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ISSN 0974-7907 (Online)
ISSN 0974-7893 (Print)
Journal of Threatened Taxa

12 March 2019 (Online & Print)

Vol. 11 | No. 4 | 13419–13510
PLATINUM
10.11609/jott.2019.11.4.13419-13510 OPEN
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Building evidence for conservation globally

SMALL WILD CATS SPECIAL ISSUE
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continued on the back inside cover

Caption: Sand Cat Felis margarita thinobius in the Kyzylkum Desert, Bukhara Province, Uzbekistan. Credit: Robert J. Burnside.
Journal of Threatened Taxa | www.threatenedtaxa.org | 12 March 2019 | 11(4): 13419–13420

Editorial
Foreword to the small wild cats special issue
Angie Appel 1 & Shomita Mukherjee 2
ISSN 0974-7907 (Online)
1
Wild Cat Network, 56470 Bad Marienberg, Germany. ISSN 0974-7893 (Print)
2
Salim Ali Centre for Ornithology and Natural History, Anaikatty, Coimbatore, Tamil Nadu 641108, India.
1
[email protected], 2 [email protected] PLATINUM
OPEN ACCESS

We are delighted to present this first special issue light upon the fate of the Caracal Caracal caracal in the
on small wild cats on a special occasion—the Journal of country. The author presents a sublime collection of re-
Threatened Taxa’s 21st anniversary! cords obtained during expeditions of over 2,000km on
Small wild cats are fascinating but elusive. Biologists dusty roads to remote areas.
are only beginning to fathom their vital ecological roles For decades, both the Fishing Cat Prionailurus viver-
in maintaining and preserving biodiversity. Yet, surveys rinus and the Clouded Leopard Neofelis nebulosa were
and conservation efforts targeting them are still under- known to inhabit Nepal’s subtropical jungles. And still,
funded. Most of them live in the long shadow cast by they show up in entirely unexpected sites, prompting
the more widely known Panthera cats that attract the the authors to contribute a new locality record for both
lion’s share of international funding. Much of what we species.
know about small wild cats is a result of by-catch data The cat of many costumes is known to occur in India
from camera trap surveys targeted at larger mammals. since the 1830s but has remained elusive in Buxa Tiger
There is still a dearth of people with passion for and ex- Reserve for more than 175 years. The reserve’s per-
pertise in research and conservation of small wild cats. sonnel discloses new records of the Asiatic Golden Cat
With this issue, we hope to raise your awareness for Catopuma temminckii.
their conservation needs and inspire you to join us in For the first time in India, a team of 14 authors pooled
unravelling their mysteries. their records on small wild cats obtained during several
The European Wildcat Felis silvestris is the only one years of camera trapping. Their contribution provides
presented in this issue that benefits from a conservation enthralling insights into the activity patterns of the Asi-
program. Running in Germany since 2004, this program atic Golden Cat, the Clouded Leopard, the Marbled Cat
aims at relinking forests and doubling the area inhabited Pardofelis marmorata, and Asia’s most successful small
by Europe’s smallest cat until 2019. The authors scruti- wild cat, the Leopard Cat Prionailurus bengalensis.
nize the relation between camera trap locations, human Asia’s smallest wild cat broke a record in Sri Lanka.
disturbance, and photographic capture success of Wild- The Rusty-spotted Cat Prionailurus rubiginosus showed
cats in an unprotected forest. up at a location where nobody would ever have expect-
Two contributions focus on small wild cats in Uz- ed it. The authors also present evidence for a breed-
bekistan’s Kyzylkum Desert. One accounts of the cryptic ing population of the species in a montane forest in the
Sand Cat Felis margarita caught red-handed in the act country.
of feeding on a large kill. The authors were fortunate to You need to know a cat’s prey to understand its hab-
happen upon amazing video footage. The other sheds its, once wrote a renowned wildlife scientist. A team

DOI: https://doi.org/10.11609/jott.4939.11.4.13419-13420

Date of publication: 12 March 2019 (online & print)

Manuscript details: #4939 | Received 08 March 2019

Citation: Appel, A. & S. Mukherjee (2019). Foreword to the small wild cats special issue. Journal of Threatened Taxa 11(4): 13419–13420; https://doi.org/10.11609/
jott.4939.11.4.13419-13420

Copyright: © Appel & Mukherjee 2019. Creative Commons Attribution 4.0 International License. JoTT allows unrestricted use, reproduction, and distribution of
this article in any medium by adequate credit to the author(s) and the source of publication.

13419
Foreword Appel & Mukherjee

of four authors took this insight to heart and into the We thank Luigi Boitani, Andrew Kitchener, Christine
field of a biosphere reserve in Thailand. For their study Thiel-Bender, Mariya Gritsina, Alexander Sliwa, Arash
on the Leopard Cat Prionailurus bengalensis, they used Ghoddousi, Anna Barashkova, Daniel Willcox, P.O.
a novel approach—they simultaneously set up camera Nameer, Anwaruddin Choudhury, Wanlop Chutipong,
traps and caught rodents—with intriguing results! Mohd Azlan Jayasilan bin Abdul Gulam Azad, Lon
A team of eight authors set out tracing records of Grassman Jr., Tawqir Bashir, Kurtis Jai-Chyi Pei, J.W.
Fishing Cat in Thailand, teasing apart authenticated Duckworth, Andreas Wilting, Carl Traeholt, Babu Ram
from alleged ones. Their update reveals that not only Lamichhane, Honnavali Kumara, and Sagar Dahal for
protected areas provide suitable habitat for the ‘queen reviewing the submitted manuscripts for this special
of the marshes’. It also indicates that the Fishing Cat issue.
found a group of champions who are committed to its We have one wish—that we will all be inspired by
protection! the curiosity and enthusiasm of those who contributed
Ten years of continuous camera trapping surveys to this issue, particularly when it comes to understand-
reveal the diversity of small wild cats in Central Kalim- ing and protecting our environment. As Alexander von
antan: the Marbled Cat, the Sunda Clouded Leopard Humboldt wrote, people only protect what they love. If
Neofelis diardi, the Bay Cat Catopuma badia, the Sunda people are to protect small wild cats, they must first and
Leopard Cat Prionailurus javanensis, and the Flat-head- foremost understand and come to have a profound af-
ed Cat P. planiceps. The authors give a comprehensive fection for nature to marvel and admire, but above all,
insight into their ecology and persistence in Borneo’s to recount gripping tales.
fire-prone peat-swamp forests. Stay fascinated and share your experiences about
small wild cats for the next issue!

Threatened Taxa

13420 Journal of Threatened Taxa | www.threatenedtaxa.org | 12 March 2019 | 11(4): 13419–13420

Journal of Threatened Taxa | www.threatenedtaxa.org | 12 March 2019 | 11(4): 13421–13431

Article
Using camera traps to study the elusive European Wildcat
Felis silvestris silvestris Schreber, 1777 (Carnivora: Felidae) in
central Germany: what makes a good camera trapping site? ISSN 0974-7907 (Online)
ISSN 0974-7893 (Print)
Harald Wening 1, Lynne Werner 2, Matthias Waltert 3 & Markus Port 4
PLATINUM
Work Group Endangered Species Conservation, University of Göttingen, Bürgerstraße 50, 37075 Göttingen,
1,3,4 OPEN ACCESS
Germany.
2
Friends of the Earth Germany, BUND Hessen, Geleitsstraße 14, 60599 Frankfurt am Main, Germany.
4
Department of Behavioral Ecology, University of Göttingen, Kellnerweg 6, 37077 Göttingen, Germany.
1
[email protected], 2 [email protected], 3 [email protected],
4
[email protected] (corresponding author)

Abstract: Camera traping is a widely used method to study the abundance and population density of elusive terrestrial animals. To make
full use of this method, it is necessary to obtain high photographic capture rates of the target species. We examine what characteristics
of camera trapping sites are associated with high photographic capture rates of European Wildcat Felis silvestris silvestris. We measured
Wildcat capture rates across 25 camera trapping sites located in a 20km² study area within an unprotected low mountain range forest in
central Germany. We measured the distance of each trapping site to the forest boundary, to the next watercourse, and to the next human
settlement, and broadly defined the type of forest structure the site was located in. None of these site characteristics, however, predicted
wildcat photographic capture success. We also examined the degree of human disturbance at the site, measured as the photographic
capture rate of humans (including vehicles). Wildcats were detected at similar rates on dirt or gravel roads (heavily used by humans) as
on soft-surfaced paths or logging trails (less frequently used by humans), and the degree of human disturbance across sites did not affect
wildcat capture success. We, therefore, suggest that trail features such as course, curvature and width, or vegetation density along the
trail are more important determinants of Wildcat capture success than habitat characteristics. We conclude that for European Wildcats,
as for many larger felids, forest roads provide suitable camera trapping sites and that Wildcats are fairly tolerant towards human traffic
on these roads.

Keywords: Capture rate, habitat selection, human disturbance, trapping success.

German Abstract: Der Einsatz von Fotofallen ist eine gängige Methode, um die Abundanz und Populationsdichte heimlicher Säugetierarten
zu untersuchen. Um diese Methode voll ausschöpfen zu können, ist eine gründliche, auf die zu untersuchende Tierart abgestimmte Auswahl
der Fotofallen-Standorte nötig. Die vorliegende Studie untersucht die Fotofrequenz der Europäischen Wildkatze (Felis silvestris silvestris)
an 25 Fotofallen-Standorten in einem 20 km² großen Untersuchungsgebiet in einem Wirtschaftswald des deutschen Mittelgebirges.
Sie geht der Frage nach, welche Charakteristiken von Fotofallen-Standorten mit einer hohen Fotofrequenz der Europäischen Wildkatze
einhergehen. Gemessen wurden die Entfernung des Fotofallen-Standorts zum Waldesrand, zum nächsten Wasserlauf und zur nächsten
menschlichen Siedlung. Außerdem wurde der Habitattyp des Fotofallen-Standortes grob bestimmt und der Grad des durch den Menschen
verursachten Störung am Fotofallen-Standort als die Foto-frequenz von Menschen (einschließlich Fahrzeugen) gemessen. Wildkatzen
wurden in ähnlichen Häufigkeiten auf Forst- und Waldwegen fotografiert wie auf Fußpfaden und Rückewegen. Jedoch hatte keine der
von uns gemessenen Variablen einen Einfluss auf die Häufigkeit, mit der Wildkatzen fotografiert wurden. Auch der Grad der durch den
Menschen verursachten Störung wirkte sich nicht nicht auf die Häufigkeit aus, mit der Wildkatzen fotografiert wurden. Diese Ergebnisse
legen nahe, dass Forst- und Waldwege für die Wildkatze ebenso gute Fotofallen-Standorte darstellen, wie für viele größere Katzen. Zudem
scheint die Wildkatze relativ tolerant zu sein gegenüber Störungen durch Menschen und Fahrzeuge auf diesen Wegen.

DOI: https://doi.org/10.11609/jott.4498.11.4.13421-13431 | ZooBank: urn:lsid:zoobank.org:pub:27953FBE-0683-45B1-A224-193230E626CF

Editor: Angie Appel, Wild Cat Network, Germany. Date of publication: 12 March 2019 (online & print)
Manuscript details: #4498 | Received 18 August 2018 | Final received 15 January 2019 | Finally accepted 24 February 2019
Citation: Wening, H., L. Werner, M. Waltert & M. Port (2019). Using camera traps to study the elusive European Wildcat Felis silvestris silvestris Schreber, 1777
(Carnivora: Felidae) in central Germany: what makes a good camera trapping site?. Journal of Threatened Taxa 11(4): 13421–13431; https://doi.org/10.11609/
jott.4498.11.4.13421-13431
Copyright: © Wening et al. 2019. Creative Commons Attribution 4.0 International License. JoTT allows unrestricted use, reproduction, and distribution of this article
in any medium by adequate credit to the author(s) and the source of publication.
Funding: Heinz Sielmann Stiftung, Duderstadt, Germany, BUND Hessen, Frankfurt, Germany.
Competing interests: The authors declare no competing interests.
For Author details, Author contribution & Acknowledgements see end of this article.

13421
Camera traps study of European Wildcat in central Germany Wening et al.

INTRODUCTION important advantage of this method is that samples

can be sexed and their taxonomic status (i.e., Wildcat,
The past few decades saw a comeback of some domestic cat, or hybrid) can be determined reliably. In
previously rare or locally extinct large and medium- addition, samples can be genotyped, allowing for DNA-
sized carnivores in central Europe (Chapron et al. 2014; based individual identification of the sampled animals.
Boitani & Linnell 2015; Thiel-Bender 2015). One of these This information can then be used for abundance
species is the European Wildcat Felis silvestris silvestris estimations based on capture-recapture approaches
Schreber 1777, even though in comparison to large (Kéry et al. 2011). For capture-recapture models to
mammalian carnivores such as Wolf Canis lupus, Lynx produce reliable abundance estimates, however, a
Lynx lynx, and Brown Bear Ursus arctos, its comeback sufficiently large number of genotyped samples are
was more secretive in nature. required, rendering the lure-stick method an expensive
The European Wildcat is a small (app. 3–6 kg), sampling method.
solitarily-hunting, and predominantly nocturnal felid, Another widely applied method for monitoring
which in size and appearance is similar to domestic cat elusive animals is the use of camera traps (O’Conell et
Felis catus (Piechocki 1990; Kitchener et al. 2005; Thiel- al. 2010; Rovero & Zimmermann 2016). If individual
Bender 2015; Image 1). The species was once widely animals can be identified on camera trap images, this
distributed across central Europe. Within the last two method, too, can be used for abundance estimations
centuries, however, its population experienced a radical based on capture-recapture models (Zimmermann
decline owing to human persecution, habitat loss, and & Foresti 2016). This method is a common tool for
hybridization with domestic cats (Piechocki 1990; Nowell abundance estimation of striped and spotted felids such
& Jackson 1996; Beaumont et al. 2001; Pierpaoli et al. as Tiger Panthera tigris (Karanth 1995), Jaguar P. onca
2003; Oliveira et al. 2008; Klar et al. 2009; Macdonald (Harmsen et al. 2017), and European Lynx (Pesenti &
et al. 2010; Hartmann et al. 2013). Today, Wildcats are Zimmermann 2013), and was also applied to estimate
distributed in fragmented populations ranging from the abundance and population density of Wildcats (Can
Scotland to the Near East, and from Belarus to Portugal et al. 2011; Anile et al. 2014; Kilshaw et al. 2015; Velli et
(Yamaguchi et al. 2015). Even though wildcat populations al. 2015). Just like any other sampling method used to
are still threatened and/ or declining in some areas of carry out capture-recapture analyses, this method, too,
Europe, particularly on the Iberian Peninsula and in requires a sufficient number of samples (in this case,
Scotland, the species is currently expanding its range in identifiable images) for reliable abundance estimation.
several European countries (e.g., France: Say et al. 2012; Yet, in comparison to the lure-stick method, camera
Germany: Thiel-Bender 2015, Steyer et al. 2016). traps have the advantage that once the initial costs of
Until recently, Wildcats in Germany were considered purchasing the cameras are paid, any sample ‘collected’
to be distributed in two isolated populations, a central by the cameras (i.e., any image taken) comes at a low
German population and a western population, the latter cost. To make full use of this advantage, it is necessary to
presumably extending into France (Birlenbach & Klar make a sensible choice of trapping sites, in other words,
2009). A recent large-scale genetic census carried out to choose trapping sites that maximize the capture
between 2007 and 2013 suggests that these formerly success of the target species.
separated populations are now connected such that the The present study examines trapping site
species appears to be continuously distributed across characteristics for the study of European Wildcats in a
large parts of western and central Germany (Steyer central European low mountain range, i.e., what site
et al. 2016). Moreover, 44% of Wildcat samples were characteristics are associated with high photographic
obtained from locations outside the previously known capture rates of European Wildcats. A first important
Wildcat distribution. These results illustrate that factor that comes into mind when choosing a suitable
German Wildcats are currently regaining large parts of camera trapping site is habitat. Habitat selection
their historic range. Estimates of Wildcat population is comparatively well-studied in European Wildcats
density, however, are often lacking, particularly from the (Okarma et al. 2002; Lozano et al. 2003; Hötzel et al. 2007;
newly colonized areas. Klar et al. 2008; Monterroso et al. 2009; Jerosch et al.
A frequently applied method for wildcat population 2017) and a number of habitat preferences of European
monitoring is the use of valerian-treated lure sticks Wildcats were identified. First of all, even though recent
(Hupe & Simon 2007; Steyer et al. 2013), which enable studies demonstrated that Wildcats can use significant
the collection of hair samples for genetic analyses. An proportions of open, agriculturally-dominated landscape

13422 Journal of Threatened Taxa | www.threatenedtaxa.org | 12 March 2019 | 11(4): 13421–13431

Camera traps study of European Wildcat in central Germany Wening et al.

Image 1. Camera trap image of a European Wildcat Felis silvestris silvestris, taken in November 2016 in the Melsunger Bergland. © University
of Göttingen Lynx Project.

(Jerosch et al. 2017; Götz et al. 2018), in central Europe, and (v) fewer Wildcats at sites located within the less-
the Wildcat is traditionally described as a species bound preferred habitats such as coniferous stands.
to forests (Piechocki 1990; Nowell & Jackson 1996; To examine whether the photographic capture rate of
Hötzel et al. 2007; Klar et al. 2008). Within forests, radio Wildcats is affected by the above habitat characteristics
tracking studies revealed that Wildcats spend more time and/ or the proximity to human settlements, we
close to the forest boundary and seem to be attracted analyzed data collected over a period of three months
also by watercourses, meadows, and open areas within at 25 camera trapping sites within a 20km² study area
the forest (Klar et al. 2008), presumably because these in central Germany. This study area is located within an
habitats are characterized by higher prey population unprotected forest that is used for timber production and
densities. A preference for such ecotone habitats was recreation activities such as hiking, mountain biking, and
also revealed by snow tracking in the Polish Carpathian hunting. Because our trapping sites varied to the extent
Mountains (Okarma et al. 2002). Moreover, Wildcats they were exposed to human disturbance (including
seem to prefer wind-throw areas and young succession vehicles), we also examined whether the photographic
stages with dense undergrowth while coniferous stands capture rate of Wildcats was affected by the degree of
tend to be avoided (Okarma et al. 2002; Hötzel et al. human disturbance at the sites.
2007). Lastly, human infrastructure, such as roads or
villages, are also usually avoided by Wildcats, though
beyond a certain distance (ca. 200m to roads and single STUDY AREA
houses, ca. 900m to villages) human infrastructure does
not seem to affect wildcat ranging pattern (Klar et al. The study area was located in a low mountain range
2008). Taking these habitat preferences into account, known as Melsunger Bergland, approximately 20km
we should thus expect the photographic capture rate southeast of the city of Kassel in central Germany (Fig. 1).
of Wildcats to increase when camera trapping sites are The study area is almost completely covered by forest,
located (i) closer to open areas within forests or to the consisting of approximately 40% broad-leaved forest,
forest boundary, (ii) closer to watercourses, and (iii) 30% mixed forest, and 30% coniferous forest. The forest
further from human settlements. We should also expect is broken up only by the village of Kehrenbach (with
(iv) more Wildcats to be camera trapped at sites located a population of 320 inhabitants) and its surrounding
within preferred habitats such as wind-thrown areas, fields, located approximately in the centre of the study

Journal of Threatened Taxa | www.threatenedtaxa.org | 12 March 2019 | 11(4): 13421–13431 13423

Camera traps study of European Wildcat in central Germany Wening et al.

Figure 1. The left-hand image shows a schematic map of Germany with the federal state of Hesse in the center. The right-hand image shows
the study area located in the north of Hesse (indicated by a red square on the left-hand image).

area, as well as by a small road (leading to the village) sites within the forest, one site located in every cell of a
in the south of the study area. Even though the forest 1kmx1km grid (omitting only the village of Kehrenbach
is used for timber production and recreation activities, and its surrounding fields). The minimum convex
it supports a diverse community of animal species, polygon encompassing all stations amounted to 20km²
including large mammals such as Roe Deer Capreolus with an average (±SD) distance between camera sites of
capreolus, Red Deer Cervus elaphus, Wild Pig Sus scrofa, 863m (±207m). Sites were located either along forest
European Badger Meles meles, and Red Fox Vulpes roads (n=9) or forest trails (n=16).
vulpes. Moreover, a small population of Eurasian Lynx All cameras used were Cuddeback® camera traps
started to recolonize the area since 2009 (Denk 2016). (Cuddeback Digital, Green Bay, USA) of the models
The elevation of the study area ranges between Ambush® and C1®. These are heat- and motion-triggered
300m in the valley of the river Fulda in the west and cameras that record colour images both at day and night
500m in the east. With an annual mean precipitation of using a white flash. We installed two camera traps per
676cm and average temperatures from 0.2°C in January station, one on each side of the road or trail, to obtain
to 17.7°C in July, the Melsunger Bergland is located in images of both flanks of a passing animal. Cameras
the transition zone between Atlantic and continental were set up 3.2–22.1 m apart from each other along
climate with mild and humid winters. the road to avoid overexposure of images by the flash
of the opposite camera. Delay time between successive
images was set to the shortest time frame possible
METHODS (1–60 s, depending on camera type and time of day).
Camera traps were secured inside metal boxes, locked
Camera trap placement with a padlock or cable lock, and attached to a tree or a
The study was carried out between 26 June and 8 pole approximately 30cm above ground. Camera traps
October 2017 as part of a project aimed at estimating were checked every four weeks to replace batteries and
Wildcat population density in the area (Werner & Port SD cards and to clear the areas in front of the cameras of
in preparation). Cameras were placed at 25 trapping overgrowing vegetation.

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Camera traps study of European Wildcat in central Germany Wening et al.

Habitat characteristics (2) Mixed/ broad-leaved forest in the succession

To define habitat characteristics of our trapping sites stages of stem exclusion and older: a very broad category
and the proximity to human infrastructure, we used that characterizes large parts of the study area. Mixed
aerial images of the study area and forest management and broad-leaved forest were not further distinguished
data generously provided by HessenForst, the forestry since their understorey structure is similar (Image 2a).
management unit of the German federal state of Hesse. (3) Pure coniferous stands in the succession stage
Geographic data were processed using ArcMap 10.5.1 stem exclusion or old: this category usually consists of
(Esri, Inc., Redlands, CA, USA). even-aged, dark stands, usually of Spruce Picea trees
To determine the distance between the trapping with poor understorey (Image 2c).
site and the closest forest boundary, we used aerial
images. As forest boundary, we defined any ‘outer’ Human disturbance
forest boundary (usually to villages, fields, or roads; Fig. The degree of human disturbance at the trapping
1), or any boundary to clearings within the forest with a site was measured as the number of camera trap images
minimum area of 20mx20m. Likewise, we measured the showing humans and/ or vehicles (e.g., bikes, cars,
distance of our trapping sites to the closest watercourse trucks). In compliance with the data protection policies
(creek, pond, or ditch) with a permanent water body. of the German federal state of Hesse, these images were
Lastly, to determine the distance between the trapping deleted afterwards.
site and human settlements, we measured the distance
to the closest house with regular human activity Wildcat detection and data analyses
(excluding, for example, barns or similar buildings). The response variable in our analyses was the number
We classified the habitat of our trapping sites based of independent Wildcat detections that occurred at a
on the type of forest stand the cameras were located trapping site. As a Wildcat detection, we defined any
in. We distinguished broadly between three categories Wildcat recorded after at least five minutes passed
of forest structure that were found to predict Wildcat since the last recorded Wildcat. Wildcats captured
ranging patterns in previous studies (Okarma et al. simultaneously by both cameras at a site were counted
2002; Hötzel et al. 2007). Owing to the small number of as only one detection. Moreover, images of a female
trapping sites, a finer subdivision of forest structure types with kittens were counted as one detection. In general,
does not appear functional. These three categories were we included all recordings of wild-living cats that
defined based on the dominating type of tree (broad- showed morphologic features characteristic for Wildcats
leaved, mixed, or coniferous) and the following four (Kitchener et al. 2005; Kilshaw et al. 2015; Thiel-Bender
succession stages (Smith et al. 1997): (1) stand initiation: 2015). We would have excluded images of wild-living
the earliest succession stage, consisting of young trees cats that showed features characteristic for domestic
with a diameter at breast height (DBH) of usually <7cm, cats (e.g., white paws, white spots on flanks, pointed tail
(2) stem exclusion: a succession stage consisting of trees tip), though no such individuals were recorded during
in early development, with DBH<20cm, (3) understorey our study.
reinitiation: the stand developed a stratification with Since it is difficult to distinguish Wildcats from hybrids
canopy, midstorey, and understorey, with older trees based on camera trap images (but see Kilshaw et al.
reaching a DBH between 21cm and 50cm, and (4) old 2015), however, we cannot exclude the possibility that
growth: the stand reached its development climax and our sample of wild-living cats contained some hybrids as
is multi-aged and multi-layered with a dense vegetation well, though we note that the degree of hybridization is
and a relatively high percentage of dead wood. Stands low (3.9%) in German Wildcats (Steyer et al. 2016).
of this succession stage were rare in the study area and We examined the relationship between our response
no trapping site was located in it. variable (number of Wildcat detections) and the various
In this way, and partially following the classifications site characteristics (distance to forest boundary, distance
used by Hötzel et al. (2007), we defined the following to water, distance to settlement, forest structure, human
three forest structure types: disturbance) using general linear models (GLMs) with
(1) Wind-throw/ stand initiation: Areas that log link function and negative binomial error structure.
experienced a recent disturbance, either by storm In order to avoid overfitting of our models, we ran two
damage or clear-cutting, and are now naturally or separate models: a first model contained the ‘habitat
artificially regenerating. Bushes such as blackberry and characteristics’, distance to forest boundary, distance
dead wood create a dense ground cover (Image 2e). to water, distance to settlement, and forest structure

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Camera traps study of European Wildcat in central Germany Wening et al.

Image 2. Trapping location types and habitat types: A - broad-leaved forest typical for the study area; B - a typical forest road; C - coniferous
stand; D - example of a forest trail; E - a wind-thrown area regrowing for approximately 10 years. © M. Port.

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Camera traps study of European Wildcat in central Germany Wening et al.

as explanatory variables. A second model contained Wildcat was recorded at each of our 25 sites.
all predictors found to significantly affect the number
of Wildcat detections in the first model, as well as the Habitat characteristics
degree of human disturbance measured at the trapping Trapping sites were located between 0m and 577m
sites. To account for the possibility that Wildcat records away from the next forest boundary (mean=226.88m,
differed between sites located on forest roads and sites SD=170.03m), between 1m and 629m away from the
located on forest trails, this model contained the location next watercourse (mean=352.44m, SD=194.35m), and
of the site (road, trail) as a further predictor. To account between 461m and 1,475m away from the next human
for differences in camera operation times between sites, settlement (mean=934.32m, SD=344.6m). Eleven sites
both models contained camera operation time (in days) were located within wind-throw areas/ stand initiations
as an offset term. Analyses were performed in R (R Core (forest structure type 1), 11 sites in broad-leaved or
Team, 2016, Vienna, Austria) using the package “Mass”. mixed forest areas of succession stages 2 or older (forest
structure type 2), while only three sites were located
in coniferous stands (forest structure type 3). None of
RESULT the habitat characteristics had a significant effect on the
number of recorded Wildcats (Table 1). On average, only
Trapping success 4 (SD=2) Wildcats were detected in coniferous stands,
Even though several cameras malfunctioned over whereas on average 6.45 (SD=5.12) and 7.36 (SD=4.8)
the course of the study, at least one camera per site Wildcats were detected in wind-throw areas and mixed
operated for 99–105 days (day=24h). Only at one site or broad-leaved forest stands, respectively, but this
both cameras broke down during the same time period, difference was statistically not significant.
such that this site had only 86 recording days. In total,
our cameras operated for 2,552 trapping days (defined Human disturbance
as the number of sites multiplied by the number of The number of human detections ranged between 0
days during which at least one camera per site was and 1058 (mean=152.84, SD=262.2) and was on average
operational). markedly higher at the nine sites located on forest roads
We recorded a total number of 164 Wildcat (mean=371.7, SD=358.99) than at the 16 sites located
detections (including possible hybrids), resulting in on trails (mean=42.93, SD=51.48). On average, 7.4
a capture rate of 6.43 records per 100 trap days. The (SD=5.01) Wildcats were recorded on forest roads, while
number of Wildcat records across sites ranged between on average 6 (SD=4.55) Wildcats were recorded on trails.
1 and 14 detections (mean=6.56, SD=4.69). At least one This difference was statistically not significant (Table 2).

Table 1. Results of the general linear model relating Wildcat capture rate to habitat characteristics. Forest structure type “Windthrow/ stand
initiation” is used as the baseline level and is represented by the intercept.

β SE z p 95% CI

Intercept -2.99 0.75 -3.95 <0.001 -4.52 -1.43

Distance water 0.001 0.001 1.06 0.29 -0.001 0.003

Distance forest boundary 0.0004 0.001 0.34 0.74 -0.002 0.003

Distance village -0.0002 0.0005 -0.4 0.69 -0.001 0.0009

Broad leaved/ mixed stand 0.1 0.38 0.27 0.79 -0.69 0.91

Coniferous stand -0.71 0.56 -1.25 0.21 -1.81 0.42

Table 2. Results of the general linear model relating Wildcat capture rate to location type and human disturbance. Location type “forest trail”
is used as the baseline level and is represented by the intercept.

β SE z p 95% CI

Intercept -2.84 0.2 -14.33 <0.001 -3.22 -2.44

Location: forest road 0.09 0.35 0.25 0.8 -0.6 0.8

Human disturbance 0.0004 0.0006 0.53 0.59 -0.0009 0.0018

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Camera traps study of European Wildcat in central Germany Wening et al.

The degree of human disturbance at the site had no radio collars can usually deliver several locations of an
effect on the number of Wildcat records. animal per day. In this way, they can draw a dense and
detailed picture of animal ranging patterns. Likewise,
snow tracking can also provide detailed insights into
DISCUSSION how frequently different habitat types are used by the
animals (Okarma et al. 2002). In contrast, camera traps
This study examined whether characteristics of can only record animal movements at the location they
camera trapping sites, such as the distance of the site are installed. Because the main aim of our survey was
to the nearest forest boundary or the forest structure to estimate Wildcat population density (Werner & Port
type the site was located in, predicted the photographic in preparation), we followed previous sampling designs
capture success of European Wildcats. Even though and placed our cameras along human-made forest
the number of Wildcat records ranged between 1 and routes (roads and trails) as we presumed that Wildcats,
14 detections per site across 25 studied sites, none of like many other felids, would preferably use such trails
the site characteristics examined in our study had a (Karanth 1995; Di Bitetti et al. 2006; Harmsen et al.
significant effect on the number of recorded Wildcats. 2010; Weingarth et al. 2015). Forest trails yield the
best capture success if they are used by individuals of
Habitat characteristics the target species regularly and repeatedly, for example,
Radio tracking studies of Wildcats revealed that they because they connect preferentially used areas of an
avoid the proximity of human infrastructure, such as individual’s home range (e.g., areas preferentially used
settlements and roads, but only within a critical distance. for hunting or resting).
This distance is reported as approximately 900m to If this is the case, however, the capture success of
settlements by Klar et al. (2008) and as approximately the trapping site is not necessarily related to habitat
500m by Hötzel et al. (2007). The minimum distance of characteristics of that site (other than the trail itself).
our trapping sites to the nearest village was 461m, and For example, a site located within a coniferous stand, a
44% of sites were located more than 900m away from habitat presumably less preferred by Wildcats, but that
the next settlement. It is thus unsurprising that in our is located at a trail connecting two preferred hunting
study the (generally large) distance of the trapping sites grounds, might still yield higher capture probabilities
from human settlements did not affect Wildcat capture than a site located close to the forest boundary (a
success. preferred habitat). For presumably the same reason,
Both radio tracking and snow tracking studies our study did also not detect any effect of the forest
revealed that Wildcats strongly prefer ecotone habitats, structure type surrounding the camera site. These results
such as forest boundaries, clearings within the forest, or are in agreement with findings from three Neotropical
riparian areas (Okarma et al. 2002; Hötzel et al. 2007; felids—the photographic capture success of Jaguars,
Klar et al. 2008), presumably because these habitats Puma Puma concolor, and Ocelot Leopardus pardalis
are characterized by high prey population densities, depended only on features of the trail where camera
particularly of small rodents. For example, Hötzel et al. traps were installed, but not on habitat characteristics
(2007) often found Wildcats hunting in open areas at such as altitude or distance to water (Harmsen et al.
night, whereas they occurred in sheltered forest areas 2010).
during the day. Okarma et al. (2002) found a large A number of other variables that were not measured
fraction of Wildcat tracks along forest edges. Given these in the present study may affect Wildcat capture success.
strong preferences of Wildcats for forest boundaries and An important variable is the distribution and population
riparian areas, it is perhaps surprising that the distance density of prey. Researchers carrying out camera trap
of our trapping sites to such habitats did not predict studies, however, do not usually have information
Wildcat capture success. This is not due to a lack of on prey distributions and population densities, and
variation in these variables—the distance of our sites can only base their choices of trapping sites on
to forest boundaries and to watercourses varied greatly, habitat characteristics likely associated with high prey
ranging between 0m and 577m for forest boundaries, abundance, in our case, for example, wind-throw areas
and between 1m and 629m for watercourses. The most (Niethammer & Krapp 1982). This is the approach we
likely reason why our study did not have similar results took in the present study but, as reported above, none
to those of radio tracking or snow tracking studies is of the studied site characteristics had a significant effect
the markedly different methodology. For example, on Wildcat capture success. It is also possible that the

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Camera traps study of European Wildcat in central Germany Wening et al.

abundance of potential competitors such as Red Foxes (as in our study area), they are more tolerant towards
affects Wildcat ranging pattern and, as a consequence, human disturbance than commonly thought.
trapping success at particular sites. Lastly, it is possible
that we obtained different results at different times of What makes a good camera trapping site?
the year. Our study was carried out in summer when the We started this article by asking “What makes a
forest floor and some of the logging trails are covered by good camera trapping site?” for the study of European
dense herbaceous vegetation. At this time of the year, Wildcats in a central European low mountain range.
Wildcats may use forest roads more often than in winter Unfortunately, we cannot provide a conclusive answer to
when ground vegetation is less dense. Likewise, females this question as none of the site characteristics examined
might be detected less often on forest roads in spring in our study affected the photographic capture success
when they have dependent offspring and prefer areas of Wildcats. We, however, photo-captured at least one
with dense vegetation cover (Piechocki 1990; Jerosch et Wildcat at all of our sites (100%) and obtained a capture
al. 2017). Any recommendation on the choice of camera rate of 6.43 detections per 100 trap days. This capture
trapping sites in European Wildcats (and other species) rate is similar to the capture rate obtained by Anile et al.
should thus take into account possible seasonal changes (2014) (6.48 detections/ 100 trap days) and at least twice
in ranging patterns. It would be interesting to compare as high as the capture rate of any other camera trapping
results of the present study with results of prospective study of European Wildcats (Can et al. 2011; Kilshaw et
future studies carried out at different times of the year al. 2015; Velli et al. 2015). These results suggest that
and/ or in areas with different population densities of either Wildcat population density in the study area
Wildcats and their competitors. was particularly high and/ or that most, if not all, of
our trapping sites were suitable for detecting Wildcats
Human disturbance by means of camera trapping. Nevertheless, there is
Photographic capture frequencies of Wildcats were a large variation with respect to the trapping success
similar on forest roads and on forest trails, despite the across sites: at some sites as many as 13 (three sites) and
fact that forest roads were more extensively used by 14 (two sites) Wildcats were detected, whereas at five
humans, including cars and trucks. Moreover, the degree sites only a single detection occurred. Clearly, therefore,
of human disturbance, measured as the photographic some sites yielded higher capture rates than others. We
capture rate of humans (including vehicles) at the site, suggest that differences in capture success are more
did not affect Wildcat capture success. In fact, one closely related to features of the trail at which cameras
of the highest capture frequencies of Wildcats (14 were installed (Trolle & Kéry 2005; Di Bitetti et al. 2006;
detections) occurred at a site with the highest frequency Harmsen et al. 2010), for example, course, curvature,
of human captures (1,058 images of humans). These and width of the trail, or the density of the vegetation
results are largely in agreement with findings on other along the trail, rather than to the habitat characteristics
felids. For example, Ocelots preferred human-made examined in the present study.
dirt roads over animal trails, both in the Pantanal Still, there are at least two conclusions we can draw
wetlands of Brazil (Trolle & Kéry 2005) and in northern from the present study. A first conclusion is that human-
Argentina (Di Bitetti et al. 2006). Moreover, Ocelots, made dirt or gravel roads are as suitable as camera
Jaguars, and Pumas in Belize preferred wider trails over trapping sites for the rather small-bodied European
narrower trails (Harmsen et al. 2010). Finally, similar Wildcat as they are for many of her larger relatives. A
to the present study, the photographic capture rate of second conclusion is that the extent of human traffic
Bobcats Lynx rufus in Virginia (Kelly & Holub 2008) and along these roads does not affect Wildcat capture
of Eurasian Lynx in central Germany (Schröder 2016) success and that even roads heavily used by humans and
was not affected by the amount of human traffic at the vehicles can still make suitable camera trapping sites for
site. These results are easily explained by the fact that the study of European Wildcats.
most human traffic takes place during the day and thus
does not interfere with the predominantly nocturnal
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techniques and evaluation of their integration. European Journal of Author details: Harald Wening is an undergraduate student of conservation
Wildlife Research 61(5): 657–668; https://doi.org/10.1007/s10344- biology at the University of Göttingen. The present research constituted Harald’s
015-0936-2 Bachelor Thesis. Lynne Werner received her MSc in behavioural biology from
Göttingen University in 2017. She joined the Friends of the Earth Germany for
Weingarth, K., T. Zeppenfeld, C. Heibl, M. Heurich, L. Bufka, K.
the present work on wildcats, and is currently working as an environmental
Daniszová & J. Müller (2015). Hide and seek: extended camera-trap consultant. Matthias Waltert is professor of conservation biology at the
session lengths and autumn provide best parameters for estimating University of Göttingen. His research focuses on wildlife biology, wildlife
Lynx densities in mountainous areas. Biodiversity and Conservation management and human-wildlife conflicts, particularly in the old-world tropics
24: 2935–2952. and the middle east.Markus Port has been working for several years as a
Yamaguchi, N., A. Kitchener, C. Driscoll & B. Nussberger (2015). behavioural ecologist, but recently discovered his new passion for the applied
Felis silvestris. In: The IUCN Red List of Threatened Species: conservation of European felids. Markus is heading the University of Göttingen
Lynx Project.
e.T60354712A50652361. Accessed on 15 August 2018; https://doi.
org/10.2305/IUCN.UK.2015-2.RLTS.T60354712A50652361.en Author contribution: HW, LW & MP carried our research, HW, LW & MP
Zimmermann, F. & D. Foresti (2016). Capture-recapture methods performed analyses. MW & MP designed research, HW, LW, MW & MP wrote
for density estimation, pp95–141. In: Rovero, F. & F. Zimmermann the paper.
(eds.). Camera-trapping for Wildlife Research. Pelagic Publishing,
Exeter, UK, 320pp. Acknowledgements: We thank HessenForst, particularly the forestry office
Melsungen, for supporting of our field work. We also thank Jann Hellmuth for his
valuable help during data collection.

Threatened Taxa

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Journal of Threatened Taxa | www.threatenedtaxa.org | 12 March 2019 | 11(4): 13432–13447
Article

Activity patterns of the small and medium felid

(Mammalia: Carnivora: Felidae) guild in northeastern India
ISSN 0974-7907 (Online) Shomita Mukherjee 1 , Priya Singh 2 , André Pinto Silva 3 , Chandan Ri 4 , Kashmira Kakati 5 ,
ISSN 0974-7893 (Print)
Binod Borah 6 , Tana Tapi 7 , Sandesh Kadur 8 , Prafull Choudhary 9 , Shikha Srikant 10 ,
PLATINUM Surabhi Nadig 11 , R. Navya 12 , Mats Björklund 13 & Uma Ramakrishnan 14
OPEN ACCESS
1,3,9
Salim Ali Centre for Ornithology and Natural History (SACON), Anaikatty, Coimbatore, Tamil Nadu 641108, India.
2,6
Researchers for Wildlife Conservation (RWC), National Centre for Biological Sciences, GKVK Campus, Bellary Road,
Bengaluru, Karnataka 560065, India.
3,13
Department of Ecology and Genetics, Animal Ecology, Evolutionary Biology Centre, Uppsala University, Norbyvägen
18D, 752 36 Uppsala, Sweden.
3,11,12,14
National Centre for Biological Sciences, TIFR, Bellary Road, Bangalore, Karnataka 560065, India.
3
Centre for Ecology, Evolution and Environmental Changes, Faculdade de Ciências, Universidade de Lisboa, Campo
Grande, 1749-016 Lisbon, Portugal.
4,7
Pakke WS & Tiger Reserve, Seijosa, Pakke Kessang, Arunachal Pradesh 790103, India.
5
Madhuban Hill, Government Press Road, P.O. Bamunimaidam, Guwahati, Assam 781021, India.
8
Felis Creations, 480, 11th Cross, Upper Palace Orchards, Sadashivnagar, Bangalore, Karnataka 560080, India.
9
Wildlife Trust of India, Flat No. 403, Imperial building, Orbital Empire Society, Jaitada, Ekatmata Nagar Road, Nagpur,
Maharashtra 440016, India.
10
Wildlife Institute of India, Chandrabani, P.O. Box 18, Dehradun, India.
1
[email protected] (corresponding author), 2 [email protected], 3 [email protected],
4
[email protected], 5 [email protected], 6 [email protected], 7 [email protected],
8
[email protected], 9 [email protected], 10 [email protected], 11 [email protected],
12
[email protected], 13 [email protected], 14 [email protected]

Abstract: Fifteen extant species of cats inhabit India, and the northeastern region of the country is among the richest with nine species. Among
these are the “standard four”, an assemblage of Clouded Leopard Neofelis nebulosa, Asiatic Golden Cat Catopuma temminckii, Marbled Cat
Pardofelis marmorata, and Leopard Cat Prionailurus bengalensis, which also occur across southeastern Asia. Within India, despite several surveys
in this region, very little information exists on the ecology of this assemblage to explain their co-occurrence. In this paper, we put together data
from several independent camera trapping studies over 10 sites across northeastern India to examine and interpret diel activity patterns of this
group. While we present results for all the four species, we focus on two species, the Marbled Cat and Leopard Cat, which are of very similar
body size and are potential competitors. We used kernel density estimates to measure diel activity patterns of all four species and overlap in
activity between Marbled Cat and Leopard Cat at the regional scale as well as the point scale. We obtained 783 captures of the standard four
from >27,500 trap nights. The Asiatic Golden Cat and Marbled Cat were strongly diurnal, Clouded Leopard largely crepuscular and nocturnal,
and Leopard Cat largely nocturnal. The degree of overlap between Marbled Cat and Leopard Cat activity was low and in consensus with other
studies across southeastern Asia. We interpret this as the differing niche spaces of the two cats due to their specific pre-existing adaptations, not
restricted to the effects of competition. The point scale analysis when both cats are captured at the same location and separately show no shift
in activity pattern, supporting our hypothesis of pre-existing differences in resources, such as food, playing a major role in facilitating co-existence.
Our study, however, is preliminary and additional information with robust analysis is required to test this finding.
Keywords: Asiatic Golden Cat, camera trap, Catopuma temminckii, Clouded Leopard, competitive exclusion, Leopard Cat, Marbled Cat, Neofelis
nebulosa, Pardofelis marmorata, Prionailurus bengalensis, standard four.
Abbreviations: CF - community forest, CL - Clouded Leopard, GC - Golden Cat, IUCN - International Union for Conservation of Nature, LC - Leopard
Cat, MC - Marbled Cat, NP - national park, TR - tiger reserve, VCR - village community reserve, WS - wildlife sanctuary.

DOI: https://doi.org/10.11609/jott.4662.11.4.13432-13447 | ZooBank: urn:lsid:zoobank.org:pub:2D56F7F4-0B18-4765-B4A3-C08232E89BD2

Editor: Angie Appel, Wild Cat Network, Germany. Date of publication: 12 March 2019 (online & print)
Manuscript details: #4662 | Received 30 October 2018 | Final received 07 January 2019 | Finally accepted 27 February 2019
Citation: Mukherjee, S., P. Singh, A.P. Silva, C. Ri, K. Kakati, B. Borah, T. Tapi, S. Kadur, P. Choudhary, S. Srikant, S. Nadig, R. Navya, M. Björklund & U. Ramakrishnan
(2019). Activity patterns of the small and medium felid (Mammalia: Carnivora: Felidae) guild in northeastern India. Journal of Threatened Taxa 11(4): 13432–13447;
https://doi.org/10.11609/jott.4662.11.4.13432-13447
Copyright: © Mukherjee et al. 2019. Creative Commons Attribution 4.0 International License. JoTT allows unrestricted use, reproduction, and distribution of this
article in any medium by adequate credit to the author(s) and the source of publication.
Competing interests: The authors declare no competing interests.
For Funding, Author details, Author contribution & Acknowledgements see end of this article.

13432
Activity patterns of small and medium felids Mukherjee et al.

INTRODUCTION al. 2013; Xiong et al. 2017). On the other hand, small
wild cats like Leopard Cat and Jungle Cat are largely
India’s geographic location at the confluence of dependent on murid rodents, and to a smaller extent
major biogeographic realms has contributed towards on birds, amphibians, and reptiles to meet their energy
its extraordinary biodiversity (Mani 1974). The family requirements (Mukherjee et al. 2004; Shehzad et al.
Felidae is particularly well-represented in India with 15 2012; Xiong et al. 2017).
extant species, constituting around 37% of the global Though similar body sizes and diet could impose
felid diversity (Kitchener et al. 2017). While speciation potential inter-species competition, some species are
seems to have played a major role in structuring the mainly found in specific habitats. For instance, the Fishing
felid assemblage of South America, the Indian felid Cat inhabits mainly the lowland swamps and wetlands,
assemblage resulted from a series of colonization events the Jungle Cat occurs in open habitat and shrubland
(Johnson et al. 2006). Consequently, of the 14 felid areas, and the Snow Leopard is restricted to higher
genera recognized by Kitchener et al. (2017), nine occur elevations (Nowell & Jackson 1996). The remaining six
in India. wild cat species tend to co-occur in the northeastern
Due to different environment conditions and species forests. In similar habitats across southeastern Asia, the
adaptations, felids are not homogenously distributed assemblage of Leopard Cat, Marbled Cat, Asiatic Golden
throughout the country. The highest species richness Cat, and Clouded Leopard is common and was named as
(nine species) is observed in the semi-arid and arid the standard four by Duckworth et al. (2014).
region to the west due to colonization from Middle- The similar body size of Leopard Cat and Marbled Cat
Eastern species, and in the northeastern forests due raises interesting questions related to co-occurrence.
to the colonization of species from southeastern Asia Asynchronous activity patterns can potentially explain
(Johnson et al. 2006). Obligate carnivory in cats along co-occurrence, and Marbled Cat and Leopard Cat were
with very similar physiologies and overall morphologies found to have low activity overlap (Lynam et al. 2013;
shared across the family raise interesting questions Singh & Macdonald 2017; Hearn et al. 2018), thereby
related to sympatry. Several studies throughout the supporting this argument. Apart from competition,
globe addressed co-occurrence patterns in the Felidae predation from a larger cat could also determine
and in most studies inferences are drawn around temporal patterns of activity and it would be expected
possible competitive interactions and avoidance thereof. for specific pairs of cats, Clouded Leopard­-Golden Cat,
Segregation over body size, diet, space, and time are Golden Cat-Marbled Cat, and Golden Cat-Leopard Cat, to
recognized as the major facilitators of co-existence in have low temporal activity overlaps. Clouded Leopard-
carnivores, including wild cats (Caro & Stoner 2003; Golden Cat and Golden Cat-Marbled Cat, however, were
Morales & Giannini 2010; Sunarto et al. 2015; Cruz et al. found to have similar activity periods (Azlan & Sharma
2018; Hearn et al. 2018). 2006; Lynam et al. 2013; Singh & Macdonald 2017).
In northeastern India, felid body mass spans almost Activity patterns can also vary regionally, but current
two orders of magnitude. The smallest are Marbled Cat information on the activity of the standard four comes
and Leopard Cat with body mass ranging from 2.4–3.7 mainly from southeastern Asia with very few studies
kg and 2.7–3.6 kg, respectively (Pocock 1939; Sunquist from India (e.g., Singh & Macdonald 2017).
& Sunquist 2002), while the largest, the Tiger Panthera In this collaborative study that collates activity
tigris, weighs up to 260kg in the Indian subcontinent (all information from ten independent camera trapping
of these were Chitwan tigers immobilised by Smith et al. studies in northeastern India, we address a knowledge
1983). The other species include the Leopard P. pardus gap pertaining to the activity of the standard four in India.
(31–63 kg), Snow Leopard P. uncia (30–50 kg), Clouded Northeastern India comprises of the northwestern edge
Leopard (18–20 kg), Asiatic Golden Cat (9–16 kg), of Clouded Leopard, Asiatic Golden Cat, and Marbled
Fishing Cat Prionailurus viverrinus (5–16 kg), and Jungle Cat distribution, and hence is of geographic significance
Cat Felis chaus (2.3–8.6 kg) (Pocock 1939; Sunquist & to these species (Grassman et al. 2016; McCarthy et al.
Sunquist 2002; Athreya & Belsare 2006; Hunter 2015). 2016; Ross et al. 2016). We estimated activity patterns
The larger species such as the Tiger tend to prey on of small to medium felids (<20kg average body mass)
large or medium-sized ungulates (Andheria et al. 2007; and compared them to patterns found throughout
Lyngdoh et al. 2014), while the Clouded Leopard and southeastern Asia. In addition, we explored potential
Asiatic Golden Cat are likely to focus on small ungulates, activity pattern variation at different elevations, where
primates, larger rodent species, and pheasants (Ross et larger cats tend to be absent or less abundant, and

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Activity patterns of small and medium felids Mukherjee et al.

under different management regimes associated with Management regimes range from legally delineated
different levels of human disturbance. NPs and WSs to locally-managed CFs and VCRs. While
Arunachal Pradesh and part of Assam constitute a part
of the eastern Himalayan biodiversity hotspot, the study
STUDY AREAS locations in Nagaland, Meghalaya, part of Assam, and
Mizoram fall within the Indo-Burma biodiversity hotspot
The northeastern region of India encompasses a (Mittermeier et al. 2004).
vast gradient of elevations (<50–>8,000 m), climate Arunachal Pradesh: This study incorporates data
conditions, and vegetation associations. As a result, the collected between May 2013 and March 2017 from
region provides a wide range of habitats and species Pakke-Eaglenest landscape and Talle Valley WS. The
adapted to each of these habitats. Pakke-Eaglenest landscape comprises of Pakke TR, which
This study incorporates camera trap data obtained covers an area of 862km2, Eaglenest WS spread over
from 10 study sites spread over five states of 217km2, and the recently proposed Singchung-Bugun
northeastern India, which include Assam, Arunachal VCR. While Pakke TR has an elevation gradient of ≥100m
Pradesh, Nagaland, Meghalaya, and Mizoram (Table to 1,500m, Eaglenest WS is located at a higher elevation,
1; Fig. 1). The predominant vegetation across most of between 500m and 3,250m. Talle Valley covers an area
these sites is tropical evergreen, subtropical evergreen, of 337km2 in the Lower Subansiri Valley, northeast of
tropical semi-evergreen, or moist deciduous forest the Pakke-Eaglenest landscape. The elevation gradient
(Champion & Seth 1968). Temperate broad-leaved of the reserve is between 1,500m and 2,825m. Though
and temperate conifers occur in the higher elevations all study sites within this landscape support most
(2,000–4,000 m) in Arunachal Pradesh, within the study mammalian species typical of this landscape, the larger
sites included in this paper (Kaul & Haridasan 1987). cats, Tiger and Leopard, are found largely in Pakke TR.

Figure 1. Sampling sites in northeastern India.

13434 Journal of Threatened Taxa | www.threatenedtaxa.org | 12 March 2019 | 11(4): 13432–13447

Activity patterns of small and medium felids Mukherjee et al.

Table 1. Sampling areas and results of camera trapping used in this study.

Approximate
Min-Max
number of Small felid species (<25kg
State Sampling areas Trap nights Trapping period elevation (m) of
camera trap average body mass) detected
records
locations
Clouded Leopard (n=40), Asiatic
Arunachal Nov 2015 to Mar Golden Cat (n=3), Marbled Cat
Pakke TR 213 ~10,260 123–732
Pradesh 2017 (n=40), Leopard Cat (n=394),
Jungle Cat (n=4), Tiger, Leopard
Clouded Leopard (n=4), Asiatic
Arunachal May 2013 to Apr
Eaglenest WS 250 8,044 1,640–3,220 Golden Cat (n=28), Marbled Cat
Pradesh 2016
(n=27), Leopard Cat (n=59)
Clouded Leopard (n=3), Asiatic
Arunachal Singchung- May 2013 to Mar
100 1,155+ 1,459–3,217 Golden Cat (n=22), Marbled Cat
Pradesh Bugun VCR 2018
(n=1), Leopard Cat (n=43)
Arunachal Mar 2016 to May Asiatic Golden Cat (n=8),
Talle-Valley WS 22 1,063 2,352–2,446
Pradesh 2016 Leopard Cat (n=14), Tiger
May 2015 to Jul
Nagaland Intanki NP 23 826 304–532 Leopard Cat (n=14)
2015
Nagaland Shatuya CF 5 ≤50 Jun 2011 ~1,300 Leopard Cat (n=1)
Clouded Leopard (n=4), Asiatic
Mar 2015 to May
Meghalaya Nongkhyllem NP 31 1,094 378–863 Golden Cat (n=2), Leopard Cat
2015
(n=12)
Balpakram- Clouded Leopard (n=4), Marbled
Jan 2013 to Jun
Meghalaya Baghmara 425 3,857 104–856 Cat (n=2), Leopard Cat (n=40),
2015
landscape Jungle Cat (n=1), Leopard
Nov 2017 to Dec Clouded Leopard (n=6), Marbled
Mizoram Dampa TR 9 ~150 567–826
2017 Cat (n=1), Leopard Cat (n=2)
Dibru-Saikhowa May 2016 to Jul
Assam 27 1,065 119–123 Leopard Cat (n=9), Leopard
NP 2016
Total 10 sites ~1105 ~27,567 (n=788)

Tiger presence was recorded in Talle Valley (André P. Silva is impacted by insurgency and encroachment (Longchar
unpublished data), while there are no recent reports of 2013). Leopard was the only big cat detected in the area.
the big cats from Eaglenest WS or Singchung-Bugun VCR. Meghalaya: We used data collected between 2013
Assam: Located in Upper Assam, Dibru-Saikhowa NP and 2015 in the Balpakram-Baghmara landscape. This
along with the adjoining reserved forest cover an area of site is in the South Garo Hills District of Meghalaya, in
340km2 in Tinsukia District. To its north, Brahmaputra proximity to the border with Bangladesh to the south,
and Lohit rivers form the park boundary, while to the covering ~600km2. While Balpakram NP covers an area
south Dibru River demarcates the reserve. Elevation of of 220km2, Baghmara Reserve Forest covers ~45km2.
the area is generally below 150m. Data were collected The two protected areas are disjunct, with community-
in this study site between May and July 2016. The owned land called ‘Aking’ separating them. Data
park faces threats from illegal logging and poaching, collected in Nongkhyllem WS between March and May
erosion by the Brahmaputra River, and military activities 2015 was also used in this study. Nongkhyllem is located
(Sekhsaria 2012). Among big cats, only Leopard was in Ri Bhoi District, close to the border with Assam. It
detected at this site. covers an area of 29km2 with an elevation ranging from
Nagaland: Data used in this study were derived 200m to 965m. Among big cats, Leopard was detected
mainly from a single study conducted in Intanki NP in the Baghmara-Balpakram landscape.
between May and July 2015. A single data point was also Mizoram: Data collected in November–December
incorporated from a survey conducted in 2011 in Shatuya 2017 from a limited area, covering ~20km2 in Dampa
CF in Phek District of eastern Nagaland (Grewal et al. TR was used in this study. Located in western Mizoram,
2012), along the international border with Myanmar. Dampa TR is contiguous to the west with the Chittagong
Intanki NP covers an area of ~200km2, located in Peren Hill tract region of Bangladesh. The protected area is
District along the Dhansiri River, and is contiguous with highly undulating with elevation ranging between 150m
the Dhansiri Reserve Forest in Assam. It is a low-lying and 1,100m. A recent study by Singh & Macdonald
protected area with elevation ranging between 200m (2017) reports the presence of the standard four felid
and 682m. The reserve faces high hunting pressure and species in the region, although no Leopard or Tiger was

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Activity patterns of small and medium felids Mukherjee et al.

1
recorded during the same period. (2009) as Δ̂1 = ʃ min {f̂(t), ĝ(t)}dt. To calculate 95% CI,
0
we generated 10,000 bootstraps of our samples and
present bootstrap bias-corrected percentiles.
MATERIALS AND METHODS To explore the variation of activity patterns, we
compared the extent of overlap in the same species
Camera trapping reported in other studies across southeastern Asia that
We used data on small and medium-sized sympatric also used the kernel density analysis. In addition, we
felids found in northeastern India, obtained via camera analysed activity patterns across an elevation gradient
traps. This data was assembled from multiple studies and management regimes for all species, depending on
(Table 1) that used camera traps as a tool to assess the data availability. For elevation comparisons, we divided
distribution and population status of the mammalian our samples into two categories, ≤1,500m and ≥1,500m,
fauna of the region. The general study design followed given variations in habitat and vegetation types across
by all contributors to this study involved placement these categories and availability of data. We had ≥20
of camera traps 20–50 cm above ground level, along samples for Leopard Cat from several different sites
animal trails, river beds, or ridge-lines, which are (n=4); hence, we compared activity cycles for the same
expected to support the highest animal movement species across different study sites.
in forested landscapes. We chose this height above Since Marbled Cat and Leopard Cat are very similar
ground to increase the detectability of small mammals in body size, we attempted to examine if competition
by the camera traps. Camera traps used in all studies influences their activity patterns. To determine this, we
were equipped with in-built wide lenses and placed analysed activity patterns for the two cats at the point
with the aim of capturing all felid species found in the location at every camera trap location, a) where both
landscape. Camera traps were regularly checked to species were recorded, b) where only one of the species
ensure uninterrupted functioning and continuous data- was recorded, and c) overlap of the activity results for
collection in 24-hour cycles. All images obtained the each of the two species, i.e., Marbled Cat captured
contained date and timestamps. with Leopard Cat at the same location and Marbled Cat
captured individually, and the same for Leopard Cat.
Activity patterns
To ensure the independence of camera trap images,
we considered consecutive records of the same species RESULTS
at the same camera trap location within an interval of
≤30min as a single capture event (Linkie & Ridout 2011; We obtained 788 records of five felid species, namely,
Sunarto et al. 2015; Singh & Macdonald 2017). If two Clouded Leopard (n=61), Asiatic Golden Cat (n=63),
consecutive images, however, were triggered by the Marbled Cat (n=71), Leopard Cat (n=588), and Jungle
same species within the 30min-period, but by differently Cat (n=5) from >27,500 trap nights in 10 sampling areas
marked individuals, we recognized each photographed across northeastern India (Images 1–4). Details about
individual as a separate event. Similarly, if a single image results from the different study areas are provided in
captured multiple individuals of a single or different cat Table 1.
species, we treated each individual as a separate event. Of these, the highest records (61%) were from Pakke
To visualize activity patterns for each species and to TR, since annual camera trapping is conducted in the
estimate activity overlaps, we used a method designed reserve, primarily to assess the Tiger population as part
by Ridout & Linkie (2009), which accounts for the of the All India Tiger Monitoring exercise followed by
circular nature of time by using a non-parametric kernel Eaglenest WS and the contiguous Singchung-Bugun VCR
density estimation approach. We used package Overlap (24%). Clouded Leopard showed weak activity during
(Meredith & Ridout 2018) available in R (R Core Team the day and was most active during dawn and dusk (Fig.
2014), to analyse species-specific activity patterns and 2a). Marbled Cat (Fig. 2b) and Asiatic Golden Cat (Fig.
overlaps after pooling data from all sites. We used a 2c) were almost entirely diurnal, while Leopard Cat (Fig.
smoothing parameter of 1.2 for sample sizes >75, and 2d) was predominantly nocturnal. The highest overlap
a smoothing parameter of 0.8 for sample sizes <50, to in diel activity patterns was seen between Marbled Cat
smoothen spikes in the graphs as recommended by and Asiatic Golden Cat with Δ̂1 of 0.82 (0.71–0.92), and
Meredith & Ridout (2018) to generate the coefficient of Clouded Leopard and Leopard Cat with of 0.79 (0.71–
overlap (Δ̂1). Δ̂1 was defined following Ridout & Linkie 0.87; Fig. 3). The lowest Δ̂1 was observed between

13436 Journal of Threatened Taxa | www.threatenedtaxa.org | 12 March 2019 | 11(4): 13432–13447

Activity patterns of small and medium felids Mukherjee et al.

Figure 2. Kernel density plots of activity patterns of four felids in northeastern India.

Leopard Cat and Marbled Cat (0.32; 0.19–0.35), and study between Leopard Cat and Marbled Cat (0.32;
Leopard Cat and Asiatic Golden Cat (0.38; 0.24–0.43; 0.19–0.35) is within that observed in other studies.
Fig. 3). At the point scale, Marbled Cat showed a slight
Comparison of activity overlap between pairs of cats variation in activity when captured on locations with
from several studies (Lynam et al. 2013; Sunarto et al. (n=34) and without (n=37) Leopard Cat captures, with
2015; Singh & Macdonald 2017; Fig. 3) reveals similar Δ̂1 of 0.81 (0.79–0.91), while Leopard Cat activity did
overall patterns, with the highest Δ̂1 between Clouded not change in the presence (n=85) or absence (n=503)
Leopard and Leopard Cat (0.73–0.90) and lowest of Marbled Cat (Fig. 4). With respect to management
between Leopard Cat and Marbled Cat (0.26–0.54). In regimes, 90% of our image-captures were obtained
the current study, however, Clouded Leopard and Asiatic from protected areas and the remaining 10% from
Golden Cat were spatially separated and hence could not community forests. We could not compare differences
be compared for overlap. The detected Δ̂1 in the current in activity patterns for Clouded Leopard (n=3, OPA)

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Activity patterns of small and medium felids Mukherjee et al.

Figure 3. Comparison of activity overlaps for the standard four felids across their geographic range, based on information obtained from other
comparable studies (Lynam et al. 2013; Sunarto et al. 2015; Singh & Macdonald 2017; Hearn et al. 2018). Asiatic Golden Cat does not occur
in Borneo and, therefore, activity estimates for the species are not present in Hearn et al. (2018). Estimates for overlaps of Asiatic Golden Cat
and Leopard Cat are not provided in Singh & Macdonald (2017).

and Marbled Cat (n=1, OPA) across protected and non- DISCUSSION
protected landscapes due to limited data, while Asiatic
Golden Cat showed the same activity pattern across For the first time, we estimated activity patterns of
differing management regimes (Fig. SD1.3). In the small wild cats across northeastern India complementing
case of altitude, we had two categories, ≥1,500m (25% the existing studies in southeastern Asia, allowing future
samples) and ≤1,500m (75% samples). Leopard Cat was analyses on the variation of activity patterns across the
the only felid captured across all sites and accounted for species range. Interestingly, we simultaneously unveiled
75% of total captures. Leopard Cat activity remained new information on the species distribution across the
mostly unchanged irrespective of altitude category (Fig. northeastern Indian landscape. Based on information
SD1.1), study site (Fig. SD1.2), or management regime used in this study, an assemblage of the four felids
(Fig. SD1.3). The Δ̂1 of 0.76 for Marbled Cat across the Clouded Leopard, Asiatic Golden Cat, Leopard Cat, and
two elevation categories, however, indicates a slight Marbled Cat occurred in three of the 10 study sites,
variation in its activity pattern (Fig. SD1.1). Above namely, Pakke TR, Eaglenest WS, and Singchung-Vugun
1,500m, Marbled Cat showed a decline in activity post VCR. Marbled Cat had very low captures (≤2; Singchung-
a pre-noon peak. Bugun VCR, Baghmara-Balpakram landscape, Dampa) or
was not captured at several sites (n=5), except for Pakke
TR and Eaglenest WS, while the Clouded Leopard had
very low captures in all but Pakke TR. This could be an
indication of the sensitivity of these species to habitat

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Activity patterns of small and medium felids Mukherjee et al.

Figure 4. Point scale analyses of activity patterns of Leopard Cat (A) and Marbled Cat (B): a - when both were photographed at the same
location (Marbled Cat, n=34; Leopard Cat, n=85); b - when only one species was photographed at a location (Leopard Cat, n=503; Marbled Cat,
n=37); c - overlap of activity of each species with and without the presence of the other.

Image 1. Asiatic Golden Cat Catopuma temminckii (Eaglenest Wildlife Sanctuary 25 March 2015). © SACON Eaglenest Small Cat Team.

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Activity patterns of small and medium felids Mukherjee et al.

change and forest loss, both being species largely competition for resources and species sorting over a
associated with primary closed-canopy forests or, more gradient of resources (Leibold et al. 2004). Competition
likely, of unequal capture efforts. The effort in Dampa TR occurs when species that require similar resources
and Shatuya CF may not have been adequate in terms of and function in an ecologically similar manner meet
camera placements as well as the number of trap nights at range boundaries and overlap zones, the outcomes
at each location to record Marbled Cat and Clouded of which are either competitive exclusion or co-
Leopard, which are intrinsically rare and arboreal. occurrence through adjustments in specific character
Our data capture one aspect of the ecology of an traits. Co-occurrence in the face of stiff competition for
assemblage of felids, and despite the relatively large limited resources is often attained through character
dataset for a group of little known, rare, and elusive displacement where species shift some characteristics of
species, our analysis remains preliminary. Although the their niches to fit into sympatry (Brown & Wilson 1956;
results of our work are not very different from other Schoener 1974; Dayan et al. 1990; Losos 2000). On the
studies on the same assemblage of cats, we suggested other hand, species sorting occurs over a gradient of
an alternative explanation for the temporal segregation resources where each species is adapted to procuring
between Marbled Cat and Leopard Cat. This needs a specific set of resources (Leibold et al. 2004). In this
to be tested through further information on diet and scenario, species from an assemblage would not require
spatial use of habitats of these two cats. Several camera shifting of their niche characters and would fit in due
trapping studies by various groups were conducted in to pre-existing differences in their requirements and
India, mostly focusing on the large cat, and by-catch adaptations.
data on the smaller cats often remains unanalyzed In the case of the similar-sized Marbled Cat and
(e.g., Borah et al. 2014). Moreover, studies at individual Leopard Cat, competition is expected, especially in
sites are often limited in time and areas covered and prime habitats, and their diel activity patterns would
hence are data-poor. Joint analysis of data through lead to that conclusion since they are largely segregated
collaborative ventures such as ours could provide over time. This contrasting pattern of the nocturnal
important information on basic ecological aspects such Leopard Cat and diurnal Marbled Cat is consistent
as activity patterns and habitat use of this poorly studied almost throughout their range and is often attributed to
group. competition due to the significant overlap in body size
Given the range and overlap of body sizes between (Gumal et al. 2014; Pusparini et al. 2014; McCarthy et
pairs of cats, we expected temporal segregation in al. 2015; Singh & Macdonald 2017; Hearn et al. 2018).
activity patterns between Asiatic Golden Cat and Clouded The Marbled Cat, though sometimes detected by on-
Leopard, and Leopard Cat and Marbled Cat. Consistent ground camera traps, is morphologically suited for an
with our expectations, Leopard Cat and Marbled Cat arboreal life, as indicated by its very long tail (longer
were mainly nocturnal and diurnal, respectively. In than its body) and perhaps also largely restricted to
our study, the Clouded Leopard and Asiatic Golden prime, closed canopy forests (Pocock 1939; Sunquist
Cat captures were spatially separated. Most Clouded & Sunquist 2002; Gumal et al. 2014; Hunter 2015,
Leopard captures were in Pakke TR and Asiatic Golden Mukherjee et al. 2016). Northeastern India has an
Cat captures in the Eaglenest landscape, and hence impressive diversity of Sciuridae as well (Pocock 1939),
were not compared for their activity overlaps. Individual and they are largely diurnal and arboreal. Therefore, the
species activity patterns of the standard four over Marbled Cat’s diurnal and arboreal activities could be a
several studies, however, are in consensus with some result of targeting arboreal prey (Hearn et al. 2018) and
exceptions. For example, Gumal et al. (2014) report not necessarily related to the presence of the Leopard
crepuscular activity of Asiatic Golden Cat and nocturnal Cat. Unfortunately, there is limited and incidental data
activity of Clouded Leopard in peninsular Malaysia. In on Marbled Cat diet. Borries et al. (2014) account of one
contrast, Zaw et al. (2014) report diurnal and nocturnal observation of a Marbled Cat attempting predation on
activity of Clouded Leopard and cathemeral activity of a juvenile Phayre’s Leaf Monkey Trachypithecus phayrei.
Asiatic Golden Cat in Myanmar. In northeastern India, It was not clear, however, whether the attempt was
we found Clouded Leopard to be largely crepuscular and made on the ground or in the canopy. Davis (1962 as
Asiatic Golden Cat diurnal, in accordance with results cited in Hearn et al. 2018) reported a species of Rattus
from Dampa TR reported by Singh & Macdonald (2017). in the stomach of a Marbled Cat. On the other hand,
The pattern of species segregation over space, time, studies on Leopard Cat diet across its global range show
or diet can be a result of various processes including a very strong dependence on murid rodents that are

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Activity patterns of small and medium felids Mukherjee et al.

Image 2. Clouded Leopard Neofelis nebulosa (Pakke Tiger Reserve, 19 December 2015). © A.P. Silva, S. Nadig & R. Navya.

Image 3. Prionailurus bengalensis (Pakke Tiger Reserve, 9 December 2015). © A.P. Silva, S. Nadig & R. Navya.

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Activity patterns of small and medium felids Mukherjee et al.

Image 4. Marbled Cat Pardofelis marmorata (Pakke Tiger Reserve, 18 December 2015). © A.P. Silva, S. Nadig & R. Navya.

largely nocturnal (Shehzad et al. 2012; Lorica & Heaney Predation from larger cats could also play a role in
2013; Mukherjee et al. 2016; Xiong et al. 2017). Future determining activity patterns and habitat use. The
studies on Marbled Cat diet are therefore a priority to Leopard appears to be absent in Eaglenest WS and
test our inference. Singchung-Bugun VCR while the Clouded Leopard was
The spatial scale of focus also matters since sympatry recorded on rare occasions by Velho et al. (2015) and
could occur at regional or local scales (Palomares et in the current study. This could be an explanation for
al. 2016). One way to test for species-sorting over the highest number of Golden Cat captures in this study
competition for explaining co-existence in these pairs being from Eaglenest WS and the adjacent Singchung-
would be to look at diel activity patterns and habitat use Bugun VCR.
in areas where each of the species occurs in allopatry This, however, is speculative and needs to be tested.
and to compare it to areas of co-occurrence to look for The combined impact of several processes acting at
character displacement and release. This can be done at different spatial scales is likely to be responsible for
various spatial scales, e.g., the point scale of camera trap structuring assemblages (Leibold et al. 2004).
locations, the grid scale in studies that use grids within
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patterns and temporal avoidance by prey in response to Sunda Funding: Science and Engineering Research Board of the Department of Science
Clouded Leopard predation risk. Journal of Zoology 290(2): 96–106; and Technology, Government of India (grant nos.: SERB/F/4937/2013-14,
https://doi.org/10.1111/jzo.12018 SERB/F/2464/2012-13 & SERB/F/5921/2014-15), Portuguese Foundation for
Science and Technology (FCT) (SFRH/BD/91190/2012), Helge Ax:son Johnson
Ross, J., J. Brodie, S. Cheyne, A. Datta, A. Hearn, B. Loken, A.
Foundation, Stiftelsen För Zoologisk Forskning, Stiftelsen Lars Hiertas Minne,
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(2016). Pardofelis marmorata. In: The IUCN Red List of Threatened
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https://doi.org/10.2305/IUCN.UK.2016-1.RLTS.T16218A97164299. Centre for Ornithology and Natural History, Coimbatore, India. Priya Singh is an
en independent researcher and member of Researchers for Wildlife Conservation
Schoener, T.W. (1974). Resource partitioning in ecological (RWC), India. Andre Pinto Silva is currently pursuing his Ph.D. at the Department
of Animal Ecology, Uppsala University, Sweden. Chandan Ri is a Research Officer
communities. Science 185(4145): 27–39; https://doi.org/10.1126/
in Pakke Tiger Reserve, Arunachal Pradesh, India. Kashmira Kakati is a wildlife
science.185.4145.27 biologist and environmental activist working the forests of Northeast India. Binod
Sekhsaria, P. (2012). Report on the State of Wildlife in northeast Borah is currently pursuing a doctoral degree in the Utah State University, USA.
India 1996–2011: A Compilation of News from the Protected Area Tana Tapi is Field Director of Pakke Tiger Reserve, Arunachal Pradesh, India.
Update. Foundation for Ecological Security, 293pp. Sandesh Kadur is a Wildlife Film Maker and Conservation Photographer from
Shehzad, W., T. Riaz, M.A. Nawaz, C. Miquel & C. Poillot (2012). Bengaluru, India. Prafull Choudhary is a Field Officer at the Wildlife Trust of
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Institute of India. Surabhi Nadig is currently enrolled in a Masters Degree
application to the Leopard Cat (Prionailurus bengalensis) in Pakistan.
program in Creative Sustainability at Aalto University, Finland. R. Navya is an
Molecular Ecology 21(8): 1951–65; https://doi.org/10.1111/j.1365- independent researcher based in Mysore, India. Mats Björklund is a Professor
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jmammal/gyx104 Author contribution: Conceptualisation: Shomita Mukherjee and Priya Singh.
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Smith, J.L.D., M.E. Sunquist, K.M. Tamang & P.B. Rai (1983). A
Mukherjee and Priya Singh. Reviews and editing: Andre Pinto Silva and Mats
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Sunarto, S., M.J. Kelly, K. Parakkasi & M.B. Hutajulu (2015). Cat Acknowledgements: The Meghalaya study conducted by KK and team in
coexistence in central Sumatra: ecological characteristics, spatial 2013–2015 was funded by the Science and Engineering Research Board,
and temporal overlap, and implications for management. Journal of Department of Science and Technology, Government of India, through grant
Zoology 296(2): 104–115; https://doi.org/10.1111/jzo.12218 nos. SERB/F/2464/2012-13 & SERB/F/5921/2014-15; institutional and logistic
support was provided by Samrakshan Charitable Trust, Delhi. KK is grateful to
Sunquist, M. & F. Sunquist (2002). Wild Cats of the World. University
the Chief Wildlife Warden, Department of Environment & Forests, Meghalaya,
of Chicago Press, Chicago, Illinois, 462pp. for permissions and P.R. Marak, DFO, Balpakram National Park Division, and all
Velho, N., U. Srinivasan, P. Singh & W.F. Laurance (2015). Large other officials and staff for their constant support. APS, SN, and NR collected
mammal use of protected and community-managed lands in a data in Nongkhyllem and Talle Valley WSs, Dibru-Saikhowa and Intanki NP, and
biodiversity hotspot. Animal Conservation 19(2): 199–208; https:// Pakke TR in 2015–2016, for which APS acknowledges the role of the Forest
doi.org/10.1111/acv.12234 Departments of Meghalaya, Nagaland, Arunachal Pradesh, and Assam for critical
Xiong, M., D. Wang, H. Bu, X. Shao, D. Zhang, S. Li, R. Wang & M. assistance on logistics and safety during fieldwork. APS is deeply grateful to all
volunteers (Satya Sarathi, Aditya Singla, Shradha Rathod, Ram Alluri, and Renuka
Yao (2017).  Molecular dietary analysis of two sympatric felids
Rajiv) and assistants (Clement, Jim, Angao, Amos, Ezembe, Sabdaby, Partha,
in the Mountains of southwest China biodiversity hotspot and Tiasa, Laxman Sowtel, Sunil Pankcho, and Shankar). Advice on logistics and
conservation implications.  Scientific Reports 7: 41909; https://doi. fieldwork assistance by Sahila Kudalkar and Krishnapriya Tamma were crucial.
org/10.1038/srep41909 APS was supported by the Portuguese Foundation for Science and Technology
Zaw, T., T. Myint, S. Htun, S.H.T. Po, K.T. Latt, M. Maung & A.J. Lynam (FCT) (SFRH/BD/91190/2012), Helge Ax:son Johnson Foundation, Stiftelsen För
(2014). Status and distribution of smaller cats in Myanmar. Cat Zoologisk Forskning, Stiftelsen Lars Hiertas Minne, Liljewalchs Stipendiestiftelse,
News Special Issue 8: 24–30. Sederholms Stiftelse, and IDEA WILD. SM and PC are grateful to the Science
and Engineering Research Board of the Department of Science and Technology,
Government of India,  for funding the study (no. SERB/F/4937/2013-14), Dr.
Ramana Athreya, IISER, Pune, for collaborating on the project, logistic support
during field and laboratory work, and for discussions throughout the study, Dr.
P.V. Karunakaran, SACON, for discussions and support, the Chief Wildlife Warden
of Arunachal Pradesh for permits, Divisional Forest Officer of Eaglenest Wildlife
Sanctuary Millo Tasser (IFS), Indi Glo, Nima Glo, Puspalal Sharma, Raju Chetri,
Munna Tamang, Wangchu Phiyang, Tsamu Rai, Bikki Gindan Marphew, Bayung
Marphew, Samir Bora, Sange Khandu Phyang, Sonam Lama, Baba Tamang, Dipu
Chetri, Raju Sharma, Changma, Nima Tsering, Suntali Tsering Monpa, Montu
Kalita, Rohan Pandit, Satyem Longchar, and Nima Majrekar for assistance and
support during fieldwork and the Bugun and Sherdukpen communities for their
hospitality. PS and SK are grateful to Pu Rosiamma, Chief Wildlife Warden,
for permits to work in Dampa TR in 2017. They also acknowledge the role of
the Field Director, Dampa TR, and other forest personnel in assistance during
fieldwork. The authors are grateful to Umesh Srinivasan and Nandini Velho for
contributing data from Singchung-Bugun VCR and to the Bugun and Sherdukpen
communities for their assistance in collecting data and conservation initiatives
in the landscape.

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Activity patterns of small and medium felids Mukherjee et al.

Supplementary Figure 1.1. Marbled Cat and Leopard Cat activity across different elevation categories (Marbled Cat below 1,500m, n=43;
Marbled Cat above 1,500m, n=28; Leopard Cat below 1,500m, n=482; Leopard Cat above 1,500m, n=106).

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Activity patterns of small and medium felids Mukherjee et al.

Supplementary Figure 1.2. Leopard Cat activity across different study sites (Eaglenest WS, n=59; Singchung-Bugun VCR, n=43; Pakke TR,
n=394; Balpakram-Baghmara, n=40).

13446 Journal of Threatened Taxa | www.threatenedtaxa.org | 12 March 2019 | 11(4): 13432–13447

Activity patterns of small and medium felids Mukherjee et al.

Supplementary Figure 1.3. Leopard Cat and Asiatic Golden Cat activity across different protection regimes (Leopard Cat IPA, n=536; Leopard Cat
OPA, n=52; Asiatic Golden Cat IPA, n=41; Asiatic Golden Cat OPA, n=22). IPA - inside protected area; OPA - outside protected area.

Threatened Taxa

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Journal of Threatened Taxa | www.threatenedtaxa.org | 12 March 2019 | 11(4): 13448–13458
Article

Estimating Leopard Cat Prionailurus bengalensis Kerr, 1792

(Carnivora: Felidae) density in a degraded tropical forest
ISSN 0974-7907 (Online)
fragment in northeastern Thailand
ISSN 0974-7893 (Print)
Wyatt Joseph Petersen 1 , Tommaso Savini 2 , Robert Steinmetz 3 & Dusit Ngoprasert 4
PLATINUM
OPEN ACCESS Conservation Ecology Program, King Mongkut’s University of Technology Thonburi, 49 Soi Thientalay 25,
1,2,4

Bangkhuntien-Chaitalay Road, Thakham, Bangkhuntien, Bangkok 10150, Thailand.

3
World Wide Fund for Nature – Thailand, 87 Phaholyothin 5, Samsen Nai, Phayathai, Bangkok 10400, Thailand.
1
[email protected] (corresponding author), 2 [email protected], 3 [email protected],
4
[email protected]

Abstract: The Leopard Cat Prionailurus bengalensis is thought to be Asia’s most abundant wild cat. Yet, the species’ status is poorly known
due to a lack of rigorous population estimates. Based on the few studies available, Leopard Cats appear to be more abundant in degraded
forests, potentially due to increased prey availability. We conducted camera trap surveys, rodent live-trapping, and spatially-explicit
capture-recapture analyses to estimate the density of Leopard Cats within a degraded tropical forest fragment (148km2) in northeastern
Thailand. A total effort of 12,615 camera trap nights across 65km2 of trapping area resulted in at least 25 uniquely identified individuals.
Average rodent biomass (the main prey of Leopard Cats) was highest in the dry evergreen forest (469.0g/ha), followed by dry dipterocarp
forest (287.5g/ha) and reforested areas (174.2g/ha). Accordingly, Leopard Cat densities were highest in the dry evergreen forest with 21.42
individuals/100km2, followed by the reforested areas with 7.9 individuals/100km2. Only two detections came from the dry dipterocarp
forest despite both an extensive survey effort (4,069 trap nights) and available prey. Although the dipterocarp supported the second
highest average rodent biomass, it lacked a key prey species, Maxomys surifer, possibly explaining low encounter rates in that habitat.
Our results provide important baseline information concerning the population status of Leopard Cat in southeastern Asia. Further, our
findings corroborate with other studies that found a tolerance among Leopard Cats for degraded forests, highlighting the potential for
forest fragments to serve as long-term conservation areas for the species.

Keywords: Camera trapping, Least Concern, predator-prey, prey availability, rodent biomass, Sakaerat Biosphere Reserve, southeastern
Asia, spatially-explicit capture-recapture.

DOI: https://doi.org/10.11609/jott.4553.11.4.13448-13458 | ZooBank: urn:lsid:zoobank.org:pub:E977B1A0-5E5E-420E-80CD-063DB1BE1838

Editor: Shomita Mukherjee, SACON, Coimbatore, India. Date of publication: 12 March 2019 (online & print)
Manuscript details: #4553 | Received 04 September 2018 | Final received 05 March 2019 | Finally accepted 08 March 2019
Citation: Petersen, W.J., T. Savini, R. Steinmetz & D. Ngoprasert (2019). Estimating Leopard Cat Prionailurus bengalensis Kerr, 1792 (Carnivora: Felidae) density in a
degraded tropical forest fragment in northeastern Thailand. Journal of Threatened Taxa 11(4): 13448–13458; https://doi.org/10.11609/jott.4553.11.4.13448-13458
Copyright: © Petersen et al. 2019. Creative Commons Attribution 4.0 International License. JoTT allows unrestricted use, reproduction, and distribution of this article
in any medium by adequate credit to the author(s) and the source of publication.
Funding: Funding was provided by Thailand’s National Science and Technology Development Agency (grant number P-17-50347).
Competing interests: The authors declare no competing interests.
Author details: Wyatt J. Petersen is a wildlife biologist interested in the ecology and conservation of small and large carnivores. He has a bachelor’s degree from
Humboldt State University and a master’s from King Mongkut’s University of Technology Thonburi. Tommaso Savini has been working for the past 15-years on the
ecology of birds and mammals in the Southeast Asian tropical forests, looking at their behavioral ecology, landscape use, and conservation. Robert Steinmetz has
conducted ecological research, training, and site-based conservation projects in Southeast Asia for over 20 years. His interests include the ecology and conservation
of bears and other large mammals. He also works closely with local people, conducting social science research on conservation-related changes in human attitudes
and behavior. Dusit Ngoprasert is interested in conservation of bears and several small carnivore species. His interests include population estimation, habitat
selection, and distribution models. In addition, he conducts field research and teaches a method course and assists students/collaborators to deploy appropriate
methodologies and undertake data analysis.
Author contribution: WJP, DN, RS, TS conceived the ideas and designed field methodology; WJP collected the data; WJP, DN analyzed the data; WJP, DN, RS, TS
contributed critically to the drafts and all authors gave final approval for publication.
Acknowledgements: We thank the Thai Department of National Parks, Wildlife and Plant Conservation, National Research Council of Thailand, Thailand Institute
of Scientific and Technological Research, and reserve directors Taksin Artchawakom and Surachit Whitethorn for supporting and permitting our work. We would
like to thank George Gale and Ronglarp Sukmasuang for both their support and feedback, Katie Oliver and Somsak Poompuang for their assistance in the field, and
the two anonymous reviewers for their helpful comments.

13448
Leopard Cat in northeastern Thailand Petersen et al.

INTRODUCTION artificial open areas such as agriculture lands and human

settlements (Chen et al. 2016).
When confronted with a lack of rigorous population An association between Leopard Cats and degraded
estimates, status assessments such as the IUCN Red habitats may reflect the species diet, which consists
List of Threatened Species must rely on expert opinion predominantly of murid rodents (e.g., Yasuma 1981;
of trends in population abundance or geographic range Rabinowitz 1990; Grassman 1998; Grassman 2000;
(Mace et al. 2008). This, however, can be problematic Grassman et al. 2005; Rajaratnam et al. 2007). Indeed,
given both the subjective nature of expert opinion rodent biomass is often elevated in degraded habitats
(Regan et al. 2005) as well as the wide variation in owing to either increased resource availability (e.g.,
population dynamics and threats faced across a species invertebrates), reduced predation pressure, or both
range. The Leopard Cat Prionailurus bengalensis, for (Lambert et al. 2006; Wells et al. 2007; Pimsai et al. 2014).
example, is thought to be the most abundant small cat Yet, in most studies on Leopard Cat habitat use and diet,
species in Asia due to its wide distribution and supposed prey distribution and abundance were not measured,
tolerance towards human-modified landscapes (Nowell despite these apparent associations between predator,
& Jackson 1996; Macdonald et al. 2010; Ross et al. 2010). prey, and habitat. Of the studies that did measure prey
Yet, little is known regarding Leopard Cat population distribution and abundance, several indicated that
status in most parts of its range. Furthermore, recent Leopard Cats may preferentially select habitats based on
studies suggest that Leopard Cat populations, in at least the availability of a specific key prey species, rather than
some areas, are being adversely affected by habitat loss, overall prey abundance (Yasuma 1981; Rabinowitz 1990;
fragmentation, and poaching (e.g., Seto et al. 2012; Rajaratnam et al. 2007).
Coudrat et al. 2014a,b; Willcox et al. 2014). As a result, In this study, we estimated Leopard Cat density using
the Leopard Cat may be threatened in many areas at a spatially-explicit capture-recapture models applied to a
local scale, despite its global status on the IUCN Red List camera trapping dataset from a degraded tropical forest
as Least Concern (Ross et al. 2010). fragment in northeastern Thailand. Our study spanned
Camera trapping is an effective approach for periods of rodent abundance and scarcity in three forest
estimating the density of uniquely marked animals such types, allowing us to observe the response of Leopard
as Leopard Cats (Balme et al. 2009; Royle et al. 2013). Cats to both spatial and temporal changes in resource
However, despite widespread use of camera traps availability. We made two predictions. First, Leopard
throughout the Leopard Cat’s extensive geographic Cat density should be highest in the forest type with
range, estimates of density are available from just a few the highest average rodent biomass (g/ha), as forest
sites in India and Malaysian Borneo (Bashir et al. 2013; types with more food should support higher Leopard
Mohamed et al. 2013; Selvan et al. 2014; Srivathsa et Cat densities compared to forest types with less food.
al. 2015). This paucity of data may be due to multiple Second, Leopard Cat movements (sigma parameter)
reasons such as 1) a lack of interest or incentive among should be larger when and where resources are scarce,
researchers towards studying a species listed as Least compared to when and where they are abundant. This
Concern by the IUCN or 2) an inherent bias in survey pattern would reflect possible increases in ranging
effort towards other species and habitats not utilized by behaviour in response to low resource availability (Fuller
Leopard Cats. Moreover, Marshall et al. (2016) clearly & Sievert 2001). We then compared overall density at
demonstrated tropical research to be heavily biased our site to density estimates from other sites throughout
towards large, charismatic, and threatened species as the species range.
well as towards large, intact, primary forests. This bias is
relevant given that several studies from primary forests
found Leopard Cats to be among the least recorded STUDY AREA
Asian felids (e.g., Ross et al. 2010; McCarthy et al. 2015).
Furthermore, studies by Mohamed et al. (2013) and Established in 1977, the Sakaerat Biosphere Reserve
Srivathsa et al. (2015) found Leopard Cat densities to (hereafter Sakaerat) in northeastern Thailand (14.5100N
be higher in more degraded habitats, which tend to be & 101.9300E ) covers 148km2 of fragmented forest with
ignored by researchers. Despite their high tolerance an elevation range of 280­–762 m (Fig. 1). Historically,
for degraded habitats, telemetry studies suggest that the areas comprising Sakaerat’s present-day reserved
Leopard Cats are still forest-dependent to some extent forest underwent periods of extensive deforestation and
(Rajaratnam et al. 2007) and may even avoid large conversion to agriculture (the 1950s–1970s), followed

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Leopard Cat in northeastern Thailand Petersen et al.

Figure 1. Sakaerat Biosphere Reserve in Thailand, including three dominant habitat types, camera trap locations, rodent live-trapping
locations, and Highway 304.

by state-run reforestation efforts using non-native tree MATERIALS AND METHODS

species (1977 onwards). This process of deforestation
and reforestation resulted in the fragmentation and Camera trap surveys
isolation of Sakaerat from nearby forests (including Dong We deployed 60 camera traps (Scout Guard SG565)
Phayayen-Khao Yai forest complex) and greatly altered spaced 1km apart in three forest types (evergreen: 28
the reserve’s current forest structure, with no primary camera traps; dipterocarp: 16 camera traps; reforested
forest remaining (Kamo et al. 2002; TISTR 2018). areas: 16 camera traps) from mid-January 2017 through
Dominant forest types today include secondary dry October 2017 (Fig. 1). One camera trap was deployed
evergreen forest (hereafter evergreen; 54%), secondary per station. Camera traps were attached to trees
dry dipterocarp forest (hereafter dipterocarp; 11%), approximately 45cm above the ground and 3m from
and reforested areas (33%) which include secondary a target zone, which was lured with fish oil scent. We
evergreen regrowth and mixed acacia (Acacia spp.) and visited camera traps once per month to replace batteries,
eucalyptus (Eucalyptus spp.) forest plantations (Fig. 1; memory cards, and the scent lure. We calculated
Ashton et al. 2014). Sakaerat has a seasonal climate trapping effort by summing the number of trap nights
with a dry season starting from November to April and (24h-periods starting from 00.00h and ending at 23.59h),
a wet season from May to October. The average annual in which camera traps were both active and functioning.
precipitation is 1,071mm, while the average annual
temperature is 26.10C (TISTR 2018).

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Leopard Cat in northeastern Thailand Petersen et al.

Rodent biomass estimation activity center and the trap is zero) and sigma (σ; a
We sampled terrestrial rodents during four periods scaling parameter reflecting animal movement) were
(February–March; April–May; July–August; September– shared among sessions. To account for potential bias in
October) in 2017 at 15 sites (seven sites in evergreen, our small mammal capture-probabilities introduced by
four in dipterocarp, and four in reforested areas) using “trap-happy” or “trap-shy” individuals, we incorporated
Sherman live traps (7.62cmx8.89cmx22.86cm). At each various behavioral responses into our models (Otis
site, we arranged 25 traps on the ground in a 5x5 grid et al. 1978; Efford 2018). Models tested included a
with 20m spacing between traps and used peanut butter constant model [D(session) g0(.) σ(.)], learned response
as bait. Each session consisted of seven consecutive model [D(session) g0(b) σ(.); “trap-happy”], site-specific
trap nights at one site, with sites being visited once per learned response model [D(session) g0(bk) σ(.); trap-
two-month period (60 sessions total). Captured animals specific “trap-happy”], transient response model
were identified to species, weighed, uniquely marked [D(session) g0(B) σ(.); “trap-shy”], and site-specific
with an ear tag (mouse ear-tag – style 1005-1), and then transient response model [D(session) g0(Bk) σ(.); trap-
released at their capture sites. Rodent live-trapping specific “trap-shy”]. Session-specific density estimates
protocol was approved by King Mongkut’s University of were then derived using the best-supported model as
Technology Thonburi’s Animal Care and Use Committee determined by AICC scores and AICC weights (Burnham
and permitted by Thailand’s Department of National & Anderson 2002).
Parks, Wildlife and Plant Conservation. Session-specific density estimates were then
To estimate the biomass of rodents, we first multiplied by the session’s corresponding mean individual
estimated density (D) using the ‘secr’ package (Efford body mass to obtain session-specific estimates of rodent
2018) in program R (version 3.42; R Development Core biomass (Chutipong et al. 2017). Session-specific rodent
Team, 2016). Due to the limited number of captures biomass estimates were then assigned as a covariate to
and recaptures at some sites, we pooled all species and the nearest four camera trap stations within the same
sites from the same forest type and sampling period for habitat type for use in estimating Leopard Cat density.
analysis, with each site designated as a separate session.
Data from different two-month sampling periods were Leopard Cat density estimation
analyzed separately. Multi-session analyses were then Spatially-explicit capture-recapture analyses were
conducted whereby the capture parameters g0 (capture used to estimate Leopard Cat densities (Efford 2018)
probability when the distance between an animal’s based on each animal’s unique spot patterns observed

Image 1. A diurnal Leopard Cat Prionailurus bengalensis investigating the fish oil scent lure placed at the centre of a camera trap’s target zone
within the dry evergreen forest of Sakaerat Biosphere Reserve, northeastern Thailand.

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Leopard Cat in northeastern Thailand Petersen et al.

in camera trap images (Image 1). Because Leopard Cats trap-happy or trap-shy), using covariates “b”, “bk”, “B”,
are patterned asymmetrically, there is a possibility of and “Bk”. Sigma (σ) was additionally modelled by the
photographing what is known as “partial identities” or trap covariate “rodent biomass”.
individuals known from only a single flank. To include We performed our model selection in three steps
these “partial” individuals into the analyses, researchers using AICC scores and AICC weights to determine the best-
typically analyze left and right flanks separately or supported model (Burnham & Anderson 2002). First, we
discard the flank with the least detections (e.g., Wang & modelled all g0 covariates individually, while leaving D
Macdonald 2009; Kalle et al. 2011; Srivathsa et al. 2015). and σ constant. Second, any individual g0 covariates
These approaches, however, result in a loss of precision performing better than the constant model [g0(.)] were
and the potential introduction of bias (Meredith 2017). then modelled together using all possible combinations.
To avoid this, we used a new method that combines data The top-ranked covariate combinations (ΔAICC<6.0)
from both flanks into the same analysis, modelling each from this second step were then considered “candidate
flank as a separate ‘session’ and estimating shared values covariate combinations”. These two steps were then
for D, g0, and σ across sessions under the assumption repeated for the D and σ parameters. Finally, we
that the latent (“true”) density and capture probabilities modelled D, g0, and σ parameters using all combinations
of both flanks are equal (Meredith 2017). of D, g0, and σ “candidate covariate combinations”. Due
After identifying individuals, we generated capture to our limited sample size, we restricted each model to
histories using daily occasions starting from 00.00h and a maximum of two covariates. All models were fit using
ending at 23.59h. Capture histories were then split the full likelihood approach.
into different sessions based on the period of resource
availability (i.e., high or low rodent biomass), habitat
type (i.e., evergreen or reforested area), and flank (i.e., RESULTS
left or right). A mask (buffer=1500m; designated using
the ‘suggest.buffer’ function) was also applied around In 12,615 camera trap nights, we obtained 115
each station, limited by the study area’s boundary. We images from 59 detections of Leopard Cats, of which
then conducted multi-session analyses, comparing a 50 detections contained images that could be used
constant model [D(.) g0(.) σ(.)] to models where D, g0, to identify individuals (Table 1). Due to the species
and σ parameters varied by the season of resource small body size, we could not identify the sex of the
availability (high rodent biomass and low rodent individuals. During the period of low rodent biomass, 14
biomass; termed “season”) and habitat type (evergreen left-flank individuals (12 in evergreen, two in reforested)
and reforested area; termed “habitat”). In addition and 13 right-flank individuals (12 in evergreen, one in
to these session-covariates, we also modelled the g0 reforested) were detected. Ten left-flank individuals (five
parameter using the trap-covariates “rodent biomass” in evergreen and five in reforested) and 11 right-flank
which corresponds to the actual rodent biomass (g/ha) individuals (six in evergreen and five in reforested) were
from the nearest rodent live-trapping site (see Rodent detected during the period of high rodent biomass. Two
biomass estimation) and behavioral responses (e.g., left-flank individuals were detected in the dipterocarp

Table 1. Summary of Leopard Cat images by period of rodent biomass from Sakaerat Biosphere Reserve, northeastern Thailand. “Identifiable
detections” refers to the number of Leopard Cat detections that contain images that are usable for individual identification. “Partial individuals”
corresponds to the number of individuals that only had a single flank photographed (either right or left). “Complete individuals” corresponds
to the number of individuals that had both flanks photographed. Dry evergreen forest: DEF; reforested areas: RFA; dry dipterocarp forest: DDF.

Low rodent biomass High rodent biomass

Survey Information
Total DEF RFA DDF Total DEF RFA DDF
Period January–May 2017 June–October 2017
Camera stations 58 28 14 16 58 28 14 16
Survey area 65km2 34km2 15km2 16km2 65km2 34km2 15km2 16km2
Trap nights 7,193 3,726 1,341 2,126 5,422 2,136 1,349 1,937
Identifiable detections 25 22 3 0 25 13 10 2
Partial individuals (right-flank) 7 6 1 0 10 6 4 0
Partial individuals (left-flank) 7 5 2 0 11 5 4 2
Complete individuals (both flanks) 7 7 0 0 1 0 1 0

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Leopard Cat in northeastern Thailand Petersen et al.

during the period of high rodent biomass. Because the evergreen (mean=469.0 g/ha) throughout our study,
of our low sample size in the dipterocarp, we did not followed by the dipterocarp (mean=287.5g/ha) and
estimate Leopard Cat density in that habitat type. reforestation areas (mean=174.2 g/ha; Fig. 2). Seasonally,
overall average rodent biomass was 3.6 times higher
Rodent biomass during the rainy season (July–October; mean=527.5g/
In total, we conducted 10,500 small mammal trap ha) compared to the dry season (February–May;
nights, during which we captured 297 unique individuals mean=145.6g/ha). Overall rodent biomass peaked in
of at least six rodent species (Table 2). Average density the months of September–October (mean=763.8g/ha),
in the evergreen was highest in September–October especially in the evergreen (mean=995.0g/ha). April–
(mean=7.3 individuals/ha; range of site-specific May had the lowest overall rodent biomass (mean 91.8g/
densities: 5.3–10.5 individuals/ha) and lowest in ha), especially in the reforested areas (0 captures). As
April–May (mean=1.3 individuals/ha; range: 0.4–1.7 such, February–May was considered the period of low
individuals/ha). In the dipterocarp, average density was rodent biomass for all habitats and July–October was
highest in July–August (mean=13.1 individuals/ha; range considered the period of high rodent biomass (Fig. 2).
of site-specific densities: 0 individuals captured–38.6
individuals/ha) and lowest in April–May (mean=2.2 Leopard Cat density
individuals/ha; range: 0 individuals captured–3.4 Our two top-ranked models based on AICC and AICC
individuals/ha). In the reforested areas, the average weights included D(.) g0(b) σ(habitat) (43% AICCwi;
density was highest in September–October (mean=5.2 Table 3) and D(habitat) g0(b) σ(.) (20% AICCwi; Table
individuals/ha; range of site-specific densities: 3.3–6.6 3). According to D(.) g0(b) σ(habitat), the Leopard Cat
individuals/ha) and lowest in April–May (0 individuals density within Sakaerat (excluding dipterocarp) was 17.7
captured). individuals/100km2 (SE 3.9; 95% CI 11.5–27.2; Table
Average rodent biomass was consistently highest in 4). The movement parameter, sigma (σ), was 480.2m

Figure 2. Rodent biomass by habitat and bimonthly period. The period of high rodent biomass corresponds with July–August and September–
October, while the period of low rodent biomass corresponds with February–March and April–May.

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Leopard Cat in northeastern Thailand Petersen et al.

Table 2. Summary of small mammal live-trapping results including trap nights by habitat, the number of unique individuals captured by both
habitat and species, and the average individual body mass by species. Live-trapping data from Sakaerat Biosphere Reserve, northeastern
Thailand, 2017.

Survey information Overall Dry dipterocarp Dry evergreen Reforested areas Average mass

Trap nights 10,500 2,800 4,900 2,800 -

Total capture events 297 137 133 27 -

Maxomys surifer 162 6 133 23 132.3g/indiv SE 2.0

Mus spp. 118 116 0 2 17.4g/indiv SE 0.5

Tupaia belangeri 10 10 0 0 177.6g/indiv SE 14.6

Menetes bermorei 4 4 0 0 198.3g/indiv SE 24.0

Crocidura attenuata 2 0 0 2 9.5g/indiv SE 0.5

Rattus rattus 1 1 0 0 79g/indiv

Table 3. Model selection for Leopard Cat density using spatially-explicit capture-recapture. ‘K’ represents the number of estimated parameters.
AICC, ΔAICC, and AICC weights ‘wi’ provide a measure of relative support for each model. Only models with a cumulative wi≤0.95 are included.
For all models, see Appendix B.

Models K AICC ΔAICC wi

D(.) g0(b) sigma(habitat) 5 961.17 0.00 0.43

D(habitat) g0(b) sigma(.) 5 962.69 1.52 0.20

D(.) g0(.) sigma(habitat) 4 963.66 2.49 0.12

D(.) g0(habitat) sigma(habitat) 5 964.69 3.52 0.07

D(habitat) g0(.) sigma(habitat) 5 965.94 4.77 0.04

D(habitat) g0(.) sigma(.) 4 966.12 4.94 0.04

D(.) g0(habitat + bk) sigma(.) 5 966.30 5.13 0.03

(SE 98.6; 95% CI 322.1–715.9) in the evergreen and Contrary to our predictions, top-ranked models did not
236.2m (SE 53.1; 95% CI 152.8–365.1) in the reforested indicate a direct effect of rodent biomass or season on
areas. The capture parameter g0 was 0.009 (SE 0.004) Leopard Cat density or movements (Table 3; Appendix
when b=0 and 0.002 (SE 0.0007) when b=1. Based on B). This result could have several explanations. First,
our second top-ranked model, D(habitat) g0(b) σ(.) although we found major differences in rodent biomass
Leopard Cat density was 21.2 individuals/100km2 (SE between seasons and within habitat types, rodents were
5.3; 95% CI 13.1–34.3; Table 3) in the evergreen and 7.9 nonetheless always available throughout the year and
individuals/100km2 (SE 2.7; 95% CI 4.1–15.0; Table 3) in the differences may not have been biologically significant
the reforested areas. Sigma (σ) was 476.0m (SE 93.4; enough to warrant detectable changes in Leopard Cat
95% CI 325.2–696.7). g0 was 0.007 (SE 0.003) when b=0 behavior. Second, although dietary studies indicate
and 0.001 (SE 0.0006) when b=1. Leopard Cats predominantly eat murid rodents, the
species also consumes other prey items including birds,
reptiles, amphibians, and invertebrates (Grassman 1998;
DISCUSSION Grassman 2000; Grassman et al. 2005; Rajaratnam et al.
2007). The availability of such alternative prey, which
Our study is among the first to simultaneously we did not sample, may have mitigated the influence of
conduct camera trapping and rodent live-trapping to rodent biomass on Leopard Cat density and behavior.
assess the relationship between small wild cat density Our raw data, though, do suggest that seasonal
and prey biomass. Specifically, we estimated Leopard differences in rodent biomass may influence Leopard
Cat density and examined the influence of rodent Cat capture probability. For example, during the low
biomass and forest type on the species density and rodent period, we obtained seven recaptures, five of
movements in a degraded tropical forest fragment. which were relocations (i.e., recaptures at different

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Leopard Cat in northeastern Thailand Petersen et al.

Table 4. Comparison of density estimates (D; individual/100km2), standard errors (SE), and coefficient of variation (CV) from our study and
other studies of Leopard Cats using both maximum likelihood (MLE) and Bayesian spatially-explicit capture-recapture. For this study, density
estimates come from two models: aD(.) g0(b) σ(habitat) and bD(habitat) g0(b) σ(.). cStandard deviation (SD) and dcoefficient of variation (CV)
are included for Bayesian estimates.

Study D SE CV Location

This study 17.7 a

3.9 0.22 Overall, Sakaerat Biosphere Reserve, Thailand

21.2b 5.3 0.25 Semi-evergreen forest, Sakaerat Biosphere Reserve, Thailand

7.9 b
2.7 0.34 Artificially reforested areas, Sakaerat Biosphere Reserve, Thailand

Srivathsa et al. 2015 10.5 3.0c 0.29d Semi-evergreen and moist deciduous forest, Bhadra Tiger Reserve, India

4.5 1.3 c
0.29 d
Various habitats, Biligiri Rangaswamy Temple Tiger Reserve, India

Selvan et al. 2014 2.9 0.2 0.07 Wet evergreen forest, Pakke Tiger Reserve, India

Mohamed et al. 2013 12.4 1.6 0.13 Mixed dipterocarp forest, Tangkulap-Pinangah Forest Reserve, Malaysian Borneo

16.5 2.0 0.12 Mixed dipterocarp forest, Segaliud Lokan Forest Reserve, Malaysian Borneo

9.6 1.7 0.18 Mixed dipterocarp forest, Deramakot Forest Reserve, Malaysian Borneo

Bashir et al. 2013 17.0 5.3 0.31 Temperate broadleaf forest, Khangchendzonga Biosphere Reserve, India

camera locations). By contrast, during the high rodent biomass and Leopard Cat density being 2.7 times higher
period, there were only four recaptures, one of which in the evergreen compared with the reforested areas.
was a relocation. We hypothesize that this reflects Only two detections (out of 4,063 trap nights), however,
Leopard Cats being easier to capture during the period came from Sakaerat’s dry dipterocarp forest, the habitat
of low prey biomass, due to being more receptive to our with the second highest estimate of rodent biomass.
scent lure and having larger home ranges, both of which Rabinowitz (1990) similarly observed Leopard Cats using
would result in higher capture probabilities. dry dipterocarp forest less than other habitat types,
Our top-ranked model, D(.) g0(b) σ(habitat), suggests noting that the dipterocarp contained lower densities of
that Leopard Cat movement was lower in reforested the Red Spiny Rat Maxomys surifer, the main prey item
areas compared to the evergreen forest. Because of Leopard Cats during that study. Results from rodent
reforested areas supported lower rodent biomass trapping in the current study had similar findings, with
compared to the evergreen, this finding is contrary to M. surifer being the most frequently captured rodent
our prediction that Leopard Cat movement would be in both evergreen (100% of captures) and reforested
negatively influenced by rodent biomass. This finding, areas (89.1% of captures), but only a few captures in the
however, may be an artefact of our data collection. dipterocarp (3% of captures).
Although we did not target dirt roads, there was an In addition to having the lowest capture rate of
abundance of dirt roads within the reforested area due a key Leopard Cat prey species, the dipterocarp also
to its artificial nature, and one camera in the reforested had relatively high encounter rates for three sympatric
area was placed (at random) near a dirt road (<10m). small carnivore species: Golden Jackal Canis aureus,
This camera alone recorded 50% of our reforested area Javan Mongoose Herpestes javanicus, and Small Indian
individuals and 100% of the reforested area’s recaptures. Civet Viverricula indica (Appendix A). Although direct
Thus, the finding that the Leopard Cat movement evidence of competition between these three species
was lower in the reforested area likely stems from the and Leopard Cats was not reported, interspecific
absence of recaptures at different camera sites. Based competition among other sympatric carnivores is well-
on this experience, we recommend that future studies documented within the literature (Palomares & Caro
targeting Leopard Cats implement stratified study 1998; Donadio & Buskirk 2006). Future studies should be
designs whereby camera traps are placed both on and careful to account for both the prey community and the
off roads to explicitly explore differences in Leopard Cat potential for interspecific competition among sympatric
density, capture probability, and movement. small carnivores when considering the suitability of a
Consistent with our predictions, our second top- specific patch of forest for Leopard Cats.
ranked model [D(habitat) g0(b) σ(.)] does indicate that
Leopard Cat density varied by habitat type in a manner
that reflects prey availability, with both average rodent

Journal of Threatened Taxa | www.threatenedtaxa.org | 12 March 2019 | 11(4): 13448–13458 13455

Leopard Cat in northeastern Thailand Petersen et al.

Leopard Cats and degraded forests Serval Leptailurus serval (Geertsema 1985), Jaguarundi
With an estimate of 17.7 individuals/100km2 Herpailurus yagouaroundi (Konecny 1989), and Guigna
overall and 21.2 individuals/100km2 in the evergreen, Leopardus guigna (Dunstone et al. 2002).
Sakaerat supports the highest recorded SECR-derived Other proposed factors influencing whether the
Leopard Cat density to date (range from other studies: habitat is suitable for Leopard Cats include habitat
2.9–17.0 individuals/100km2; Table 3). These findings structure and its effect on prey ‘catchability’ (Rajaratnam
corroborate other studies which also found Leopard Cat et al. 2007). In addition to supporting high murid
densities to be higher in more degraded environments densities, oil palm plantations are notable for being
(Mohamed et al. 2013; Srivathsa et al. 2015). Srivathsa relatively free of understory vegetation, potentially
et al. (2015), for example, compared Leopard Cat facilitating successful predation events (Rajaratnam et
densities in four protected areas in India, finding higher al. 2007). This may be relevant to our study’s findings
densities clustered around secondary, disturbed, or due to the ubiquity of tall dense bamboo Arundinaria
partially modified forests. Bhadra Tiger Reserve, which pusilla within Sakaerat’s dipterocarp forest. If the
supported the study’s highest Leopard Cat density presence of this grass reduced the catchability of prey
(10.5 individuals/100km2), consists predominantly in the dipterocarp, it may potentially explain why so few
of semi-evergreen forest still recovering from the Leopard Cats were recorded in this habitat. Although we
voluntary resettlement of 26 forest villages in 2002 lack evidence to directly support this hypothesis, studies
and currently adjoins large tracts of coffee plantations of other felids demonstrated a preference for habitats
and several unprotected forest reserves (Srivathsa et with high prey catchability over areas with high prey
al. 2015). Similarly, Mohamed et al. (2013) recorded a availability (e.g., Cheetah Acinonyx jubatus, Broomhall
higher Leopard Cat density in more intensively logged et al. 2003; Lion Panthera leo, Hopcraft et al. 2005).
commercial forest reserves (12.4 individuals/100km2 Leopard Cats tolerate habitat degradation and even
and 16.5 individuals/100km2) compared to a more associate with specific human land uses such as oil palm
sustainably logged reserve (9.6 individuals/100km2) in (Ross et al. 2010) and sugar cane plantations (Lorica
Sabah, Malaysian Borneo. & Heaney 2013) and logged forests (Ross et al. 2010;
It is hypothesized that the Leopard Cat’s association Mohamed et al. 2013). Yet, despite a growing body of
with degraded environments is related to elevated literature, their habitat requirements, population sizes,
rodent populations within such areas (Lambert et al. and long-term viability within both degraded forests
2006; Rajaratnam et al. 2007; Wells et al. 2007; Pimsai and agriculture habitats remain unclear. Our study not
et al. 2014), given the tendency for murid rodents to only contributes to the current need for information
be key prey items of Leopard Cats (Rabinowitz 1990; on population density and habitat selection within
Grassman et al. 2005; Rajaratnam et al. 2007; Shehzad forest fragments but also supports the idea that forest
et al. 2012). Researchers and conservationists, however, fragments may be crucial for the conservation of Leopard
should be careful when interpreting these general Cats in human-dominated landscapes. More research,
findings as not all degraded areas are alike. Oil palm however, is needed concerning the long-term viability
plantations, for example, support abnormally high murid of these populations. To fill this crucial knowledge gap,
rodent densities, which in some cases can exceed 100 we recommend future studies leave the confines of
individuals/ha, whereas during our study the highest protected areas in order to investigate the role their
recorded density within the evergreen was only 10.5 surroundings play both as habitat and as facilitators of
individuals/ha (Wood & Fee 2003; Scott et al. 2004). connectivity.
Variation in prey communities among degraded habitats
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D(.) g0(season) sigma(season) 4 976.18 15.01 0.00

D(.) g0(B) sigma(.) 4 976.26 15.09 0.00

D(.) g0(.) sigma(rodent biomass) 4 976.31 15.14 0.00

D(.) g0(rodent biomass) sigma(.) 4 976.42 15.25 0.00

D(.) g0(Bk) sigma(.) 4 976.49 15.32 0.00

Threatened Taxa

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Journal of Threatened Taxa | www.threatenedtaxa.org | 12 March 2019 | 11(4): 13459–13469

Article
An update on the status of Fishing Cat
Prionailurus viverrinus Bennett, 1833
(Carnivora: Felidae) in Thailand ISSN 0974-7907 (Online)
ISSN 0974-7893 (Print)
Wanlop Chutipong 1 , Anucha Kamjing 2 , Worata Klinsawat 3 , Dusit Ngoprasert 4 ,
Kitipat Phosri 5 , Niti Sukumal 6 , Pongnapa Wongtung 7 & Naruemon Tantipisanuh 8 PLATINUM
OPEN ACCESS
Conservation Ecology Program, King Mongkut’s University of Technology Thonburi, Bang Khun Thien,
1,2,4–8

Bangkok, 10150 Thailand.

3
School of Bioresources and Technology, King Mongkut’s University of Technology Thonburi, Bang Khun Thien,
Bangkok, 10150 Thailand.
1
[email protected], 2 [email protected], 3 [email protected], 4 [email protected],
5
[email protected], 6 [email protected], 7 [email protected], 8 [email protected]
(corresponding author)

Abstract: Fishing Cat Prionailurus viverrinus is threatened throughout its range by habitat loss, persecution, and non-targeted hunting; it
is listed as Vulnerable on the IUCN Red List of Threatened Species. Even basic distribution data are still lacking in many parts of its range,
particularly in southeastern Asia where most wildlife surveys focus on large charismatic carnivores in protected habitats, typically inland
blocks of evergreen or semi-evergreen and deciduous forests. This report aims to update on distribution and status of Fishing Cat in
Thailand. Historic (the 1980s) and current (2007–2017) records from Thailand were compiled based on personal communications, local
news agencies, social media pages, and publications. The current Thai Fishing Cat distribution seems to be highly fragmented and mostly
in coastal wetlands of the Inner Gulf of Thailand and the Thai-Malay Peninsula with one confirmed record from a riverine habitat in central
Thailand. No confirmed records came from protected forested areas—perhaps these are marginal habitat for Fishing Cat. Nevertheless,
there were no targeted surveys in those areas. Fishing Cat was so far not detected from on-going otters’ targeted camera trap surveys
along Thailand’s Andaman coast. Future surveys should focus on coastal and inland wetlands to expedite the discovery of remaining
populations before these are extirpated.

Keywords: Coastal wetland, human-dominated habitat, inland wetland, mangrove forest, persecution, riverine habitat, species distribution,
threat assessment.

Abbreviations: DNP - Department of National Park, Wildlife, and Plant Conservation; SRY - Khao Sam Roi Yot National Park; TSL - Thung
Salaeng Luang National Park.

DOI: https://doi.org/10.11609/jott.4557.11.4.13459-13469 | ZooBank: urn:lsid:zoobank.org:pub:DE815A67-7ED6-4BC8-B804-DCE3F572E5C7

Editor: Angie Appel, Wild Cat Network, Germany. Date of publication: 12 March 2019 (online & print)

Manuscript details: #4557 | Received 11 November 2018 | Final received 30 December 2018 | Finally accepted 04 March 2019

Citation: Chutipong, W., A. Kamjing, W. Klinsawat, D. Ngoprasert, K. Phosri, N. Sukumal, P. Wongtung & N. Tantipisanuh (2019). An update on the status of
Fishing Cat Prionailurus viverrinus Bennett, 1833 (Carnivora: Felidae) in Thailand. Journal of Threatened Taxa 11(4): 13459–13469; https://doi.org/10.11609/
jott.4557.11.4.13459-13469

Copyright: © Chutipong et al. 2019. Creative Commons Attribution 4.0 International License. JoTT allows unrestricted use, reproduction, and distribution of this
article in any medium by adequate credit to the author(s) and the source of publication.

Funding: This research was (partially) supported by the Asahi Glass Foundation, National Science and Technology Development Agency [P-15-51209], and King
Mongkut’s University of Technology Thonburi.

Competing interests: The authors declare no competing interests.

For Author details & Author contribution see end of this article.

Acknowledgements: We thank Angie Appel for her effort in encouraging, managing, and editing our manuscript. Comments and suggestions made by two
reviewers (JW Duckworth and Daniel Willcox) help improving the manuscript significantly. We also thank Shomita Mukherjee for co-editing our manuscript and
Vanessa Herranz Muñoz for clarification regarding Fishing Cat from Cambodia. We are grateful to George A. Gale for reading earlier version of our manuscript. In
DNA barcoding protocol, we appreciate helpful discussion, financial support and technical assistance from Assoc. Prof. Dr. Worawidh Wajjwalku and Ms. Nongnid
Kaolim from Faculty of Veterinary Medicine, Kasetsart University. We are grateful to Parinya Padungthin for pointing out records of Fishing Cat in Thailand.

13459
Status of Fishing Cat in Thailand Chutipong et al.

INTRODUCTION particularly along the west coast of southern Thailand.

Given such a paucity of surveys in suitable habitats,
Fishing Cat Prionailurus viverrinus has a wide Thailand might hold a large Fishing Cat population or,
distribution across southern and southeastern Asia. equally, the species might be close to extinction (Appel
Most populations, however, appear to have declined & Duckworth 2016). This very wide range of possibilities
significantly due to habitat loss, non-targeted hunting, clearly indicates that there is an urgent need for a
and retaliatory killing associated with livestock conservation status assessment in the country.
depredation and/or damage to aquaculture (e.g., fish This article compiles evidence of Fishing Cat
ponds; Mukherjee et al. 2016). Known strongholds are occurrence that was not included in the previous review
Sri Lanka, Bangladesh, West Bengal in India, and the by Tantipisanuh et al. (2014). Some records in this article
Terai-Duar belt of the Himalayan foothills in India and date to before 2014, but these records are not exhaustive
Nepal (Mukherjee et al. 2016). In southeastern Asia, as attempts were not made to review all historic records,
recent records dating to 2000–2016 are scarce, and e.g., examining specimens in museums. This remains a
the species occurrence is extremely patchy in Vietnam, priority because it has the possibility to indicate a longer-
Cambodia, Thailand, Myanmar, and Indonesia (Java and term change in range. Results from a camera trap survey
perhaps Sumatra) (Mukherjee et al. 2016 and references in a coastal mangrove site where wetland-associated
therein). Fishing Cat is listed as Vulnerable on the IUCN species such as otters, Lutrinae, were targeted are also
Red List of Threatened Species (Mukherjee et al. 2016). summarized. This present article provides an update on
Thailand may be one of the important strongholds Fishing Cat distribution in Thailand and therefore helps
for Fishing Cat and a regional priority range country for to identify areas for further surveys and conservation
its conservation—it has few degraded habitats, at least efforts in the country. It is also in line with one of the
those that are potentially suitable for Fishing Cat, e.g., objectives of the Fishing Cat conservation strategy that
in coastal mangroves, within large protected areas with aims to close information gaps on Fishing Cat distribution
a high protection level and law enforcement measures, and status in range countries (Appel & Duckworth 2016).
and populations of ecologically similar species like otters.
Nevertheless, between 1996 and 2011, there were
only a few targeted surveys for Fishing Cat that yielded METHODS
confirmed records and these were mainly in and around
Khao Sam Roi Yot (SRY) and Thale Noi Non-hunting Area We compiled Fishing Cat evidence from various
(Cutter & Cutter 2009; Tantipisanuh et al. 2014; Fig. 1). sources—personal communications, newspaper articles,
Results of radio telemetry research on 23 radio-collared social media pages, and publications. We attempted
Fishing Cats in an area of approximately 35km2 suggested to verify each report by (1) circulating images among
that SRY was a stronghold for the Fishing Cat in Thailand experts for confirmation of species identification when
(Cutter & Cutter 2009; Cutter 2015; Patumrattanathan images were available, or (2) visiting and taking images
2015). In the same area, however, negative interaction and recording narration from owners of the stuffed
with people on livestock-raiding led to retribution killings mounted specimens, reports, and then circulating
of at least five out of 16 Fishing Cats monitored during images for confirmation. Records date from the late
this study (Cutter 2015). 1980s to the late 2010s. Some records are from the
In a review of the status of small cats in Thailand same review period in Tantipisanuh et al. (2014) but
(Tantipisanuh et al. 2014), Fishing Cat rarely occurred were overlooked at the time and therefore not included
in protected areas with no significant wetland habitats in it.
where most of camera trap surveys were conducted,
although none of these surveys had specifically targeted
Fishing Cat, except that of Cutter & Cutter (2009). RESULTS AND DISCUSSION
Wetland habitats such as mangrove and peat swamp
which were largely under-surveyed may still hold some Records came from Muang District of Phitsanulok
remaining Fishing Cat populations and other threatened Province in north-central Thailand, Bangkhuntian
small carnivores and therefore require immediate District of Bangkok Province, Muang District of Samut
attention for surveys (Chutipong et al. 2014). These Sakorn Province, Laem Phak Bia Subdistrict in Ban Laem
wetlands were heavily used for aquaculture in the past District of Phetchaburi Province located in the Inner
several decades but in many parts are still little degraded, Gulf, and Singhanakorn District of Songkhla Province,

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Status of Fishing Cat in Thailand Chutipong et al.

Muang District and Mai Kaen District of Pattani Province subterranean streams in the Chao Phraya basin and Nan
in peninsular Thailand (Fig. 1). Ongoing surveys in River sub-catchment, and 29% mixed deciduous forest.
mangrove habitats in Ranong, Phang-nga, and Krabi This habitat is unlikely to harbour Fishing Cats. Since
provinces along Thailand’s western coast so far failed to there were no targeted surveys for Fishing Cat anywhere
detect any evidence of Fishing Cat. near or in the park, the absence of Fishing Cat there
remains uncertain. The park, however, may be of low
A. Upper Central Thailand priority for a Fishing Cat targeted survey due to a lack of
A.1. Sa Khlo Village, Hua Raw Subdistrict, Muang significant wetlands.
District, Phitsanulok Province This inland record in Sa Khlo Village would be a really
On 24 December 2012, a local news agency reported significant one if it is genuinely of a wild animal since
a male Fishing Cat captured by a group of people at it is almost 400km from the coast where most suitable
Sa Khlo Village (Anonymous 2012; Fig. 1). The report, habitats lie. It would also indicate that Fishing Cat once
however, did not elaborate details on the type of habitat occurred far away from coastal wetlands. One inland
and the animal’s capture. It further documented that record of ‘Fishing Cat’ is known from northwestern
the cat was reported to a local politician who appeared Thailand, in Tak Province (Duckworth et al. 2010) and
to recognize the animal as a Fishing Cat. He then asked to another one, a camera trap record from Kulen Promtep
keep the animal instead of releasing it to the wild, in fear Wildlife Sanctuary in northern Cambodia (Rainey & Kong
of the cat being killed and eaten by locals. He convinced 2010). In the Indian subcontinent, there are many more
the locals that he would report to the authorities, inland records, e.g., from Terai Arc landscape, Chitwan
the Department of National Park, Wildlife, and Plant and Badia NPs, Nepal (Dahal & Dahal 2011; Yadav et
Conservation (DNP), for permission to keep the animal al. 2018), India (e.g., Palei et al. 2018), and Sri Lanka
in captivity. There was no further information to validate (Thudugala 2016). Duckworth et al. (2010) did not,
if the cat was a resident caught in the mentioned area however, examine the specimen from northwestern
and not an escaped pet. The set of images published Thailand, and thus they could not be sure whether the
with this newspaper article was circulated among six specimen was correctly identified and of certain origin
experienced field conservationists/ researchers, and (J.W. Duckworth in litt. 1 July 2017). Due to the intense
all agreed that the animal was a Fishing Cat, based on use of inland wetlands in southeastern Asia, this type of
its features: small ears in relation to head, short tail in habitat is now much reduced (Davidson 2014). As Fishing
relation to the body, and protruding claws of the front Cat is highly adaptable and can persist in degraded
paws. Examination of the habitat (using Google Earth, habitats (e.g., SRY: Cutter 2015; Pathumratanatarn
16 October 2017; Table 1) where the cat was claimed 2015), all sizes and conditions of inland wetlands need
to be caught, included a small low-lying river, named to be identified and surveyed.
Sa Khlo, which runs through paddy fields and has a 3–5
m wide scrubby strip dotted with clumps of bamboo B. Inner Gulf of Thailand
situated along both sides. Small and scattered human B.1. Bangkhuntian District, Bangkok Province
settlements are situated on both sides of the river in On 10 July 2014, W. Chutipong and D. Ngoprasert
this relatively flat area. Satellite image of the riverine interviewed a local school teacher, a former hunter,
habitat along the Sa Khlo River matches no other habitat regarding the historic occurrence of Fishing Cat in
where Fishing Cat populations were found in low-lying Bangkhuntian, a suburb of Bangkok. The person said
riverbeds but perhaps similar to those in Pakistan (see that he had stuffed 14 Fishing Cats, which were killed
Roberts 1977 cited by Appel 2016). by either him or his sibling or sourced from locals in
A literature search (published in English) traced no and around his community. He also mentioned that
records of Fishing Cat in the nearby Thung Salaeng Luang Fishing Cat was opportunistically hunted for meat or as
National Park (TSL; 1,262km2), located about 20km retribution for killing livestock. We traced one stuffed
farther east. TSL is part of the Phu Miang-Phu Thong mount stored at a local museum held by the Klong
Conservation Corridor (9,944km2), which also includes Pittayalongkorn School where he is currently working
Khao Kho National Park and Wang Pong-Chon Daen (Figs. 1; Image 1). This specimen, dated to the late
Non-hunting Area (Tordoff et al. 2012). According to 1980s, was obtained from a local villager who trapped
the GIS database of the Land Development Department the cat accidentally with a fish trap (Kriangsak Rukngam
2018 (LDD 2018), TSL consists mainly of evergreen forest pers. comm. 10 July 2014).
(59%) with patches of limestone caves and associated

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Status of Fishing Cat in Thailand Chutipong et al.

Figure 1. Locations of traced Fishing Cat records. Letters and numbers represent records described in the text and in Table 1. All coordinates are
approximate locations based on available information from each record unless otherwise stated.

Table 1. Summary of Fishing Cat and Leopard Cat records in Thailand from the late 1980s to 2016. All coordinates are approximated locations
based on available information unless otherwise stated.
Date of image
Locality Coordinates Evidence Source Remarks
taken
A. Upper Central Thailand
A.1. Sa Khlo, Muang, Local news
16.8830N, 100.2830E Images [not indicated]
Phitsanulok (Anonymous 2012)
B. Inner Gulf of Thailand
The animal was stuffed in the early 1980s
Image of stuffed Interviewed the
B.1. Bangkhuntian, Bangkok 13.5170N, 100.4170E 10 July 2014 and kept at Pittayalongkorn Pittayakhom
mount specimen hunter
School; Image 1.
A road-kill female Fishing Cat found on a
B.2. Phantai Norasing, Khok Image of stuffed Wanlop Chutipong,
13.5000N, 100.3170E 10 July 2014 road near a school on 14 December 2013 at
Kham, Samut Sakorn mount specimen Dusit Ngoprasert
5.30h; Image 2.
It is uncertain whether the photographed
B.2. Phantai Norasing, Khok Image of a cat in a cat is from SRY/ Phantai, but it is certain that
13.4830N, 100.3170E Kitipat Phosri 28 May 2016
Kham, Samut Sakorn breeding centre some Fishing Cats were trapped in Phantai
and taken to the breeding centre; Image 3.
Coordinates were taken at the restaurant
B.3. Laem Pak Bia, Ban Laem, Jonathan Murray
13.0170N, 100.0670E Skull 2010 where the skull of a Leopard Cat was
Phetchaburi (deceased)
retrieved; Image 4.
Image of a dead
B.3. Laem Pak Bia, Ban Laem, Internet search for
13.0330N, 100.0830E animal posted March 2011 Image 5; the URL is no longer available.
Phetchaburi Fishing Cat
online
C. Peninsular Thailand
https://www.youtube.com/watch?v=aBj7_R-
07.2500N, 100.4670E
Report with GuxA (YouTube video)
(Pa Khat), Suppakorn
C.1. Singhanakorn, Songkhla images and VDO 2013–2015 The VDO shows the release of a female
07.2500N, 100.4330E Patumrattanathan
link. Fishing Cat as part of the study mentioned in
(Pak Ro)
the report (Ramsuti 2014).
Image of live- Wanchamai
C.2. Bangplamor, Pattani 06.8500N, 101.2000E 2007 Reported in Buatip et al. 2013; Image 6.
captured animal Karnthanut
The wild-caught Fishing Cat from Mai Kaen
Image of caged
C.3. Mai Kaen, Pattani 06.6330N, 101.6670E Niti Sukumal 2014 District, Pattani Province, was kept as a pet;
animal
Image 7.

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Status of Fishing Cat in Thailand Chutipong et al.

Figure 2. Survey grids used for completed, on-going, and planned otter occupancy surveys in southern Thailand. Approximated locations of
records (C.1.–C.3.), as reported in the main text, are shown.

B.2. Phantai Norasing, Khok Kham Subdistrict, A. Kamjing (Figs. 1; Image 2) at the Marine and Coastal
Muang District, Samut Sakorn Province Resource Conservation Center located close to where
In December 2013, during an occupancy survey for the animal was found. The road-killed cat was found
Smooth-coated Otter Lutrogale perspicillata and Asian dead on a road close to a patch of mangrove forest next
Small-clawed Otter Aonyx cinereus in the Inner Gulf to a school by W. Chantong who then reported it to the
of Thailand, A. Kamjing heard of a road-killed Fishing mangrove conservation centre nearby. The reporter was
Cat at Phantai Norasing from local landowners when convinced that the cat was of wild origin since no Fishing
he surveyed their ponds for otter signs. In July 2014, Cat was kept as a pet in the area that he was aware of.
W. Chutipong and D. Ngoprasert examined a stuffed In 2012, S. Patumrattanathan, a researcher from DNP,
mount of a female road-killed Fishing Cat reported by live-trapped four adult Fishing Cats, comprising one male

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Status of Fishing Cat in Thailand Chutipong et al.

a b

Image 1. A stuffed mount specimen of a Fishing Cat dated back to the 1980s from Bangkhuntian District, Bangkok: a - top view; b - lateral view.
© W. Chutipong, 10 July 2014.

and three females close to Phantai Norasing, Khok Kham Jonathan Murray who obtained at least one skull of a
Subdistrict, Muang District, Samut Sakorn Province. This Fishing Cat (Figs. 1; Image 4, but see discussion below)
trapping was part of a three-year project with the aim to from a restaurant close to Laem Phak Bia in 2010. He
increase the genetic diversity of Fishing Cat in Thailand questioned the owner to confirm the origin of the skull.
by breeding Fishing Cats from SRY and Phantai Norasing The cause of death of this specimen was not reported.
and then releasing the captive-bred individuals at Bueng Based on DNA analysis of its nasal bone fragments,
Boraphet, a wetland of international importance in however, we confirmed that the skull was of a Leopard
Nakhon Sawan Province (Suppakorn Patumrattanathan Cat Prionailurus bengalensis. The analysis was done using
pers. comm. 13 August 2018). All four trapped Fishing multilocus DNA barcoding (the combined sequences of
Cats from Phantai and the unknown number from SRY partial mtDNA Cytochrome b,  16S,  ND5  gene,  Control
were relocated to Khao Prathab Chang Wildlife Breeding Region, and four Y-chromosome introns including
Center in Pak Chong Subdistrict, Chom Bueng District, SMCY3,  SMCY7, DBY7, and UTY11; Luo et al. 2014),
Ratchaburi Province, and gave birth to two kittens. The coupled with reference databases of genetic diversity of
project, however, was suspended in 2013 before the Fishing Cat and Leopard Cat in Thailand (Klinsawat et al.
cats were released. Now only 1–2 pairs of these Fishing unpublished data) and other range countries (Luo et al.
Cats remain at the breeding centre. Kitipat Phosri visited 2014; Patel et al. 2017). With this example, we strongly
the centre in early 2018 and saw the animals (Image recommend that when recovering animal parts without
3). No precise information was available about which accompanying images of the entire body, one should
individuals were caught in SRY and Phantai Norasing. conduct DNA analysis to verify species identification.
Since S. Patumrattanathan studied Fishing Cats in SRY Another record from this area appeared in March
and Singhanakorn District, Songkhla Province (see C1 2011. An image of a dead Fishing Cat was posted on the
in Fig. 1) using radio telemetry and in captivity for 6–7 internet (Image 5) by a person who observed the carcass,
years (Patumrattanathan 2015; Pathumratanathan et al. apparently shot by a worker from a nearby construction
2015), we are confident that he indeed captured Fishing site. The cat was shot by a modified slingshot used
Cats in this area. for fishing. The reporter, however, mistook the animal
B.3. Laem Phak Bia Subdistrict, Ban Laem District, as a Black Leopard Panthera pardus. A description of
Phetchaburi Province the post (in Thai) indicated that the cat was shot when
Two records came from Laem Phak Bia in Ban Laem the poacher was searching for wild game for a special
District of Phetchaburi Province, located in south-central feast. It was also mentioned that wild game hunting
Thailand. The first record was reported by the late was a common practice among the group of people

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Status of Fishing Cat in Thailand Chutipong et al.

Image 2. A stuffed mount specimen of a road-killed female Fishing Image 3. A wild-caught Fishing Cat, presumably from either Sam
Cat from Samut Sakorn Province. © W. Chutipong, 10 July 2014. Roi Yot National Park or Phantai, Khok Kham, kept at Khao Prathab
Chang Wildlife Breeding Center. © K. Phosri, 28 May 2016.

mentioned in the post. This latter evidence suggested garzetta nests and mentioned Fishing Cat as a possible
the continued presence of the Fishing Cat in Laem Phak nest predator. A follow-up revealed a confirmed record
Bia area (at least up to March 2011). On the other hand, of Fishing Cat occurrence in the area (Image 6). “[In
threats such as illegal opportunistic hunting were also 2007] this cat was trapped in a patchy mangrove around
documented. the village named Bangplamor of Muang District, Pattani
Province. It is about 15 minutes’ drive and close to the
C. Peninsular Thailand [Prince Songkhla] University’s [Pattani] campus where
C.1. Pa Khat and Pak Ro subdistricts, Singhanakhon the locals earn their living on small scale fishery,” said
District, Songkhla Province W. Karntanut, one of the article’s author. W. Karntanut
Pathumratanathan et al. (2015) conducted an also reported that two cats were trapped at the same
ecologic study of Fishing Cat in three subdistricts of time but one died shortly after. The remaining cat was
Singhanakhon District, Songkhla Province, covering an released after being photographed since it did not take
area of approximately 42km2, between 2013 and 2015 any of the provided food. Further correspondence
(Fig. 1). Ten Fishing Cats were captured, comprising seven revealed that Fishing Cats thrived, at least at the time
males and three females, in the two subdistricts Pa Khat of the report in late 2013, in coastal mangrove habitat
and Pak Ro. Five males and one female were fitted with close to the campus and local communities. There were
VHF radio transmitters. Habitats in these subdistricts also reports of persecution due to loss of livestock.
consist mainly of paddy fields (52%), degraded peat Sometimes, however, “trouble-making” Fishing
swamp and mangrove (24%), agriculture areas including Cats were trapped and kept as pets instead of being
shrimp farms (10%) and oil palm plantations (8%), and persecuted (Wanchamai Karntanut in litt. 1 October
settlements (6%) (Pathumratanathan et al. 2015). The 2013). This record appears to be the southernmost
authors mentioned active hunting of Fishing Cat for meat confirmed record in mainland southeastern Asia (Angie
and persecution due to perceived loss of aquaculture Appel in litt. 22 December 2013).
stock such as fish and shrimps, and highly valued C.3. Mai Kaen Village, Mai Kaen Subdistrict, Mai
fighting roosters that are specially selected and raised Kaen District, Pattani Province
for cockfight or gamecocks. The authors, however, do Another report of Fishing Cat from Pattani Province
not quantify the intensity of such killings and hunting came from Mai Kaen District in 2014 (Fig. 1). This cat
pressures. (Image 7) was caught in mangrove forest close to Saiburi
C.2. Bangplamor Village, Rusamilae Subdistrict, River, but no precise date and location were reported.
Muang District, Pattani Province The person who caught the cat lived there for more
In late 2013, evidence of Fishing Cat was uncovered than 60 years. He and N. Sukumal, who was born in
in Pattani Province (Fig. 1), where Buatip et al. (2013) this area, reported that Fishing Cat was not commonly
conducted research on predation of Little Egret Egretta encountered in the past 20 years, but it was present in

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Status of Fishing Cat in Thailand Chutipong et al.

a
b
c

Image 4. A skull of Leopard Cat retrieved from a restaurant near Laem Phak Bia by the late Jonathan Murray in 2010: a - top view; b - lateral
view; c - front view. © A.J. Pierce, 2018.

Image 5. A poached Fishing Cat from Laem Phak Bia area. Image
posted on the internet in March 2011 by a worker from a nearby Image 6. A wild-caught Fishing Cat in captivity in Pattani Province.
construction site. © W. Karntanut, 5 May 2007.

the area of mangrove forest along Saiburi River and the where Asian Small-clawed Otters were found during a
canals that connect to the main Saiburi River. In the past preliminary survey. In each grid cell, we set 3–6 camera
two decades, Fishing Cats opportunistically entered the trap stations at locations where we found evidence
areas in this district to prey on poultry (Niti Sukumal of otter presence like spraints and footprints. Some
personal observation). Some cats were caught and kept cameras were also deployed in sites without evidence
as pets, and some were killed (Niti Sukumal personal of otters but which exhibit similar characteristics as
observation). Fishing Cat is rare in the area at present, habitat used by otters, e.g., the presence of mounds that
although villagers reported in 2016 that two kittens were are well above the highest tide. To avoid inundation,
found in the forested area dominated by Melaleuca cameras were set 1–3 m above ground but still aimed at
cajuputi. This report, however, should be treated with the focal areas on the ground, which was large enough
caution as species identification could not be validated; to capture large otter groups (Image 8). Cameras were
the animals were already sold by the time we visited the kept at locations for approximately 20 days. Fish oil was
area in 2016. used as a lure to attract the focal species to the focal
C.4. Southwestern coast, Ranong, Phang-nga, and area of cameras, i.e., approximately 3m in front of the
Krabi provinces camera traps. Fish oil lure appears to attract Fishing
Camera trap surveys targeting Smooth-coated Cats to camera traps as observed in a concurrent survey
Otter and Small-clawed Otter in coastal mangroves in of Fishing Cat in SRY (Kitipat Phosri & Dusit Ngoprasert
southwestern Thailand have been running since August personal observations).
2016 (Fig. 2; Tantipisanuh et al. 2018). Grid cells of Eighteen months of surveys in Ranong, Phang-nga,
5km×5km for camera trapping were initially selected and Krabi provinces covered an area of approximately
on a basis of a minimum area of mangrove (10%) 2,825km2 and totalled 11,563 camera trap days across

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Status of Fishing Cat in Thailand Chutipong et al.

Fishing Cats were recorded in coastal mangroves

in other range countries, e.g., the deltaic mangrove
forest of Coringa WS in Andhra Pradesh (Mukherjee et
al. 2012) and Odisha, both in eastern India (Palei et al.
2018), mangrove forests of southern Cambodia (Thaung
et al. 2017), and Ayeyarwady Delta of Myanmar (Naing
Lin & Than Zaw, WCS Myanmar Program, in litt. 11 May
2018). Fishing Cat, however, was not detected during
surveys conducted along coastal Kerala in southwestern
India (Janardhanan et al. 2014). Currently, it is unclear
whether Fishing Cat occurs in this part of Thailand and
was simply missed in the current surveys or whether it is
not present there at all. Despite the non-detection, the

entire southwestern coastal wetlands are still priority
Image 7. A wild-caught Fishing Cat in captivity in Mai Kaen District,
Pattani Province. © N. Sukumal, 2014. sites, as they provide potentially suitable habitat, and
need targeted surveys for Fishing Cat.

558 camera trap stations. Smooth-coated Otter and MANAGEMENT IMPLICATIONS AND CONCLUSIONS
Asian Small-clawed Otter were detected in 165 of 558
camera trap stations (30%) and in 71 stations (13%), These new localities show that Fishing Cat is more
respectively. Meanwhile, Leopard Cat (0.7%), Large- widely distributed along Thailand’s coastal wetlands
spotted Civet Viverra megaspila (0.2%), Greater Hog than previously reported—eight records traced were
Badger Arctonyx collaris (0.2%), Small Asian Mongoose from outside protected areas. The single confirmed
Herpestes javanicus (0.3%), and Common Palm Civet record (accepting that the cat’s origin is not known)
Paradoxurus hermaphroditus (27%) were also recorded, traced from inland and its surrounding habitat, appears
but not a single Fishing Cat (Tantipisanuh et al. to match known inland records from elsewhere in Fishing
unpublished data). Assuming that other small carnivore Cat range. Combined with the previous confirmed
species that are present were readily detected during records in Tantipisanuh et al. (2014), this evidence
the surveys, Fishing Cat either occurs at a very low strongly suggests that Fishing Cat populations largely
abundance or does not occur at all in the surveyed area. occur outside Thailand’s protected area system. These
It is possible that Fishing Cat has very specific habitat results highlight the need to conduct targeted surveys
requirements, which our otter targeted survey failed to for Fishing Cat in both coastal zones and inland areas
cover. with suitable habitat outside protected areas in human-

a b

Image 8. Camera trap images of family groups from Phang-nga Province in 2017: a - Smooth-coated Otter from Kuraburi; b - Asian Small-clawed
Otter from Takuapa. © Tantipisanuh et al. 2018.

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Steinmetz, K.E. Jenks, L.I. Grassman Jr., M. Tewes, S. Kitamura,
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of the species. Due to the difficulty in distinguishing Gale, F.K. Harich, A.C. Treydte, P. Cutter, P.B. Cutter, S. Suwanrat,
between Fishing Cat and the co-occurring Leopard Cat, K. Siripattaranukul, Hala-Bala Wildlife Research Station, Wildlife
Research Division & J.W. Duckworth (2014). Current distribution
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Marienberg, Germany & Saltford, Bristol, United Kingdom, 73pp. Peafowl (Pavo muticus)”. He is interested in conservation and ecology
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Valley of Bardia National Park, Nepal. Cat News 67: 31–33. surveys of small carnivores in coastal wetlands and mangroves in
southern Thailand.

Author contribution: All authors contributed equally to the conception,

Threatened Taxa
design, data collection, writing, and revising the manuscript.

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Journal of Threatened Taxa | www.threatenedtaxa.org | 12 March 2019 | 11(4): 13470–13477
Communication

The Caracal Caracal caracal Schreber, 1776

(Mammalia: Carnivora: Felidae) in Uzbekistan
ISSN 0974-7907 (Online) Mariya Alexeevna Gritsina
ISSN 0974-7893 (Print)

PLATINUM Institute of Zoology, Academy of Sciences of the Republic of Uzbekistan, Tashkent 100047, Uzbekistan.
[email protected]
OPEN ACCESS

Abstract: This article provides information about 27 records of the Caracal Caracal caracal in Uzbekistan, particularly in the Kyzylkum Desert
and the Ustyurt Plateau. The data collected between 2011 and 2017 were based on information from literary sources, field research, and
interviews with local people. At least 11 individuals of the species were killed intentionally and one was run over by a car. Basic threats to
the species in Uzbekistan are negative interactions between herders and Caracal, lack of knowledge about its protected status among local
people, and lack of conservation measures. The preconditions for the protection of Caracal are the existence of remote unpopulated areas
close to state borders between adjacent countries and socio-economic factors that prompt people to move from rural to urban areas. Caracal
habitats are protected in Kyzylkum State Reserve, Saigachiy Landscape Sanctuary, and six wildlife sanctuaries. To conserve Caracal, it is
necessary to strengthen the network of protected areas in deserts and raise the awareness of local communities and decision-makers in the
national government. It is important to continue research on Caracal and develop a government-approved action plan for its conservation.

Keywords: Central Asia, interview survey, Kyzylkum Desert, negative interaction, threat assessment, threatened felid, Ustyurt Plateau.
Russian Abstract: В публикации представлена информация о 27 встречах каракала Caracal caracal в пустыне Кызылкум и на плато
Устюрт (Республика Узбекистан). Приведены сведения за период с 2011 по 2017 гг., которые основаны на имеющихся литературных
данных, собранном во время экспедиционных выездов материале и сообщениях местного населения. По крайней мере 11
особей каракала были преднамеренно убиты и еще одно животное было сбито машиной. Основными угрозами в Узбекистане
представляются существующий конфликт между скотоводами и каракалами, отсутствие знаний об охранном статусе вида среди
местного населения и отсутствие должных мер по его охране. Предпосылками для сохранения каракала являются наличие
удаленных незаселенных территорий вблизи государственных границ между соседними странами и социально-экономические
факторы, которые побуждают людей переезжать из сельской местности в города. Места обитания каракала охраняются в
Кызылкумском государственном заповеднике, ландшафтном заказнике «Сайгачий» и еще в шести природных заказниках. Для
сохранения каракала необходимо укрепить сеть пустынных охраняемых природных территорий и повысить осведомленность
местного населения и лиц, принимающих решения на правительственном уровне. Представляется важным продолжить
исследования по каракалу и в дальнейшем разработать одобренный правительством план действий по его сохранению.

DOI: https://doi.org/10.11609/jott.4375.11.4.13470-13477 | ZooBank: urn:lsid:zoobank.org:pub:08E55358-2C11-48AD-9D2F-E2B382D448B7

Editor: Angie Appel, Wild Cat Network, Germany. Date of publication: 12 March 2019 (online & print)
Manuscript details: #4375 | Received 01 July 2018 | Final received 08 December 2018 | Finally accepted 02 March 2019
Citation: Gritsina, M.A. (2019). The Caracal Caracal caracal Schreber, 1776 (Mammalia: Carnivora: Felidae) in Uzbekistan. Journal of Threatened Taxa 11(4):
13470–13477; https://doi.org/10.11609/jott.4375.11.4.13470-13477
Copyright: © Gritsina 2019. Creative Commons Attribution 4.0 International License. JoTT allows unrestricted use, reproduction, and distribution of this article in
any medium by adequate credit to the author(s) and the source of publication.
Funding: Data collected during expeditions aimed on different biodiversity study topics and supported by Central Asian Biodiversity Network, German Academic
Exchange Service, Manfred Hermsen Foundation, German Federal Ministry of Environment, Nature Conservation and Nuclear Safety, World Wide Fund for Nature,
Central Asian Desert Initiative, Michael Succow Foundation, Saiga Conservation Alliance, and Mohamed bin Zayed Species Conservation Fund.
Competing interests: The author declares no competing interests.
Author Details: The author, a graduate of the biological faculty, Samarkand State University, Uzbekistan, has a master degree in zoology. She worked as a
researcher at the Zarafshan National Reserve and later Samarkand Museum of Regional Studies. Since 2015 she is a researcher at the Institute of Zoology,
Academy of Sciences (Uzbekistan).
Acknowledgements: The author of this article is deeply thankful to Natalia Marmazinskaya, Denis Nuridjanov, Timur Abduraupov, Valentin Soldatov, and Alexander
Korshikov—her colleagues and constant companions in research expeditions, assisting her in collecting material and supporting her in every way. Special thanks
to Jim Sanderson and Anna Barashkova, initiators and supporters of various projects associated with small cats implemented in Uzbekistan, and to Angie Appel
who encouraged us to prepare this publication. The author thanks the government agencies of Uzbekistan, Central Asian Biodiversity Network, German Academic
Exchange Service, Manfred Hermsen Foundation, German Federal Ministry of Environment, Nature Conservation and Nuclear Safety, World Wide Fund for Nature,
Central Asian Desert Initiative, Michael Succow Foundation, Saiga Conservation Alliance, and Mohamed bin Zayed Species Conservation Fund for their support.

13470
Caracal in Uzbekistan Gritsina

INTRODUCTION Tien Shan and Pamir-Alai ranges in the southeast. Most

of the desert lies in Uzbekistan and Kazakhstan, with a
The Caracal Caracal caracal is one of the rarest few small portions in Turkmenistan (Fig. 1). The basic
vertebrates in central Asia, where it occurs in Uzbekistan, landscape of the Ustyurt Plateau is clay desert covered
Kazakhstan, Turkmenistan, northern Iran, and Afghanistan with Wormwood Artemisia or mixed Wormwood and
(Heptner & Sludskii 1972). Its range also covers arid Glasswort Salicornia communities, with occasional sand
regions in Africa, the Arabian Peninsula, the Middle East, areas. The plateau is edged by ‘chinks’, a regional name
Pakistan, and India (Avgan et al. 2016). for escarpments, which are up to 200m high in some
The Turkmen Caracal is included as Critically places. Most of the plateau is covered with vegetation,
Endangered in the Red Data Books of Uzbekistan varying from Wormwood and Glasswort in the northern
(Abdunazarov 2009) and Kazakhstan (Bekenov & desert subzone to ephemeral plants and Wormwood in
Kasabekov 2010) and as Endangered in the Red Data the southern desert subzone. The summer is hot and
Book of Turkmenistan (Hodzhamuradov & Imamov 2011). long, lasting from May to September. Average daily
The species is included in Appendix I of the CITES (Avgan temperature in July is 26–28 °С, reaching up to 40–60 °С
et al. 2016). in some years. In winter it drops to -26°С and even to
In the 20th Century, the Caracal was recorded on -41°С.
Uzbekistan’s Ustyurt Plateau, in Lower Amudarya area, The Kyzylkum Desert largely consists of fixed and
northwestern and southwestern parts of Kyzylkum Desert, semi-fixed dunes, with occasional patches of unvegetated
and the plains adjoining Surkhan and Zeravshan rivers sand. It includes extensive areas of clay, gypsum, takyr,
(Heptner & Sludskii 1972; Mitropolsky 1979; Lesnyak et and saline soils and low sky islands. The plant species
al. 1984; Bogdanov 1992; Abdunazarov 2009). It inhabits prevailing on sandy patches are Sand Sedge Carex
bumpy, well-fixed sands along the Ustyurt Plateau arenaria, White Saxaul Haloxylon persicum, Calligonum,
escarpments and on gypsum and stony plains (Sapojnikov and Richter’s Saltwort Salsola arbuscula; the ones on clay
1962; Heptner & Sludskii 1972). Its diet consists primarily are Wormwood often mixed with shrubs, and Anabasis
of Tolai Hare Lepus tolai, Gerbils Gerbellidae, Jerboas salsa mixed with Saltwort. Average temperature by day
Dipodidae, birds, reptiles, and insects. Sometimes it in July is between 26°C and 29°C, reaching up to 51°С, and
hunts Goitered Gazelle Gazella subgutturosa, Red Fox in January between 0°C and 9°С.
Vulpes vulpes, and lambs of Domestic Sheep Ovis aries
(Sapojnikov 1960, 1962).
Only fragmentary data are available on its current MATERIAL AND METHODS
distribution and population size in Uzbekistan. Studies
on Caracal were not conducted since the 1980s. Most Information was obtained through the collection
of the information on recent encounters with Caracal and analysis of data from literature, field surveys, and
was published in Russian (Lim 2009; Lim & Klichev 2009; interviews. Data collection in the field was carried out
Gritsina 2012; Marmazinskaya еt al. 2012; Bykova et al. during expeditions to the Ustyurt Plateau and Kyzylkum
2015; Gritsina et al. 2016; Marmazinskaya & Mardonova Desert (Fig. 1). During the expeditions, we observed the
2016). Thus, this information is poorly available for the terrain from elevated points using binoculars or telescope
international audience and is expedient to provide in this and installed camera traps (Covert UV562HD, Bushnell
article. This article summarizes all the data on Caracal HD Trophy Camera Camo 119547, and Covert UV552).
currently available in Uzbekistan and specifies the existing Using GPS based on WGS 84 datum, we recorded our
threats, their causes, and conservation measures that car and walking routes as well as the points where we
need to be taken. encountered wild animals.
For interviews with local people, we designed a
Study area questionnaire in Russian. It comprised 10 questions
Ustyurt Plateau lies between the Mangyshlak regarding the sex, age, and occupation of respondents,
Peninsula, Kara-Bogaz-Gol Depression, Aral Sea, their knowledge about the presence of the species in
and Amudarya Delta. The plateau is an important their environs, their attitudes to wild carnivores, and their
transboundary region shared by Uzbekistan, Kazakhstan, perception of interaction with carnivores and of threats
and Turkmenistan. The Kyzylkum Desert covers the to wildlife.
landscape between the Amudarya and Syrdarya rivers, In the villages, we interviewed people who were
which is bordered by the Aral Sea in the north and the outside their homes and asked them to recommend

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Caracal in Uzbekistan Gritsina

Figure 1. Surveyed areas in Uzbekistan between 2011 and 2017. Expedition routes are marked in red, and international border of Uzbekistan
by a solid line.

hunters who were knowledgeable about wildlife. Outside RESULTS

villages, we interviewed all the shepherds that we met
during our transits across the desert. We also collected The analysis of published Caracal records was an
data from the staff of a Bukhara regional nature protection important task for determining routes of our expeditions.
organization and of a compressor station on the Ustyurt Table 1 shows all available information on Caracal
Plateau. We recorded the GPS location of each interview presence in Uzbekistan published between 2000 and
and wrote down the responses of the informants. During 2016.
interviews, we showed them images of wild cats that We did not find any record of Caracal in the area of
possibly occur in the area (Asiatic Wildcat Felis lybica the Surkhan River published later than 2008 (Fig. 1).
ornata, Jungle Cat F. chaus, Sand Cat F. margarita, Manul Our expeditions took place in the southern and central
Otocolobus manul, and Caracal), of species that certainly parts of the Ustyurt Plateau between 2011 and 2015,
do not occur in Uzbekistan (Andean Cat Leopardus and in 2017. In 2011, we travelled in the southwestern
jacobita and Fishing Cat Prionailurus viverrinus), and of Kyzylkum Desert. Between 2014 and 2015, we carried
species that historically occurred in the area (Cheetah out surveys in the northwestern part of this desert, and
Acinonyx jubatus and Tiger Panthera tigris). We noted between 2014 and 2016 in the central part. During the
all the data on Caracal records and attacks of Caracals expeditions, we conducted 1,865.5km transect routes
and other predators on livestock to gain a general by car and 428km on foot and accumulated about 350
understanding of the situation in the region. We assessed observations. Camera traps were set up in 50 locations,
threats to Caracal based on previous literature and results including 14 locations on Ustyurt Plateau on 1,080
of the interview surveys. camera trap days and 36 locations in the Kyzylkum Desert
on 3,741 camera trap days.
We interviewed 104 local people from seven villages

13472 Journal of Threatened Taxa | www.threatenedtaxa.org | 12 March 2019 | 11(4): 13470–13477

Caracal in Uzbekistan Gritsina

Table 1. Caracal records in Uzbekistan published between 2000 and 2016.

Date Location name Type of record Source

Kyzylkum Reserve in northwestern Kyzylkum,
Summer 2000 riparian forests of the Tiksuat Section, bank of Sightings of two individuals. Salimov 2004
the Amudarya River in Ostrov Island section
Zeravshan-Uchkuduk Road, 30km from the A.S. Nuridjanov pers. comm. June
June 2005 An individual killed by a car.
town of Uchkuduk in central Kyzylkum 2005; Gritsina et al. 2016
An individual killed a few lambs in a sheep
herd. Local people killed this individual,
probably in retaliation for sheep losses.
3.iv.2009 Sura Well, 35km from the Kyzylkum Reserve Lim 2009; Lim & Klichev 2009
Later, the dead individual was stuffed and is
now exhibited in the Museum of Nature at
the Kyzylkum State Reserve (Image 1).
2009 Ustyurt Plateau One individual sighted. Abdunazarov 2009
Karnabchul, southwestern Kyzylkum, 8km
2011 One individual sighted. Marmazinskaya & Mardonova 2016
southeast of Igrichi
2013 16km southeast of Igrichi One individual sighted. Marmazinskaya & Mardonova 2016

2014 10km northeast of Igrichi One individual sighted. Marmazinskaya & Mardonova 2016

July 2015 Northwestern chink of lake Sarykamysh One individual captured on a camera trap. Bykova et al. 2015

and 220 shepherds in the desert, including 278 in group

sessions with 2–12 people and the remaining individually.
Respondents were shepherds, hunters, farmers,
housewives, village elders, and local authorities aged
between 14 and 73 years.
Caracal occurrence reported by local people and
camera trap records obtained during our field surveys
are summarised in Tables 2 and 3. Information on
observations and killed Caracals was published in Russian
by Gritsina (2012), Marmazinskaya et al. (2012), and
Gritsina et al. (2016).
Of the 324 respondents, 28 people (8.5% of total)
pointed to the facts of killing Caracal in the past 17
years, either intentionally or as a result of trapping in
Image 1. Stuffed Caracal exhibited in the Museum of Nature at the
snares set up for wild predators in general. Local people
Kyzylkum State Reserve in Uzbekistan. © N. Marmazinskaya.
are completely unaware that Caracal is a law-protected
species. Only 58 people identified Caracal correctly from
the images. We recorded that the local population were
on the whole less aggressive to Caracal than to Wolf. village. Later, we found the carcass of a young female in
Wolf is considered the main threat to livestock. Eighteen the place indicated by the local people (Image 2). The
people from the respondents were aggressive towards cat was caught in a leg-snare near a sheep pen. Also, the
wild cats and believed that they caused significant villagers informed us of another Caracal killed by herders
damage to sheep and chicken. in 2012, which used to attack lambs in the calving period,
Though we did not find any Caracal skins at local killing up to 10 lambs at one time but leaving them
markets, we did find the skins and products of other small uneaten. This information was confirmed by herders
wild cats, mostly that of Asiatic Wildcat (14 skins and 21 from the village of Jankeldy, 14km from Kalaata. They
hats) and Jungle Cat (four skins and two hats). told us that people from Kalaata killed three individual
Caracals between 2012 and 2013. An official of the
Kyzylkum Desert Bukhara Regional Department of the State Committee for
In the Kyzylkum area, we interviewed 245 people in Nature Protection, F. Salimov, observed Caracals several
five villages and in more than 40 locations. Villagers of times near the above-mentioned village of Jankeldy and
Kalaata in central Kyzylkum reported a Caracal killed by on the shore of lake Zamonbobo in the past 10–15 years.
herders on 20 March 2014 about 20km southwest of the He accounted for the sighting of one individual “several

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Caracal in Uzbekistan Gritsina

Table 2. Caracal records collected in Kyzylkum Desert during expeditions between 2011 and 2017.

Date Location name Type of record

Between 2000 and 2005 Village of Jankeldy and lake Zamonbobo Several sightings of individuals.

Autumn 2006 Surroundings of Turtkul One individual sighted.

March 2011 Shore of lake Dengizkul Caracal footprints found by M. Gritsina.

Between 2012 and 2013 Surroundings of Kalaata Village Three individuals killed by herders.
One individual sighted by F. Salimov, an official of the
October 2013 Near the town of Gazli
Bukhara Regional Committee for Nature Protection.
Autumn 2013 Lake Dengizkul One individual sighted by F. Salimov.

20.iii.2014 20km southwest of Kalaata One individual killed by herders.

Image 2. The carcass of a young female found in Kyzylkum Desert in
Uzbekistan on 20.iii.2014. © M. Gritsina. Image 3. Caracal captured by a camera trap on 22.ix.2017 on Ustyurt
Plateau in Uzbekistan. © M. Gritsina & D. Nuridjanov.

dozen kilometres” from the village of Turtkul in 2006. In In autumn 2010, herders killed an individual that took
October 2013, he also saw a Caracal by day near the town five lambs from a sheep pen near Churuk Well. According
of Gazli, and another one in the autumn of the same year to local people, Caracals frequent the well area. In the
not far from lake Dengizkul (Gritsina et al. 2016). same winter, a hunter from Kubla-Ustyurt killed a Caracal
near the village. This was not the hunter’s first Caracal,
Ustyurt Plateau as in 1976 this person had killed another one (Gritsina et
In the Ustyurt Plateau area, we interviewed 79 people al. 2016).
in four villages. On 22 September 2017 at 07.06h, one individual was
Footprints of Caracals were recorded in seven places in captured in a camera trap on the eastern chink of Ustyurt
the southern and central parts of the Karakalpak portion Plateau, opposite the western bank of Sarykamysh Lake
of the Ustyurt Plateau (Marmazinskaya et al. 2012). (Image 3).
In 2014, a hunter from the village of Kubla-Ustyurt Thus, in the years from 2000 to 2017, Caracal was
killed a Caracal whose skin still remains in the village. We documented in at least 27 localities in Uzbekistan,
could not estimate the number of animals caught. In late including 13 dead individuals and 14 live ones. At least
April to early May 2015, the manager of the Kubla-Ustyurt 11 individuals were killed intentionally and one was run
Compressor Plant encountered an individual 5km from over by a car. Of the dead and live animals, the presence
the Raushan Ascent on the chink, when he was returning of Caracal in the area was recognised by its footprints in
to the village from Kungrad during daytime. The Caracal eight cases.
was lying on the road and only when the driver stopped
the car and honked several times did the animal rise
unhurriedly and go towards the chink (Gritsina et al.
2016).

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Caracal in Uzbekistan Gritsina

Table 3. Caracal records collected in Ustyurt Plateau during expeditions between 2011 and 2017.

Date Location name Type of record

1976 Kubla-Ustyurt Village One individual killed.

Winter 2005 40km from Kirkkiz Village towards the chink of the Aral Sea One individual killed by a hunter.
One individual observed by workers of Microwave Relay
May 2010 Near Cape Aktumsyk in the Aral Sea village Karateren
Station 23.
Autumn 2010 Churuk Well Village One individual killed by herders.

Winter 2010 Amudarya River near the city of Urgench One individual killed.

Winter 2010 Surroundings of Kubla-Ustyurt Village One individual killed by a hunter.

Winter 2011 Between the cities of Kungrad and Muynak, Aral Sea area One individual caught in a trap.
Southern and central parts of the Karakalpak portion of the
May 2012 Caracal tracks in seven places identified by the survey team.
Ustyurt Plateau
Spring 2013 Near lake Sudochye One individual sighted by a respondent.

Winter 2014 Village of Kubla-Ustyurt One individual killed.

Between 2000 and 2014 Eastern chink of the Aral Sea village Individuals often caught in traps set for Wolves.
Late April to early May
5km from the Raushan Ascent on the chink village One individual sighted by respondent.
2015
22.ix.2017 Near Sarykamysh Lake on the eastern chink (Image 3) One individual captured on a camera trap.

DISCUSSION authors demonstrated that predation by Caracal in the

vicinity of West Coast National Park depended on the
Threats to Caracal in Uzbekistan availability of wild prey and husbandry techniques;
Analysis of interview surveys revealed that all predation on small livestock was limited to the lambing
respondents with the exception of local authorities did season in spring when rodent densities decreased.
not know that Caracal is included as a protected species Although Caracal is a non-migratory species, it covers
in the Red Data Book of Uzbekistan (Abdunazarov 2009). long distances of up to 90km during dispersals and in
Five Caracals were hunted in retaliation for killing search of food and mates (Avenant & Nel 1998). For this
livestock. Another five individuals were killed intentionally reason, the fence along the border between Uzbekistan,
by poachers, while two others were caught in traps set for Turkmenistan, and Kazakhstan poses a certain threat.
other species. Caracal habitats are interspersed by heavy-traffic asphalt
The main reason for negative interactions between roads and a network of dirt roads. In view of the Caracal
local people and mammalian predators including Caracal killed in a traffic accident, the road network passing
is the lack of knowledge of the people on how to properly through Caracal habitat is a potential threat. Another
guard their small livestock. They use traditional, long- potential threat is railways, with a few tracks crossing the
established, outdated grazing and breeding methods. feline’s habitat.
The pens built by herders have low walls without solid
roofs. Enclosures on grazing grounds are mostly made of Ecologic factors possibly impacting Caracal distribution
thin mesh tied to ordinary wooden sticks and are roofless. Abdunazarov (2009) suggested that the quantity of
In spring and summer, sheep and goats graze and breed available food impacts Caracal distribution and population
without any structures for their protection. Some herds size. We do not have any data on Caracal’s food resources
are escorted by a few dogs Canis familiaris, usually of and its dependency on prey density. A specialized study is
Tazy breed, but these dogs are rarely trained to guard required to assess these ecologic variables.
and protect the livestock from predators. In addition, Bekenov & Kasabekov (2010) suggested that cold
they are poorly fed by herders and therefore hunt small and snowy winters have a negative effect on Caracal
wildlife. Due to these circumstances, small livestock is a population because the species is not adapted to low
helpless and easy prey for predators. We did not find any temperatures and thick snow cover. Data on the death
evidence of dogs attacking Caracal. of Caracals as a result of cold and snowy winters in
Results of studies conducted in South Africa indicate Uzbekistan for the last 15 years are lacking.
that Caracal preys foremost on rodents (Grobler 1981; Some parts of Caracal habitats are remote and poorly
Palmer & Fairall 1988; Avenant & Nel 2002). The latter accessible, which makes patrolling by nature protection

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Caracal in Uzbekistan Gritsina

agencies problematic and ineffective. The main obstacles natural environment if there is a sanctioned quota and
are insufficient funding, lack of technically-skilled human a permission approved by the Cabinet of Ministers and
resource, and the absence of anti-poaching brigades. the Academy of Sciences of the Republic of Uzbekistan,
Owing to these circumstances, environment protection in compliance with the Law of the Republic of Uzbekistan
measures are currently not implemented. On Protection and Use of Fauna (No. 408, 19 September
2016) (Mirzeev 2016), and the Decree by the Cabinet of
Ecologic factors favouring the conservation of Caracal in Ministers of the Republic of Uzbekistan On Regulation of
Uzbekistan Use of Biological Resources and On Permission Issuance
Caracal occurs only in the desert regions of Procedures in the Use of Natural Resources (No. 290, 20
Uzbekistan. The climate in deserts is characterised by low October 2014) (Mirzeev 2014).
winter and high summer temperatures alongside other Caracal habitats are conserved in two protected areas,
hard natural conditions such as strong winds, poorly Kyzylkum State Reserve (IUCN Ia) covering 10,311ha, and
accessible areas, and deficiency of fresh water. Some Saigachiy Wildlife Sanctuary (IUCN Ib) with 219,800ha.
of these areas were brought into use during the Soviet The Kyzylkum State Reserve is a well-protected area,
period, but most of them remain unpopulated and are mostly because it is situated next to the state border with
rarely visited by people. Among these unpopulated Turkmenistan. Visits to the reserve and its neighbourhoods
and unvisited territories are the southern portion of the are only allowed to local residents and holders of a special
Ustyurt Plateau, the section of northwestern Kyzylkum permit. The protection of the Saigachy Wildlife Sanctuary
next to the border with Kazakhstan, and some parts founded in 2016 on the Ustyurt Plateau is unsatisfactory
of central Kyzylkum. These conditions determined by because of its vast area that is difficult to manage and is
ecogeographic and socioeconomic factors leave many remote from settlements. There are 10 rangers who have
suitable habitats intact and are therefore favourable for four vehicles at their disposal but insufficient funding
Caracal conservation in Uzbekistan. (fuel and daily allowances) for the continued patrol
of the protected area. In addition, there are several
Socioeconomic factors favouring the conservation of smaller protected areas that include Caracal habitats,
Caracal in Uzbekistan such as the Sudochy Ornithological Wildlife Sanctuary
In the Soviet period, many desert areas were on the western side of the Amudarya Delta (50,000ha),
intensively used for large-scale construction of roads, Dengizkul Wildlife Sanctuary (50,000ha), Karakir Wildlife
boring of wells, and establishment of rural communities. Sanctuary (30,000ha), Sichankul Wildlife Sanctuary in the
By now, some villages built in the Soviet times are fully or Sundukli sands, southwestern Kyzylkum (70,375ha), and
partly abandoned, and most areas are depopulated due Karnabchul Wildlife Sanctuary in southwestern Kyzylkum
to the migration of people to towns within the country (25,000ha). All these reserves must be considered ‘paper
or abroad. parks’ without real protection and management.
Some of these territories lie close to the borders with We think it is necessary to implement the following
Kazakhstan and Turkmenistan, and special permissions activities for Caracal conservation in Uzbekistan: 1)
must be obtained from border police for visiting these strengthen the protection of Caracal habitats in the
areas. existing protected areas and establish new protected
Formerly, there were a number of small villages in the areas in the Kyzylkum Desert and on the Ustyurt Plateau,
southern part of the Ustyurt Plateau where people used 2) raise local people’s awareness and knowledge and
existent wells as water sources and lands as pastures for inform them about the need for Caracal research and
their livestock. Currently, no villages exist here any longer protection, 3) improve the protection of livestock
and livestock grazing is not practised. by establishing predator-proof corrals and herding
These demographic factors lower human pressures practices, and 4) raise the awareness and knowledge
upon wildlife and favour the conservation of biodiversity, of nature conservation agencies, hunters, and hunters’
including that of Caracal, in the region. associations. Much more attention should be paid to
scientific research on Caracal, particularly in terms of
Existing conservation measures and measures that need estimating its current status and range and of developing
to be developed a functional population monitoring system. It is also
Hunting Caracal is prohibited, as it is included in important to design, approve, and implement an action
the Red Data Book of Uzbekistan (Abdunazarov 2009). plan for Caracal conservation in Uzbekistan and to secure
Species listed therein can only be taken out of their support from conservation authorities.

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Caracal in Uzbekistan Gritsina

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Threatened Taxa

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Journal of Threatened Taxa | www.threatenedtaxa.org | 12 March 2019 | 11(4): 13478–13491
Communication

Small cat surveys: 10 years of data from Central Kalimantan,

Indonesian Borneo
ISSN 0974-7907 (Online) Karen Anne Jeffers 1 , Adul 2 & Susan Mary Cheyne 3
ISSN 0974-7893 (Print)

PLATINUM
1–3
Borneo Nature Foundation, Jalan Bukit Raya Induk No. 17, Palangka Raya, 73112, Central Kalimantan, Indonesia.
3
Oxford Brookes University, Department of Social Sciences, Faculty of Humanities and Social Sciences, Gibbs Building,
OPEN ACCESS Headington Campus, Oxford, OX3 0BP, UK.
1
[email protected], 2 [email protected], 3 [email protected] (corresponding author)

Abstract: We present an update on the photographic detections from camera traps and the activity patterns of Borneo’s four small
cats, namely, Sunda Leopard Cat Prionailurus javanensis, Flat-headed Cat P. planiceps, Marbled Cat Pardofelis marmorata, and Bay Cat
Catopuma badia, at two sites in Central Kalimantan, Indonesia. Camera trap survey data of 10 years (2008–2018) from the first site in
Sebangau provide details about the temporal partitioning of these small cats from each other but overlap with Sunda Clouded Leopard
Neofelis diardi. The activity of Flat-headed Cat was higher after midnight and that of Leopard Cat at night with no clear preference before
or after midnight. The Marbled Cat is predominantly diurnal, but the remaining three cats have flexible activity periods. While limited
data are available from Rungan, the second site, we confirmed the presence of all four small cat species found on Borneo, though we have
insufficient data to comment on the Bay Cat. The cat sightings, however, are intermittent and may reflect the unprotected status of this
forest. Leopard Cats appear relatively unaffected by habitat disturbance based on encounter rates on camera traps. Conservationists,
both NGOs and the government, must pay particular attention to specialists like Flat-headed Cats and Bay Cats when assessing habitat
suitability for long-term cat conservation.

Keywords: Activity patterns, camera traps, Catopuma badia, diversity, felids, fire, peat-swamp forest, Pardofelis marmorata, Prionailurus
javanensis, Prionailurus planiceps.

Bahasa Indonesia Abstract: Kami menyajikan pembaruan pada pendeteksian fotografi dari perangkap kamera dan pola aktivitas empat
kucing kecil Borneo yaitu Kucing Kuwuk Prionailurus bengalensis, Kucing Batu Pardofelis marmorata, Kucing Tandang Prionailurus planiceps
dan Kucing Merah Cat Catopuma badia di dua lokasi di Kalimantan Tengah, Indonesia. Data survei perangkap kamera 10 tahun (2008–
2018) dari situs pertama di Sebangau memberikan rincian tentang partisi sementara kucing kecil ini dari satu sama lain tetapi tumpang
tindih dengan Macan Dahan Neofelis diardi. Aktivitas kucing kepala datar lebih tinggi setelah tengah malam, dan Kucing Kuwuk di malam
hari tanpa preferensi yang jelas sebelum atau setelah tengah malam. Kucing Batu didominasi diurnal, tetapi ketiga kucing memiliki periode
aktivitas yang fleksibel. Sementara data terbatas tersedia dari situs kedua (Rungan), kami telah mengkonfirmasi keberadaan keempat
spesies kucing kecil yang ditemukan di Borneo, meskipun kami tidak memiliki cukup data untuk mengomentari Kucing Merah. Namun,
penampakan kucing berselang-seling dan mungkin mencerminkan status hutan yang tidak terlindung. Kucing Kuwuk kembali muncul
relatif tidak terpengaruh oleh gangguan habitat berdasarkan pada tingkat pertemuan pada perangkap kamera. Konservasionis (LSM dan
pemerintah) harus memberi perhatian khusus kepada spesialis misalnya Kucing Tandang dan Kucing Merah ketika menilai kecocokan
habitat untuk konservasi kucing jangka panjang.

DOI: https://doi.org/10.11609/jott.4466.11.4.13478-13491 | ZooBank: urn:lsid:zoobank.org:pub:0EA7C81F-E24F-4E17-B8F8-E45620396EEB

Editor: Angie Appel, Wild Cat Network, Germany. Date of publication: 12 March 2019 (online & print)

Manuscript details: #4466 | Received 02 August 2018 | Final received 04 February 2019 | Finally accepted 27 February 2019

Citation: Jeffers, K.A., Adul & S.M. Cheyne (2019). Small cat surveys: 10 years of data from Central Kalimantan, Indonesian Borneo. Journal of Threatened Taxa
11(4): 13478–13491; https://doi.org/10.11609/jott.4466.11.4.13478-13491

Copyright: © Jeffers et al. 2019. Creative Commons Attribution 4.0 International License. JoTT allows unrestricted use, reproduction, and distribution of this article
in any medium by adequate credit to the author(s) and the source of publication.

Funding: SMC was funded through a grant to David W. Macdonald from the Kaplan family and by the Clouded Leopard Project. SMC and DWM’s work on Bornean
felids is part of the WildCRU/Panthera collaboration and was carried out within the BNF-CIMTROP multi-disciplinary research project in the northern Sabangau
forest, Central Kalimantan, Indonesia. SMC gratefully thanks the Centre for the International Cooperation in Management of Tropical Peatlands (CIMTROP) for
sponsoring her research and providing invaluable logistical support. We gratefully acknowledge the contribution of all the researchers who assisted with the
project. SMC and KAJ thanks the Indonesian Ministry for Science and Technology for providing research permissions for this work.

Competing interests: The authors declare no competing interests.

For Author details & Author contribution see end of this article.

Acknowledgements: We thank CIMTROP, University of Palangka Raya and University Muhammadiyah Palangka Raya, for their collaboration in making this work
possible. We thank the Indonesian Ministry of Science and Technology (RISTEK) and Director General of Nature Conservation (PHKA) for permission to carry out
research in Indonesia. 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, The Clouded Leopard Project of Point Defiance Zoo and Aquarium, and Fresno Chafee Zoo USA. Three anonymous
reviewers provided invaluable comments on the previous drafts of this manuscript.

13478
Small cats survey data from Central Kalimantan, Indonesia Jeffers et al.

INTRODUCTION in the Rungan Forest, which covers about 1,440km2

between the Kahayan and Rungan rivers. The forest
In the absence of Tiger Panthera tigris, Borneo’s is a lowland forest mosaic comprising peat-swamp,
cats represent a fascinating guild, and all are in need of ‘kerangas’ (heathland) and the dominant canopy trees are
conservation attention. Five species occur on Borneo: Palaquium sp. (nyatoh), Syzygium sp. (jambu) and Shorea
the Sunda Clouded Leopard Neofelis diardi (Endangered (meranti), (Dipterocarpaceae family). BNF and the Wildlife
(EN) on the IUCN Red List; Hearn et al. 2016c), the Bay Conservation Research Unit (WildCRU) of the University of
Cat Catopuma badia (EN and endemic to Borneo; Hearn Oxford initiated the Sebangau Felid Project in May 2008,
et al. 2016a), the Flat-headed Cat Prionailurus planiceps and BNF initiated the Rungan work in 2016.
(EN; Wilting et al. 2016a), the Marbled Cat Pardofelis
marmorata (Near Threatened; Ross et al. 2016a), and the
Sunda Leopard Cat Prionailurus javanensis (Least Concern; MATERIAL AND METHODS
Ross et al. 2016). They are all protected under Indonesian
law (P.20/MENLHK/SETJEN/KUM.1/6/2018). As they are Since 2008, a total of 210 camera locations were
rare and elusive, it is difficult to study them in the wild. surveyed across both forest areas with an average of 30
Thus, there is limited knowledge about their ecology units in each forest area at any one time. Between 2008
despite increased scientific interest. and 2012, cameras were set up in pairs in Sebangau and
Since 2008, Borneo Nature Foundation (BNF) and subsequently as single units. In the Rungan site, cameras
University of Palangka Raya, Centre for the International were all set up as single units. Cameras were set in a
Management of Tropical Peatlands (UPR-CIMTROP), stratified random survey design. Cameras were placed
have been conducting long-term monitoring of the 500m to 1,000m apart and were in each location for a
Sunda Clouded Leopard Neofelis diardi in Indonesia and, minimum of six months; some cameras were in the same
specifically, N. d. borneensis in Central Kalimantan. During location since May 2008. Locations were selected to cover
this time, we also opportunistically obtained images of a range of habitats and disturbances within the forests,
three of the four small cats in the Sebangau catchment avoiding streams and slopes wherever present. Camera
area. Given the disturbed mosaic nature of Sebangau, traps were placed along established human-made trails
we also hope to determine which areas are unsuitable for (more than four years old) and, where possible, watering
small cats, either due to habitat changes and/or due to areas, to maximise the success rate of photographic
human disturbance. Live Leopard Cats are more common captures. A combination of camera models were used,
in Kalimantan markets for sale as pets than Sunda Clouded including Cuddeback Expert®, Cuddeback Capture IR®
Leopards, and their skins are more often found in homes (Cuddeback Digital, Non-Typical Inc, WI, USA) Maginon,
(Rabinowitz et al. 1987). Far less is known about small cat Crenova, and Bushnell. Cameras were checked every
movements, habitat preferences, seasonal movements, 40 days when batteries were changed and SD cards
breeding patterns, and effects of anthropogenic exchanged. Data were managed in a custom Microsoft
disturbance across their range. These data are particularly Access database. Active behaviour times were calculated
lacking from tropical peat-swamp forests. We present using the kernel density method (‘href’ bandwidth for
here updated information on temporal activity pattern kernel smoothing; Ridout & Linkie 2009; Meredith &
of four small cats from the Central Kalimantan region of Ridout 2016) to account for average dawn and dusk times
Indonesia. in the sites, which are situated almost on the equator (for
more information on the Sebangau study site, see Cheyne
Study areas & Macdonald 2011; Cheyne et al. 2016b). Detection rate
The Sebangau catchment, Central Kalimantan, was estimated as number of detections/100 trap nights.
Indonesia (Fig. 1), is a peat-swamp forest (mixed-swamp Weather data were collected daily at each research site
forest sub-type) covering an area of ~5,600km2. This and fire data was obtained from the Indonesian Agency
study took place in the 50km2 research forest located in of Meteorology, Climatology, and Geophysics. A 30-min
the northeast. The area was logged under a concession interval between photos of the same species was used to
system between 1991 and 1997 followed by illegal logging determine if photos of were an independent event at the
between 1997 and 2004. The site is at an altitude of about same location and date.
10m. The area was significantly affected by the forest fires
that impacted Indonesia in 2015.
The second research site was established in 2016

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Small cats survey data from Central Kalimantan, Indonesia Jeffers et al.

Figure 1. Study sites: circle - Sebangau; star - Rungan.

RESULTS in 2008, there was a decline in the detections of Marbled

Cats and Flat-headed Cats (Fig. 2). From 2014 to July
The number of camera trap (CT) stations at each site 2018, there was an average of 3.9 independent Marbled
varied annually due to broken units (Table 1). Cat images/month (min=0, max=16). No Flat-headed Cats
Detailed descriptions of CT locations are in Appendices were recorded by camera traps between January 2014 and
1 & 2, including descriptions of the microhabitats, February 2018 (Fig. 2), which coincided with a significant
number of trap nights, number of detections, and careful fire event from September to November 2015.
descriptions of the setup around the CT stations. CTs were Of the three small cat species, the Sunda Leopard Cat
placed in different habitats across the two sites (Table 2). is predominantly nocturnal with no clear preference for
The small cats were recorded by 56 of 83 camera traps time of night. Flat-headed Cats also showed nocturnal
in Sebangau (67%) and by 16 of 37 camera traps in Rungan activity but with a slight preference for post-midnight
(43%) (Table 3). hours. Marbled Cats are strongly diurnal. Interestingly,
All small cats in Sebangau were photographed in all 65% of 115 nocturnal records (18.00–05.59 h) of Sunda
main habitat types in the interior forest, <20m from the Clouded Leopards were between 01.00h and 05.59h (Fig.
forest edge and in disturbed areas. All four small cats in 3), thus overlapping with the preferred active time for Flat-
Rungan were recorded in the interior forest; only the Flat- headed Cats.
headed Cat was recorded near the lake.
Rungan
Sebangau Three of the small cats were confirmed in Rungan in
Since the cameras were first placed in May 2008, we the first few months of the study but it took 12 months
captured 157 independent images of Sunda Clouded to confirm the presence of the Flat-headed Cat (Table 4).
Leopards (Image 1), but only 109 of Sunda Leopard Cats,
54 of Marbled Cats (Image 2), and 33 of Flat-headed Cats Bay Cat
(Image 3). Compared to the average detection rates of With over eight years of long-term camera trap
small cats since the inception of the camera trap study surveying in the peat-swamp forest of the Sebangau

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Table 1. Number of camera trap stations per year in the study site. Table 2. Number of camera trap (CT) locations at each site (NA -
habitat type not present in the study site).
Year Sebangau Rungan

2008 40 0 Habitat type Sebangau Rungan

2009 40 0 Burned area 3 NA

2010 40 0 Kerangas/ heath NA 30

2011 36 0 Low interior forest 1 4

2012 34 0 Mixed swamp forest 74 3

2013 30 0 Tall interior forest 5 NA

2014 25 0 Total CT locations 83 37

2015 30 0 Total trap nights 65,261 14,642

2016 30 36

2017 25 50

2018 28 30 catchment, the Bay Cat was not detected, and ongoing
work suggests that it is not found in peat-swamp forests.
The Bay Cat was confirmed in mosaic heath/ peat-swamp
forest habitat for the first time (Sastramidjaja et al. 2015;
Cheyne et al. 2016a, 2017). Through the use of camera
traps, we present new location information on the
distribution of Bay Cat in Kalimantan. This new location
is approximately 64km southeast outside the range
depicted by Hearn et al. (2016a). Our record of Borneo
Bay Cat from the new habitat (heath/ peat-swamp forest)
warrants further surveys in different habitat types to fully
understand Bay Cat distribution and ecologic needs.

DISCUSSION

The small cats are appearing evenly across the habitat

types in both Sebangan and Rungan, with the exception of
the Bay Cat that likely does not exist in deep ombrogenous
peat-swamp forest (Sebangau). Additionally, we have
evidence of breeding in Flat-headed Cat and Marbled Cat
in Sebangau (images of kittens) (Images 4 & 5).
Flat-headed Cats have a more irregular capture rate and
though they are active throughout the day, more captures
Figure 2. Mean detection of small wild cat species from the inception are obtained at night and therefore they are predominantly
of the camera trap surveys in Sebangau since 2008. nocturnal. Leopard Cats have a more regular capture rate

Table 3. Camera trap (CT) locations with detections and non-detections of small cats with mean occupancy estimates (Ψ) in the study sites.
NA indicates occupancy cannot be calculated due to no small cats being photographed at these locations.

Sebangau Rungan
Number of CT Number of CT
Ψ Ψ
locations locations
Leopard Cat 20 21.05 10 26.32

Marbled Cat 20 13.68 2 10.26

Flat-headed Cat 16 11.58 3 6.50

Bay Cat 0 0 1 1.28

No small cats 36 NA 23 NA

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Image 1. Sunda Leopard Cat Prionailurus javanensis in Sebangau Forest, Central Kalimantan, Indonesian Borneo. © Borneo Nature Foundation,
28 July 2008.

Image 2. Marbled Cat Pardofelis marmorata in Sebangau Forest, Central Kalimantan, Indonesian Borneo. © Borneo Nature Foundation,
20 September 2013.

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Small cats survey data from Central Kalimantan, Indonesia Jeffers et al.

Image 3. Flat-headed Cat Prionailurus planiceps in Sebangau Forest, Central Kalimantan, Indonesian Borneo. © Borneo Nature Foundation, 22
February 2013.

Table 4. Capture rate of small cat species from the inception of the camera trap surveys in Rungan between June 2016 and May 2018.

Jun–Aug Sep–Nov Dec 2016–Feb Mar–May Jun–Aug Sep–Nov Dec 2017–Feb Mar–May
2016 2016 2017 2017 2017 2017 2018 2018
Flat-headed Cat 0 0 0 0 1 1 0 0

Sunda Leopard Cat 2 2 2 1 0 0 0 0

Marbled Cat 1 4 0 0 1 0 0 0

Bay Cat 1 0 0 0 0 0 0 0

and appear to be active both during the day and night, suggested to be poor or marginal habitat for Sunda
though they appear to avoid the hottest time of the day Leopard Cat (Mohamed et al. 2016), but our work suggests
(11.00–13.00 h). Marbled Cats have a regular capture rate that Sunda Leopard Cat are far more common (Cheyne &
with the majority of images taken during the day (05.00– Macdonald 2011; Adul et al. 2015; Cheyne et al. 2016b).
16.00 h), suggesting they are diurnal. There is only one The infrequent capture of the small cats in both sites
image of a Bay Cat taken at 11.17h. These data are similar is likely an artefact of the placing of the cameras (±1km
to those of Hearn et al. (2018), though these authors did apart) to focus on the wide-ranging Sunda Clouded
not obtain sufficient images of Flat-headed Cats to make a Leopard. By moving the cameras closer (±500m) we
detailed analysis. hope to determine the population density for the small
Peat-swamp and associated lowland wetlands are cats, determine if the Bay Cat is indeed absent from this
postulated to be an important habitat for Flat-headed Cats forest, and to continue our monitoring of the Sunda
(Cheyne et al. 2009; Wilting et al. 2010, 2016b; Cheyne Clouded Leopard population. The long period of time
& Macdonald 2011; Adul et al. 2015). Marbled Cats are required to obtain images of small cats, possibly due to
not believed to frequent roads or plantations (Hearn et al. the placement of the cameras targeting Sunda Clouded
2016c) and prefer intact forests, though data are lacking Leopard, highlights the importance of long-term data and
on this cat (Rustam et al. 2016). monitoring to avoid false-negative presence data. Sunda
Peat-swamp and associated lowland wetlands were Leopard Cat is the most commonly recorded species in the

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Small cats survey data from Central Kalimantan, Indonesia Jeffers et al.

restore. These data could be an artefact of survey effort (a

high number of broken cameras) or a genuine reflection of
this species behaviour.
Large parts of the peat-swamp are naturally seasonally
flooded for up to nine months per year. The flooded nature
of Sebangau does not always prevent the animals using
the ground; indeed we have evidence of male Orangutans
Pongo pygmaeus wading through water (Ancrenaz et al.
2014). Keeping a selection of camera trap locations the
same over several months or years allows for variations in
detection to be accounted for; given the regular flooding of
the forest, it is likely that the wildlife is accustomed to this.
We do notice animals using our boardwalks more regularly
in the wet season. Peatlands and associated forest fires
are a crucial conservation concern in Kalimantan (Gaveau
et al. 2016; Miettinen et al. 2017). This is especially true
during dry years such as in 2015 when a strong El Niño
event led to particularly dry conditions. From August to
November of that year, MODIS satellites detected over
50,000 fire hotspots in Kalimantan, 53% of which were on
peatland (Gaveau et al. 2016; Miettinen et al. 2017). Since
2006, 17.35% of forest in the core Sebangau research area
burned down (9.63% in 2015 alone—5.3km2 of 55km2). Of
particular threat to the Flat-headed Cat are peat drainage
and drying out due to logging canals, the loss of permanent
water, and increased hydrologic instability (Page et al.
2009; Vanthomme et al. 2013).

Conservation impact
This work represents the first-ever comprehensive
and long-term survey of small cats in Central Kalimantan.
There is a severe lack of data on these species in non-
protected or small forest areas that may also contain
viable populations. It is crucial to remember that, while
Figure 3. Kernel density estimates of activity patterns of species using
alpha of 0.1 smoothing parameter: a - Flat-headed Cat; b - Marbled these surveys indicate the continued presence of these
Cat; c - Sunda Leopard Cat. cats, habitat loss, wildlife trade, and likely presence of
populations in non-protected areas means that more work
is needed to understand the impacts of anthropogenic
study site. Marbled Cat is hard to study and, as many are activities on these cats. As detailed in Appendix 1, this
arboreal, having cameras mainly on the forest floor means project provides extensive and detailed data about many
we could be missing out on key aspects of their behaviour. wildlife species in Sebangau and Rungan forests in addition
Flat-headed Cat is a wetland specialist and prefers forests to the cats—an additional 7,959 images (2,765 videos) of
with water (Wilting et al. 2010). Its diet likely consists of 74 species. Of these, two are IUCN Red Listed as Critically
fish, frogs, and small mammals, and it may fill a niche on Endangered, five as Endangered, 14 as Vulnerable, 12 as
Borneo filled by the Fishing Cat Prionailurus viverrinus Near Threatened, and 41 as Least Concern.
elsewhere in Asia (Iwaniuk et al. 2001). Due to this
dependence on wetlands, we think that the devastating
fires of 2015 may have severely impacted the Flat-headed
Cat population, pushing it away from fire-affected areas.
Our preliminary results suggest that Flat-headed Cats are
returning to these areas, which BNF is actively working to

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Small cats survey data from Central Kalimantan, Indonesia Jeffers et al.

Image 4. Flat-headed Cat Prionailurus planiceps kitten

Image 5. Marbled Cat Pardofelis marmorata and kitten

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Appendix 1. Summary of camera trap (CT) locations in Sebangau, Central Kalimantan Indonesian Borneo. LC - Leopard Cat, MC - Marbled Cat,
FHC - Flat-headed Cat.

Additional habitat
No. of CT days Location of camera Habitat class Altitude (m) LC MC FhC
information
372 T 1.3E x TY 2015 Burned area Interior forest 16–20 X    

375 T 1B East 975m 2015 Burned area Forest edge (<20m) 16–20   X  

180 T SC East x TY 2015 Burned area Forest edge (<20m) 16–20 X    

1475 JE1 Mixed swamp forest Forest edge (<20m) 16–20     X

2399 Km2 x Railway Mixed swamp forest Interior forest 16–20      

2399 Km3 x Railway Mixed swamp forest Interior forest 16–20 X    

2399 Km4 x Railway Mixed swamp forest Interior forest 16–20 X    

2447 Old Railway 400m Mixed swamp forest Disturbed logging railway 16–20   X X

241 Old Railway x T2E Mixed swamp forest Disturbed logging railway 16–20      

724 Old Railway x TX Mixed swamp forest Disturbed logging railway 16–20   X  

849 OR x T0.8E 2013 Mixed swamp forest Disturbed logging railway 16–20      

346 OR1150m Mixed swamp forest Disturbed logging railway 16–20      

382 Ottercam T1B Canal Mixed swamp forest Interior forest 16–20      

375 P.Jelotung x TD 2015 Mixed swamp forest Interior forest 16–20   X X

52 Railway 1450m Mixed swamp forest Disturbed logging railway 16–20      

835 Secret Transect Mixed swamp forest Forest edge (<20m) 16–20   X  

262 T 0.4 East End 2015 Mixed swamp forest Interior forest 16–20   X  

740 T 0.4 X TD 2015 Mixed swamp forest Interior forest 16–20 X    

374 T 0.8 x TB 2015 Mixed swamp forest Interior forest 16–20 X X  

465 T 0.8 x TE 2015 Mixed swamp forest Interior forest 16–20 X   X

372 T 0.8E x ORW 2015 Mixed swamp forest Interior forest 16–20      

2395 T 1.6 x P.owa-owa 2015 Mixed swamp forest Interior forest 16–20   X X

436 T 1.6 x T E Mixed swamp forest Interior forest 16–20   X  

372 T 1.6E x TW 2015 Mixed swamp forest Interior forest 16–20   X  

375 T 16 x TB 2015 Mixed swamp forest Interior forest 16–20      

375 T 1A x Railway 2015 Mixed swamp forest Disturbed logging railway 16–20      

375 T 2 700m 2015 Mixed swamp forest Interior forest 16–20   X  

375 T 2 x TE 2015 Mixed swamp forest Interior forest 16–20      

372 T 2E x ORW 2015 Mixed swamp forest Disturbed logging railway 16–20     X

372 T 2E x TX 2015 Mixed swamp forest Interior forest 16–20      

649 T 2km 700m Mixed swamp forest Interior forest 16–20      

2450 T CC 25m di atas pohon Mixed swamp forest Canopy 10m 10 X    

802 T DD 400m Mixed swamp forest Forest edge (<20m) 16-20 X    

248 T FF 125m 2015 Mixed swamp forest Forest edge (<20m) 16–20 X   X

261 T SC 1412m 2015 Mixed swamp forest Forest edge (<20m) 16–20      

64 T SC 530m 2016 Mixed swamp forest Forest edge (<20m) 16–20      

965 T SC 610m 2015 Mixed swamp forest Forest edge (<20m) 16–20     X

841 T SC East 275m Mixed swamp forest Forest edge (<20m) 16–20   X  

935 T SC East 275m 2015 Mixed swamp forest Forest edge (<20m) 16–20     X

843 T.Secret 1412m Mixed swamp forest Forest edge (<20m) 16–20   X  

880 T.Secret 610m Mixed swamp forest Forest edge (<20m) 16–20   X  

566 T0 950m di atas pohon Mixed swamp forest Canopy 10m 10   X  

436 T0 x T F Mixed swamp forest Interior forest 16–20   X  

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Small cats survey data from Central Kalimantan, Indonesia Jeffers et al.

Additional habitat
No. of CT days Location of camera Habitat class Altitude (m) LC MC FhC
information
2154 T0 x TC Mixed swamp forest Interior forest 16–20     X

907 T0 x TC 2013 Mixed swamp forest Interior forest 16–20 X    

907 T0 x TG 2013 Mixed swamp forest Interior forest 16–20     X

437 T0 x TH Mixed swamp forest Interior forest 16–20     X

379 T0.4E END Mixed swamp forest Interior forest 16–20      

904 T0.8 x TG 2013 Mixed swamp forest Interior forest 16–20     X

427 T0.8 x THH Mixed swamp forest Interior forest 16–20 X    

2154 T0.8E x TX Mixed swamp forest Interior forest 16–20      

616 T0.8E x TY Mixed swamp forest Interior forest 16–20      

841 T1.3E x TY 2013 Mixed swamp forest Interior forest 16–20   X  

55 T1.6 375m Mixed swamp forest Interior forest 16–20 X    

2395 T1.6 x Pondok Owa-Owa Mixed swamp forest Interior forest 16–20   X X

2395 T1.6 x Railway Mixed swamp forest Interior forest 16–20      

881 T1.6 x TC 2013 Mixed swamp forest Interior forest 16–20      

435 T1.6E x TZ Mixed swamp forest Interior forest 16–20   X  

2395 T1A x Railway Mixed swamp forest Interior forest 16–20 X    

436 T1A x TD Mixed swamp forest Interior forest 16–20      

83 T1B x Railway Mixed swamp forest Interior forest 16–20      

649 T2 700m 2013 Mixed swamp forest Interior forest 16–20      

5 T2 x TA Mixed swamp forest Interior forest 16–20      

2446 T2 x TB Mixed swamp forest Interior forest 16–20      

699 T2 x TD Mixed swamp forest Interior forest 16–20 X    

62 T2 x THH Mixed swamp forest Interior forest 16–20      

905 T2E x OR 2013 Mixed swamp forest Interior forest 16–20      

435 T2E x TY Mixed swamp forest Interior forest 16–20      

2450 TD x Jelutong Pondok Mixed swamp forest Interior forest 16–20   X X

1007 Tower Path Mixed swamp forest Interior forest 16–20 X    

260 TREE Railway 1350m 2015 Mixed swamp forest Canopy 10m 10      

258 TREE T 0.8 412m 2015 Mixed swamp forest Canopy 10m 10      

116 TREE T 0.8E x TX 2015 Mixed swamp forest Canopy 10m 10      

247 TREE T 1B 350m 2016 Mixed swamp forest Canopy 10m 10 X    

257 TREE T SC 685m 2015 Mixed swamp forest Canopy 10m 10      

81 TS x TBB 525m Mixed swamp forest Forest edge (<20m) 16–20 X    

81 TS x TCC Mixed swamp forest Forest edge (<20m) 16–20      

131 TP 0 650m Tall interior forest Interior forest 16–20 X   X

131 TP 1 1200m Tall interior forest Interior forest 16–20      

130 TP A 800m Tall interior forest Interior forest 16–20      

130 TP A x TP 1 Tall interior forest Interior forest 16–20      

131 TP B 1700m Tall interior forest Interior forest 16–20      

2399 Km5 x Railway Low interior forest Interior forest 16–20 X    

13488 Journal of Threatened Taxa | www.threatenedtaxa.org | 12 March 2019 | 11(4): 13478–13491

Small cats survey data from Central Kalimantan, Indonesia Jeffers et al.

Appendix 2. Summary of camera trap (CT) locations in Rungan, Central Kalimantan Indonesian Borneo. LC - Leopard Cat, MC - Marbled Cat,
FHC - Flat-headed Cat.

Additional habitat Altitude

No. of CT days Location of camera Habitat class LC MC FhC BC
information (m)
107 Mungku Baru Cam 13 Frank Kerangas/ heath Interior forest 50 X      

107 Mungku Baru Cam 15 Frank Kerangas/ heath Interior forest 50        

107 Mungku Baru Cam 16 Frank Kerangas/ heath Interior forest 50 X      

106 Mungku Baru Cam 18 Frank Kerangas/ heath Interior forest 50        

106 Mungku Baru Cam 19 Frank Kerangas/ heath Interior forest 50        

682 Mungku Baru Cam 2 Frank Kerangas/ heath Interior forest 50        

316 Mungku Baru Cam 3 Frank Kerangas/ heath Interior forest 50 X      

274 Mungku Baru Cam 4 Frank Kerangas/ heath Interior forest 50        

682 Mungku Baru Cam 5 Frank Kerangas/ heath Interior forest 50 X X    

682 Mungku Baru Cam 6 Frank Kerangas/ heath Interior forest 50        

64 Mungku Baru Cam 7 Frank Kerangas/ heath Interior forest 50 X      

63 Mungku Baru Cam 8 Frank Kerangas/ heath Interior forest 50        

63 Mungku Baru Cam 9 Frank Kerangas/ heath Interior forest 50     X  

661 Mungku Baru CAM BNF 1+2 Kerangas/ heath Interior forest 50 X      
Mungku Baru CAM BNF
661 Kerangas/ heath Interior forest 50 X      
11+12
Mungku Baru CAM BNF
661 Kerangas/ heath Interior forest 50 X     X
13+14
Mungku Baru CAM BNF
660 Kerangas/ heath Interior forest 50 X      
17+18
Mungku Baru CAM BNF
660 Kerangas/ heath Interior forest 50        
19+20
Mungku Baru CAM BNF
659 Kerangas/ heath Interior forest 50        
21+22
Mungku Baru CAM BNF
659 Kerangas/ heath Interior forest 50        
23+24
Mungku Baru CAM BNF
660 Kerangas/ heath Interior forest 50        
25+26
Mungku Baru CAM BNF
497 Kerangas/ heath Interior forest 50        
27+28
Mungku Baru CAM BNF
660 Kerangas/ heath Interior forest 50     X  
29+30
498 Mungku Baru CAM BNF 3+4 Kerangas/ heath Interior forest 50        
Mungku Baru CAM BNF
660 Kerangas/ heath Interior forest 50        
31+32
Mungku Baru CAM BNF
660 Kerangas/ heath 12m in canopy 70        
33 TREE
Mungku Baru CAM BNF
355 Kerangas/ heath 10m in canopy 70        
34 TREE
296 Mungku Baru CAM BNF 5+6 Kerangas/ heath Interior forest 70   X    

498 Mungku Baru CAM BNF 7+8 Kerangas/ heath Interior forest 70 X      

62 Mungku Baru Cam 1 Frank Kerangas/ heath Interior forest 70        

63 Mungku Baru Cam 10 Frank Low interior forest Interior forest 70        

109 Mungku Baru Cam 11 Frank Low interior forest Interior forest 70        

109 Mungku Baru Cam 12 Frank Low interior forest Interior forest 70        

107 Mungku Baru Cam 17 Frank Low interior forest Interior forest 70        

107 Mungku Baru Cam 14 Frank Mixed swamp forest Interior forest 70        
Mungku Baru CAM BNF
660 Mixed swamp forest Interior forest 70        
15+16
Mungku Baru CAM BNF
661 Mixed swamp forest Edge of a lake 40     X  
9+10 Danau

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Small cats survey data from Central Kalimantan, Indonesia Jeffers et al.

Appendix 3. Scientific, common, and Indonesian names and IUCN Red List status of species from Central Kalimantan, Indonesian Borneo.

Scientific name Common name Local name Recent IUCN status

Pongo pygmaeus Bornean Orangutan Kahiu Critically Endangered

Manis javanica Sunda Pangolin Trenggiling, Peusing Critically Endangered

Prionailurus planiceps Flat-headed Cat Kucing dampak Endangered

Cynogale bennettii Otter Civet Musang air Endangered

Catopuma badia Bay Cat Kucing merah Endangered

Hylobates albibarbis Bornean White-bearded Gibbon Kalaweit/ Owa-owa Endangered

Ciconia stormi Storm’s Stork Unknown Endangered

Neofelis diardii Sunda Clouded Leopard Macan dahan/ Harimau dahan Vulnerable

Lophura erythrophthalma Malay Crestless Fireback Manok himba Vulnerable

Helarctos malayanus Sun Bear Beruang Vulnerable

Arctictis binturong Binturong Binturong Vulnerable

Aonyx cinerea Asian Small-clawed Otter Unknown Vulnerable

Sus barbatus Bearded Pig Babi hutan Vulnerable

Cervus unicolor Sambar Rusa Vulnerable

Presbytis rubicunda Red langur Kelasi/ Lutung merah Vulnerable

Macaca nemestrina Pig-tailed Macaque Beruk Vulnerable

Tarsius bancanus borneanus Horsfield’s Tarsier Inkir/ Binatang hantu Vulnerable

Petinomys setosus Temminck's Flying Squirrel Unknown Vulnerable

Mulleripicus Pulverulentus Great Slaty Woodpecker Balatuk Vulnerable

Setornis criniger Hook-billed Bulbul Unknown Vulnerable

Melanoperdix niger Black Partridge Unknown Vulnerable

Pardofelis marmorata Marbled Cat Kuwuk Near Threatened

Hemigalus derbyanus Banded Civet Musang Near Threatened

Herpestes semitorquatus Collared Mongoose Unknown Near Threatened

H. brachyurus Short-tailed Mongoose Unknown Near Threatened

Muntiacus atherodes Bornean Yellow Muntjac Kijang/ Kidang, Muncak Near Threatened

Anthracoceros malayanus Black Hornbill Tingang/ Enggang Near Threatened

Carpococcyx radiatus Bornean Ground-cuckoo Unknown Near Threatened

Strix leptogrammica Brown Wood-owl Unknown Near Threatened

Lophura ignita Bornean Crested Fireback Unknown Near Threatened

Harpactes diardii Diard's Trogon Unknown Near Threatened

Trichixos pyrropygus Rufous-tailed Shama Unknown Near Threatened

Trichastoma rostratum White-chested Babbler Unknown Near Threatened

Prionailurus javanensis Leopard Cat Kucing hutan, Meong congkok Least Concern

Trichys fasciculata Long-tailed Porcupine Landak Least Concern

Hystrix brachyura Malayan Porcupine Landak Least Concern

Prionodon linsang Banded Linsang Musang congkok Least Concern

Paradoxurus hermaphroditus Common Palm Civet Unknown Least Concern

Viverra tangalunga Malay Civet Unknown Least Concern

Arctogalidia trivirgata Small-toothed Palm Civet Civet Least Concern

Martes flavigula Yellow-throated Marten Unknown Least Concern

Mustela nudipes Malay Weasel Unknown Least Concern

Muntiacus muntjak Southern Red Muntjac Unknown Least Concern

Macaca fascicularis Long-tailed Macaque Monyet ekor panjang (Kra) Least Concern

13490 Journal of Threatened Taxa | www.threatenedtaxa.org | 12 March 2019 | 11(4): 13478–13491

Small cats survey data from Central Kalimantan, Indonesia Jeffers et al.

Scientific name Common name Local name Recent IUCN status

Nannosciurus melanotis Black-eared Squirrel Hantitik Least Concern

Sundasciurus lowii Low's Squirrel Unknown Least Concern

Echinosorex gymnura Moonrat Unknown Least Concern

Ptilocercus Lowii Pen-tailed Treeshrew Unknown Least Concern

Callosciurus notatus Plantain Squirrel Unknown Least Concern

C. prevostii Prevost's Squirrel Unknown Least Concern

Maxomys surifer Indomalayan Maxomys Unknown Least Concern

Tupaia glis Common Treeshrew Tupai Least Concern

T. splendidula Ruddy Treeshrew Tupai Least Concern

T. picta Painted Treeshrew Tupai Least Concern

T. tana Large Treeshrew Tupai Least Concern

T. gracilis Slender Treeshrew Tupai Least Concern

T. minor Lesser Treeshrew Tupai Least Concern

Exilisciurus whiteheadi Tufted Pygmy Squirrel Hantitik Least Concern

Pellorneum capistratum Black-capped Babbler Unknown Least Concern

Pitta moluccensis Blue-winged Pitta Unknown Least Concern

Ketupa ketupu Buffy Fish-owl Unknown Least Concern

Stachyris erythroptera Chestnut-winged Babbler Unknown Least Concern

Phaenicophaeus curvirostris Chestnut-breasted Malkoha Unknown Least Concern

Centropus bengalensis Lesser Coucal Unknown Least Concern

Accipiter trivirgatus Crested Goshawk Unknown Least Concern

Spilornis cheela Crested Serpent-eagle Unknown Least Concern

Rhipidura javanica Pied Fantail Unknown Least Concern

Caprimulgus affinis Savanna Nightjar Unknown Least Concern

Pelargopsis capensis Stork-billed Kingfisher Bakaka Least Concern

Sitta frontalis Velvet-fronted Nuthatch Unknown Least Concern

Amaurornis phoenicurus White-breasted Waterhen Baburak Least Concern

Copsychus malabaricus White-rumped Shama Murai Least Concern

Spizaetus melanoleucus Black-and-white Hawk-eagle Antang Least Concern

Varanus salvator Common Water Monitor Biawak Least Concern

Author details: Karen Jeffers is the Sebangau Research Project Manager of the Borneo Nature Foundation. She has over 10 years’ experience in Indonesia studying
biodiversity, policy and has been researching small felids in Sebangau for some years, providing unique insights into their behaviour. Mr. Adul is the Senior Camera Trap
Coordinator for BNF. Adul has worked with the camera trap project since its inception and has travelled to several other conservation projects to share his knowledge
and skills in running camera trap projects. Adul presented BNF’s work at the Indonesian Carnivore Conference in 2017. Susan Cheyne has worked in Southeast Asia
since 1997 and in Indonesia since 2002. She is a co-director of BNF and has carried out long-term gibbon and mammal population monitoring in eight sites across
Indonesian Borneo. She is a vice-chair of the IUCN Primate Specialist Group Section on Small Apes and a member of the IUCN Cat Specialist Group.

Author contribution: Karen Jeffers carried out the field research under RISTEK permit number 8/TKPIPA/E5/Dit.KI/VIII/2018. She contributed to data collection with
Adul and analysis and writing with SMC. Adul designed the camera study grid with KAJ and SMC and assisted the fieldwork. He contributed to data analysis with KAJ
and SMC. Susan Cheyne conceived of the study and assisted in the field design. Along with KAJ and Adul she analysied the data and wrote the paper.

Threatened Taxa

Journal of Threatened Taxa | www.threatenedtaxa.org | 12 March 2019 | 11(4): 13478–13491 13491

Journal of Threatened Taxa | www.threatenedtaxa.org | 12 March 2019 | 11(4): 13492–13496
Short Communication

Insights into the feeding ecology of and threats to

Sand Cat Felis margarita Loche, 1858 (Mammalia: Carnivora:
ISSN 0974-7907 (Online)
Felidae) in the Kyzylkum Desert, Uzbekistan
ISSN 0974-7893 (Print)
Alex Leigh Brighten 1 & Robert John Burnside 2
PLATINUM
OPEN ACCESS 1,2
School of Environmental Sciences, University of East Anglia, Norwich Research Park, Norwich, NR2 7TJ, UK.
1
[email protected] (corresponding author), 2 [email protected]

Abstract: Little is known about the ecology of the Sand Cat Felis Felis margarita is classified as Least Concern in the
margarita throughout its range in the deserts of northern Africa to
central Asia. We present observations of the Sand Cat in the southern
IUCN Red List (Sliwa et al. 2016). In Uzbekistan, while
Kyzylkum Desert, Uzbekistan, potentially preying upon a large bird and F. m. thinobius is not listed in the Red Data Book of the
returning to the kill on subsequent nights. This record contributes to country (Khassanov 2009), it has been recommended for
the knowledge about the feeding ecology and varied diet of the Sand
Cat and its opportunistic hunting strategy.
inclusion in the next edition of the book, which is yet to
be published (Gritsina pers. comm. 18 April 2018). Apart
Keywords: Asian Houbara, Bukhara region, camera trapping, from local knowledge and anecdotal evidence, nothing
Chlamydotis macqueenii, conservation needs, opportunistic feeding,
scavenging. is known about the ecology, distribution, population
sizes, or trends of the species in Uzbekistan (Gritsina
2014) nor anywhere in central Asia. Therefore, any new
observation contributes to the knowledge base on this
The distribution of the Sand Cat Felis margarita species. Here we present opportunistic observations on
ranges from northern Africa to central Asia across which the feeding ecology of a Sand Cat in Uzbekistan.
it almost exclusively inhabits sandy and stony deserts
(Schauenberg 1974). Very little is known about its Materials and Methods
ecology and while some aspects have been studied in As part of long-term research into the ecology of
Israel (Abbadi 1991), Morocco (Sliwa et al. 2013; Breton Asian Houbara Chlamydotis macqueenii in the southern
et al. 2016; Sliwa et al. 2017; Breton & Sliwa 2018), and Kyzylkum Desert west of Bukhara, field research teams
Iran (Ghafaripour et al. 2017), the Central Asian Sand have been spending 3.5 months in the study area each
Cat F. m. thinobius remains particularly understudied. year from 2012–2018 as described in Burnside et al.
Burnside et al. (2014) confirmed a breeding population 2014. During this fieldwork, they occasionally observe
to be still present in the southern Kyzylkum Desert, Sand Cat, but as the work is diurnal it does not overlap
Uzbekistan (Fig. 1), aligning modern data with the species well with the nocturnal activity of Sand Cat, which
distribution reported by Heptner & Sludskii (1992). reduces the probability of detecting the species. The

DOI: https://doi.org/10.11609/jott.4445.11.4.13492-13496 | ZooBank: urn:lsid:zoobank.org:pub:85CFD5B8-ECAE-4652-8E58-A4A9BD34BAD2

Editor: Shomita Mukherjee, SACON, Coimbatore, India. Date of publication: 12 March 2019 (online & print)

Manuscript details: #4445 | Received 27 July 2018 | Final received 14 January 2019 | Finally accepted 25 February 2019

Citation: Brighten, A.L. & R.J. Burnside (2019). Insights into the feeding ecology of and threats to Sand Cat Felis margarita Loche, 1858 (Mammalia: Carnivora:
Felidae) in the Kyzylkum Desert, Uzbekistan. Journal of Threatened Taxa 11(4): 13492–13496; https://doi.org/10.11609/jott.4445.11.4.13492-13496

Copyright: © Brighten & Burnside 2019. Creative Commons Attribution 4.0 International License. JoTT allows unrestricted use, reproduction, and distribution of
this article in any medium by adequate credit to the author(s) and the source of publication.

Funding: The Ahmed bin Zayed Charitable Foundation.

Competing interests: The authors declare no competing interests.

Acknowledgements: The authors are funded by the Ahmed bin Zayed Charitable Foundation. We are grateful to Angie Appel for her insightful knowledge and
valuable feedback, to Erasil Khaitov for his tracking expertise and extensive knowledge of the desert, and to the Emirates Bird Breeding Center for Conservation
(EBBCC) for their support. We also thank the anonymous reviewers for their comments that helped improve the manuscript. Field research into Asian Houbara
ecology is conducted under permissions from Gosbiokontrol, Uzbekistan.

13492
Feeding ecology of Sand Cat Brighten & Burnside

Figure 1. Study area, C. macqueenii Research Project, Kyzylkum Desert, Uzbekistan, showing Sand Cat observation (green dot) and the different
vegetation zones. © R. Burnside.

data is thus generally limited to opportunistic diurnal Results

observations. This equivalent field effort in each field The carcass of C. macqueenii was found on a small
season resulted in an average of one Sand Cat report hill of consolidated sand with low shrubs dominated
per year, with the exception of 2014 (Burnside et al. by Astragalus villosissimus and Salsola spp. and sparse
2014) and 2018 (this article) when the species was also grass cover. The discovery was made after sighting
recorded through camera traps. feathers distributed in shrubs up to 10m around the
Part of our work is to monitor the survival of released kill at 40.4230N & 63.9860E. Feathers did not show
captive-bred C. macqueenii and establish causes of its signs of chewing but had been plucked. The pectoral
mortality in the field. In the period after their release, muscles were partially eaten and the entire head and
captive-bred birds are more susceptible to predation neck were missing. The legs and wings were intact
than their wild counterparts (Burnside et al. 2016). and undamaged. This is unlike a kill by Red Fox Vulpes
On 26 March 2018, we located a freshly-killed and vulpes, another common predator of released captive-
partially-eaten C. macqueenii, identified by its leg rings bred C. macqueenii in the area (Burnside et al. 2016). A
as a recently released captive-bred yearling male. The Red Fox usually chews the limbs, removing and caching
evidence found around the carcass suggested that the them, while leaving chewed feather quill tips as opposed
predator was a cat. We deployed a trail camera (Bushnell to plucking them at kill sites (Robert J. Burnside,
Trophy Cam HD Essential, model #119836) 2m from the unpublished data).
kill, low to the ground and concealed in a shrub, for three Pounce marks found close to the carcass,
nights. We set it to record motion-triggered, 15s-long approximately 4m away, were identified as that of a
videos both during the day and night (PIR sensor) and Sand Cat. Erasil Khaitov, an experienced tracker in
then returned to collect it three days later. the research team who has worked extensively in the

Journal of Threatened Taxa | www.threatenedtaxa.org | 12 March 2019 | 11(4): 13492–13496 13493

Feeding ecology of Sand Cat Brighten & Burnside

Image 1. Camera trap footage of a Sand Cat feeding on the carcass of a captive-bred Asian Houbara in the Kyzylkum Desert, Uzbekistan. © A. Brighten.

Image 2. Sand Cat pounce marks left during a hunt, with four paw-prints (highlighted with white dashed boxes) in the Kyzylkum Desert,
Uzbekistan. © A. Brighten & R. Burnside.

Kyzylkum Desert, identified the species’ prints without the kill once more at 21.24h on 28 March 2018, after
hesitation. Tracks leading up to the kill site showed a which there were no more observed visits by the cat or
slow, creeping approach indicating that the cat was other vertebrate scavengers. The footage showed the
moving low to the ground. In two areas, the tracks Sand Cat to be a male. The morning after the second
deepened with the force of a pounce (Image 2), with all visit by the cat, however, the carcass had been removed.
four paws visible; drag marks of approximately 6m were There was a single night trigger on 28 March, the second
seen nearby leading to where the carcass was found. visit by a cat, suggesting there to be a battery failure
The camera trap recorded footage of a Sand Cat limiting the firing of the infra-red flash. The removal
returning to the carcass on 26 March 2018 (Image 1). It of the carcass was, therefore, not caught on camera as
arrived after dark at 20.55h and spent 15min at the kill the next trigger was 29 March at 10.32h showing the C.
where it was seen to feed on the C. macqueenii (Video macqueenii to have been taken away. We were unable to
1). confirm which scavenger removed the carcass.
A Sand Cat was recorded on the camera trap visiting

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Feeding ecology of Sand Cat Brighten & Burnside

Discussion more than what they could eat, later returning to feed
Very little is known about the Sand Cat’s feeding on the carcasses. Returning to kills and scavenging has
ecology. Components of its diet were described in been documented in only a few small wild cat species.
Uzbekistan in the 1960s from stomach contents of Sliwa (1994) observed Black-footed Cats Felis nigripes
hunted cats (Schauenberg 1974), which mainly consisted killing and caching Southern Black Bustards Afrotis afra
of small burrowing rodents. Other studies from central in South Africa and a scavenging event on a Springbok
Asia summarised by Heptner & Sludskii (1992), using Antidorcas marsupialis lamb, while Avenant & Nel
stomach contents, faecal samples, or a combination of (2002) reported Caracal Caracal caracal feeding on the
both, found Sand Cat diet dominated by gerbils Gerbillus carcasses of Springbok that it had presumably killed.
and jerboa species like Allactaga, Dipus, and Paradious; In Uzbekistan and other parts of central Asia, the
however, this also varied, comprising of other mammals Sand Cat is likely threatened by increasing degradation
such as Tolai Hare Lepus tolai and Souslik Spermophillus and encroachment of its desert habitat through
leptodactilus, reptiles such as snakes Spalerosophis anthropogenic activities, both industrial and private.
diodema and Coluber karelini and gecko Teratoscincus, Particularly in the Bukhara region, this encroachment
birds such as Turtle Dove Streptopelia turtur, Crested Lark includes expanding industry and infrastructure, mainly
Galerida cristata, Saxaul Jay Podoces panderi, and Desert construction of railways, roads, and pipelines, as well as
Sparrow Passer simplex, a single observation of a Pallas mining for natural resources such as gas, oil, gold, sand,
Sand Grouse Syrrhaptes paradoxus, and arthropods such and gravel. Unlike large businesses, local communities
as Coleoptera, Phalangids, and Scorpiones. Sand Cats in Uzbekistan still have limited access to reliable fuel
were observed preying on gecko Stenodactylus in Israel resources due to the government’s policies on gas export.
(Abbadi 1991), and on Cape Hare Lepus capensis, viper The result is that the gathering of fuelwood continues
Cerastes, Greater Hoopoe Lark Alaemon alaudipes, and on a large scale in the Bukhara region. Historically, this
Domestic Fowl in the Sahara (Dragesco-Joffé 1993). was limited to Saxaul Haloxylon persicum, but recently
The southern Kyzylkum Sand Cat diet likely comprises we have seen the collection become less discriminating,
fauna found in our Bukhara study area, such as small uprooting other woody shrubs. We encountered tractors
rodents, including several Gerbillus species, Long-clawed undertaking such activities on an almost daily basis in
Souslik Spermophillus leptodactilus and Yellow Souslik S. the spring between 2016 and 2018 in the region. This
fulvus, 30 reptile species including Toad-headed Agama resulted in changes to the shrub structure and increase
Phrynocephalus, and four amphibian species (Showler in drifting sand (Robert J. Burnside, unpublished data).
2017). Small bird species are also numerous in the area, Pastoralism is the most widespread anthropogenic
Alaudidae in particular, including the abundant Crested activity in the desert. In general, it seems to have a low
Lark Galerida cristata, which are also probable prey impact on vegetation communities and structure at the
of the Sand Cat. The accounts of Abbadi (1991) and landscape-level and is at a stable level in the Bukhara
Dragesco-Joffé (1993) both describe Sand Cat hunting region (Koshkin et al. 2014). As with other rangeland
strategy as opportunistic and our observations in the systems, however, there is a general mistrust of
Kyzylkum Desert presented here support this assertion. mammalian predators among the local people. We have
From the evidence presented, it seems likely that the first-hand reports of the prevalent negative perceptions
C. macqueenii was killed by the Sand Cat. The average and direct persecution of cats in general, both of Sand Cat
weight of a captive-bred C. macqueenii yearling male and Asian Wildcat Felis lybica ornata, by the rangeland
is 1.5–2 kg, whereas Central Asian Sand Cats weigh inhabitants in our study area. One recent account
on average 3.125kg for males (2.65–3.40 kg, n=6) and (Erasil Khaitov, pers. comm. 20 May 2018) involved the
2.194kg for females (1.35–3.10 kg, n=5) (Heptner & destruction of a Sand Cat den and killing of kittens by a
Sludskii 1992). The C. macqueenii was, therefore, shepherd in retaliation for the loss of a lamb, supposedly
large prey for a small cat. The species may not form a killed by a Sand Cat. The evidence was that the lamb
significant part of Sand Cat diet, but this predator-naïve, was killed by a bite to the neck, which is indicative of a
recently-released C. macqueenii may have offered an cat, although other cats and carnivores inhabit the area
easy opportunity for the Sand Cat. (Caracal, Asian Wildcat, Jungle Cat Felis chaus, Red Fox,
Our record is the first of a Sand Cat returning to a kill Corsac Fox Vulpes corsac, and Grey Wolf Canis lupus).
in the Kyzylkum Desert, and it did not cover the carcass. Another threat to Sand Cat is human-introduced
In Niger’s Ténéré Desert, Dragesco-Joffé (1993) observed mammals such as the Domestic Dog Canis familiaris,
Sand Cats burying their prey in the sand when they killed which are potential predators of cats (Cole & Wilson

Journal of Threatened Taxa | www.threatenedtaxa.org | 12 March 2019 | 11(4): 13492–13496 13495

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Threatened Taxa

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Journal of Threatened Taxa | www.threatenedtaxa.org | 12 March 2019 | 11(4): 13497–13501

Short Communication
First photographic evidence of Fishing Cat
Prionailurus viverrinus Bennett, 1833 and
Clouded Leopard Neofelis nebulosa Griffith, 1821 ISSN 0974-7907 (Online)
(Carnivora: Felidae) in Parsa National Park, Nepal ISSN 0974-7893 (Print)

PLATINUM
Shashank Poudel 1 , Babu Ram Lamichhane 2 , Santosh Bhattarai 3 , Dipendra Adhikari 4 , OPEN ACCESS
Chiranjibi Prasad Pokheral 5 , Tek Raj Bhatt 6 , Sailendra Raj Giri 7 , Saneer Lamichhane 8 ,
Amir Sadaula 9 , Ashish Gurung 10 , Laxman Prasad Poudyal 11 , Uba Raj Regmi 12 &
Naresh Subedi 13

National Trust for Nature Conservation, P.O. Box 3712, Khumaltar, Lalitpur, 44600, Nepal.
1–3,5,8–10,13

4,6,7
Zoological Society of London, Nepal Office, Bhatbhateni, Kathmandu, Nepal.
11,12
Department of National Parks and Wildlife Conservation, Government of Nepal, Babarmahal, Kathmandu, Nepal.
1
[email protected] (corresponding author), 2 [email protected], 3 [email protected],
4
[email protected], 5 [email protected], 6 [email protected], 7 [email protected],
8
[email protected], 9 [email protected], 10 [email protected], 11 [email protected],
12
[email protected], 13 [email protected]

Abstract: Twelve cat species were recorded in Nepal including the Nepal is home to 12 cat species including both the
largest, Tiger Panthera tigris, and the smallest, Rusty-spotted Cat
Prionailurus rubiginosus. There is more research on the Panthera
largest, Tiger Panthera tigris, and the smallest, Rusty-
species than on small wild cats; consequently, the conservation status, spotted Cat Prionailurus rubiginosus, of the world
distribution, and ecology of small cat species are poorly known. In (Lamichhane et al. 2016; Lama et al. 2019). Distribution
this article, we report on the first photographic evidence of Clouded
Leopard Neofelis nebulosa and Fishing Cat Prionailurus viverrinus in
and ecology of large charismatic cats like Tiger, Snow
Parsa National Park in southern central Nepal during a camera trap Leopard Panthera uncia, and Leopard Panthera pardus
survey targeted at the tiger between 2014 and 2016. There were are researched considerably in Nepal (see Jackson 1996;
only single detections of each species; this does not give enough
information to establish distribution or conservation status of either Smith et al. 1998; Thapa et al. 2014; Karki et al. 2015) in
of the species in Parsa National Park. Further targeted surveys are comparison to small felids. There are huge information
needed to establish the significance of this protected area for the gaps on the distribution and status of smaller felids in
conservation of these two species.
Nepal. Most of the existing information on small cat species
Keywords: Camera trapping survey, small wild cats, southern central are based on historic references and specimens, anecdotal
Nepal, Terai.

DOI: https://doi.org/10.11609/jott.4549.11.4.13497-13501 | ZooBank: urn:lsid:zoobank.org:pub:DDF127AC-3B59-49F4-A4E4-D91EFE3A1387

Editor: Angie Appel, Wild Cat Network, Germany. Date of publication: 12 March 2019 (online & print)

Manuscript details: #4549 | Received 11 November 2018 | Final received 02 March 2019 | Finally accepted 09 March 2019

Citation: Poudel, S., B.R. Lamichhane, S. Bhattarai, D. Adhikari, C.P. Pokheral, T.R. Bhatt, S.R. Giri, S. Lamichhane, A. Sadaula, A. Gurung, L.P. Poudyal, U.R. Regmi &
N. Subedi (2019). First photographic evidence of Fishing Cat Prionailurus viverrinus Bennett, 1833 and Clouded Leopard Neofelis nebulosa Griffith, 1821 (Carnivora:
Felidae) in Parsa National Park, Nepal. Journal of Threatened Taxa 11(4): 13497–13501; https://doi.org/10.11609/jott.4549.11.4.13497-13501

Copyright: © Poudel et al. 2019. Creative Commons Attribution 4.0 International License. JoTT allows unrestricted use, reproduction, and distribution of this article
in any medium by adequate credit to the author(s) and the source of publication.

Funding: Funding for this work was provided by Zoological Society of London (ZSL) Nepal Office and Panthera.

Competing interests: The authors declare no competing interests.

Acknowledgements: We would like to thank the Department of National Parks and Wildlife Conservation and Parsa National Park office for coordinating the
survey and allowing us to study in the protected area. We are obliged for the support extended by all the field technicians of the National Trust for Nature
Conservation. In addition, we would also like to extend our gratitude towards the Nepal office of the Zoological Society of London and Panthera for supporting
the camera trap survey in Parsa National Park.

13497
Fishing Cat and Clouded Leopard in Parsa NP Poudel et al.

records, and sign surveys carried out in protected areas dry months (Lamichhane et al. 2017). The protected
for large felids (Lamichhane et al. 2016; Taylor et al. 2016). area supports diverse mammalian fauna including Tiger,
These efforts detected common and large felids but may Leopard, Dhole Cuon alpinus, Striped Hyena Hyaena
have missed rare and elusive smaller felids (Tempa et al. hyaena, Golden Jackal Canis aureus, Bengal Fox Vulpes
2013). Increased use of camera trapping in recent years bengalensis, and Honey Badger Mellivora capensis (Thapa
aided in the discovery of rare species like Clouded Leopard et al. 2014). The major prey species are Spotted Deer Axis
Neofelis nebulosa and Fishing Cat Prionailurus viverrinus in axis, Barking Deer Muntiacus muntjak, Gaur Bos gaurus,
new areas and provided verifiable records (see Appel et al. Nilgai Bosephalus tragocamelus, Wild Boar Sus scrofa, and
2012; Lamichhane et al. 2016; Taylor et al. 2016; Ghimirey Sambar Rusa unicolor. The combined ungulate density
& Acharya 2018; Yadav et al. 2018; Lama et al. 2019). The was estimated to be 25.33 (SE±3.9) ungulates/km2 in 2013
latest addition is the discovery of Marbled Cat Pardofelis (Dhakal et al. 2014).
marmorata in the eastern Himalayas of Nepal (Lama et al.
2019). Materials and Methods
Among the cat species, Clouded Leopard receives less The study was primarily designed for tiger monitoring,
attention from the conservation community in Nepal and deploying camera traps in continuous cells of 2kmx2km
is threatened by illegal wildlife trade and degrading habitat during the cool and dry seasons of November 2014 to
(Ghimirey & Acharya 2018). Though Hodgson (1853) January 2015 and February to April 2016, covering the
reported its presence in Nepal already in the mid-18th core area of Parsa NP during the first survey period and
Century, it was recorded in different parts of the country the new extension area during the 2016 survey period
only since the late 1980s (Dinerstein & Mehta 1989), such (Fig. 1). A pair of camera traps (model: Panthera V5) was
as in Dhanusa, Nawalparasi, and Kaski districts (Dinerstein placed in each cell. All the camera traps were active for
& Mehta 1989), Annapurna Conservation Area (Appel 24 hours for a minimum of 21 days during both survey
et al. 2012; Ghimire et al. 2019), Shivapuri Nagarjun NP periods. Following completion of the field survey, images
(Pandey 2012), and Chitwan NP (Ghimirey et al. 2014; were checked manually for the species recorded.
Lamichhane et al. 2014). In recent years, Fishing Cat was
recorded in Chitwan NP (Dahal & Dahal 2011; Mishra et Results
al. 2018), Koshi Tappu Wildlife Reserve (Taylor et al. 2016), Camera traps were deployed in 130 and 167 locations
Bardia NP (Yadav et al. 2018), Shuklaphanta NP (NTNC during the first and second survey period, with a total
survey records, unpublished), and Jagadishpur Reservoir survey effort of 7,230 trap nights, including 3,549 and
in Kapilvastu area of southwestern Nepal (Dahal 2016). 3,681 trap nights, respectively. We obtained a single image
We report Clouded Leopard and Fishing Cat recorded of Clouded Leopard and four images of a Fishing Cat.
for the first time in Parsa NP in southern central Nepal
during a monitoring survey targeting tiger. These are the Clouded Leopard
first confirmed records for the presence of both species in A single Clouded Leopard was photographed on 30
Parsa NP. November 2014 at 00.26h at 27.312°N & 84.961°E (Fig. 1)
on the eastern edge of Parsa NP. The species was identified
Study area by comparing the pelt pattern with the image of a Clouded
Parsa NP is located in southern central Nepal (27.3060N Leopard provided in the IUCN Red List (Grassman et al.
& 84.7810E) (Fig. 1), covering an area of 627km2 (Thapa et 2016). The age and sex of the individual, however, could
al. 2014; Lamichhane et al. 2018). It is contiguous with not be determined. The Clouded Leopard was camera
Chitwan NP in the west and is a critical region in the trapped in Sal-dominated mixed forests (Image 1).
Chitwan-Parsa-Valmiki trans-boundary forest complex
in the Terai Arc landscape (Chanchani et al 2014). The Fishing Cat
park is dominated by subtropical dry deciduous forest Four images of Fishing Cat were obtained from three
with colonizing Saccharum spontaneum and Imperata camera trap stations (27.235°N & 84.892°E; 27.234°N &
cylindrica on the dry riverbeds and floodplains to a climax 84.914°E; 27.246°N & 84.946°E) on the southeastern edge
Sal Shorea robusta forest on Bhabhar and hillsides (Thapa of Parsa NP during the 2016 survey (Fig. 1). Comparison
et al. 2014). The streams running off the Churia Hills of the pelage on both flanks from the paired cameras
permeates the porous sediment and flows underground, confirmed that all three stations recorded the same
reappearing south of the park and restricting water individual. We could not confirm its sex from the images.
availability in more than 70% of its area throughout the These three camera trap stations were in a Sal-dominated

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Fishing Cat and Clouded Leopard in Parsa NP Poudel et al.

Figure 1. Study area with camera trap locations in Parsa National Park, Nepal.

forest (Image 2). Our protocol of checking camera traps every alternate day
may have impacted encounter rates of shy and elusive
Discussion small wild cats.
Our records confirm that both Clouded Leopard and Large intact and interconnected forest patches of
Fishing Cat occur in Parsa NP. Thus, both species were this complex (ca. 3,000km2) might have provided an
documented in all three protected areas within the opportunity for the dispersal of the Clouded Leopard. Such
Chitwan-Parsa-Valmiki trans-boundary forest complex interconnected forest habitats are important for sustaining
(Clouded Leopard: Ghimirey et al. 2014; Lamichhane et viable populations of large carnivores such as Tiger,
al. 2014; Kamlesh Maurya pers. comm. 2017; Fishing Cat: Leopard, and Clouded Leopard, which in turn also protect
Dahal & Dahal 2011; Mishra 2016; Mukherjee et al. 2016). the functionality of the ecosystem (Borah et al. 2014;
Borah et al. (2014) reported frequent captures of Chanchani et al. 2014). To maintain its ecological integrity,
the Clouded Leopard by camera traps set up on paths the Chitwan-Parsa-Valmiki protected area complex should
and animal trails in Manas NP. Also, Lamichhane et al. also be kept intact by avoiding the construction of linear
(2014) reported records of the species on the forest floor infrastructure such as roads and railways that fragment
in Chitwan NP. The camera locations in Parsa NP were the forests, or by establishing wildlife-friendly corridors.
selected after an intensive search for signs such as scratch Across its range, the Fishing Cat is associated with
marks to maximize the probability of capturing the tiger. wetlands such as coastal and inland wetlands, rivers

Journal of Threatened Taxa | www.threatenedtaxa.org | 12 March 2019 | 11(4): 13497–13501 13499

Fishing Cat and Clouded Leopard in Parsa NP Poudel et al.

Image 1. Clouded Leopard Neofelis nebulosa in a dry riverbed in Parsa Image 2. Fishing Cat Prionailurus viverrinus on a forest road in Parsa
National Park, Nepal. © Department of National Parks and Wildlife National Park, Nepal. © Department of National Parks and Wildlife
Conservation, National Trust for Nature Conservation, Zoological Conservation, National Trust for Nature Conservation, Zoological
Society of London, 16 November 2014. Society of London, 08 March 2016.

Table 1. Details of Clouded Leopard and Fishing Cat capture events

in Parsa National Park, Nepal.

The location where the Fishing Cat was recorded is
a dry forest road in Bhabar Forest, which contrasts with
Particulars Fishing Cat Clouded Leopard records in wetlands across southern and southeastern
K17 Asia (e.g., Cutter & Cutter 2009 in Thailand; Mukherjee et
(27.2350N & 84.8920E),
Grid ID L17 N13
al. 2012 in West Bengal, India; Islam et al 2015 in Pakistan;
(GPS Locations) (27.2340N & 84.9140E), (27.3120N & 84.9610E) Taylor et al. 2016 in eastern Nepal; Palei et al. 2018 in
M16
(27.2460N & 84.9460E)
Odisha, India; Thaung et al. 2018 in coastal Cambodia). A
Elevation 201–255 m 458m targeted survey for Fishing Cat by Sharma (2016) did not
Number of record the species in Parsa NP. This survey covered only
detections 3 1 a small area (~20km2), targeting two sites close to rivers
Date of capture 8.iii.2016 30.xi.2014
and streams. Our record is at a distance of ~20km from
04.48h (K17), 05.37h the survey area of Sharma (2016) and well away from
Time of capture (L17) & 19.43h (M16) 00.26h wetland sites or water sources (>5km). Looking at the
Flank image Both Right linear movement and the single record of the species, we
Forest road Riverbed assume that the individual might have arrived incidentally
Habitat surrounding
Sal forest Sal forest at the location or been passing through it while migrating
the location
Distance to nearest
to another habitat.
4.5–5.6 km 5.7km
settlement While the global ranges of Fishing Cat and Clouded
Distance to nearest Leopard are declining due to various anthropogenic factors
perennial water 4.2–5.7 km 0.5km
source (Grassman et al. 2016; Mukherjee et al. 2016), the current
records in an additional protected area would help in their
conservation.
These records, however, also indicate that more in-
and streams, marsh areas, reed beds, tidal creeks, and depth surveys on their distribution, abundance, and
mangrove forests (Mukherjee et al. 2016). Parsa NP is a habitat use are necessary and warranted. In addition, such
relatively dry area with limited water sources, especially targeted surveys will also contribute to understanding the
during the dry rainless season between October and dynamics between larger carnivores, like tiger and leopard,
April. Perennial water sources here are confined to small and small wild cats. Hence, we recommend a targeted
streams coming from the Churia Hills and flowing partly survey covering the current capture locations to ascertain
underneath the surface, which limits the availability of whether these individuals were transiting or are resident.
water in the park (Israil et al. 2006; Thapa et al. 2014).

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Fishing Cat and Clouded Leopard in Parsa NP Poudel et al.

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Threatened Taxa

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Journal of Threatened Taxa | www.threatenedtaxa.org | 12 March 2019 | 11(4): 13502–13505
Short Communication

First photographic evidence of polymorphic

Asiatic Golden Cat Catopuma temminckii Vigors & Horsfield,
ISSN 0974-7907 (Online)
1827 (Mammalia: Carnivora: Felidae) in Buxa Tiger Reserve,
ISSN 0974-7893 (Print) West Bengal, India
PLATINUM
OPEN ACCESS Mayukh Ghose 1 , Deepak Sharma 2 & Nandavaram Subbarao Murali 3
1–3
Buxa Tiger Reserve, Buxa Forest, Buxa, Alipurduar District, West Bengal 736122, India.
1
[email protected] (corresponding author), 2 [email protected], 3 [email protected]

Abstract: During a camera trap survey in Buxa Tiger Reserve in West (Protection) Act 1972 (Menon 20014).
Bengal, India, individuals of Asiatic Golden Cat Catopuma temminckii
were photographed between 13 and 26 February 2018. The images
The Asiatic Golden Cat is the largest wild cat among
provide the first photographic evidence of the species presence in this the oriental Felinae (Bashir et al. 2011) It is remarkably
protected area. Both golden and spotted individuals were recorded. polymorphic in its pelage and is also known as “a feline
Keywords: Camera trapping, eastern Himalaya, spotted morph,
of many costumes” (Dhendup 2016). The most common
subtropical wet hill forest, Temminck’s Cat. coat colour is golden or red-brown, less frequently also
dark brown (Jutzeler et al. 2010), grey, or black (Jigme
2011). The spotted morph was previously known only
from China (Smith & Xie 2008) and Bhutan (Wang 2007;
The Asiatic Golden Cat Catopuma temminckii, also Wang & Macdonald 2009).
known as the Temminck’s Cat, is a medium-sized elusive
wild cat distributed from eastern Nepal (Ghimirey & Pal Study area
2009) to southeastern Asia (Holden 2001; Johnson et al. Buxa Tiger Reserve is situated in Alipurduar District,
2009; Gray et al. 2014; Tantipisanuh et al. 2014; Willcox West Bengal, India, covering an area of 760.87km2. It
et al. 2014; Zaw et al. 2014). In India, it was recorded lies between 26.500–26.702 0N and 89.333–89.860 0E,
in the protected areas of Sikkim, Assam, Arunachal nestled between the international boundary to Bhutan
Pradesh, Meghalaya, and Mizoram (Choudhury 2007; in the north, the state boundary to Assam in the east,
Bashir et al. 2011; Lyngdoh et al. 2011; Gouda et al. Jaldapara Wildlife Division in the west, and Cooch Behar
2016; Nadig et al. 2016; Mukherjee et al. 2018). It is District in the south (Fig. 1). The reserve is located at
listed as Near Threatened by the IUCN (McCarthy et al. the confluence of three major biogeographic zones,
2015) and as Scheduled I species in the Indian Wildlife namely, central Himalaya, Brahmaputra Valley, and

DOI: https://doi.org/10.11609/jott.4254.11.4.13502-13505 | ZooBank: urn:lsid:zoobank.org:pub:95415006-5494-48D1-88CB-4FE2B630DC80

Editors: Angie Appel, Wild Cat Network, Germany & Shomita Mukherjee, SACON, Coimbatore, India. Date of publication: 12 March 2019 (online & print)

Manuscript details: #4254 | Received 15 May 2018 | Final received 03 July 2018 | Finally accepted 15 December 2018

Citation: Ghose, M., D. Sharma & N.S. Murali (2019). First photographic evidence of polymorphic Asiatic Golden Cat Catopuma temminckii Vigors & Horsfield,
1827 (Mammalia: Carnivora: Felidae) in Buxa Tiger Reserve, West Bengal, India. Journal of Threatened Taxa 11(4): 13502–13505; https://doi.org/10.11609/
jott.4254.11.4.13502-13505

Copyright: © Ghose et al. 2019. Creative Commons Attribution 4.0 International License. JoTT allows unrestricted use, reproduction, and distribution of this article
in any medium by adequate credit to the author(s) and the source of publication.

Funding: Buxa Tiger Conservation Foundation Trust, Buxa Tiger Reserve, Department of Forests Government of West Bengal.

Competing interests: The authors declare no competing interests.

Acknowledgements: Firstly we would like to thank Kalyan Rai, IFS, Deputy Field Director, Buxa Tiger Reserve (West Division), for his kind suggestions and sharing
required resources. This work would have been incomplete without Dr. Pradip Bauri, IFS, Deputy Field Director, Buxa Tiger Reserve (East Division), for his help in
conducting the field study. We are thankful to staff for their support, especially to Karna Bahadur Chettri, Wangal Drukpa, Bishwajit Rava, Mrinmoy Dutta Roy, and
Bhakte Chettri, for their immense help during the field study and for setting up of camera traps in remote locations.

13502
Asiatic Golden Cat in Buxa TR Ghose et al.

Figure 1. Buxa Tiger Reserve, West Bengal, India (Source: Tiger Conservation Plan 2013).

Figure 3. Locations where Asiatic Golden Cats were recorded in
Figure 2. Boundary of Buxa Tiger Reserve in West Bengal, India. February 2018 (yellow pins).

lower Gangetic plains. Altitude ranges between 53m Table 1. Details of camera trap images of Asiatic Golden Cat in Buxa
Tiger Reserve, West Bengal, India.
and 1,735m and constitutes the eastern Himalayan
subtropical wet hill forest (Tiger Conservation Plan Image Location Date Time Altitude Morph
2013). Notable tree species in the study area are 1, 2
26.770⁰N &
17.ii.2018 13.19h 1,025m Golden
89.577⁰E
Quercus, Acer, Castanopsis, Magnolia cathcartii, Alnus
26.781⁰N & 09.22– Golden and
nepalensis, Phoebe attenuate, Betula cylindrostachys, 3, 4, 5
89.611⁰E
13.ii.2018
09.23 h
1,355m
spotted
and various bamboo Bambusoideae species (Tiger 6
26.781⁰N &
26.ii. 2018 15.28h 1,355m Golden
89.611⁰E
Conservation Plan 2013).
This camera trap survey was conducted between
January and March 2018 using 182 Cuddeback C series
camera traps with colour strobe module. We recorded

Journal of Threatened Taxa | www.threatenedtaxa.org | 12 March 2019 | 11(4): 13502–13505 13503

Asiatic Golden Cat in Buxa TR Ghose et al.

Images 1 & 2. Golden morph of Asiatic Golden Cat recorded at 1,025m on 17 February 2018. © Camera Trap Buxa Tiger Reserve, placed by
Mayukh Ghose.

Images 3, 4 & 5. Spotted and golden morphs of Asiatic Golden Cat recorded at 1,355m on 13 February 2018. © Camera Trap Buxa Tiger Reserve,
placed by Mayukh Ghose.

cameras were placed opposite each other, keeping a

minimum distance of 0.7km to the next pair. The 91
camera trap pairs were active all throughout 24 hours.

Results
Camera traps were active on 26 days for a total of
2,366 station days in 195 locations. Asiatic Golden Cats
were recorded in six images on three different days in
two locations, both located in the high altitude zone of
the Buxa Tiger Reserve. These locations are separated
by 450sqkm. All six images show Asiatic Golden Cats
Image 6. Golden morph of Asiatic Golden Cat recorded at 1,355m
on 26 February 2018. © Camera Trap Buxa Tiger Reserve, placed by during the day (Images 1–6).
Mayukh Ghose. Golden and spotted morphs were recorded at the
same location, interacting with each other in a single
coordinates using a Garmin E-trex 10 GPS, based on WGS frame (Images 3 & 4).
84 datum, and a digital altimeter for recording elevation
of camera trap locations. Discussion
The study area was divided into a grid with cells of Our records provide the first photographic evidence
1.414×1.414 sqkm each in two blocks of 240km2. The for the presence of Asiatic Golden Cat in the Buxa Tiger

13504 Journal of Threatened Taxa | www.threatenedtaxa.org | 12 March 2019 | 11(4): 13502–13505

Asiatic Golden Cat in Buxa TR Ghose et al.

Reserve. This is also the first instance of golden and in Johor, peninsular Malaysia. Cat News Special Issue 8: 10–18.
Haidir, I.A., Y. Dinata, M. Linkie & D.W. MacDonald (2013). Asiatic
spotted morphs of Asiatic Golden Cat being recorded Golden Cat and Sunda Clouded Leopard occupancy in the Kerinci
together in the reserve. Seblat landscape, west-central Sumatra. Cat News 59: 7–10.
The spotted morph, also known as ocelot morph, Holden, J. (2001). Small cats in Kerinci Seblat National Park, Sumatra,
Indonesia. Cat News 35: 11–14.
of the Asiatic Golden Cat was long thought to be more Jigme, K. (2011). Four colour morphs of and the altitudinal record of
common in China than elsewhere in the species global Asiatic Golden Cat in Bhutan. Cat News 55: 12–13.
range (Jutzeler et al. 2010). In the Himalaya, this morph Johnson, A., C. Vongkhamheng & T. Saithongdam (2009). The
diversity, status and conservation of small carnivores in a montane
was first recorded in Bhutan’s Jigme Singye Wangchuck tropical forest in northern Laos. Oryx 43(4): 626–633.
National Park at an elevation of 3,738m (Wang 2007). Jutzeler, E., Y. Xie & K. Vogt (2010). Asiatic Golden Cat in China. Cat
Four colour morphs of Asiatic Golden Cat were recorded News Special Issue 5: 40–41.
Lyngdoh, S., K.M. Selvan, G.V. Gopi & B. Habib (2011). First
in the eastern Himalaya. Ghimirey & Pal (2009) reported photographic evidences of two rare cats from Pakke Tiger Reserve,
a melanistic morph in a Schima-Castanopsis forest in western Arunachal Pradesh. Current Science 101(10): 1284–1286.
Nepal’s Makalu-Barun National Park. Bashir et al. (2011) McCarthy, J., S. Dahal, T. Dhendup, T.N.E. Gray, S. Mukherjee, H.
Rahman, P. Riordan, N. Boontua & D. Willcox (2015).  Catopuma
reported melanistic and dark grey morphs co-occurring temminckii. In: The IUCN Red List of Threatened Species:
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Menon, V. (2014). Indian Mammals: A Field Guide. Hachette Boo
black morphs were recorded above 1,500m in Bhutan Publishing India Pvt. Ltd., Gurgaon, India, 528pp.
(Jigme 2011; Vernes et al. 2015; Dhendup 2016), in Mukherjee, S., P. Choudhary, R. Arthreya & P.V. Karunakaran (2018).
A tale of tails — an attempt to decipher small cat distributions by
Myanmar (Zaw et al. 2014), and in and around Eaglenest tail lengths, pp11–12. In: Appel, A., S. Mukherjee & S.M. Cheyne
Wildlife Sanctuary, Arunachal Pradesh, India (Mukherjee (eds.). Proceedings of the First International Small Wild Cat
et al. 2018). Conservation Summit, 11–14 September 2017, United Kingdom.
Wild Cat Network, Bad Marienberg, Germany, Sálim Ali Centre for
In contrast, only golden morphs were reported south Ornithology and Natural History, Coimbatore, India, and Borneo
of the Isthmus of Kra in peninsular Malaysia (Gumal Nature Foundation, Oxford, United Kingdom, 40pp.
et al. 2014) and in Sumatra (Holden 2001; Haidir et al. Nadig, S., R. Navya & A.P. Silva (2016). Small cats in the Himalayan
foothills: the Asian Golden Cat of Nongkhyllem Wildlife Sanctuary,
2013; Pusparini et al. 2014). The occurrence of diverse India. Small Wild Cat Conservation News 2: 23.
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to the eastern Himalaya, China, and Indo-China (Patel L. Werner, D. Lenz, H. Pfestorf, S. Kramer-Schadt, V. Radchuk, J.
Fickel & A. Wilting (2016). Two species of southeast Asian cats in
et al. 2016). The latter authors suggested that this the genus Catopuma with diverging histories: an island endemic
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Open Science 3(10): 160350; https://doi.org/0.1098/rsos.160350
diverse habitats in the species northern range while
Pusparini, W., H.T. Wibisono, G.V. Reddy, Tarmizi & P. Bharata (2014).
only golden morphs colonized the Sunda shelf during its Small and medium-sized cats in Gunung Leuser National Park,
southward expansion. Sumatra, Indonesia. Cat News Special Issue 8: 4–9.
Smith, A.T. & Y. Xie (2008). A Guide to the Mammals of China. Princeton
University Press, New Jersey, 544pp.
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Threatened Taxa

Journal of Threatened Taxa | www.threatenedtaxa.org | 12 March 2019 | 11(4): 13502–13505 13505

Journal of Threatened Taxa | www.threatenedtaxa.org | 12 March 2019 | 11(4): 13506–13510
Short Communication

First photographic record of the Rusty-spotted Cat

Prionailurus rubiginosus (I. Geoffroy Saint-Hilaire, 1831)
ISSN 0974-7907 (Online)
(Mammalia: Carnivora: Felidae) in Horton Plains National
ISSN 0974-7893 (Print) Park, Sri Lanka
PLATINUM
OPEN ACCESS Thilina Sudarshana Nimalrathna 1* , Yan Ru Choo 2* , Enoka P. Kudavidanage 3 ,
Thakshila Ravindra Amarasinghe 4 , Udamulle Gedara Sumith Indika Bandara 5 ,
Wanninayaka Aarahchilage Charitha Lakmali Wanninayaka 6 , Piyal Ravindrakumar 7 ,
Marcus Aik Hwee Chua 8 & Edward L. Webb 9
1,3,4
Tropical Ecosystem Research Network, 30c, Temple Road, Hadigama, Piliyandala, Sri Lanka.
2,9
Department of Biological Sciences, National University of Singapore, Singapore 117558.
3
Department of Natural Resources, Sabaragamuwa University of Sri Lanka, Belihuloya 70140, Sri Lanka.
5,6,7
Department of Wildlife Conservation, 811A, Jayanthipura, Battaramulla, Sri Lanka.
8
Lee Kong Chian Natural History Museum, National University of Singapore, Singapore 117558.
8
Department of Environmental Science and Policy, George Mason University, 4400 University Dr, Fairfax, VA 22030,
USA.
1,3,4
30c, Temple Road, Hadigama, Piliyandala, Sri Lanka.
2,9
14 Science Drive 4, Department of Biological Sciences, National University of Singapore, Singapore 117558.
5,6,7
Horton Plains National Park, Department of Wildlife Conservation, Ohiya, Sri Lanka.
8
2 Conservatory Drive, Singapore 117377.
1
[email protected], 2 [email protected], 3 [email protected] (corresponding author),
4
[email protected], 5 [email protected], 6 [email protected],
7
[email protected], 8 [email protected], 9 [email protected] (corresponding author)
* These authors contributed equally to the work.

Abstract: The Rusty-spotted Cat Prionailurus rubiginosus is thought The Rusty-spotted Cat Prionailurus rubiginosus  (I.
to be present in most forested areas of Sri Lanka. Though it was
suggested that the species may occur in montane regions, there
Geoffroy Saint-Hilaire, 1831) is native to India, Nepal,
was no photographic evidence to date. Here we present the first and Sri Lanka (Nekaris 2003; Mukherjee et al. 2016).
photographic record of the Rusty-spotted Cat in Horton Plains National It is currently listed by the IUCN as Near Threatened
Park. Individuals including cubs were photo-captured on 15 separate
occasions during a 5,538 camera trap days study. These photo-
(Mukherjee et al. 2016) and is considered nationally
captures were made both during the day and night, and indicate the Endangered in Sri Lanka (Ministry of Environment 2012).
presence of a breeding population in this protected area. Its main threats are habitat loss and fragmentation
Keywords: Camera trap, carnivore, Felidae, montane forest, protected (Mukherjee et al. 2016).
area. In Sri Lanka, the Rusty-spotted Cat is thought to be

DOI: https://doi.org/10.11609/jott.4094.11.4.13506-13510 | ZooBank: urn:lsid:zoobank.org:pub:1287B562-C3A1-4787-A3FA-CD7239A3A1F4

Editor: Angie Appel, Wild Cat Network, Germany. Date of publication: 12 March 2019 (online & print)

Manuscript details: #4094 | Received 26 February 2018 | Final received 31 August 2018 | Finally accepted 04 December 2018

Citation: Nimalrathna, T.S., Y.R. Choo, E.P. Kudavidanage, T.R. Amarasinghe, U.G.S.I. Bandara, W.A.C.L. Wanninayaka, P. Ravindrakumar, M.A.H. Chua & E.L. Webb
(2019). First photographic record of the Rusty-spotted Cat Prionailurus rubiginosus (I. Geoffroy Saint-Hilaire, 1831) (Mammalia: Carnivora: Felidae) in Horton Plains
National Park, Sri Lanka. Journal of Threatened Taxa 11(4): 13506–13510; https://doi.org/10.11609/jott.4094.11.4.13506-13510

Copyright: © Nimalrathna et al. 2019. Creative Commons Attribution 4.0 International License. JoTT allows unrestricted use, reproduction, and distribution of this
article in any medium by adequate credit to the author(s) and the source of publication.

Funding: The Singapore Ministry of Education Tier 1 grant to ELW and the Department of Biological Sciences, National University of Singapore.

Competing interests: The authors declare no competing interests.

Acknowledgements: The Leopard Conservation and Ecology Project operated under permit W/L/3/2/67/15 from the Deparatment of Wildlife Conservation, Sri
Lanka. Funding was by a Singapore, Ministry of Education AcRF Tier 1 grant to ELW. The authors acknowledge the assistance of the Deparatment of Wildlife
Conservation for this project.

13506
Rusty-spotted Cat in Horton Plains Nimalrathna et al.

present in most forested areas (Philips 1980 cited in and terraces indicate areas of former farmland (Padmalal
Nekaris 2003). Direct evidence of its presence below & Kikuchi 1993). HPNP is surrounded by natural forests,
170m in arid scrub forests, dry forests, and monsoon pine and eucalyptus plantations, and tea plantations
forests was presented by Nekaris (2003) and Kittle & with an associated village.
Watson (2004). Indirect or anecdotal evidence suggests
its presence in wet forests near montane regions (Nekaris Material and Methods
2003). Photographic evidence, however, is scarce, During the initial phase of a camera trapping project
particularly in the montane region. Here we report the focusing on Sri Lankan Leopard Panthera pardus kotiya
first photographic record of the Rusty-spotted Cat above in HPNP, we established 18 survey points in the core of
2,000m in Horton Plains National Park (HPNP), located the national park, next to and surrounding the grassland
in Sri Lanka’s montane zone, from a series of photo- (Fig. 1). Survey points were located within each cell of
captures made over a 15-month period. a 1km2 systematic grid. At each survey point, paired
ReconyxTM HC500 camera traps were set up facing each
Study area other 45cm above the target level. The cameras were
HPNP is a protected area covering 31.6km2 of deployed from 3 December 2015 to 3 March 2017, for
grassland and montane forest; it was designated as a total of 5,538 camera trap days. Each camera was in
a national park in 1988 (IUCN 1990) and is part of the operation for 24 hours daily in RapidfireTM mode, with
Central Highlands of Sri Lanka World Heritage Site no delay between image sets, taking three images per
(World Heritage Convention 2018). It is located within trigger.
the wet and cool highlands (Fernanado 1968) between The retrieved images were sorted into incidents; we
6.783–6.833 °N and 80.767–80.850 °E at an altitude of define an incident as an image or a series of images of a
2,100–2,300 m (Fig. 1). It receives an annual rainfall of species or individual separated by intervals of less than
2,000–5,000 mm (Werner 1988). The vegetation consists one hour. The timings of the incidents were analysed
of tropical montane forest interspersed with a mosaic of using the overlap package (Ridout & Linkie 2009) in R
large grasslands and forest patches; extensive grasslands v. 3.5.1 (R Core Team 2018). The Rusty-spotted Cat was

Figure 1. The study site in Horton Plains National Park (HPNP; marked in red on the map of Sri Lanka), located within Nuwara Eliya District
(orange). The circles on the satellite image of HPNP represent camera trap locations; Rusty-spotted Cat was recorded in locations marked in
red. Administrative map of Sri Lanka obtained from GADM v. 2.8 (2015).

Journal of Threatened Taxa | www.threatenedtaxa.org | 12 March 2019 | 11(4): 13506–13510 13507

Rusty-spotted Cat in Horton Plains Nimalrathna et al.

identified by its reddish-grey fur in daytime images,

reddish-brown spots that ran longitudinally along its
body and head, and by the dark stripes on the inner sides
of its front legs (Sunquist & Sunquist 2009). Because of
the poor contrast between the cat’s spots and coat in
our infrared images, we were not able to distinguish
among individuals.

Results
Over the survey period, the Rusty-spotted Cat was
captured in 15 incidents at five locations, of which two
incidents (13.3%) occurred during the day (Table 1;
Image 1). These locations were separated by distances Figure 2. Patterns in photo-capture timings (n=15) of the Rusty-
of 0.8–5.1 km. In the first incident on 26 March 2016, spotted Cat in Horton Plains National Park, Sri Lanka
two adult individuals were recorded in the same image.
The last incident was recorded on 16 October 2016. One
notable incident shows an adult female with a kitten Watson 2004). Further, the record of an adult female
during the day on 16 June 2016. On 22 July 2016, an Rusty-spotted Cat with a kitten indicates the presence
adult individual was photographed with a rodent in its of a breeding population in Horton Plains National Park.
jaws. Overall, most photo-captures occurred between This also indicates that the Rusty-spotted Cat possibly
sunset and sunrise (Fig. 2). maintains a reproductive population in montane forest
regions.
Discussion Given the nationally Endangered status of the species,
Our findings make an important extension to the these findings highlight the conservation value of HPNP
documented range of the species, as previously it was and may have implications for managing tourism in the
only documented below 2,100m (Nekaris 2003; Kittle & national park. HPNP is one of the most highly visited
national parks in Sri Lanka (Sri Lanka Tourist Board 2017)
with strict legal protection and enforcement; therefore,
Table 1. Records of Rusty-spotted Cat in Horton Plains National Park, the wildlife populations therein are likely adequately
Sri Lanka, in 2016. protected. Outside HPNP, however, the montane forest
Camera trap station Date and time of is one of the most threatened habitats in Sri Lanka (Kittle
Remarks
(CT) and elevation records et al. 2017). More research needs to be carried out in
6.vi.2016, 01.37h
high-altitude habitats throughout Sri Lanka to determine
13.vi.2016, 23.37h the importance of montane forest habitats and to assess
26.vi.2016, 23.59h the impact of forest fragmentation and land use change
CT02 (2,135m)
27.vi.2016, 20.29h for the cat’s survival.
27.vi.2016, 23.30h

30.vi.2016, 23.24h References

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Threatened Taxa

13510 Journal of Threatened Taxa | www.threatenedtaxa.org | 12 March 2019 | 11(4): 13506–13510

Dr. Okan Külköylüoğlu, Abant Izzet Baysal University, Bolu, Turkey (Crustacea)  Dr. Oguz Turkozan, Adnan Menderes University, Aydın, Turkey
Dr. Jesse Leland, Southern Cross University, New South Wales, Australia (Crustacea) 
Dr. George Mathew, Kerala Forest Research Institute, Peechi, India  Birds 
Dr. Mohilal Meitei, Manipur University, Camchipur, Manipur, India 
Dr. John C. Morse, Clemson University, Long Hall, Clemson, USA  Dr. Hem Sagar Baral, Charles Sturt University, NSW Australia 
Late Dr. T.C. Narendran, (Retired) Professor, University of Calicut, Kerala, India  Dr. Chris Bowden, Royal Society for the Protection of Birds, Sandy, UK 
Dr. John Noyes, Natural History Museum, London, UK Dr. Priya Davidar, Pondicherry University, Kalapet, Puducherry, India 
Dr. Albert G. Orr, Griffith University, Nathan, Australia (Odonata)  Dr. J.W. Duckworth, IUCN SSC, Bath, UK 
Dr. Renkang Peng, Charles Darwin University, Darwin, Australia (Heteroptera)  Dr. Rajah Jayapal, SACON, Coimbatore, Tamil Nadu, India 
Dr. Nancy van der Poorten, Toronto, Canada  Dr. Rajiv S. Kalsi, M.L.N. College, Yamuna Nagar, Haryana, India 
Dr. C. Raghunathan, Zoological Survey of India, Andaman and Nicobar Islands  Dr. Crawford Prentice, Nature Management Services, Jalan, Malaysia 
Dr. R. Ramanibai, Guindy Campus, Chennai, Tamil Nadu, India  Dr. V. Santharam, Rishi Valley Education Centre, Chittoor Dt., Andhra Pradesh, India 
Dr. Brett C. Ratcliffe, University of Nebraska, Lincoln, USA  Dr. C. Srinivasulu, Osmania University, Hyderabad, India 
Dr. Klaus Ruetzler, Smithsonian Institution, Washington, DC  Dr. K.S. Gopi Sundar, International Crane Foundation, Baraboo, USA 
Dr. Kareen Schnabel, NIWA, Wellington, New Zealand (Crustacea)  Dr. Gombobaatar Sundev, Professor of Ornithology, Ulaanbaatar, Mongolia 
Dr. R.M. Sharma, (Retd.) Scientist, Zoological Survey of India, Pune, India (Lepidoptera, Prof. Reuven Yosef, International Birding & Research Centre, Eilat, Israel
Coleoptera)  Dr. Taej Mundkur, Wetlands International, Wageningen, The Netherlands
Dr. Peter Smetacek, Butterfly Research Centre, Bhimtal, India (Lepidoptera)  Dr. Carol Inskipp, Bishop Auckland Co., Durham, UK
Dr. Manju Siliwal, WILD, Coimbatore, Tamil Nadu, India (Araneae)  Dr. Michael Hutchins, American Bird Conservancy, Washington, USA.
Dr. G.P. Sinha, Botanical Survey of India, Allahabad, India (Lichens)  Dr. V. Gokula, National College, Tiruchirappalli, Tamil Nadu, India
Dr. K.G. Sivaramakrishnan, Madras Christian College, Chennai, Tamil Nadu, India 
Dr. K.A. Subramanian, Zoological Survey of India, New Alipore, Kolkata, India  Mammals 
Dr. P.M. Sureshan, Zoological Survey of India, Kozhikode, Kerala, India (Hymenoptera) 
Dr. Martin B.D. Stiewe, The Natural History Museum, UK (Mantodea)  Dr. Giovanni Amori, CNR - Institute of Ecosystem Studies, Rome, Italy 
Dr. R. Varatharajan, Manipur University, Imphal, Manipur, India  Dr. Daniel Brito, Federal University of Goiás, Goiânia, Brazil 
Dr. Eduard Vives, Museu de Ciències Naturals de Barcelona, Terrassa, Spain  Dr. Anwaruddin Chowdhury, Guwahati, India 
Dr. John Veron, Coral Reef Foundation, Townsville, Australia  Dr. P.S. Easa, Kerala Forest Research Institute, Peechi, India 
Dr. Hui Xiao, Chinese Academy of Sciences, Chaoyang, China  Dr. Colin Groves, Australian National University, Canberra, Australia 
Dr. James Young, Hong Kong Lepidopterists’ Society, Hong Kong Dr. Cecília Kierulff, Victorville, California 
Dr. R. Sundararaj, Institute of Wood Science & Technology, Bengaluru, India (Isoptera) Dr. Kristin Leus, Copenhagen Zoo, Annuntiatenstraat, Merksem, Belgium 
Dr. M. Nithyanandan, Environmental Department, La Ala Al Kuwait Real Estate. Co. K.S.C., Dr. David Mallon, Zoological Society of London, UK 
Kuwait Dr. Antonio A. Mignucci-Giannoni, Universidad Interamericana de Puerto Rico, Puerto
Dr. George O. Poinar, Oregon State University, Corvallis, USA Rico 
Dr. S. Arularasan, Annamalai University, Parangipettai, India (Molluscs) Dr. Sanjay Molur, WILD/ZOO, Coimbatore, India 
Dr. Himender Bharti, Punjabi University, Punjab, India (Hemiptera) Dr. Shomita Mukherjee, SACON, Coimbatore, Tamil Nadu, India 
Mr. Purnendu Roy, London, UK (Lepidoptera)  Dr. P.O. Nameer, Kerala Agricultural University, Thrissur, Kerala, India 
Dr. Saito Motoki, The Butterfly Society of Japan, Tokyo, Japan Dr. Jill Pruetz, Iowa State University, Ames, USA 
Dr. Sanjay Sondhi, TITLI TRUST, Kalpavriksh, Dehradun, India (Lepidoptera)  Dr. Ian Redmond, UNEP Convention on Migratory Species, Lansdown, UK 
Dr. Nguyen Thi Phuong Lien, Vietnam Academy of Science and Technology, Hanoi, Vietnam Dr. Heidi S. Riddle, Riddle’s Elephant and Wildlife Sanctuary, Arkansas, USA 
(Hymenoptera) Dr. Kumaran Sathasivam, Marine Mammal Conservation Network of India, India 
Dr. Xiaoli Tong, South China Agricultural University, Guangzhou, China (Thysanoptera) Dr. Karin Schwartz, George Mason University, Fairfax, Virginia. 
Dr. Nitin Kulkarni, Tropical Research Institute, Jabalpur, India (Orthoptera) Dr. Christoph Schwitzer, University of the West of England, Clifton, Bristol, BS8 3HA 
Dr. Robin Wen Jiang Ngiam, National Parks Board, Singapore (Odonata) Dr. Jodi L. Sedlock, Lawrence University, Appleton, USA
Dr. Lional Monod, Natural History Museum of Geneva, Genève, Switzerland. Dr. Lala A.K. Singh, Bhubaneswar, Orissa, India 
Dr. Asheesh Shivam, Nehru Gram Bharti University, Allahabad, India Dr. Mewa Singh, Mysore University, Mysore, India 
Dr. Rosana Moreira da Rocha, Universidade Federal do Paraná, Curitiba, Brasil Dr. Meena Venkataraman, Mumbai, India 
Dr. Kurt R. Arnold, North Dakota State University, Saxony, Germany (Hemiptera) Dr. Erin Wessling, Max Planck Institute for Evolutionary Anthropology, Germany 
Dr. James M. Carpenter, American Museum of Natural History, New York, USA Dr. Dietmar Zinner, German Primate Center, Göttingen, Germany
(Hymenoptera: Dr. A.J.T. Johnsingh, Nature Conservation Foundation, Mysuru and WWF-India, India
Dr. David M. Claborn, Missouri State University, Springfield, USA (Diptera) Dr. Paul Racey, University of Exeter, Devon, UK
Dr. Kareen Schnabel, Marine Biologist, Wellington, New Zealand Dr. Honnavalli N. Kumara, SACON, Anaikatty P.O., Coimbatore, Tamil Nadu, India
Dr. Amazonas Chagas Júnior, Universidade Federal de Mato Grosso, Cuiabá, Brasil Dr. Ashwin Naidu, University of Arizona, Tucson, USA
Dr. Marc W. Holderied, University of Bristol, Bristol, UK
Fishes  Dr. H. Raghuram, The American College, Madurai, Tamil Nadu, India
Dr. David Olson, Zoological Society of London, UK
Dr. Neelesh Dahanukar, IISER, Pune, Maharashtra, India  Dr. Paul Bates, Harison Institute, Kent, UK
Dr. Carl Ferraris, Smithsonian Institution, Portland, USA Dr. Hector Barrios-Garrido, James Cook University, Townsville, Australia
Dr. M. Afzal Khan, Department of Zoology, Aligarh Muslim University, Aligarh, India  Dr. Jim Sanderson, Small Wild Cat Conservation Foundation, Hartford, USA
Dr. Topiltzin Contreras MacBeath, Universidad Autónoma del estado de Morelos, México  Dr. Dan Challender, University of Kent, Canterbury, UK
Dr. Heok Hee Ng, National University of Singapore, Science Drive, Singapore  Dr. David Mallon, Manchester Metropolitan University, Derbyshire, UK
Dr. Rajeev Raghavan, St. Albert’s College, Kochi, Kerala, India   
Dr. Lukas Rüber, Department of Vertebrates, Natural History Museum, Switzerland  Other Disciplines 
Dr. Anjana Silva, Rajarata University of Sri Lanka, Saliyapura, Sri Lanka 
Dr. Robert D. Sluka, Chiltern Gateway Project, A Rocha UK, Southall, Middlesex, UK  Dr. Aniruddha Belsare, Columbia MO 65203, USA (Veterinary)
Dr. Kevin Smith, IUCN, Cambridge, UK  Dr. S. Jayakumar, Pondicherry University, Puducherry, India (Climate Change) 
Dr. E. Vivekanandan, Central Marine Fisheries Research Institute, Chennai, India  Dr. Jeff McNeely, IUCN, Gland, Switzerland (Communities) 
Dr. W. Vishwanath, Manipur University, Imphal, India Dr. Stephen D. Nash, Scientific Illustrator, State University of New York, NY, USA (Scientific
Dr. J. Jerald Wilson, King Abdulaziz University, Jeddah, Saudi Arabia  Illustrator) 
Dr. Davor Zanella, University of Zagreb, Zagreb, Croatia Dr. Mandar S. Paingankar, University of Pune, Pune, Maharashtra, India (Molecular) 
Dr. A. Biju Kumar, University of Kerala, Thiruvananthapuram, Kerala, India Dr. Jack Tordoff, Critical Ecosystem Partnership Fund, Arlington, USA (Communities)
Dr. Frederic H. Martini, University of Hawaii at Manoa, Hanolulu, Hawaii Dr. Ulrike Streicher, University of Oregon, Eugene, USA (Veterinary)
Dr. Hari Balasubramanian, EcoAdvisors, Nova Scotia, Canada (Communities) 
Amphibians 
 
Dr. Indraneil Das, Sarawak, Malaysia  Reviewers 2016–2018
Dr. Sushil K. Dutta, Indian Institute of Science, Bengaluru, Karnataka, India  Due to pausity of space, the list of reviewers for 2016–2018 is available online.
Dr. Annemarie Ohler, Muséum national d’Histoire naturelle, Paris, France

Reptiles 

Late Dr. S. Bhupathy, SACON, Coimbatore, Tamil Nadu, India 

Dr. Llewellyn D. Densmore, Texas Tech University, Lubbock, USA 
Dr. Eric Smith, University of Texas, Arlington, USA 
Dr. Gernot Vogel, Heidelberg, Germany 
Dr. Anders G.J. Rhodin, Chelonian Research Foundation, Lunenburg, USA
Dr. Raju Vyas, Vadodara, Gujarat, India
Dr. Pritpal S. Soorae, Environment Agency, Abu Dubai, UAE.
Dr. Olivier S.G. Pauwels, Royal Belgian Institute of Natural Sciences, Brussels, Belgium The opinions expressed by the authors do not reflect the views of the
Dr. Anders G.J. Rhodin, Chelonian Research Foundation, Lunenburg, USA Journal of Threatened Taxa, Wildlife Information Liaison Development Society,
Zoo Outreach Organization, or any of the partners. The journal, the publisher,
the host, and the partners are not responsible for the accuracy of the political
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tematic Mycology, Biological Abstracts, BIOSIS Previews, CAB Abstracts,
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ISSN 0974-7907 (Online) | ISSN 0974-7893 (Print)

March 2019 | Vol. 11 | No. 4 | Pages: 13419–13510

Date of Publication: 12 March 2019 (Online & Print)
www.threatenedtaxa.org DOI: 10.11609/jott.2019.11.4.13419-13510

SMALL WILD CATS SPECIAL ISSUE

Editorial Short Communications

Foreword to the small wild cats special issue Insights into the feeding ecology of and threats to Sand Cat
– Angie Appel & Shomita Mukherjee, 13419–13420 Felis margarita Loche, 1858 (Mammalia: Carnivora: Felidae) in the
Kyzylkum Desert, Uzbekistan
– Alex Leigh Brighten & Robert John Burnside, Pp. 13492–13496
Articles
First photographic evidence of Fishing Cat Prionailurus viverrinus
Using camera traps to study the elusive European Wildcat Bennett, 1833 and Clouded Leopard Neofelis nebulosa Griffith, 1821
Felis silvestris silvestris Schreber, 1777 (Carnivora: Felidae) in (Carnivora: Felidae) in Parsa National Park, Nepal
central Germany: what makes a good camera trapping site? – Shashank Poudel, Babu Ram Lamichhane, Santosh Bhattarai,
– Harald Wening, Lynne Werner, Matthias Waltert & Markus Port, Dipendra Adhikari, Chiranjibi Prasad Pokheral, Tek Raj Bhatt,
Pp. 13421–13431 Sailendra Raj Giri, Saneer Lamichhane, Amir Sadaula, Ashish Gurung,
Laxman Prasad Poudyal, Uba Raj Regmi & Naresh Subedi, Pp. 13497–
Activity patterns of the small and medium felid (Mammalia: 13501
Carnivora: Felidae) guild in northeastern India
– Shomita Mukherjee, Priya Singh, André Pinto Silva, Chandan Ri, First photographic evidence of polymorphic Asiatic Golden Cat
Kashmira Kakati, Binod Borah, Tana Tapi, Sandesh Kadur, Catopuma temminckii Vigors & Horsfield, 1827 (Mammalia: Carnivora:
Prafull Choudhary, Shikha Srikant, Surabhi Nadig, R. Navya, Felidae) in Buxa Tiger Reserve, West Bengal, India
Mats Björklund & Uma Ramakrishnan, Pp. 13432–13447 – Mayukh Ghose, Deepak Sharma & Nandavaram Subbarao Murali,
Pp. 13502–13505
Estimating Leopard Cat Prionailurus bengalensis Kerr, 1792
(Carnivora: Felidae) density in a degraded tropical forest fragment in First photographic record of the Rusty-spotted Cat Prionailurus
northeastern Thailand rubiginosus (I. Geoffroy Saint-Hilaire, 1831) (Mammalia: Carnivora:
– Wyatt Joseph Petersen, Tommaso Savini, Robert Steinmetz & Felidae) in Horton Plains National Park, Sri Lanka
Dusit Ngoprasert, Pp. 13448–13458 – Thilina Sudarshana Nimalrathna, Yan Ru Choo, Enoka P.
Kudavidanage, Thakshila Ravindra Amarasinghe, Udamulle Gedara
An update on the status of Fishing Cat Prionailurus viverrinus Bennett, Sumith Indika Bandara, Wanninayaka Aarahchilage Charitha Lakmali
1833 (Carnivora: Felidae) in Thailand Wanninayaka, Piyal Ravindrakumar, Marcus Aik Hwee Chua &
– Wanlop Chutipong, Anucha Kamjing, Worata Klinsawat, Dusit Edward L. Webb, Pp. 13506–13510
Ngoprasert, Kitipat Phosri, Niti Sukumal, Pongnapa Wongtung &
Naruemon Tantipisanuh, Pp. 13459–13469

Communications

The Caracal Caracal caracal Schreber, 1776 (Mammalia: Carnivora:

Felidae) in Uzbekistan
– Mariya Alexeevna Gritsina, Pp. 13470–13477

Small cat surveys: 10 years of data from Central Kalimantan,

Indonesian Borneo
– Karen Anne Jeffers, Adul & Susan Mary Cheyne, Pp. 13478–13491
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