Morphological diversities and evolutionary implications of the late Middle Pleistocene hominins in China

doi:10.16359/j.1000-3193/AAS.2022.0024

Abstract

Abstract:

The hominin fossils have been found in more than 20 late Middle Pleistocene sites in present China. For many years, these fossils have been classified as archaic Homo sapiens, intermediate between Homo erectus and early modern humans, and the ancestors of modern humans in East Asia. However, such an opinion has never been widely accepted in paleoanthropological community. There have been debates on the evolution and taxonomy of late Middle Pleistocene hominins in China and around the world. Since 21^st century, some noticeable progresses on the late Middle Pleistocene hominin evolution in China have been achieved. The discoveries and studies on the hominin fossils of Penghu, Xuchang, Hualongdong, Xiahe and Harbin greatly enriched the hominin fossil record in China. Studies on the late Middle Pleistocene hominin fossils in China revealed complicated morphological diversities indicating simply classifying all the hominins of this time period into archaic Homo sapiens cannot accurately reflect the evolutionary patterns of Middle Pleistocene hominins in China. According to the studies of the hominin fossils from Xujiayao, Xuchang, Hualongdong, Xiahe and Harbin, some new opinions on the evolutionary pattern and taxonomy of the late Middle Pleistocene hominins in China have been proposed and triggered different understandings.

In this study, with analyzing the morphological diversities of late Middle Pleistocene hominin fossils in China, four types of the diversities were identified. Xuchang and Xujiyao are characterized with huge-sized crania and cranial capacity (1800 mL and 1700 mL respectively) with some Neanderthal features. These features occurred together in Xuchang and Xujiayao crania constitute unique morphological combinations which have not been found in all other contemporaneous hominin fossils. The other types of morphological diversities are characterized with 1) dominant common sharing features of late Middle Pleistocene hominins; 2) mainly primitive features; and 3) mainly derived or modern features. These morphological diversities suggest that not all the late Middle Pleistocene hominins in China contributed the formation of modern humans equally. Regarding the opinions that put some of the hominin fossils into different Homo members or taxa, the authors believe that at the present with no fully understanding these morphological diversities, treating the related late Middle Pleistocene hominin fossils in China with combined or mosaic morphological features as populations of unclear taxonomic status is a proper way.

Key words: Late Middle Pleistocene, archaic Homo sapiens, morphological diversity, taxonomy, evolution

CLC Number:

Q981

LIU Wu, WU Xiujie. Morphological diversities and evolutionary implications of the late Middle Pleistocene hominins in China[J]. Acta Anthropologica Sinica, 2022, 41(04): 563-575.

Figures/Tables 5

References 52

[1]	刘武, 吴秀杰, 邢松, 等. 中国古人类化石[M]. 北京: 科学出版社, 2014, 1-378
[2]	吴新智, 张银运. 中国古人类综合研究[A]. 见:中国科学院古脊椎动物与古人类研究所(编).古人类论文集[C]. 北京: 科学出版社, 1978, 28-42
[3]	吴新智. 大荔颅骨在人类进化中的位置[J]. 人类学学报, 2014, 33: 405-426
[4]	Rightmire GP. Human evolution in the Middle Pleistocene: the role of Homo heidelbergensis[J]. Evolutionary Anthropology, 1998, 6: 218-227 doi: 10.1002/(SICI)1520-6505(1998)6:6<218::AID-EVAN4>3.0.CO;2-6 URL
[5]	Rightmire GP. Homo in the Middle Pleistocene: hypodigms, variation and species recognition[J]. Evolutionary Anthropology, 2008, 17: 8-21 doi: 10.1002/evan.20160 URL
[6]	Hublin JJ, Ben-Ncer A, Bailey S, et al. New fossils from Jebel Irhoud, Morocco and the pan-African origin of Homo sapiens[J]. Nature, 2017, 546: 289-292 doi: 10.1038/nature22336 URL
[7]	Reich D, Green RE, Kircher M, et al. Genetic history of an archaic hominin group from Denisova Cave in Siberia[J]. Nature, 2010, 468: 1053-1060 doi: 10.1038/nature09710 URL
[8]	Roksandic M, Radovic P, Wu XJ, et al. Resolving the “muddle in the middle”: The case for Homo bodoensis sp. nov[J]. Evolutionary Anthropology, 2022, 31: 20-29 doi: 10.1002/evan.21929 URL
[9]	Arsuaga JL, Martínez I, Arnold LJ, et al. Neandertal roots: cranial and chronological evidence from Sima de los Huesos[J]. Science, 2014, 344: 1358-1363 doi: 10.1126/science.1253958 pmid: 24948730
[10]	Stringer C. The status of Homo heidelbergensis (Schoetensack 1908)[J]. Evolutionary Anthropology, 2012, 21: 101-107 doi: 10.1002/evan.21311 pmid: 22718477
[11]	Chang CH, Kaifu Y, Takai M, et al. The first archaic Homo from Taiwan[J]. Nature Communication, 2015, 6: 6037 doi: 10.1038/ncomms7037 URL
[12]	吴新智, 同号文. 关于澎湖人类下颌骨化石意义和年代的讨论[J]. 人类学学报, 2015, 34: 281-287
[13]	Li ZY, Wu XJ, Zhou LP, et al. Late Pleistocene archaic human crania from Xuchang, China[J]. Science, 2017, 355: 969-972 doi: 10.1126/science.aal2482 URL
[14]	Wu XJ, Pei SW, Cai YJ, et al. Archaic human remains from Hualongdong, China, and Middle Pleistocene human continuity and variation[J]. Proceedings of the National Academy of Sciences, 2019, 116: 9820-9824 doi: 10.1073/pnas.1902396116 URL
[15]	Chen FH, Frido W, Shen CC, et al. A late Middle Pleistocene Denisovan mandible from the Tibetan plateau[J]. Nature, 2019, 569: 409-412 doi: 10.1038/s41586-019-1139-x URL
[16]	Ni XJ, Ji Q, Wu WS, et al. Massive cranium from Harbin in northeastern China establishs a new Middle Pleistocene human lineage[J]. The Innovation, 2021, 2: 100130 doi: 10.1016/j.xinn.2021.100130 URL
[17]	Ji Q, Wu WS, Ji YN, et al. Late Middle Pleistocene Harbin cranium represents a new Homo species[J]. The Innovation, 2021, 2: 100132 doi: 10.1016/j.xinn.2021.100132 URL
[18]	刘武, 吴秀杰, 邢松. 更新世中期中国古人类演化区域连续性与多样性的化石证据[J]. 人类学学报, 2019, 38: 473-490
[19]	Wu XJ, Pei SW, Cai YJ, et al. Morphological description and evolutionary significance of 300 ka hominin facial bones from Hualongdong, China[J]. Journal of Human Evolution, 2021, 161: 103052 doi: 10.1016/j.jhevol.2021.103052 URL
[20]	吴新智. 大荔中更新世人类颅骨[M]. 北京: 科学出版社, 2020, 1-205
[21]	Wu XZ, Athreya S. A description of the geological context, discrete traits, and linear morphometrics of the Middle Pleistocene hominin from Dali, Shaanxi Province, China[J]. American Journal of Physical Anthropology, 2013, 150: 141-157 doi: 10.1002/ajpa.22188 URL
[22]	刘武, 惠家明, 何嘉宁, 等. 门齿孔位置在中国古人类化石与现代人群的表现及其演化意义[J]. 人类学学报, 2021, 40: 739-750
[23]	吴汝康. 辽宁营口金牛山人化石头骨的复原及其主要性状[J]. 人类学学报, 1988, 7: 97-101
[24]	何嘉宁. 金牛山人化石牙齿初步研究[J]. 人类学学报, 2000, 19: 216-225
[25]	Xing S, Wu XJ, Liu W, et al. Middle Pleistocene human femoral diaphyses from Hualongdong, Anhui Province, China[J]. American Journal of Physical Anthropology, 2021, 174: 285-298 doi: 10.1002/ajpa.24121 URL
[26]	Shao QF, Ge JY, Ji Q, et al. Geochemical provenancing and direct dating of the Harbin archaic human cranium[J]. The Innovation, 2021, 2: 100131 doi: 10.1016/j.xinn.2021.100131 URL
[27]	吴汝康, 彭如策. 广东韶关马坝发现的早期古人类型人类化石[J]. 古脊椎动物与古人类, 1959, 1: 159-163
[28]	Wu XJ, Bruner M. The endocranial anatomy of Maba 1[J]. American Journal of Physical Anthropology, 2016, 160: 633-643 doi: 10.1002/ajpa.22974 URL
[29]	Wu XJ, Crevecoeur I, Liu W, et al. Temporal labyrinths of eastern Eurasian Pleistocene humans[J]. Proceedings of the National Academy of Sciences, 2014, 111: 10509-10513 doi: 10.1073/pnas.1410735111 URL
[30]	Wu XJ, Trinkaus E. The Xujiayao 14 mandibular ramus and Pleistocene Homo mandibular variation. Comptes Rendus Palevol[J]. 2014, 13: 333-341
[31]	Xing S, Martinón-Torres M, Bermúdez de Castro JM, et al. Hominin teeth from the early Late Pleistocene site of Xujiayao, Northern China[J]. American Journal of Physical Anthropology, 2015, 156: 224-240 doi: 10.1002/ajpa.22641 URL
[32]	Wu XJ, Maddx S, Pan L, et al. Nasal floor variation among eastern Eurasian Pleistocene Homo[J]. Anthropological Science, 2012, 120: 217-226 doi: 10.1537/ase.120709 URL
[33]	Wu XJ, Bae CJ, Friess M, et al. Evolution of cranial capacity revisited: A view from the late Middle Pleistocene cranium from Xujiayao, China[J]. Journal of Human Evolution, 2022, 163: 103119 doi: 10.1016/j.jhevol.2021.103119 URL
[34]	贾兰坡. 长阳人化石及共生的哺乳动物群[J]. 古脊椎动物与古人类, 1957, 1: 247-257
[35]	许春华, 张银运, 陈才弟, 等. 安徽巢县发现的人类枕骨化石和哺乳动物化石[J]. 人类学学报, 1984, 3: 202-209
[36]	许春华, 张银运, 方笃生. 安徽巢县人类化石地点的新材料[J]. 人类学学报, 1986, 5: 305-310
[37]	Bailey S, Liu W. A comparative dental metrical and morphological analysis of a Middle Pleistocene hominin maxilla from Chaoxian (Chaohu), China[J]. Quaternary International, 2010, 211: 14-23 doi: 10.1016/j.quaint.2009.01.008 URL
[38]	Xing S, Martinón-Torres M, Bermúdez de Castro JM. Late Middle Pleistocene hominin teeth from Tongzi, southern China[J]. Journal of Human Evolution, 2019, 130: 96-108 doi: 10.1016/j.jhevol.2019.03.001 URL
[39]	Liu W, Schepartz L, Xing S, et al. Late Middle Pleistocene hominin teeth from Panxian Dadong, South China[J]. Journal of Human Evolution, 2013, 64: 337-355 doi: 10.1016/j.jhevol.2012.10.012 pmid: 23465337
[40]	Weidenreich F. The Skull of Sinanthropus pekinensis: A Comparative Study of A Primitive Hominid Skull. Palaeontologica Sinica N.S. D10, The Geological Survey of China[M]. 1943, 1-484
[41]	Wood BA. The origin of Homo erectus[J]. Cour Forsch Inst Senkenberg, 1984, 69: 99-111
[42]	Antón SC. Evolutionary significance of cranial variation in Asian Homo erectus[J]. American Journal of Physical Anthropology, 2002, 118:301-323 doi: 10.1002/ajpa.10091 URL
[43]	Xing S, Tafforeau P, O’Hara M, et al. First Systematic Assessment of Dental Growth and Development in an Archaic Hominin (genus, Homo) from East Asia[J]. Science Advance, 2019, 5: eaau0930
[44]	Hublin JJ, Ben-Ncer A, Bailey S, et al. New fossils from Jebel Irhoud, Morocco and the pan-African origin of Homo sapiens[J]. Nature, 2017, 546: 289-292 doi: 10.1038/nature22336 URL
[45]	Reich D, Green R, Kircher M, et al. Genetic history of an archaic hominin group from Denisova Cave in Siberia[J]. Nature, 2010, 468: 1053-1060 doi: 10.1038/nature09710 URL
[46]	Bergström A, Stringer C, Hajdinjak M, et al. Origins of modern human ancestry[J]. Nature, 2021, 590: 229-237 doi: 10.1038/s41586-021-03244-5 URL
[47]	Brown P, Sutikna T, Morwood M, et al. A new small-bodied hominin from the Late Pleistocene of Flores, Indonesia[J]. Nature, 2004, 431: 1055-1061 doi: 10.1038/nature02999 URL
[48]	Berger L, Hawks J, Ruiter D, et al. Homo naledi, a new species of the genus Homo from the Dinaledi Chamber, South Africa[J]. eLife, 2015, 4: e09560
[49]	Détroit F, Mijares A, Corny J, et al. A new species of Homo from the Late Pleistocene of the Philippines[J]. Nature, 2019, 568: 181-186 doi: 10.1038/s41586-019-1067-9 URL
[50]	Arsuaga JL, Martínez I, Arnold LJ, et al. Neandertal roots: cranial and chronological evidence from Sima de los Huesos[J]. Science, 2014, 344: 1358-1363 doi: 10.1126/science.1253958 pmid: 24948730
[51]	Hershkovitz I, May H, Sarig R, et al. A Middle Pleistocene Homo from Nesher Ramla, Israel[J]. Science, 2021, 372: 1424-1428
[52]	Derevianko AP. Upper Paleolithic in Africa and Eurasia and the Formation of a Human of the Modern Anatomical Type[M]. Novosibirsk:Instiute of Archaeology and Ethnography,Siberian Branch of the Russian Academy of Sciences, 2011, 1-560

地点Site	人类化石Human fossils	年代Age	来源Source
大荔Dali	头骨（模型）	30-26万年前[19]	[3, 20, 21, 22]
金牛山Jinniushan	头骨、牙齿（模型）	26万年前[1]	[22, 23, 24]
许昌Xuchang	头骨	12.5-10.5万年前[13]	[13]
华龙洞Hualongdong	头骨、下颌骨、牙齿	30-27万年前[14]	[14, 19, 22, 25]
哈尔滨Harbin	头骨、牙齿	30.9-13.8万年前[26]	[16, 17]
马坝Maba	头骨	30-13万年前[1]	[27, 28]
许家窑Xujiayao	头骨、下颌骨、牙齿	20-16万年前[1]	[29, 30, 31, 32, 33]
澎湖Penghu	下颌骨、牙齿（模型）	19万年前[11]	[11, 12]
夏河Xihe	下颌骨、牙齿	17万年前[15]	[15]
长阳Changyang	上颌骨、牙齿	19.5万年前[1]	[22, 32, 34]
巢县Chaoxian	头骨、上颌骨、牙齿	30-20万年前[1]	[22, 35, 36, 37]
丁村Dingcun	牙齿	20-16万年前[1]	本文
桐梓Tongzi	牙齿	24-17.2万年前[1, 38]	[38]
盘县大洞Panxian Dadong	牙齿	28 -13万年前[1]	[39]

地点Site	人类化石Human fossils	年代Age	来源Source
大荔Dali	头骨（模型）	30-26万年前[19]	[3, 20, 21, 22]
金牛山Jinniushan	头骨、牙齿（模型）	26万年前[1]	[22, 23, 24]
许昌Xuchang	头骨	12.5-10.5万年前[13]	[13]
华龙洞Hualongdong	头骨、下颌骨、牙齿	30-27万年前[14]	[14, 19, 22, 25]
哈尔滨Harbin	头骨、牙齿	30.9-13.8万年前[26]	[16, 17]
马坝Maba	头骨	30-13万年前[1]	[27, 28]
许家窑Xujiayao	头骨、下颌骨、牙齿	20-16万年前[1]	[29, 30, 31, 32, 33]
澎湖Penghu	下颌骨、牙齿（模型）	19万年前[11]	[11, 12]
夏河Xihe	下颌骨、牙齿	17万年前[15]	[15]
长阳Changyang	上颌骨、牙齿	19.5万年前[1]	[22, 32, 34]
巢县Chaoxian	头骨、上颌骨、牙齿	30-20万年前[1]	[22, 35, 36, 37]
丁村Dingcun	牙齿	20-16万年前[1]	本文
桐梓Tongzi	牙齿	24-17.2万年前[1, 38]	[38]
盘县大洞Panxian Dadong	牙齿	28 -13万年前[1]	[39]