The Current Distribution and Status of Leopards Panthera Pardus in China

The current distribution and status of leopards Panthera pardus in China

A LICE L AGUARDIA,JAN F. KAMLER,SHENG L I ,CHENGCHENG Z HANG Z HEFENG Z HOU and K UN S HI

Abstract The leopard Panthera pardus, categorized globally Introduction as Near Threatened on the IUCN Red List, has the widest distribution of any wild felid species, although in Asia he leopard Panthera pardus has the widest it has declined dramatically and five subspecies are Tdistribution of any wild felid species, ranging from Endangered or Critically Endangered. In China at least western and southern Africa to the Russian Far East and  three subspecies have been reported to occur throughout Java (Stein & Hayssen, ). This is largely a result of the ’ much of the country, and in  the population was esti- species high adaptability, as it can occupy diverse ecosys- mated to be ,. However, recent studies have indicated tems ranging from tropical rainforests to boreal forests  that leopards have disappeared from large areas, probably and arid savannahs (Bertram, ) and take a large variety  as a result of habitat loss, a low prey base and poaching, in- of prey species (Hayward et al., ). Despite these charac- dicating this species may not be as common in China as pre- teristics, the leopard has declined dramatically worldwide  viously believed. To examine this we reviewed recent (Stein & Hayssen, ), and disappeared from at least literature and interviewed specialists to determine the cur- one-third of its historical range in Africa (Ray et al.,  rent status and distribution of the leopard in China. Our ). In Asia, leopards have exhibited dramatic declines   findings indicate that the species has declined dramatically, in the Middle East (Khorozyan, ; Mallon et al., ),  with confirmation of presence at only  sites in  pro- Russian Far East (Jackson & Nowell, ), Sri Lanka (Kittle   vinces, despite extensive surveys. Current populations are & Watson, ) and Java (Ario et al., ), and small and fragmented, and occur mainly in isolated nature consequently subspecies occupying these regions are cate- reserves. We estimate a total population of only – gorized as Endangered or Critically Endangered on the P. pardus japonensis (the north Chinese leopard), which is IUCN Red List. Although the leopard still occupies much  endemic to China, and ,  individuals for each of the of its historical distribution in India (Karanth et al., ), ’ other subspecies whose distributions extend beyond the species status is unknown across large regions of Asia, China. We recommend that a separate IUCN assessment particularly in China where data on leopards are outdated be made for P. pardus japonensis, and that this subspecies and limited. be categorized as Critically Endangered. Our findings are Historically, the leopard was distributed throughout the first reliable estimates of the current distribution and China, with the exception of the arid Gobi desert and moun- .   status of the leopard in China, and provide valuable infor- tainous western regions at elevations , m. At present  mation that will help guide conservation efforts. the species reportedly occurs in at least provinces (Bao et al., ) and is on the list of fauna for many protected Keywords Amur leopard, China, conservation, Indochin- areas. Despite these optimistic reports, there are few recent ese leopard, north Chinese leopard, Panthera pardus,Red records in most of these areas (Li et al., ; Jutzeler et al., List status ). Comparisons of several recent leopard surveys with a – To view supplementary material for this article, please visit nationwide mammal assessment carried out during  http://dx.doi.org/./S concluded that most of the existing habitat is no longer suitable and local extinctions have occurred in several regions across the leopard’s range (Ran & Chen, ; Liu et al., a,b, ). Therefore it is highly unlikely that

ALICE LAGUARDIA,CHENGCHENG ZHANG and KUN SHI (Corresponding author) The the only available estimate of the leopard population in Wildlife Institute, Beijing Forestry University, Tsinghua-east Rd 35, Beijing China (,;Ma,) is still reliable. 100083, China. E-mail [email protected] Lack of information on the current distribution and JAN F. KAMLER Wildlife Conservation Research Unit, Department of Zoology, population of a species can undermine conservation efforts, The Recanati-Kaplan Centre, University of Oxford, Abingdon, UK; Panthera, New York, USA as such information is essential for developing appropriate strategies to maintain viable populations. The aim of our SHENG LI School of Life Sciences, Peking University, Beijing, China study therefore was to conduct a review of records of the ZHEFENG ZHOU Wocheng Institute of Ecology and Environment, Taiyuan,  Shanxi, China leopard in China from onwards, to determine the spe- ’ Received  April . Revision requested  June . cies current status and distribution and to make recom- Accepted  July . First published online  October . mendations for its conservation.

Oryx, 2017, 51(1), 153–159 © 2015 Fauna & Flora International doi:10.1017/S0030605315000988 Downloaded from https://www.cambridge.org/core. IP address: 170.106.33.14, on 27 Sep 2021 at 20:36:34, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0030605315000988 154 A. Laguardia et al.

Subspecies dilemma subject to bias (Farrell et al., ; Davison et al., ; Prugh & Ritland, ; Harrington et al., ; Laguardia According to phylogenetic classifications (Miththapala et al., ) and therefore extreme caution was adopted   et al., ; Uphyrkina et al., ) there are at least three, when categorizing records (e.g. consideration of the experi- and probably four, subspecies of leopard in China but ence of the person making the observation, presence of sym- their geographical boundaries are unclear. The Amur leop- patric carnivores). ard P. pardus orientalis occupies the north-east, as far south The results confirmed the presence of the leopard in  as the area north of Beijing. The Indochinese leopard provinces (Fig. ; Supplementary Table S), in a total of  P. pardus delacouri occurs in the south-east, presumably in- locations where the species has been detected using camera cluding all of Yunnan Province and as far north as the Pearl traps, carcasses of poached individuals were recovered, or River (Zhu Jiang in Chinese) in Guangxi and Guangdong sightings or scat identifications were made by experienced  provinces (Miththapala et al., ). The most widespread researchers. Thirty-three of the records were in nature re- subspecies in China is the north Chinese leopard P. pardus serves or county forests and  in unprotected sites japonensis, commonly referred to as P. pardus fontanierii in (Supplementary Table S). An additional six provinces the Chinese literature. This subspecies is endemic to China had potential (Fig. ; Supplementary Table S) areas of leop- and occurs in the central and eastern regions, presumably as ard occurrence, based on interviews with local staff, ques- far north as the Beijing area, and as far south as the Pearl tionnaires to resident communities, livestock depredation River, including Sichuan and Guizhou provinces. The sub- reports or habitat quality assessments. In  locations the specific status of leopards in Xizang (Tibet) Autonomous leopard was considered absent (Supplementary Table S) Region (hereafter, Tibet) is uncertain. Although this popu- because no signs were found, although there has been insuf-  lation could be P. pardus fusca (Miththapala et al., ; ficient survey effort in some parts of the historical distribu-  Uphyrkina et al., ), this subspecies has been considered tion. Although most camera-trap surveys occurred within synonymous with P. pardus delacouri in southern China protected areas, this is unlikely to bias our results because  (Smith & Xie, ). However, the Tibet population is sepa- remaining forest habitat and potential prey species are pri- rated from the main distribution of P. pardus delacouri by marily restricted to these areas. high mountain ranges, although the Tibet population occu- We summarize the records (details in Supplementary pies similar habitat and shares a continuous distribution Table S) and distributions for each subspecies (Fig. ): with P. pardus japonensis. Genetic research is necessary to clarify the subspecies of the Tibet population; here, we consider it to be P. pardus japonensis. Leopards from south- Amur leopard P. pardus orientalis central Tibet, however, such as those near Mount Everest, are almost certainly P. pardus fusca. In Jilin and Heilongjiang provinces leopards were probably extirpated by the s (Jutzeler et al., ) but recent efforts to create large nature reserves along the Chinese– Current distribution Russian border have made it possible for individuals to cross into China. Camera traps have recorded leopards in Searches were conducted for newspaper articles, scientific several areas of Jilin Province (Huang et al., ; Xiao reports and peer-reviewed publications, in English or et al., a; Beijing Normal University, ), and recent Chinese, published from  onwards, which contained sign surveys and camera traps have recorded leopards in records of the leopard in China. We searched Google south-eastern Heilongjiang Province (Jutzeler et al., ; Scholar () for the terms ‘panthera pardus AND china Beijing Normal University, ). OR japonensis OR delacouri OR orientalis OR fusca’ for –, resulting in c. , items. As this database does not include all Chinese journals we also searched the North Chinese leopard P. pardus japonensis China National Knowledge Internet () with the key- word ‘jinqianbao’ (‘leopard’ in Chinese) for –, re- In eastern and central China leopard populations are greatly sulting in , items. The titles of items, then the abstracts reduced and highly fragmented. Recent camera-trap surveys and finally the full articles were filtered by relevance to our and other evidence confirmed the presence of the subspecies search objectives, reducing the number of items consider- in only eight provinces (from north to south): northern ably. The corresponding authors of key publications and Hebei, Shanxi, Shaanxi, Ningxia, northern Henan, western key felid experts were contacted for additional information. Sichuan, southern Qinghai, and Tibet. Most populations in A final total of  publications were used (Supplementary these provinces are small, and occur in isolated protected Table S) and seven experts provided additional information areas, and it is unknown whether these subpopulations are vi- as personal communications or unpublished data. Identifi- able in the long term. Although some records were from out- cation of species records from sightings, scats and signs is side protected areas, it is probable that these subpopulations

FIG. 1 The current distribution of subspecies of the leopard Panthera pardus in China: O, P. pardus orientalis;J,P. pardus japonensis;D,P. pardus delacouri; F, P. pardus fusca.

function as sinks as a result of the higher human disturbance and s leopards were recorded in Wolong Nature and lower prey numbers, similar to the situation reported for Reserve (Schaller et al., ; Johnson et al., ), one of tigers Panthera tigris in India and Nepal (Karanth et al., ). the largest protected areas in Sichuan, where intensive re- In Hebei Province camera traps recorded leopards in  search on giant pandas Ailuropoda melanoleuca has taken and , ,  km from Beijing (T. Song, unpubl. data); the place. However, multiple camera trapping studies during status in other regions of the province is unknown but num- – did not record leopards (Li et al., ) and bers are likely to be low. Most records were attributed to they are probably extirpated from the reserve. Camera- Shanxi Province, where extensive camera trapping during trapping studies in an additional nine protected areas in – identified leopard populations in  protected central Sichuan during – also failed to record leo- areas across the province (Sohu News, ; Song et al., pards, even though they were on the official mammal lists of ; Sina News, ; Xinhuanet News, ; Z. Zhou, un- the reserves, indicating this species might be extirpated from publ. data). In the neighbouring Shaanxi Province there large areas of the province (Li et al., ; Xiao et al., b). were leopard records from six nature reserves in addition In Tibet leopards were recently recorded in the east (S. Li, to two verified attacks on people and two leopards killed pers. comm.). There are also several recent records of leo- by poachers (Sohu News, , ; Li et al., ;Wu pards from south-central Tibet (Hu et al., ), including et al., ; Liu et al., ; Xinhuanet News, ; S. Li, the forest zone of Mt Everest (Hou, ). The records pers. comm.). In Ningxia Autonomous Region there were from south-central Tibet are probably subpopulations of  recent records of leopards (Gao et al., ; CCTV, P. pardus fusca, however. In Qinghai Province, data from ; China Economic Net, ). Leopards also occur in local interviews suggested leopards occur in the south the northern part of Henan Province, where eight leopards (D. Wang, pers. comm.). were killed during –, in addition to multiple sight- There are no recent confirmed records from the other pro- ings and reports of livestock predation by leopards (Wang vinces in central or eastern China. In Gansu Province Liu et al., ). Habitat restoration efforts have increased the et al. (a) did not find any leopard sign during an prey base (mostly hare Lepus sp. and wild boar Sus scrofa) -day field survey in key areas during December – and could have had a positive effect on the leopard popula- March . However,  unconfirmed leopard occurrences tion in this area (Wang et al., ; Hou, ). (reports of leopard tracks and leopard attacks on livestock) In Sichuan Province the current status of leopards is un- were recorded in the last  years. The leopard may be extir- clear, and all recent records are from the west, including pated from most of its former range in Gansu and its poten- camera trap images and fresh scats. Intensive camera trap- tial distribution limited to areas on the border with Shaanxi ping is underway (S. Li, pers. comm.) to identify the subspe- and Sichuan. Leopard tracks were reported in  in south- cies’ current distribution in the province. During the s central Inner Mongolia but there are no other records despite

camera traps having been used extensively in this province for the number of presumed P. pardus fusca in south-central studies of Eurasian lynx Lynx lynx and other carnivores. Tibet is probably small, also with ,  individuals. Leopards appear to have been extirpated throughout all There are an estimated  P. pardus japonensis in Henan other provinces in central and eastern China, including Province (Wang et al., ; Hou., ), – in southern Hunan, Hubei, Zhejiang, Fujian, Guangxi and Jiangxi. In Ningxia (Gao et al., ), – in southern Shanxi Jiangxi Province one leopard was recorded by a security (Z. Zhou, unpubl. data),  in a county forest in northern camera in  or  (S. Li, pers. comm.) but afterwards Shanxi (Song et al., ),  in Fujian (Liu et al., ), no further leopards were recorded there despite extensive  in Guizhou (Ran & Chen, ) and ,  in Zhejiang camera trapping and therefore this population is probably (Liu et al., ). There are no estimates available for any now extirpated (Tilson et al., ). Recent camera trapping of the other provinces or nature reserves. The information surveys did not detect leopards in Hunan (Tilson et al., collected from articles, reports and personal communica- ; Dahmer et al., ; Liu et al., ) or Guangxi tions is not sufficient to provide a population estimate for (Wang et al., ). Based on records of livestock losses, in- P. pardus japonensis. Therefore, we estimated a population terviews with witnesses and habitat quality, some research- size range for this subspecies based on the number of con- ers have suggested that potential leopard populations firmed leopard locations ( sites), the mean area of the sites   existed in nature reserves in Zhejiang, Fujian and Guizhou ( km ) and densities of  and  leopard per  km .We  provinces (Ran & Chen, ; Liu et al., b; Liu et al., chose  leopard per  km as a minimum density because ). However, recent camera trapping in several nature that was the estimated leopard density in a  study of leo- reserves in these provinces failed to detect leopards pards in Lishan Nature Reserve in southern Shanxi Province (Tilson et al., ; D. Song, pers. comm.) and the species (A. Laguardia, unpubl. data.) and the density reported for is probably extirpated from these provinces. leopards in Bhutan (Wang & Macdonald, ) in similar  forested mountain habitat. We chose  per  km as a max- imum density to account for the possibility of higher leopard Indochinese leopard P. pardus delacouri densities in some areas with more favourable habitat and prey numbers. Leopard densities are unlikely to be .  per In the south-east this subspecies has recently been recorded   km anywhere in China. Our estimates gave a total popu- in camera traps in two nature reserves in south-western lation of – for confirmed areas. If we also include potential Yunnan Province near the border with Myanmar (The  leopard locations ( sites, mean area =  km ) then the total Wildlife Institute, Beijing Forestry University, unpubl. population estimate is –. data; Jutzeler et al., ) but the population is low (prob- The reasons for the decline of leopard populations and ably ,  individuals in each reserve) and is unlikely to re- their disappearance from many areas in China have yet to cover because of the high levels of habitat fragmentation and be investigated. Leopards are known to be successful at poaching, and low prey numbers. There are no other recent adapting to altered habitat and can persist as long as there records of P. pardus delacouri in south-eastern China and is an adequate prey base, but they cannot withstand intense this subspecies might be on the verge of extirpation in the persecution. Retaliatory killings as a result of human–wild- country. The distribution and status of this subspecies is life conflicts and poaching for the wildlife trade are reported currently being assessed in the remaining countries of its in several provinces (Gao et al., , Liu et al., a,Wang distribution (J.F. Kamler, unpubl. data). et al., , Xinhuanet News, ). In addition, low prey numbers, especially of wild ungulates, are reported even in Population status and major threats many nature reserves. Loss and fragmentation of habitat through logging, farming, mining, expanding settlements The previous nationwide estimate of , leopards (Ma, and road construction are also consistently described as ) is now outdated. The available information that exists the main threats. Consequently, leopard subpopulations is limited to specific areas or nature reserves. Particular ef- are now more vulnerable because they are small and fort has been dedicated to surveying for P. pardus orientalis isolated, with unsuitable habitat between them. in north-eastern China, using both camera traps and sign surveys, and initial results have suggested a population of Conservation actions – (Huang et al., ; Xiao et al., a; Jilin Forestry Department, WCS, & WWF, unpubl. data). Additionally, Panthera pardus is categorized as Near Threatened on the  individual Amur leopards were identified during – IUCN Red List (Henschel et al., ), although five subspe-  (Beijing Normal University, ; Wang et al., ), cies, all from Asia, are categorized as Endangered or although many of these might be cross-border individuals Critically Endangered. For China assessment of intraspecific and not resident leopards. The total number of P. pardus de- taxa is available only for P. pardus orientalis, which is cate- lacouri in south-eastern China is probably , . Similarly, gorized as Critically Endangered (Jackson & Nowell, ).

As a result of the immediate risk of extinction highlighted by University), Charlotte Whitham (Beijing Forestry our review, we recommend that P. pardus japonensis should University) and the staff of Wocheng Institute of Ecology have a separate subspecies assessment. With its recent and and Environment. dramatic range reduction in China, an estimated population of –, and with no subpopulation .  individuals, we recommend P. pardus japonensis is categorized as Critically References Endangered based on criteria Ab,c and Ca(i) (IUCN, ). The China Species List has already categorized ARIO, A., SUNARTO,S.&SANDERSON,J.() Panthera pardus   Panthera pardus as Critically Endangered and it is a Class melas.InThe IUCN Red List of Threatened Species v. . . Http:// www.iucnredlist.org [accessed  September ]. I protected species and among  particularly important spe- BAO, W., XU, J., CUI,G.&FRISINA, M.R. () Cat research in China. cies targeted for conservation and restoration (Wang & Xie, Cat News,S, –. ; Lu et al., ). BEIJING NORMAL UNIVERSITY () Academic News: Monitoring Compared to other Panthera species, such as the snow and protection go hand in hand for decades, the Siberian tiger  leopard Panthera uncia and tiger, leopards receive little at- population returns to growth. Http://www.bnu.edu.cn/xzdt/ . html [accessed  March ]. [In Chinese] tention and limited funding (Jutzeler et al., ). The inter- BERTRAM, B.C.B. () Leopard. In The Encyclopedia of Mammals national conservation community in particular has not yet (ed. D.W. Macdonald), pp. –. Andromeda Oxford Limited, recognized the dramatic decline of this species (Stein & Oxford, UK. Hayssen, ), especially in China. To ensure the long-term CCTV () Guyuan, Ningxia: ran into leopard while on patrol,    survival of leopards in China, information about their popu- expert confirmed identification. Http://news.cntv.cn/ / / / VIDE.shtml [accessed  February ]. [In lation dynamics and habitat requirements in relatively small Chinese] isolated reserves is needed so that effective conservation ac- CHINA ECONOMIC NET () Netizen photographed leopard again in tion can be implemented. Liupanshan. Http://www.ce.cn/celt/wyry///t_ Similar to other large carnivore species, leopards have .shtml [accessed  March ]. [In Chinese] large home ranges and naturally occur at relatively low dens- CHINA NATIONAL KNOWLEDGE INTERNET () Http://www.ckin.   ities, requiring large areas to persist. However, their persist- net [accessed March ]. DAHMER, T.D., GUI,X.&TIAN,S.() Camera-trapping for south ence in China will probably be limited to nature reserves as a China tiger in Hupingshan National Nature Reserve, Hunan result of extensive habitat loss and low prey numbers outside Province, China. Chinese Journal of Wildlife, , –. these areas. Therefore, to avoid problems related to small DAVISON, A., BIRKS, J.D.S., BROOKES, R.C., BRAITHWAITE, T.C. & and fragmented populations (e.g. inbreeding depression), MESSENGER, J.E. () On the origin of faeces: morphological habitat and leopard prey in nature reserves will need to be versus molecular methods for surveying rare carnivores from their scats. Journal of Zoology, , –. restored and expanded. To foster connection between na- DUTTA, T., SHARMA, S., MALDONALDO, J.E., WOOD, T.C., PANWAR, ture reserves, corridors containing suitable habitat need to H.S. & SEIDENSTICKER,J.() Gene flow and demographic be established to allow movement and increase gene flow history of leopards (Panthera pardus) in the central Indian between otherwise isolated populations (Dutta et al., ). highlands. Evolutionary Applications, , –.  Priority should also be given to increasing patrols against FARRELL, L.E., ROMAN,J.&SUNQUIST, M.E. ( ) Dietary separation of sympatric carnivores identified by molecular analysis poaching. Although it is has been illegal to kill leopards in of scats. Molecular Ecology, , –.  China since , poaching incidents are still reported, indi- GAO, E., HU, D., WANG,Z.&BAI,Q.() An investigation of cating that persecution of the species has yet to be elimi- leopard in Liupan Mountain Nature Reserve. Forest Resources nated. National surveys for wildlife are ongoing in China, Management, , –. [In Chinese]   and our findings, together with new data, will help prioritize GOOGLE SCHOLAR ( ) Http://scholar.google.com [accessed March ]. conservation efforts and ensure appropriate measures are HARRINGTON, L.A., HARRINGTON, A.L., HUGHES, J., STIRLING,D.& taken to secure the leopard in China. MACDONALD, D.W. () The accuracy of scat identification in distribution surveys: American mink, Neovison vison, in the northern highlands of Scotland. European Journal of Wildlife Research, , –. Acknowledgements HAYWARD, M.W., HENSCHEL, P., O’BRIEN, J., HOFMEYR, M., BALME, G. & KERLEY, G.I.H. () Prey preferences of the leopard We thank all the researchers that contributed valuable (Panthera pardus). 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Guangguangshan ValleyofLongxi-Hongkou National Nature Reserve, species distribution modelling. J AN F. KAMLER is interested in the south-west China. Biodiversity Science, , –.[InChinese] conservation of large carnivores and their prey. He is especially inter- XINHUANET NEWS () Another rare leopard killed in China. Http:// ested in how large carnivores affect the ecology of smaller carnivores news.xinhuanet.com/english/china/-//c_.htm and their prey, and what effects this has on ecosystems. S HENG L I is [accessed  October ]. a wildlife researcher working on the ecology and conservation of large XINHUANET NEWS () Leopard was photographed for the first time forest mammals, especially carnivores and ungulates, in south-western in Pangquangou National Nature Reserve. Http://www.sx.xinhuanet. China. C HENGCHENG Z HANG is interested in using genetic methods com/newscenter/-//c_.htm [accessed  March for a deeper understanding of carnivores and their conservation. ]. [In Chinese] Z HEFENG Z HOU is founder of the Wocheng Institute for Ecology and Environment, focusing on biodiversity monitoring and conserva- Biographical sketches tion in Shanxi province. KUN SHI, as Secretary-general of the China Cats Specialist Group, is leading a feline research group at the A LICE L AGUARDIA’s research interests include the conservation Wildlife Institute of Beijing Forestry University, focusing on conserva- of fragmented populations, non-invasive genetic techniques and tion of the snow leopard, leopard and tiger.