Journal of Threatened Taxa |
www.threatenedtaxa.org | 14 December 2020 | 12(16): 17222–17228
ISSN 0974-7907 (Online) | ISSN 0974-7893
(Print)
doi: https://doi.org/10.11609/jott.6487.12.16.17222-17228
#6487 | Received 29 July 2020 | Final
received 17 October 2020 | Finally accepted 29 October 2020
Male residency of Sunda Clouded Leopard Neofelis
diardi (Cuvier, 1823) (Mammalia: Carnivora:
Felidae) in a peat swamp forest, Indonesian Borneo
Lynn Pallemaerts
1, Adul 2, Ici
P. Kulu 3, Karen Anne Jeffers 4,
David W. Macdonald 5 & Susan
Mary Cheyne 6
1 Department of Biology, Lund
University, Sölvegaten 37, 22362 Lund, Sweden.
2, 4, 6 Borneo Nature Foundation, Jalan Bukit Raya 82, Bukit Raya, Palangka
Raya 73112, Central Kalimantan, Indonesia.
3 Deceased, UPT LLG-CIMTROP, Jl. Yos Sudarso, Kampus
UNPAR Tunjung Nyahu, Palangka Raya, Central Kalimantan, Indonesia.
5 WildCRU,
Department of Zoology, University of Oxford, The Recanati-Kaplan
Centre, Tubney House, Tubney,
OX13 5QL UK.
6 Gibbs Building G2.23, Department
of Social Sciences, Oxford Brookes University, Headington, Oxford, OX3 0BP,
United Kingdom.
1 lynn.pallemaerts@gmail.com (corresponding
author), 2 adul@borneonature.org, 4 k.jeffers@exeter.ac.uk.,
5 david.macdonald@zoo.ox.ac.uk, 6
s.cheyne@borneonature.org
Editor: Angie Appel, Wild Cat Network,
Bad Marienberg, Germany. Date
of publication: 14 December 2020 (online & print)
Citation: Pallemaerts,
L., Adul, I.P. Kulu, K.A.
Jeffers, D.W. Macdonald & S.M. Cheyne (2020). Male residency of Sunda Clouded Leopard Neofelis
diardi (Cuvier, 1823) (Mammalia: Carnivora:
Felidae) in a peat swamp forest, Indonesian Borneo. Journal of Threatened Taxa 12(16): 17222–17228. https://doi.org/10.11609/jott.6487.12.16.17222-17228
Copyright: © Pallemaerts
et al 2020. Creative Commons Attribution
4.0 International License. JoTT allows unrestricted use, reproduction, and
distribution of this article in any medium by providing adequate credit to the
author(s) and the source of publication.
Funding:
The Robertson Foundation, Panthera, Borneo Nature
Foundation, The Clouded Leopard Project/Point Defiance Zoo and Aquarium and Fresno Chafee Zoo.
Competing interests: The authors
declare no competing interests.
Author contribution: Study design: LP, SMC. Fieldwork:
A, KAJ, SMC. Data analysis: LP. Writing: LP. Feedback: SMC, DWM.
Acknowledgements: The authors’ sincerest thanks go
to the Centre for International Cooperation in Sustainable Management of
Tropical Peatland (CIMTROP) and to the University of Palangka
Raya (UPR), as well as RISTEKDIKTI for supporting the research of KAJ (Surat Izin Penelitian
360/SIP/DIT.KI/IX/2015, 47/EXT/SIP/FRP/E5/Dit.KI/2016 and
55/EXT/SIP/FRP/E5/Dit.KI/IX/2017), who provided data for this paper. LP wishes
to thank the Borneo Nature Foundation (BNF), and Ola Olsson and Yann Clough at
the Department of Biology at Lund University. The Robertson Foundation provided
funding for the majority of the survey work presented here. Additional funding
for different stages of this work was kindly provided by Panthera,
Borneo Nature Foundation, The Clouded Leopard Project/Point Defiance Zoo and
Aquarium and Fresno Chafee Zoo. These data were collected by the Borneo Nature
Foundation in collaboration with CIMTROP. SMC oversaw the work with RISTEK
permit number 203/E5/E5.4/SIP/2019.
Abstract: The Sunda
Clouded Leopard Neofelis diardi is the apex predator on the island of Borneo,
yet little is known of its ecology. We
document the length of residency of male Sunda
Clouded Leopards in Central Kalimantan, Indonesian Borneo. Over 10 years, camera trap data have been
obtained in the Sebangau peat swamp forest in a study
area of ~ 105km2. We
identified 11 individuals (eight males, one female, and two with unknown sex),
from 152 notionally independent photographs.
On average, males remained in the study area for 39.3 months (SE 8.3),
or 3.3 years (SE 0.7), ranging from less than a month of residency up to 71
months. Females were infrequently
recorded, possibly as a result of human disturbance and/or high male densities. Our results reveal that even a 10-year
dataset is inadequate to answer some basic ecological questions, emphasising
the importance of long-term monitoring of this species.
Keywords: Camera trap, Kalimantan, male
sex-bias, social organisation.
introduction
The rainforests of Sumatra
and Borneo harbour one of the least known cat species
(Allen et al. 2016): the Sunda Clouded Leopard Neofelis diardi
(Wilting et al. 2006). This medium-sized
cat weighs 11–25 kg and is thought to be adept to climbing because of its broad
paws and long tail (Wilting et al. 2006).
On Borneo, it occupies the role of the apex predator, being the largest
of five cat species on the island (Cheyne & Macdonald 2011).
The Sunda
Clouded Leopard inhabits primary and selectively logged dipterocarp forest,
peat swamp forest, and mangroves (Hearn et al. 2016). Borneo’s forests have been heavily affected
by logging, conversion to plantations and fire since the early 1970s (Gaveau et al. 2014).
This habitat loss coupled with illegal hunting of wild prey is
considered a serious threat to the Sunda Clouded
Leopard (Wilting et al. 2006), resulting in a decreasing population (Hearn et
al. 2015). Since 2015, the Sunda Clouded Leopard is listed as Vulnerable on the IUCN
Red List (Hearn et al. 2015), and rapid and efficient conservation measures are
necessary.
Research
effort has already increased, but studying the Sunda
Clouded Leopard remains a challenge (Hearn et al. 2013). It is rare, elusive, and
predominantly nocturnal (Adul et al. 2015). The increase in camera trapping for
ecological studies, however, has somewhat alleviated this problem.
The
Sunda Clouded Leopard is solitary, and assumed to be
territorial (Allen et al. 2016), however, this assumption is based on little
empirical evidence, and results of recent research indicate that home ranges of
males overlap (Allen et al. 2016; Pallemaerts et al.
2019). A similar social organisation was
identified in some other felid species such as Leopard Cat Prionailurus
bengalensis (Grassman
et al. 2005), Leopard Panthera pardus (Odden & Wegge 2005), Jaguar P. onca
(Harmsen et al. 2009; Guilder et al. 2015), and Tiger
P. tigris (Goodrich et al. 2010). This spatial organisation involves a
combination of temporal avoidance and tolerance towards conspecifics (Grassman et al. 2005; Harmsen et
al. 2009; Núñez-Pérez 2011).
We
describe the residency of male Sunda Clouded Leopards
that were recorded on camera trap between 2008 and 2018 in a peat swamp forest
in Kalimantan, Indonesia.
Study area
We conducted this study in
a portion of the 550km2 Natural Laboratory for Peat Swamp Forest
(NLPSF), nested within the tropical peat swamp forest of Sebangau
in Central Kalimantan, Indonesia (Fig.
1). This area of approx. 5,600km2 is
the largest area of contiguous forest remaining in Kalimantan (Cheyne et al.
2013). The NLPSF is seasonally flooded
from October to June, and is one of the deepest peat swamp forests in the world
with a peat depth of 3–26 m (Page et al. 1999).
The forest has a history of logging concessions, illegal logging,
drainage and wildfire (Cheyne 2010; Cheyne & Macdonald 2011), but has been
protected as a national park since 2004 (Cheyne 2010). Due to past exploitation and disturbance, it
is a patchwork of pristine and recovering forest (Page et al. 1999).
Material and methods
In
May 2008, an extensive and continuous camera trap survey was started within the
NLPSF. It is still running and is,
therefore, the longest-running camera trap survey on Borneo. Over the years, we used six camera
trap models including Cuddeback Expert, Cuddeback IR, Cuddeback Ambush
IR, Maginon WK 3 HD, Crenova
RD100, and Bushnell NatureView HD.
For
this study, only data between May 2008 and February 2018 were used. At the start of the survey, we planned to
place camera traps in a systematic grid in the study area, and this was
achieved in the forest immediately surrounding the base camp (Fig. 1). Due to the difficult terrain, however, such a
grid was not practical in the areas further away from base camp, and so, over
the years, short expeditions were undertaken to set up several camera trap
stations in areas of interest within the NLPSF, resulting in a clumped trapping
grid (Fig. 1). As a result, we placed
camera traps at 94 different locations between 2008 and 2018 and sampled an
area of 104.8km2 (i.e. 19.1%) of the NLPSF, as determined by the
polygon formed by the outermost locations (Fig. 1). Most locations were sampled
for two or three months, but sometimes 1–2 years, within this 10-year period,
while other locations have been sampled almost continuously since 2008, with an
average sampling duration of approximately 21 months for a single
location. On average, the cameras were
located at a distance of 2.7 ± 0.6 km from each other. Because of the close proximity between
stations, the clumped array of cameras, and uneven distribution of sampling
across our study area, we checked the independent photographs for spatial
autocorrelation using Moran’s index I (Sokal &
Oden 1978) in ArcMap 10.3 with weights for pairs of neighbouring stations
computed using the inverse distance between the respective camera trap
stations. Camera trap photographs were
spatially independent at all stations, with Moran’s index over the entire
survey period (2008–2018) being non-significant (I = 0.043, p = 0.87).
Where
possible, we placed the camera traps in pairs at a distance of 7–10 m from each
other, henceforth termed station, to allow for simultaneous photographs of both
sides of Sunda Clouded Leopards. We installed the stations along transects,
animal trails, boardwalks, natural bridges, or watering holes to maximize
photographic rates. If insufficient cameras
were available, they were placed singly.
We deployed each camera at about 50cm above ground and protected them
with a simple plastic box from rainfall and potential destruction by
wildlife. In the early years of the
camera trap survey, the stations were checked every two weeks to change
batteries and retrieve memory cards; later, due to improvements in battery
life, they were checked every 40 days.
We
identified individual Sunda Clouded Leopards based on
their unique coat patterns and, when possible, sexed them based on secondary
sexual traits. For each individual, a
capture history was made based on the number of notionally independent photographs
each year (Fig. 2), where a photograph was considered notionally independent if
it was taken at a different station than the previous one, or if there was at
least a 10-minute interval between two photographs at the same station. We defined the residency of
individual Sunda Clouded Leopards as the number of
continuous months and years each individual was recorded during the entire
study period and estimated it using the dates of first and last detection
within the study area. We categorised an
individual as resident if it was recorded for at least six continuous months, a
length of time that was judged to reflect site attachment by Hemker et al. (1984) and Zimmermann et al. (2005). We categorised individuals present in the
area for less than this period as transient individuals.
Results
During the entire study
period, we obtained 152 notionally independent photographs of Sunda Clouded Leopards in more than 55,075 traps
nights. Eight of these photographs were
not clear enough to be attributed to any known individual, and were excluded
from further analyses. From the
remaining 144 independent photographs, we were able to identify 11 individuals
(Table 1). Eight were male, one was
female, and two could not be sexed. The
number of individual photographs for individuals ranged from one to 32, with a
mean at 13.1 (SE 3.7) records per individual.
Time spent living in the
study area ranged from less than a month up to 71 months. We identified eight individuals that were
present for at least six months in the study area, and three transient
individuals. The mean residency of the
eight resident individuals was 39.3 months (SE 8.3 months) or 3.3 years (SE 0.7
years) with a range of nine to 71 months.
All but one residents were males.
The number of individuals
present within the study area each year varied from one to five, and averaged
2.8 (SE 0.5). During one year,
individuals were recorded between once and 19 times across the study area (Fig.
2). Several camera traps were visited by
different males, with an average of 1.5 (SE 0.1) males visiting the same
location in a six-month period.
Discussion
On
average, resident males seem to remain in our study area for approximately
three years, before ceasing to be detected by camera traps. Short gaps in an individual’s detection
record may indicate that it evaded camera traps (M1 and M4), whereas longer gaps
(M3 and M5) could indicate genuine periods of absence (Fig. 2). The reason for these absences are
unknown. As our study area is located
close to a hard border of the overall habitat, the Sebangau
River, it may be possible that our camera traps were positioned at the edge of
the individuals’ home ranges, and short gaps in detection could indicate that
the individuals spent some time in their core range.
In
our study area, four males appeared to overlap (M2, M3, M4, and M5 all overlap
amongst themselves; Pallemaerts et al. 2019). This suggests some combination of temporal
partitioning and tolerance, as also reported for the Leopard Cat (Grassman et al. 2005) and Jaguar (Harmsen
et al. 2009; Guilder et al. 2015). We
did not record any instance of wounds that indicate fighting. Several photographs revealed instances of
olfactory communication like urine spraying, cheek rubbing and tree scratching,
all of which have been interpreted as territorial behaviour by Allen et al.
(2016). They could equally function in
the maintenance of temporal avoidance.
Another,
more striking, result of our camera trap survey was the low detection rate of
female Sunda Clouded Leopards. Over the course of 10 years, we were able to
identify only one female, namely F1 (Image 2).
This male-biased detection is recurrent in studies on both Sunda Clouded Leopard and mainland Clouded Leopard Neofelis nebulosa
(Cheyne & Macdonald 2011; Wilting et al. 2012; Cheyne et al. 2013; Sollmann et al. 2014; Mohamad et al. 2015; Hearn et al. 2017),
and has been noted for the Jaguar (Harmsen et al.
2009) and the Leopard (Gray & Prum
2012) as well. To date, only Borah et
al. (2014) reported a more balanced detection of both Leopard and mainland
Clouded Leopard females and males, based on a camera trapping design that
covered the 300km2 large study area homogeneously. The lack of photographs of females may be due
to their spending more time in trees (Wilting et al. 2012) and avoiding trails
and forest edges, which may affect their detectability (Cheyne et al.
2013). Adult female Sunda
Clouded Leopards weigh only half as much as adult males (Nájera
et al. 2017), adding to the likelihood of niche separation (Quentin Phillipps pers. comm. 10 September 2018). While there is no evidence to date that Sunda Clouded Leopards move in the canopy, there have been
reports of them hunting in trees (Matsuda et al. 2008; Wilcox et al. 2016). We recommend to place camera traps in a
systematic grid to enhance the chances for a more balanced representation of
both sexes of the Sunda Clouded Leopard in future
studies. The
possibility of spatial segregation between females and males, and their
different exposure and reactions to disturbance, are all factors relevant for
planning adequate conservation measures, and thus merit further investigation.
Table 1. Summary
of all adult Sunda Clouded Leopard detections within
the study area from May 2008 until February 2018. Individuals were classified
as residents if they were recorded in the study area for longer than six
months. One captive-born individual released in the study area was excluded
from this analysis.
|
ID |
Sex |
No. of notionally independent
photographs |
First detection |
Last detection |
Residence (months) |
Status |
|
M1 |
M |
9 |
3.vii.2008 |
28.vii.2011 |
36 |
Resident |
|
M2 |
M |
29 |
14.viii.2008 |
25.xii.2011 |
40 |
Resident |
|
M3 |
M |
20 |
4.xi.2009 |
29.x.2015 |
71 |
Resident |
|
M4 |
M |
32 |
3.xi.2009 |
20.viii.2015 |
69 |
Resident |
|
M5 |
M |
29 |
26.x.2013 |
25.i.2018 |
51 |
Resident |
|
M6 |
M |
3 |
14.ii.2014 |
14.ii.2014 |
0 |
Transient |
|
M8 |
M |
11 |
6.xi.2014 |
5.xi.2016 |
24 |
Resident |
|
M9 |
M |
3 |
9.x.2016 |
8.xii.2017 |
14 |
Resident |
|
F1 |
F |
2 |
8.x.2009 |
9.x.2009 |
0 |
Transient |
|
ID1 |
? |
1 |
5.x.2014 |
5.x.2014 |
0 |
Transient |
|
ID2 |
? |
5 |
25.xii.2013 |
8.ix.2014 |
9 |
Resident |
M-—male | F—female | ?—unknown sex.
For figures
& images - - click here
References
Adul, B. Ripoll, S.H. Limin & S.M. Cheyne (2015). Felids of Sebangau:
camera trapping to estimate activity patterns and population abundance in
Central Kalimantan, Indonesia. Biodiversitas
16(2): 151–155. https://doi.org/10.13057/biodiv/d160208
Allen, M.L., H.U. Wittmer, E. Setiawan, S. Jaffe & A.J. Marshall (2016). Scent marking in Sunda Clouded Leopards (Neofelis
diardi): novel observations close a key gap in
understanding felid communication behaviours. Scientific
Reports 6: e35433. https://doi.org/10.1038/srep35433
Borah, J., T. Sharma, D. Das, N. Rabha, N. Kakati, A. Basumatary, M.F. Ahmed
& K. Vattakaven (2014). Abundance and density
estimates for common Leopard Panthera pardus and Clouded Leopard Neofelis
nebulosa
in Manas National Park, Assam, India. Oryx 48(1):
149–155. https://doi.org/10.1017/S0030605312000373
Cheyne, S.M. (2010). Behavioural
ecology of Gibbons (Hylobates albibarbis) in a degraded peat-swamp forest. Pp. 121–156
In: Gursky, S. & J. Supriatna
(eds.). Indonesian Primates. Springer New York, New York, NY, 410pp. https://doi.org/10.1007/978-1-4419-1560-3_8
Cheyne, S.M. & D.W. Macdonald (2011). Wild felid diversity and
activity patterns in Sabangau peat-swamp forest,
Indonesian Borneo. Oryx 45(1): 119–124.
https://doi.org/10.1017/S003060531000133X
Cheyne, S.M., D.J. Stark, S.H. Limin
& D.W. Macdonald (2013). First estimates of population ecology and threats to Sunda Clouded Leopards Neofelis
diardi in a peat-swamp forest, Indonesia. Endangered
Species Research 22: 1–9. https://doi.org/10.3354/esr00525
Gaveau, D.L.A., S. Sloan, E. Molidena, H. Yaen, D. Sheil, N.K. Abram, M. Ancrenaz,
R. Nasi, M. Quinones, N. Wielaard & E. Meijaard (2014). Four decades of forest persistence,
clearance and logging on Borneo. PLoS One
9(7): e101654. https://doi.org/10.1371/journal.pone.0101654
Goodrich, J.M., D.G. Miquelle, E.N.
Smirnov, L.L. Kerley, H.B. Quigley, M.G. Hornocker
(2010).
Spatial structure of Amur (Siberian) Tigers (Panthera
tigris altaica) on Sikhote-Alin Biosphere Zapovednik,
Russia. Journal of Mammalogy 91(3): 737–748. https://doi.org/10.1644/09-MAMM-A-293.1
Grassman, L.I., M.E. Tewes, N.J. Silvy & K. Kreetiyutanont (2005). Spatial organization and diet of the
Leopard Cat (Prionailurus bengalensis) in north-central Thailand. Journal of
Zoology 266(1): 45–54. https://doi.org/10.1017/S095283690500659X
Gray, T.N.E. & S. Prum (2012). Leopard density in
post-conflict landscape, Cambodia: evidence from spatially explicit capture–recapture.
The Journal of Wildlife Management 76(1): 163–169. https://doi.org/10.1002/jwmg.230
Guilder, J., B. Barca, S. Arroyo-Arce, R. Gramajo
& R. Salom-Pérez (2015). Jaguars (Panthera onca)
increase kill utilization rates and share prey in response to seasonal
fluctuations in nesting Green Turtle (Chelonia
mydas mydas) abundance
in Tortuguero National Park, Costa Rica. Mammalian
Biology 80: 65–72. https://doi.org/10.1016/j.mambio.2014.11.005
Harmsen, B.J., R.J. Foster, S.C.
Silver, L.E.T. Ostro & C.P. Doncaster
(2009).
Spatial and temporal interactions of sympatric Jaguars (Panthera
onca) and Pumas (Puma concolor)
in a Neotropical forest. Journal of Mammalogy 90(3): 612–620. https://doi.org/10.1644/08-MAMM-A-140R.1
Hearn, A.J., J. Ross, D. Pamin, H.
Bernard, L. Hunter & D.W. Macdonald (2013). Insights into the spatial and
temporal ecology of the Sunda Clouded Leopard Neofelis diardi. Raffles
Bulletin of Zoology 61(2): 871–875.
Hearn, A.J., J. Ross, D.W. Macdonald, G. Bolongon,
S.M. Cheyne, A. Mohamed, H. Samejima, J.F. Brodie, A.
Giordano, R. Alfred & R. Boonratana (2016). Predicted distribution of
the Sunda Clouded Leopard Neofelis
diardi (Mammalia: Carnivora: Felidae) on Borneo. Raffles
Bulletin of Zoology Supplement 33: 149–156.
Hearn, A.J., J. Ross, H. Bernard, A.S. Bakar, B. Goossens, L.T.B. Hunter & D.W. Macdonald (2017). Responses of Sunda Clouded Leopard Neofelis
diardi population density to anthropogenic
disturbance: refining estimates of its conservation status in Sabah. Oryx
53(4): 643–653. https://doi.org/10.1017/S0030605317001065
Hearn, A.J., J. Ross, J. Brodie, S.M. Cheyne, I.A. Haidir, B. Loken, J. Mathai, A.
Wilting & J. McCarthy (2015). Neofelis diardi.
The IUCN Red List of Threatened Species 2015: e.T136603A97212874. Downloaded on
16 October 2020. https://doi.org/10.2305/IUCN.UK.2015-4.RLTS.T136603A50664601.en
Hemker, T.P., F.G. Lindzey & B.B. Ackerman (1984). Population
characteristics and movement patterns of Cougars in southern Utah. Journal
of Wildlife Management 48(4): 1275–1284. https://doi.org/10.2307/3801788
Matsuda, I., A. Tuuga & S.
Higashi (2008). Clouded Leopard (Neofelis diardi) predation on Proboscis Monkeys (Nasalis larvatus)
in Sabah, Malaysia. Primates 49(3): 227–231. https://doi.org/10.1007/s10329-008-0085-2
Mohamad, S., D. Rayan, W. Christopher, M. Hamirul,
A. Mohamed, C. Lau & E. Siwan (2015). The first description of
population density and habitat use of the mainland Clouded Leopard Neofelis nebulosa within
a logged-primary forest in South East Asia. Population Ecology 57:
495–503. https://doi.org/10.1007/s10144-015-0494-1
Nájera, F., A.J. Hearn, J. Ross,
D.A. Ramírez Saldivar, M.N. Evans, S. Guerrero-Sánchez, S.K.S.S. Nathan, I. De
Gaspar Simón, D.W. Macdonald, B. Goossens & L.R.
Rueda (2017). Chemical immobilization of free-ranging and captive Sunda
Clouded Leopards (Neofelis diardi) with two anesthetic protocols:
medetomidine-ketamine and tiletamine-zolazepam. Journal of Veterinary
Medical Science 79(11): 1892–1898. https://doi.org/10.1292/jvms.17-0259
Núñez-Pérez, R. (2011). Estimating Jaguar
population density using camera-traps: a comparison with radio-telemetry
estimates. Journal of Zoology 285(1): 39–45. https://doi.org/10.1111/j.1469-7998.2011.00812.x
Odden, M. & P. Wegge (2005). Spacing and activity patterns of Leopards Panthera pardus in
the Royal Bardia National Park, Nepal. Wildlife
Biology 11(2): 145–152. https://doi.org/10.2981/0909-6396(2005)11[145:SAAPOL]2.0.CO;2
Page, S.E., J.O. Rieley, Ø.W. Shotyk & D. Weiss (1999). Interdependence of peat and
vegetation in a tropical peat swamp forest. Philosophical Transactions of
the Royal Society of London. Series B: Biological Sciences 354: 1885–1897. https://doi.org/10.1142/9781848160125_0014
Pallemaerts, L., Adul,
I.P. Kulu, K.A. Jeffers, D.W. Macdonald & S.M.
Cheyne (2019). Bornean Clouded Leopard minimum home range analysis, Indonesian Borneo. Cat
News 70: 32–35.
Sokal, R.R. & N.L. Oden
(1978).
Spatial autocorrelation in biology 1. Methodology. Biological Journal of the
Linnean Society 10(2): 199–228. https://doi.org/10.1111/j.1095-8312.1978.tb00013.x
Sollmann, R., M. Linkie, I.A. Haidir & D.W.
Macdonald (2014). Bringing clarity to the Clouded Leopard Neofelis
diardi: first density estimates from Sumatra. Oryx
48(4): 536–539. https://doi.org/10.1017/S003060531400043X
Wilcox, C.H., Supiansyah, A. Azis K., J. Zainuddin, Rahman
& S.M. Cheyne (2016). Predator mobbing and interspecies cooperation: an interaction
between gibbons, langurs and a Clouded Leopard. Asian Primates Journal 6(1):
20–26.
Wilting, A., F. Fischer, S. Abu Bakar & K.E. Linsenmair (2006). Clouded Leopards, the secretive
top-carnivore of South-East Asian rainforests: their distribution, status and
conservation needs in Sabah, Malaysia. BMC Ecology 6(16): 1–13. https://doi.org/10.1186/1472-6785-6-16
Wilting, A., A. Mohamed, L.N. Ambu,
P. Lagan, S. Mannan, H. Hofer & R. Sollmann
(2012). Density
of the Vulnerable Sunda Clouded Leopard Neofelis diardi in
two commercial forest reserves in Sabah, Malaysian Borneo. Oryx 46(3):
423–426. https://doi.org/10.1017/S0030605311001694
Zimmermann, F., C. Breitenmoser-Würste &
U. Breitenmoser (2005). Natal dispersal of Eurasian Lynx (Lynx lynx) in Switzerland. Journal
of Zoology 267(4): 381–395. https://doi.org/10.1017/S0952836905007545