A General Guide to

Camera-trapping
Large Mammals in
Tropical Rainforests,
with Particular
Reference to Tigers

Shariff Wan Mohamad

& Mark Rayan Darmaraj

© WWF-Malaysia / Mark Rayan Darmaraj

A General Guide to Camera-trapping Large Mammals in Tropical Rainforests, with Particular
Reference to Tigers
Shariff Wan Mohamad & Mark Rayan Darmaraj
May 2009

Published May 2009 by WWF – World Wide Fund for Nature (formerly World Wildlife Fund),
Petaling Jaya, Malaysia. Any reproduction of this publication must mention the title and credit the
above-mentioned publisher as the copyright owner. © Text 2009 WWF. All rights reserved.
 CONTENTS

INTRODUCTION 1
A BRIEF OVERVIEW ON WILDLIFE MONITORING METHODS 3
CAMERA-TRAPPING 6
1. Introduction 7
2. Survey design considerations 7
2.1 Sampling duration 7
2.2 Camera-trap spacing 8
2.3 Number of camera-trap locations 9
3. Conducting the study 9
3.1 Choosing a camera-trap 9
3.2 Reconnaissance surveys 10
3.3 Choosing a suitable camera-trap location 11
3.4 Camera-trap settings 12
3.5 Camera-trap deployment 13
3.6 Maintenance/monitoring of camera-traps in the field 13
3.7 Community involvement 14
4. Analysis of results 14
4.1 Identification of individual tigers 14
4.2 Calculating population size 16
4.3 Estimating the effective area sampled 17
4.4 Calculating density 18
5. Camera-trapping other species 18
REFERENCES 20
Appendix I: Tracks of large mammals found in Peninsular Malaysia 26
Appendix II: Example of a basic camera-trap data form 33
 INTRODUCTION
Protected areas are currently inadequate to conserve the biological diversity
found within tropical forests due to their limited size, number, distribution,
composition and protection status (Fimbel et al., 2001). Production forests
represent an opportunity to complement
the existence of protected areas in
providing critical habitat for wildlife;
hence it is important to ensure that
production forests are managed wisely
through wildlife monitoring and better
management practices. Worldwide,
numerous studies have been conducted
on the impacts of logging on wildlife in
tropical forests (Grieser Johns, 2001).
© WWF-Malaysia / Shariff Mohamad
These studies consistently report that
many animals show a decline in observed abundance, some are totally
eliminated and others are known to increase in abundance (Fimbel et al., 2001).
Two main factors influencing the variability of results obtained from studies done
on the impacts of logging are different assessment techniques and different
species variability in response to logging. Thus, it is not easy to make any
generalisation on the impacts of forestry practices on wildlife mainly because of
the variations in a spatial and temporal context and the seemingly poor
predictability of the long-term responses of each wildlife species.

Acknowledging the need for a simple, cost-effective and easily implemented

method of large mammal assessment for the forestry sector to conduct during
pre-felling and post-felling surveys in production forests, at this point of time, we
could suggest using tracks to identify the presence of large terrestrial mammal
species and to quantify the amount of effort used during these surveys to derive
a Track Encounter Rate (TER) index. Such surveys have previously been
conducted in Peninsular Malaysia and with a considerable amount of data
collected over time, could possibly be used to gauge relative population trends
(Grieser Johns, 1997). However, due to the inherent problems of TER indices
relating to detection probability which could possibly effect interpretation of
results from such surveys, we would recommend camera-trapping to estimate
the absolute abundance of individually-identifiable large mammals (such as
tigers) for areas that encompass forest compartments equivalent to at least 100
km2.

1
WWF-Malaysia is currently exploring the use of a combination of sign surveys
and camera-trapping to estimate occupancy of tigers and their prey in several
Permanent Reserved Forests (PRFs) designated for logging. Using occupancy
as a state variable to monitor changes in wildlife populations is a relatively new
concept. Nevertheless, the fact that
these methods are being explored in
Malaysia lends credence to the
possibility of a paradigm shift from just
using ad-hoc based surveys, towards
incorporating a robust methodological
framework to gain information on wildlife
populations, especially with regards to a
sustainable forest management agenda.
Thus, it is anticipated that this © WWF-Malaysia / Mark Rayan Darmaraj
pioneering study will be able to provide
recommendations for better management practices as well as improved and
validated methodologies in relation to monitoring large mammals, particularly
tigers and their prey in PRFs. In this document, we provide a general guide on
the use of camera-traps to estimate the abundance of tigers within a particular
study area. A more comprehensive guideline is expected to be made available
after the aforementioned study has been completed.

2
 A BRIEF OVERVIEW ON
WILDLIFE MONITORING METHODS

3
Monitoring animal populations can be described as the estimation of abundance
(relative or absolute) to draw inferences about the change in abundance over
space (spatial) or time (temporal). This can be used to assess animal distribution
patterns as well as the relationship between animal abundance and factors such
as habitat or management effectiveness (Nichols & Karanth, 2002).

Estimations of animal populations are almost always based on count statistics.

Count statistics are generally conducted via direct counts of individual animals, or
based on animal sign such as tracks or dung (Nichols & Karanth, 2002). Taking
C as the count statistic (number of animals counted) and p as the sampling
fraction (proportion of area sampled), the actual number of animals (N) can be
estimated by the following equation (Nichols, 1992):

N = C/ p

However, there are two main problems

with count statistics - observability and
spatial sampling. Observability in this
context refers to the inability to detect all
animals or animal signs in the study
area, whereas spatial sampling is
conducted when it is not feasible to
survey an entire area. In this case, a
representative subset of the area is
surveyed instead. The results of this © WWF-Malaysia / Oriane Ansel
can then be used to draw inferences
about the entire area (Nichols & Karanth, 2002; Karanth et al., 2003).

Comparisons of count statistics over spatial or temporal units, however, need to

take into consideration the detection probability, which is the probability that an
animal is detected during the survey period (Karanth et al., 2002). Monitoring
abundance of large mammals in tropical rainforests remains a challenge because
detection probability is severely reduced due to physical constraints such as
dense vegetation, undulating terrain and climatic conditions.

In many regions, distance sampling (Buckland et al., 1993) is used to estimate

densities of terrestrial mammals through line transects. This method is commonly
used for animals which are fairly easy to sight, such as large or medium-sized
herbivores in relatively open habitats. As such it is not well-suited for sampling
animals which occur at low densities or in dense habitats, as is often the case in
tropical countries. In Peninsular Malaysia, line transect sampling has not been
met with much success (Kawanishi, 2002), and an unreasonably large sampling
effort would be required to obtain sufficient samples in order to produce reliable
density estimates.

4
 Surveys using indirect signs such as tracks and dung are also commonly used,
as in most cases secondary signs are
easier to come across than the animal
itself. However, estimating relative or
true abundance from secondary signs
can be complicated due to several
additional assumptions, although
recently-developed techniques have
made dung surveys one of the preferred
methods for monitoring elephant
Wong
Wong © WWF-Malaysia / Christopher Wong
populations (Hedges & Lawson, 2006).
Abundance estimations using dung
surveys have successfully been obtained for sika deer in Scotland (Marques et
al., 2001), and for Asian elephants in Sumatra (Hedges et al., 2005), among
others. In southern Peninsular Malaysia, The Wildlife Conservation Society
(WCS) has conducted occupancy surveys in the Endau-Rompin Forest Complex
using secondary signs (tracks and dung) to obtain occupancy estimates and
possibly abundance estimates of tiger prey. Occupancy surveys are based on
the underlying principle that changes in the percentage of area occupied by a
species could be related to changes in population size (Linkie et al., 2007).
Recently, a new model using an occupancy approach to estimate abundance
(Royle & Nichols, 2003) has also been proposed and is currently being
investigated by numerous researchers especially because it does not require
individual recognition of animals. WWF-Malaysia is currently exploring this
method with the use of camera-traps and sign surveys in selectively logged
forests.

Camera-trapping has been favored for abundance estimation of individually-

identifiable mammals, which allow population densities to be obtained through a
mark-recapture framework (Karanth 1995; Karanth & Nichols 1998; Henschel &
Ray, 2003; Silver et al., 2004). It is important to note that at this moment, if the
objective of a sampling survey is to monitor changes of absolute abundance of
large mammals through camera-trapping, then this method is only applicable to
animals with unique natural markings. This holds true until and unless camera-
trapping data which incorporates the use of the Royle and Nichols (2003) model
or any other methods is proven to provide a reliable absolute abundance
estimate for animals without unique natural markings. In a nutshell, the question
of which method is suitable and practical to be implemented on the ground to
assess and monitor large mammals will ultimately depend on the research
question being asked, and the amount of resources available.

5
CAMERA-TRAPPING

6
1. Introduction
In the past, large-scale censuses of tracks have been conducted in order to
obtain total counts of tigers throughout India. However, such “census” techniques
have been shown to be flawed due to the assumption that all animals have been
counted, and is thus an unreliable method for obtaining population estimates
(Karanth, 1999; Karanth et al., 2003). In the early 1990s, Ullas Karanth
pioneered the use of remote cameras in obtaining estimates of tiger densities
using the capture-recapture method (Karanth, 1995) based on the fact that
individual tigers can be identified by their stripes. Since then the methodologies
have been further refined, and camera-trapping has now become an effective
and widely-used tool for obtaining density estimates of individually-identifiable
animals.

Camera-trapping for the purpose of estimating abundance of individually-

identifiable animals basically involves setting remote cameras within an area of
interest, configured in such a way that the target species does not have a zero-
probability of being detected. Based on the number of captures (photographs)
and recaptures of each individual, the abundance is estimated using statistical
software such as CAPTURE. The density estimate is then obtained by dividing
the population estimate by the effective area sampled.

2. Survey design considerations

Successful surveys require careful planning and preparation. In particular, the
purpose and objectives of your survey needs to be thought through as this will
determine the information that you need to obtain, and thus the methodology that
is most appropriate. These goals should be achievable; taking into account the
skills and resources available. In addition, before you start surveying you need to
know how the data will be analyzed. This is vital in order to develop an
appropriate sampling design. If data on abundance is required, then the survey
design has to take into consideration several key factors to obtain good density
estimates. Here, we will elaborate on these factors using tigers as a case study.

2.1 Sampling duration

One assumption of the mark-recapture framework is that the population is
closed during the sampling period (no births, deaths, immigration or
emigration occur). Thus, camera-trapping should be limited to a relatively
short period. According to Nichols & Karanth (2002a) sampling is ideally
limited to two weeks. Several studies in India have utilised a 15-day sampling
period (Karanth et al., 2004; Chauhan et al., 2005), but in regions with lower
tiger densities sampling can take place over a much longer period in order to
obtain sufficient data. Although Nichols & Karanth (2002a) proposed that
sampling be conducted over a maximum of 2-3 months, several other studies
conducted in South-east Asia have exceeded this whilst not violating
statistical closure as calculated by program CAPTURE (Kawanishi &
Sunquist, 2004; Simcharoen et al., 2007; Rayan & Shariff, 2009).

7
 Nevertheless, CAPTURE is unlikely to reject closure when dealing with small
samples (Otis et al., 1978). Taking all these factors into account, a
biologically-realistic sampling time frame should be limited to between three
and six months.

2.2 Camera-trap spacing

Typically, camera-traps will either have certain spacing between locations or
be set via an imaginary grid system (Figure 1), in order to cover the whole
study area without leaving any significant gaps – this is because the target
species needs to have a non-zero probability of being detected (Silver, 2002).
This means that the gaps between camera-trap locations should not be big
enough for a tiger to travel within its home range and not have any probability
of being detected. To estimate a minimum gap size, the smallest known home
range of tigers from that particular region and habitat should be obtained. For
example, in Peninsular Malaysia the smallest estimated (minimum) home
range of an adult female obtained was 90 km2 from a disturbed secondary
forest (Darmaraj, 2007). If the home range is assumed to be circular, the
radius would work out to approximately 5.4 km, which should be used as the
maximum distance between camera-traps locations. However, it would be
better to have about 2-4 camera-trap locations encompassed within the home
range of each individual (White et al., 1982; Karanth & Nichols, 2002a) to
increase the probability that the tiger will be photographed during each
sampling occasion. On the other hand, our goal is to catch as many individual
tigers as possible, with a limited number of camera-traps. Therefore, a
compromise has to be found between these two factors (Karanth et al.,
2002a). Keeping this in mind, an inter-trap distance of approximately 2-4 km
should be appropriate for studies conducted within tropical rainforests.

20 km

10 km

2 km

2 km
Figure 1: A hypothetical 200 km study site (camera-traps represented by dots)

8
 2.3 Number of camera-trap locations
Generally, the greater the number of individuals captured and recaptured, the
more robust your density estimate will be. Since a larger study area would
theoretically cover the home range of more individuals, the larger the study
area the better! Realistically though, this will likely be limited by the number of
camera-traps available, manpower and also logistics. It would be good to aim
for an area of at least 200 km2 (although this is not always feasible), as we
would expect an area of this size to hold about 3-6 tigers (O’Brien et al., 2003;
Kawanishi & Sunquist, 2004; Rayan & Shariff, 2009). Assuming that an
imaginary 2 km x 2 km grid system is used whereby one camera-trap location
is contained within each 4 km2 grid square, then a total of 50 camera-trap
locations would be needed for an area covering 200 km2 (Figure 1). Since
each location would have one camera-trap on each side of the trail, this works
out to a total of 100 camera-traps (not including spares!). This can work out to
a hefty investment, so an area of around 100-150 km2 will probably be more
financially practical. If the only limiting factor is the number of camera-traps
available (rather than manpower or logistics), then another option would be to
divide a large area into two or more smaller sections and then sampling each
one in succession by rotating the camera-traps between each section. Further
details on this are provided by Nichols & Karanth (2002a).

3. Conducting the study

3.1 Choosing a camera-trap
There are two types of camera-traps which are normally used - active and
passive. An active camera-trap consists of the camera itself, a transmitter and
a receiver which is placed on the opposite side of the trail. The transmitter
emits a narrow infra-red beam, which triggers the camera to take a photo
each time it is broken. Passive camera-traps however, detect heat-in-motion
and take a photo whenever an animal passes within the range of its sensor.
Although both types of camera-traps have their respective pros and cons,
passive camera-traps seem to be favoured by many researchers, mainly due
to their ease of transport and setup.

Previously only film camera-traps were available; more recently digital

camera-traps have also been developed. These digital units offer a big
advantage – the ability to store far more photos than the conventional 36
exposures that was limited by the usage of film. For example, a 4 megapixel
camera-trap with a 1 GB memory card has the potential to store
approximately 500 photos. This can allow for less frequent visits (to change
film) and also enable camera-traps to be set at sites with a higher level of
activity than typical forest trails, such as saltlicks and active logging roads.
Another advantage is that the photos can usually be viewed while checking
the camera-trap, enabling the user to judge on the spot whether a suitable
location was chosen based on the photos taken. If the location is deemed to

9
be unsuitable, the camera-trap can be moved somewhere else immediately
without having to wait for the film to be processed. Digital camera-traps do
have a couple of drawbacks however – the costs can be higher and the delay
time (lag time between detection of the animal and the photo being taken) is
typically slower than analog camera-traps. This is not true in all cases
however; these details will vary according to the specifications of each model.

Even though the initial cost of digital camera-traps may be high, opting for this
does eliminate film processing costs. Having the photos in digital format also
makes them far more accessible and enables them to be organized in a more
efficient manner. As many camera-trapping studies utilize some sort of
database for organizing camera-trap data, having digital photos does absolve
you from having to scan hundreds or even thousands of film photographs!

Deciding on which camera-trap model to purchase can be a risky venture,

simply because of the large cost involved in purchasing numerous camera-
traps. Nobody wants to make a large investment only to find out later that it is
unsuitable for their study for some unforeseen reason. It helps to compare the
specifications of what is on the
market and narrow the choices down
based on your specific needs (and
costs!). Also try and find out what
other researchers doing similar
studies are using, and if possible get
their feedback regarding their choice
of camera-traps. Try and explore
your possibilities from various
© WWF-Malaysia / Shariff Mohamad
angles; even something simple such
as reading user reviews of camera-
traps on a website can provide invaluable information. Some good online
resources for camera-trap reviews include Trailcampro
(http://www.trailcampro.com), Cabela’s (http://www.cabelas.com) and
Amazon (http://www.amazon.com). After you have decided on a suitable
model, it is advisable to purchase one or two units initially to test out in the
field. If you are satisfied with its features and performance, you can then
proceed with ordering more camera-traps.

If funds permit, it is recommended to purchase at least 20% more camera-

traps than you are planning to deploy in the field. This is to cover the loss of
camera-traps which will inevitably be stolen, vandalized or experience
malfunctions. For example, if your study requires 50 camera-traps then a
minimum of 60 should be purchased, although having even more than that
would be much preferred.

10
3.2 Reconnaissance surveys
The general choice of study area is often dependent on the research
questions that need to be addressed. It has to be noted that the proposed
study area should be representative of the larger area which is intended to be
assessed. This can be in terms of disturbance levels, elevation classes, land
use or other features. Potential study sites can be identified from a
combination of topography maps, satellite imagery and land use in the area;
but it is still vital to conduct reconnaissance surveys before embarking upon a
full-blown study. A couple of initial surveys can be conducted to gauge the
suitability of the area, and subsequent ones to familiarize oneself with the
study area, field conditions and to identify potential difficulties in carrying out
the study. GPS tracking can be done during this period to map out road
networks and other trails, which is useful in logistical planning.

It is advisable to train field assistants during the reconnaissance survey

period, if they are not familiar with the scope of camera-trapping studies. This
period also provides time for the
researcher to field test the camera-
traps before the sampling actually
begins. Single camera-traps can be
deployed at promising sites, to
ascertain travel routes of tigers and
to identify the placement of
permanent camera-trap locations
during the sampling period. Also,
mark any tiger sign which you come © WWF-Malaysia / Roshan Guharajand
across using a handheld GPS. A
record of tiger presence obtained through a combination of GPS waypoints
and preliminary camera-trapping will give you a good idea of where to set
your camera-traps during the actual sampling period.

3.3 Choosing a suitable camera-trap location

Because our goal is to obtain as many tiger captures/recaptures as possible,
it is important to choose optimal sites in order to maximize capture
probabilities (Karanth et al., 2002a). Tigers and other medium- to large-sized
terrestrial mammals are known to regularly travel along wildlife trails, man-
made paths (e.g. logging roads), ridges, and also along riverbeds/streams.
However, animals might avoid these areas if there is a high level of
disturbance such as logging or other human activities. As resident tigers often
use the same trails repeatedly, an ideal camera-trap location would be along
a trail which has recent signs of tiger usage. These locations can be readily
identified by the presence of tiger pugmarks, although other signs such as
scratch marks and scats are also occasionally found. If no tiger sign is found
in a particular area, then the camera-trap should preferably be set where
there is prey activity (typically ungulates) or along suitable trails. It is possible
that tigers do use these routes, but the pugmarks may not be visible due to

11
 the ground substrate or they may have been washed away by heavy rain. A
guide to large mammal tracks from Peninsular Malaysia along with
photographic examples is included in Appendix I for reference purposes.
Tracks of other mammal species from the region can be obtained from
sources such as van Strien (1983) and Kanjanavanit (1997).

It is recommended to use a reasonably wide trail to reduce the chances of

obtaining a partial photo capture or even missing the animal entirely, due to
the tiger passing too close to the
camera. If there are only narrow
trails, the camera-trap can be set on
a tree slightly off the path to enable
a wider coverage to be obtained.
However, make sure the camera’s
field of view is not blocked by other
trees and clear any vegetation which
might obscure the photo. When
© WWF-Malaysia / Shariff Mohamad
appropriate, obstructions such as
logs or fallen branches can be
placed strategically to discourage animals from passing too close to the
camera.

3.4 Camera-trap settings

Settings can typically be changed on two components of the camera-trap: the
camera itself and the control board. The settings to be used will vary
according to what camera-trap model is used and the objectives of the study,
which will ultimately determine how it will be set in the field.

Cameras should be set to run for 24 hours a day, as tigers aren’t active only
during the night (or day!). It is recommended to use a high ISO (sensitivity to
light) of 400, to reduce the chance of obtaining blurred pictures. This option
can either be selected via the film used (for analog cameras) or through the
camera settings (for digital cameras). A delay time of several minutes
between photos has been commonly used in previous camera-trapping
studies, to prevent wastage of film by animals lingering in front of the camera-
trap. For digital cameras, the minimum delay time can be selected as the
storage capacity is much higher. However, this also depends on the capacity
n 
of the memory card which is used and the length of time you plan to leave it in
d  the field. Also make sure that the correct date and time are set on the
camera; this is vital as it will ultimately be used in the analysis of camera-trap
data.

3.5 Camera-trap deployment

For the purpose of monitoring tigers, two camera-traps are typically set on
either side of a trail, to enable unambiguous identification of individual tigers
(Karanth et al., 2002a) - more of which is covered in section 4.1. Each

12
camera-trap should be assigned a unique number or code (such as C1, C2,
and so on); this can be marked on the external casing using a permanent
marker. It is also essential to take down the coordinates of each camera-trap
location using a GPS unit.

Whenever camera-traps are used as a research tool, some sort of datasheet

should be used to record the information required for analysis in the later
stages. This datasheet should preferably be simple, whilst retaining all the
essential information required for the study. An example of a basic camera-
trapping datasheet can be found in Appendix II; additional information can be
included according to the specific objectives of the study. This form has to be
filled up each time a camera-trap is set and retrieved.

Before setting the camera-trap, it is advisable to insert a packet of silica gel

inside the camera-trap housing to absorb any
moisture. This is especially relevant in tropical
countries, where high levels of rain and humidity
often result in moisture seeping into the camera-
trap unit. After turning on the camera-trap, fasten
it to a suitable tree using a cable-lock, about 40-
50 cm above the trail (around knee height) which
is a suitable height to photograph tigers (Karanth
et al., 2002a). Then conduct a walk-test by
crawling at different distances from the camera-
trap, making sure it detects you throughout the
width of the trail. The angle of the camera-trap
might have to be adjusted accordingly; this can
be done by wedging twigs or small pieces of
wood strategically between the camera-trap and © WWF-Malaysia / Oriane Ansel
tree. After you are satisfied with the placement,
then activate the camera-trap to start taking photos. Make sure to take a test
shot before leaving the area!

3.6 Maintenance/monitoring of camera-traps in the field

Even though the battery life and storage capacity of modern digital camera-
traps are greater than that of analog cameras, it is still recommended to
check each camera-trap on a monthly basis. This is done mainly to minimize
the loss of data due to possible theft, damage or malfunction of camera-traps.
If the camera-trap detects no tigers after a couple of months or if the location
is found to have little wildlife movement, it is recommended to move the
camera-traps to a more suitable location. During each monthly visit, retrieve
and label the memory card / film roll and store it in a safe place. Also replace
the old batteries with new ones. Make sure the camera-trap takes a retrieval
test shot upon retrieval; if you suspect it is faulty then replace it with a new
one (assuming you have spares!).

13
 3.7 Community involvement
If there are settlements situated in or around the study area (indigenous
villages, logging camps, etc.), it is
recommended to have an informal
chat with the people in charge to
inform them of your plans to conduct
a camera-trapping study in the area.
Give a brief introduction of what you
plan to do and ask pertinent
questions which can help in planning
your study. Indigenous people can
© WWF-Malaysia / Shariff Mohamad
be an invaluable source of
information, and can help identify
wildlife routes, saltlicks and other areas of interest.

The help of field assistants are typically needed throughout the course of the
study. It is recommended to hire people familiar with the study area, such as
indigenous people or locals living within the vicinity. Finding reliable people
can sometimes be rather tricky, so try and gauge the candidate’s experience
and interest in conducting this kind of work to the best of your ability. Seeking
the support of the local community can also help to reduce incidences of
camera-trap theft or vandalism.

4. Analysis of results
4.1 Identification of individual tigers
Like a human fingerprint, the stripe pattern of each individual tiger is unique.
Individuals can be readily identified as long as a clear photo of the flank is
obtained. Do keep in mind that the left and right flank of each individual tiger
differs – this is why two opposing camera-traps are typically set in each set
location. A relatively efficient way of identifying individuals would be to focus
on just two or three distinctive stripe patterns, instead of looking at the entire
flank (Figure 2). Keep aside a pair of complete photos (both left and right
flank) of each individual as a main reference, for easy comparison with any
new photos that are obtained from the study. In most cases even partial
photos can be used to identify individuals, as long as there is a complete
reference photo available.

14
 © WWF-Malaysia / Mark Rayan Darmaraj

© WWF-Malaysia / Mark Rayan Darmaraj

Figure 2: Identifying individual tigers through comparison of the same flank; confirmation
of the same individual is obtained through the matching stripe pattern

15
4.2 Calculating population size
For estimating the population size of tigers using a mark-recapture
framework, the program CAPTURE (White et al., 1982; Rexstad & Burnham,
1991) is commonly used. The software and manual can downloaded for free
at http://www.mbr-pwrc.usgs.gov/software.html. CAPTURE generates
estimates of abundance using several different models, which vary in their
assumed sources of variation in capture probability such as individual
heterogeneity (Mh), behavioural response (Mb), time (Mt), and a combination
of these factors (Mbh, Mth, Mtb and Mtbh). A constant parameter model (M0) is
also calculated (and is often ranked as the highest model), but this model
does not take into account the various underlying assumptions that affect
capture probability (Karanth et al., 2006). Nichols & Karanth (2002a) suggest
Mh as the most suitable model on the basis that heterogeneous capture
probabilities exist among individual tigers due to their social organization and
unequal access to camera-traps.

To prepare for data input into CAPTURE, a capture history of each individual
tiger has to be compiled in an ‘X matrix’ format. This is constructed in the form
of a table with each row representing an individual tiger, and each column
corresponding to one sampling occasion (refer to Table 1 below). An
occasion will typically range from one day to a couple of weeks, depending on
the duration of the entire study. The number of sampling occasions can be set
by the researcher after the study, with more occasions usually leading to an
increased precision (Henschel & Ray, 2003). The matrix is then filled with 1’s
and 0’s, according to which occasion each tiger was detected (1 = detected, 0
= not detected). After inputting this data into CAPTURE, it will then calculate
the population estimate (N), associated confidence intervals and capture
probabilities, in addition to the mean and standard error. Do note that
CAPTURE does not actually calculate density; it only provides estimates of
abundance.

Table 1: Example of a capture history table

Occasion
Individual
1 2 3 4 5 6 7 8
Male 1 1 0 1 1 0 1 0 1
Male 2 0 1 1 0 1 0 1 0
Female 1 1 0 0 0 0 1 0 0
Female 2 0 0 0 0 0 1 0 1
Female 3 0 1 0 1 1 0 1 0

X matrix format

16
4.3 Estimating the effective area sampled
Because not all animals ‘captured’ have their entire home range within the
intensive sampled area (also known as the edge effect), using the actual area
which was sampled (A) will lead to severe overestimation of density estimates
(Otis et al., 1978). To account for this, a buffer (W) has to be added to the
polygon connecting the outermost camera-trap locations. An example of this
taken from a camera-trapping study in Gunung Basor, Kelantan, is shown in
Figure 3. Several methods can be used to calculate the buffer width. The
‘half’ mean maximum distance moved (MMDM) method (Wilson & Anderson,
1985) is currently one of the most widely used for estimating the density of
individually-recognizable felids (Karanth & Nichols, 1998; Trolle & Kery, 2003;
Silver et al., 2004; Chauhan et al., 2005a), although other researchers have
also used the ‘full’ MMDM (Soisalo & Cavalcanti, 2006) and AMDM (absolute
maximum distance moved; Kawanishi & Sunquist, 2004) methods in their
studies. Using the ‘half’ MMDM method, the buffer width is calculated as half
of the mean maximum distance between captures of each tiger photographed
at more than one location, averaged for all individuals (Nichols & Karanth,
2002a). The intensive sampled area plus the added buffer is then taken as
the effective sampled area, A(W).

Figure 3: Example of a camera-trapping study area showing a polygon connecting

the outermost camera-trap locations, with an added buffer (Darmaraj, 2007)

17
 4.4 Calculating density
Assuming we have already calculated the population size (N) and effective
sampled area, A(W), we can then obtain the estimated population density (D)
with the following equation:

D = N / A(W)

The density estimate derived is only applicable for the area sampled; it is not
advisable to extrapolate this value to a larger scale unless it possesses
similar attributes to the area sampled such as habitat type, topographic
features, prey abundance, and disturbance levels. However as you might
expect, this is rarely the case!

5. Camera-trapping other species

Although most large scale camera-trapping studies have been conducted on
tigers, density estimates for other individually-identifiable species have also been
obtained using this method, such as jaguars (Silver et al., 2004; Soisalo &
Cavalcanti, 2006), leopards (Chauhan et al., 2005a) and ocelots (Trolle & Kery,
2003; Maffei et al., 2005). Recently camera-trapping has also been used to
obtain density estimates of animals which lack unique coat patterns, but are able
to be distinguished by other identifiable features and markings on the body (Noss
et al., 2003; Trolle et al., 2007; Trolle et al., 2008). Aside from obtaining
abundance or density estimates, camera-traps can also be used to obtain
biological data such as diversity, relative abundances, occupancy, activity
patterns, and also presence of species (Kawanishi et al., 1999; Srbek-Araujo &
Chiarello, 2005; De Luca & Rovero, 2006; Grassman et al., 2006; Linkie et al.,
2007).

As camera-traps are capable of recording the date and time of each

photographic capture, the activity patterns of photo-captured animals can
subsequently be determined. A simple activity pattern graph can be constructed
for any given species by plotting the percentage of captures obtained during
each hour, over a 24-hour period (Figure 4). This information is useful to
ascertain the activity class of animals which haven’t been well-studied due to
their elusiveness or occurrence at low densities; scenarios which are especially
prevalent in dense tropical forests (van Schaik & Griffiths, 1996; Grassman et al.,
2006). This is not only important from a natural history viewpoint, but also in
terms of assisting research and management efforts (Gomez et al., 2005).

18
 18.0

Activity level (%)

16.0
14.0
12.0
10.0
8.0
6.0
4.0
2.0
0.0
0000 - 0100

0200 - 0300

0400 - 0500

0600 - 0700

0800 - 0900

1000 - 1100

1200 - 1300

1400 - 1500

1600 - 1700

1800 - 1900

2000 - 2100

2200 - 2300
Tim e in hours (hrs)

Barking deer Wild boar

Figure 4: Activity patterns of two ungulate species: barking deer and wild boar
(Darmaraj, 2007)

If the objective of the monitoring study is to monitor changes in abundance rather

than changes in absolute numbers, then relative abundance indices can be used.
In the case of camera-trap surveys, the sampling effort is measured in terms of
the number of trap nights, which refers to a 24-hour period where the camera-
trap was functional. If more than one camera-trap is used then the trap nights are
accumulated to derive the total sampling effort. A simple quantitative index of
relative abundance can thus be derived, consisting of the number of camera-trap
photos (detections) of a given species per unit sampling effort. This index can be
summarized by the following equation:

RAI = Σ detections x 100 / Σ trap-nights

This method can be used to monitor abundance changes on a temporal and

spatial scale, as long as the data is collected and treated in a similar manner
(Kawanishi et al., 1999). This relative abundance index is used in accordance
with Carbone et al. (2001), which substantiates the correlation between
photographic rate and density. However, there are statistical and ecological
arguments against the use of such indices as good predictors of absolute
abundance in an area (Jenelle et al., 2002; Nichols & Karanth, 2002b) so these
values should be treated with caution.

Camera-trapping can also be used to obtain more general information such as

the biodiversity of medium- to large-sized mammals in a given area, which can
be useful for inventory studies. However, the number of species detected will
generally depend on the area of coverage and the length of time camera-trapping
is conducted; a large sampling effort is thus required in order to survey an area
thoroughly. In addition, the presence of rare and elusive species can be detected
via camera-trapping, as demonstrated by the possible new records of fishing cat
from Peninsular Malaysia (Kawanishi & Sunquist, 2003). This could prove
valuable in confirming the presence of other scarce or thinly-distributed species
such as the Sumatran rhino, otter civet or flat-headed cat.

19
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25
 APPENDIX I: TRACKS OF LARGE MAMMALS FOUND IN
PENINSULAR MALAYSIA* (ADAPTED FROM VAN STRIEN, 1983)

30-50 cm

 
 
 

© WWF-Malaysia / Christopher Wong

ASIAN ELEPHANT
Elephas maximus

> 6 cm

© WWF-Malaysia / Ahmad Zafir Abdul Wahab

> 17 cm
SUMATRAN RHINOCEROS
Dicerorhinus sumatrensis

26
 < 17 cm

 
 

© WWF-Malaysia / Shariff Mohamad

MALAYAN TAPIR
Tapirus indicus

> 10 cm

© WWF-Malaysia / Lau Ching Fong

GAUR
Bos gaurus

27
 6-9 cm

 
 
 
 

© WWF-Malaysia / Lau Ching Fong

SAMBAR DEER
Rusa unicolor

4-6 cm

© WWF-Malaysia / Christopher Wong

SOUTHERN SEROW
Capricornis sumatraensis

28
< 7 cm

© WWF-Malaysia / Christopher Wong

WILD PIG
Sus scrofa

< 5 cm

© WWF-Malaysia / Christopher Wong

BARKING DEER
Muntiacus muntjak

29
> 9 cm

© WWF-Malaysia / Mark Rayan Darmaraj

> 7 cm

TIGER
Panthera tigris

7-9 cm

© WWF-Malaysia / Christopher Wong

5-7 cm

LEOPARD
Panthera pardus

30
6-8 cm

© WWF-Malaysia / Azlan Mohamed

4-5 cm

CLOUDED LEOPARD
Neofelis nebulosa

~ 5-6 cm

© WWF-Malaysia / Christopher Wong

WILD DOG
Cuon alpinus

31
 ~ 7-8 cm

© WWF-Malaysia / Lau Ching Fong

MALAYAN SUN BEAR

Helarctos malayanus

* Notes:

¾ Here, large mammals are defined as terrestrial mammals with a body weight exceeding 20 kg
(Morrison et al., 2007). Bearded pig (Sus barbatus) is excluded from this list due to its patchy
distribution and range limited mostly to the southern region of Peninsular Malaysia.

¾ These track measurements are meant only as a rough guide; actual measurements taken in
the field may differ slightly from the ranges provided due to differences among individual
animals as well as substrate conditions.

32
 APPENDIX II: EXAMPLE OF A BASIC CAMERA-TRAP DATA FORM

Location ID: Study area:

Elevation: Site description:

Longitude:

Latitude:

Date deployed:

Time deployed:

Camera-trap ID Date Time

Comments
L R checked checked

33
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