First discovery of Panthera spelaea cranium from Salawusu, northern China

doi:10.19615/j.cnki.2096-9899.250704

Vertebrata Palasiatica ›› 2025, Vol. 63 ›› Issue (4): 323-334.DOI: 10.19615/j.cnki.2096-9899.250704

First discovery of Panthera spelaea cranium from Salawusu, northern China

JIANGZUO Qi-Gao¹^,^*(), LI Hong²^,^*, YAMAGUCHI Nobuyuki³, Joan MADURELL-MALAPEIRA⁴, ZHANG Jian-Sheng², MA Hui-Min², GUO Ding-Ge¹^,⁶, LI Shi-Jie¹^,⁶, FU Jiao¹^,⁶, ZHANG Xiao-Xiao⁵, LI Chun-Xiao¹^,⁶, XIE Kun⁷, TONG Hao-Wen¹, LIU Jin-Yi¹, WANG Shi-Qi¹, DENG Tao¹

1 Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences Beijing 10044, China
2 Inner Mongolia Museum Hohhot 750306, China
3 Wildlife Conservation Research Unit, Department of Biology, University of Oxford Abingdon OX13 5QL, UK
4 Earth Sciences Department, Paleo[Fab]Lab, Università di Firenze Firenze 50121, Italy
5 Tianjin Natural History Museum Tianjin 300201, China
6 University of Chinese Academy of Science Beijing 100049, China
7 State Key Laboratory of Loess Science, Institute of Earth Environment, Chinese Academy of Sciences Xi’an 710061, China

Received:2025-03-14 Online:2025-10-20 Published:2025-10-28
Contact: jiangzuo@ivpp.ac.cn
About author:First author contact:
^*These authors contribute equally to the paper.

中国北方萨拉乌苏动物群首次发现洞狮头骨

江左其杲¹^,^*(), 李虹²^,^*, 山口诚之³, 琼安·马杜雷利-马拉佩拉⁴, 张建升², 马慧敏², 郭丁歌¹^,⁶, 李世杰¹^,⁶, 付娇¹^,⁶, 张晓晓⁵, 李春晓¹^,⁶, 谢坤⁷, 同号文¹, 刘金毅¹, 王世骐¹, 邓涛¹

1 中国科学院古脊椎动物与古人类研究所北京 100044
2 内蒙古博物院呼和浩特 750306
3 英国牛津大学生物学系野生动物保护研究所阿尔宾 OX13 5QL
4 意大利佛罗伦萨大学地球科学系古生物实验室佛罗伦萨 50121
5 天津自然博物馆天津 300201
6 中国科学院大学北京 100049
7 中国科学院西安地球环境研究所黄土科学实验室西安 710061

基金资助:
国家重点研发计划(2023YFF0804501);国家自然科学基金重点项目和青年基金(42430207);国家自然科学基金重点项目和青年基金(42102001)

Abstract

Abstract:

Cave lion (Panthera spelaea) remains have been described in all the Holarctic domain, however, its records in China are very rare. Here we describe an almost complete cranium from Salawusu, Northern China. Even though the dentition is largely broken or missing, the cranium shows key characters with clear cave lion affinity, particularly from the basicranium region. These features include wide and short nasals, wide rostrum without post-canine constriction, enlarged ectotympanic, flatted entotympanic, separated stylomastoid foramen and hyoid fossa and medially located hypoglossal foramen. The cranium is larger than the known cranial size range of Beringia cave lion P. spelaea vereshchagini, suggesting that the geographical delimitation of cave lion subspecies may be more complex than previously understood. The coexistence of cave lion with Palaeoloxodon in Salawusu indicates a high plasticity in diet and environmental adaptation for this species.

Key words: eastern Asia, Pleistocene, cave lion, tiger, biogeography

摘要：

洞狮(Panthera spelaea)是中晚更新世冰期动物群的典型代表。该类群的化石在整个全北界均有记录，广布于欧洲，北亚，北美洲，但中国的发现却极为稀少。报道了发现于中国北方萨拉乌苏动物群的一具近乎完整的洞狮头骨化石。尽管其牙齿大部分已破损或缺失，但该头骨，尤其是在颅基区域，显示出了洞狮的关键特征。其中比较重要的有：宽阔而短的鼻骨、宽阔的吻部且无犬齿后的收缩、膨大的外鼓骨、扁平的内鼓骨、分离的茎乳孔与舌骨窝以及向内侧延伸扩展的舌下神经孔。该颅骨尺寸超出了已知北亚和北美洲的白令洞狮(P. spelaea vereshchagini)的变异范围。而形态测量学分析显示，该头骨完全落入欧洲洞狮的范畴，这表明洞狮亚种的地理区划可能较既往认知更为复杂。在萨拉乌苏动物群，洞狮与古菱齿象属(Palaeoloxodon)动物群共存，这一现象较为罕见(一般与猛犸象共存), 揭示了洞狮在食性选择和环境适应方面具有高度的可塑性。

关键词: 东亚, 更新世, 洞狮, 虎, 生物地理

CLC Number:

Q915.874

JIANGZUO Qi-Gao, LI Hong, YAMAGUCHI Nobuyuki, Joan MADURELL-MALAPEIRA, ZHANG Jian-Sheng, MA Hui-Min, GUO Ding-Ge, LI Shi-Jie, FU Jiao, ZHANG Xiao-Xiao, LI Chun-Xiao, XIE Kun, TONG Hao-Wen, LIU Jin-Yi, WANG Shi-Qi, DENG Tao. First discovery of Panthera spelaea cranium from Salawusu, northern China. Vertebrata Palasiatica, 2025, 63(4): 323-334.

江左其杲, 李虹, 山口诚之, 琼安·马杜雷利-马拉佩拉, 张建升, 马慧敏, 郭丁歌, 李世杰, 付娇, 张晓晓, 李春晓, 谢坤, 同号文, 刘金毅, 王世骐, 邓涛. 2025, 63(4): 323-334, 中国北方萨拉乌苏动物群首次发现洞狮头骨. 古脊椎动物学报.

Figures/Tables 5

Fig. 1 Cranial measurements of Panthera used in this study from Jiangzuo et al. (2022) Abbreviations: C. canine; L. length; L1. total length; L2. condylobasal length; L3. basilar length; LDP. diastema length between canine and P3; LT. toothrow length from canine to P4; W. width; W1. zygomatic width; W2. rostrum width across P4/M1; W3. rostrum width across canine; W4. cranial width at mastoid; W5. condyle width

Fig. 2 Cranium of Panthera spelaea from the Salawusu Fauna, IMM 035389 A. dorsal view; B. ventral view; C. lateral view; D. anterior view; E. posterior view

Fig. 3 Cranial comparison in lateral (A1-F1), dorsal (A2-F2) and ventral (A3-F3) views of Panthera from Salawusu, and selected fossil and modern Panthera A. Panthera spelaea, IMM 035389, Salawusu, China; B. P. spelaea, MB.MA.50948, (type of P. spelaea according to Kempe and Döppes (2009), Diedrich (2011)), Zoolithen Cave, Germany; C. P. spelaea, AMNH F:AM30460, Cleary Creek, Alaska, USA; D. P. atrox, LACMHC2900-7, La Brea, USA; E. P. leo AMNH M85149, modern; F. P. tigris AMNH M113743, modern. Not to scale

Fig. 4 Basicranial comparison of selected Panthera in ventral views Note on the separated hyoid fossa (H) and stylomastoid foramen (S), and medially extended hypoglossal foramen (Hy) in lion lineages A. P. spelaea Salawusu IMM 035389, B. P. spelaea Alaska AMNH F:AM30460, C. P. spelaea Zoolithen Cave MB.MA.50948; D. P. atrox La Brea LACMHC2900-7; E. P. leo AMNH M85149, modern; F. P. tigris AMNH M113743, modern

Fig. 5 PCA plot of three cranial measurements (L2, W2 and W3) of cave lion and related taxa The data sources see materials and methods

References 37

[1]	Barnett R, Shapiro B, Barnes I et al., 2009. Phylogeography of lions (Panthera leo ssp.) reveals three distinct taxa and a late Pleistocene reduction in genetic diversity. Mol Ecol, 18: 1668-1677 DOI PMID
[2]	Barnett R, Mendoza M L Z, Soares A E R et al., 2016. Mitogenomics of the extinct cave lion, Panthera spelaea (Goldfuss, 1810), resolve its position within the Panthera cats. Open Quat, 2: 1-11
[3]	Baryshnikov G, Boeskorov G, 2001. The Pleistocene cave lion, Panthera spelaea (Carnivora, Felidae) from Yakutia, Russia. Cranium, 18: 7-23
[4]	Boeskorov G, Begolyubskyi I N, Valerievich P et al., 2012. New finds of fossil cave lion in territory of Yakutia. Nauk Obraz, 2(66): 45-51
[5]	Boule M, 1928. Le paléolithique de la Chine. Paris: Archives de l'Institut de Paleontologie Humanie. 1-136
[6]	Christiansen P, 2008. Distinguishing skulls of lions (Panthera leo) and tigers (Panthera tigris). Mamm Biol, 73: 451-456
[7]	de Manuel M, Barnett R, Sandoval-Velasco M et al., 2020. The evolutionary history of extinct and living lions. Proc Natl Acad Sci USA, 117: 10927-10934 DOI PMID
[8]	Diedrich C G, 2011. The largest European lion Panthera leo spelaea (Goldfuss 1810) population from the Zoolithen Cave, Germany: specialised cave bear predators of Europe. Hist Biol, 23: 271-311
[9]	Diedrich C G, 2014. Late Pleistocene Eemian hyena and steppe lion feeding strategies on their largest prey—Palaeoloxodon antiquus Falconer and Cautley 1845 at the straight-tusked elephant graveyard and Neanderthal site Neumark-Nord Lake 1, Central Germany. Archaeol Anthropol Sci, 6: 271-291
[10]	Guthrie R D, 1968. Paleoecology of the large-mammal community in interior Alaska during the late Pleistocene. Am Mid Nat, 79: 346-363
[11]	Haltenorth T, 1936. Die verwandtschaftliche Stellung der Großkatzen zueinander. I. Beschreibung der Schädelknochen. Z Säugetierkd, 11: 32-105
[12]	Hammer Ø, Harper D, Ryan P, 2001. PAST: paleontological statistics software package for education and data analysis. Palaeontol Electron, 4: 1-9
[13]	Harington C R, 1969. Pleistocene remains of the lion-like cat (Panthera atrox) from the Yukon Territory and northern Alaska. Can J Earth Sci, 6: 1277-1288
[14]	Harington C R, 2011. Pleistocene vertebrates of the Yukon Territory. Quat Sci Rev, 30: 2341-2354
[15]	Hemmer H, 2011. The story of the cave lion-Panthera leo spelaea (Goldfuss, 1810) - a review. Quaternaire, suppl: 201-208
[16]	Huang G, Rosowski J, Ravicz M et al., 2002. Mammalian ear specializations in arid habitats: structural and functional evidence from sand cat (Felis margarita). J Comp Physiol, A188: 663-681
[17]	Jiangzuo Q G, Liu J Y, 2020. First record of the Eurasian jaguar in southern Asia and a review of dental differences between pantherine cats. J Quat Sci, 35: 817-830
[18]	Jiangzuo Q G, Li L, Madurell-Malapeira J et al., 2022. The diversification of the lynx lineage during the Plio-Pleistocene—evidence from a new small Lynx from Longdan, Gansu Province, China. Biol J Linn Soc, 136: 536-551
[19]	Jiangzuo Q G, Madurell-Malapeira J, Li X et al., 2025. Insights on the evolution and adaptation towards high altitude and cold environments in the snow leopard lineage. Sci Adv, 11: eadp5243
[20]	Kahlke R D, 2014. The origin of Eurasian mammoth faunas (Mammuthus-Coelodonta faunal complex). Quat Sci Rev, 96: 32-49
[21]	Kempe S, Döppes D, 2009. Cave bear, cave lion and cave hyena skulls from the public collection at the Humboldt Museum in Berlin. Acta Carsol, 38: 2-3
[22]	Marciszak A, Gornig W, 2024. From giant to dwarf: a trend of decreasing size in Panthera spelaea (Goldfuss, 1810) and its likely implications. Earth Hist Biodivers, 1: 100007
[23]	Marciszak A, Schouwenburg C, Darga R, 2014. Decreasing size process in the cave (Pleistocene) lion Panthera spelaea (Goldfuss, 1810) evolution-a review. Quat Int, 339: 245-257
[24]	Marciszak A, Lipecki G, Pawłowska K et al., 2021. The Pleistocene lion Panthera spelaea (Goldfuss, 1810) from Poland-a review. Quat Int, 605: 213-240
[25]	Marciszak A, Ivanoff D V, Semenov Y A et al., 2023. The Quaternary lions of Ukraine and a trend of decreasing size in Panthera spelaea. J Mamm Evol, 30: 109-135
[26]	Nedoluzhko A, Boeskorov G, Tikhonov A et al., 2024. 20th anniversary of the history of genetic research on cave lions: a short review. Earth Hist Biodivers, 2: 100013
[27]	Persico D, 2021. First fossil record of cave lion (Panthera (Leo) spelaea intermedia) from alluvial deposits of the Po River in northern Italy. Quat Int, 586: 14-23
[28]	Puzachenko A Y, Argant A, Baryshnikov G F et al., 2024. Distribution history of the cave lion (Panthera spelaea (Goldfuss, 1810)). Earth Hist Biodivers, 1: 100006
[29]	Qi G Q, 1975. Quaternary mammalian fossils from Salawusu River district, Nei Mongol. Vert PalAsiat, 13: 239-249
[30]	Sabol M, 2011a. Masters of the lost world: a hypothetical look at the temporal and spatial distribution of lion-like felids. Quaternaire, 4: 229-236
[31]	Sabol M, 2011b. A record of Pleistocene lion-like felids in the territory of Slovakia. Quaternaire, 4: 215-228
[32]	Sabol M, Tomašových A, Gullár J, 2022. Geographic and temporal variability in Pleistocene lion-like felids: implications for their evolution and taxonomy. Palaeontol Electron: 1-27
[33]	Sherani S, Perng L, Sherani M, 2022. Evidence of cave lion (Panthera spelaea) from Pleistocene Northeast China. Hist Biol, 35: 1-9
[34]	Sotnikova M, Nikolskiy P, 2006. Systematic position of the cave lion Panthera spelaea (Goldfuss) based on cranial and dental characters. Quat Int, 142: 218-228
[35]	Stanton D W, Alberti F, Plotnikov V et al., 2020. Early Pleistocene origin and extensive intra-species diversity of the extinct cave lion. Sci Rep, 10: 12621 DOI PMID
[36]	Tong H W, Li H, Xie J Y, 2008. Revisions of some taxa of the Salawusu fauna from Sjara-Osso-Gol area, Nei Mongol, China. Quat Sci, 28: 1106-1113
[37]	Tong H W, Li H, Xie J Y, 2017. Fossil Vertebrates. In: Dong G R, Li B S, Cheng Y Z eds. Comprehensive Study of the Late Quaternary Geology and Paleoanthropology of the Xarusgol River Region. Beijing: Science Press. 157-204

First discovery of Panthera spelaea cranium from Salawusu, northern China

中国北方萨拉乌苏动物群首次发现洞狮头骨

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