Animal Conservation (2004) 7, 285290

C 2004 The Zoological Society of London. Printed in the United Kingdom DOI:10.1017/S1367943004001477

Estimation of tiger densities in the tropical dry forests of Panna,

Central India, using photographic capturerecapture sampling

K. Ullas Karanth1, *, Raghunandan S. Chundawat2 , James D. Nichols3 and N. Samba Kumar4

1 Wildlife Conservation Society (International Programs), 2300, Southern Boulevard, Bronx, New York 10460, USA
2 Wildlife Conservation Society (India Program), 26-2, Aga Abbas Ali Road (Apt: 430), Bangalore 560 042, India
3 US Geological Survey, Patuxent Wildlife Research Center, Laurel, Maryland 20708, USA
4 Centre for Wildlife Studies, 26-2, Aga Abbas Ali Road (Apt: 430), Bangalore 560 042, India
(Received 4 September 2003; accepted 22 January 2004)

Abstract
Tropical dry-deciduous forests comprise more than 45% of the tiger (Panthera tigris) habitat in India. However,
in the absence of rigorously derived estimates of ecological densities of tigers in dry forests, critical baseline
data for managing tiger populations are lacking. In this study tiger densities were estimated using photographic
capturerecapture sampling in the dry forests of Panna Tiger Reserve in Central India. Over a 45-day survey
period, 60 camera trap sites were sampled in a well-protected part of the 542-km2 reserve during 2002. A
total sampling effort of 914 camera-trap-days yielded photo-captures of 11 individual tigers over 15 samp-
ling occasions that effectively covered a 418-km2 area. The closed capturerecapture model Mh , which
incorporates individual heterogeneity in capture probabilities, fitted these photographic capture history data
well. The estimated capture probability/sample, p = 0.04, resulted in an estimated tiger population size and
standard error ( N (S E N )) of 29 (9.65), and a density ( D(S E D)) of 6.94 (3.23) tigers/100 km2 . The estimated
tiger density matched predictions based on prey abundance. Our results suggest that, if managed appropriately,
the available dry forest habitat in India has the potential to support a population size of about 9000 wild tigers.

INTRODUCTION support adequate densities of ungulate prey species

Although surviving wild tiger populations are distributed
across 13 Asian countries, India is thought to harbour
substantially more tigers than any other range country
(Seidensticker, Christie & Jackson, 1999). In spite of the
greater habitat fragmentation and intense anthropogenic
pressures prevalent in India, the status of tigers appears
to be comparatively better because of two factors: (1) the
inherently high potential carrying capacities of tropical
forests in India (Karanth, 1995; Karanth & Nichols, 1998)
and (2) the presence of a reasonably effective protected
area system (Karanth, 2003).
Recent estimates suggest that more than 300 000 km2
of forests are potentially suitable for tigers in India
(Wikramanayake et al., 1999), but tigers have been
extirpated or occur only at low densities over most
of this area (Karanth & Chundawat, 2002; Karanth,
2003). Tropical dry-deciduous forests (Meher-Homji,
1990; Wikramanayake et al., 1999) form the bulk (> 45%)
of such potential tiger habitat (Chundawat, Gogate &
Johnsingh, 1999), but breeding populations of tigers
probably occur only in a few protected reserves that
(Karanth & Chundawat, 2002; Karanth, 2003). However,
because of the lack of rigorously derived estimates of
tiger densities, critical baseline data for managing tiger
populations in India are lacking (Karanth et al., 2003).
We used photographic capturerecapture sampling
methodology (Karanth & Nichols, 1998, 2002; Karanth,
Kumar & Nichols, 2002) to estimate the density of tigers
in a well-protected part of the Panna Tiger Reserve,
Madhya Pradesh State of Central India (Fig. 1) to better
understand tiger ecology in tropical dry forests. Our
specific objectives were:
1. To design and implement a rigorous sample survey
to estimate tiger population size and density using
capturerecapture modelling of the photo-capture
data.
2. To examine how tiger densities are related to abundance
of ungulate prey species.
3. To examine the implication of our results for the future
management of tigers in Panna and other dry forests of
Asia.

STUDY AREA
All correspondence to: K. Ullas Karanth, 26-2, Aga Abbas Ali Road
(Apt: 403) Bangalore, Karnataka-560 042, India. Tel: 91-80-2671- The landscape of Panna Reserve (542 km2 ) consists of
5364; Fax: 91-80-2671-5255; E-mail: ukaranth@wcs.org two extensive, step-like plateaus separated by 3080 m
286
N The spacing between camera traps and their schematic
Camera trap-site
Effectively sampled tiger habitat
Panna tiger reserve
Reserve boundary
Camera trap polygon
0 4 8 both flanks of the animal. To prevent possible theft and
Kilometers
Fig. 1. Map of the Panna study area showing the reserve boundary,
the camera trap locations, the trap polygon and the effectively
sampled tiger habitat.
high escarpments that run parallel to the river Ken, a
perennial water source, which flows through the central
Indian plains. There are 14 villages located inside the
reserve consisting of about 6000 people and 9500 head
of livestock. The dominant vegetation type is tropical
dry-deciduous forest (Meher-Homji, 1990), designated
as dry forest tiger habitat (Chundawat et al., 1999;
Wikramanayake et al., 1999). Structurally, the tiger
habitats include riparian gallery forests, open grasslands,
open woodlands with tall grasses, closed woodlands with
short or tall grasses and thorny woodlands. The area
receives an average annual rainfall of 1100 mm, mostly
during the monsoon between July and September, which
is followed by the cool season between October and
February. An increasingly dry summer sets in thereafter,
when the maximum temperature may frequently exceed
45 C. Due to the steep topography, rapid drainage and the
short wet season, the availability of surface water becomes
a severe limiting factor during the summer.
The reserve supports a diverse mammalian fauna be-
sides tigers, including large carnivore species such as leo-
pard (Panthera pardus), striped hyena (Hyaena hyaena),
sloth bear (Melursus ursinus), dhole (Cuon alpinus) and,
rarely, the wolf (Canis lupus). The principal wild prey
species of tigers are: sambar (Cervus unicolor), chital
K. U. KARANTH ET AL.
(Axis axis), nilgai (Boselephas tragocamelus), chinkara
(Gazella bennetti), chousinga (Tetraceros quadricornis)
and wild pig (Sus scrofa).
We carried out a reconnaissance of the entire reserve to
select a study area that had a relatively high abundance
of ungulate prey and encompassed the areas used by four
tigers that had been radio-collared earlier in the study
(Chundawat et al., 1999). This study area, situated in the
central portion of the reserve (Fig. 1), was relatively less
prone to anthropogenic pressures such as hunting, wood-
fuel collection and livestock grazing.
METHODS
Field methods
Based on the presence of tracks, scats and other evidence
indicative of frequent tiger activity, we identified 60
camera-trap sites spread out across the study area (Fig. 1).
India layout were based on survey design considerations
described elsewhere (Karanth & Nichols, 1998; Karanth
et al., 2002; Nichols & Karanth, 2002).
We used commercially available TrailmasterTM camera
traps. Each trap consisted of an electronic tripping device
activated by animal movement that simultaneously fired
two cameras through a multi-camera trigger to photograph
vandalism we used protective shells (Karanth & Nichols,
2002: 184186).
Because we had only 20 camera traps to cover 60 trap-
sites, we divided the study area into 3 trapping-blocks and
set traps at 20 sites within each block, for 15 successive
days. Thus, each sampling occasion (Otis et al., 1978)
combined captures from 3 days of trapping (1 day drawn
from each block), covering one pass over the entire area
(Nichols & Karanth, 2002: 133). There were 15 sampling
occasions during the 45 days of trapping, involving a total
effort of 914 trap-days. We used a standard data collection
format (Karanth & Nichols, 2002: 183) that allowed us to
assign each tripping and associated photographic event to
the correct sampling occasion.
We gave unique identification numbers to each roll
of film and to each photograph within it, enabling
us to correctly match the time, location and picture
resulting from each tripping, thereby enabling accurate
identification of individual tigers. Every photo-captured
tiger was given a unique identification number (e.g. PAT-
102) after examining the position and shapes of several
stripes on its flanks, fore-quarters and limbs (Fig. 2) as
described by Karanth et al. (2002). It was also often
possible to identify the sex of the animals from the
photographs.
Analytical methods
We constructed individual capture histories for the
identified tigers using a standard X-matrix format (Otis
et al., 1978; Nichols, 1992), in which 1 indicates capture
 Tiger density in dry forests
PAT- 101
PAT- 102
Fig. 2. Camera trap photographs of tigers PAT-101 (top) and
PAT-102 (bottom) obtained in Panna, showing differences in stripe
patterns.
of a particular animal during a specific sampling occasion,
while a 0 indicates that the animal was not captured
during that occasion. For example, a capture history
of 001110000000000 indicates that a particular tiger
was caught only on the third, fourth and fifth sampling
occasions in a survey with 15 occasions.
Because tigers are long-lived animals (Sunquist,
1981; Smith, 1993), we assumed that the sampled
population was demographically closed (Otis et al., 1978;
Karanth, 1995; Karanth & Nichols, 1998) during our
45-day survey. We analysed our capture history data
using CAPTURE (Rexstad & Burnham, 1991) free-
ware from http://www.mbr-pwrc.usgs.gov/software.html,
which provides a statistical test of the assumption of
population closure. The program also facilitates the
objective comparison of several probabilistic models of
the underlying capturerecapture process that are likely to
have generated the observed capture histories (Otis et al.,
1978; Nichols, 1992).
The analysis of capture history data requires com-
parisons between possible capturerecapture models using
a series of hypothesis tests and the results of an overall
discriminant function test, in order to select the most
appropriate abundance estimation model for a given
data set. The various capturerecapture models offered
consider the potential effects of the behavioural response
of tigers to camera trapping (e.g. trap-avoidance), time-
specific variation (e.g. weekly weather changes) and
heterogeneity between individual animals (e.g. caused by
287
factors such as territorial status or trap access), on capture
probabilities. If required, various combinations of these
three factors on capture probabilities can also be modelled
using the program CAPTURE.
We considered the following models (Otis et al., 1978;
Nichols, 1992) and estimation options implemented in
CAPTURE:
Mo , capture probability is the same for all tigers and is not
influenced by behavioural response, time, or individual
heterogeneity.
Mh , capture probabilities are heterogeneous for each
individual tiger, but not affected by trap response or
time.
Mb , capture probabilities differ between previously caught
and uncaught tigers due to trap-response behaviour, but
are not influenced by heterogeneity or time.
Mt , capture probability is the same for all individual tigers,
but varies during the survey only due to time-specific
factors.
The model selection process also considered more
complex models such as Mbh , Mth , Mtb and Mtbh , which
incorporate the effects of heterogeneity, trap response
and time, in various combinations. The overall model
selection function (Otis et al., 1978) scores potential
models between 0.01.0, with a higher score indicating
a better relative fit of the model to the specific capture
history data generated by the survey. For each analysis,
the program CAPTURE estimated capture probabilities
per sample ( p) and the tiger population size, N (i.e. the
number of tigers in the sampled area, inclusive of animals
that were not photo-captured at all).
We estimated the area effectively sampled ( A) as the
convex polygon connecting the outermost camera traps
plus a buffer area whose width (W ) is an estimate of
the half home range length, averaged for tigers in the
sampled area. This width was estimated as the average of
the maximum distances between photo-captures for each
individual tiger caught more than once during the study
(for details, see Wilson & Anderson, 1985; Karanth &
Nichols, 1998; Nichols & Karanth, 2002).
RESULTS
Photographic captures of tigers
We camera-trapped for 45 days during FebruaryApril
2002, partitioning the survey duration into 15 sampling
occasions, each covering 3 days of effort. The total
sampling effort realized was 914 trap-days. We obtained
36 photographs (17 right flanks, 19 left flanks) of
11 individual tigers (8 females, 3 males). Individual
tigers could be clearly identified by comparing their
stripe patterns (Karanth et al., 2002; Fig. 2). The
capture histories generated from the sample survey
(Table 1) reveal that sample size (number of individuals
captured) was small, as expected for low-density (< 10 in-
dividuals/100 km2 ) populations (Karanth & Chundawat,
2002).
288 K. U. KARANTH ET AL.

Table 1. Capture histories of individual tigers photographed in Estimates of capture probabilities and tiger
Panna Tiger Reserve, Central India, on 15 sampling occasions population size
during FebruaryApril 2002
To generate parameter estimates under the Mh model we
Individual used the jackknife estimator (Burnham & Overton, 1978;
identification Otis et al., 1978) implemented in CAPTURE, which had
number 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 performed well in earlier photographic capture studies
of tigers (Karanth, 1995; Karanth & Nichols, 1998).
PAT-101 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0
In our survey, eight individual tigers were captured only
PAT-102 0 0 1 1 1 0 0 0 0 0 0 0 0 0 0
PAT-103 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0
once, one animal was caught twice, one animal thrice and
PAT-104 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 one animal four times (Table 1). Using the Mh jackknife
PAT-105 0 0 0 1 0 0 0 0 0 0 0 1 1 0 1 estimator, the average capture probability per sample
PAT-106 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 ( p) was 0.0391 and the corresponding population size
PAT-107 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 estimate ( N ) was 29 with a standard error (S E N ) of 9.65.
PAT-108 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Thus, the overall probability of capturing a tiger present
PAT-109 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 in the sampled area (Mt+1 / N ) was only 38%. Therefore, it
PAT-110 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 was critically important for us to have used a population
PAT-111 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 estimation method that took into account the fact that
not all of the animals present in the study area would be
detected (Nichols, 1992; Thompson et al., 1998; Williams,
Nichols & Conroy, 2002).

Tests for population closure and model selection

The statistical test for population closure in CAPTURE
(Otis et al., 1978; Rexstad & Burnham, 1991) supported
our assumption that the sampled population was closed
for the study interval (z = 1.494, P = 0.07).
The goodness of fit test for model Mh (as opposed to any
alternative model) showed a reasonable fit ( 2 = 15.156;
d.f. = 14; P = 0.37), as did a similar test for Model Mb
( 2 = 24.143; d.f. = 19; P = 0.19). Because of sample size
constraints, we could not test the fits of models Mt or Mh
against model Mo .
The test of the behavioural response did not provide
evidence in support of model Mb ( 2 = 1.614; d.f. = 1;
P = 0.20). Similarly, the test for time-specific variation in
capture probabilities during the survey did not provide
support for model Mt ( 2 = 3.597; d.f. = 14; P = 0.997).
Further comparisons showed that Mh was a more likely
model than Mbh . The overall model selection test in
CAPTURE scored the competing models as follows:
Mo = 1.00, Mh = 0.91, Mb = 0.53, Mbh = 0.79, Mt = 0.0,
Mth = 0.46, Mtb = 0.49, Mtbh = 0.78.
Although the null model, Mo , ranked higher, the
estimator based on this model is not robust to violations
of the underlying assumption that capture probabilities do
not vary between individual tigers. Tigers have a complex
social organisation consisting of resident breeders that
maintain home ranges that overlap between the two sexes.
In addition, a proportion of the population consists of non-
breeding floaters that do not retain a fixed home range
(Sunquist, 1981; Smith, 1993; Karanth & Sunquist, 2000).
These spatial patterns, as well as the locations of traps in
relation to tiger movements and home range boundaries,
were likely to cause capture probabilities to vary between
individual tigers. Considering both ecological factors
and the model comparison results reported above, we
selected Mh as the most likely model for representing
the underlying capturerecapture process that generated
the tiger capture histories we observed (Table 1).
Estimates of effectively sampled area and tiger densities
The polygon formed by the outermost camera traps
measured 222.1 km2 . Using the approach based on
distances between photo-captures described in the
Methods, above, we estimated a buffer width (W (S E W ) of
2.94 (1.88) km) and an effectively sampled area A(S E A)
of 418.14 (136.47) km2 . Tiger density was obtained
by dividing the estimated population size ( N ) by the
The estimated density
effectively sampled area ( A).
( D(S E D)) for the Panna study area was 6.94 (3.23) tigers/
100 km2 .
DISCUSSION
Applicability of camera trap surveys of tigers
This study further supports earlier findings (Karanth &
Nichols, 1998; OBrien, Kinnaird & Wibisono, 2003;
Trolle & Kery, 2003; Karanth et al., 2004) that
photographic capturerecapture sampling is a reliable
technique for estimating the abundances of tigers and
other secretive animal species that can be identified
individually from their natural markings.
A comparison of our camera trapping results in Panna
with similar studies conducted at other sites (OBrien
et al., 2003; Karanth et al., 2004) shows that estimated
capture probabilities/sample ( p) can vary substantially
among sites (0.030.22) for tigers. Consequently, camera-
trapping studies that derive only raw trapping rates
(number of animal photographs/trap-day e.g. some studies
cited by Carbone et al., 2001) could have been improved by
employing capturerecapture survey designs and analyses
(also see Jennelle, Runge & Mackenzie, 2002). Given the
large number of camera trap surveys now being conducted
across the world, we believe there is room for improvement
in the survey designs and analytical protocols used in many
 Tiger density in dry forests
cases. As pointed out by Karanth & Nichols (2002), such
scaling up of camera trap studies requires more careful
planning, but no additional effort or resources.
Implications for tiger ecology and conservation
Our survey area within Panna Tiger Reserve had been
reasonably well protected from adverse anthropogenic
pressures such as forest fires, livestock grazing and
illegal hunting for about 5 years prior to our study.
Wild prey species of tigers were reasonably abundant at
31 ungulates/km2 or a biomass density of 3117 kg/km2
(R. S. Chundawat et al., unpublished data). The mean
tiger density at Panna was lower than at another dry forest
site at Ranthambore (11.46 tigers/100 km2 ), which had a
25-year history of protection. However, the mean density
(6.94 tigers/100 km2 ) in our study area, which covered
77% of the reserve, was reasonably high compared to
other human-impacted sites in India for which reliable
data on tiger densities are available (Karanth et al.,
2004).
Assuming an annual average prey requirement of
3000 kg of live prey/tiger (Schaller, 1967; Sunquist, 1981)
and that tigers take about 10% of the standing prey
numbers each year (Karanth et al., 2004), the estimated
tiger density of 6.94 tigers/100 km2 at Panna is close to the
prediction based on prey abundance (6.18 tigers/100 km2 ).
There are an estimated 150 000 km2 of tropical dry
forests persisting in India (Chundawat et al., 1999). If this
entire area can be as effectively protected as the Panna site
was prior to our study, these dry forests can potentially
support a population of about 9000 wild tigers. Based
on our observations at Panna, such a recovery of wild
tiger populations is potentially feasible with enhanced
anti-poaching patrols, restrictions on livestock grazing
and other biomass removals and voluntary relocations
of settlements from critical tiger habitats. Rigorous
monitoring, as conducted in this study, can help identify
which conservation actions could be most effective in
restoring prey and tiger populations.
Acknowledgements
This study was funded by the Wildlife Conservation
Society, New York, Save the Tiger Fund of the National
Fish and Wildlife Foundation, USA and Global Tiger
Patrol, UK. K.U.K., R.S.C. and N.S.K. were supported by
the Wildlife Conservation Society, New York, while J.D.N.
was supported by the US Geological Service-Patuxent
Wildlife Research Center, Laurel, MD. We thank all of the
above agencies for their sustained support of our work. We
are grateful to B. Sundaram and J. Hines who assisted us
with data collection and analysis, respectively. We thank
the Chief Wildlife Warden of Madhya Pradesh State, the
Field Director of Panna Tiger Reserve, their staff and the
Directorate of Project Tiger, Government of India, for
facilitating this study.
289
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