Population density estimates and conservation concern for clouded leopards Neofelis nebulosa, marbled cats Pardofelis marmorata and tigers ...
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Population density estimates and conservation
concern for clouded leopards Neofelis nebulosa,
marbled cats Pardofelis marmorata and tigers
Panthera tigris in Htamanthi Wildlife Sanctuary,
Sagaing, Myanmar
HLA NAING, JOANNA ROSS, DAWN BURNHAM
S A W H T U N and D A V I D W . M A C D O N A L D
Abstract The clouded leopard Neofelis nebulosa is a potent Keywords Ambassador species, camera trapping, clouded
ambassador species for conservation, occurring from the leopard Neofelis nebulosa, density estimate, marbled cat
Himalayan foothills eastwards to Indochina, between Pardofelis marmorata, spatial capture–recapture, tiger
which Myanmar is a biogeographical land bridge. In Panthera tigris
Myanmar’s Northern Forest Complex, the species co-occurs
with the tiger Panthera tigris, leopard Panthera pardus,
marbled cat Pardofelis marmorata, golden cat Catopuma Introduction
temminckii and leopard cat Prionailurus bengalensis. We
deployed cameras within the Htamanthi Wildlife
Sanctuary over consecutive years. In – we
deployed camera stations around the Nam Pa Gon stream
A guild of wild species of Felidae comprising various
combinations of up to eight species (Macdonald
et al., ) is distributed across South-east Asia, with
(Catchment ) for , trap days. In – we deployed species ranging in size from the tiger Panthera tigris to the
camera stations around the Nam E Zu stream (Catchment flat-headed cat Prionailurus planiceps. Little is known of the
) for , trap days. In Catchment we identified five tigers ecology of most of these species, and less of their guilds.
from detections, five clouded leopards from detections Amongst the least known is the clouded leopard Neofelis
( photographs) and marbled cats from detections. nebulosa, a potent ambassador species for conservation
Using Bayesian-based spatial capture–recapture we esti- (Macdonald et al., unpubl. data) that occurs from the
mated the densities of tigers and clouded leopards to be Himalayan foothills and eastwards to Indochina, between
. ± SD . and . ± SD . individuals per km, which Myanmar serves as a biogeographical land bridge.
respectively. In Catchment we identified two tigers from The species occupies areas undergoing some of the most
three detections, nine clouded leopards from detections rapid deforestation (Hansen et al., ), and is threatened
and marbled cats from detections. Densities of clouded leo- by poaching and wildlife trafficking (D’Cruze &
pards and marbled cats were . ± SD . and . ± SD . Macdonald, ; Nijman & Shepherd, ; Min et al., in
individuals per km, respectively. These differences sug- press). Clouded leopards are the apex predators in many
gest that human activities, in particular gold mining, are af- South-east Asian rainforests, although where they co-occur
fecting felid populations, and these are a paramount concern with larger predators such as tigers their density and habitat
in Htamanthi. We demonstrate the importance of use may vary (Sunquist & Sunquist, ; Sunarto et al.,
Htamanthi within the Northern Forest Complex and high- ). Although there have been discoveries regarding the
light the Yawbawmee corridor as a candidate for protection. felid guilds and habitat use of the Sunda clouded leopard
Neofelis diardi (Haidir et al., ; Sollmann et al., ;
Hearn et al., ; Macdonald et al., unpubl. data), and the
threat to the species from habitat loss (Cushman et al., ),
little is known for the mainland clouded leopard and the fe-
HLA NAING* and SAW HTUN Wildlife Conservation Society Myanmar, Yangon,
lids with which it is sympatric.
Myanmar In Myanmar the density of clouded leopards has not
JOANNA ROSS, DAWN BURNHAM and DAVID W. MACDONALD (Corresponding been estimated; however, of the areas of the country sur-
author) Wildlife Conservation Research Unit, Department of Zoology, veyed (by camera trapping) for tigers during –,
University of Oxford, The Recanati-Kaplan Centre, Tubney House, Tubney,
OX13 5QL, UK. E-mail david.macdonald@zoo.ox.ac.uk
clouded leopards were found in areas and tigers in
three (Myanmar Forest Department, ). In clouded
*Also at: Wildlife Conservation Research Unit, Department of Zoology,
University of Oxford, The Recanati-Kaplan Centre, Tubney, UK leopards were recorded in three mountainous regions of
Received May . Revision requested September . northern Myanmar where they had not been recorded pre-
Accepted August . First published online November . viously (Zaw et al., ), and at a fourth site in the south
Oryx, 2019, 53(4), 654–662 © 2017 Fauna & Flora International doi:10.1017/S0030605317001260
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https://doi.org/10.1017/S0030605317001260Wild felids in Myanmar 655
hills to the western and south-western part of the
Sanctuary, and most from the eastern side can be navigated
by boat throughout the year, except for their upper reaches
in the driest parts of summer. There are three main footpaths
connecting villages on the west and east sides of the
Sanctuary. There are four management zones, Nam Phi
Lin, Nam E Zu, Nam Pa Gon and Nam Yan Yin; this study
was conducted in an area of km in Nam Pa Gon and of
km in Nam E Zu.
Methods
Camera trapping
We used two models of passive infrared digital camera traps
(Cuddeback, Non Typical Inc., De Pere, USA, and
ScoutGuard, HCO Outdoor Products, Norcross, USA).
During December – March we deployed cam-
era stations in Catchment , Nam Pa Gon, with .–. km
FIG. 1 Locations of camera traps in Catchments and of between camera stations. During December –
Htamanthi Wildlife Sanctuary, in north-western Myanmar. March we deployed camera stations in Catchment
, Nam E Zu, with similar spacing. At each station we used
unbaited, paired camera traps, set at c. cm height from
(WCS, , unpubl. data). Clouded leopards have also been ground level and c. . m either side of the trail.
confirmed in several locations in the east of the country
(Moo et al., ). Among the northern sites, Htamanthi
Analysis
Wildlife Sanctuary is particularly important because of its
proximity to the Northern Forest Complex. This vast con- We estimated population densities of tigers, clouded
tiguous forest landscape (, km), probably the largest leopards and marbled cats, using a spatially explicit
in Asia, lies in a transition zone of three biodiversity hot- capture–recapture model implemented within a Bayesian
spots, all of which contain clouded leopards and tigers: framework using the package SPACECAP v. ..
Himalaya, Indo-Burma and the Mountains of Southwest (Gopalaswamy et al., ) in R v .. (R Development
China (Myers et al., ; Conservation International, ). Core Team, ). Individual identification was carried
As part of a range-wide camera-trapping study of the out by at least two people, and detection histories for each
clouded leopard and members of its guild we deployed cam- individual were then constructed. We considered each
era traps in two catchments within the Htamanthi Wildlife -hour period to be a sampling occasion, as short sampling
Sanctuary over consecutive years. Htamanthi Wildlife intervals can improve the precision of estimates in spatially
Sanctuary is the southernmost Key Biodiversity Area of explicit capture–recapture analyses (Goldberg et al., ).
the Northern Forest Complex (also Tiger Conservation To assume we were surveying a demographically closed
Landscape , GTRP, ). population we used a -day survey period for tigers and
clouded leopards. Marbled cats are smaller and therefore
Study area may have a higher population turnover, so we subsampled
a -day period that maximized detections for marbled cats.
Htamanthi Wildlife Sanctuary (, km) is one of the For clouded leopards we explored the consequences of
largest protected areas in the region, located between the adopting a -day period, again subsampled to maximize
Chindwin and Uru Rivers in north-western Myanmar detections. These are similar or shorter durations compared
(Fig. ). The area is characterized by tropical evergreen with other estimates of population density, justifying the as-
forest, with some mixed deciduous forest in the western sumed sampling of a closed population (e.g. Brodie &
portion and dry mixed deciduous forest types along the east- Giordano, ; Wilting et al., ; Mohamed et al., ;
ern boundary (Arino et al., ). The area was gazetted for its Hearn et al., ). We constructed the state space by adding
megafauna, including the Asian elephant Elephas maximus, a buffer to the coordinates of the outermost camera stations,
tiger, gaur Bos gaurus, Asiatic black bear Ursus thibetanus, using ArcMap . (ESRI, Redlands, USA). We then added
sun bear Helarctos malayanus and clouded leopard. Seven potential activity centres by generating regularly spaced
streams flow in parallel from the eastern and north-eastern points with a resolution of . km (tigers and clouded
Oryx, 2019, 53(4), 654–662 © 2017 Fauna & Flora International doi:10.1017/S0030605317001260
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https://doi.org/10.1017/S0030605317001260656 H. Naing et al.
leopards) or . km (marbled cats) within this buffer. Each
independent detections of clouded leopards Neofelis nebulosa, tigers Panthera tigris and marbled cats Pardofelis marmorata, numbers of individuals detected and number of detections per
TABLE 1 Survey effort and number of sampling occasions at the two study sites (Catchments and ) in Htamanthi Wildlife Sanctuary, in north-western Myanmar (Fig. ), with numbers of
6(3), 5(2), 4(3), 1(2), 1(1), 1(1), 1(1), 1(1), 1(1), 1(1), 1(1), 1(1)
activity centre was designated either habitat or non-habitat,
based on local knowledge of the area. As land outside the
boundaries of the park is largely agricultural and human
2(2), 2(1), 2(1), 1(1), 1(1), 1(1), 1(1), 1(1), 1(1), 1(1)
presence is high, we designated all activity centre points
11(8), 10(9), 4(4), 4(2), 3(2), 3(2), 2(2), 2(1), 1(1)
10(7), 5(5), 4(4), 3(2), 2(2), 2(2), 2(2), 2(1), 1(1)
No. of detections per individual (no. of camera
falling outside the park as non-habitat. We increased the
stations at which each individual was detected)
size of the state space during preliminary analyses until
the probability of detection at the edge of the state space
became negligible. We determined that a buffer of km
was sufficient for clouded leopards and tigers, and a buffer
of km was appropriate for marbled cats. For all analyses
19(13), 1(1), 13(15), 1(1), 1(1)
16(12), 2(2), 1(1), 1(1), 1(1)
6(7), 6(6), 1(5), 1(4), 1(2)
we ran SPACECAP with trap response absent, half normal
detection function, , iterations and a thinning rate
of . In SPACECAP parameter convergence is assessed
using Geweke z scores; values between –. and . are
considered to be acceptable. We increased the burn-in
(the number of initial iterations discarded during the
analysis) during preliminary runs until the Geweke z scores
fell within this range. The upper limit to the population
size within the state space is set by the data augmentation
(sampling period)
value. We increased this value until ψ (the ratio of the
of individuals
estimated abundance within the state space to the maximum
defined by the data augmentation value) was # .. For
Total no.
11 (10)
12 (12)
Catchment we ran the clouded leopard data with a burn-in
5 (5)
5 (5)
5 (5)
9 (9)
9 (9)
of , for the -day peirod and , for the day
2
period. The tiger data were run with a burn-in of ,.
detections (sampling period)
The data augmentation value was set to for both
Total no. of independent
species. For Catchment we ran the clouded leopard
data with a burn-in of , and data augmentation of
for both the - and -day periods. The marbled cat
data were run with a burn-in of , and data augmenta-
tion of . We compared clouded leopard densities
49 (41)
49 (24)
27 (26)
24 (13)
54 (43)
54 (32)
37 (25)
between catchments, and following Sollmann et al. ()
we considered a difference to be significant if the % high-
3
est posterior density of one did not include the mean of the
No. of sampling
other.
As Asiatic golden cats do not have patterned coats they
occasions
are not reliably identifiable to individual, and therefore
population densities cannot be estimated using these meth-
90
60
90
60
90
60
60
ods. For golden cats, leopard cats and potential felid prey
detected by the cameras we calculated the number of inde-
trap-days (closed period)
pendent detections and naïve occupancy, which was calcu-
lated as the proportion of camera stations at which the
Total survey effort,
species was detected.
7,354 (7,104)
7,354 (4,534)
7,354 (7,104)
7,354 (7,104)
7,192 (6,861)
7,192 (4,534)
7,192 (4,727)
7,192
Results
We detected all three focal species in both catchments, but
Clouded leopard
Clouded leopard
Clouded leopard
Clouded leopard
detection frequencies and naïve occupancies varied greatly
Catchment 1
Catchment 2
(Tables & ). In Catchment we were able to identify
Marbled cat
Marbled cat
individual.
% of the tiger photographs, .% of clouded leopard
Species
photographs and .% of marbled cat photographs to
Tiger
Tiger
individual. Given the limited number of re-detections of
Oryx, 2019, 53(4), 654–662 © 2017 Fauna & Flora International doi:10.1017/S0030605317001260
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https://doi.org/10.1017/S0030605317001260Wild felids in Myanmar 657
TABLE 2 The total number of independent detections, detection rate, and naïve occupancies for felid species and potential prey species of
tigers and clouded leopards at the two study sites (Catchments and ) in Htamanthi Wildlife Sanctuary, in north-western Myanmar
(Fig. ).
Total no. of independent detections
(detection rate*) Naïve occupancy
Catchment 1 Catchment 2
Species (7,354 trap days) (7,192 trap days) Catchment 1 Catchment 2
Tiger 27 (0.37) 3 (0.04) 0.23 0.04
Clouded leopard 49 (0.67) 54 (0.75) 0.44 0.39
Asiatic golden cat Catopuma temminckii 104 (1.41) 62 (0.86) 0.41 0.28
Marbled cat 24 (0.33) 37 (0.51) 0.16 0.23
Leopard cat Prionailurus bengalensis 75 (1.02) 84 (1.17) 0.26 0.3
Leopard Panthera pardus 0 (0) 1 (0.01) 0 0.01
Gaur Bos gaurus 67 (0.91) 12 (0.17) 0.34 0.15
Wild boar Sus scrofa 141 (1.92) 122 (1.70) 0.61 0.56
Northern red muntjac Muntiacus vaginalis 563 (7.66) 491 (6.83) 0.94 0.85
Sambar deer Rusa unicolor 0 (0) 1 (0.01) 0 0.01
Chinese serow Capricornis milneedwardsii 2 (0.03) 0 (0) 0.02 0
Malayan porcupine Hystrix brachyura 121 (1.65) 138 (1.92) 0.30 0.45
Stump-tailed macaque Macaca arctoides 35 (0.48) 43 (0.60) 0.34 0.33
Northern pig-tailed macaque Macaca leonina 11 (0.15) 12 (0.17) 0.11 0.14
*The number of independent detections per trap days
marbled cats it was not possible to model these data, and the Numbers and population densities of tigers
values are included for comparison only.
The spatially explicit capture–recapture posterior sum- Given the substantial difference in tiger numbers detected
maries of the model parameter values are in Table . The between the surveys of Catchments and , although only
Bayesian p-values indicated that the models were of ad- a year apart and separated by , km, it is not useful to cal-
equate fit, and the Geweke z scores indicated that all culate a mean from the aggregated data. However, had we
model parameters converged. The estimated density for extrapolated from the estimated population density in
clouded leopards in Catchment derived from the -day Catchment this would have yielded an estimate of tigers
period is . ± SD . individuals per km and from (range – tigers) in the Htamanthi Wildlife Sanctuary, simi-
the -day period . ± SD . individuals per km. lar to the estimate of by Rabinowitz et al. () and within
For tigers the density estimate is . ± SD . individuals the wide range of densities (.–. tigers per km)
per km. estimated in the Hukaung Valley (Lynam et al., ).
In Catchment we were able to identify % of the tiger Such an extrapolation might have seemed warranted in
photographs, % of clouded leopard photographs and % the light of observations of field signs and prey, and the rela-
of marbled cat photographs to individual. The clouded tive abundance of the tiger’s preferred prey, the Eurasian
leopard population density was significantly higher than in wild pig Sus scrofa, the gaur and the barking deer
Catchment , with estimates of . ± SD . and . ± SD Muntiacus vaginalis (Hayward et al., ; Ngoprasert
. individuals per km from the - and -day peri- et al., ). During January –January the
ods, respectively. The population density of marbled cats Wildlife Conservation Society’s biological monitoring
was . ± SD . individuals per km. There were too team (monitoring the eastern hoolock gibbon Hoolock leu-
few tiger detections to estimate population density (Table ). conedys), SMART (Spatial Monitoring and Reporting Tool)
patrol teams and a community-based natural resource
management team confirmed that tigers were present
Discussion from the southern buffer zone to the northern boundary
of the Sanctuary (– km). However, the results from
We present the first population density estimates for the Catchment suggest that the tiger population density may
clouded leopard and marbled cat in Myanmar. We found be significantly lower than in Catchment , and we consider
that - and -day periods for clouded leopards yielded possible explanations for this below. Our estimates of tiger
almost identical results, and we are confident that surveying population densities are presented in the context of meth-
for days is appropriate to assume the population is odologically comparable estimates elsewhere in South-east
closed. Asia in Table .
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https://doi.org/10.1017/S0030605317001260658 H. Naing et al.
Abundance and population densities of clouded
*σ, movement parameter; λo, baseline trap encounter rate; Ψ, the ratio of the estimated abundance within the state space to the maximum allowable number set by the data augmentation value; N, number of
95% highest posterior
leopards
3,051.80–8,082.48
706.41–1,377.14
density interval With no previous data on clouded leopard numbers in the
0.001–0.02
0.003–0.02
0.04–0.41
0.32–1.55
0.20–0.70
4.0–13.75
Sanctuary we cannot assess whether their abundance has
44–151
changed, but we present our data for comparison with
5–24
methodologically comparable studies for other populations
of N. nebulosa and N. diardi in Table .
5,247.34 ± 1,438.47
1,014.65 ± 185.28
0.011 ± 0.005
96.11 ± 28.47
Population density of marbled cats, and naïve occupancy
0.009 ± 0.01
0.20 ± 0.10
12.58 ± 6.18
0.81 ± 0.40
0.43 ± 0.13
8.80 ± 2.60
of other felids and potential prey
Mean ± SD
0.57
0.74
Marbled cats are little studied and there are only two other
Tigers
estimates for comparison. Hearn et al. () found dens-
ities of .–. individuals per km, depending on
study area, and Singh & Macdonald () report densities
of . individuals per km. Our results are in line with
95% highest posterior
these estimates. Our naïve occupancy estimates for felids are
3,823.74–7,676.21
1,949.91–4,416.69
comparable to those reported in previous studies; for ex-
density interval
0.0017–0.006
ample, Haidir et al. () found that the naïve occupancy
0.005–0.042
0.04–0.30
0.32–1.09
0.12–0.63
1.04–5.34
of golden cats in Sumatra was ., and for Sunda clouded
TABLE 3 Posterior summaries of the model parameters from SPACECAP for each analysis from Catchments and .
19–97
leopards was .. The occupancy of ungulates has been
5–17
little studied in South-east Asia, but Gray & Prum ()
Clouded leopards, 60 days
reported a naïve occupancy of . for gaur in Cambodia.
This is similar to our result of . in Catchment . Gray
5,632.55 ± 1,020.07
() also reported that tiger abundance was also very
3,045.93 ± 737.76
0.0037 ± 0.001
57.30 ± 20.55
low in the same area of Cambodia, which suggests that
0.020 ± 0.01
0.16 ± 0.07
9.67 ± 3.72
0.62 ± 0.24
0.36 ± 0.13
3.2 ± 1.13
low numbers of gaur may be limiting tiger populations in
Mean ± SD
0.42
0.69
these areas.
Guild compositions and explanations for differences
between Catchments 1 and 2
95% highest posterior
Felid species and other mammalian carnivores occur
3,975.97–8,061.17
1,848.01–3,645.31
density interval
throughout South-east Asia in various combinations.
0.002–0.006
These guilds offer the opportunity to study the ecological
0.004–0.02
individuals in the state space; D, density (individuals per km)
0.04–0.30
0.32–1.09
0.14–0.60
1.27–5.06
processes of guild dynamics and, specifically, potential com-
23–92
petition between the member species. A plausible hypothesis
5–17
for wild felids, and one for which there is strong evidence
Clouded leopards, 90 days
amongst the Canidae (Macdonald & Sillero-Zubiri, ),
is that competition from larger species of felids will be inimi-
5,906.68 ± 1,087.64
cal to somewhat smaller species within a guild, and that this
2,689.11 ± 499.88
0.01 ± 0.005
0.004 ± 0.001
55.39 ± 18.71
in turn could result in mesopredator release for yet smaller
0.15 ± 0.07
9.35 ± 3.70
0.60 ± 0.24
0.35 ± 0.12
3.05 ± 1.03
species. This does not appear to be the case on Sumatra,
Mean ± SD
0.47
0.86
where the Sunda clouded leopard co-occurs with tigers.
Density estimates from Sumatra (Sollmann et al., )
and Borneo (Brodie & Giordano, ; Wilting et al., ;
Hearn et al., ) are broadly similar, suggesting that
tigers are not a limiting factor for clouded leopards on
Catchment 1
Catchment 2
Sumatra.
Parameter*
Bayesian p
Bayesian p
Our comparison of Catchments and provides the op-
portunity to evaluate the number of clouded leopards
λo
λo
where tigers were relatively abundant and where they
Ψ
Ψ
N
N
D
D
σ
σ
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https://doi.org/10.1017/S0030605317001260Wild felids in Myanmar 659
TABLE 4 Published estimates of tiger population density based on the use of spatially explicit techniques.
Density 95% CI/highest Spatially explicit
(individuals posterior density capture–recapture
Site per 100 km2) interval method Source
Ussuriiskii Nature Reserve, Russia 0.112–0.586 * Bayesian Hernandez-Blanco
et al. (2013)
Rajaji National Park, India 3.31 ± SD 1.51 1.56–7.03 Bayesian Harihar et al. (2011)
2.67 ± SD 0.97 1.24–4.96 Bayesian
5.17 ± SD 1.94 2.07–9.50 Bayesian
5.81 ± SD 2.26 1.86–9.92 Bayesian
Mudumalai Tiger Reserve, India 8.31 ± SE 2.56 * Bayesian Kalle et al. (2011)
Ranthambhore Tiger Reserve, India 6.1 ± SE 1.2 * Maximum likelihood Singh et al. (2014b)
Kalakad-Mundanthurai Tiger Reserve, India 2.9 ± SE 1.40 * Bayesian Ramesh et al. (2012)
2.2 ± SE 1.6 * Maximum likelihood
Pakke Tiger Reserve, India 1.86 ± SE 0.7 * Maximum likelihood Selvan et al. (2014)
Hunchun National Nature Reserve 0.33 ± SD 0.10 0.18–0.56 Bayesian Xiao et al. (2016)
0.40 ± SD 0.12 0.20–0.68 Bayesian
0.30 ± SE 0.10 0.16–0.55 Maximum likelihood
0.24 ± SE 0.09 0.12–0.48 Maximum likelihood
Sundarbans, India 4.08 ± SE 1.51 0.18–0.56 Maximum likelihood Roy et al. (2016)
5.81 ± SE 1.24 Maximum likelihood
Suklaphanta Wildlife Reserve, Nepal 2.10 ± SE 0.8 * Maximum likelihood Karki (2011)
Chitwan National Park, Nepal 2.21 ± SD 0.42 1.41–3.01 Bayesian Thapa & Kelly (2017)
2.08 ± SD 1.13 0.31–4.39 Bayesian Thapa & Kelly (2017)
Pakke Tiger Reserve, India 0.97 ± SE 0.23 * Maximum likelihood Singh et al. (2014a)
*Data are not available.
TABLE 5 Published estimates of clouded leopard population density based on the use of spatially explicit techniques.
Density 95% CI/highest Spatially explicit
(individuals per posterior density capture–recapture
Site 100 km2) interval method Source
Sunda clouded leopard Neofelis diardi
Tangkulap-Pinangah Forest Reserve, Sabah, 0.84 ± SD 0.42 0.25–1.83 Bayesian Wilting et al. (2012)
Malaysia
Segaliud Lokan Forest Reserve, Sabah, 1.04 ± SD 0.58 0.29–2.55 Bayesian Wilting et al. (2012)
Malaysia
Maliau Basin Conservation Area, Sabah, 1.90 ± SE* 0.70–5.40 Maximum Brodie & Giordano (2012)
Malaysia likelihood
Renah Kayu Embun, Sumatra, Indonesia 1.57 ± SD 0.69 0.58–3.27 Bayesian Sollmann et al. (2014)
Sipurak, Sumatra, Indonesia 0.77 ± SD 0.52 0.15–2.10 Bayesian Sollmann et al. (2014)
Bungo, Sumatra, Indonesia 1.62 ± SD 0.73 0.58–3.37 Bayesian Sollmann et al. (2014)
Ipuh, Sumatra, Indonesia 1.11 ± SD 0.47 0.42–2.24 Bayesian Sollmann et al. (2014)
Danum Valley Conservation Area, Sabah, 1.73 ± SD 0.54 0.81–2.78 Bayesian Hearn et al. (2017)
Malaysia
Tawau Hills Park, Sabah, Malaysia 2.23 ± SD 0.52 1.35–3.27 Bayesian Hearn et al. (2017)
Crocker Range Park, Sabah, Malaysia 1.39 ± SD 0.41 0.77–2.21 Bayesian Hearn et al. (2017)
Ulu Segama Forest Reserve, Sabah, Malaysia 3.10 ± SD 1.11 1.26–5.32 Bayesian Hearn et al. (2017)
Tabin Wildlife Sanctuary, Sabah, Malaysia 2.66 ± SD 1.11 0.79–4.74 Bayesian Hearn et al. (2017)
Kinabatangan Wildlife Sanctuary, Sabah, 1.54 ± SD 0.70 0.41–2.90 Bayesian Hearn et al. (2017)
Malaysia
Mainland clouded leopard Neofelis
nebulosa
Manas National Park, India 4.73 ± SE 1.43 * Maximum likelihood Borah et al. (2014)
Temengor, Malaysia 3.46 ± SE 1.00 1.98–6.04 Maximum likelihood Mohamad et al. (2015)
Belum, Malaysia 1.83 ± SE 0.61 0.97–3.48 Maximum likelihood Mohamad et al. (2015)
Dampa Wildlife Reserve, India 5.14 ± SD 1.80 2.05–8.72 Bayesian Singh & Macdonald (2017)
*Data are not available.
Oryx, 2019, 53(4), 654–662 © 2017 Fauna & Flora International doi:10.1017/S0030605317001260
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https://doi.org/10.1017/S0030605317001260660 H. Naing et al.
FIG. 2 Location of Htamanthi
Wildlife Sanctuary in relation
to Hukaung Valley Wildlife
Sanctuary and other protected
areas in the north of
Myanmar.
were less so. The difference in estimated population density Human presence could influence the mammalian
of clouded leopards (. individuals per km in community structure in several ways. People may engen-
Catchment compared to . individuals per km in der fear amongst both felids and their prey (e.g.
Catchment ) is in line with the prediction of the intra-guild Oriol-Cotterill et al., a,b), and may affect tigers dir-
hostility hypothesis. ectly by killing them, and indirectly by killing their prey
These comparisons raise the question of why there were (the absence of gaur and serow may be a case in point
markedly fewer tigers in Catchment than in Catchment , and, being big prey, may affect the biggest felids); another
and what other factors might underlie the different guild dy- possibility is that the domestic stock trafficked through
namics observed between these surveys. Three obvious, and the protected area could transmit disease to the wild
not mutually exclusive, hypotheses are that () there was a ungulates.
difference in habitat or prey availability, or some other en- The anthropogenic hypothesis is sufficiently compelling,
vironmental variable, between the two sites, () conditions and has such serious implications for conservation, that it
changed between the two survey periods, or () some other merits further investigation as a priority. Rabinowitz et al.
factor, such as an anthropogenic impact, caused the differ- () cautioned that the populations of tigers and gaur
ences in felid guild structure, either directly or indirectly. were at risk of elimination if threats prevailing at the time
Given that the two catchments are separated by , km were not controlled, and we suspect this warning is now
and seem generally similar, the habitat hypothesis is un- even more pressing.
promising. However, two large prey species, gaur and
serow Capricornis milneedwardsii, were detected less fre-
quently in Catchment (Table ). The temporal change The Yawbawmee Corridor
hypothesis is also unpromising, given that the two surveys
were separated by barely months. However, the third The National Tiger Survey (–) confirmed the pres-
hypothesis, that Catchment was subject to damaging ence of tigers in Hukaung Valley, Upper Chindwin,
human activity, is strongly supported. SMART patrols and Htamanthi in northern Myanmar, and Tenasserim Hills
incidental observations in indicated that although there in the south. Our findings emphasize the importance of
were human incursions in both areas, incidents of gold Htamanthi within the northern Myanmar landscape. To
mining were times higher in Catchment than in the immediate north of Htamanthi lies the Yawbawmee
Catchment , and . snares were removed from Corridor, , km of currently unprotected forest, which
Catchment compared with in Catchment for a similar could link Htamanthi with Hukaung Valley Wildlife
patrol effort. Sanctuary (Fig. ). The gazettement of this corridor would
Oryx, 2019, 53(4), 654–662 © 2017 Fauna & Flora International doi:10.1017/S0030605317001260
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https://doi.org/10.1017/S0030605317001260Wild felids in Myanmar 661
substantially increase the connectivity of the protected areas C U S H M A N , S.A., M AC D O N A L D , E.A., L A N D G U T H , E.L., M A L H I , Y. &
of the Northern Forest Complex and would deliver a sub- M AC D O N A L D , D.W. () Multiple-scale prediction of forest loss
risk across Borneo. Landscape Ecology, , –.
stantial contribution to landscape-level conservation of fe-
D’C R U Z E , N. & M A C D O N A L D , D.W. () Clouded in mystery: the
lids and other threatened species in the region. global trade in clouded leopards. Biodiversity and Conservation, ,
–.
G O L D B E R G , J.F., T E M P A , T., N O R B U , N., H E B B L E W H I T E , M., M I L L S , L.
Acknowledgements S., W A N G C H U K , T.R. & L U K AC S , P. () Examining temporal
sample scale and model choice with spatial capture–recapture
The survey was funded jointly by a grant to DWM from models in the common leopard Panthera pardus. PLoS ONE, (),
Partnership for Change and by Wildlife Conservation e.
Society (WCS) Myanmar, with support and coordination G O P A L A S WA M Y , A.M., R OY L E , J.A., H I N E S , J.E., S I N G H , P., J AT H A N N A ,
from Htamanthi Wildlife Sanctuary, Nature Wildlife D., K U M A R , N.S. & K A R A N T H , K.U. () Program SPACECAP:
software for estimating animal density using spatially explicit
Conservation Division, Myanmar Forest Department. We capture–recapture models. Methods in Ecology and Evolution, ,
thank Dr Nyi Nyi Kyaw, Director General of Myanmar –.
Forest Department, and U Win Naing Thaw, Director of G R A Y , T.N.E. & P R U M , S. () Leopard density in post-conflict
the Nature Wildlife Conservation Division under the landscape, Cambodia: evidence from spatially explicit capture–
Ministry of Natural Resources and Environmental recapture. The Journal of Wildlife Management, , –.
GTRP (G LO B A L T I G E R R E C O V E R Y P R O G R A M ) () Global Tiger
Conservation, for granting permission and supporting the
Recovery Program –. Http://documents.worldbank.org/
project; Htamanthi Park Warden U Khin Maung Hla and curated/en//pdf/WPBox
park rangers for logistical support to our field team; the sur- FinalVersionEng.pdf [accessed September ].
vey team members for their tireless effort and dedication; H A I D I R , I.A., D I N ATA , Y., L I N K I E , M. & M AC D O N A L D , D.W. ()
WCS Myanmar Country Director U Than Myint, Northern Asiatic golden cat and Sunda clouded leopard occupancy in
the Kerinci Seblat landscape, West-Central Sumatra. Cat News, ,
Forest Complex Coordinator U Than Zaw and staff for
–.
support throughout the project; and Cedric Tan and two an- H A N S E N , M.C., P O TA P O V , P.V., M O O R E , R., H A N C H E R , M.,
onymous reviewers for their valuable inputs. DWM gratefully T U R U B A N O VA , S.A., T Y U K AV I N A , A. et al. () High-resolution
acknowledges grants from the DASSK Trust and the global maps of st-century forest cover change. Science, ,
Irrawaddy Policy Exchange for the training of Myanmar con- –.
servationists and support of our clouded leopard programme. H A R I H A R , A., P A N D AV , B. & G OY A L , S.P. () Responses of leopard
Panthera pardus to the recovery of a tiger Panthera tigris population.
We thank the Recanati-Kaplan Foundation, Panthera and Journal of Applied Ecology, , –.
Robertson Foundation for their wider support of WildCRU. H A Y WA R D , M.W., J Ę D R Z E J E W S K I , W. & J Ê D R Z E J E W S K A , B. ()
Prey preferences of the tiger Panthera tigris. Journal of Zoology, ,
Author contributions –.
H E A R N , A.J., R O S S , J., B E R N A R D , H., B A K A R , S.A., G O O S S E N S , B.,
DWM, DB and SH conceived the study. DWM, SH and HN H U N T E R , L.T.B. & M AC D O N A L D , D.W. () Responses of Sunda
clouded leopard density to anthropogenic disturbance and refining
designed the study and HN collected the data. JR and HN
estimates of their conservation status in Sabah. Oryx, https://doi.
conducted data analysis. DWM, DB, JR, SH and HN wrote org/./S.
the article. H E A R N , A.J., R O S S , J., B E R N A R D , H., B A K A R , S.A., H U N T E R , L.T.B. &
M AC D O N A L D , D.W. () The first estimates of marbled cat
Pardofelis marmorata population density from Bornean primary
References and selectively logged forest. PLoS ONE, (), e.
H E R N A N D E Z -B L A N C O , J.A., R O Z H N O V , V.V., L U K A R E V S K I Y , V.S.,
A R I N O , O., R A M O S P E R E Z , J.J., K A LO G I R O U , V., B O N T E M P S , S., N A I D E N KO , S.V., C H I S T O P O LOVA , M.D., S O R O K I N , P.A. et al. ()
D E FO U R N Y , P. & V A N B O G A E R T , E. () Global Land Cover Map Spatially explicit capture–recapture method (SECR, SPACECAP): a
for (GlobCover ). European Space Agency & Université new approach to determination of the Amur tiger (Panthera tigris
catholique de Louvain. Http://dx.doi.org/./PANGAEA. altaica) population density by means of camera-traps. Doklady
[accessed September ]. Biological Sciences, , –.
B O R A H , J., S H A R M A , T., D A S , D., R A B H A , N., K A K AT I , N., K A L L E , R., R A M E S H , T., Q U R E S H I , Q. & S A N K A R , K. () Density of
B A S U M AT A R Y , A. et al. () Abundance and density estimates tiger and leopard in a tropical deciduous forest of Mudumalai Tiger
for common leopard Panthera pardus and clouded leopard Reserve, southern India, as estimated using photographic capture–
Neofelis nebulosa in Manas National Park, Assam, India. Oryx, , recapture sampling. Acta Theriologica, , –.
–. K A R K I , J.B. () Occupancy and abundance of tigers and their prey in
B R O D I E , J. & G I O R D A N O , A.J. () Density of the Vulnerable Sunda the Terai Arc Landscape, Nepal. PhD thesis. Forest Research
clouded leopard Neofelis diardi in a protected area in Sabah, Institute University, Dehradun, India.
Malaysian Borneo. Oryx, , –. L Y N A M , A.J., R A B I N OW I T Z , A., M Y I N T , T., M A U N G , M., L A T T , K.T. &
C O N S E R VAT I O N I N T E R N AT I O N A L () Biodiversity Hotspots P O , S.H.T. () Estimating abundance with sparse data: tigers in
Revisited. Conservation International, Washington, DC, USA. northern Myanmar. Population Ecology, , –.
Http://www.biodiversityhotspots.org/xp/Hotspots/resources/maps. M AC D O N A L D , D.W. & S I L L E R O -Z U B I R I , C. () Dramatis personae.
xml [accessed October ]. In The Biology and Conservation of Wild Canids (eds
Oryx, 2019, 53(4), 654–662 © 2017 Fauna & Flora International doi:10.1017/S0030605317001260
Downloaded from https://www.cambridge.org/core. IP address: 46.4.80.155, on 21 Feb 2021 at 15:18:55, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms.
https://doi.org/10.1017/S0030605317001260662 H. Naing et al.
D.W. Macdonald & C. Sillero-Zubiri), pp. –. Oxford University conventional population estimation methods in a unique ecosystem
Press, Oxford, UK. Population Ecology, , −.
M AC D O N A L D , D.W., L O V E R I D G E , A.J. & N OW E L L , K. () Dramatis S E LVA N , K.M., L Y N G D O H , S., H A B I B , B. & G O P I , G.V. ()
personae: an introduction to the wild felids. In The Biology and Population density and abundance of sympatric large carnivores
Conservation of Wild Felids (eds D.W. Macdonald & A.J. Loveridge), in the lowland tropical evergreen forest of Indian Eastern Himalayas.
pp. –. Oxford University Press, Oxford, UK. Mammalian Biology – Zeitschrift für Säugetierkunde, , –.
M I N , S., D’C R U Z E , N. & M AC D O N A L D , D.W. (in press) A note on felid S I N G H , P. & M AC D O N A L D , D.W.M. () Populations and activity
trade at local markets in Myanmar. Cat News, . patterns of clouded leopards and marbled cats in Dampa Tiger
M O H A M A D , S.W., R A Y A N , D.M., C H R I S T O P H E R , W.C.T., H A M I R U L , Reserve, India. Journal of Mammalogy, https://doi.org/./
M., M O H A M E D , A., L A U , C.F. & S I WA N , E.S. () The first jmammal/gyx.
description of population density and habitat use of the mainland S I N G H , R., C H A U H A N , D.S., M I S H R A , S., K R A U S M A N , P.R. & G O Y A L ,
clouded leopard Neofelis nebulosa within a logged-primary forest in S.P. (a) Tiger density in a tropical lowland forest in the Eastern
South East Asia. Population Ecology, , –. Himalayan Mountains. SpringerPlus, , .
M O H A M E D , A., S O L L M A N N , R., B E R N A R D , H., A M B U , L.N., L A G A N , P., S I N G H , R., Q U R E S H I , Q., S A N K A R , K., K R A U S M A N , P.R. & G O YA L , S.P.
M A N N A N , S. et al. () Density and habitat use of the leopard cat (b) Evaluating heterogeneity of sex-specific capture probability
(Prionailurus bengalensis) in three commercial forest reserves in and precision in camera-trap population estimates of tigers. Wildlife
Sabah, Malaysian Borneo. Journal of Mammalogy, , –. Society Bulletin, , –.
M O O , S.S.B., F R O E S E , G.Z.L. & G R AY , T.N.E. () First structured S O L L M A N N , R., L I N K I E , M., H A I D I R , I.A. & M AC D O N A L D , D.W. ()
camera-trap surveys in Karen State, Myanmar, reveal high diversity Bringing clarity to the clouded leopard Neofelis diardi: first density
of globally threatened mammals. Oryx, https://doi.org/./ estimates from Sumatra. Oryx, , –.
S. S U N A R T O , S., K E L LY , M.J., P A R A K K A S I , K. & H U TA J U L U , M.B. ()
M Y A N M A R F O R E S T D E P A R T M E N T () A National Tiger Action Cat coexistence in central Sumatra: ecological characteristics, spatial
Plan for the Union of Myanmar. Myanmar Forest Department, and temporal overlap, and implications for management. Journal of
Ministry of Forestry, Yangon, Myanmar. Zoology, , –.
M Y E R S , N., M I T T E R M E I E R , R.A., M I T T E R M E I E R , C.G., D A F O N S E C A , S U N Q U I S T , M. & S U N Q U I S T , F. () Wild Cats of the World.
G.A.B. & K E N T , J. () Biodiversity hotspots for conservation University of Chicago Press, Chicago, USA.
priorities. Nature, , –. T H A P A , K. & K E L LY , M.J. () Density and carrying capacity in the
N G O P R A S E R T , D., L Y N A M , A.J., S U K M A S U A N G , R., T A N T I P I S A N U H , N., forgotten tigerland: tigers in the understudied Nepalese Churia.
C H U T I P O N G , W., S T E I N M E T Z , R. et al. () Occurrence of three Integrative Zoology, , –.
felids across a network of protected areas in Thailand: prey, W I LT I N G , A., M O H A M E D , A., A M B U , L.N., L A G A N , P., M A N N A N , S.,
intraguild, and habitat associations. Biotropica, , –. H O F E R , H. & S O L L M A N N , R. () Density of the Vulnerable Sunda
N I J M A N , V. & S H E P H E R D , C.R. () Trade in tigers and other wild clouded leopard Neofelis diardi in two commercial forest reserves in
cats in Mong La and Tachilek, Myanmar—a tale of two border Sabah, Malaysian Borneo. Oryx, , –.
towns. Biological Conservation, , –. X I AO , W., F E N G , L., M O U , P., M I Q U E L L E , D.G., H E B B L E W H I T E , M.,
O R I O L -C O T T E R I L L , A., M AC D O N A L D , D.W., V A L E I X , M., E K WA N G A , G O L D B E R G , J.F. et al. () Estimating abundance and density of
S. & F R A N K , L.G. (a) Spatiotemporal patterns of lion space Amur tigers along the Sino–Russian border. Integrative Zoology, ,
use in a human-dominated landscape. Animal Behaviour, , –.
–. Z AW , T., M Y I N T , T., H T U N , S., P O , S.H.T., L AT T , K.T., M A U N G , M. &
O R I O L -C O T T E R I L L , A., V A L E I X , M., F R A N K , L.G., R I G I N O S , C. & L Y N A M , A.J. () Status and distribution of smaller cats in
M AC D O N A L D , D.W. (b) Landscapes of coexistence for Myanmar. Cat News, , –.
terrestrial carnivores: the ecological consequences of being
downgraded from ultimate to penultimate predator by humans.
Oikos, , –.
R D E V E LO P M E N T C O R E T E A M () R: A Language and Environment Biographical sketches
for Statistical Computing. R Foundation for Statistical Computing,
Vienna, Austria. H L A N A I N G is field coordinator of WildCRU’s clouded leopard
R A B I N O W I T Z , A., S C H A L L E R , G.B. & U G A , U. () A survey to assess programme in Myanmar. J O A N N A R O S S ’s research focuses on the
the status of Sumatran rhinoceros and other large mammal species conservation of members of the Bornean felid guild, and their spatial
in Tamanthi Wildlife Sanctuary, Myanmar. Oryx, , –. ecology. D A W N B U R N H A M is a member of WildCRU’s conservation
R A M E S H , T., S R I D H A R A N , N., S A N K A R , K., Q U R E S H I , Q., S E LVA N , K. geopolitics research group, with a particular focus on conservation
M., G O K U L A K K A N N A N , N. et al. () Status of large carnivores and ethics. S A W H T U N is WCS’s Deputy Country Director in Myanmar.
their prey in tropical rainforests of South-western Ghats, India. D A V I D W. M A C D O N A L D has a background in behavioural
Tropical Ecology, , –. ecology, and is running several long-term and wide-ranging con-
R O Y , M., Q U R E S H I , Q., N A H A , D., S A N K A R , K., G O P A L , R. & J H A L A , Y. servation programmes with an emphasis on carnivores and
V. () Demystifying the Sundarban tiger: novel application of interdisciplinarity.
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