Geographical and chronological distribution of Chinese Pleistocene large canids: Current status and prospects

Abstract

Large canids (here referrering to Canis, Cuon, Lycaon, and Lupulella and their fossil relatives) are some of the most iconic elements in Pleistocene fauna. They play an important role in biostratigraphy inference and paleoecology reconstruction. Large canids are also a very important element in Chinese Pleistocene faunas. Here we review the Pleistocene large canid fossil record in China and summarize the geographic and chronological distribution of this group. Our review suggests that there are more than ten large canid species known for the entire Pleistocene. The early Early Pleistocene of northern China is represented by Canis antonii from Yangshaocun, Mianchi, Anhui Province and Fancun, Taigu Shanxi Province; Canis teilhardi (and possibly Canis longdanensis, Canis brevicephalus) from Longdan, Dongxiang, Gansu Province; Canis aff. mosbachensis from Ningyang, Shandong Province; Sinicuon dubius from Huiyu, Beijing and Bajiazui, Qingyang, Gansu Province. The early Early Pleistocene of southern China is represented by Sinicuon cf. dubius from Longgupu, Wushan Chongqing and Baeryan, Bijie, Guizhou Province and Gigantopithecus Cave, Liucheng, and Baikong Cave, Chongzuo, Guangxi. The middle Early Pleistocene of northern China is represented by Canis chihliensis from Huailai and Nihewan Basin, Hebei Province; the middle Early Pleistocene of southern China is represented by Sinicuon yuanmouensis from Madahai, Yuanmou, Yunnan Province, Sinicuon sp. from Longgudong Cave, Jianshi, Hubei Province. The late Early Pleistocene of northern China is represented by Canismosbachensis from Gongwangling, Lantian, Shaanxi Province; the late Early Pleistocene of southern China is represented by Sinicuon sp. from Xiyuanhekou, Yunxian, Hubei Province. The Middle Pleistocene is represented by Canis mosbachensis variabilis, Xenocyon lycaonoides, and Cuon alpinus from several localities of Zhoukoudian, Beijing, Canis lupus from Loc. 3 of Zhoukoudian, Beijing. Cuon alpinus is also recorded from various localities in southern China, including Yanjiugou. The Late Pleistocene is represented only by living species Canis lupus and Cuon alpinus. Notably, Canis lupus appeared late in southern China, and was restricted to high altitude area during the Pleistocene. Northern China shows a greater diversity of large canids than that of southern China through the entire Pleistocene, suggesting the large canids are more adaptive to open environments. Similar to the situation in Europe, the early Pleistocene has the highest diversity for large canids, especially in northern China. The Middle and Late Pleistocene large canid guild shows great similarity to that of Europe, with almost all species shared by both regions. While the Middle and Late Pleistocene large canids are mostly well-defined and their taxonomy is clear, there is still much uncertainty about the taxonomy and evolution of Early Pleistocene large canids. This is due to the more abundant material from Middle and Late Pleistocene, and more specialization of the Middle and Late Pleistocene large canids. In contrast, the Early Pleistocene large canids are more diverse, less specialized and many species has very few specimens. The Early Pleistocene large canids in China need further work, especially in three aspects: (1) The possibility of coexistence of multiple similar large canid in the single locality, e.g., Longdan; (2) the possibility of presence of multiple large canid lineages (Lycaon lineage, Cuon lineage, grey wolf lineage and extinct lineages) in China; (3) the possibility of presence of ancestor the stock of African hunting dog lineage (Xenocyon) and how can it be distinguished from the species of the early dhole lineage.