Eur J Wildl Res (2011) 57:627–637
DOI 10.1007/s10344-010-0473-y
ORIGINAL PAPER
Prey preference of large carnivores in Anamalai Tiger
Reserve, India
Arumugam Kumaraguru & R. Saravanamuthu &
K. Brinda & S. Asokan
Received: 3 July 2010 / Revised: 11 November 2010 / Accepted: 12 November 2010 / Published online: 26 November 2010
# Springer-Verlag 2010
Abstract Prey preferences of large carnivores (tiger (Panthera
tigris), leopard (Panthera pardus) and dhole (Cuon alpinus))
in the tropical forest of Anamalai Tiger Reserve (ATR) were
evaluated. This was the first study in ATR to estimate the
density of prey and the food habits of these large carnivores.
The 958-km2 intensive study area was found to have a high
mammalian prey density (72.1 animals per square kilometre)
with wild boar (20.61 animals per square kilometre) and chital
(20.54 animals per square kilometre) being the most common
species, followed by nilgiri tahr (13.6 animals per square
kilometre). When the density figures were multiplied by the
average weight of each prey species, a high biomass density
of 14,204 kg km−2 was obtained for the intensive study area.
Scat analysis and incidental kill observation were used to
determine the dietary composition of these predators. During
the study from the period of March 2001 to April 2004, 1,145
tiger scats, 595 leopard scats and 2,074 dhole scats were
collected and analysed. Kill data were based on direct
observation of 66 tiger kills and 39 leopard kills. Sambar,
with a density of 6.54 kg km−2 was the preferred prey for
Communicated by C. Gortázar
A. Kumaraguru (*) : R. Saravanamuthu : S. Asokan
PG Research and Development of Wildlife Biology,
Division of Zoology, AVC College,
Mayiladuthurai 609001, India
e-mail: wildlife_guru@yahoo.com
K. Brinda
Project Trainee, National Facility for Marine Cyanobacteria,
Bharathidasan University,
Tiruchirappalli 621 024, India
Present Address:
A. Kumaraguru
Centre for Cellular and Molecular Biology,
Hyderabad 500 007, India
these carnivores. Sambar constitutes 35% of the overall diet of
tiger, whereas it constitutes 17% and 25% in leopard and
dhole diets, respectively. Chital was utilized less than sambar
in the range of about 7%, 11% and 15% by tiger, leopard and
dhole, respectively. Predator diet was estimated more accu-
rately by scat analysis, which reveals 30% of smaller prey
species in leopard’s diet, which was not observed by kill data.
This study reveals that ATR harbours high prey density, and
these large carnivores seem mostly dependent on the wild
prey rather than on domestic livestock as in some other areas
in the subcontinent. These factors make ATR a potential area
for long-term conservation of these endangered carnivores.
Keywords Prey preference . Food habits . Tiger . Leopard .
Dhole . Anamalai Tiger Reserve (ATR)
Introduction
Food habits of large carnivores are central to the ecological
niche they occupy and play an important role in explaining
their social systems, behaviour and factors affecting predator
density. It may also have important implications in the life
history of their prey. Knowledge of food selection is critical in
understanding the life history strategies and developing sound
conservation recommendations (Miquelle et al. 1996). The
larger carnivores play a vital role as predators in regulating
and perpetuating the ecological processes and maintaining
the ecosystem as a whole (Sunquist and Sunquist 1989;
Terborgh et al. 2002; Kumaraguru 2002). Predatory strate-
gies are shaped and refined by natural selection to maximize
nutrient intake within the bounds of a wide range of
biologically relevant ecological constraints (Sunquist and
Sunquist 1989; Clutton-Brock and Harvey 1983). The
scenario gets complicated when several predatory species
hunt in the same area, resulting in a joint demand for a
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limited prey source. Ultimately, competition for a limited
prey resource leads to increased extinction risk of sympatric
carnivores (Hayward and Kerley 2008).
Studies on food habits are of conservation significance in
two ways, i.e. primarily, competition can reduce the popula-
tion size of an endangered carnivore (Caro and Stoner 2003)
(Hayward and Kerley 2008); secondarily, competition
between carnivores can affect the population of other species
at lower tropic levels (Jedrzejewska and Jedrzejewski 2005).
For example, the absence of carnivore species may increase
either herbivore population (Sinclair et al. 1990) or other
medium-sized carnivores (Sovada et al. 1995) or both
(Sinclair et al. 1990). Carnivores often regulate or limit the
numbers of their prey, thereby altering the structure and
function of entire ecosystems (Schaller 1972; Estes et al.
1998; Berger et al. 2001; Terborgh et al. 2002), and large
carnivores themselves are limited by the abundance of their
prey (Hayward et al. 2007). Prey selection of large carnivore
is a complex phenomenon (Bekoff et al. 1984; Kruuk 1972;
Sunquist and Sunquist 1989). The hypotheses so far
proposed to explain that prey selection by predators indicate
that the energetic benefits for the predator and proximate
mechanisms of selection shape their overall prey selection
(Griffiths 1975; Taylor 1976; Stephens and Krebs 1987;
Temple 1987; Karanth and Sunquist 1995).
Studies of feeding ecology of large carnivores rely on one or
a combination of many methods. There are varieties of
techniques including fecal analysis (Johnsingh 1983; Karanth
and Sunquist 1995; Swaminathan et al. 2002; Kumaraguru
2002) spoor tracking (Eloff 1984; Stander et al. 1997), radio
tracking (Seidensticker 1976) and direct observation of
animals hunting (Mills 1984). Scat analysis is non-invasive
and has been extensively applied in the studies of the
carnivore food habits, either alone (Karanth and Sunquist
1995; Hersteinsson and Macdonald 1996; Ranawana et al.
1998; Ramakrishnan et al. 1999; Khorozyan and Malkhasyan
2002) or in combination with data from predator kills
(Sunquist 1981; Johnsingh 1983). The analysis of food habits
provides practical and immediately accessible information for
the management of a particular species and occasionally aids
law enforcement and management needs (Korschgen 1971).
With the intention of collecting baseline information on
large carnivores and its prey species at ATR, the present study
was designed to estimate the density and biomass of major
prey species of large carnivores in the study area and to study
the food habits of large carnivores with reference to its prey
availability and utilization pattern. This study was the first
report in ATR to understand the ecological parameters, such as
availability of prey, which support the survival of these
endangered carnivores. Thus, increasing knowledge of prey
preference and food habits of these carnivores will enable us to
recognize the plasticity in the predator's ability to use the
available resources and conservation of these resources.
Eur J Wildl Res (2011) 57:627–637
Study area
The present study was a long-term investigation (from
March 2001 to April 2004) planned to provide information
on the large carnivores in ATR. This reserve was located in
the Coimbatore district of Tamil Nadu adjoining Kerala,
India (10° 12′ and 10° 54′ N and 76° 44′ and 77° 48′ E),
and was spread over 958 km2 (Fig. 1). Minimum and
maximum temperature ranges from 10°C in December to
37°C in April. Average annual precipitation varied between
1,178 and 2,268 mm. The landscape was highly undulating,
and accordingly the rainfall varies from an annual average
of 500 mm in the eastern part of the sanctuary to about
3,000 mm in the Western Plateau and slopes. ATR has a
heterogeneous nature of vegetation which ranges from
evergreen to tropical scrub and thorn forest. These habitats
varied in their extent of cover and also experience two wet
seasons, a winter season and a dry season, in a year, each
lasting for 3 months. The dominant vegetation types were
tropical evergreen forests, tropical semi-evergreen forests,
moist deciduous forests and teak plantation forests, inter-
spersed with patches of dry-deciduous forests, scrub forests
and grassland. The ATR was a home to teeming biodiver-
sity and supports a diverse assemblage of large mammal
prey species. Endemism was quite high, and the area was
the last stronghold of remnant populations of the nilgiri tahr
(Hemitragus hylocrius) and the lion-tailed macaque
(Macaca silenus). Sympatric carnivore species include the
tiger (Panthera tigris), leopard (Panthera pardus), stripped
hyena (Hyaena hyaena), sloth bear (Melursus ursinus) and
dhole (Cuon alpinus). The larger mammalian prey species
include gaur (Bos frontalis), sambar (Cervus unicolor),
nilgiri tahr (H. hylocrius), wild boar (Sus scrofa) and sloth
bear (M. ursinus). Medium prey species like chital (Axis
axis), barking deer (Muntiacus muntjak), lion-tailed macaque
(M. silenus), common langur (Semnopithecus entellus),
nilgiri langur (Presbytis johni), porcupine (Hystrix indica)
and other mesopredatory species such as jungle cat (Felis
chaus), leopard cat (Felis bengalensis) and fishing cat (Felis
viverrina) were found in the area. The smaller prey species
were mouse deer (Tragulus meminna), black-naped hare
(Lepus nigricollis), peafowl (Pavo cristatus), civets, mon-
gooses, hedgehog (Paraechinus micropus) and honey badger
(Mellivora capensis).
Methodology
Estimation of density, biomass and group size distribution
of prey species
The line transect method (Eberhardt 1968; Burnham et al.
1980; Buckland et al. 1993) was used to estimate densities of
Eur J Wildl Res (2011) 57:627–637
Fig. 1 Map showing the study area Anamalai tiger reserve along with grids for laying transects and carnivore scat collected roads
prey species in the study area. This method had been
consistently and efficiently used as a standard method to
determine animal densities under similar tropical conditions
(Karanth and Sunquist 1992, 1995; Varman and Sukumar
1995; Khan et al. 1996; Biswas and Sankar 2002). The data
were analysed using Gajah version 1.0 which was based on
Fourier Series estimation. Forty two transects which varied in
length between 2 and 5 km were laid randomly in the study
area. The total transect length of 127.4 km was monitored
eight times in the early morning (0600–1000 hours) and in
the late afternoon (1500–1900 hours), resulting in 1,020 km
of transect walked. The following sighting parameter such as
sighting angle (with a field compass); sighting distance
(ocular estimated); and group size, sex and age class of
individuals (whenever it was possible to classify them) were
recorded for each sighting in the transect. The major habitat
and microhabitat types were sampled in proportion to their
availability in the study area. The sampling effort was
uniform for different seasons of the year.
Stratified random sampling (Buckland et al. 1993) was
used. The stratified vegetation map of the study area was
divided into number of grids comprising 4 km2 in 1:50,000
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survey of India’s toposheet. The maps were based on recent
satellite images and aerial photographs of the vegetation. A
total of 250 grids were drawn, out of which 64 were
randomly selected (Fig. 1). The total number of grids
(samples) selected for each habitat was in proportion to the
area of availability. In the selected grids, one transect was
laid with a length of 2 km. Prey species populations were
estimated using direct sighting method. In the selected grid
(s), transects were laid randomly with respect to the
distribution of animals and topography of the area. Trans-
ects of known distance were marked, keeping in view that
changes in vegetation, if any, within the transect are kept
minimum in order to avoid attraction or repulsion of the
animals (prey species).
Carnivore sample collection
Scat samples were collected from well-defined sampling
areas along 18 different roads within the protected area of
the reserve. Attempts were made to select roads which
represented a particular vegetation type. Generally, large
carnivores have extensive home ranges; hence, roads
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located well within a particular habitat were considered for
this study, but not those which cut across two or more
different types of habitats.
Scat identification
Identification of dhole scat was fairly simple, as they tend
to defecate in the middle of the roads and paths unlike the
large cats, which defecate along the edge of the road or
path. Additionally, the entire pack of dhole defecates at the
same spot. Identification of tiger and leopard scats in the
areas where they co-exist has been largely based on size.
For instance, tiger scat was differentiated from leopard scat,
as a full-grown leopard was about one fourth the size of a
full-grown tiger (Seidensticker 1976), thus producing
identifiably smaller scat. Though complete scat of large
adult tigers can be easily differentiated from those of
leopards, there can be mistakes in identification when shape
alone was taken into account. The scats were identified
based on the shape as well as the associated signs such as
pugmarks or the size of the scrape in the present study. This
allowed greater accuracy as pug marks allowed more
accurate identification of the predator.
Scat analysis
A reference key was developed for the identification of prey
species on the basis of hair structure/morphology. All scat
samples were sun-dried and preserved in tagged polythene
bags for further analysis. Each scat was carefully broken and
soaked in water to separate prey remains, such as hair, bones,
hooves, teeth, feathers, etc. All these parts were observed and
analysed with a magnifying glass and under a light micro-
scope. They were identified against the reference collection
taken from captive animals by comparing features such as
structure, colour and medullary configuration to identify prey
species (Kopikar and Sabins 1976; Amerasinghe 1983;
Karanth 1993; Kitsos et al. 1995; Ranawana et al. 1998;
Ramakrishnan et al. 1999). The remains of one prey species
in one scat were scored as 1. If there were prey remains of
two species in a scat, each prey species was scored as 1.
Prey selectivity index
The most preferred prey of the these sympatric carnivores
estimated by Ivelv’s selectivity index (1961) was represented
in Fig. 3. Selectivity index value, which ranges from +1 to −1,
denotes preference for a particular prey with a positive value
and avoidance with a negative value. Electivity index was
calculated using the following formula.
ðri  niÞ
Ei ¼
ðri þ niÞ
Eur J Wildl Res (2011) 57:627–637
where Ei is Ivelv’s electivity measure of species i, ri is the
percentage of species i in the diet, and ni is the percentage of
species i in the environment.
Kill study
The tiger, leopard and wild dog kills were studied in order
to assess their prey selection. Clues like odour, alarm calls
of prey, carnivore signs and calls were useful to locate the
kill (Karanth and Sunquist 1995). Whenever the kill was
found relatively intact, the age and health of the killed
individual were recorded on the basis of the size and colour
of the animal, sexual characters, etc. Whenever possible,
the colour and texture of femur marrow fat were examined
in order to record the health condition of the kill as
suggested by Schaller (1967).
Reconstruction of predator diets
Frequency occurrence of mammalian prey in carnivore scat
was commonly used as a parameter in predation studies.
The frequency occurrence of different prey species in the
scat of tiger was converted to the relative biomass (Floyd et
al. 1978; Ackerman et al. 1984; Karanth and Sunquist
1995). The equation used is as follows:
Y ¼ 1:980 þ 0:035x ðTiger and LeopardÞ
Y ¼ 0:035 þ 0:020x ðDholeÞ
where Y is the kilogramme of prey consumed per field
collectible scat and x is the average weight of an individual
of a particular prey type (Ackerman et al. 1984). Multiply-
ing each Y by the number of scats found to contain a
particular prey species gave the relative weight of each prey
type consumed. These values were used to estimate per cent
biomass contribution of different prey species to the large
carnivore diet (Biswas and Sankar 2002).
Results
Density of prey species
Density estimation of individuals and groups of nine
potential prey species present in the study area was
summarized in Table 1. The study area harboured high
mammalian prey density of 72.1 animals per square
kilometre with chital and wild boar constituting about
50% of it. Among the different prey species, wild boar and
chital were the most abundant prey species with the highest
density (20 km−2). Although the next predominant prey
species was the nilgiri tahr (13.67 km−2), it was restricted in
its distribution to high-altitude grasslands, cliff areas of
Eur J Wildl Res (2011) 57:627–637 631

Table 1 Individual density, group density and group size of different prey species in ATR

Species name Group density (km−2) Group size Individual density (D/km2) 95% Confidence interval

Dg SE GS SE Lower Upper

Wild boar 1.79 0.168 11.21 0.46 20.61 11.48 28.56

Chital 0.94 0.1 22.24 1.87 20.54 8.09 22.25
Nilgiri tahr 0.66 0.15 19.79 3.29 13.67 2.1 23.26
Gaur 1.26 0.11 10.02 0.72 12.34 5.6 16.43
Sambar 1.59 0.14 4.19 0.17 6.54 4.31 8.38
Elephant 0.43 0.07 6.38 0.5 2.79 1.49 3.42
Black-naped hare 0.78 0.13 1 0 0.78 0.69 0.88
Barking deer 0.28 0.05 1.04 0.2 0.28 0.23 0.34
Mouse deer 0.15 0.04 1.14 0.36 0.18 0.12 0.22

scrub forest and adjoining evergreen habitat. Whereas other
larger prey species such as gaur and sambar were found
with relatively lower density values of 12.34 and
6.54 km−2, respectively, with wide distribution all over
the sanctuary. Nilgiri tahr, barking deer and mouse deer
were present at relatively lower densities than chital.
Wild boar showed the highest group density value of
1.79±0.17 km−2. Certain prey species such as chital and
nilgiri tahr showed higher density value but with lower
group density. The ecological density varied with group
size of different prey species. For example, prey species
such as chital and nilgiri tahr showed relatively lower group
density than gaur and sambar. However, because of higher
group size, they showed higher density value, which was
more than 13 km−2. Among the different prey species,
chital and nilgiri tahr showed the highest group size of 22±
1.87 and 19.79± 3.29, respectively. The solitary prey
species such as hare, barking deer and mouse deer did not
show any variation between the group density and actual
density. The largest herbivore, calves of elephant (Elephas
maximus), showed mean group size of 6.38±0.5 with
density value of 2.79 km−2. The density figures of ungulates
when multiplied with the average weight of the respective
species gave a biomass density of 14,204 kg km−2 for the
intensive study area which was compared with density of
other tropical forest in Table 2. The estimate of the prey
biomass density showed that the bulk of prey biomass of
gaur was 69% of the total standing prey biomass (Fig. 2).
Four mammalian prey species, spotted deer, wild boar, nilgiri
tahr and sambar, together form 99.9% of standing biomass.
Prey preference and diet composition of tiger
The average prey size of tiger was 92 kg with sambar,
nilgiri tahr and gaur as preferred prey. Based on Ivelv’s
selectivity index, tiger showed avoidance for other medium
prey such as chital and wild boar (Fig. 3). Analysis of 1,145
tiger scats revealed the presence of eight prey species with a
high preponderance of large-sized ungulates in the tiger’s
diet (Fig. 4). Sambar and gaur constituted 66% of the
overall diet composition of tiger followed by nilgiri tahr
and chital which constituted about 11% and 7%, respec-
tively. Other prey species such as wild boar, porcupine and
black-naped hare constituted less than 10% of the total diet
composition of tiger. Comparison with the prey availability
and the consumption of prey by these predators was
depicted in Fig. 6. Black-naped hare and porcupine
contributes 1.34% and 2.21% of tiger’s diet, which was
represented only in scat data. Analysis of 66 kills of tiger
revealed the presence of seven prey species (Fig. 5).
Sambar and gaur constituted 22% and 21%, respectively.
Chital formed 12% of the diet by kill data which was high
when compared to that by scat analysis. The per cent
occurrence of cattle by kill data was high for tiger with
21%, when compared to 5% by scat data.
Prey preference and diet composition of leopard
Black-naped hare and mouse deer were the most frequently
taken prey of leopards, followed by sambar and nilgiri tahr.
With an average prey size of 37 kg, leopard showed
avoidance for wild boar, chital and gaur (Fig. 3). Analysis
of 595 leopard scats reveals that approximately 40%
composition of its diet was of medium-sized prey species
(Fig. 4). Sambar, wild boar and black-naped hare formed
the major composition of leopard’s diet with 45%. The per
cent occurrence of black-naped hare (15%), nilgiri langur
(8%), mouse deer (3%) and mongoose (0.77%) as leopard’s
prey was revealed only from scat data analysis. Leopard
had a more varied diet and regularly consumed at least 12
different prey species. Similarly, chital, gaur, goat and
nilgiri langur contributed 37% of its overall diet. Analysis
of 39 kills of leopard revealed the presence of eight prey
species (Fig. 5). Sambar constituted 23% of its overall diet.
632 Eur J Wildl Res (2011) 57:627–637

Table 2 Comparison of wild biomass density of herbivores at tropical sites

Region Area Habitat type Biomass density (kg km−2) References

Asia Nagarhole Deciduous forest 14,744 Karanth and Sunquist (1992)

Bandipur Dry forest–woodland 14,520 Johnsingh (1983)
ATR Deciduous forest 14,204 Present study
Kanha Moist forest-meadows 1,592 Schaller (1967)
Pench Deciduous forest 6,013 Biswas and Sankar (2002)
Wilpattu Dry forest-meadows 766 Eisenberg and Lockhart (1972)
Bardia Moist forest-grasses 3,101 Dinerstein (1980)
Chitwan Moist forest-grasses 2,581 Tamang (1982)
Africa Rwenzori Swamps-savanna 21,373 Eltringham (1979)
Manyara Dry savanna 19,259 Eltringham (1979)
Serengeti Dry savanna 6,840 Eltringham (1979)
Mara Dry savanna 19,200 Stelfox (1986)
Nairobi Dry savanna 4,470 Eltringham (1979)
Gabon Evergreen forest 1,020 Prins and Reitsma (1989)
South America Masaguaral Dry forest-pasture 711 Eisenberg (1980)
Barro Colorado Evergreen forest 3,553 Eisenberg (1980)
Manu Evergreen forest 1,220 Terborgh (1986)
Pantanal Moist forest pasture 295 Eisenberg (1980)

Chital and gaur constituted 21% and 5%, respectively.
Chital formed 12% of the diet by kill data which was high
then compared to that by scat analysis. The per cent
occurrence of cattle by kill data was high for leopard with
23%, when compared to 6% by scat data.
Prey preference and diet composition of dhole
Barking deer, sambar and mouse deer were the most
frequently taken prey of dhole. It showed avoidance for
wild boar, chital, gaur, mouse deer and nilgiri tahr (Fig. 3)
with an average prey size of 36 kg. The diet spectrum of
dhole included about 14 species (Fig. 4). Their specific
territorial marking behavior combined with their use of
latrine site for defecation enabled us to collect large
numbers of scats for this study. Two thousand and seventy
four scats were collected and analysed to understand the
food habits of dhole in ATR. Sambar (26%), chital (15%),
black-naped hare (13%), gaur (12%) and wild boar (10%)
together constituted more than 75% of prey composition of
the dhole’s diet (Fig. 6).

Discussion

Over most of their range, tigers co-exist with other

Fig. 2 Proportion of the standing biomass of different prey species in ATR
predatory carnivores such as leopards and dholes. The
densities of different predator species within such guilds
appear to be greatly influenced by the relative abundance of
different size classes of prey species in the assemblage
(Karanth and Sunquist 1995; Karanth and Sunquist 2000;
Karanth et al. 2004). We examine our results in an attempt
to provide a more reliable measure of dietary patterns of
these three sympatric carnivores in ATR.
Prey densities estimated in the present study, when
compared with that of other tropical forests from other species
revealed that ATR harbours a high density of chital and nilgiri
tahr. ATR is dominated by fairly open canopy with a
heterogeneous habitat, ranging from evergreen to tropical
thorn forest. This condition of high habitat heterogeneity
Eur J Wildl Res (2011) 57:627–637
Fig. 3 Overall prey preference
index (Ivelv’s) of tiger, leopard
and dhole for different prey
species in ATR
probably favoured the high density of grazer (Eisenberg and
Seidensticker 1976) such as chital. ATR also harbours large-
sized prey such as sambar and gaur. Contrary to intuition,
three smaller species such as barking deer, mouse deer and
black-naped hare were lower in densities when compared to
that of chital and sambar. These prey species considered to
be selective feeders on rich but scarce food items such as
shoots and fruits might be a reason for their low density.
Their solitary nature and territorial spacing mechanism may
also contribute to their relatively low densities.
ATR ranks third when compared with other tropical
sites, with a biomass density of 14,204 kg km−2. In places
Fig. 4 Proportion of prey occur-
rence in scats (%) of tiger
(n=1,145), leopard (n=595) and
dhole (n=2,074) in ATR
633
where overall prey biomass densities were greater than
1,000 kg km−2, the tiger preferentially takes larger prey
(Karanth and Nichols 1998). The leopard was morpholog-
ically adapted to kill large prey but may depend heavily on
locally abundant small prey in difficult times (Hayward et
al. 2006). Hence, the density and distribution of all the
three carnivores were often shaped by the dominant prey
species because dominant prey species provides the bulk of
available prey biomass (Kumaraguru 2002). Thus, identi-
fying the dominant prey was often the first step towards
understanding the carrying capacity for large carnivores in a
particular area. Based on the preferences of these three
634
Fig. 5 Proportion of prey oc-
currence in kills (%) of tiger
(n=66) and leopard (n=39)
in ATR
carnivores, sambar deer was the critical prey species in
ATR. Whereas in most of the other tropical south Indian
forest, chital was also a preferred prey species of leopards
(Hayward et al. 2006) and for these three large carnivores
as reported from Bandipur (Johnsingh 1983), Nagarahole
(Karanth and Sunquist 1995) and Mudumalai (Swaminathan
et al. 2002). This trend of chital was not only reported from
South India but also from neighbouring countries such as
Chitwan, Nepal (McDougal 1977). Similarly, other species
such as red deer in Sikhote Alin, Russia (Miquelle et al.
1996), and barking deer in Thailand (Rabinowitz and
Nottingham 1986) were the dominant prey species for large
carnivores especially for tigers.
The prey species was also selected based on the prey
size which tend to support the concept of optimal foraging
Fig. 6 Relation between per
cent avilable and consumption
of different prey species by
tiger, leopard and dhole in ATR
Eur J Wildl Res (2011) 57:627–637
which was based on the premise of a maximization of
energy return to the predator for each prey encounter
(Werner and Hall 1974; Krebs and Davies 1979). The mean
and range of prey size can increase with an increase in
predator size (Wilson et al. 1995). Prey was categorized
into three size classes distinguishable in the field based on
species and weight. Black-naped hare, porcupine, nilgiri
langur, mouse deer, mongoose and bird species were
classified as small prey (5–30 kg). Chital, nilgiri tahr, wild
boar and goat were considered as medium-sized prey (31–
100 kg). Sambar, gaur and cattle were categorized as large
(>100 kg) prey. In ATR, the average weight of a wild prey
species of tiger and leopard was 92 and 37 kg, respectively.
Similar to that of leopard, the prey size for dhole was also
36 kg. Our results showed that large-sized prey (sambar,
Eur J Wildl Res (2011) 57:627–637
gaur and cattle) comprised 70%, medium-sized prey (chital,
nilgiri tahr, wild boar and goat) comprised 25% and small
prey (porcupine, black-naped hare, nilgiri langur, mouse
deer and mongoose) provided only 3.5% of tiger’s diet
based on scat analysis. On the other hand, medium-sized
prey dominated leopard diet, contributing to 37%. Large
prey and small prey provided about 31% and 27% based on
scat analysis. Dhole depend even more predominantly on
medium-sized prey (~50% of overall diet).
Sambar was the most important prey species for tigers,
leopard and dhole in our study. In the site where predation on
sambar was more, a correspondingly lower degree of chital
predation by tigers is recorded (Biswas and Sankar 2002). In
ATR, chital occurred in high densities, which might have
increased their predation rate, but their gregarious nature was
supposed to be one of the factors that reduced the chance of
predation by these large carnivores (Karanth and Sunquist
1995). Predation of wild boar in ATR suggests high overlap
of habitat use between these large carnivores and wild boar
which might have facilitated a high level of predation on the
wild boar (Miquelle et al. 1996). The selectivity for wild
boar as prey suggests that these predators were making
foraging decisions on the basis of energy gain and not on
injury risk (Sunquist and Sunquist 1989).
Tigers depend on concealment and ambush to capture
prey. In ATR, grasslands were interspersed with islands of
shola which enables tigers to capture tahr and that could be
the reason for the higher proportion of tahr in tiger scat.
The prime factor influencing the predation of gaur in the
study area was probably due to their high density
(12 km−2). Presence of porcupine remains in the scats of
tiger reflects the ability of tiger to hunt this aggressive prey
species. Similar observations have also been reported from
Mudumalai on tiger which fed upon sloth bear (Swaminathan
et al. 2002). The restricted distribution of chital and nilgiri
tahr in specific habitats may have compelled the tiger to go
for different prey selection.
The higher proportion of cattle in the diet of these large
carnivores could be attributed to the considerable behavioural
plasticity of these predators which could enable them to
survive in diverse forest environments. Supplementary prey
such as cattle become more important due to low prey
availability in specific size class and competition between co-
predators for food and space. The role of livestock in the
feeding habits of these large carnivores was relevant not only
to the survival of these predators but also to understanding and
thus managing their conflict with farmers.
In contrast to the preference of tiger, leopard and dhole
showed preference for medium- and small-sized prey species.
Occurrence of smaller species such as black-naped hare,
mouse deer, porcupine and mongoose in their diet might be
due the weight of these species that are within the preferred
prey weight range of leopard and dhole. The presence of
635
arboreal prey such as langur in the scats of leopard could be
linked to the leopard’s greater arboreality and crypticity in
comparison to the other two carnivores. The dietary prefer-
ences of dhole are concluded based only on the scat analysis in
this study. Since prey such as chital and sambar are eaten
almost entirely, only the lower jaws remain, making detection
of such kills unlikely (Venkataraman et al. 1995). The
presence of black-naped hare and bird species in the diet of
dhole could be due to pack hunting and the ability of the
dholes to flush out and hunt the smaller and cryptic prey
species. While being too small for the pack as a whole, such
prey was sufficient for an individual dog. Black-naped hare
when flushed out by the dogs was usually grabbed by the
nearest dog and eaten alone or shared with another dog while
the hunt continues for a larger prey more suitable for the
entire pack. It was likely that risk of injury during prey
capture may be the reason underlying the lower per cent
composition of large prey in the scats of leopard (Hayward et
al. 2006) and dhole.
In the present study, the data from scat analysis showed
that smaller prey species constituted more than 30% of the
leopard’s diet, which was not clear from the kill observa-
tions. This indicates that scat analysis give a more accurate
estimate, whereas kill samples underestimated the propor-
tion of smaller prey in the carnivore’s diet. Hence in the
present investigation, the scat of predators and kill data
were judiciously taken into consideration to determine the
prey composition of predator’s diet.
ATR was one of the areas that still harbour high prey
density with predators mostly depending on wild prey
rather than on domestic livestock for food as in many other
areas of the Indian subcontinent. From the present study, it
can be concluded that ATR, because of its high wild prey
density, has the potential to accommodate a higher density
of predators (Hayward et al. 2007; Kumaraguru 2002),
making it comparable to few of the best remaining tiger
habitats of the Indian subcontinent. Thus, protection of the
habitat along with regular monitoring of endangered species
such as tigers, leopard and dhole along with their prey
population using comparable scientific methods was essen-
tial for ATR to emerge as one of the most important areas
for wildlife conservation in India.
Acknowledgement We acknowledge PCCF and CCF, Tamil Nadu
Forest Department (TNFD), for permitting us to enter and to carry out
research in protected areas of the forest. We extend our sincere thanks to the
TNFD for funding part of this project. We thank all forest rangers, foresters
and field staff of ATR without whom the project could not be completed.
We express our sincere thanks to principal, HOD and all staff members of
the Department of Wildlife Biology, AVC College, for providing us full
freedom to carry out the research. We thank Dr. J.C. Daniel, BNHS, and
Ajay A Desai., Asian Elephant Co-chair person, IUCN, for his valuable
guidance to carry out this research and for useful suggestion during
manuscript preparation. We express our sincere gratitude to the two
anonymous authors for their valuable suggestions.
636
References
Ackerman BB, Lindzey FG, Hemker TP (1984) Cougar food habits in
Southern Utah. J Wildl Manage 48:147–155
Amerasinghe FP (1983) The structure and identification of the hair of
the mammals of Sri Lanka. Ceylon J Sci Biol Sci 16:76–125
Bekoff M, Daniels TJ, Gittleman JL (1984) Life history patterns and
the comparative social ecology of carnivores. Ann Rev Ecolog
Syst 15:191–232
Berger J, Stacey-Peter B, Bellis L, Johnson MP (2001) A mammalian
predator–prey imbalance: grizzly bear and wolf extinction affect
avian neotropical migrants. Ecol Appl 11:947–960
Biswas S, Sankar K (2002) Prey abundance and food habit of tigers
(Panthera tigris tigris) in Pench National Park, Madhya Pradesh,
India. J Zool 256:411–420
Buckland ST, Anderson DR, Burnham KP, Laake JL (1993) Distance
sampling: estimating abundance of biological populations.
Chapman & Hall, London
Burnham KP, Anderson DR, Laake JL (1980) Estimation of density
from line transect sampling of biological populations. Wildl
Monogr 72:1–202
Caro TM, Stoner CJ (2003) The potential for interspecific competition
among African carnivores. Biol Conserv 110:67–75
Clutton-Brock TH, Harvey PH (1983) The functional significance of
variation in body size among mammals. In: Eisenberg JF,
Kleiman DG (eds) Advances in the study of mammalian
behaviour. Allen, Lawrence, pp 632–663
Dinerstein E (1980) An ecological survey of the Royal Karnali-Bardia
Wildlife Reserve, Nepal. Part III: ungulate populations. Biol
Conserv 18:5–38
Eberhardt LL (1968) A preliminary appraisal of line transect. J Wildl
Manage 32:82–88
Eisenberg JF, Lockhart M (1972) An ecological reconnaissance of
Wilpattu National Park, Ceylon. Smithsonian contribution to
Zoology 101:1–118
Eisenberg JF, Seidensticker J (1976) Ungulates in Southern Asia: a
consideration of biomass estimates for selected habitats. Biol
Conserv 10:293–307
Eisenberg JF (1980) The density and biomass of tropical mammals. In
Conservation Biology, an Evolutionary-Ecological Perspective
(eds M. E. Soule and B. A. Wilcox). Sinauer Press, Sunderland
Eloff FC (1984) Food ecology of the Kalahari lion (Panthera leo).
Koedoe 27:249–258
Eltringham SK (1979) The ecology and conservation of large African
mammals. Macmillan, London
Estes JA, Tinker MT, Williams TM, Doak DE (1998) Killer whale
predation on sea otters linking oceanic and near shore ecosystems.
Science 282:473–476
Floyd TJ, Mech LD, Jordan PJ (1978) Relating wolf scat contents to
prey consumed. J Wildl Manage 42:528–532
Griffiths D (1975) Prey availability and the food of predators. Ecology
56:1209–1214
Hersteinsson P, Macdonald DW (1996) Diet of Arctic foxes (Alopex
lagopus) in Iceland. J Zool Lond 240:457–474
Hayward MW, Kerley GIH (2008) Prey preferences and the
conservation status of Africa's large predators. S Afr J Wildl
Res 38:93–108
Hayward MW, Henschel P, Brien J, Hofmeyr M, Balme G, Kerley
GIH (2006) Prey preferences of the leopard (Panthera pardus). J
Zool Lond 270:298–313
Hayward MW, Brien J, Kerley GIH (2007) Carrying capacity of large
African predators: predictions and tests. Biol Conserv 139:219–
229
Jedrzejewska B, Jedrzejewski W (2005) Large carnivores and
ungulates in European temperate forest ecosystems: bottom-up
Eur J Wildl Res (2011) 57:627–637
and top-down control. In: Ray JC, Redford KH, Steneck RS,
Berger J (eds) Large carnivores and the conservation of
biodiversity. Island, Washington, pp 230–246
Johnsingh AJT (1983) Large mammalian prey–predators in Bandipur.
J Bombay Nat Hist Soc 80:1–57
Karanth KU (1993) Predator–prey relationships among large mammals of
Nagarhole National Park. Mangalore University, Bangalore, India.
Ph.D. dissertation
Karanth KU, Nichols JD (1998) Estimation of tiger densities in India
using photographic captures and recaptures. Ecology 79:2852–
2862
Karanth KU, Sunquist ME (1992) Population structure, density and
biomass of large herbivores in the tropical forests of Nagarhole,
India. J Trop Ecol 8:21–35
Karanth KU, Sunquist ME (1995) Prey selection by tiger, leopard and
dhole in tropical forests. J Anim Ecol 64:439–450
Karanth KU, Sunquist ME (2000) Behavioural correlates of predation
by tiger (Panthera tigris), leopard (Panthera pardus) and dhole
(Cuon alpinus) in Nagarhole, India. J Zool Lond 250:255–265
Karanth KU, Nicholas JD, Kumar NS, Link WA, Hines JE (2004)
Tigers and their prey: predicting carnivore densities from prey
abundance. Proc Natl Acad Sci 101:4854–4858
Khan JA, Chellam R, Rodgers WA, Johnsingh AJT (1996) Ungulate
densities and biomass in the tropical dry deciduous forests of Gir,
Gujarat, India. J Trop Ecol 12:149–162
Khorozyan I, Malkhasyan A (2002) Ecology of the leopard (Panthera
pardus) in Khosrov Reserve, Armenia: implications for conserva-
tion. Societa Zoologica ‘La Torbiera’, Italy, scientific report no. 6
Kitsos AJ, Hunter MJ, Sabnis JH, Mehta A (1995) A guide to the
identification of some Indian mammal hairs. In: Berwick SH,
Saharia VB (eds) The development of international principles and
practices of wildlife research and management, Asian and American
approaches. Oxford University Press, New Delhi, pp 125–130
Kopikar BR, Sabins JH (1976) Identification of hairs of some Indian
mammals. J Bombay Nat Hist Soc 73:5–20
Korschgen LJ (1971) Procedures for food-habits analysis. In: Giles
RH (ed) Wildlife management technique. Wildlife Society,
London, pp 233–258
Krebs JR, Davies NB (1979) Behavioral ecology and evolutionary
approach. Sinauer, Sunderland
Kruuk H (1972) The spotted hyena: a study of predation and social
behavior. University of Chicago Press, Chicago
Kumaraguru A (2002) The influence of prey species diversity and
densities in different vegetation types on the foraging ecology
and community structure of large carnivores in Indira Gandhi
Wildlife Sanctuary and National Park, South India. Report
submitted to Tamil Nadu Forest Department, Tamil Nadu, India
McDougal C (1977) The face of the tiger. Rivington Books, London
Mills MGL (1984) Prey selection and feeding habits of the large
carnivores in the Southern Kalah. Koedoe 27:281–294
Miquelle DG, Smirnov EN, Quigley HB, Hornocker MG, Nikolaev
IG, Matyukshin EN (1996) Food habits of Amur tigers in
Sikhote-Alin Zapovednik and the Russian Far East and implications
for conservation. J Wildl Res 1:138–147
Prins HHT, Reitsma JM (1989) Mammalian biomass in an African
equatorial rain forest. J Anim Ecol 58:851–861
Rabinowitz A, Nottingham BG (1986) Ecology and behaviour of the
jaguar (Panthera onca) in Belize, Central America. J Zool
210:149–159
Ramakrishnan U, Coss RG, Pelkey NW (1999) Tiger decline caused
by the reduction of large ungulate prey: evidence from a study of
leopard diets in Southern India. Biol Conserv 89:113–120
Ranawana KB, Bambaradeniya CNB, Bogahawatte TD, Amerasinghe
FP (1998) A preliminary survey of the food habits of the Sri
Lanka leopard (Panthera pardus fusca) in three montane wet
zone forests of Sri Lanka. Ceylon J Sci Biol Sci 25:65–71
Eur J Wildl Res (2011) 57:627–637
Schaller GB (1967) The deer and the tiger. University of Chicago
Press, Chicago
Schaller GB (1972) The Serengeti lion: a study of predator–prey
relations. University of Chicago Press, Chicago, p 504
Seidensticker J (1976) On the ecological separation between tigers and
leopards. Biotropica 8(4):225–234
Sinclair ARE, Olsen PD, Redhead TD (1990) Can predators regulate
small mammal populations? Evidence from house mouse outbreaks
in Australia. Oikos 59:382–392
Sovada MA, Sargeant AB, Grier JW (1995) Differential effects of
coyotes and red foxes on duck nest success. J Wildl Manage 59:1–9
Stander PE, II Ghau, Tsisaba D, II OMA, VI (1997) Tracking and the
interpretation of spoor: a scientifically sound method in ecology.
J Zool 242:329–341
Stelfox JB (1986) Effects of livestock enclosures (bomas) on the
vegetation of the Athi Plains, Kenya. Afr J Ecol 24:41–45
Stephens DW, Krebs JR (1987) Foraging theory. Princeton University
Press, Princeton
Sunquist ME (1981) The social organization of tigers (Panthera tigris) in
Royal Chitwan National Park, Nepal. Smithsonian contributions to
zoology no. 336. Smithsonian Institution Press, Washington, DC,
p 98
Sunquist ME, Sunquist FC (1989) Ecological constraints on predation by
large felids. In: Gittleman JL (ed) Carnivore behaviour, ecology and
evolution. Cornell University Press, Ithaca, pp 283–301
Swaminathan S, Desai AA, Daniel JC (2002) Large carnivores in
Mudumalai Wildlife Sanctuary and National Park. Report
submitted to Bombay Natural History Society, Bombay, 80
637
Tamang KM (1982) The status of tiger and its impact on principal
prey populations in the Royal Chitawan National Park, Nepal.
Ph.D Thesis, Michigan State University, East Lansing
Taylor RJ (1976) Value of clumping to prey and the evolutionary
response of ambush predators. Am Nat 110:13–29
Temple SA (1987) Do predators always capture sub standard
individuals disproportionately from prey populations? Ecology
68:669–674
Terborgh J (1986) Keystone plant resources in the tropical forest.
Sinauer Associates, Sunderland
Terborgh J, Lopez L, Nunez P, Rao M, Shahabudin G, Orihuela G,
Riveros M, Ascanio R, Adler GH, Lambert TD, Balbas L (2002)
Ecological meltdown in predator-free forest fragments. Science
294:1923
Varman KS, Sukumar R (1995) The line transect method for
estimating densities of large mammals in a tropical deciduous
forest: an evaluation of models and field experiment. J Biosci
20:273–287
Venkataraman AB, Arumugam R, Sukumar R (1995) The foraging
ecology of dhole (Cuon alpinus) in Mudumalai Sanctuary,
southern India. J Zool 237:543–561
Werner EE, Hall D (1974) Optimal foraging and the size selection of
prey by the bluegill sunfish (Lepomis macrochirus). Ecology
55:1042–1052
Wilson RP, Putz K, Gremillet D, Culik BM, Kierspel M, Regel J,
Charles AB, Lage J, Cooper J (1995) Reliability of stomach
temperature changes in determining feeding characteristics of
seabirds. J Exp Biol 198:1115–1135