Is the Clouded Leopard Neofelis Nebulosa Extinct in Taiwan, and Could It Be Reintroduced? an Assessment of Prey and Habitat

Home , Formosan black bear, Neofelis

Is the clouded leopard Neofelis nebulosa extinct in Taiwan, and could it be reintroduced? An assessment of prey and habitat

P O -JEN C HIANG,KURTIS J AI-CHYI P EI,MICHAEL R. VAUGHAN C HING-FENG L I ,MEI-TING C HEN,JIAN-NAN L IU,CHUNG-YI L IN L IANG-KONG L IN and Y U -CHING L AI

Abstract During – we conducted a nationwide become over-abundant. Thus, it is important to address camera-trapping survey and assessed the availability of the cascading effect of the disappearance of top-down pred- prey and habitat for the clouded leopard Neofelis nebulosa ator control. Our assessment indicated that, with proper in Taiwan. We surveyed , camera-trap sites over regulation of hunting, habitat restoration and corridor im- , camera-trap days, from the seashore to an altitude provement, it may be possible to reintroduce the clouded of , m and covering various types of vegetation. No leopard. clouded leopards were photographed during , cam- Keywords Camera-trapping, extinct, habitat assessment, era-trap days, including at  sites in other studies, con- Neofelis nebulosa, prey distribution, reintroduction, Taiwan firming the presumed extinction of clouded leopards in Taiwan. Assessment of the prey base revealed altitudinal distribution patterns of prey species and prey biomass. Areas at lower altitudes and with less human encroachment and hunting supported a higher prey biomass and more of Introduction the typical prey species of clouded leopards. Habitat analysis he clouded leopard Neofelis nebulosa is categorized as    revealed , km of suitable habitat but this was reduced to Vulnerable on the IUCN Red List and is listed on    T , km when adjacent areas of human encroachment Appendix I of CITES (). It is the largest felid in were subtracted. In the absence of hunting and large mam- Taiwan and has been categorized as Endangered under malian carnivores the major prey of clouded leopards in the Wildlife Conservation Act since . Lee & Lin () Taiwan, such as Formosan macaques Macaca cyclopis, suggested that clouded leopards were nearly or already ex- ’ Reeves s muntjacs Muntiacus reevesi, Formosan serow tinct in Taiwan; no direct records of occurrence (e.g. sight- Capricornis swinhoei and sambar Rusa unicolor, could ings, photographs, carcasses) had been reported since  and pre- records were based on interviews. However, for a species to be categorized as Extinct, IUCN stipulates PO-JEN CHIANG* (Corresponding author) and KURTIS JAI-CHYI PEI Institute of that exhaustive surveys must be conducted in its range Wildlife Conservation, National Pingtung University of Science and ’ Technology, Pingtung, Taiwan. E-mail [email protected] over a time frame appropriate to the taxon s life cycle  MICHAEL R. VAUGHAN Department of Fisheries and Wildlife Sciences, Virginia (IUCN, ). No field surveys of clouded leopards have Polytechnic Institute and State University, Blacksburg, Virginia, USA been conducted in Taiwan and information from local peo-  CHING-FENG LI Department of Botany and Zoology, Masaryk University, Brno, ple is anecdotal and unsubstantiated (Rabinowitz, ). Czech Republic The population status of the species cannot be ascertained MEI-TING CHEN Institute of Biological Resources, National Pingtung University without field surveys but if any clouded leopards remain of Science and Technology, Pingtung, Taiwan they can only be in very small numbers and in remote areas. JIAN-NAN LIU Taiwan Endemic Species Research Institute, Nantou, Taiwan The clouded leopard’s prey species have experienced CHUNG-YI LIN Institute of Ecology and Evolutionary Biology, National Taiwan pressure from habitat loss as a result of human encroach- University, Taipei, Taiwan ment and from hunting, although commercial hunting LIANG-KONG LIN Department of Life Sciences, Tunghai University, Taichung, was banned in  (Lee & Lin, ). However, populations Taiwan of some prey species, including Formosan macaques YU-CHING LAI Department of Environmental and Hazards-Resistant Design, Huafan University, New Taipei, Taiwan Macaca cyclopis and some mammalian herbivores, have reportedly increased, causing conflict on farmlands and in *Also at: Formosan Wild Sound Conservation Science Centre, 2F, No. 335, Yongmei Road, Yangmei City, Taoyuan County, Taiwan forests. Although these increases could be a result of Received  November . Revision requested  January . the hunting ban, reduced predation pressure from the disap- Accepted  April . First published online  November . pearance of clouded leopards cannot be ruled out (Crooks &

Oryx, 2015, 49(2), 261–269 © 2014 Fauna & Flora International doi:10.1017/S003060531300063X Downloaded from https://www.cambridge.org/core. IP address: 170.106.40.40, on 30 Sep 2021 at 04:28:55, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S003060531300063X 262 P. Chiang et al.

Soule, ; Chiang et al., ). Disappearance of large apex (Kansas, USA) in some of the surveys, and by  film predators can impose trophic cascades on an ecosystem, camera-traps had been replaced completely by digital. The with effects on biodiversity (Terborgh et al., ; Estes digital camera-traps were set at c. . m height for horizontal et al., ). Thus, it is important to assess the population detection, to accommodate their sensor design. Such horizon- status of the clouded leopard and locate any surviving indi- tal detection may result in higher variations in the detection viduals. Reintroduction of clouded leopards to Taiwan range, given the variable terrain. With film camera-traps tilted could be considered if extirpation is confirmed. downwards, the detection area was more consistent for prey The success of any conservation action and/or reintro- comparison across sites. Thus, only film camera-traps were duction of clouded leopards would depend on the avail- used to assess the prey base, to reduce bias from the variable ability of suitable prey and habitat. The discovery of detection areas at sites with digital cameras. clouded leopards would necessitate conservation actions A more extensive camera-trap survey was conducted in such as habitat improvement and/or restocking the popu- the Tawu Mountain area during – to study prey lation to increase genetic diversity (Roelke et al., ). If populations under the most natural conditions, without it were confirmed that no clouded leopards remain, suitable human disturbance and hunting, using stratified sampling reintroduction sites should be identified and habitat quality according to altitude (Chiang et al., ). Several camera- and connectivity improved. Our objectives were () to inves- trap sites were baited with live chickens and other olfactory, tigate whether there are any remaining clouded leopards in visual or auditory lures to increase the chances of photo- Taiwan, () to document the altitudinal distribution pat- graphing clouded leopards. terns of major potential prey species, () to assess the abundance of prey species, () to test whether prey populations  are being reduced by anthropogenic hunting, and ( )to Prey assessment identify suitable habitat for restoration and for the reintroduction of clouded leopards. We used the number of photographic events per camera- trap day (O’Brien et al., ) as an index of relative abundance of prey because of its correlation with population Study area densities of mammalian herbivores (O’Brien et al., ;  Rowcliffe et al., ; Rovero & Marshall, ). Taiwan is an orogenic island of c. , km , with a large Consecutive photographs of the same species within  altitudinal range and a maximum altitude of c. , m. The hour were counted as one event and multiple consecutive vegetation gradient changes from coastal plains and lowland pictures of groups of animals (e.g. Formosan macaques) tropical and subtropical rainforest to temperate coniferous were also counted as single events. The relative abundance forest and alpine grassland at the highest altitudes. The index of each prey species was multiplied by the edible per- Tawu Mountain area in southern Taiwan (Fig. ) encom- centage of mean adult body weight (g) and summed across passes two protected areas: Tawu Nature Reserve and all prey species, for use as a prey biomass index. Edible per- Twin Ghost Lake Important Wildlife Area. It contains the centage was determined as % for prey . kg, % for largest remaining lowland primary forest and is the location prey . kg and % for prey , kg (Emmons, ; in which, if clouded leopards still occur on the island, they Pedersen et al., ; Mills et al., ). However, we esti- are most likely to be found (Rabinowitz, ). There is mated that the maximum amount of meat a clouded leopard minimal human disturbance in the area and it is primarily could obtain from large prey (assuming multiple feeding covered by pristine Ficus–Machilus, Machilus–Castanopsis events) was  kg, based on the daily meat consumption and Quercus forests and Tsuga rainforests along the altitudi-  and kill rates of other wild felids. nal gradient –, mover km . To investigate altitudinal patterns of prey distribution and abundance under natural conditions, without hunting, Methods we compared relative abundance indices for each prey and carnivore species and prey biomass indices between four –  Camera-trap surveys altitudinal zones in the Tawu Mountain area ( , , ,–,, ,–, and ,–, m). These four We conducted camera-trap surveys in national parks, pro- zones reflect the four major vegetation types in the area. tected areas and fragmented lowlands throughout Taiwan We used Kruskal–Wallis tests to identify if there were sig- during –, mostly using film cameras developed in nificant differences between relative abundance indices Taiwan. Film camera-traps were set at c.  m height and tilted and prey biomass indices between the zones. We used at –° to face the trail or intersection of trails. In  Jonckheere–Terpstra tests to identify if there were mono- we started using digital camera-traps from Cuddeback tonic patterns of relative abundance indices and prey bio- (Wisconsin, USA), Reconyx (Wisconsin, USA) and Bushnell mass indices along the altitudinal gradient. We performed

FIG. 1 Locations of camera-trap sites across Taiwan. The inset shows the location of Taiwan off the coast of China.

one-sided Wilcoxon rank–sum tests to examine the effects accounted for almost %ofthepreybiomassindex.Wehy- of hunting on relative abundance indices and prey biomass pothesized that prey biomass could be influenced by human indices in hunted and non-hunted areas at altitudes ,, activity, distance to central Taiwan and altitude. We used m in the Tawu Mountain area. We used Rv.. (R multiple linear regression and the information-theoretic ap-  Development Core Team, ) to conduct statistical analy- proach (Akaike information criterion, AICc) to compare the sis. Jonckheere–Terpstra tests were performed in SPSS v. full model with three independent variables and models with (SPSS, Chicago, USA). each variable removed in turn. We also conducted a meta-analysis of prey biomass index, In our models the value for human activity was based on including data from other camera-trap studies in Taiwan three factors: accessibility from roads and villages ( or ), (Wang, , ; Wu et al., ; Wang & Hsu, ; hunting pressure (–), and history of forest practices or agri- Wang & Huang, ;Hwang&Chian,), to understand cultural use (–). These three factors were each scaled to factors influencing prey biomass across the country. The prey  andsummedtogiveanoverallscoreforhumanactivity biomass index used in this meta-analysis was based on the five (–). Accessibility from roads and villages was assigned largest prey species (macaques and ungulates) because limited avalueof if the area was within  km of major roads or data were available for some areas. These five species well-maintained logging roads or within  km of aboriginal

villages, and a value of  if the area was outside these ranges. prey were photographed (Table ). We excluded other car- Hunting pressure was assigned a value of  if there was no nivores as prey because clouded leopards have not been ob- hunting,  if there was occasional hunting,  if there was per- served killing carnivores, nor have carnivores been found in sistent seasonal hunting (i.e. every year during October–April) clouded leopard scats (Griffiths, ; Nowell & Jackson, and  if there was persistent hunting all year round. The history ; Grassman et al., ). of forest practices was determined from literature and field ob- In the Tawu Mountain area we identified a significant servations and was based on the estimated percentages of forest decreasing trend in prey biomass index with increasing alti- alteration for agriculture, plantations and clear-cutting. tude (Jonckheere–Terpstra test, P , .; Table ). Reeves’s muntjacs Muntiacus reevesi had the highest relative abundance index (Table )at,, m and accounted for Habitat assessment .% of the prey biomass index at ,, m. Formosan ma- We identified areas of suitable habitat for clouded leopards caques were the second most important contributor to the ,  based on the species’ habitat requirements (Nowell & relative abundance index at , m but contributed less Jackson, ; Grassman et al., ). Natural broadleaf forest, than Formosan serow Capricornis swinhoei and sambar mixed broadleaf–conifer forest (primary or secondary) and Rusa unicolor to the prey biomass index because they are old-growth cypress forest were considered potential suitable not as heavy. Although wild pigs Sus scrofa were the second habitats. We excluded coniferous forest, which usually oc- largest prey, because of their low relative abundance index curs at ., m in Taiwan, non-forested areas, agricultural their contribution to the prey biomass index was lower ,  land, bamboo forest and clear-cut plantation forest (mostly than that of the other four prey species at , . conifers). We identified potentially suitable habitat using a m. Macaques and the four ungulates contributed %of digitized vegetation map produced by Taiwan Forestry the total prey biomass index. Bureau in . Based on data from other studies (home- We detected significant decreasing trends in relative range and core-area sizes and distances covered daily; abundance index with increasing altitude for macaques, Grassman et al., ; Austin et al., ) we included in muntjacs, Chinese pangolins Manis pentadactyla,  ’ our analyses forest patches . km and fragmented Swinhoe s pheasants Lophura swinhoii, red-bellied tree  patches – km that were within  km of contiguous pri- squirrels Callosciurus erythraeus, and spinous country rats – ,   mary habitat. Given that clouded leopards may hunt at for- Niviventer coninga (Jonckheere Terpstra test, all P . ;  est edges (Grassman et al., ) we considered a  m Table ). Although there were significant altitudinal differ- buffer along the boundaries of areas of suitable habitat. ences, no monotonic linear trends were observed for serow Human encroachment and hunting are common and per- and sambar. Only Taiwan white-faced flying squirrels sistent near villages and along roads, and therefore we con- Petaurista alborufus lena, Formosan white-bellied rats sidered that suitable habitat must be at least  km from Niviventer culturatus and Taiwan partridges Arborophila villages and  km from major roads. If clouded leopards crudigularis showed increasing relative abundance indices – used areas adjacent to villages and major roads they would along the altitude gradient (Jonckheere Terpstra test, all ,    probably have been observed. P . ; Table ); however, these are not important prey species nor do they weigh . . kg. At altitudes ,, m in the Tawu Mountain area the Results prey biomass index was significantly lower in hunted than non-hunted areas (P , .; Table ) because of Camera-trap surveys the significantly reduced abundance of ungulate and pri- mate species (except wild pigs) in hunted areas (Table ). –   During we established , camera-trap sites In contrast, none of the smaller prey species, which were   over , camera-trap days in various parts of Taiwan, not targeted by hunters, had significantly lower relative    from the coast to , m altitude. These included abundance indices in hunted areas than in non-hunted   sites ( , camera-trap days) in the Tawu Mountain areas (Table ).  area. The total effort, including sites from other surveys, Based on camera-trap data from across the country, the      was , sites (Fig. ) and , camera-trap days. No prey biomass indices of the five largest prey species decreased camera traps recorded clouded leopards.   ,   as human activity increased (F, = . ,P . ). The full model, which included all three variables (human ac- Prey assessment tivity, distance to central Taiwan and mean altitude), best ex- plained countrywide variations in the prey biomass indices Our camera-trap survey yielded a comprehensive record of of these five species (Table ) and all three factors were sig- potential mammalian and avian prey. Twelve mammal spe- nificant. In summary, reduced hunting pressure and human cies and two phasianid bird species identified as potential encroachment, lower altitudes and closer proximity to

https://www.cambridge.org/core TABLE 1 Prey species in descending order by weight, with maximum edible weight, mean adult weight, relative abundance index (no. of photographic events per camera-trap day) in four zones of altitude in Tawu Mountain area, Taiwan, during – (Fig. ), P for Kruskal–Wallis test for altitudinal differences, and P for Jonckheere–Terpstra test for monotonic patterns among the four zones. –

269 Relative abundance index © . Maximum edible Mean adult 150–1,200 m 1,200–2,000 m 2,000–2,500 m 2,500–3,100 m https://doi.org/10.1017/S003060531300063X 04Fua&FoaItrainldoi:10.1017/S003060531300063X International Flora & Fauna 2014 Prey species weight (g)1 weight (kg)2 (n = 72) (n = 49) (n = 43) (n = 22) Kruskal–Wallis P Jonckheere–Terpstra P Prey . 4kg

. IPaddress: Sambar 50,000 165 0.0251 0.0342 0.0291 0.0016 0.0032 0.1594 Rusa unicolor Wild pig 28,470 43.8 0.0093 0.0040 0.0074 0.0089 0.1644 0.4988

170.106.40.40 Sus scrofa Formosan serow 18,200 28 0.0792 0.0868 0.0403 0.0858 0.0040 0.5522 Capricornis swinhoei Reeves’s muntjac 8,000 10 0.6006 0.1962 0.1200 0.0069 , 0.0001 , 0.0001 , on Muntiacus reevesi

30 Sep2021 at04:28:55 Formosan macaque 7,200 9 0.0931 0.0984 0.0532 0.0374 0.0087 0.0023 Macaca cyclopis Chinese pangolin 3,600 4.5 0.0056 0.0046 0.0000 0.0000 0.0025 0.0003 Manis pentadactyla Prey , 2kg White-faced flying squirrel 1,370 1.522 0.0002 0.0000 0.0016 0.0105 , 0.0001 0.0004 , subjectto theCambridgeCore termsofuse,available at Petaurista alborufus lena Swinhoe’s pheasant 990 1.1 0.0244 0.0363 0.0069 0.0027 0.0007 0.0007 Lophura swinhoii Red-bellied tree squirrel 324 0.36 0.0107 0.0157 0.0000 0.0000 0.0007 0.0002 Callosciurus erythraeus Long-nosed tree squirrel 324 0.36 0.0024 0.0028 0.0010 0.0044 0.5639 0.8893 Dremomys pernyi owstoni Taiwan partridge 281 0.312 0.0048 0.0083 0.0134 0.0115 0.0694 0.0132

Arborophila crudigularis 265 leopard clouded of Reintroduction Striped tree squirrel 63 0.07 0.0000 0.0000 0.0013 0.0017 0.2222 0.0697 Tamiops maritimus Formosan white-bellied rat 61 0.068 0.0000 0.0062 0.0182 0.0088 , 0.0001 , 0.0001 Niviventer culturatus Spinous country rat 61 0.068 0.0650 0.0949 0.0004 0.0000 , 0.0001 , 0.0001 Niviventer coninga Total relative abundance index 0.9204 0.5879 0.2926 0.1801 Total prey biomass index 8491.1 5750.0 3758.8 2241.7 , 0.0001  Minimum of % of body weight and  kg for large prey and % of body weight for small prey  Based on our own field data or literature 266 P. Chiang et al.

TABLE 2 Prey species in descending order by weight, relative abun- TABLE 3 Linear-regression models of prey biomass index, based on dance indices for non-hunted and hunted areas at altitudes data for macaques, sambar, Reeves’s muntjacs, Formosan serow ,, m in the Tawu Mountain area, Taiwan (Fig. ), during and wild pigs from camera-trapping studies conducted at  sites –, and Wilcoxon rank-sum P for hunting impacts. across Taiwan during –. Δ 2 Relative abundance index Model* AICc Akaike weight R Non-hunted Hunted Wilcoxon HA + DCT + ALT 0.00 0.887 0.714 area area rank-sum HA + DCT 5.48 0.057 0.604 Prey species (n = 121) (n = 22) P HA + ALT 5.51 0.056 0.604 Prey . 4kg ALT + DCT 27.18 0.000 0.141 Sambar 0.0288 0.0011 0.0004 *HA, index of human activity; DCT, distance to central Taiwan; ALT, mean Wild pig 0.0071 0.0151 0.8334 altitude Formosan serow 0.0823 0.0374 0.0082 Reeves’s muntjac 0.4368 0.1452 ,0.0001 Formosan 0.0953 0.0703 0.0385 clouded leopard is concentrated in southern and eastern macaque Taiwan. Chinese pangolin 0.0052 0.0000 0.0479 Prey ,2kg White-faced flying 0.0001 0.0000 0.3420 Discussion squirrel Swinhoe’s 0.0292 0.0196 0.3276 The mean number of camera-trap days required to record a pheasant clouded leopard or Sunda clouded leopard Neofelis diardi in Red-bellied tree 0.0128 0.0088 0.4832 other South-east Asian countries is –, with as few as squirrel – camera-trap sites (Lynam et al., ; Kawanishi & Long-nosed tree 0.0026 0.0000 0.1095 Sunquist, ; Rao et al., ; Azlan & Lading, ; squirrel Azlan & Sharma, ; Johnson et al., ; Cheyne & Taiwan partridge 0.0062 0.0125 0.9200   Striped tree 0.0000 0.0011 0.9910 Macdonald, ). Carbone et al. ( ) suggested that   squirrel , camera-trap days was sufficient time to detect tigers Formosan white- 0.0025 0.0024 0.7050 Panthera tigris at low population densities (.–. per  bellied rat  km ). The area of suitable habitat for clouded leopards  Spinous country 0.0771 0.1344 0.4261 in Taiwan is c. , km . If there were only one tiger in this rat area the predicted camera-trapping effort to detect its pres- , Prey biomass index 7381.1 2871.2 0.0001 ence would be c. , trap days. Given our camera- trapping effort of , camera-trap days without success, central Taiwan supported higher biomass of clouded leopard it is unlikely that clouded leopards still exist in Taiwan.  prey. Carbone et al. ( ) suggested that carnivores weighing ,. kg feed mostly on prey that is #% of their own weight. Adult Formosan macaques and Reeves’s muntjacs   – Habitat assessment have a mean weight of . kg, which is c. %ofa clouded leopard’s body weight (– kg) and is approxi- The total area of suitable habitat for clouded leopards, ex- mately equal to the mean weight of confirmed prey of  cluding small, isolated fragments, was c. , km (%of clouded leopards, based on scat analyses and field observa- the total land area of Taiwan). After applying the  m buf- tions (Griffiths, ; Grassman et al., ). Formosan fer and excluding areas around roads and villages the area of serow, which are similar in weight to clouded leopards,  potential high-quality habitat was reduced to , km could also be an important prey species as clouded leopards  (Fig. a). The largest continuous block is , km , in cen- have been observed feeding on goats cached in trees tral/eastern Taiwan and the second largest block, which en- (Hazarika, ). Although clouded leopards are reportedly compasses the Tawu Mountain area in southern Taiwan, is capable of killing sambar weighing  kg (Swinhoe, ;  , km (Fig. a). However, a larger portion of the Tawu Kano, ; Nowell & Jackson, ), a skull analysis sug- Mountain area is at altitudes ,, m compared to other gested that large prey need to be partially restrained for patches (Fig. b). Areas of suitable habitat that are unen- clouded leopards to deliver a powerful bite at the nape of croached by humans are fragmented and isolated by the neck (Therrien, ). Eurasian lynx Lynx lynx regularly roads, agricultural lands and coniferous plantation forests, prey on reindeer Rangifer tarandus up to – times their particularly at altitudes ,, m and more so at altitudes body weight but the reindeer are generally those in poor ,, m(Fig. c), where the most abundant prey are body condition (Pedersen et al., ). We speculate that found. In summary, the most suitable habitat for the sambar preyed upon by clouded leopards are mostly

FIG. 2 (a) Distribution of suitable habitat for the clouded leopard Neofelis nebulosa in Taiwan, excluding areas close to roads and villages, (b) suitable habitat at , , m altitude, and (c) suitable habitat at , , m altitude.

smaller, weaker or younger individuals. Thus, the import- the past for commercial purposes, and hunting was wide- ance of sambar in terms of prey biomass index may be in- spread before the Wildlife Conservation Act of  was en- flated. Wild pigs had low relative abundance indices and forced. Prey depletion would have been a threat to the may be too aggressive to be targeted even by large felids survival of clouded leopards in Taiwan over the past decades such as leopards (Hayward et al., ). or even centuries (Karanth & Stith, ). Potential clouded leopard prey in Taiwan can be divided Three of the larger prey species (muntjacs, macaques and into two categories by weight (Table ): $  kg (macaques pangolins) were more abundant at lower altitudes. Sambar, and ungulates) and # kg (birds and rodents). Chinese serow and wild pig did not show altitudinal differences in pangolin, arboreal flying squirrels and smaller carnivores abundance. However, as altitude increased, the three largest are excluded because they had low relative abundance indi- prey species (i.e. heavier than a clouded leopard) comprised ces or there were no confirmed records of them being preyed a higher percentage of total available prey (Table ). This upon by clouded leopards (Griffiths, ; Nowell & may not be good for clouded leopards because these larger Jackson, ; Grassman et al., ). For prey in the # prey may be difficult to catch (Griffiths, ; Carbone et al., kg category only Swinhoe’s pheasant weighs c.  kg; all the ). As the prey base showed a decreasing trend with alti- others weigh ,. kg. Their contributions to the total prey tude, we suggest that the primary habitat for clouded leo- biomass index was, on average, ,.% of that of the five lar- pards is at lower altitudes, particularly ,, gest prey species for all altitudes in the Tawu Mountain area. m. However, both clouded leopards and their major prey Clouded leopards may not spend much time travelling in populations have suffered habitat loss and hunting pressure rugged and steep terrain to search for prey ,. kg as this as a result of timber harvesting and human encroachment. would not be optimal for maximizing their energy intake Thus, habitat loss and prey depletion are the two main fac- (Griffiths, ). Therefore, larger prey species, particularly tors driving the extinction of the clouded leopard in Taiwan. macaques, muntjacs and serow, would be of greatest import- As a result of increasing awareness of the need for en- ance to the survival of clouded leopards in Taiwan. vironmental protection, logging of natural forests in However, these major prey species are targeted by hun- Taiwan was halted in . Forests have been undergoing ters. The prey biomass index of hunted areas at altitudes of regeneration and succession, becoming suitable habitats ,, m could be as low as that at the highest altitudes, for clouded leopards and their prey again. If habitat around where there is no hunting and clouded leopards would areas of human encroachment could be better managed rarely occur. Therefore, hunting could reduce the prey and/or restored to a less fragmented condition, the best- base to an unfavourable level for clouded leopards. The quality habitat for clouded leopard in Taiwan could cover  five largest prey species, plus Chinese pangolins and flying c. , km or more. Such an area could be inhabited by squirrels, were hunted both extensively and intensively in – clouded leopards, based on population density

 estimates from Thailand (.  per  km ; Austin & Tewes, for species-level distinctions in clouded leopards. Current Biology,  ), or up to  ( per  km , %CI–; Wilting , –. et al., ), based on density estimates for the Sunda CARBONE, C., CHRISTIE, S., CONFORTI, K., COULSON, T., FRANKLIN, N., GINSBERG, J.R. et al. () The use of photographic rates to clouded leopard in Borneo. However, these are conservative estimate densities of tigers and other cryptic mammals. Animal estimates, because leopards would also utilize nearby areas Conservation, , –. in addition to the best-quality habitat assessed. CARBONE, C., MACE, G.M., ROBERTS, S.C. & MACDONALD, D.W. Populations of the clouded leopard’s prey species are () Energetic constraints on the diet of terrestrial carnivores.  – also recovering as a result of the ban on commercial hunt- Nature, , . CHEYNE, S.M. & MACDONALD, D.W. () Wild felid diversity and ing, and thus, given also the recovery and protection of habi- activity patterns in Sabangau peat-swamp forest, Indonesian tat, the reintroduction of clouded leopards in Taiwan may Borneo. Oryx, , –. potentially be considered. The Tawu Mountain area, CHIANG, P.J., PEI, K.J.C., VAUGHAN, M.R. & LI, C.F. () Niche where vegetation coverage and prey levels are suitable and relationships of carnivores in a subtropical primary forest in  – human activities are limited, would be the ideal place for in- southern Taiwan. Zoological Studies, , . CITES () Convention on International Trade in Endangered itiating the re-establishment of the clouded leopard in Species of Wild Fauna and Flora. Appendices I, II and III. Http://  Taiwan. As genetic (Buckley-Beason et al., ; Wilting cites.org/eng/app/appendices.php [accessed  November ]. et al., ) and morphological research (Kitchener et al., CROOKS, K.R. & SOULE, M.E. () Mesopredator release ) did not identify clouded leopards in Taiwan as a dis- and avifaunal extinctions in a fragmented system. Nature, , – tinct subspecies, animals from mainland Asia could be uti- . EMMONS, L.H. () Comparative feeding ecology of felids in a lized for reintroduction. As wildlife conservation is a neotropical rainforest. Behavioral Ecology and Sociobiology, , priority issue there, and considering the ongoing recovery –. of habitat and prey, Taiwan could become a global refuge ESTES, J.A., TERBORGH, J., BRASHARES, J.S., POWER, M.E., BERGER, J., for clouded leopards. BOND, W.J. et al. () Trophic downgrading of planet earth. Science, , –. GRASSMAN, L.I., TEWES, M.E., SILVY, N.J. & KREETIYUTANONT,K. () Ecology of three sympatric felids in a mixed evergreen forest Acknowledgements in north-central Thailand. Journal of Mammalogy, , –. GRIFFITHS,D.() Foraging costs and relative prey size. 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