Volume 30 Issue 9   20 September 2022
China is one of the countries harboring the highest Felidae diversity throughout the world, with 12 wild cat species occurred in the country. Front, from left to right: marbled cat (Pardofelis marmorata), wild cat (Felis silvestris), Chinese mountain cat (F. bieti), Pallas’s cat (Otocolobus manul), Asiatic golden cat (Catopuma temminckii). Back, from left to right: clouded leopard (Neofelis nebulosa), Eurasian lynx (Lynx lynx), snow leopard (Panthera uncia), tiger (P. tigris), jungle cat (Felis chaus), leopard (Panthera pardus), leopard cat (Prionailurus bengalensis). (Illustrated by Xiaozhao Mao)
    • Progress in the research and conservation of China’s Felidae species
      Sheng Li Tianming Wang
      Biodiv Sci. 2022, 30 (9):  22560.  doi:10.17520/biods.2022560
      Abstract ( 163 )   PDF (760KB) ( 188 )   Save
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      Restoring tiger population in Asia: Challenges, opportunities and future prospects
      Yixiao Zhu Dawei Wang Zhilin Li Jiawei Feng Tianming Wang
      Biodiv Sci. 2022, 30 (9):  22421.  doi:10.17520/biods.2022421
      Abstract ( 104 )   PDF (6572KB) ( 85 )   Save
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      Background & Aim: The tiger (Panthera tigris) may be the most charismatic and well-recognized flagship species in the world. As an ecological umbrella species and apex predator, the species symbolizes the well-being of the forest ecosystem. Tigers have lost 93% of their historical range and are experiencing rapid population declines. To enact effective conservation, it is important to understand the ecology and natural history of this globally endangered species. In this paper, we review previous articles related to tiger ecology and conservation research, summarizing the population dynamics and major conservation challenges in Asia in order to outline the actions required to conserve tigers and their ecosystems. Review Results: We found that, while overall research about tigers was increasing, efforts focused primarily on the subspecies with the most remaining range (e.g., P. t. tigris and P. t. altaica) and neglected subspecies requiring urgent attention. Tiger population has increased over the past decade, with estimates increasing from 3,200 to 4,500 during 2010–2021. However, stressors such as habitat fragmentation and loss, hunting of tigers and their prey, illegal trade, and human-tiger conflicts have isolated wild tigers in small populations across their ancestral range. Isolation, small territories, and disease further threaten the extant subspecies. Perspectives: We suggest that establishing a long-term monitoring network is critical for the conservation of tigers. To achieve the goal of a large tiger meta-population across Asia we recommend extensive land use planning, restoring native ungulates, reducing anthropogenic disturbances, improving connectivity of tiger habitats, controlling disease, and extensive cooperation across territories. In landscapes lacking breeding females, we also recommend reintroduction of tigers as a means of increasing recovery speeds.
      Special Feature: Research and Conservation of China's Felidae Species
      Competition and coexistence between tigers and leopards in Asia
      Zhilin Li Tianming Wang
      Biodiv Sci. 2022, 30 (9):  22271.  doi:10.17520/biods.2022271
      Abstract ( 116 )   PDF (1192KB) ( 156 )   Save
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      Background & Aim: As apex carnivores, tigers (Panthera tigris) and leopards (P. pardus) play critical roles in maintaining the structural and functional stability of Asian forest ecosystems. The populations and ranges of these two felids have declined due to human disturbance. To mitigate anthropogenic threats to endangered species in these ecosystems, it is essential that we understand how tigers and leopards interact. Although competition between tigers and leopards has been studied for several decades, most studies have taken place in the famous reserves like Nagarahole Natural Park and Chitwan National Park in South Asia, so it is unknown how their interactions differ elsewhere in their overlapping range. Here, we review 36 previous papers on competitive interactions between the two big cats from 1976 to 2021. We summarize the nature of both interference and exploitation competition between tigers and leopards in their current overlapping range, as well as how prey and human disturbance shapes this competition.

      Review Results: We found that the most important factors affecting tiger-leopard competition and coexistence included species richness at various size classes of prey and also the spatial distribution of human disturbance. Local habitat, prey composition, and disturbance factors affect tiger-leopard interactions by shaping the trade-off between ecological opportunities (e.g., easy prey) and the risk of escalating conflict with humans and other competitors across spatial, temporal, and dietary niches.

      Perspectives: There is currently a significant regional bias in the study of tiger-leopard competition and coexistence, with most research focusing on the overlapping range in South Asia and neglecting these in Northeast and Southeast Asia. To more fully understand how tigers and leopards interact, future research should occur longitudinally throughout their shared range, with a focus on multiple ecological niches and fine spatial scales. Research on the biotic and abiotic factors affecting tiger-leopard competition should aim to identify their ecological thresholds and the regulation mechanisms by which these factors affect the intensity and types of competition.

      The evolutionary genetics, taxonomy, and conservation of the Chinese mountain cat
      Ke Liu Sicheng Han He Yu Shujin Luo
      Biodiv Sci. 2022, 30 (9):  22396.  doi:10.17520/biods.2022396
      Abstract ( 94 )   PDF (938KB) ( 63 )   Save
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      Background & Aims: The Chinese mountain cat (Felis bieti) is the only wild felid endemic to China, and it is one of the world’s least studied felids. It is listed as Vulnerable in the IUCN Red List of Threatened Species and is listed as a nationally first-class protected species in China. Here, we reviewed recent advances in the distribution, range, taxonomy, genetic diversity, evolutionary history, and conservation threats to the Chinese mountain cat to provide a much needed scientific basis for conservation efforts.

      Review Results: Scientists have not reached a unanimous agreement on the taxonomic status of the Chinese mountain cat (Felis bieti) since its inception as a species in 1892. Morphological distinctiveness supports its independent species status, yet recent genetic studies revealed that it is equidistant with other currently recognized wildcat subspecies and hence should be considered a wildcat conspecific (Felis silvestris bieti). However, because of the divergent evolution that occurred over one million years between the wildcat subspecies, an alternative approach to resolve the dispute over the Chinese mountain cat’s taxonomy would be to elevate all F. silvestris lineages to the species level, thus retaining the Chinese mountain cat as F. bieti. Nevertheless, this proposition would require a comprehensive analysis of the entirety of the genome data from all wildcat taxa. In addition, a complex admixture scenario was depicted, including an ancient introgression from the Asiatic wildcat (F. s. ornata) to the Chinese mountain cat, as well as a widespread signal of contemporary genetic introgression from F. s. bieti to domestic cats across, but not beyond, the range of F. s. bieti. Regional socioeconomic change in the Tibetan region since the mid-20th century may have facilitated an expansion of domestic cats that had likely recently arrived to the Plateau, setting the stage for their close contact, frequent interaction, and interbreeding with sympatric Chinese mountain cats. This raises concern for conservationists about the opposite direction of gene flow that may pose a threat to the Chinese mountain cat and jeopardize its genetic integrity as well as its evolutionary adaptation to high altitude, an issue with profound conservation implications. The IUCN Red List assessment confirmed the Chinese mountain cat’s range to be restricted to the eastern Qinghai-Tibet Plateau, encompassing the eastern Qinghai, north-western Sichuan, south-western Gansu, and a small area in south-eastern Tibet adjacent to Qinghai. The free-ranging population of the Chinese mountain cat is likely threatened by indirect poisoning by rodenticide, illegal trade, poaching for furs, habitat loss, fragmentation, and road kills caused by grazing and infrastructure development, as well as potential introgression from local domestic cats.

      Conclusion: There is an urgent need to assess the efficacy of current conservation measures in place for the protection of the Chinese mountain cat, including the effectiveness of the management of current protected areas, to safeguard China’s only endemic felid.

      An update on the current distribution and key habitats of the clouded leopard (Neofelis nebulosa) populations in China
      Ziyu Ma Zai-Xin HE Yiqing Wang Dazhao Song Fan Xia Hongxin Su Jian-lin DENG Ping Li Sheng Li
      Biodiv Sci. 2022, 30 (9):  22349.  doi:10.17520/biods.2022349
      Abstract ( 146 )   PDF (1270KB) ( 165 )   Supplementary Material   Save
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      Aims: The clouded leopard (Neofelis nebulosa) is the most dependent species on forest ecosystems among all large felids native to China, its distribution is therefore sensitive to deforestation and its survival needs particular attention in conservation. China hosted the majority of N. nebulosa’s historical range, however with the impact of poaching, deforestation and land use change, the range and population of N. nebulosa had undergone severe contraction and decline throughout the past decades. In addition, relevant research and effective conservation actions were still lacking both in and out China. As the result of the drastic changes in recent decades, the current distribution and survival outlook of this charismatic large cat in China requires urgent and rigorous reassessment in order to raise more awareness and conservation investment to ensure the survival of this species in the future. Specifically, this study has 3 objectives: (1) to review the historical range of N. nebulosa in China since 1950; (2) to assess the current distribution of N. nebulosa in China from 2010 to 2020; (3) to identify the existing potential habitats of N. nebulosa in China and evaluate their habitat suitability in terms of area, quality, deforestation and protected area.

      Methods: (1) To review the historical range of N. nebulosa in China, we searched and examined the occurrence records of N. nebulosa with solid evidence in specimen collections, peer-reviewed publications, local gazetteers and media reports since 1950, and summarized the confirmed years of attainable presence record in each provincial administrative division of China. (2) As for the current distribution of N. nebulosa from 2010 to 2020, we conducted camera-trapping surveys in 55 sites across China, and reviewed contemporary camera-trapping studies in China for confirmed captures of N. nebulosa. (3) To identify the existing potential habitats of N. nebulosa, we joined the results obtained in this study with results from previous studies to generate a presences/absence localities dataset of N. nebulosa in China, and then calibrated the habitat suitability models that was recently published with this dataset to determine the potential habitat patches within China. Lastly, potential habitat patches identified in this manner were overlaid with protection area and administrative divisions, and habitat suitability change was assessed by forest cover change in 2010–2020.

      Results: Back to the 20th century, the distribution of N. nebulosa in China spanned 17 provincial administrative divisions. However, the once widely distributed N. nebulosa can now only be confirmed to continuously occur in 10 study sites of 2 divisions: Yunnan Province and Tibetan Autonomous Region. There has been no evidence for the presence of N. nebulosa in 12 of its historically distributed divisions for more than 20 years. The species’ potentially suitable habitats in mainland China are now restricted to 9 patches with a total area of 64,093 km2, and in only 2 trans-border patches in the southwestern can we confirm the presence of the species from 2010 to 2020: the Himalaya-Western Hengduan-Arakan Mountains and South Wuliangshan-Annamite regions. Although in these 2 patches, the contiguous area abroad (428,511 km2) was much more extensive than that in China (41,373 km2), the extent of annual forest cover loss in 2010‒2020 was slighter in China (0.84%) comparing to abroad (1.57%). In addition, the percentage of protected area cover was higher in China (34.33%) than abroad (22.02%) in these 2 patches.

      Conclusion: From 2010 to 2020, N. nebulosa occurrence in China was confirmed from southwest Tibet, west Yunnan, and south Yunnan, where populations were likely only sustained in transborder habitats with domestic challenges and ongoing severe threats abroad. Relevant departments should strengthen domestic anti-poaching law enforcement, restore habitat suitability and connectivity, and foster transborder collaboration among protected areas, research institutes, governmental organizations and local communities, in order to ensure the long-term survival and growth of N. nebulosa populations in such border regions. In particular, special attention and efforts should be put to Mengla County, Xishuangbanna, Yunnan, where N. nebulosa still exists and 68.36% of its potential habitat is already protected.

      Camera-trapping study of Marbled Cat (Pardofelis marmorata) in Tengchong area of Mt. Gaoligongshan, Yunnan
      Fei Li Xiangyuan Huang Zhang Xingchao Tsz Kin Au Bosco Pui Lok Chan
      Biodiv Sci. 2022, 30 (9):  22089.  doi:10.17520/biods.2022089
      Abstract ( 147 )   PDF (1081KB) ( 130 )   Save
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      Background & Aim: The marbled cat (Pardofelis marmorata) has a highly restricted distribution range in China and is rarely recorded, and very little is known about this elusive wild felid. There has been no published work to analyze the spatial distribution and daily activity pattern of the species in China, which will enhance our understanding of this nationally endangered and protected species, and provide scientific information for conservation management.

      Methods: Gaoligongshan is located in the western part of Yunnan Province bordering Myanmar, extending from the Tibetan Plateau to Indochina. We conducted camera-trap survey at 34 camera-trap stations in the Tengchong Section of Yunnan Gaoligongshan National Nature Reserve and its surroundings between April 2014 and May 2021. We analyzed the spatial distribution of marbled cat in Tengchong Gaoligongshan based on the data acquired. Daily activity patterns of marbled cat and the sympatric and similarly sized leopard cat (Prionailurus bengalensis) were also analyzed using kernel-density estimation.

      Results: During the study period, a total of 230 independent records of marbled cat were obtained in 31,462 camera trap nights, with an encounter rate many-folded higher compared with data from tropical regions. Our study yielded significantly higher encounter rate of marbled cat in the northern portion of Tengchong (i.e. north of 25°30' N), and the species mainly occurred at well preserved evergreen broad-leave forest at 2,300‒3,100 m. Daily activity pattern analysis reveals that marbled cat was mainly diurnal and has bimodal activity peaks at 6:00‒11:00 and 14:00‒19:00, respectively. The sympatric leopard cat was predominantly nocturnal in the northern portion of Tengchong while it tended to be more crepuscular in the southern portion where marbled cat was rare.

      Recommendations: The existence of a stable population of marbled cat in Tengchong Gaoligongshan provides a golden opportunity to enhance our understanding on different aspects of this elusive species, for the benefit of its conservation management. A study using spatially explicit capture-recapture modelling derived from camera-trapping data to estimate its population density would be desirable, and further study should focus on habitat selection, diet and relationships with other sympatric carnivorous species.
      Population dynamic study of snow leopard (Panthera uncia) in Yunta Village, Sanjiangyuan National Nature Reserve, China
      Moyan Chu Shujie Liang Peiyun Li Ding Jia Abudusaimaiti Maierdiyali Xueyang Li Nan Jiang Xiang Zhao Li Faxiang Zhi Lü
      Biodiv Sci. 2022, 30 (9):  22157.  doi:10.17520/biods.2022157
      Abstract ( 97 )   PDF (1346KB) ( 93 )   Supplementary Material   Save
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      Aims: Wildlife population monitoring projects provide important insight and basis for species research and conservation efforts. The snow leopard (Panthera uncia), as the top predator and flagship species of Asian mountainous ecosystems, is of great scientific and conservation interest. However, due to its remote habitat, elusive behavior and large home ranges, long-term population monitoring study on snow leopards is rarely reported. Here we report a long-term population monitoring project of snow leopards in Sanjiangyuan National Nature Reserve (Yunta Village, Haxiu Township, Yushu Prefecture, Qinghai Province) from Oct. 2013 to Jan. 2019.
      Methods: We monitored the population using infrared cameras maintained by the local community. We estimated population size and density as well as population growth rate using spatially explicit capture-recapture model (SECR). We calculated population turnover rates and analyzed territory replacement.
      Results: In total, we identified 35 snow leopard individuals. From SECR, population estimation based on 3-month high-quality data for three consecutive years (2015–2017) suggested stable population dynamic with a growth rate of 1.02. However, individual turnover was obvious with a rate of 0.44, and territory displacement happened around core utilization area of snow leopards. We proposed that the snow leopard population might locate on a potential dispersal pathway of snow leopards, or the camera array only covered a fraction of the population.
      Conclusion: As the first long-term population monitoring project of snow leopards reported in China, our work highlights the importance of long-term monitoring, and demonstrates the feasibility of mammal population monitoring maintained by local communities.

      Population density and activity patterns of the leopard cat (Prionailurus bengalensis) in southern China: Estimates based on camera-trapping data
      Jianhuan Yang King Wa Li Ho Yuen Yeung Tsz Kin Au Xi Zheng Anthony J.Giordano Bosco Pui Lok Chan
      Biodiv Sci. 2022, 30 (9):  21357.  doi:10.17520/biods.2021357
      Abstract ( 523 )   PDF (1332KB) ( 480 )   Save
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      Aim: Reliable estimates of population density are fundamental to wildlife conservation and management. Although the leopard cat is the most common and widespread wild felid in China, little is known about the ecology and population biology of this species in the country. Using spatially explicit capture-recapture (SECR) modelling derived from extensive camera-trapping data, we estimated the population density and activity patterns of leopard cats in a well- protected private nature reserve in southern China.
      Methods: Between June 2020 and May 2021, we conducted a camera-trap survey across a pre-determined grid system in Kadoorie Farm and Botanic Garden (KFBG), Hong Kong. KFBG was established on a barren hillside following sustained anthropogenic disturbances. After six decades of protection, secondary forest currently covers approximately 80% of the site. We deployed a total of 19 camera trap stations across our small 1.5 km2 study area, with two opposite-facing cameras at each station to obtain images of both flanks of leopard cats. The consecutive 12-month survey yielded 113 independent capture events of the leopard cat, of which 61 events were clear enough to facilitate individual identification. Based on closed population assumptions for capture-recapture models, we also divided the 12-month survey into six two-month sampling periods, and estimated the population density for each sampling period using Maximum Likelihood SECR modelling. We also examined activity kernel densities to estimate the difference in diel activity patterns between wet and dry seasons.
      Results: Our analyses revealed that results from two sampling periods were robust and precise enough (i.e., low standard error and narrow 95% confidence intervals) to estimate leopard cat density. We estimated leopard density between June and July 2020 as D = 0.64 ± 0.31 (0.26–1.55) individuals/km2, and between February and March 2021 as D = 0.87 ± 0.48 (0.31–2.40) individuals/km2, which are among the highest density estimates for this species reported in any region. We also found that the diel activity pattern of leopard cats in the study site is arrhythmic during the wet season, but became more nocturnal-crepuscular during the dry season, though they also exhibited some diurnal activity. Kernel density analyses however suggested no significant differences in diel activity patterns occurred between wet and dry seasons.
      Conclusions: Our study provides important early data on the population density of leopard cats in southern China, the results of which allow for comparisons with other studies elsewhere using capture-recapture modelling approaches. We further demonstrate the utility of SECR methods for estimating population density over short and long sampling periods across necessarily small sampling areas. We further provide practical recommendations for conducting camera trap surveys to enhance the success rates of individual identification and “recapture” of leopard cats. In accordance with the conclusion of other studies, our results show that leopard cats are highly adaptable, exhibit great plasticity in daily activity, and thrive well in human-modified landscapes.

      Gut bacterial composition of four leopard cat subpopulations in Beijing
      Fan XIA Jing YANG Jian Li Yang SHI Lixin GAI Wenhua Huang Jingwei ZHANG Nan YANG Fuli Gao Yingying HAN Weidong Bao
      Biodiv Sci. 2022, 30 (9):  22103.  doi:10.17520/biods.2022103
      Abstract ( 80 )   PDF (1218KB) ( 62 )   Save
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      Aims: The composition of gut microbiota of wild animals is most influenced by food types and genetic background. This study aimed to investigate gut bacterial composition of leopard cat (Prionailurus bengalensis) and its influencing factors in Beijing.
      Methods: In this study, fecal samples of leopard cat collected from Yunmengshan, Yunfengshan, Songshan and Baihuashan were analyzed by high-throughput sequencing on bacterial 16S rRNA hypervariable region of V3–V4.
      Results: The results showed that the dominant flora of gut bacteria of leopard cat in phyla included Firmicutes (relative abundance 52.40%), Proteobacteria (25.18%), Actinobacteria (9.07%), Bacteroidetes (8.17%), and Fusobacteria (4.74%). The top 5 genera of bacteria with the highest abundance accounted for about 50% of the total abundance, which were Pseudomonas (13.37%), Blautia (11.20%), Clostridium_sensu_stricto_1 (9.10%), Peptoclostridium (8.62%), and Lactobacillus (6.08%). There was no significant difference in gut bacterial abundance and β diversity among the leopard cat subpopulations in four sampling areas, but the ACE index and Chao 1 index in Songshan subpopulation were different from those in Yunmengshan and Yunfengshan.
      Conclusion: Considering there was a disparity trend in genetic structure in Songshan subpopulation, and the climate types of the four areas and food composition of the leopard cat were highly similar, we postulated that the gut bacteria of leopard cat was mainly influenced by the genetic traits for local adaptation.

      Seasonal variation of daily activity rhythm between leopard cats (Prionailurus bengalensis) and their potential prey in Neixiang Baotianman National Nature Reserve of Henan Province, China
      Xueqin Deng Tong Liu Tianshi Liu Kai Xu Song Yao Xiaoqun Huang Zhishu Xiao
      Biodiv Sci. 2022, 30 (9):  22263.  doi:10.17520/biods.2022263
      Abstract ( 70 )   PDF (1044KB) ( 45 )   Save
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      Aim: The daily activity rhythm of wild animals is an adaptive response to food availability, predation risk, and environmental constraints. Therefore, elucidating the driving factors of daily activity rhythms are crucial in understanding wildlife fitness and conservation. Knowledge about the daily activity rhythms of most wild animals is rudimentary and occasionally inaccurate due to limited sampling and anthropogenic environmental changes. Leopard cats (Prionailurus bengalensis)—formerly believed to be nocturnal—are found to have some diurnal activity, along with their primary crepuscular activity. This illustrates the necessity of investigating how daily activity rhythm in wild animals, such as leopard cats, changes over time and space.

      Methods: During 2016, we deployed 55 infrared-triggered cameras to monitor leopard cats and their potential prey in the Neixiang Baotianman National Nature Reserve, Henan Province. We compared daily activity patterns by employing a kernel density estimation and using overlap coefficient to quantify the temporal overlap between leopards cat and their potential prey across warm and cold seasons.

      Results: We identified 1,343 independent images of leopard cats and their potential prey over 14,972 camera trapping days. We found leopard cats, Tolai hares (Lepus tolai), and nocturnal rats mainly active at night, with the leopard cat’s activity peaking at dusk and dawn. Conversely, Pallas’s squirrels (Callosciurus erythraeus), Pere David’s rock squirrels (Sciurotamias davidianus), golden pheasants (Chrysolophus pictus) and koklass pheasants (Pucrasia macrolopha) were all diurnal. More diurnal activity of leopard cats and their potential prey (except nocturnal rats) occurred during warm seasons. Overlap analysis showed that leopard cats had a higher overlap coefficient with Tolai hares and nocturnal rats across warm and cold seasons (Δ ≥ 0.50) than with any diurnal prey (Δ ≤ 0.40).

      Conclusions: Our study demonstrates that daily activity rhythm for leopard cats and their potential prey may be relatively conserved, but with some seasonal plasticity. This plasticity may be caused by seasonal variation in environment and prey distribution. Our results indicate that broad-scale monitoring and research are essential to elucidate the causes and seasonal variations in daily activity rhythms of wild animals.

      Spatiotemporal overlap among sympatric Pallas’s cat (Otocolobus manul), Tibetan fox (Vulpes ferrilata) and red fox (V. vulpes) in the source region of the Yangtze River
      Dong Wang Gaowa Saiqing Zihan Wang Hongxiu Zhao Xinming Lian
      Biodiv Sci. 2022, 30 (9):  21365.  doi:10.17520/biods.2021365
      Abstract ( 464 )   PDF (2007KB) ( 275 )   Save
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      Aim: Pallas’s cat (Otocolobus manul), Tibetan fox (Vulpes ferrilata) and red fox (V. vulpes) are small sympatric carnivores inhabiting the Sanjiangyuan region on the Qinghai-Tibet Plateau. The spatiotemporal overlaps between these species were analyzed in this study.
      Methods: From June 2014 to September 2019, we set up 208 infrared cameras along the Tuotuo River and the Tongtian River in the source region of the Yangtze River to collect the daily activity rhythm and spatial distribution data of three carnivores. The spatial overlap coefficient was used to evaluate spatial niche differentiation and the kernel density estimation method was utilized to evaluate daily activity rhythm. Then, the overlap coefficient of rhythms was applied to analyze the temporal niche differentiation among the three carnivores.
      Results: The spatial overlap coefficients between Pallas’s cat and Tibetan fox, Pallas’s cat and red fox, and Tibetan fox and red fox were 0.25, 0.48 and 0.17, respectively. This indicated that there were differences in space utilization among the three carnivores. The results of daily activity rhythm demonstrated that Pallas’s cats and Tibetan foxes displayed diurnal behavioral patterns and red foxes were primarily nocturnal. The overlap coefficients of daily activity rhythms of Pallas’s cats, Tibetan foxes, and red foxes between the cold and warm seasons were 0.83, 0.78 and 0.88, respectively. Between any two species, the highest overlap coefficient (0.84) of daily activity rhythm was between the Pallas’s cats and Tibetan foxes, and the lowest overlap coefficient (0.48) was between the Tibetan foxes and red foxes. A lower coefficient meant more significant temporal niche differentiation. Additionally, the overlap coefficients of daily activity rhythm between the two species were smaller in the warm season than those in the cold season.
      Conclusion: In the source region of the Yangtze River, the three small sympatric carnivores could reduce interference and competition through spatial or/and temporal niche differentiation and thus achieve coexistence.

      Overview of monitoring methods for Tigers, Leopards and Ungulate prey
      Wei Yi Guangshun Jiang
      Biodiv Sci. 2022, 30 (9):  21551.  doi:10.17520/biods.2021551
      Abstract ( 150 )   PDF (1038KB) ( 112 )   Save
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      Background & Aim: Species conservation efforts require an understanding of population sizes. For predators such as tigers and leopards, it is also vital to monitor populations of their ungulate prey. In recent years, monitoring methods for tigers and leopards mainly entailed derivations from capture-recapture models based on data from noninvasive sampling and camera traps. Meanwhile,  ungulate prey populations were monitored using line transects, belt transects, sample plots, camera trapping, and also using the capture-recapture model based on data from noninvasive genetic methods. Each monitoring technique has different assumptions and is based on varying ecological principles, and, as a consequence, accuracy also varies. Thus the applicability of each method depends on the biological characteristics and population distribution of study species, monitoring targets, and spatial scales of interest. We present details about common monitoring methods, including specific procedures and statistical principles. We also analyze advantages and disadvantages of these methods to provide reference for selecting the appropriate one for monitoring projects going forth.

      Review Results: While monitoring population of tiger, leopard, and ungulate populations, it is necessary to select appropriate methods based on the population status of the target species in the focal area (i.e., considering distribution law and dynamic characteristics) and monitoring targets. For monitoring of tigers and leopards, for instance, excessive intensity in the study area may result in pseudoreplication, which may lead to habitat disturbance and waste of resources. Meanwhile, using automated-camera traps set for monitoring tigers and leopards to assess ungulate population size may be inappropriate.

      Perspectives: Population surveys on tigers, leopards and ungulates can benefit from combining monitoring methods to achieve more accurate population estimations in the future. Recent advances in artificial intelligence technologies, in particular, are helpful for increasing monitoring accuracy by automatically identificating individual tigers and leopards and prevent repeated counts. Not only can this save labor costs, but it is also much higher in accuracy compared to traditional methods.
      Determining the daily activity pattern of Chinese mountain cat (Felis bieti): A comparative study based on camera-trapping and satellite collar tracking data
      Yueqiao Kong Yanlin Liu Chengwu He Tianti Li Quanliang Li Cunxin Ma Dajun Wang Sheng Li
      Biodiv Sci. 2022, 30 (9):  22081.  doi:10.17520/biods.2022081
      Abstract ( 168 )   PDF (1281KB) ( 129 )   Save
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      Aims: For elusive and difficult to observe wild cats, camera trap and satellite tracking collar are widely used sampling methods to determine daily activity patterns. While satellite tracking collars can continuously monitor a limited number of captured individuals, camera traps have the potential to monitor all population members in a given area. The detection probability of camera traps for wild cat activity patterns is likely influenced by the selected spatial placement of the cameras. The activity patterns are constructed based on the binary classification of target animals’ active status (i.e., active or inactive). Although the different methods of monitoring wild cats may lead to different activity patterns, few empirical studies have examined the difference between the patterns. The Chinese mountain cat (Felis bieti), a small felid endemic to China, is distributed in the eastern and northern edge of the Qinghai-Tibet Plateau at low density. The Chinese mountain cat was uplisted to Class-I National Key Protected Wildlife of China in 2021, is listed as Vulnerable (VU) by the IUCN Red List, and is considered Critically Endangered (CR) in China’s Red List. However, the existing surveys and data are insufficient to carry out conservation of Chinese mountain cat. Therefore, urgent field investigations about the ecological activities of the Chinese mountain cat are needed. 
      Methods: From June 2020 to December 2021, we investigated activity patterns of Chinese mountain cats in Qilian Mountains of Qinghai Province using camera traps and satellite-tracking collars. We compared daily activity patterns derived from camera traps placed at den sites (2, 173 independent detections) and animal trails (23, 423 independent detections), to satellite-tracking collars fitted on Chinese mountain cats (10 individuals, 62,942 fixed locations).
      Results: Results show Chinese mountain cats were active all day long, with peak activity from 17:00 to 19:00. A daily activity curve produced by camera traps showed a high level of overlap with results produced by the activity sensor implanted in the collars; the coefficient of overlap is 0.89. However, the camera traps showed significantly higher levels of activity from 16:00 to 21:00. Detection of activity from 16:00 to 21:00 was primarily concentrated at a small range of den sites. Concurrently, activity records of the collars were relatively lower. Based on the ethogram of Chinese mountain cat, activity patterns recorded from the camera traps placed at den sites predominantly represents social behaviors related to reproduction and nursing while activity records from the trail camera traps generally represent movement and foraging. The satellite-tracking collar was limited to distinguishing just the movement and still of an individually tracked animal.
      Conclusions: Our results indicate considerable differences between the activity patterns of wild cats obtained via camera-trapping and a satellite collar tracking. The binary classification of “active” or “inactive” used to construct activity patterns reflect different behaviors in data derived from different methods. Caution should be taken for researchers when comparing the activity patterns derived from different studies. Proper and accurate interpretation of data should be made on the basis of in-depth and comprehensive understanding of the target animal’s behaviors.

      Habitat use of the North Chinese leopard (Panthera pardus japonensis) in the Liupanshan Mountains and its implications for conservation planning
      Shuanggui Wang Zhihong Guo Bojian Gu Tianti Li Yubing Su Bocheng Ma Hongxin Guan Qiaowen Huang Fang Wang Zhuojin Zhang
      Biodiv Sci. 2022, 30 (9):  22342.  doi:10.17520/biods.2022342
      Abstract ( 62 )   PDF (1095KB) ( 56 )   Save
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      Aims: Large carnivores play important roles in ecosystem functions yet are globally threatened and require urgent research and conservation actions. The North China leopard (Panthera pardus japonensis) is a leopard subspecies that is endemic to China. It is the only remaining large carnivore in many forest ecosystems in northern China, and the North China leopard facing severe threats such as habitat degradation and fragmentation. This study aims to assess the habitat use of North China leopards in the Liupanshan Mountains.
      Methods: This study was conducted in the Liupanshan Mountains. Infrared cameras were used to survey North China leopard distribution. We constructed occupancy models to analyze species habitat use. Based on species-environmental associations obtained from occupancy modeling, we identified suitable habitat patches, measured species association with human interference, and evaluated habitat fragmentation patterns.
      Results: Occupancy modeling revealed that North China leopards occupied habitat patches with mature forest in ragged terrain. The occurrence probability was high in landscapes that were distant from croplands and roads, but the species did not demonstrate significant avoidance to farmland edges and residential areas. We identified 35 suitable habitat patches that were primarily distributed along the east and west ridges of the Liupanshan Mountains. The largest suitable habitat patch had an area of 214 km2, and the average area of suitable habitat patches were 16 km2. Approximately 55% of all suitable habitat patches were located within the Liupanshan National Nature Reserve boundary.
      Conclusions: The primary conclusion of this study is that while the Liupanshan National Nature Reserve effectively conserves existing suitable habitat of North China leopards, the risks of habitat fragmentation and anthropogenic interference still exist. We suggest conservation actions such as habitat capacity building and a restriction of anthropogenic activity in protected area be taken to ensure the long-term persistence of North China leopards. Conservation efforts beyond provincial boundaries should be reinforced to promote the dispersal of North China leopards as well as population recovery. This study fills in the knowledge gaps left by the North China leopards ecological study in the Ningxia Hui Autonomous Region, and has the potential to support conservation planning in the Liupanshan Mountains as well as other regions.

      Spatiotemporal patterns of cattle depredation by the North Chinese Leopard in Taihang Mountains and its management strategy: A case study in Heshun, Shanxi Province
      Yiqing Wang Ziyu Ma Gang Wang Yanlin Liu Dazhao Song Bei-bei LIU Lu Li Xinguo Fan Qiaowen Huang Sheng Li
      Biodiv Sci. 2022, 30 (9):  21510.  doi:10.17520/biods.2021510
      Abstract ( 269 )   PDF (1414KB) ( 246 )   Supplementary Material   Save
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      Background & Aim: Human‒wildlife conflict is a global challenge facing wildlife conservation today, and requires case-specific solutions for mitigation. To effectively manage the conflicts and prevent future incidents, thorough understandings of its patterns and causes, the locals’ attitudes toward the conflicts and factors that may affect people’s behaviors are needed. Large carnivores can be perceived as problematic as they may cause severe human-wildlife conflicts through livestock depredation and even direct human injury or death. Livestock depredation is one of the main causes of human‒carnivore conflicts. If managed inefficiently, it will lead to retaliatory killings toward the “problematic” large carnivores, thus imposing a threat to the species’ survival. Elimination of these apex predators will subsequently lead to the losses of critical functions they play in the ecosystem. Taihang Mountains is the core distribution area of the North Chinese leopard (Panthera pardus japonensis), the leopard subspecies endemic to China. Cattle depredation by leopards frequently occurs in this area and is therefore a challenge facing North Chinese leopard conservation.
      Methods: Our study area is in Heshun County, Shanxi Province, locating in the middle part of the Taihang Mountains. Since 2015, Shanxi Tieqiaoshan Provincial Nature Reserve, Heshun Ecological Conservation Association, Government of Mafang Town and Chinese Felid Conservation Alliance jointly launched a compensation project for cattle depredation by leopards in Heshun. Information for each cattle depredation event was recorded during this project from 2015 to 2019. A camera-trap network was also set up in the same area (from 2016 to 2019) to examine the spatial distribution of the relative abundance of cattle, North Chinese leopard and leopard’s primary wild prey, i.e. Siberian roe deer (Capreolus pygargus) and wild boar (Sus scrofa). In 2020, we interviewed 107 households who received money from compensation about their attitudes toward leopards, cattle depredation and the compensation project. With these data, we examined the spatiotemporal patterns of cattle depredation events and conducted a spatial risk model to evaluate what factors that may increase the probability of depredation incidents. We then mapped the risk of cattle depredation by leopards to the entire study area. We also investigated the relationship between herd management and the number of cattle injured/dead per household. As for the economic and social aspects, we calculated the economic impacts of cattle depredation on Heshun’s livestock husbandry industry and on individual households, and examined people’s attitudes toward leopards and their perceptions towards the risk of cattle depredation. At last, we provided specific suggestions for the mitigations of human-wildlife conflicts in Heshun.
      Results: We identified 195 cattle depredation events from 2015 to 2019 and compensated 116 households with a total of 270.5 thousand Yuan RMB. We found that the frequency of depredation was higher during the summer, during which the cattle were ranged freely in the mountain. The risk of cattle depredation decreased with the increase in the relative abundance of Siberian roe deer and increased with distance to villages, significantly (P < 0.05). The relative abundance of cattle was the third most important factor in predicting the risk. The boundary of protected area and the relative abundance of leopards are the two least important factors, both of which were not present in the 15 models with ΔAIC ≤ 2. Besides, the practice of herd management may also have an impact on this risk, as herders with loose management tend to lose more cattle from depredation (P = 0.056). We found that the impact of cattle depredation on Heshun’s animal husbandry industry was minor as a whole. However, individual households who have experienced cattle depredation might bear a remarkable economic loss. This had led to individuals’ negative attitudes towards leopards, as well as complaints about insufficient compensation.
      Recommendations: In order to mitigate the impacts of livestock depredation by leopards as well as reduce future incidents, we proposed the following recommendations. Firstly, strengthening cattle management by, for example, constraining the free-ranging cattle within a certain distance away from villages and enclosing cattle in captivity at night during the free-ranging season to reduce the risk of depredation by leopards. Secondly, designating non-grazing zones where livestock grazing is strictly prohibited, thus reducing the risk of livestock depredation and allowing the recovery of the leopard’s wild prey population. Thirdly, continuing the cattle depredation compensation project to mitigate the economic burden incurred. In addition, carrying out regular visits to the local communities to alleviate the emotional impacts suffered by the locals, and conducting natural education to promote positive attitudes towards leopards and other wildlife. Finally, conducting wildlife population monitoring, especially for leopard’s natural prey, and implementing conservation measures when necessary to maintain their population stability. We believe these management strategies may effectively reduce the risk of livestock depredation and help promoting the coexistence between human and wildlife in the long term.

      Why do we believe in Darwin's theory of evolution——On the 25 folds of aesthetic parsimony of On the Origin of Species
      Yajun Sun
      Biodiv Sci. 2022, 30 (9):  22243.  doi:10.17520/biods.2022243
      Abstract ( 61 )   PDF (850KB) ( 87 )   Save
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      Background & Aim: Essentially, a contest between scientific theories is a contest for aesthetic parsimony, which makes scientific revolutions possible. By analyzing Darwin’s On the Origin of Species, I demonstrate the inner rationality of the greatest revolution in the history of life sciences.
      Progress: The analysis shows the 25 parsimonious advantages of Darwin’s theory of evolution over creationism, reflected in explaining (1) the appearance of domestic races, (2) that the extent of the difference in some external character tends to be larger than that in some internal part among domestic races, (3) the morphological changes of offspring especially produced by crossing, (4) the differences in vigour/fertility between hybrids and mongrels, (5) the differences in the extent of morphological change among offspring by different kinds of crossing, (6) the clustered distribution of some morphological character among taxonomic categories, (7) that the extent of morphological differentiation is unequal among taxonomic categories, (8) that the extent of morphological differentiation is unequal among different morphological characters, (9) the gradual variation of traits among species, (10) the morphological changes accompanying environmental changes, (11) the morphological inertia relative to environmental changes, (12) the unequal distribution of competitive intensity among organisms, (13) the morphological homology among species, (14) that the morphological differences among individuals tend to increase with individual development, (15) the biological vestiges or rudiments, (16) the systematical disparity (cross taxonomic ranks) of biogeographic distribution, (17) the relation between the “hierarchical status” of a species and its distributional range, (18) the “island phenomenon” in biogeographic distribution, (19) the relatively lower species richness and higher ratio of endemic species in an island compared to those in a similar-sized area of a mainland, (20) the specific types of organisms that can be found or not in an island compared to that in a mainland, (21) the similarity between the organisms of an island and that of the neighbouring mainland, (22) the geographic heterogeneity of species migration and species extinction, (23) the gradual process of species extinction, (24) the morphological relation between extinct and extant organisms, and (25) the morphological similarity between the embryo of an organism and its ancestor.
      Conclusion: It is these 25 folds of aesthetic parsimony that have cast the essence of On the Origin of Species as a greatest scientific work, and have provided the modern life-science research with the universal methodology and these general directions, i.e., the paradigm. This analysis also helps us understand the genuine spirit of scientific development and manage the contemporary scientific research.

      On the methodology of Darwinism and Non-Darwinism
      Ruiwu Wang
      Biodiv Sci. 2022, 30 (9):  22482.  doi:10.17520/biods.2022482
      Abstract ( 47 )   PDF (748KB) ( 89 )   Save
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      Turbid evolutionary theories and their tidy textbook reconstruction
      Huajie Liu
      Biodiv Sci. 2022, 30 (9):  22535.  doi:10.17520/biods.2022535
      Abstract ( 31 )   PDF (689KB) ( 45 )   Save
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