THE RAFFLES BULLETIN OF ZOOLOGY 2010

THE RAFFLES BULLETIN OF ZOOLOGY 2010 58(1): 145–156

Date of Publication: 28 Feb.2010
© National University of Singapore

MAMMAL DIVERSITY AND CONSERVATION IN A SMALL

ISOLATED FOREST OF SOUTHERN THAILAND

S. Kitamura
Laboratory of Animal Ecology, Department of Life Sciences, Faculty of Science, Rikkyo University, Tokyo 171-8501, Japan;
Thailand Hornbill Project, Department of Microbiology, Faculty of Science,
Mahidol University, Rama 6 Rd., Bangkok 10400, Thailand
Present address: Division of Nature and Environmental Management, The Museum of Nature and
Human Activities, Hyogo, Yayoigaoka 6, Sanda, Hyogo, 669-1546, Japan
Email: [email protected] (Corresponding author)

S. Thong-Aree and S. Madsri

National Park, Wildlife and Plant Conservation Department, 61 Phahonyothin Rd.,
Chatuchak, Bangkok 10900 Thailand

P. Poonswad
Department of Microbiology, Faculty of Science, Mahidol University, Rama 6 Rd., Bangkok 10400, Thailand
Email: [email protected]

ABSTRACT. – Knowledge of the presence and distribution of species is crucial for designing and evaluating
conservation strategies within a region. We conducted a camera-trapping survey of terrestrial mammal and
bird diversity in a small isolated forest of southern Thailand over 3 yr. A total of 15 camera traps, which
accumulated 11,106 camera-days, were set in three forest types: primary, logged, and hill forests. Despite its
small size, isolation, and surrounding agricultural areas, a total of 35 mammal species, eight bird species, and
one reptile species were recorded in the forest system. The total number of species photographed was similar
among forest types (26-30 species), and rarefaction curves of each forest did not indicate any differences in
the relationship between sampling effort and recorded species richness. As the activity period of these animals
does not appear to be affected by human activities, we believe that the effect of direct poaching on large
mammals in the study area is negligible. Although we did not observe any previously unrecorded animals
in our study site, our findings are very valuable and point to the importance of biodiversity conservation
efforts in these small fragmented and human-modified forest landscapes.

KEY WORDS. – Camera trapping, dipterocarp forest, Hala-Bala Wildlife Sanctuary, rarefaction curve.

INTRODUCTION Research into the presence and distribution of species is

crucial for planning and evaluating conservation strategies
Tropical forests are becoming increasingly fragmented, which within a region (Tobler et al., 2008). Even though studies
in turn threatens the survival of their resident species (Sodhi on mammals in tropical forests are often difficult, a great
et al., 2004b; Primack & Corlett, 2005; Laurance, 2007). deal of comprehensive work has focused on mammals in
Changes in the size, shape, or configuration of habitat that Southeast Asia (Carbone et al., 2001; Lynam et al., 2001;
result from human disturbances strongly affect populations Fernando et al., 2003; Kinnaird et al., 2003; O’Brien et al.,
of various animals. Species extirpation is driven by the 2003; Wong et al., 2004; Fredriksson et al., 2006; Wilting
direct effects of hunting and illegal trading, by the indirect et al., 2006; Meijaard & Sheil, 2008). Many of these species
effects of habitat loss (deforestation and fragmentation), and are charismatic and/or flagship species for the conservation
by interactions among these factors (Peres, 2001; Dirzo et of habitats or ecosystems.
al., 2007; Wright et al., 2007). No tropical forests are more
threatened and no tropical fauna are more endangered than Studies of large mammals in tropical forests have benefited
those of Southeast Asia (Sodhi et al., 2004a; Corlett, 2007; greatly from recent technological improvements that provide
Laurance, 2007). various ready-made camera traps with tiny infrared-motion
sensors, built-in-flashes, and data packs at a reasonable cost
(Swann et al., 2004; Yasuda, 2004). In fact, infrared-triggered

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camera systems are now widely used in vertebrate ecology, MATERIALS AND METHODS
with applications to nest ecology, population estimation,
behavioural ecology, mammal inventories, and studies of Study site. – The Bala forest is part of the Hala-Bala Wildlife
animal damage (Carthew & Slater, 1991; Cutler & Swann, Sanctuary (hereafter, Bala), on the Thai-Malaysian border
1999). Even in tropical forests, recent studies have determined (5°44'N 101°46'E–5°57'N 101°51'E; Fig. 1). The Bala
that sampling medium- to large-sized mammals using camera portion of the sanctuary is 111.5 km2 in area and is isolated
traps is a practical, non-invasive method that requires little from other forests by agricultural lands on the Thai side of
effort and is particularly useful when compared to alternative the border (the 314 km2 Hala portion of the sanctuary lies
methods, such as line transects, direct observations, track and approximately 22 km to the west) and a mix of forest and
faeces identification, trapping, and interviews of local people agriculture on the Malaysian side. Bala ranges in elevation
(Silveira et al., 2003; Srbek-Araujo & Chiarello, 2005). from 50 to 960 m above sea level and is broadly classified as
tropical lowland evergreen forest (Bird Conservation Society
In Southeast Asia, camera traps have been used in a number of Thailand, 2004). The forests primarily comprise species
of studies for a variety of research purposes, e.g., mammal typical of the Sundaic or Malesian region (Malay Peninsula,
inventories of specific areas (Numata et al., 2005; Mohd- Borneo, Sumatra, Java), many of which are uncommon in
Azlan & Engkamat, 2006; Giman et al., 2007; Dinata et al., Thailand and have their primary home ranges farther south
2008); monitoring animals visiting a particular site, such as on the Malay Peninsula. During 1987–1992, Bala was partly
natural licks (Matsubayashi et al., 2007a; Matsubayashi et logged along the paved road that bisects the lower one-third of
al., 2007b), natal dens (Lim & Ng, 2008), or fruiting trees the site and was sporadically (and illegally) logged elsewhere
(Miura et al., 1997; Kitamura et al., 2004; Kitamura et al., in the sanctuary.
2006; Suzuki et al., 2007; Yasuda et al., 2007; Van der
Meer et al., 2008; Kitamura et al., 2009); recording animal Bala forest receives an average annual rainfall of 3,500 mm
activity periods (Griffiths & van Schaik, 1993b; van Schaik (in some years nearly 5,000 mm), mostly from southeast
& Griffiths, 1996; Grassman et al., 2005a; Grassman et al., monsoons between October and January. The clear, dry
2005b; Grassman et al., 2006; Mohd-Azlan, 2006; Mohd- season lasts from February to April, and the mean monthly
Azlan & Sharma, 2006; Suzuki et al., 2006; Kawanishi temperature is 23°C. Fruit production is highly seasonal, and
& Sunquist, 2008); identifying seed dispersers/predators an abundance of ripe fruits peaks during July and September,
(Yasuda et al., 2000; Kitamura et al., 2004; Kitamura et al., while periods of fruit scarcity strongly differ across years (S.
2006; Kitamura et al., 2008); and estimating the relative Kitamura, unpublished data). Most field studies conducted in
and absolute abundance of a particular species, such as the Bala forest have focused on describing specific taxa, such
tigers (Mohd-Azlan & Sharma, 2003; O’Brien et al., 2003; as figs (Chantarasuwan & Thong-Aree, 2006), ants (Noon-
Kawanishi & Sunquist, 2004; Linkie et al., 2006; Mohd- Anant, 2003), and bats (Bumrungsri et al., 2006), with the
Azlan & Sharma, 2006; Lynam et al., 2007; Simcharoen et exception of several ecological studies on hornbills (Gale &
al., 2007), leopards (Ngoprasert et al., 2007), Malayan sun Thong-Aree, 2006; Kemp et al., 2007; Thong-Aree, 2007).
bears (Wong et al., 2004; Linkie et al., 2007), or Malayan
tapirs (Holden et al., 2003). Camera trapping. – Each remote camera was equipped with
a built-in infrared motion sensor, a built-in flash, and a data
Relatively undisturbed lowland forest is now rare in the pack that stamped each photograph with the time and date of
Sundaic region (Lambert & Collar, 2002), particularly in
southern Thailand, where more than 95% of the natural
forest has been destroyed (Round, 1988). Most of the
remaining forests in southern Thailand are highly fragmented
and isolated as a result of human activities, including
illegal logging, rubber plantations, and fruit orchards (Bird
Conservation Society of Thailand, 2004). With such a rapid
destruction of natural habitats, it is important to document
the presence, diversity, and distribution of large mammals
occurring in various forest patches in southern Thailand and
to study mammal species that may not be able to survive in
small isolated forests. Such research will provide essential
information for conservation and management efforts.
Moreover, the data can be used to assess loss of diversity
elsewhere if isolation/fragmentation occurs in a similar
manner. Here, we present data on mammalian fauna obtained
from a 3-year camera-trap study in an isolated forest of
southern Thailand.

Fig. 1. Hala-Bala Wildlife Sanctuary (Bala forest) in southern

Thailand, and the road cutting through the forest and associated
trails on either side, as used by Gale and Thong-Aree (2006).

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 THE RAFFLES BULLETIN OF ZOOLOGY 2010

the event (Sensor Camera Fieldnote, Marif Co., Ltd., Iwakuni, Data analysis. – Camera-trapping data are often difficult to
Yamaguchi, Japan). Power was supplied by a CR123A interpret when a series of photographs depict the same species,
lithium battery, which lasted approximately three months. which leads to issues of self-dependence and unsuitability
Colour print film (ISO 400) was used in each camera, and for statistical analysis (O’Brien et al., 2003; Yasuda, 2004).
no photographic delay interval was available for the model One way to minimise self-dependence in camera-trapping
used. If an animal or a group of animals remained at the results is to consider a series of photographs of the same
site, the camera was triggered every 10–15 sec. Each camera species taken within a certain period of time as a single event
was wrapped tightly in a transparent polypropylene bag and (Otani, 2001; O’Brien et al., 2003). In this study, we treated
encased in an unsealed plastic box for waterproofing. a picture as indicating a single visit by a given species if that
picture was taken >30 min after the previous picture of that
Cameras were deployed at 15 locations in three different species (O’Brien et al., 2003; Yasuda, 2004).
forest types (Fig. 1). In each forest type, cameras were
deployed 300–500 m apart along an existing trail: primary Researchers and conservation planners are often interested
forest (trail #4, southern part of the paved road, P1-5: 323 in the number of species supported by a particular habitat,
± 11 m a.s.l., mean ± SD), logged forest (trail #8, northern region, or protected area. In our study, unequal sampling
part of the paved road, L1-5: 301 ± 44 m a.s.l.), and hill effort (36 mo for both primary and logged forests versus 18
forest (trail #12, western part of Bala, H1-5: 715 ± 95 m mo for the hill forest) limited our ability to compare species
a.s.l.). The sampling effort varied among forests. In the richness among habitat types. When animal abundance varies
primary and logged forests, the same camera locations were between sites, comparisons of diversity will favour the more
maintained for 36 months (Oct.2004 to Oct.2007), whereas abundant site with more animals because of higher species
those in the hill forest were maintained for 18 months (May detection rates (Gotelli & Colwell, 2001). To control for
2006 to Sep.2007). this issue, we implemented the rarefaction approach with
EstimateS Win 8.0.0 (Colwell, 2005), and we used species
In the primary and logged forests, several cameras were accumulation curves to compare richness levels among
also set up along animal trails and around common fruiting habitats.
trees, such as Anisophyllea cornei (Anisophylleaceae),
Baccaurea parviflora (Euphorbiaceae), Canarium littorale In our study area, sunrise annually varies between 0557
(Burseraceae), Cheilosa malayana (Euphorbiaceae), and 0629 hrs and sunset varies between 1756 and 1834
Elaeocarpus stipularis (Elaeocarpaceae), Garcinia sp.1 hrs. According to the standard time near Bala, we divided
(Guttiferae), Irvingia malayana (Irvingiaceae), Palaquium the day into two time zones, i.e., daytime (0600 to 1800
impressinervium (Sapotaceae), and Sterculia macrophylla hrs) and nighttime (1800 to 0600 hrs). Time periods were
(Sterculiaceae). Ten fruits were collected from the ground pooled in 1-hour intervals. We assumed that the numbers
around fruiting trees and were placed at the base of the same of photographs taken were correlated with animal activity,
trees as animal bait. Cameras were set approximately 2 m and we classified the activity patterns of captured animals
away from the bait. When fruits were consumed by animals based on previous studies in Southeast Asia (van Schaik &
or damaged by insects, they were replaced with new samples. Griffiths, 1996; Grassman et al., 2005a; Grassman et al.,
Although the photographing duration was limited by the 2006). For species with > 10 captures, we defined diurnally
length of fruit availability, each session continued for at active species as those with < 10 percent of captures at night,
least one month. All camera-trapping stations were marked and nocturnally active species as those that had > 90 percent
using a global positioning system (GPS; GARMIN eTrex). captures at night. We considered species with between 10 and
Cameras were checked every month to reload new film. In 90 percent nocturnal captures as arrhythmic, i.e. they showed
several instances, the film had finished prior to checking; no clear activity pattern (Grassman et al., 2006).
therefore, some records may have been missed.

Species identification and their conservation status in RESULTS

Thailand. – After the film was developed, the photographs
were examined for images of animals. Animal nomenclature Species diversity of photographed animals in Bala. – The
follows (Lekagul & Round, 1991) for birds, and (Corbet & total number of camera-days was 11,106 over 36 months
Hill, 1992) for mammals. The identification of some small (4,428 camera-days in the primary forest, 3,965 in the logged
mammals was difficult when based solely on photographs. In forest, 1,891 in the hill forest, 613 along animal trails, and
addition to photographing, we also conducted live trapping of 389 around fruiting trees). In total, 8,414 photographs were
small mammals in the primary and logged forests of Bala (S. exposed, 3,750 of which (44.6%) contained no animal image.
Kitamura, unpublished data). Thus, we were able to identify The remaining 4,664 photographs (55.4%) showed images of
small mammals by comparing body sizes and colours of those 44 species, including 35 mammal species, eight bird species,
in the pictures to those that had been trapped. Based on the and one reptile species (Table 1). Of these, 2,087 photographs
Thailand Red Data Book (Nabhitabhata & Chan-Ard, 2005; (44.7%) were considered independent visits and were used
Sanguansombat, 2005), we determined the conservation status for further analysis.
in Thailand of the photographed animals.
Some species of bats and birds were photographed several
times but could not be identified from the photographs.

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Table 1. Summary of animals recorded in the Hala-Bala Wildlife Sanctuary, Thailand, from November 2004 to October 2007. Species
with > 10 photo-captures are underlined. Site abbreviations: P: primary forest; L: logged forest; H: hill forest; A: animal trail; and F:
fruiting tree. TRD: Thailand Red Data (Nabhitabhata & Chan-Ard, 2005; Sanguansombat, 2005), EN: endangered, VU: vulnerable, NT:
near threatened, LC: least concern.

Family Scientific name English name N Site TRD

Mammalia
Tupaiidae Ptilocercus lowii Feather-tailed treeshrew 2 F VU
Tupaia glis Common treeshrew 55 P, L, A, F
Erinaceidae Echinosorex gymnurus Moonrat 47 P, L, F VU
Cercopithicidae Macaca nemestrina Pig-tailed macaque 237 P, L, H, A, F NT
Trachypithecus obscurus Dusky langur 12 P, L, H VU
Manidae Manis javanica Sunda pangolin 5 P, L, F NT
Sciuridae Callosciurus caniceps Grey-bellied squirrel 1 L
C. notatus Palntain squirrel 5 P, L, H, F
Lariscus insignis Three-striped ground squirrel 9 P, L, F VU
Rhinosciurus laticaudatus Shrew-faced ground squirrel 1 F VU
Muridae Leopoldamys sabanus Noisy rat 83 P, L, F
Maxomys surifer Red spiny rat 293 P, L, H, A, F
M. whiteheadi Whitehead’s rat 6 P, L, F VU
Sundamys muelleri Muller’s rat 1 P
Hystricidae Atherurus macrourus Bush-tailed porcupine 5 L, H, A LC
Hystrix brachyura Malayan porcupine 160 P, L, H, A, F LC
Mustelidae Martes flavigula Yellow-throated marten 17 P, L, H, F
Viverridae Arctictis binturong Binturong 7 P, L, H
Hemigalus derbyanus Banded palm civet 13 P, L, H, A, F EN
Paguma larvata Masked palm civet 21 P, L, H, A
Paradoxurus hermaphroditus Common palm civet 1 L
Prionodon linsang Banded linsang 7 P, L, H, A VU
Herpestidae Herpestes javanicus Javan mongoose 1 H
Felidae Catopuma temminckii Asian golden cat 43 P, L, H, A, F VU
Panthera pardus Leopard 2 P, L VU
P. tigris Tiger 5 P, H EN
Pardofelis nebulosa Clouded leopard 4 H VU
Prionailurus bengalensis Leopard cat 13 P, L, H, A
Tapiridae Tapirus indicus Malayan tapir 99 P, L, H, A EN
Suidae Sus scrofa Common wild pig 426 P, L, H, A, F
Tragulidae Tragulus kanchil Lesser mouse deer 58 P, L, A, F
T. napu Greater mouse deer 12 P, L, F EN
Cervidae Muntiacus muntjak Common barking deer 396 P, L, H, A, F
Bovidae Naemorhedus sumatraensis Southern serow 1 H NT
Bat spp. Bat spp. 5 L, H, A
Mammal total 35 spp. 2,053
Aves
Accipitridae Spilornis cheela Crested serpent eagle 1 H
Phasianidae Argusianus argus Great argus 18 L, H VU
Gallus gallus Red junglefowl 1 A
Columbidae Chalcophaps indica Emerald dove 2 F
Bucerotidae Anorrhinus galeritus Bushy-crested hornbill 1 H NT
Turdidae Zoothera citrina Orange-headed thrush 1 P
Copsychus malabaricus White-rumped shama 1 F
Bird spp. Bird spp. 3 P, H
Bird total 8 spp. 28
Reptilia
Varinidae Varanus sp.1 Monitor lizard 6 L, H, A, F
Reptile total 1 spp. 6
2,087

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Mammals accounted for 98.3 percent of all animals identified first year than during the second or third years (Fig. 3). In
from the photographs. Of the identified species, wild boar, contrast, the number of species photographed during the
Sus scrofa, was the most frequently photographed (426 second compared to the third year did not significantly
times, 20.4% of the total), followed by common barking differ in either the primary or logged forest. Most animal
deer, Muntiacus muntjak (396 times), red spiny rat, Maxomys species were photographed in the first year (93% of recorded
surifer (293 times), pig-tailed macaque, Macaca nemestrina species in both the primary and logged forests). Similarly,
(237 times), and Malayan porcupine, Hystrix brachyura (160 the number of species photographed within the first 12 mo
times). These five species accounted for 72.5 percent of all accounted for 85% of the total species recorded for 18 mo
identified photographs. In contrast, six species of mammals in the hill forest.
and five species of birds were photographed only once during
the study (Table 1). Most animals were photographed alone Activity period. – The activity periods were based on filtered
or in pairs, except for the Malayan porcupine (up to three data and were determined for only 18 species (Fig. 4). Six
individuals), pig-tailed macaque (up to six individuals), and species of mammals (Echinosorex gymnurus, Leopoldamys
wild boar (up to 17 individuals). Of the 44 animal species sabanus, Maxomys surifer, Hystrix brachyura, Hemigalus
photographed, 21 are listed in the Thailand Red Data derbyanus, and Prionailurus bengalensis) were classified as
Book, including four endangered, 11 vulnerable, four near nocturnal, whereas three mammal species and one bird species
threatened, and two species of least concern (Table 1). (Macaca nemestrina, Trachypithecus obscurus, Tupaia glis,
and Argusianus argus) were considered diurnal.
Species accumulation curves. – In total, 30, 28, and 26
species were photographed in the primary, logged, and hill
forests, respectively. The identity of species substantially
overlapped in the primary and logged forests (24 species in
common) but overlapped to a lesser extent in the logged and
hill forests (19 species) and the hill and primary forests (17
species). Rarefaction of the source pool of species sampled
in each forest indicated no differences among forests in the
relationship between sampling effort and recorded species
richness (Fig. 2). The number of photographs per 100
camera-days also did not differ among forest types: 20.2 ±
4.3 (mean ± SD) in the primary forest, 17.1 ± 14.4 in the
logged forest, and 16.6 ± 11.4 in the hill forest. Similarly,
the number of photographed species per 100 camera-days
Fig. 3. Rarefaction curves for photo-captured animal assemblages
did not substantially differ among forests: 1.9 ± 0.6 in the in primary and logged forests over 3 years (thick line: first year;
primary forest, 2.1 ± 0.9 in the logged forest, and 3.1 ± 1.7 open circles: second year; solid circles: third year) in the Hala-Bala
in the hill forest. Wildlife Sanctuary, Thailand. The dotted line represents the 95%
CI for the first year. For clarity, the 95% CI for the second and
In both the primary and logged forests, the number of third years are not presented.
photographed species was significantly higher during the

Fig. 2. Rarefaction curves for photo-captured animal assemblages

in three forest types (thick line: primary forest; open circles: logged
forest; solid circles: hill forest) in the Hala-Bala Wildlife Sanctuary, Fig. 4. Activity periods of photo-captured animals (N>10) in
Thailand, from Nov.2004 to Oct.2007. The dotted line represents the Hala-Bala Wildlife Sanctuary, Thailand, from Nov.2004 to
the 95% CI for the primary forest. For clarity, the 95% CI for the Oct.2007. Species are listed in order of decreasing frequency of
logged and hill forests are not presented. diurnal activity. Numbers in parentheses indicate sample size.

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DISCUSSION Davison, 2006; Dinata et al., 2008). In Bala, the call of

this species is also the most commonly heard birdsong;
Species diversity of mammals and birds photographed in thus, camera trapping would not be necessary for a general
Bala. – Our study revealed that at least 44 species of animals, survey. Although camera trapping provides some baseline
of which 21 (47.7%) are listed in the Thailand Red Data information for species records (Mohd-Azlan & Davison,
Book, were found in an isolated forest of Bala in southern 2006) and can sometimes detect very rare species (Dinata
Thailand. One method of assessing the relative conservation et al., 2008), we did not record any new birds for the Bala
value of Bala is to compare its species richness as assessed forest. For certain mammal species, camera trapping is
by photography to that evaluated by photography elsewhere, a promising survey technique that generates useful data;
assuming that higher species richness implies higher quality however, camera trapping used solely for the study of birds
habitat. The species richness of large- and medium-sized is impractical in Bala.
mammals photographed in the Jerangau Forest Reserve,
Peninsular Malaysia, was 27 species over 5,972 camera-days Activity periods of mammals. – Research into animal activity
(Mohd-Azlan, 2006), and that of three study sites in Taman periods using camera trapping in tropical forests in Southeast
Negara National Park, Peninsular Malaysia, was 27 species Asia is increasing (Griffiths & van Schaik, 1993a; b; van
over 4,336 camera-days, 25 over 4,847 camera-days, and 26 Schaik & Griffiths, 1996; Miura et al., 1997; Mohd-Azlan
over 4,871 camera-days (Kawanishi, 2002). Compared to & Sharma, 2003; Grassman et al., 2005a; Grassman et al.,
these studies, the species richness of large- and medium-sized 2006; Mohd-Azlan, 2006; Mohd-Azlan & Engkamat, 2006;
mammals (not including that of small mammals belonging Mohd-Azlan & Sharma, 2006; Suzuki et al., 2006; 2007).
to Tupaiidae, Erinaceidae, Sciuridae, Muridae, and bats) in For 11 animal species, we found documentation of activity
Bala was relatively low (21 species). periods for the same species in previous studies from the Phu
Kheio Wildlife Sanctuary (Grassman et al., 2005a; Grassman
Despite the huge sampling effort in our study, we did et al., 2006) and Khao Yai National Park (Suzuki et al., 2006;
encounter certain limitations that deterred us from examining 2007) in Thailand and Jerangau Forest Reserve (Mohd-Azlan,
the relative importance of the Bala forest in greater depth. 2006), Taman Negara National Park (Kawanishi & Sunquist,
Because we continuously maintained our camera trappings at 2004), and Pasoh Forest Reserve (Miura et al., 1997) in
the same locations for three years, and the minimum distance Peninsular Malaysia (Table 2). There were no differences
between each camera location was short (300–500 m), the between the activity periods determined in our study and
effective sampling area in our study was rather limited. those reported in the previous studies, with the exception of
Although the rarefaction curve for photo-captured animal the leopard cat, which we classified as nocturnal but others
assemblages continued to increase throughout the study categorised as arrhythmic (Table 2). The activity periods
period (Fig. 2), the number of species photographed in the first of mammals common to Bala generally do not differ from
year included most of the photographed species. Some species those at other study sites, despite its small isolated habitat
may actually be more abundant or even restricted to one surrounded by agricultural areas, such as fruit orchards and
habitat type. To increase the probability of capturing species rubber plantations. In areas severely disturbed by humans,
that rarely use trails or are habitat specialists, it is important some large game mammals shift their activity periods from
to ensure that all major habitat types are sampled. Indeed, diurnal to nocturnal (Griffiths & van Schaik, 1993a; b; van
clouded leopard and southern serow were not recorded in Schaik & Griffiths, 1996). We did not observe this trend at
the primary or logged forests over the three study years but our sites, nor did we record any hunting of large animals
were observed in the hill forest during one year. Even in the during our study, although we did witness the collection of
primary and logged forests, several rare species in Bala, such non-forest timber products (e.g., wild fruits and mushrooms)
as the Asian elephant Elephas maximus and the Malayan and the trapping of small passerines (e.g., bulbuls and leaf
sun bear Ursus malayanus went undetected by our camera birds) by local peoples. Thus, we believe that the effect of
trappings. To prevent camera malfunction, no cameras were direct poaching on large mammals in Bala is negligible, and
deployed near streams due to flooding or near steep rocky the activity period of these animals is likely not affected by
slopes. Together, these factors indicate that a comprehensive such human activities.
mammal list has yet to be compiled for this area, particularly
for secretive animals that are strictly arboreal or associated The protected area of Bala is only 111.5 km2, a threshold
with aquatic habitats. To obtain a complete list of the below which tigers and some other large mammals may
mammals of Bala, other methods of observation must be rapidly become extirpated from this region (Laidlaw, 2000).
combined with camera-trapping studies. Most large mammals photographed in this study have large
home ranges (Table 3) and require large areas of contiguous
In our photographs, bird species accounted for only 1.7 forest for their long-term survival (Karanth & Nichols, 2002).
percent of all animals identified. In general, the frequency Habitats for large mammals are highly fragmented and are
of photographed mammals is much greater than that of lost through the construction of roads, power networks,
birds in tropical forests of Southeast Asia (Miura et al., plantations, urban expansion, dams, irrigation, and other
1997; Mohd-Azlan & Davison, 2006; Dinata et al., 2008). development in this region (Bird Conservation Society
The great argus Argusianus argus, a true ground dweller, of Thailand, 2004; Lynam et al., 2007). Although we did
was the most commonly photographed bird species (Table not record any animals that were previously unknown to
1), as previously reported in this region (Mohd-Azlan & Bala, we did confirm the existence of some endangered

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 Table 2. Comparison of activity periods of 11 mammal species photographed in this study at five different sites in Thailand and Peninsular Malaysia. Activity abbreviations: D: diurnal; N:
nocturnal; A: arrhythmic.

Thailand Peninsular Malaysia

Species This study (Grassman et al., (Suzuki et al., (Mohd-Azlan, 2006) (Kawanishi & (Miura et al., 1997)
2005a; 2006) 2006; 2007) Sunquist, 2004)
Common treeshrew D D D
Pig-tailed macaque D D D D D
Noisy rat N N N
Red spiny rat N N N

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Malayan porcupine N N N N
Asian golden cat A A
Leopard cat N A A A
Malayan tapir A A A
Common wild pig A A A A A A
Lesser mouse deer A A A A A A
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Common barking deer A A A A A

 Table 3. Home-range sizes of various mammals photographed in the Hala-Bala Wildlife Sanctuary, Thailand. Only studies conducted in tropical Asia are cited. NA: not available.

Family Scientific name Home range (ha) References

Mammalia
Tupaiidae Ptilocercus lowii 1.9–6.4 (Emmons, 2000)
Tupaia glis 0.33 (Walker & Rabinowitz, 1992)
Erinaceidae Echinosorex gymnurus NA
Cercopithicidae Macaca nemestrina 62–828 (Caldecott, 1986)
Trachypithecus obscurus 17–33 (Curtin, 1980)
Manidae Manis javanica 5.6–7.0 (Lim & Ng, 2008)
Sciuridae Callosciurus caniceps 0.7–2.4 (Saiful et al., 2001)
C. notatus 0.7–1.8 (Saiful et al., 2001)
Lariscus insignis 0.2–3.7 (Saiful et al., 2001)
Rhinosciurus laticaudatus NA
Muridae Leopoldamys sabanus 0.25–0.32 (Wells et al., 2008)
Maxomys surifer 0.12–0.84 (Walker & Rabinowitz, 1992)
M. whiteheadi 0.02–0.14 (Nakagawa et al., 2007)
Sundamys muelleri NA
Hystricidae Atherurus macrourus NA

152
Hystrix brachyura NA
Mustelidae Martes flavigula 350–1180 (Grassman et al., 2005c)
Viverridae Arctictis binturong 470–650 (Grassman et al., 2005c)
Hemigalus derbyanus NA
Paguma larvata 60–145 (Rabinowitz, 1991)
Paradoxurus hermaphroditus 210–630 (Rabinowitz, 1991)
Prionodon linsang NA
Herpestidae Herpestes javanicus NA
Kitamura et al.: Mammal diversity of a Thai forest

Felidae Catopuma temminckii 3260–4770 (Grassman et al., 2005e)

Panthera pardus 500–4800 (Odden & Wegge, 2005)
P. tigris 7800 (Lynam et al., 2001)
Pardofelis nebulosa 2290–4510 (Grassman et al., 2005e)
Prionailurus bengalensis 220–3710 (Grassman et al., 2005d)
Tapiridae Tapirus indicus 1275 (Williams, 1978)
Suidae Sus scrofa NA
Tragulidae Tragulus kanchil 3.5–6.2 (Matsubayashi et al., 2003)
T. napu NA
Cervidae Muntiacus muntjak 647–592 (Heydon, 1994)
Bovidae Naemorhedus sumatraensis NA
 THE RAFFLES BULLETIN OF ZOOLOGY 2010

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ACKNOWLEDGMENTS University of California Press, London, UK.
Fernando, P., T. N. C. Vidya, J. Payne, M. Stuewe, G. W. H.
We thank the National Research Council of Thailand and Davidson, R. J. Alfred, P. Andau, E. Bosi, A. M. Kilbourn
the Thailand Department of National Park, Wildlife and & D. J. Melnick. 2003. DNA analysis indicates that Asian
Plant Conservation for collaboration and granting of permits. elephants are native to Borneo and are therefore a high priority
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Hala-Bala Wildlife Sanctuary. This study was supported by Fredriksson, G. M., S. A. Wich & Trisno. 2006. Frugivory in
the Mahidol University Government Research Grant, the sun bears (Helarctos malayanus) is linked to El Nino related
National Center for Genetic Engineering and Biotechnology fluctuations in fruiting phenology, East Kalimantan, Indonesia.
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Research Fellowships for S.K. Gale, G. A. & S. Thong-Aree. 2006. Density estimates of nine
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