of modern anatomy: By speciation or intraspecific evolution? Evolutionary Anthropology, vol. 17 (1): 22–37.
11. Browning S.R., Browning B.L., Zhou Y., Tucci S., Akey M.J. 2018 Analysis of human sequence data reveals two pulses of archaic Denisovan admixture. Cell, vol. 173: 53–61.
12. Callaway E. 2024 How the mysterious Denisovans thrived on top of the World. Nature, vol. 631: 262–263.
13. Chang C.-H., Kaifu Y., Takai M., Kono R., Grün R., Matsuura S., Lin L.-K. 2015 The first archaic Homo from Taiwan. Nature Communications, vol. 6: 6037.
14. Chen F.-H., Frido W., Shen C.-C., Bailey S.E., Bergmann I., Davis S., Xia H., Wang H., Fischer R., Freidline S.E., Yu T.L., Skinner M.M., Stelzer S., Dong Guangrong, Fu Q., Dong Guanghui, Wang J., Zhang D., Hublin J.- J. 2019 A late Middle Pleistocene Denisovan mandible from the Tibetan Plateau. Nature, vol. 569: 409–412.
15. Chen T.M., Yang Q., Wu E. 1994 Antiquity of Homo sapiens in China. Nature, vol. 368: 55–56.
16. Chen T.M., Yuan S., Gao S. 1984 Using the uranium method to investigate important Palaeolithic dates in northern China. Acta Anthropologica Sinica, vol. 3 (3): 259–269.
17. Chen T.M., Yuan S., Gao S., Wang L., Zhao G. 1982 Uranium-series dating of Xujiayao (Hsu-Chia-Yao) site. Acta Anthropologica Sinica, vol. 1: 91–95.
18. Chen T.M., Yuan S.X. 1988 Uranium-series dating of bones and teeth from Chinese Palaeolithic sites. Archaeometry, vol. 30 (1): 59–76.
19. Chen T.M., Yuan S.X., Gao S.J., Hu Y.Q. 1987 Uranium-series dating of fossil bones from Hexian and Chaoxian human fossil sites. Acta Anthropologica Sinica, vol. 6 (3): 249–254. (In Chinese with English abstract).
20. Chia L.P., Wei Q. 1976 A Palaeolithic site at Hsue-chia-yao in Yangkao County, Shansi Province. Acta Archaeologica Sinica, vol. 2: 97–114.
21. Chia L.P., Wei Q., Li C. 1979 Report on the excavation of Hsue-chia-yao man site in 1976. Vertebrata PalAsiatica, vol. 17 (4): 277–293.
22. Demeter F., Zanolli C., Westaway K.E., Joannes-Boyau R., Duringer P., Morley M.W., Welker F., Rüther P.L., Skinner M.M., McColl H., Gaunitz C., Vinner L., Dunn T.E., Olsen J.V., Sikora M., Ponche J.-L., Suzzoni E., Frangeul S., Boesch Q., Antoine P.-O., Pan L., Xing S., Zhao J.-X., Bailey R.M., Boualaphane S., Sichanthongtip P., Sihanam D., Patole-Edoumba E., Aubaile F., Crozier F., Bourgon N., Zachwieja A., Luangkhoth T., Souksavatdy V., Sayavongkhamdy T., Cappellini E., Bacon A.-M., Hublin J.-J., Willerslev E., Shackelford L. 2022 A Middle Pleistocene Denisovan molar from the Annamite Chain of northern Laos. Nature Communications, vol. 13 (1). doi:10.1038/s41467-022-29923-z
23. Dennell R.W. 2020 From Arabia to the Pacifi c: How our Species Colonised Asia. [S.l.]: Routledge.
24. Derevianko A.P. 2001 The Middle to Upper Paleolithic transition in the Altai (Mongolia and Siberia). Archaeology, Ethnology and Anthropology of Eurasia, No. 3: 70–103. (In Russian and English).
25. Derevianko A.P. 2011 The Upper Paleolithic in Africa and Eurasia and the Origin of Anatomically Modern Humans. Novosibirsk: Izd. IAET SO RAN. (In Russian and English).
26. Derevianko A.P. 2014 Bifacial Industry in East and Southeast Asia. Novosibirsk: Izd. IAET SO RAN. (In Russian and English).
27. Derevianko A.P. 2019 Three Global Human Migrations in Eurasia. Vol. IV: The Acheulean and Bifacial Lithic Industries in China, Korea, Mongolia, Kazakhstan, Turkmenistan, Uzbekistan and in the Caucasus. Novosibirsk: Izd. IAET SO RAN. (In Russian and English).
28. Derevianko A.P. 2022 Three Global Human Migrations in Eurasia, vol. VI. Pt. 1: Denisovan Human: Origins, Material and Spiritual Culture. Novosibirsk: Izd. IAET SO RAN. (In Russian and English).
29. Derevianko A.P. 2024a Origin of Neanderthals. Neanderthals of the Altai: Myth or reality? Archaeology, Ethnology and Anthropology of Eurasia, vol. 52 (1): 3–34. (In Russian and English).
30. Derevianko A.P. 2024b The peopling of Tajikistan by Homo sapiens denisovan. Archaeology, Ethnology and Anthropology of Eurasia, vol. 52 (4): 3–28. (In Russian and English).
31. Derevianko A.P. 2024c Three Global Human Migrations in Eurasia, vol. VI. Pt. 2: Dispersal of Denisovans and Neanderthals in Southern Siberia. Novosibirsk: Izd. IAET SO RAN. (In Russian and English).
32. Derevianko A.P. 2024d Who were the Chagyrskaya Neanderthals? The possibility of their dispersal across Central Asia and Southern Siberia. Archaeology, Ethnology and Anthropology of Eurasia, vol. 52 (2): 3–19. (In Russian and English).
33. Derevianko A.P. 2024e The origin of H. s. denisovan and their dispersal across Iran. Archaeology, Ethnology and Anthropology of Eurasia, vol. 52 (3): 3–18. (In Russian and English).
34. Derevianko A.P. 2025a The peopling of Uzbekistan by Homo sapiens denisovan. Archaeology, Ethnology and Anthropology of Eurasia, vol. 53 (1): 3–24. (In Russian and English).
35. Derevianko A.P. 2025b The peopling of Mongolia by Homo sapiens denisovan. Archaeology, Ethnology and Anthropology of Eurasia, vol. 53 (2): 3–28. (In Russian and English).
36. Derevianko A.P. 2025c The Origin of Anatomically Modern Humans in China. Archaeology, Ethnology and Anthropology of Eurasia, vol. 53 (3): 107–130. (In Russian and English).
37. Derevianko A.P. 2025d Three Global Human Migrations in Eurasia, vol. VII. Dispersal of Denisovans in Southern Siberia. Novosibirsk: Izd. IAET SO RAN. (In Russian and English).
38. Derevianko A.P., Markin S.V. 1992 Mustye Gornogo Altaya (po materialam peshchery im. Okladnikova). Novosibirsk: Nauka.
39. Derevianko A.P., Markin S.V., Kolobova K.A., Chabai V.P., Rudaya N.A., Buzhilova A.P., Mednikova M.B., Vasiliev S.K., Zykin V.S., Zykina V.S., Zazhigin V.S., Volvakh A.O., Roberts R.G., Jacobs Z., Bo Li. 2018 Mezhdistsiplinarniye issledovaniya Chagyrskoy peshchery – stoyanki srednego paleolita Altaya. Novosibirsk: Izd. IAET SO RAN.
40. Derevianko A.P., Shunkov M.V., Agadjanian A.K., Baryshnikov G.F., Ulyanov V.A., Kulik N.A., Postnov A.V., Anoikin A.A. 2003 Prirodnaya sreda i chelovek v paleolite Gornogo Altaya. Usloviya obitaniya v okrestnostyakh Denisovoy peshchery. Novosibirsk: IAET SO RAN.
41. Derevianko A.P., Shunkov M.V., Kozlikin M.B. 2020 Who were the Denisovans? Archaeology, Ethnology and Anthropology of Eurasia, vol. 48 (3): 3–32. (In Russian and English).
42. Derevianko A.P., Shunkov M.V., Kozlikin M.B., Fedorchenko A.Y., Pavlenok G.D., Belousova N.E. 2016 Kostyanaya igla nachala verkhnego paleolita iz tsentralnogo zala Denisovoy peshchery (po materialam raskopok v 2016 godu). In Problemy arkheologii, etnografi i, antropologii Sibiri i sopredelnykh territoriy, vol. XXII. Novosibirsk: Izd. IAET SO RAN, pp. 72–75.
43. Derevianko A.P., Shunkov M.V., Tsybankov A.A., Ulyanov V.A., Volkov P.V. 2008 Izucheniye verkhnepaleoliticheskikh sloev v vostochnoy galereye Denisovoy peshchery. In Problemy arkheologii, etnografi i, antropologii Sibiri i sopredelnykh territoriy, vol. XIV. Novosibirsk: Izd. IAET SO RAN, pp. 60–65.
44. Douka K., Slon V., Jacobs Z., Ramsey C.B., Shunkov M.V., Derevianko A.P., Mafessoni F., Kozlikin M.B., Li B., Grün R., Comeskey D., Devièse T., Brown S., Viola B., Kinsley L., Buckley M., Meyer M., Roberts R.G., Pääbo S., Kelso J., Higham T. 2019 Age estimates for hominin fossils and the onset of the Upper Palaeolithic at Denisova Cave. Nature, vol. 565: 640–644.
45. Feng X., Yin Q., Gao F., Lu D., Fang Q., Fen Y., Huang X., Chen T., Zhou H., Li Q., Zhang Chi, Stringer C., Ni X. 2025 The phylogenetic position of the Yunxian cranium elucidates the origin of Homo longi and the Denisovans. Science, vol. 389 (6767): 1320–1324. doi:10.1126/science.ado9202
46. Foley R., Lahr M.M. 2003 On stony ground: Lithic technology, human evolution, and the emergence of culture. Evolutionary Anthropology, vol. 12 (3): 109–122.
47. Fu Q., Li H., Moorjani P., Jay F., Slepchenko S.M., Bondarev A.A., Johnson P.L.F., Aximu-Petri A., Prüfer K., Filippo C., de, Meyer M., Zwyns N., Salazar-Garcira D.C., Kuzmin Y.V., Keates S.G., Kosintsev P.A., Razhev D.I., Richards M.P., Peristov N.V., Lachmann M., Douka K., Higham T.F.G., Slatkin M., Hublin J.-J., Reich D., Kelso J., Viola B., Pääbo S. 2014 Genome sequence of a 45,000-year-old modern human from Western Siberia. Nature, vol. 514: 445–450.
48. Gao B., Shen G.J., Qiu L.C. 2007 Preliminary U-series dating of southern branch cave of Maba hominid site. Journal of Jinan University. Natural Science, vol. 28: 308–311.
49. Gao X., Zhang X., Yang D., Shen C., Wu X. 2010 Revisiting the origin of modern humans in China and its implication for global human evolution. Science China, Earth Sciences, vol. 53: 1927–1940.
50. Glantz M., Viola B., Chikisheva T.A. 2004 Noviye ostanki gominina iz grota Obi-Rakhmat. In Grot Obi-Rakhmat. Novosibirsk: Izd. IAET SO RAN, pp. 75–92.
51. Glantz M., Viola B., Wrinn P., Chikisheva T., Derevianko A., Krivoshapkin A., Islamov U., Suleimanov I., Ritzman T. 2008 New hominin remains from Uzbekistan. Journal of Human Evolution, vol. 55 (2): 223–237.
52. Goren-Inbar N., Alperson-Afi l N., Sharon G., Herzlinger G. 2018 The Acheulian Site of Gesher Benot Ya’aqov. Vol. 4: The Lithic Assemblages. Dordrecht: Springer. (Vertebr. Paleobiol. and Paleoanthropol. Ser.).
53. Grot Obi-Rakhmat. 2004 Novosibirsk: Izd. IAET SO RAN.
54. Groves C.P. 1994 The origin of modern humans. Interdisciplinary Science Reviews, vol. 19 (1): 23–34.
55. Grün R., Huang P.H., McDermott W.P., Thorne F., Stringer C.B., Yan G. 1998 ESR and U-series analyses of teeth from the palaeoanthropological site of Hexian, Anhui Province, China. Journal of Human Evolution, vol. 34 (6): 555–564.
56. Howell F.C. 1999 Paleo-demes, species clades, and extinctions in the Pleistocene hominin record. Journal of Anthropological Research, vol. 55: 191–243.
57. Huang W., Ciochon R., Gu Y.M., Larick R., Fang Q.R., Schwarcz H., Yonge C., Vos J., de, Rink W. 1995 Early Homo and associated artefacts from Asia. Nature, vol. 378 (6554): 275–278.
58. Huang W., Otte M., Hu Y., Hou Y. 2015 Panxian Dadong and the Chinese Levallois. In The 20th (2) International Symposium of Celebration of the 30th Anniversary of the 1st Site Excavation: Suyanggae and Her Neighbors in Korea. [S.n.]: Inst. of Korean Prehistory, pp. 69–92.
59. Hublin J.-J. 2001 Northwestern African Middle Pleistocene hominids and their bearing on the emergence of Homo sapiens. In Human Roots. Africa and Asia in the Middle Pleistocene. Bristol: West. Acad. and Spec. Press, pp. 99–121.
60. Hublin J.-J. 2013 The Middle Pleistocene record. On the origin of Neanderthals, modern humans and others. In A Companion to Paleoanthropology, D. Begun (ed.). Hoboken: WileyBlackwell, pp. 517–537.
61. Huerta-Sánchez E., Jin X., Asan, Bianba Z., Peter B.M., Vinckenbosch N., Liang Y., Yi X., He M., Somel M., Ni P., Wang B., Ou X., Huasang, Luosang J., Cuo Z.X.P., Li K., Gao G., Yin Y., Wang W., Zhang X., Xu X., Yang H., Li Y., Wang Jian, Wang Jun, Nielsen R. 2014 Altitude adaptation in Tibetans caused by introgression of Denisovan-like DNA. Nature, vol. 512: 194–197.
62. Hun L.Y., Xu Q.H., Zhang S.R., Wang F.G., Liu L.Q., Li J.Y., Li M.Y. 2011 Paleoenvironment and paleoclimate changes of Houjiayao site in Yangyuan county, Hebei province based on pollen analysis. Quaternary Science, vol. 31 (6): 951–961.
63. Jacobs Z., Li B., Shunkov M.V., Kozlikin M.B., Bolikhovskaya N.S., Agadjanian A.K., Uliyanov V.A., Vasiliev S.K., O’Gorman K., Derevianko A.P., Roberts R.G. 2019 Timing of archaic hominin occupation of Denisova Cave in southern Siberia. Nature, vol. 565: 594–599.
64. Jacobs Z., Zavala E.I., Li B., O’Gorman K., Shunkov M.V., Kozlikin M.B., Derevianko A.P., Uliyanov V.A., Goldberg P., Agadjanian A.K., Vasiliev S.K., Brink F., Peyrégne S., Slon V., Pääbo S., Kelso J., Meyer M., Roberts R.G. 2025 Pleistocene chronology and history of hominins and fauna at Denisova Cave. Nature Communications, vol. 16 (1). doi:10.1038/s41467-025-60140-6
65. Ji Q., Wu W., Ji Y., Li Q., Ni X. 2021 Late middle Pleistocene Harbin cranium represents a new Homo species. The Innovation, vol. 2 (3). doi:10.1016/j.xinn.2021.100132
66. Kaifu Y. 2006 Advanced dental reduction in Javanese Homo erectus. Anthropological Science, vol. 114 (1): 35–43.
67. Kaifu Y., Baba H., Aziz F., Indriati E., Schrenk F., Jacob T. 2005 Taxonomic affinities and evolutionary history of the early Pleistocene hominins of Java: Dentognathic evidence. American Journal of Physical Anthropology, vol. 128 (4): 709–726.
68. Keates S.G. 2000 Early and Middle Pleistocene Hominid Behaviour in Northern China. Oxford: BAR. (BAR Intern. Ser.; No. 863).
69. Kozlikin M.B. 2017 Paleoliticheskiye kompleksy vostochnoy galerei Denisovoy peshchery. Cand. Sc. (History) Dissertation. Novosibirsk.
70. Krause J., Fu Q., Good J.M., Viola B., Shunkov M.V., Derevianko A.P., Pääbo S. 2010 The complete mitochondrial DNA genome of an unknown hominin from Southern Siberia. Nature, vol. 464: 894–897.
71. Le site de Lʼhomme de Yunxian. Quyuanhekou, Quingqu, Yunxian, Province du Hubei. 2008 Paris: CNRS Ed.
72. Li F., Kuhn S.L., Chen F., Wang W., Southon I., Peng F., Shan M., Wang C., Ge I., Wang X., Yun T., Gao X. 2018 The easternmost Middle Paleolithic (Mousterian) from Jinsitai Cave, North China. Journal of Human Evolution, vol. 114: 76–84.
73. Li Tianyuan, Ai Dan, Feng Xiaobo. 1996 Povtornoye obsuzhdeniye morfologicheskikh kharakteristik cherepa yunsyanskogo cheloveka. Jianghan kaogu, No. 1: 40–44. (In Chinese).
74. Li Tianyuan, Wang Zhenghua, Li Wensen, Feng Xiaobo, Wu Xianzhu. 1994 Morfologicheskiye kharakteristiki cherepa gominida iz Tsyuiyuanhekou uyezda Hubei i yego mesto v evolyutsionnom protsesse. Renleixue xuebao, No. 13 (2): 104–116. (In Chinese).
75. Li Z.-Y., Wu X.-J., Zhou L.-P., Liu W., Gao X., Nian X.-M., Trinkaus E. 2017. Late Pleistocene archaic human crania from Xuchang, China. Science, vol. 355 (6328): 969–972.
76. Liu C., Su P., Jin Z.X. 1992 Discovery of Blake Episode in the Xujiayao Paleolithic site, Shanxi, China. Scientia Geologica Sinica, vol. 1: 87–95.
77. Liu W., Jin C.-Z., Zhang Y.-Q., Cai Y.-J., Xing S., Wu X.-J., Cheng H., Edwards R.L., Pan W.-S., Qin D.-G., An Z.S., Trinkaus E., Wu X.-Z. 2010a Human remains from Zhirendong, south China and modern human emergence in East Asia. PNAS, vol. 107: 19201–19206.
78. Liu W., Schepartz L.A., Xing S., Miller-Antonio S., Wu X., Trinkaus E., Martinón-Torres M. 2013 Late Middle Pleistocene hominin teeth from Panxian Dadong, South China. Journal of Human Evolution, vol. 67: 337–355.
79. Liu W., Si X. 1997 The human teeth discovered in Dadong, Panxian County, Guizhou Province. Acta Anthropologica Sinica, vol. 16: 193–200.
80. Liu W., Wu X.-Z., Pei S.-W., Wu X.-J., Norton C.-J. 2010b Huanglong cave: A late Pleistocene human fossil site in Hubei Province, China. Quaternary International, vol. 211: 29–41.
81. Martinón-Torres M., Wu X., Bermudes de Castro J.M., Xing S., Liu W. 2017 Homo sapiens in the Eastern Asia Late Pleistocene. Current Anthropology, vol. 17 (Suppl.): S444–S448.
82. Meyer M., Arsuaga J.-L., Filippo C., de, Nagel S., Aximu-Petri A., Nickel B., Martinez I., Gracia A., Bermudez de Castro J.M., Carbonell E., Viola B., Kelso J., Prüfer K., Pääbo S. 2016 Nuclear DNA sequences from the Middle Pleistocene Sima de los Huesos Hominins. Nature, vol. 531: 504–507.
83. Meyer M., Fu Q., Aximu-Petri A., Glocke I., Nickel B., Arsuaga J.-L., Martínez I., Gracia A., Bermudez de Castro J.M., Carbonell E., Pääbo S. 2014 A mitochondrial genome sequence of a hominin from Sima de los Huesos. Nature, vol. 505: 403–406.
84. Meyer M., Kircher M., Gansauge M.-T., Li H., Racimo F., Mallick S., Schraiber J.G., Jay F., Prüfer K., Filippo C., de, Sudmant P.H., Alkan C., Fu Q., Do R., Rohland N., Tandon A., Siebauer M., Green R.E., Bryc K., Briggs A.W., Stenzel U., Dabney J., Shendure J., Kitzman J., Hammer M.F., Shunkov M.V., Derevianko A.P., Patterson N., Andrés A.M., Eichler E.E., Slatkin M., Reich D., Kelso J., Pääbo S. 2012 A high-coverage genome sequence from an archaic Denisovan individual. Science, vol. 338 (6104): 222–226.
85. Miller-Antonio S., Shepartz L.A., Karkanas P., Hou Y., Wei W., Bekken D. 2004 Lithic raw material use at the Late Middle Pleistocene site of Panxian Dadong. Asian Perspectives, vol. 43: 314–332.
86. Nagatomo T., Shitaoka Y., Namioka H., Sagawa M., Wei Q. 2009 OSL dating of the strata at Palaeolithic sites in the Nihewan Basin, China. Acta Anthropologica Sinica, vol. 28: 276–284.
87. Ni Xijun, Ji Qiang, Wu Wensheng, Shao Qingfeng, Ji Yannan, Zhang Chi, Liang Lei, Ge Junyi, Guo Zhen, Li Jinhua, Li Qiang, Grün R., Stringer C. 2021 Massive cranium from Harbin in northeastern China establishes a new Middle Pleistocene human lineage. The Innovation, vol. 2 (3). doi:10.1016/j.xinn.2021.100130
88. Okladnikov A.P. 1949 Issledovaniye mustyerskoy stoyanki i pogrebeniya neandertaltsa v grote Teshik-Tash, Yuzhniy Uzbekistan. In Teshik-Tash. Paleoliticheskiy chelovek. Moscow: Izd. Mosk. Gos. Univ., pp. 7–87.
89. Ongaro L., Huerta-Sánchez E. 2024 A history of multiple Denisovan introgression events in modern humans. Nature Genetics, vol. 56: 2612–2622.
90. Pope G.G. 1992 Craniofacial evidence for the origin of modern humans in China. American Journal of Physical Anthropology, vol. 35: 243–298.
91. Prüfer K., Racimo F., Patterson N., Jay F., Sankararaman S., Sawyer S., Heinze A., Renaud G., Sudmant P.H., Filippo C., de, Heng Li, Mallick S., Dannemann M., Qiaomei Fu, Kircher M., Kuhlwilm M., Lachmann M., Meyer M., Ongyerth M., Siebauer M., Theunert C., Tandon A., Moorjani P., Pickrell J., Mullikin J.C., Vohr S.H., Green R.E., Hellmann I., Johnson Ph.L.F., Blanche H., Cann H., Kitzman J.O., Shendure J., Eichler E.E., Lein E.S., Bakken T.E., Golovanova L.V., Doronichev V.B., Shunkov M.V., Derevianko A.P., Viola B., Slatkin M., Reich D., Kelso J., Pääbo S. 2014 The complete genome sequence of a Neanderthal from the Altai Mountains. Nature, vol. 505 (7481): 43–49.
92. Qiu Zhonglang. 1989 Srednepaleoliticheskaya kultura v Kitaye. In Pervobytniy chelovek v Kitaye (Zhonggo yuanshide renlei). Beijing: Kexue chubanshe, pp. 195–220. (In Chinese).
93. Razhev D.I., Vasilyev S.V., Korost D.V., Borutskaya S.B. 2024 Bony labyrinth in Upper Paleolithic individuals buried at Sungir. Archaeology, Ethnology and Anthropology of Eurasia, vol. 52 (3): 118–126. (In Russian and English).
94. Reich D., Green R.E., Kircher M., Krause J., Patterson N., Durand E.Y., Viola B., Briggs A.W., Stenzel U., Johnson P.L.F., Maricic T., Good J.M., Marques-Bonet T., Alkan C., Fu Q., Mallick S., Li H., Meyer M., Eichler E.E., Stoneking M., Richards M., Talamo S., Shunkov M.V., Derevianko A.P., Hublin J.-J., Kelso J., Slatkin M., Pääbo S. 2010 Genetic history of an archaic hominin group from Denisova Cave in Siberia. Nature, vol. 468: 1053–1060.
95. Rightmire G.Ph. 1996 The human cranium from Bodo, Ethiopia: Evidence for speciation in the Middle Pleistocene? Journal of Human Evolution, vol. 31: 21–39.
96. Rightmire G.Ph. 1998a Evidence from facial morphology for similarity of Asian and African representatives of Homo erectus. American Journal of Physical Anthropology, vol. 106: 61–85.
97. Rightmire G.Ph. 1998b Human evolution in the Middle Pleistocene: The role of Homo heidelbergensis. Evolutionary Anthropology, vol. 6: 218–227.
98. Rightmire G.Ph. 2001a Diversity in the earliest “modern” populations from South Africa, Northern Africa and Southwest Asia. In Humanity from African Naissance to Coming Millennia, P.V. Tobias et al. (eds.). Florence: Firenze Univ. Press, pp. 231–236.
99. Rightmire G.Ph. 2001b Comparison of Middle Pleistocene hominids from Africa and Asia. In Human Roots: Africa and Asia in the Middle Pleistocene, L. Barham, K. Robson-Brown (eds.). Bristol: West. Acad. and Spec. Press, pp. 123–133.
100. Rightmire G.Ph. 2004a Affinities of the Middle Pleistocene crania from Dali and Jinniushan. American Journal of Physical Anthropology, vol. 38 (Suppl.): 167.
101. Rightmire G.Ph. 2004b Brain size and encephalization in early to Mid-Pleistocene Homo. American Journal of Physical Anthropology, vol. 124 (2): 109–123.
102. Rightmire G.Ph. 2008 Homo in the Middle Pleistocene: Hypodigms, variation, and species recognition. Evolutionary Anthropology, vol. 17 (1): 8–21.
103. Sawyer S., Renaud G., Viola B., Hublin J.-J., Gansauge M.-T., Shunkov M.V., Derevianko A.P., Prüfer K., Kelso J., Pääbo S. 2015 Nuclear and mitochondrial DNA sequences from two Denisovan individuals. PNAS, vol. 112 (51): 15696–15700.
104. Shen G.J., Tu H., Xiao D.F., Qiu L.C., Feng Y.X., Zhao J.X. 2014 Age of Maba hominin site in southern China: Evidence from U-series dating of Southern Branch Cave. Quaternary Geochronology, vol. 23: 56–62.
105. Shoaee M.J., Breeze P.S., Drake N.A., Hashemi S.M., Nasab H.V., Beifenback S.F.M., Stevens T., Bevin N., Petraglia M.D. 2023 Defining paleoclimatic routes and opportunities for hominin dispersals across Iran. PLoS ONE: 1–31.
106. Shoaee M.J., Nasab H., Petraglia M.D. 2021 The Paleolithic of the Iranian Plateau: Hominin occupation history and implication for human dispersals across southern Asia. Journal of Anthropological Archaeology, vol. 62: 1–30.
107. Silaev V.I., Slepchenko S.M., Bondarev A.A., Smoleva I.V., Kiseleva D.V., Shanina S.N., Martirosyan O.V., Tropnikov E.M., Khazov A.F. 2017 Ust-ishimskaya kost: Mineralogo-geokhimicheskiye svoistva kak istochnik paleontologicheskoy, paleoantropologicheskoy i paleoekologicheskoy informatsii. Vestnik Permskogo universiteta. Geologiya, vol. 16 (1): 1–25.
108. Skoglund P., Jakobsson M. 2011 Archaic human ancestry in East Asia. PNAS, vol. 108: 18301–19306.
109. Slon V., Hopfe C., Weiß C.L., Mafessoni F., Rasilla M., de la, Lalueza-Fox C., Rosas A., Soressi M., Knul M.V., Miller R., Stewart J.R., Derevianko A.P., Jacobs Z., Li B., Roberts R.G., Shunkov M.V., Lumley H., de, Perrenoud C., Gušić I., Kućan Ž., Rudan P., Aximu-Petri A., Essel E., Nagel S., Nickel B., Schmidt A., Prüfer K., Kelso J., Burbano H.A., Pääbo S., Meyer M. 2017 Neandertal and Denisovan DNA from Pleistocene sediments. Science, vol. 356 (6338): 605–608.
110. Slon V., Mafessoni F., Vernot B., Filippo C., de, Grote S., Viola B., Hajdinjak M., Peyrégne S., Nagel S., Brown S., Douka K., Higham T., Kozlikin M.B., Shunkov M.V., Derevianko A.P., Kelso J., Meyer M., Prüfer K., Pääbo S. 2018 The genome of the offspring of a Neanderthal mother and a Denisovan father. Nature, vol. 561 (7721): 113–116.
111. Stringer Ch.B. 1983 Some further notes on the morphology and dating of the Petralona hominid. Journal of Human Evolution, vol. 12 (8): 731–742.
112. Stringer Ch.B. 2002 Modern human origins: Progress and prospects. Philosophical Transactions of the Royal Society. Ser. B, vol. 357: 563–579.
113. Tattersall I., Schwartz J.H. 2000 Extinct Humans. Boulder: Westview Press.
114. Tu H., Shen G., Li H., Xie F., Granger D.E. 2015 26Al/10Be burial dating of Xujiayao-Houjiayao Site in Nihewan Basin, Northern China. PLoS ONE. URL: https://doi.org/10.1371/journal.pone.0118315
115. Vincenzo F., Manzi G. 2023 Homo heidelbergensis as Middle Pleistocene common ancestor of Denisovans, Neanderthals and modern humans. Journal of Mediterranean Earth Sciences, vol. 15: 303–315.
116. Viola B.T., Gunz P., Neubauer S., Slon V., Kozlikin M.B., Shunkov M.V., Meyer S., Pääbo S., Derevianko A.P. 2019 A parietal fragment from Denisova cave. American Journal of Physical Anthropology, vol. 168: 258.
117. Viola B., Seidler H., Nadden D. 2004 Izucheniye verkhnikh kraev pyramid visochnykh kostey OR-1 s pomoshchyu kompyuternoy grafiki. In Grot Obi-Rakhmat. Novosibirsk: Izd. IAET SO RAN, pp. 100–106.
118. Wall J.D., Yang M.A., Yay F., Kim S.K., Durand E.Y., Stevison L.S., Gignoux C., Woerner A., Hammer M.F., Slatkin M. 2013 Higher levels of Neanderthal ancestry in East Asians than in Europeans. Genetics, vol. 194: 199–209.
119. Wang X., Wei J., Chen Q., Tang Z., Wang C. 2010 A preliminary study on the excavation of the Jinsitai Cave site. Acta Anthropologica Sinica, vol. 29: 15–32. (In Chinese).
120. Wei Qi. 2004 Paleolit kotloviny Nikhevan. In Jiushiqi shidai kaogu juan: Zhonggo kaoguxue yanjiude shiji huigu [A CenturyLong Retrospective of Chinese Archaeology: Archaeology of the Paleolithic]. Beijing: Kexue chubanshe, pp. 83–110. (In Chinese).
121. Weidenreich F. 1947 Facts and speculations concerning the origin of Homo sapiens. American Anthropology, vol. 49: 187–203.
122. Wolpoff M.H. 1999 Paleoanthropology. Boston: McGraw-Hill.
123. Woo R.K., Peng R.C. 1959 Fossil human skull of early paleoanthropic stage found at Mapa, Shaoquan, Kwantung Province. Vertebrata Palasiatica, vol. 3: 176–182.
124. Wu M. 1984 New discovery of human fossil in Tongzi, Guizhou. Acta Anthropologica Sinica, vol. 3: 195–201.
125. Wu Maolin. 1986 Issledovaniye visochnoy kosti cheloverka iz Syuitszyayao. Renleixue xuebao, No. 5: 220–226. (In Chinese).
126. Wu M., Wang L., Zhang Y. 1975 The human fossils and associated cultural relics found in Tongzi, Guizhou. Vertebrata Palasiatica, vol. 13: 14–53.
127. Wu Xinzhi. 1988 O proiskhozhdenii sovremennykh kitaitsev po kraniologicheskim i odontologicheskim kharakteristikam pozdnikh Homo sapiens Kitaya. Renleixue xuebao, No. 17 (4): 276–288. (In Chinese).
128. Wu X. 2006 Evidence of multiregional human evolution hypothesis from China. Quaternary Sciences, No. 26 (5): 702–709. (In Chinese).
129. Wu X., Athreya S. 2013 A description of the geological context, discrete traits, and linear morphometrics of the Middle Pleistocene hominin from Dali, Shaanxi Province, China. American Journal of Physical Anthropology, vol. 150: 141–157.
130. Wu X.-J., Bae C.J. 2024 Xujiayao Homo: A new form of large brained hominin in Eastern Asia. PaleoAnthropology. Special Issue: What’s in a Name? Late Middle and Early Late Pleistocene Hominin Systematics. URL: https://doi.org/10.48738/202x.issx.xxx
131. Wu X.-J., Bae C.J., Friess M., Xing S., Athreya Sh., Liu W. 2022 Evolution of cranial capacity revisited: A view from the late Middle Pleistocene cranium from Xujiayao, China. Journal of Human Evolution, vol. 163: 1–11.
132. Wu X.-J., Crevecoeur I., Liu W., Xing S., Trinkaus E. 2014 Temporal labyrinths of eastern Eurasian Pleistocene humans. PNAS, vol. 111: 11509– 11513.
133. Wu X.-J., Maddux S.D., Pan L., Trinkaus E. 2012 Nasal floor variation among eastern Eurasian Pleistocene Homo. Anthropological Science, vol. 20: 217–226.
134. Wu X.-J., Pei S.-W., Cai Y.-J., Tong H.-W., Li Q., Dong Z., Cheng J.-C., Jin Z.-T., Ma D.-D., Xing S., Li X.-I., Cheng X., Cheng H., de la Torrek I., Edwards R.L., Gong X.-C., An Z.-S., Trinkaus E., Liu W. 2019 Archaic human remains from Hualongdong, China, and Middle Pleistocene human continuity and variation. PNAS, vol. 116: 9820–9824.
135. Wu X.-J., Pei Sh., Cai Y., Tong H., Zhang Z., Yan Y., Xing S., Martinón-Torres M., Bermúdes de Castro J.M., Liu W. 2023 Morphological and morphometric analyses of a late Middle Pleistocene hominin mandible from Hualongdong, China. Journal of Human Evolution, vol. 182. doi:10.1016/ jhevol.2023.103411
136. Wu X.-J., Trinkaus E. 2014 The Xujiayao 14 mandibular ramus and Pleistocene Homo mandibular variation. Comptes Rendus Palevol, vol. 13: 333–341.
137. Wu X.-J., Xing S., Trinkaus E. 2013 An enlarged parietal foramen in the late Archaic Xujiayao 11 neurocranium from northern China, and rare anomalies among Pleistocene Homo. PLoS ONE. URL: https://doi.org/10.1371/journal.pone.0059587
138. Wu X.Z. 1988 Comparative study of early Homo sapiens from China and Europe. Acta Anthropologica Sinica, vol. 7: 292–299.
139. Wu X.Z. 2004 On the origin of modern humans in China. Quaternary International, vol. 117: 131–140.
140. Wu X.Z., Poirier F.E. 1995 Human Evolution in China. A Morphometric Description of the Fossils and a Review of the Sites. New York: Oxford Univ. Press.
141. Xia H., Zhang D., Wang J., Fagernäs Z., Li T., Li Y., Yao J., Lin D., Troché G., Smith G.M., Chen X., Cheng T., Shen X., Han Y., Olsen J.V., Shen Z., Pei Z., Hublin J.-J., Chen F., Welker F. 2024 Middle and Late Pleistocene Denisovan subsistence at Baishiya Karst Cave. Nature, vol. 632: 108–113. URL: https://doi.org/10.1038/s41586-024-07612-9
142. Xing S., Martinón-Torres M., Bermudes de Castro J.M. 2019a Late middle Pleistocene hominin teeth from Tongzi, southern China. Journal of Human Evolution, vol. 130: 96–108.
143. Xing S., Martinón-Torres M., Bermudes de Castro J.M., Wu X.-J., Liu W. 2015 Hominin teeth from the early Late Pleistocene site of Xujiayao, Nothern China. American Journal of Physical Anthropology, vol. 156 (2): 224–240.
144. Xing S., Tafforeau P., O’Hara M., Modesto-Mata M., Martin-Francés L., Martinón-Torres M., Zhang I., Schepartz L.A., Bermúdez de Castro J.M., Guatelly-Steinberg D. 2019b First systematic assessment of dental growth and development in an archaic hominin (genus, Homo) from East Asia. Science Advances, vol. 5 (1). URL: https://doi:10.1126/sciadv.aau0930
145. Xu C., Zhang Y., Chen C., Fang D. 1984 Human occipital bone and mammalian fossils from Chaoxian, Anhui. Acta Anthropologica Sinica, vol. 3: 202–209.
146. Yuan S., Chen T., Gao S. 1986 The dating of southern Chinese Palaeolithic sites by uranium method. Acta Anthropologica Sinica, vol. 9: 179–190.
147. Zanolli C. 2013 Additional evidence for morpho-dimensional tooth crown variation in a new Indonesian H. erectus sample from the Sangiran Dome (Central Java). PLoS ONE. doi:10.1371/journal.pone.0067233
148. Zavala E.I., Jacobs Z., Vernot B., Shunkov M.V., Kozlikin M.B., Derevianko A.P., Essel E., de Fillipo C., Nagel S., Richter J., Romagné F., Schmidt A., Bo Li, O’Gorman K., Slon V., Kelso J., Pääbo S., Roberts R.G., Meyer M. 2021 Pleistocene sediment DNA reveals hominin and faunal turnovers at Denisova Cave. Nature, vol. 595: 399–403.
149. Zhang D., Xia H., Chen F., Li B., Slon V., Cheng T., Yang R., Jacobs Z., Dai Q., Massilani D., Shen X., Wang J., Feng X., Cao P., Yang M.A., Yao J., Yang J., Madsen D.B., Han Y., Ping W., Liu F., Perreault C., Chen X., Meyer M., Kelso J., Pääbo S., Fu Q. 2020 Denisovan DNA in Late Pleistocene sediments from Baishiya Karst Cave on the Tibetan Plateau. Science, vol. 370 (6516): 584–587.
150. Zhang X.L., Ha B.B., Wang S.J., Chen Z.J., Ge J.Y., Long H., He W., Da W., Nian X.M., Yi M.J., Zhou X.Y., Zhang P.Q., Jin Y.S., Bar-Yosef O., Olsen J.W., Gao X. 2018 The earliest human occupation of the high-altitude Tibetan Plateau 40 thousand to 30 thousand years ago. Science, vol. 362: 1049–1051.
151. Zwyns N. 2012 Laminar Technology and the Onset of the Upper Paleolithic in the Altai, Siberia (Studies in Human Evolution): Doctoral Thesis. Leiden: Univ. Press.
152. Zubova A.V., Chikisheva T.A., Shunkov M.V. 2017 The morphology of permanent molars from the Paleolithic layers of Denisova Cave. Archaeology, Ethnology and Anthropology of Eurasia, vol. 45 (1): 121–134. (In Russian and English).
Review
For citations:
Derevianko A.P. The Dispersal of Denisovans and Neanderthals in China. Archaeology, Ethnology & Anthropology of Eurasia. 2025;53(4):03-27. https://doi.org/10.17746/1563-0110.2025.53.4.003-027
Views:
30
JATS XML
Email this article 