Anonymous. 2004. Status, Ecology and Conservation of Tigers in Peninsular Malaysia. Report: 1-54. Department of Wildlife and National Parks, DWNP), Wildlife Conservation Society - International Programme, (WCS).

Keywords: 4MY/camera-trapping/conservation/ecology/Panthera tigris/status/survey/tiger

Abstract: During 1997-2001 a national tiger conservation program was conducted by the Department of Wildlife and National Parks (DWNP) with the support of international conservation agencies Wildlife Conservation Society (WCS), WWF – Malaysia, and the University of Florida. The objectives of the programme were to assess the current conservation status of tigers in Peninsular Malaysia, study tiger ecology, use the results to make management recommendations, and develop staff capacity to manage tigers. This report covers fieldwork conducted by DWNP and WCS during 1997-1999. Nine sites were chosen for tiger survey across four states: Pahang, Trengganu, Kelantan and Perak. Eleven variables describing microhabitat structure were measured at each sampling location. Principal components analysis (PCA) was used to identify two major gradients in habitat disturbance and identify relatively disturbed sites (Lepar, Gunong Tebu, Ulu Temiang) from relatively less disturbed sites (Temenggor, Taman Negara, Bintang Hijau). Surveys involving 6,259 trapnights of sampling using 174 camera-trap setups revealed the presence of tiger and 38 other mammals, almost half of which were globally threatened species. Also recorded were twelve species of large reptile and bird including five globally threatened or near-threatened species. Ten individual tigers were recorded from 51 photographs at six of the nine sites. Tigers were detected at a quarter of all sampling locations (range=0- 2 48% per site), and at a frequency of 4.8±0.5 detections/100 trapnights (range=0-5.9 detections/100 trapnights per site). Activity patterns were inferred for tiger, other carnivores, and prey species from the time imprinted on photographs. Tigers showed a crepuscular pattern of activity and apparently avoided periods when human traffic was greatest. Human activity was recorded at all sites except for Bintang Hijau. Overall human traffic occurred across 20% of the sampling areas. Photographs from camera-traps were used to identify tiger individuals from their unique stripe patterns. The photographs were used to generate capture histories for tigers. These were used along with a capture-recapture approach to determine statistically the probability of capture for tigers and sample estimates of abundance at different sites. Coefficients of variation for abundance estimates ranged from 2 to 392% reflecting low numbers of recaptures. Tiger density was estimated from the sample abundance estimates and by assuming a sampling effective area for each site. This area was calculated by adding a buffer to the configuration of camera-traps equal to the absolute maximum movement distance (AMDM) of tigers. Tiger density varied from extremely low (2 tigers/100km2) at Temenggor. Despite intensive sampling, because the capture rates for tigers were low, and captures took place over a one-year period, extrapolating a statistical estimate of tiger abundance across Peninsular Malaysia is difficult at best. Nonetheless, population estimates for tigers were attempted for individual reserves, for Tiger Conservation Units (TCUs) and for all areas in Peninsular Malaysia by assessing (1) population density, (2) forest size and (3) a correction factor to take into account avoidance of habitat due to roads, powerlines, settlements and other human infrastructure. While these estimates are very rough indeed, they provide an estimate of the national population of 184-305 tigers, most of which occur in one Level I TCU (TCU#129; 173-257 tigers). This is less than half of the previous estimate for Peninsular Malaysia. 13. Tigers are threatened by a range of factors chiefly habitat loss and fragmentation, hunting of prey, commercial trade in live animals, dead animals and their parts, and harassment and displacement. Immediate action is needed to reduce these threats in the short-term (next five years) and to recover populations of wild tigers in Peninsular Malaysia in the longer-term (5-20 years from now

FINAL REPORT

Status, Ecology and Conservation of Tigers in Peninsular Malaysia

For

Department of Wildlife and National

Parks (DWNP)

By

Wildlife Conservation Society – International Programme

(WCS)

January 2004

ii

Table of Contents

1. Executive Summary of Findings 1 2. Recommended short-term actions 2 3. Recommended longer-term actions 3 4. Abstract 4 5. Introduction 5 6. Personnel 6 7. Study sites 7 8. Methods

a. Vegetation measurements 9 b. Sign based surveys versus camera-trapping 10 c. Survey objectives 11 d. Survey design - location of surveys 12 e. Sampling units – camera-traps 12 f. Location of camera-traps 12 g. Arrangement of camera-traps 13 h. Analysis 13 i. Relative abundance/activity 14 j. Absolute abundance 14 k. Activity patterns 15 l. Human impacts 15

9. Results a. Vegetation structure 15 b. Sampling efforts 17 c. Species richness 17 d. Tiger captures 19 e. Tiger abundance 19 f. Activity patterns 22 g. Human use 22

10. Discussion a. Significance of tigers 25 b. Tiger distribution 25 c. Tiger abundance 26 d. Relationship between index of abundance and density 26 e. Population size 27 f. Threats to tigers 29 g. Suggested priorities for tiger conservation 32

11. References 35 List of figures

1. Location of nine survey sites in Peninsular Malaysia (Dec ’97-Dec ’99) 8 2. Plot showing distribution of study sites along PC1 and PC2 axes. 17 3. Species accumulation curves for large mammals at nine sites in 18

Peninsular Malaysia. 4. Activity levels (%) of carnivores; (A) tigers and leopards (B),

clouded leopard/Asiatic golden cat, (C) Malayan sunbear/Dhole. 23 5. Activity levels (%) of tigers potential prey species;

(A) Common muntjak, Wild boar and mouse deer spp., (B) Sambar and Malayan tapir. 24

6. Activity levels (%) of tigers and humans. 24

iii

List of tables 1. Landscape context of nine tiger study sites 9 2. Component matrix showing loadings of 11 habitat structure

variables on four principal components. 16 3. Sampling effort for tiger survey at nine sites in Peninsular Malaysia. 18 4. Habitat use and relative abundance for tigers at nine sites. 19 5. Estimates for tiger abundance at sites in Peninsular Malaysia 21 6. Maximum distances moved and sample density estimates for tigers

at sites in Peninsular Malaysia 21 7. Human traffic at nine sites in Peninsular Malaysia. 25 8. Estimated numbers of tigers in reserves and TCUs in Peninsular Malaysia 29 9. Targets for tiger conservation with various time and spatial scales 34

Appendices

I. Site descriptions 40 II. Records from 174 camera-trap set-ups at nine sites in Peninsular Malaysia 45 III. List of reptiles, birds and mammals recorded by camera-trap survey at

nine sites in Peninsular Malaysia (December 1997-December 1999) 46 IV. Estimation of tiger abundance for sites where single tigers were caught 50

Cover photo; Tiger of Taman Negara National Park. Courtesy of Dr Kae Kawanishi

1

Executive Summary of Findings

1. Malaysia is one of fourteen range countries for the Tiger (Panthera tigris) and extensive tracts of montane forests and fragmented lowland forests in Peninsular Malaysia form high potential habitat for the species. These same tiger areas are recognized as ecoregions of global importance for biodiversity conservation.

2. In the last two decades, tigers have become a species of increasing national

concern due to the conflicting issues of increasing frequency of livestock depredation and human injury, but also increasing concern over the conservation future of the species.

3. With increasing demands on public lands to support economic development,

habitats for tigers and other wildlife continue to be compromised. As part of Malaysia’s strategy of balancing development with conservation, there is an urgent need to incorporate tiger conservation into national development strategies.

4. The Tiger is a wide-ranging species with relatively large area requirements,

but also relatively high resilience to human disturbance. Therefore it is a useful indicator of ecosystem health since only where sufficient habitat, prey, and water occur can the Tiger survive. Also, in Malaysia, the Tiger is a nationally revered species that is Totally Protected by law. For these reasons it provides a convenient flagship and target species for monitoring.

5. During 1997-2001 a national tiger conservation program was conducted by the

Department of Wildlife and National Parks (DWNP) with the support of international conservation agencies Wildlife Conservation Society (WCS), WWF – Malaysia, and the University of Florida. The objectives of the programme were to assess the current conservation status of tigers in Peninsular Malaysia, study tiger ecology, use the results to make management recommendations, and develop staff capacity to manage tigers. This report covers fieldwork conducted by DWNP and WCS during 1997-1999.

6. Nine sites were chosen for tiger survey across four states: Pahang, Trengganu,

Kelantan and Perak. Eleven variables describing microhabitat structure were measured at each sampling location. Principal components analysis (PCA) was used to identify two major gradients in habitat disturbance and identify relatively disturbed sites (Lepar, Gunong Tebu, Ulu Temiang) from relatively less disturbed sites (Temenggor, Taman Negara, Bintang Hijau).

7. Surveys involving 6,259 trapnights of sampling using 174 camera-trap setups

revealed the presence of tiger and 38 other mammals, almost half of which were globally threatened species. Also recorded were twelve species of large reptile and bird including five globally threatened or near-threatened species.

8. Ten individual tigers were recorded from 51 photographs at six of the nine

sites. Tigers were detected at a quarter of all sampling locations (range=0-

2

48% per site), and at a frequency of 4.8±0.5 detections/100 trapnights (range=0-5.9 detections/100 trapnights per site).

9. Activity patterns were inferred for tiger, other carnivores, and prey species

from the time imprinted on photographs. Tigers showed a crepuscular pattern of activity and apparently avoided periods when human traffic was greatest. Human activity was recorded at all sites except for Bintang Hijau. Overall human traffic occurred across 20% of the sampling areas.

10. Photographs from camera-traps were used to identify tiger individuals from

their unique stripe patterns. The photographs were used to generate capture histories for tigers. These were used along with a capture-recapture approach to determine statistically the probability of capture for tigers and sample estimates of abundance at different sites. Coefficients of variation for abundance estimates ranged from 2 to 392% reflecting low numbers of recaptures.

11. Tiger density was estimated from the sample abundance estimates and by

assuming a sampling effective area for each site. This area was calculated by adding a buffer to the configuration of camera-traps equal to the absolute maximum movement distance (AMDM) of tigers. Tiger density varied from extremely low (2 tigers/100km2) at Temenggor.

12. Despite intensive sampling, because the capture rates for tigers were low, and

captures took place over a one-year period, extrapolating a statistical estimate of tiger abundance across Peninsular Malaysia is difficult at best. Nonetheless, population estimates for tigers were attempted for individual reserves, for Tiger Conservation Units (TCUs) and for all areas in Peninsular Malaysia by assessing (1) population density, (2) forest size and (3) a correction factor to take into account avoidance of habitat due to roads, powerlines, settlements and other human infrastructure. While these estimates are very rough indeed, they provide an estimate of the national population of 184-305 tigers, most of which occur in one Level I TCU (TCU#129; 173-257 tigers). This is less than half of the previous estimate for Peninsular Malaysia.

13. Tigers are threatened by a range of factors chiefly habitat loss and

fragmentation, hunting of prey, commercial trade in live animals, dead animals and their parts, and harassment and displacement. Immediate action is needed to reduce these threats in the short-term (next five years) and to recover populations of wild tigers in Peninsular Malaysia in the longer-term (5-20 years from now).

Recommended short-term actions 1. Ecological monitoring. Central to a recovery plan would be the establishment

of tiger and prey monitoring programs at key sites, landscapes and Level I and II TCUs, along with training to build research capacity for key staff. Monitoring would focus on existing breeding populations of female tigers. In addition, protected area databases need to be established, and staff assigned to

3

manage the databases, so that data from research and monitoring activities may be used to detect population trends and made available for management planning.

2. Tiger protection. Monitoring programs would be combined with enforcement

training and the design of patrolling and anti-poaching systems to maintain and expand tiger populations at key priority sites. These programs would involve recruiting people in local communities who will provide information about who is poaching in the forest.

3. Trade research. Live and dead tigers from Peninsular Malaysia are potentially

traded across borders with neighbouring countries, especially those with relatively lax domestic laws and high demand. Better collaboration is needed between agencies concerned with enforcement to investigate and suppress this illegal trade.

Recommended longer-term actions 4. Creation of new protected areas. Only one area Taman Negara National Park

is large enough (>3,000km2) to support viable populations of tigers in the long-term. Many protected areas are smaller than 500km2. New protected areas need to be established in key tiger areas and existing areas increased in size.

5. Landscape-level planning for tigers. Tiger habitat throughout Asia is

continually being lost. Through use of a GIS database, the rate of habitat loss in Malaysia can be estimated, future loss may be projected and effects of creation of forest edges predicted. Identification of large blocks of habitat or potential habitat which are under threat is a potentially useful management tool, in particular since the loss of certain areas, such as the forest corridor between Taman Negara and Krau Wildlife Reserve, may have a negative impact on protected areas and the ability of tigers to move about the landscape.

6. Arresting habitat degradation. In some key sites habitat degradation is largely

a result of activities of local communities. Specific actions need to be taken to stabilize blocks of tiger current and potential tiger habitat.

7. Monitoring human-tiger conflicts. Information on tiger mortality must be

assembled to quantify the rate and reasons why tigers are getting killed in Peninsular Malaysia. This information can be used to increase awareness to garner support for conservation programmes.

8. Increasing public awareness. WCS and DWNP have assembled a considerable

amount of information and camera-trap photographs. These could be used creatively to increase public awareness and motivate field staff.

9. Tiger area management. Management Plans such as that developed for Krau

Wildlife Reserve need to be developed for other key tiger sites so that habitats may be protected and tiger populations recovered.

4

10. Revising national tiger action plan. Information contained in this report together with separate reports on tiger ecology for Taman Negara provides the foundation for revising the National Tiger Action Plan.

Abstract

Across its global range the tiger is declining due to factors understood by researchers

and managers. However, effective management of tiger populations suffers from a

lack of information on their status and distribution. In Peninsular Malaysia, tigers are

a national emblem and Totally Protected species. Recognizing this importance and

their potential as an umbrella for the protection of forested landscapes and contained

wildlife, and the need for developing staff capacity for managing wildlife, the

Department of Wildlife and National Parks (DWNP) embarked on an ambitious

program to ascertain the health of tiger populations in representative habitats across

Peninsular Malaysia. From 1997-2001 surveys for tigers and other large mammals

were executed at twelve sites across four states, Perak, Terengganu, Kelantan and

Pahang. Camera-traps were the primary tool used in surveys. Surveys were jointly

designed and implemented by central and state level staff of DWNP, the Wildlife

Conservation Society, and the University of Florida, Gainesville. Results from

surveys at nine sites conducted between December 1997 and December 1999 are

reported here. Surveys revealed 39 species of mammals almost half of which were

globally threatened species. Tigers were confirmed from six sites in the Main Range

and Greater Taman Negara landscape, including multiple locations inside a Level I

Tiger Conservation Unit (TCU#129), inside Level II TCU#130, and near other Level

II and III TCUs. Tiger abundance was estimated using photographic records of tiger

individuals, capture–recapture statistical analysis, and a sample area defined based on

the maximum distance that individual tigers move during the sample period. The tiger

population for Peninsular Malaysia is conservatively estimated as 235 (range=184-

305). Key threats facing tigers are habitat loss and fragmentation, hunting of prey,

commercial tiger trade and harassment and displacement. A framework for

organizing priority targets for tiger conservation is described.

5

Introduction

One hundred years ago when Peninsular Malaysia was extensively covered with

forest, tigers were hunted regularly for sport (Wallace 1869; Whitney 1905). The

combination of this plus loss of habitat meant that by the early 1950’s tigers had

disappeared from Penang (Locke 1952), and later from Melaka and Perlis. Up until

1976 the Indochinese tiger was considered a pest, with hunting combining with loss

of habitat to cause serious declines in Peninsular Malaysia (Blanchard 1977). After

1976, the tiger became a totally protected species (Samsudin & Elagupillay 1994). In

spite of this change in legal status, little was known about the status of wild

Indochinese tiger in Peninsular Malaysia (Elagupillay & Wan Shahruddin 1999;

Topani 1990). While surveys revealed tigers in some places (Ratnam et al. 1995)

until the end of the last century most information was derived from unconfirmed

reports (Rabinowitz 1999).

To plan for the future conservation of tigers, knowledge of their current status and

threats is required. In the late 1990’s an attempt was made to identify potentially

viable tiger habitat across the tiger’s entire range across Asia. Natural vegetation was

divided into three grades of tiger habitat, using current available information on

habitat integrity, poaching pressure and population status (Dinerstein et al. 1997).

These graded habitats are known as Level 1, or Level 2 or Level 3 Tiger

Conservation Units (TCUs). Level 1 TCUs are thought to offer the highest probability

of persistence of tiger populations over the long-term (Dinerstein et al. 1997). One

Level 1 TCU (#129) was described for Peninsular Malaysia. It covers the northern

part of the Peninsular and encompasses parts of Kedah, Perak, Selangor, Kelantan,

Terengganu and Pahang, and includes Taman Negara and the forested part of

southern Thailand, a total of 27,469km2 of potential tiger habitat. Two Level 2 TCUs

(2,472km2) and 13 Level 3 TCUs (4,344 km2) were also identified in the Peninsular.

A striking feature of this ‘framework analysis’ was that for Peninsular Malaysia, and

some other countries in Mainland Southeast Asia, there existed large gaps inside the

TCUs for which no recent information on tiger status was available. Added to this

was the fact that most records of tigers compiled by the Malaysian Department of

Wildlife and National Parks (DWNP) came from forest-farmland interfaces

(Elagupillay et al. 2001) and at least some of these were unconfirmed reports

6

(Rabinowitz 1999). This reflects the historical priorities for tiger management which

focussed on human-tiger conflict resolution (DWNP 1992; Elagupillay 1983, 1984;

Elagupillay et al. 2001; Elagupillay & Wan Shahruddin 1999; Jasmi 1986; Jasmi &

Wan Shahruddin 1998; Khan 1987; Kitchener 1961; Samsudin & Elagupillay. 1994;

Topani 1990) and the need for staff training in tiger survey methods.

Recognizing the paucity of information on tigers from areas inside forest reserves,

and at the same time desiring to mitigate the effects on tiger survival of future

intrusions into these habitats by national development programmes, the DWNP

embarked on an ambitious programme with partners Wildlife Conservation Society,

the University of Florida, Gainesville, and WWF Malaysia. The programme had

three broad goals:

1. To determine the distribution and status of tigers throughout Peninsular

Malaysia.

2. To collect information on tiger numbers, movements, behaviour, home range

size and habitat use in a key tiger site identified from the survey programme.

3. To transfer technology and train Malaysian wildlife biologists in techniques

for surveying and studying tigers to allow for the future management and

conservation of tiger populations.

This report details the findings of a field survey and training programme conducted by

DWNP with support from the Wildlife Conservation Society from December 1997 to

December 1999. Recommendations for future research and management action are

provided. Results of an ecological study mounted by the University of Florida,

Gainesville are reported elsewhere (Kawanishi 2002; Kawanishi et al. 1999) but are

referenced in this report. Together, these two studies provide current baseline

information on the status, distribution and threats facing tigers in Peninsular Malaysia,

and provide a basis for updating the Malaysia Tiger Action Plan (DWNP 1995).

Personnel

The lead agency for the program was the DWNP. All work was done in full

collaboration with DWNP. The tiger programme was implemented for DWNP by;

1. En. Musa Nordin, Director-General

7

2. En. Sahir Othman, Director of Research and Wildlife Conservation

3. En. Sivananthan Elagupillay, Head of Wildlife Management Unit

4. En. Jasmi Abdul, Director of Perak and local staff

5. En. Mohd. Shamsuddin Saari, Director of Terengganu and local staff

6. En. Rahim Ahmad, Director of Kelantan and local staff

7. Mr Wan Shaharuddin Wan Noordin, Tiger Coordinator

At the time the project was commenced, DWNP lacked sufficient trained staff to

conduct tiger survey. Thus during the two year project, three foreign experts were

recruited to assist with training and data collection;

1. Dr Ruth Laidlaw. Responsible for direct field supervision of surveys in target

areas, and for detailed training of DWNP staff in tiger and prey survey

techniques e.g. rapid survey techniques, camera-trapping, data recording and

management, and assimilation of information into a database.

2. Ms Kae Kawanishi. Conducted an ecological study of tigers at Taman

Negara and trained DWNP staff in ecological study methods, e.g., camera-

trapping, collection, management and analysis of data.

3. Dr Antony Lynam. Jointly responsible, with Dr Laidlaw, for overall project

design and initial training of DWNP staff. He provided advice on survey

implementation during the programme, assembled and analyzed data, and

compiled this final report.

In addition to project personnel, Drs Elizabeth Bennett, Joshua Ginsberg, and

Timothy O’Brien provided expert comments on earlier drafts of this report.

Study Sites

In this study, tiger surveys were conducted in forest blocks with greatest potential for

conserving tigers in the long term (DWNP unpublished data; Dinerstein et al. 1997;

(Wikramanayake et al. 2002). Surveys were completed in a total of nine sampling

plots in Perak, Terengannu, Kelantan and Pahang (Fig. 1). Descriptions of the sites

are given in Appendix I. Seven sites were located inside Level 1 TCU (#129), one in

Level II TCU (#130) and one adjacent to Level II TCU (#134) to determine in which

parts of these potential habitat tigers persist and why. These sites were chosen since

8

such knowledge is critical in allowing DWNP to take appropriate action to conserve

tigers. The information would also provide invaluable baseline data for future

monitoring.

Fig. 1. Location of nine tiger survey sites in Peninsular Malaysia (Dec ’97-Dec ’99).

Within the TCUs, study sites were selected to cover a range of different forest types

and management histories, e,g. riverine forest, lowland, hill and montane forest;

logged and unlogged forest; protected forest and forest reserves; forest fragments,

edge forest and core forest and tiger-human conflict zones and non-conflict zones

9

(Table 1). One site was located inside a protected area, Taman Negara National Park.

Totally protected areas (national parks and wildlife reserves) constitute approximately

5.7% of the land area in Peninsular Malaysia. All other sites were in Forest Reserves.

Table 1. Landscape context of nine tiger study sites (see also Appendix I).

Study site Location Area Logging history

Temenggor central * continuous unlogged

Bintang Hijau central >1,400 km2 **unlogged/logged

Jengai central * continuous **unlogged/logged

Gunung Tebu edge * continuous **unlogged/logged

Ulu Temiang edge

10

4. DISTANCE MEDIUM TREE - Average distance to nearest tree < 30 cm gbh

and >1.3 m in height, measured in four quadrants at the point

5. CIRCUMFERENCE MEDIUM TREE - Average circumference of nearest

tree < 30 cm gbh and > 1.3 m height, measured in four quadrants at the point

6. UNDERSTORY 5m - Median understory density at 5m (scored out of 10,

with 1 = very poor visibility), measured in four quadrants at the point

7. UNDERSTORY 10m - Median understory density at 10m (scored out of 10;

with 1 = very poor visibility), measured in four quadrants at the point

8. WOODY CLIMBERS – Number of woody climbers ≥ 3cm diametre within

20m (scored out of 4)

9. BAMBOO – Number of bamboo clumps within 20m (scored out of 4)

10. RATTAN – Number of rattans ≥ 2cm diametre within 20m (scored out of 4)

11. PALM – Number of palms ≥1.3m height within 20m (scored out of 4)

Principal components analysis (Pielou 1984; Tabachnick & Fidell 1983) was used to

identify independent patterns of covariation among the habitat structure variables.

PCA constructs new, synthetic variables that are linear combinations of the raw

variables. Each sampling point can be scored on each PCA axis based upon the values

of its raw variables.

Sign based surveys versus camera-trapping

The Indochinese tiger occurs at low densities and is difficult to detect in the humid

forest conditions of Peninsular Malaysia (Elagupillay & Wan Shahruddin 1999) and

forest habitats elsewhere in its range (Duckworth and Hedges 1998; Rabinowitz

1993). Until recently, sign counting was the most commonly used and readily

available method for assessing tigers in Malaysia (Topani 1990). However, tiger

census using indirect sign alone can be fraught with difficulties as it depends on the

substrate, the skill of the tracker, and the long-term knowledge of an area and of the

tigers within (McDougal 1999). Furthermore, tiger tracks are seldom visible on

substrates covered with leaf litter, they can be difficult to distinguish from leopard

tracks and it is rarely possible under these conditions to distinguish one individual

tiger from another by using tracks. Countrywide total censuses using tiger tracks

were first introduced in India in the 1970’s (Panwar 1979; Karanth 1987). The

methods used (pug-mark identification) have since been tested and proved not to be

11

valid (Karanth 1987, 1995; Karanth & Nichols 1998; Karanth et al. 2003). In

addition, surveying using other animal signs is problematic. For example, prey

carcasses and scats decompose rapidly in hot, humid conditions and can also be

difficult to distinguish from those of leopards (Duckworth and Hedges 1998;

Elagupillay & Eh Pat 1997).

A more reliable method both for detecting tigers and estimating their abundance

involves the use of remote camera-photography (Cutler & Swann. 1999). For

example, camera-traps have been used to establish tiger presence-absence (Griffiths &

Schaik 1993a). Each tiger has its own unique coat pattern (Schaller 1967) thus

permitting their individual identification (Franklin et al. 1999). Thus camera-traps

can be used for estimating tiger relative abundance (Lynam et al. 2001) and absolute

abundance (Karanth 1995; Karanth and Nichols 1998; O'Brien et al. 2003). Although

there are relatively high initial costs of investment involved in camera-trapping it

appears to be the most appropriate technique for large cat inventory over a wide range

of environmental conditions (Silveira et al. in press).

Survey objectives

Field managers desire to know several important pieces of information about tigers;

(1) the distribution of tigers and prey species across a reserve or region, (2) how tiger

abundance compares between areas where they are present, and (3) information on

specific threats to tigers, and how to mitigate those threats. In this study we addressed

the first objective by determining the pattern of presence or absence of tigers and

other large mammals at sampling points inside representative habitats. A second

objective was to compare the frequency of use of different habitats between sites, and

also to achieve sample estimates of tiger numbers for areas of forest at each site.

Finally, information on threats to tigers was gleaned from records of human intrusions

at each site via camera-trap records and other field observations. This information

was supplemented by general appraisals of habitat suitability from other studies

(Laidlaw 2000; Kawanishi 2002).

Survey design - location of surveys

Since 1976, the DWNP Tiger Research Unit has collected information on tigers from

field observations of researchers and other staff, and via reports by farmers and other

12

local people. The latter reports were associated with occurrences of livestock

depredation or attacks on humans. While this has provided valuable information on

tiger occurrence, the collection of field data was done in response to calls from the

public and was non-systematic in its coverage. In the current study, DWNP staff and

local people were interviewed to determine potential tiger areas in each State.

Selection of survey sites was therefore non-random but based on the best available

information.

Sampling units – camera-traps

Data were collected using camera-traps as they can provide non-invasive unbiased

information on animal movements, activity patterns (van Schaik & Griffiths 1996)

and abundance (Karanth 1995). The camera-trap devices used (CamTrak South Inc.

Georgia, U.S.A.) consisted of a weatherproof Infinity Mini DLX Olympus camera

with a 35mm lens and flash, and a passive infrared sensor, all housed in a waterproof

casing. The cameras are triggered when the sensor detects a differential in heat and

motion across a conical beam in front of the camera-trap, and a photograph is taken.

The cameras included a databack and could be programmed to record date and time

(24 hour clock) on the film (Fig. 2). Fuji 400ASA, 36 exposure film was used with the

camera-traps. A heavy-duty cycle lock was used to secure each trap to a tree. In most

cases this prevented theft. The cameras were programmed with a 3-minute time delay

so that animals lingering in front of the camera for short periods of time would not

cause the camera to take an entire roll of film.

Location of camera-traps

Camera-traps were placed to maximize the probability of detection (p) for tigers

(Karanth and Nichols 1998, 2002). Camera-traps were set at ‘optimal’ locations on

animal trails and roads, and mounted 40cm above the ground, and 3-5m (the focal

length of the camera) from the middle of the trail. Locations for traps were places

where either tiger sign, or sign of tiger prey species was detected. Surveys were

constrained by remoteness with most sites accessible only on foot. Traps could not be

checked on a daily basis and were left in the forest for 30-35 days set for operation

day and night. One “trap-night” in this paper, therefore, refers to one 24 hr period

(Kawanishi et al. 1999). Information on details of camera-trap setup and retrieval

were recorded on a standardized datasheet and the number of trap-nights over which a

13

trap was functional was determined. Surveys were conducted from December 1997 to

December 1999 inclusive.

Arrangement of camera-traps

At each study site, thirty camera-traps were established at 25 ‘optimal’ locations on a

10 x 4 km grid. Each grid was divided into 40 subunits of 1 km2. At twenty alternate

1 km2 grids, single camera-traps were positioned at ‘optimal’ locations. At five

opportunistic locations, usually places where tiger sign was detected, at junctions of

trails, trails on ridgelines or water-crossings, or on saddles between hills, pairs of

traps were positioned on opposite sides of the trail/road in "check-points". Check-

points allowed the identification of individual tigers by allowing both sides of an

animal to be photographed simultaneously. Field crews attempted to distribute the

“check-point” pairs across the grid to achieve maximum coverage. Additional details

on procedures for selecting camera-traps, and deployment of traps are given in

(Laidlaw 1998).

Analysis

Films retrieved from traps were processed and information on time, date and species

recorded from each frame was entered into a database and summary information

extracted onto a standardized datasheet (see also Kawanishi et al. 1999). Independent

events or ‘detections’ were defined as (1) consecutive photographs of different

individuals of the same species, (2) consecutive photographs of individuals of the

same species taken more than 0.167 hrs apart. Sometimes, the time record on certain

photographs was illegible. This was due to flash overexposure in the corner of the

photograph where the time was marked. Occasionally, also, the time was not printed

on the photograph. This was largely due to human error setting up and programming

the camera. These photographs without inscribed times, were not used in our

analyses.

Relative abundance/activity

For each species, two relative abundance/activity indices (RAI) were calculated and

follows O’Brien et al. (2003);

14

1. RAI1 = number of days from the start of trapping to the first record of a tiger.

This measured the effort required to record a tiger from survey and should

decrease with increasing population density (Carbone et al. 2001).

2. RAI2 = number of detections divided by the number of trap-nights during

which the trap was functional at a location. This is the reverse of RAI1 and

should increase with population density. RAI2 was scaled to detections per

100 trapnights.

Absolute abundance

Photos of tigers from camera-traps and capture-recapture statistical models were used

to obtain sample estimates of tiger density at each study site (Karanth 1995; Karanth

and Nicols 1998). The procedures followed the recommendations of Karanth and

Nichols (2002) and were analyzed using Program CAPTURE (Otis et al. 1978;

Rexstad & Burnham. 1991; White et al. 1982). CAPTURE assumes that populations

are ‘closed’, i.e., that there are no gains or losses of individuals e.g from births,

deaths, immigration or emigration during the surveys. Since most sampling locations

employed only one camera-trap, so photos from only one side of an animal were

available, analyses were done using either right side or left side photos only. This

lowers capture probability (p) but it is still possible to use the method for estimation

(Karanth 1995; J.D. Nichols pers. comm.).

Data analysis was done as follows;

1. individual tigers were identified from photographs and given unique codes

2. a capture history – a row of ‘0’s and ‘1’s - was developed for each coded

animal with ‘0’s indicating no detection (no photograph) or ’1’ indicating

detection (photograph taken) for a given sampling interval i.

3. each sampling interval was a group of 3 trapnights. Up to a maximum of 17

sampling intervals were used at each of the nine study sites

4. following Karanth and Nichols (1998), Model Mh was used to model capture

probability. This model allows for variability in capture probability among

individuals since tigers in different age classes i.e., cubs, juveniles, transients

and adults, might have different capture probabilities due to different

movement behaviour (Karanth and Nichols 2002). It also assumes that an

individual tiger’s capture probability stays unchanged during the survey.

15

Activity patterns

To assess the ecological overlap or separation of carnivores and prey, activity budgets

were compared among species using camera-trap photographs imprinted with times.

Patterns of activity were represented by frequency histograms with time divided into

hourly intervals.

Human impacts

Human activity can have significant impacts on tigers and other wildlife (Griffiths &

Schaik 1993b). In order to document this for each site, human traffic was measured

as number of photographs of humans/100 camera-trapnights (i.e. RAI2), and human

activity patterns were compared with those of tigers.

Results

Vegetation structure

The 11 habitat structure variables were measured at 151 points where camera-traps

were established. These variables can be summarized by 4 composite principal

components variables (Table 2). The four composite variables explained 65% of the

variation in the dataset. The loadings for each variable described what each of the

four new variables described in terms of gradients in vegetation structure:

1. PC1. This variable described a disturbance gradient with increasing canopy

cover and decreasing distance to large trees (i.e. increasing closure of forest),

decreasing distance of medium-size trees (increasing density of understory),

and increasing density of woody climbers and palms.

2. PC2. This variable also described a disturbance gradient of increasing

visibility of understory vegetation and decreasing density of medium-sized

trees (i.e. increasing openness of understory).

3. PC3. This variable described a gradient of increasing size of medium-sized

trees and increasing density of rattan.

4. PC4. This variable described a gradient of increasing density of bamboo.

Principal components can be used to distinguish sites along the continuum from least

disturbed to most disturbed. When mean values for PC1 and PC2 were plotted on two

axes, sites that were least disturbed (Temenggor, Taman Negara, Bintang Hijau) were

16

identified from more disturbed sites (Gunong Tebu, Lepar and Ulu Temiang)(Fig. 2).

Other sites (Jengai, Ayer Ngah) lie in between these two sets of points.

Table 2. Component matrix showing loadings of 11 habitat structure variables on four principal components. Significant principal components have eigenvalues >1.0. Bold denotes factor loadings >0.5. These variables described the gradients represented by the principal components.

Components Variable measured

PCA1 PCA2 PCA3 PCA4

CANOPY COVER -.850 .094 **** .006

DISTANCE LARGE TREE -.812 .236 .264 .175

CIRCUM LARGE TREE .319 -.154 -.279 .326

DISTANCE MEDIUM TREE -.525 .595 .239 .119

CIRCUM MEDIUM TREE -.339 -.455 .583 .0019

UNDERSTORY 5m .459 .635 -.298 ****

UNDERSTORY 10m .280 .578 .287 -.286

WOODY CLIMBERS .743 .041 .324 ****

BAMBOO .128 .152 **** .873

RATTAN .453 .082 .655 .180

PALM .674 **** .199 .101

Eigenvalue 3.4 1.4 1.3 1.0

% variance explained 30.8 13.0 11.7 9.5

Cumulative variance explained 30.8 43.8 55.5 65.0

17

-0.5

-0.4

-0.3

-0.2

-0.1

0

0.1

0.2

0.3

0.4

0.5

-0.6 -0.4 -0.2 0 0.2 0.4 0.6 0.8 1

More disturbance - PC1 - Less disturbance

Fig. 2. Plot showing distribution of study sites along PC1 and PC2 axes. These axes describe gradients in forest disturbance. One site (Cameron Highlands) could not be

portrayed because of missing habitat data.

Sampling efforts

A total of 174 camera-trap units were deployed at the 151 sampling locations during

the surveys. These units recorded a total of 2,979 photographs from 6,259 trapnights,

at the nine study sites (Table 3, Appendix II). The overall rate for camera-trapping

was, therefore, roughly one photograph taken every three days. When test

photographs (taken during setup or takedown) were excluded, camera-traps failed at a

rate of 6% (Appendix II).

Species richness

Tigers and 38 other species of mammals, as well as humans were recorded by camera-

traps at the nine sites (Appendix III). Almost half of the mammals (16 species) are

globally threatened or data deficient species (Hilton-Taylor 2000), and two are near

threatened species. Twelve were CITES listed species, 5 of them Appendix I species.

Because sampling efforts varied between sites, large mammal richness cannot be

compared directly. Also species numbers did not asymptote at any site during the

periods of sampling (Fig. 3). However, the rates of species accumulation were

relatively higher at three sites (Ulu Temiang, Gunong Tebu and Lepar), relatively

medium at three sites (Ayer Ngah, Temenggor and Taman Negara), and relatively

lower at three sites (Bintang Hijau, Jengai and Cameron Highlands).

Jengai

Taman Negara

Temenggor

Bintang HijauAyer Ngah

Ulu Temiang

Gunong Tebu

Lepar

18

0

2

4

6

8

10

12

14

16

18

1 4 7 10 13 16 19 22 25 28 31 34 37

Time (days)

Ayer NgahBintang HijauCameron HighlandsGunong TebuJengaiLeparTemenggorTaman NegaraUlu Temiang

Fig. 3. Species accumulation curves for large mammals at nine sites in

Peninsular Malaysia.

Camera-traps also recorded 12 species of reptiles and birds. Three birds are globally

threatened and two are near threatened species. Six species were CITES Appendix II

or III species.

Table 3. Sampling effort for tiger survey at nine sites in Peninsular Malaysia.

Site Start End # tigers

(recaps) Camera-trap

setups

Trap

nights

1. Temenggor 10.12.97 2.3.98 2(4) 22 785

2. Bintang Hijau 12.2.98 26.3.98 2(4) 19 646

3. Jengai 28.4.98 4.6.98 0 15 467

4. Gunong Tebu 26.6.98 27.8.98 1(11) 27 807

5. Ulu Temiang 30.8.98 30.9.98 3(11) 21 557

6. Ayer Ngah 9.10.98 13.11.98 1 18 562

7. Cameron Highlands 7.2.99 22.3.99 0 13 495

8. Taman Negara 26.5.99 14.8.99 0 16 768

9. Lepar 28.9.99 23.12.99 1(4) 23 1172

Totals 10(34) 174 6,259

19

Tiger captures

A total of 51 photographs of ten individual tigers were taken at 36 sampling locations

across the nine sites. Tigers were recorded at a quarter of all locations (Table 4) or, at

each site, at 18 ± 6% of sampling locations. Tigers were not recorded at Taman

Negara, Cameron Highlands and Jengai but occurred at almost half of the locations at

Ulu Temiang. On average one tiger photograph was taken for every 122 trapnights of

sampling. Average RAI1 at locations where tigers were detected was just over 15

trapnights/first capture (Table 4). Overall RAI2 was 0.82 photographs/100 trapnights,

with average per point RAI2 less than 5 photographs/100 trapnights.

Table 4. Habitat use and relative abundance for tigers at nine sites.

Site %

locations detected a

RAI1b

( x ± SE)(n)

RAI2 c

( x ± SE)(n)

1. Gunong Tebu 26 9.1±3.2 (7) 5.9±0.6 (7)

2. Ulu Temiang 48 11.8±2.6 (10) 5.7±0.9 (10)

3. Bintang Hijau 32 12.7±2.4 (6) 5.3±1.9 (6)

4. Temenggor 16 12.4±3.3 (5) 4.3±0.6 (5)

5. Ayer Ngah 6 7 (1) 3.1 (1)

6. Lepar 26 37.8±6.2 (6) 2.3±0.3 (6)

7. Cameron Highlands 0

8. Taman Negara 0

9. Jengai 0

All sites 25 15.8 ± 2.3 (34) 4.8 ± 0.5 (34) a Naïve estimate of occupancy i.e. no. locations detected/total no. locations. b Mean/standard error (number of locations) days to first capture (number of locations). c Mean/standard error (number of locations) number of detections/100 trapnights.

Tiger abundance

Between 1 and 3 individual tigers were detected at six sites. For sites where

recaptures of tigers were made, Program CAPTURE was used to obtain sample

abundance estimates for tigers. Captures/recaptures were divided into 11 to 20

sampling occasions for analysis. Tigers were detected with a range of probability

values (Table 5) indicating possible differences between sites. Estimates of tiger

numbers had wide variance reflecting low numbers of recaptures at sites (Table 5).

20

For sites where only single tigers were recorded Program CAPTURE could not

calculate a detection probability ( ˆ p ) or SE for ˆ N . To estimate tiger density, these

parameters were first derived using a method described in Appendix IV.

Three northern sites (Temenggor, Ulu Temiang, and Gunong Tebu) that are part of

the Northern Forest Complex were all sampled within a 10 months period (Table 3;

December 1997-September 1998). During the same period, an estimate of tiger

density was made using the same method across the border in contiguous forest at a

site inside Thailand’s Halabala Forest Complex (Bang Lang National Park). If it is

assumed that the populations were demographically closed during this time (i.e., no

births, deaths, or movements in and out), then combining the recaptures of tigers from

these sites gives a sample estimate for tiger abundance for the transboundary northern

Peninsular Malaysia/southern Thailand forest complex. Because these estimates

assume a closed model for estimation, and we assume the population was indeed

closed over the timeframe analyzed, the results are indicative but not definitive.

Mean sample density ˆ D and range of density values at each site were calculated using

the sample estimates of tiger numbers ˆ N and by assuming a sampling effective area ˆ A at each site. Translation of the abundance estimate into density was done by:

ˆ D ' =ˆ N '

ˆ A (W ) ,

where )(ˆ WA is the sampling area with buffer width W. )(ˆ WA was calculated by

linking the outermost locations of camera-traps and adding a buffer with radius W

equivalent to half the absolute maximum distance moved (AMDM) between

recaptures for individual tigers (Wilson & Anderson. 1985; Kawanishi 2002) For

example, at the Bintang Hijau site, two tigers were captured and recaptured by

camera-traps. One female tiger ranged a maximum 9.75km between successive

points of capture/recapture. A male tiger ranged a maximum 5.25km between

locations. Buffer width (W) was then equal to 9.75km. The area of the plot plus

buffer was then (4 + 9.75km) x (10 + 9.75km) = 272km2 (Table 6).

21

Table 5. Estimates for tiger abundance at sites in Peninsular Malaysia*.

Site T Mt+1 p a ˆ N (SE) b CI CV (%)

1. Temenggor 12 2 0.0833 3(1.38) 3-9 46

2. Ulu Temiang 11 3 0.333 3(0.05) 3-3 2

3. Gunong Tebu 11 1 1

4. Bintang Hijau 14 2 0.1429 2(7.85) 2-2 392

5. Ayer Ngah 12 1 1

6. Lepar 20 1 1

Northern sites (1-3) 11 6 0.2468 7(1.38) 7-13 20

Halabala (Thailand) 8 3 0.2812 4(1.35) 4-10 34

Northern sites +

Halabala

11 9 0.2314 11(1.96) 10-19 18

* T=number of sampling occasions; Mt+1 = Number of animals captures; CI=95% confidence interval of estimate; CV (%)= coefficient of variation; a Detection probability estimated using Program CAPTURE; b Single sided M-R estimates using Program CAPTURE

Table 6. Maximum distances moved and sample density estimates for tigers at sites in Peninsular Malaysia.

Site Plot

size

(km2)

AMDM

a (km) )(ˆ WA ˆ D b ˆ D min -

ˆ D max

1. Temenggor 40 4.0 112 2.68 1.44-3.92

2. Ulu Temiang 40 8.2 154 d 1.95 1.92-1.98

3. Gunong Tebu 40 7.0 188 0.60 e 0.45-0.75

4. Bintang Hijau 40 9.8 272 0.74 0-3.63

5. Ayer Ngah 40 7.0 c 188 0.60 d 0.45-0.75

6. Lepar 40 7.3 195 0.58 d 0.43-0.72

Northern sites (1-3) 120 8.2 368 1.90 1.53-2.28

Halabala (Thailand) 40 10.5 297 1.35 0.89-1.80

Northern sites +

Halabala

160 10.5 543 2.02 1.66-2.39

a AMDM=Absolute maximum movement distance; b Sample estimate of tiger density; no. tigers/100km2; buffer width=AMDM/2; c No recaptures at this site, assumed same AMDM as Gunong Tebu (nearest site); d Calculated sampling area was larger than size of the reserve so the latter was used to estimate density; e No recaptures or single individuals. Density (D) = No. tigers (N) /Area, where N = No. tigers detected/p, and p=0.888 (p calculated from entire Malaysia study - Appendix IV).

22

Sample density estimates ranged from 0.58-0.60 tigers/100km2 at Lepar, Gunong

Tebu and Ayer Ngah to 2.68 tigers/100km2 at Temenggor (Table 6). A sample

estimate for tiger density in the northern Peninsular region was 1.90 tigers/100km2

(Range=1.53-2.28). If the northern Malaysia sites and Halabala, Thailand are

combined, an estimate of tiger density for the transboundary forest complex that

comprises TCU#129 was 2.02 tigers/100km2 (Range=1.66-2.39).

Activity patterns

Camera-trap photographs revealed that tigers were active at all times of the day and

night but had bi-modal peaks of activity from 06:00-11:00hrs and 17:00-21:00hrs

(Fig. 4 A). This crepuscular pattern was significant (P

23

0

5

10

15

20

25

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24

Hour

Tiger

Leopard

A

0

2

4

6

8

10

12

14

16

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24

Hour

Clouded leopard

Golden cat

B

0

5

10

15

20

25

30

35

40

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24

Hour

Sun bear

Dhole

C

Fig. 4. Activity levels (%) of carnivores; (A) tigers and leopards (B), clouded leopard and Asiatic golden cat, and (C) Malayan sunbear and Dhole.

24

0

2

4

6

8

10

12

14

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24

Hour

Muntjac

Wild boar

Mouse deer

A

0

5

10

15

20

25

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24

Hour

Sambar

Tapir

B

Fig. 5. Activity levels (%) of tigers potential prey species; (A) Common muntjak, Wild boar and mouse deer spp., (B) Sambar and Malayan tapir.

0

5

10

15

20

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24

Hour

Tiger

Human

Fig. 6. Activity levels (%) of tigers and humans.

25

Table 7. Human traffic at nine sites in Peninsular Malaysia.

Site %

locations detected a

% total traffic b

RAI2 c

( x ± SE)(n)

1. Gunong Tebu 26 6 28.7±23.4(7)

2. Ulu Temiang 33 6 37.4±26.6(7)

3. Bintang Hijau 0 0 0

4. Temenggor 23 4 14.4±10.5(5)

5. Ayer Ngah 33 8 45.9±15.9(6)

6. Lepar 22 3 21.2±14.5(5)

7. Cameron Highlands 8 3 52.6±47.4(2)

8. Taman Negara 12 2 2.4±2.4(2)

9. Jengai 1 4 55.9±31.4(2)

All sites 20 4 25.6±4.6(36) a Naïve estimate of occupancy i.e. no. locations detected/total no. locations. b No. human photographs x 100/total no. photographs. c Mean/standard error (no. locations) no. detections/100 trapnights.

Discussion

Significance of tigers

The tiger is a critically endangered species whose range has contracted to 95% of its

original extent in the past century (Dinerstein et al. 1997; Kawanishi 2002; Nowell &

Jackson 1996). Tigers have large area requirements and are particularly sensitive to

habitat fragmentation and edge effects (Woodroffe & Ginsberg 1998) including those

induced by roads (Miquelle et al. 1999). The tiger is therefore an indicator of

ecosystems in crisis (J. Seidensticker in Ginsberg 2001) especially since their absence

can potentially lead to the collapse of mammalian food chains (Seidensticker 2002).

As Asian landscapes become ever-more developed to accommodate humans, the tiger

will help us to monitor the impacts on wildlife and wilderness.

Tiger distribution

In this study, surveys across nine forest areas revealed the presence of tigers in

multiple parts of a Level I TCU (#129: Temenggor, Ayer Ngah, Gunong Tebu, Ulu

Temiang) as well as one Level II TCU (#130: Bintang Hijau) and adjacent to one

26

Level III TCU (#134: Lepar). Tigers were not detected at three sites, specifically a

site in the Cameron Highlands, Ulu Sat in the eastern part of Taman Negara, and

Jengai which lies to the southeast of Taman Negara. In contrast Kawanishi (2002)

found tigers to occur the central and western parts of Taman Negara National Park at

Merapoh, Kuala Terengan, and Kuala Koh.

Between sites the probability of detecting tigers ranged an order of magnitude (Table

4). Within sites, tigers were also evenly detected across survey plots and on average

occurred at around 25% of sampling locations (Table 4). That tigers were not

consistently found across available habitat suggests that factors that vary both within

and between sites influence their distribution. Clearly such factors need to be

identified and their effects understood in order to effectively manage tigers across

sites and landscapes.

Tiger abundance

In parts of its range in South Asia, tiger populations have been intensively surveyed in

open grasslands and deciduous forest habitats (Karanth 1995; Karanth and Nichols

1998; Karanth and Nichols 2000). However, few estimates of tiger abundance exist

for rainforest habitats where the species is difficult to see. In Bukit Berisan Selatan

National Park, Sumatra tigers occur at a density of 1.6 tigers/100 km2 (95% CI=1.2-

3.2) at (O’Brien et al. 2003). In Taman Negara, tigers ranged in density from 1.10

tigers/100km2 (95% CI=1.1-4.4) to 1.98 tigers/100km2 (95% CI=1.7-4.0)(Kawanishi

2002). In Thailand’s Halabala Forest Complex, contiguous with Malaysia’s northern

forest complex, tigers ranged in density from 1.2 tigers/100km2, to 1.4 tigers/100km2

(95% CI=1.3-3.4) (Table 4; Lynam unpublished data). At Kaeng Krachan, a

rainforest site in Peninsular Thailand, tigers occurred at slightly higher densities 3.5

tigers/100km2 (95% CI=3.5-9.2)(Lynam unpublished data). At Badhra, India also an

evergreen forest site, tigers occurred at similar density to Kaeng Krachan i.e. 3.4 ± 0.8

tigers/100km2 (Karanth and Nichols 2000).

Relationship between index of relative abundance and tiger density

Of particular interest to conservation managers is the relationship between the

accumulation rate of tiger photographs from camera-trapping and tiger density

(Carbone et al. 2001; O’Brien et al. 2003). If a monotonic relationship exists between

27

the two variables, then an abundance index could be used to infer density. Using data

from six sites where tiger density could be estimated, a regression of RAI1 with

Ln(density) was non-significant (r2=0.05, p>0.05). Therefore, for this particular

dataset, the number of photos does not provide a reliable density index.

Population size

Population estimates based on mark-recapture data have been made for large cats in

reserves similar in size to Taman Negara (tigers in Indonesia: O’Brien et al. 2003;

tigers in Myanmar: Myanmar Forest Department 2003) and reserves that approximate

the size of all potential tiger habitat in Peninsular Malaysia (Maffei et al. 2003; Silver

et al. submitted). In each of these studies survey efforts were similar to or less than

those in this study, and the study of Kawanishi (2002). This suggests the possibility

for meaningful population estimates of tigers in Malaysian forests.

Recent estimates of tiger numbers in Peninsular Malaysia have apparently increased

over time: from 250 in the early 1980’s (Khan et al. 1983), then fluctuating between

500 (Topani 1990) and 600-650 (Khan 1987; Samsudin & Elagupillay 1996).

However, these estimates were partly based on track counts, a method which assumes

that individual tigers can be uniquely identified from tracks, and that tracks of every

tiger present in an area can be detected (Panwar 1979). In Southeast Asia there are no

long-term sites where workers are able to distinguish individual tigers by the shape of

their tracks. Also in tropical rainforests sign is not always 100% detectable due to

year-round rainfall so these assumptions are violated, and results of such surveys

unreliable (Karanth 1987, 1995, 2003).

In this study, as an alternative, photographs that identified tigers with 100%

confidence were used along with a mark-recapture approach that takes into account

the probability of detecting individual tigers. The results were combined to obtain

sample estimates of tiger numbers for each site. In Peninsular Malaysia, tiger density

varies across forested landscapes (Table 6). However, this variance can be estimated

by sub-sampling density at multiple sites. If the variance is assumed for blocks of

habitat, density can be extrapolated to estimate numbers in those blocks with some

level of confidence. Estimates of tiger numbers are given for individual reserves and

for the 16 TCUs described by Dinerstein et al. (1997)(Table 8). For the purpose of

28

the analysis, for TCUs that were not surveyed (i.e. those in the southern Peninsula

region), tiger density was assumed from adjacent sites or sites with similar

characteristics that were surveyed (see Table 6). We acknowledge that such

extrapolation may not yield precise population estimates, but consider this the best

alternative available at this time for estimating overall population numbers.

In estimating the proportion of habitat occupied by tigers, roads, forest edges, gaps

between forest patches, powerlines, human settlements and other infrastructure were

taken into account since they all pose barriers to dispersal and reproduction (Goosem

1997; Rabinowitz 1993; Woodroffe & Ginsberg 1998). In the absence of more

detailed information, it was assumed that tigers might use only 50% of available

habitat, so that populations might be 50% lower than maximum estimates. For

example, Kawanishi (2002) estimated a maximum range for tigers in Taman Negara

but actual numbers in the park may be less. Thus we arrived at a set of conservative

estimates of tiger populations.

The number of tigers estimated to be in individual reserves in this study ranged from

1-18 tigers. To estimate tiger numbers in TCUs, estimates of tiger density from sites

contained in the TCUs, or from sites near the TCUs were used to extrapolate tiger

numbers (Table 8). For Selama (TCU#130) the density for Bintang Hijau was used.

For the 13 Level III TCUs in the southern Peninsula region, and Endau (TCU#139),

the density for Lepar was assumed. Tigers were confirmed or possibly present in and

near these 14 southern TCU’s since 1987 (Rabinowitz 1999; DWNP unpublished

data), and for the purposes of this analysis tigers were assumed to be still present in

the TCUs. Tiger estimates for TCUs ranged from 3 in Endau to 278 in the Halabala-

Belum-Taman Negara complex (TCU#129). A conservative estimate of the national

tiger population was generated by adding the estimates for all TCUs, assuming 82%

of the estimate for TCU#129 (18% of which lies in Thailand). This resulted in an

estimate of the total tiger population of Peninsular Malaysia of 235 (range=184-305).

29

Table 8. Estimated numbers of tigers in reserves and TCUs in Peninsular Malaysia.

7. Taman Negara d 1.66 1.45-1.87 Reserve 4,343 68(52-84) 34(26-42)

8.Halabala-Belum-

T. Negara

2.02 1.66-2.39 129 a 27,469 555

(456-656)

278

(228-328)

* Area for these sites not defined; a Level I TCU, b Level II TCU, c Level III TCU; d Original data from Kawanishi 2002. Estimate in far right column is interpreted. e Minimum sampling area (222km2) was larger than the size of the reserve so used the latter to calculate density; f Assumes 82% of area of TCU#129 lies inside Peninsular Malaysia

Threats to tigers

Habitat loss and fragmentation are related threats that have direct and indirect

influences on tigers in Peninsular Malaysia. They occur with the creation of road and

power networks, conversion of forests for rubber, coconut and oil plantation,

expansion of urban areas, dams, irrigation, and other planned or unplanned landscape

development (Wikramanayake et al. 2002). Tigers are affected because they are

wide-ranging species that need relatively large (>3,000km 2) areas to sustain long-

term viable populations (Karanth and Nichols 2002). Tigers go rapidly extinct in

small habitat fragments due to conflict with humans at forest edges (Woodroffe and

Site ˆ D

ˆ D min - ˆ D max

Reserve

/ TCU#

Area

(km2)

Raw no.

tigers

(min-max)

50% use

(min-

max)

1. Temenggor 2.68 1.44-3.92 Reserve 1,336 36(19-52) 18(10-26)

2. Ulu Temiang 1.95 1.92-1.98 Reserve 154 e 3(3-3) 2

3. Gunong Tebu 0.60 0.45-0.75 Reserve 255 2(1-2) 1

4. Bintang Hijau 0.74 0-3.63 Reserve 1,167 9(0-42) 5(0-21)

5. Ayer Ngah 0.60 0.45-0.75 Reserve *

6. Lepar 0.58 0.43-0.72 Reserve *

9.Belum-T. Negara 1.90 1.53-2.28 129 a 22,524 f 428

(345-514)

214

(173-257)

10. Selama 0.74 0-3.63 130 b 1,684 12 (0-61) 6(0-30)

11. Endau 0.58 0.43-0.72 139 b 788 5(3-6) 3(2-3)

12. S. Peninsular

Level III TCUs

0.58 0.43-0.72 131-

144 c

4,163 24(18-30) 12(9-15)

P. Malaysia

(sum of 9-12)

235

(184-305)

30

Ginsberg 1998). In Peninsular Malaysia 80% of lowland forests have been lost, with

almost all protected areas less than 500km2 in size, many less than 10km2

(Wikramanayake et al. 2002). Tigers and their largest prey species are mostly absent

from forest fragments smaller than 100km2 (Laidlaw 2000). Tigers also have some

ecological requirements which render them vulnerable to certain types of habitat loss.

Specifically, they depend on access to rivers, so disturbance of or along rivers (e.g. by

roads, dams) can wipe out tigers from areas of otherwise suitable habitat. Logging

alters forest structure and succession, and degrades tiger habitat.

Habitat fragmentation increases accessibility to the remaining forested areas, which in

turn become degraded through a range of human activities. The increased access due

to logging allows for increased hunting (Robinson et al. 1999). Hunting in turn

affects tigers by removing their prey base which can lead to population declines

(Madhusudan and Karanth 2002; Ramakrishnan et al. 1999). Access paths created for

other purposes also facilitate increased hunting. At Temenggor, for example, well-

used trails are traversed by aloewood collectors who hunt illegally to subsist in the

forest. Some of these people were Thai and Lao, as evidenced by carvings in trees in

these languages. Information from former poachers suggests these people spend

several weeks to a month collecting aloewood (P. Klinklay pers. comm.) which is

prevalent in primary and secondary lowland and hill dipterocarp forests in Malaysia

(Barden et al. 2000). Evidence of hunting was found at several sites in this study

(Wan Shahruddin 1998). In general, while there exist a few examples where

populations of tropical wildlife species can be sustained when hunted, there exist

many more examples where hunting is unsustainable and leads to depletion or loss

(e.g case studies in (Robinson & Bennett 1999). The sustainability of hunting has not

been studied in Peninsular Malaysia, but the level of hunting of most species in

tropical forests is known to be low if it is to be sustainable (Robinson 2000).

The increasing frequency of reports of depredation and human injury from tigers in

Peninsular Malaysia (Elagupillay et al. 2001) can either be seen to reflect an increase

in tiger numbers or is evidence that the processes of forest loss and fragmentation, and

over-hunting of prey, are affecting tiger populations. Because our overall estimates

suggest a sharp decline in tiger numbers, we believe the most parsimonious

explanation is that increased conflict reflects increased habitat fragmentation and

31

forest loss. For example, between 1988 and 1997, there were 108 cases of reported

tiger incidents with humans in four out of six states, and 503 reported tiger-livestock

incidents in six states (Elagupillay et al. 2001). Tigers increasingly come into contact

with people as their habitats are encroached upon (Woodroffe & Ginsberg 1998), and

hunting inside forest reserves may deplete prey populations forcing tigers to take

livestock. The effect of tiger mortality through conflict with humans has not been

estimated for tiger populations in Peninsular Malaysia.

Commercial trade in wild tigers and tiger parts has occurred in Asia and across the

world for a very long time (Nowell 2000). Official records indicate it continues today

in Malaysia (Elagupillay et al. 2001), as it does in other countries (Bennett & Rao

2002). The trade is difficult to monitor because it is illegal and involves large

amounts of money, so traders use clandestine methods to disguise it. However,

monitoring is needed to assess impacts on wild populations, and enforcement

measures need to be stepped up to suppress the trade.

Aside from hunting, tiger reproduction and other behaviour may be disrupted and

prey populations displaced by harassment from humans. Eight of nine survey areas

showed evidence of human intrusions in this study (Table 7). Human traffic occurred

in up to a third of the study areas, and was locally intense in some places especially at

Ayer Ngah, Cameron Highlands and Jengai. Malaysian tigers apparently avoid

humans by altering their activity patterns (Fig. 6), as is evidenced in other parts of

their range (Griffiths and van Schaik 1993b).

Habitat loss combined with forest fragmentation due to landscape development

potentially could result in genetic erosion of large mammal populations, such as

tigers, if development creates barriers to dispersal of individuals between forest

patches.

Finally, perceptions that typify tigers as pests are a threat to their survival.

Education programs will be needed to change awareness of officials and the general

public to tigers.

32

Suggested priorities for tiger conservation

In order to plan for the tiger’s future in Peninsular Malaysia, each of the threats

described above need to be considered. Additionally, a set of targets need to be

identified that allow realistic progress to be made in reducing these threats at

appropriate scales of time and space (Ginsberg 2001). Tiger conservation in

Peninsular Malaysia could target the following;

• TCUs that encompass several landscapes;

• Landscapes that contain several populations of females and habitat

connections between the populations;

• Sites that contain at least several breeding females.

Priority Tiger Conservation Units - The Halabala-Belum-Taman Negara Level I

TCU (#129) (Dinerstein et al. 1997) is clearly of highest priority A secondary priority

would be Level I TCU#130 which is small enough (10,000km2) and

connectivity with important tiger habitats in southern Thailand. A second priority is

the Greater Taman Negara Landscape given it’s size (>10,000km2) and the status

of the core part of the landscape as a National Park. Both landscapes contain low-

density tiger populations with supporting prey base in several places.

Priority sites might include;

1. Main Range

a. Belum State Park-Temenggor Forest Reserve complex. The

original Temenggor site surveyed in this study has now been logged

(Abdul Kadir 1998; Wan Shahruddin 1998) but the area may still

support tigers. Currently the east-west Grik Highway divides the area

into two. If connectivity could be re-established by retrofitting the

road with multiple culverts to permit north-south tiger dispersal, and if

systematic wildlife enforcement patrolling can be done, this site would

33

become even more important.

b. Bintang Hijau might be a secondary target given the apparent low

level of human use of the forest.

2. Taman Negara Landscape

a. Taman Negara National Park. Merapoh in the western part of the

park supported the highest density of tigers and lowest human traffic of

three sites surveyed (Kawanishi 2002; p.67), and clearly is a priority

core area within the site. Tigers are threatened by aloewood collectors

who poach prey animals in the park (Barden et al. 2000; Wan

Shahruddin 1998)

3. Krau Wildlife Reserve. A secondary priority site given the problem of

habitat connectivity with the Taman Negara Landscape, and a dwindling tiger

population (DWNP/DANCED 2002). The area is small (531km2) but together

with adjacent forest tracts contains potential tiger habitat. Tiger populations

inside Krau and in adjacent areas could be restored with appropriate

management of prey populations, especially controlling hunting, and habitat

management.

A possible framework which identifies these areas and suggests appropriate short

term and long-term goals for tiger conservation is shown in Table 9. This could form

the basis of future discussions on tiger conservation priorities in Peninsular Malaysia.

34

Table 9. Targets for tiger conservation with various time and spatial scales (adapted

from Ginsberg, 2001).

TARGETS

SHORT TERM (2 – 5 YEARS)

LONG TERM (10 – 20 YEARS)

SITE (An area containing several breeding females) e.g. Belum State Park-Temenggor Forest Reserve complex, Taman Negara NP, Bintang Hijau

• Maintain occupancy of tiger habitat

• Define critical areas within sites

• Stabilize present tiger populations

• Prevent loss of tigers

• Maintain potentially breeding populations of tigers at maximum density

• Maintain expanding population (at r>1)

• Strictly protect core areas

LANDSCAPE (A larger area containing several populations of breeding females) e.g. Main Range, Greater Taman Negara Landscape

• Maintain potential for dispersal between sites

• Maintain ecologically functioning viable tiger populations

• No human intervention required to achieve stable/growing populations

• Recolonization of empty habitat

TIGER CONSERVATION UNIT (An area containing several landscapes) e.g. Halabala-Belum-Taman Negara TCU#129, Selam TCU#130

• Maintain integrity of intact habitat

• Maintain sufficient prey base

• Maintain multiple landscapes including transboundary landscapes in each TCU

• Coordinate establishing protected areas across boundaries

• Promote tiger friendly conservation in each country in TCU

• Re-establish connections between sites and landscapes to ensure genetic exchange

• Maintain heterogeneity of ecoregion

35

36

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