Is the clouded leopard Neofelis nebulosa extinct in Taiwan, and could it be reintroduced? An assessment of prey and habitat
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Is the clouded leopard Neofelis nebulosa extinct in
Taiwan, and could it be reintroduced? An assessment
of prey and habitat
PO-JEN CHIANG, KURTIS JAI-CHYI PEI, MICHAEL R. VAUGHAN
CHING-FENG LI, MEI-TING CHEN, JIAN-NAN LIU, CHUNG-YI LIN
L I A N G - K O N G L I N and Y U - C H I N G L A I
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
revealed , km of suitable habitat but this was reduced to
, km when adjacent areas of human encroachment
T he clouded leopard Neofelis nebulosa is categorized as
Vulnerable on the IUCN Red List and is listed on
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 neckrikulau@gmail.com 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
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. https://doi.org/10.1017/S003060531300063X262 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 abun-
dance 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 reintro-
duction of clouded leopards. We used the number of photographic events per camera-
trap day (O’Brien et al., ) as an index of relative abun-
dance 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 –, m over 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
Oryx, 2015, 49(2), 261–269 © 2014 Fauna & Flora International doi:10.1017/S003060531300063X
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. https://doi.org/10.1017/S003060531300063XReintroduction of clouded leopard 263
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 % of the prey biomass index. We hy-
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 R v. . (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 and summed to give an overall score for human activity
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 a value of 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
Oryx, 2015, 49(2), 261–269 © 2014 Fauna & Flora International doi:10.1017/S003060531300063X
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. https://doi.org/10.1017/S003060531300063X264 P. Chiang et al.
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
Oryx, 2015, 49(2), 261–269 © 2014 Fauna & Flora International doi:10.1017/S003060531300063X
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. https://doi.org/10.1017/S003060531300063X. https://doi.org/10.1017/S003060531300063X
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Oryx, 2015, 49(2), 261–269 © 2014 Fauna & Flora International doi:10.1017/S003060531300063X
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.
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
Prey species weight (g)1 weight (kg)2 (n = 72) (n = 49) (n = 43) (n = 22) Kruskal–Wallis P Jonckheere–Terpstra P
Prey . 4 kg
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
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
Muntiacus reevesi
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 , 2 kg
White-faced flying squirrel 1,370 1.522 0.0002 0.0000 0.0016 0.0105 , 0.0001 0.0004
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
Reintroduction of clouded leopard
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
265266 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 –.
Relative abundance index Model* ΔAICc Akaike weight R2
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 . 4 kg 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 ,2 kg
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. –% of a
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
Oryx, 2015, 49(2), 261–269 © 2014 Fauna & Flora International doi:10.1017/S003060531300063X
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. https://doi.org/10.1017/S003060531300063XReintroduction of clouded leopard 267
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
Oryx, 2015, 49(2), 261–269 © 2014 Fauna & Flora International doi:10.1017/S003060531300063X
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