Tuesday, August 24, 2021

[PaleoMammalogy • 2021] First Report of Leopard Panthera pardus Fossils from A Limestone Cave in Kenting Area, southern Taiwan


Panthera pardus  
fossil leopard teeth uncovered from Longshia-dong Cave

in Chi​, Gan​, Yang et Chang,. 2021. 
Illustrated by Chien-Hsing Lee

Abstract
Longshia-dong Cave, a limestone cave located in the Kenting area within the Kenting National Park of southern Taiwan, yields numerous terrestrial mammalian fossils. Many of them were not reported in historical literature and are neither present in Taiwan. For instance, no historical literature mentioned leopards inhabited in Taiwan, and thus their existence remained unknown. This study describes three fossil leopard (Panthera pardus) teeth uncovered from Longshia-dong Cave. Two isolated lower premolars and one lower molar, respectively p3, p4 and m1, were discovered in a very small area (11 × 6 cm) and show a series of progressive increase in size. Thus, the three teeth should have been belonging to the same individual from the subfamily of Pantherinae. Traditional linear measurements and two-dimensional geometric morphometric analysis for the occlusal surface outlines were conducted on the fossil teeth and extant pantherines inhabited in Asia such as clouded leopards (Neofelis nebulosa), leopards (Panthera pardus), and tigers (Panthera tigiris). Results show that the fossil teeth are similar both in size and morphology to the teeth of extant leopards, suggesting the assignment of the fossil teeth to leopards. This study, for the first time, reported the presence of leopards in the Late Pleistocene of Taiwan. In addition, the smaller size of the fossil teeth in comparison with Chinese fossil leopards is putatively attributed to insular dwarfism or individual size variability, yet more studies are required.

The three felid lower cheek teeth from Longshia-dong Cave,
including p3 (a, F056584), p4 (b, F056585), and m1 (c, F056585),
and their buccal views (1), lingual views (2), and occlusal views (3).

Family Felidae Fischer von Waldheim (1817)
Subfamily Pantherinae Pocock, 1917

Genus Panthera Oken, 1816

Panthera pardus (Linnaeus, 1758)  


Illustrated by Chien-Hsing Lee
李建興繪製

Figure 10: Discovery sites of Pleistocene leopard fossils in East Asia.
Early Pleistocene 1, Zhen’an, Shaanxi Province, China (Li & Deng, 2003); 2, Bailong Cave, Yunxi, Hubei Province, China (Wu et al., 2009); 3, Longgupou, Wushan County, Chongqing, China (Jin et al., 2008); 4, Cili County, Hunan Province, China (Wang et al., 1982); 5, Hezhang County, Guizhou Province, China (Zhao et al., 2013); 6, Yuanmou, Yunnan Province, China (You & Qi, 1973);
Middle Pleistocene 7, Chaoxian, Anhui Province, China (Xu et al., 1984); 8, Yanjinggou (formerly Yenchingkou), Chongqing, China (Young, 1935); 9, Panlong Cave, Yunfu, Guangdong Province, China (Wang et al., 1990);
Late Pleistocene 10, Niuyan Cave ( = Bull Eye Cave), Mentougou, Beijing, China (Deng, Huang & Wang, 1999); 11, Locality 1, Zhoukoudien (formerly Choukoutien), Beijing, China (Pei, 1934); 12, Locality 13, Zhoukoudien, Beijing, China (formerly Choukoutien) (Teilhard de Chardin & Pei, 1941); 13, Anyang, Henan Province, China (Teilhard de Chardin & Young, 1936); 14, Gongwangling, Lantian County, Shaanxi Province, China (Hu & Qi, 1978); 15, Yang’er Cave, Huatan County, Hunan Province, China (Wu, Deng & Zheng, 2008); 16, Fuyan Cave, Daoxian, Hunan Province, China (Li et al., 2013); 17, Mawoukou Cave, Bijie, Guizhou Province, China (Zhao et al., 2016); 18, Heshang Cave, Fumin County, Yunnan Province, China (but possibly Middle Pleistocene; (Zhang et al., 1989)); 19, Liucheng Gigantopithecus Cave, Guangxi Province, China (Pei, 1987); 20, this study. 21, Lower Hamakita Formation, Honshu Island, Japan (Suzuki, 1966) 22, Chongphadae Cave, Hwangju County, Democratic People’s Republic of Korea (Choe et al., 2020).

Conclusions:
 Leopards are a group of carnivores widely distributed throughout Asian and African continents, yet they are no longer found in eastern Asia due to civilization. Fossils thus represent the only clue indicating prehistoric leopards. While their teeth are significantly smaller than the other leopard teeth compared and thus are easily identified, poorly preserved fossils hinder further investigations. This study, based on an integration of morphological and geometric morphometric analyses, reveals the assignment of the fossil teeth excavated from Longshia-dong Cave to Panthera pardus, which is currently absent on Taiwan Island and suggests the presence of leopards in the Late Pleistocene of Taiwan. Such a record of prehistoric leopards in Taiwan thus adds up to the carnivore biodiversity of Taiwan. However, whether the Panthera individual is aboriginal or migrated is still uncertain. Our study also found that the teeth from Longshia-dong Cave belonged to a smaller individual in comparison with the fossil record from China indicates the smaller size of Taiwanese leopard fossil teeth than Chinese ones. Such a smaller dental size was possibly a result of individual size variability or insular dwarfism. To conclude, the discovery of the leopard fossil in Taiwan opens shed some light on the origin of the Kenting Fauna. Nevertheless, the insular dwarfism of, and the migration history of, the prehistoric leopards in Taiwan are still speculations and thus require more specimens and studies.


Tzu-Chin Chi​, Yi Gan​, Tzu-Ruei Yang and Chun-Hsiang Chang​. 2021. First Report of Leopard Fossils from A Limestone Cave in Kenting Area, southern Taiwan. PeerJ. 9:e12020. DOI: 10.7717/peerj.12020