Journal of Threatened Taxa |
www.threatenedtaxa.org | 26 April 2020 | 12(5): 15535–15546
ISSN 0974-7907 (Online) | ISSN 0974-7893
(Print)
doi: https://doi.org/10.11609/jott.5607.12.5.15535-15546
#5607 | Received 09 December 2019 | Final
received 07 April 2020 | Finally accepted 18 April 2020
Prey selection and food habits of
the Tiger Panthera tigris
(Mammalia: Carnivora: Felidae) in Kalakkad-Mundanthurai
Tiger Reserve, southern Western Ghats, India
Bawa Mothilal
Krishnakumar 1, Rajarathinavelu
Nagarajan 2 & Kanagaraj Muthamizh Selvan 3
1,2 PG and Research Department of Zoology and Wildlife Biology, A.V.C.
College (Autonomous), (Affiliated to Bharathidasan University,
Tiruchirappalli), Mannampandal, Mayiladuthurai, Tamil
Nadu 609305, India.
1,3 Department of Ecology & Environmental Sciences, School of Life
Sciences, Pondicherry University, R.V. Nagar, Kalapet,
Puducherry 605014, India.
2 Centre
for Research in Animal Behaviour (CRAB), Department of Psychology, Washington
Singer Laboratories, University of Exeter,
Perry Road, Exeter, EX4 4QG, UK.
3 Scientist
D/Joint Director, Project Elephant, Ministry of Environment Forest and Climate
Change, Indira Paryavaran Bhawan,
New Delhi, 110003, India.
1 maduraisauf@gmail.com, 2 oystercatcher@rediffmail.com,
3 tamildove@gmail.com
(corresponding author)
Editor:
Mewa Singh, University of Mysore, Mysuru, India. Date of publication: 26 April
2020 (online & print)
Citation: Krishnakumar, B.M., R. Nagarajan & K.M. Selvan (2020).
Prey selection and food habits of
the Tiger Panthera tigris
(Mammalia: Carnivora: Felidae) in Kalakkad-Mundanthurai
Tiger Reserve, southern Western Ghats, India. Journal of Threatened
Taxa 12(5): 15535–15546. https://doi.org/10.11609/jott.5607.12.5.15535-15546
Copyright:
© Krishnakumar et al. 2020. Creative Commons
Attribution 4.0 International License. JoTT allows unrestricted use, reproduction, and
distribution of this article in any medium by providing adequate credit to the
author(s) and the source of publication.
Funding: This research was funded by DST-INSPIRE (Faculty Award/IFA13-LSBM84), Ministry of Science & Technology, Department of Science & Technology, Government of India.
Competing interests: The authors declare no competing interests.
Author details: B.M. Krishnaumar is a PhD student in PG and Research Department of
Zoology & Wildlife Biology at A.V.C. College (Autonomous) (Affiliated to
Bharathidasan University, Tiruchirappalli). He has previously worked in
Department of Ecology and Environmental Sciences, Pondicherry University, under
the DST-INSPIRE Project. Dr. K. Muthamizh
Selvan graduated in MSc Wildlife Biology from Bharathidasan University,
then entered India’s Wildlife Institute, where he worked on big sympathetic
carnivores at Pakke Tiger Reserve, Arunachal Pradesh.
He holds a PhD in wildlife sciences from the Saurashtra University, Gujarat.
Upon completing his PhD, he received the DSTI-INSPIRE Faculty Award from the
Science and Technology Department and Joined in Department of Ecology and
Environmental Sciences, Pondicherry University as a DST-INSPIRE Faculty
Awardee. He has also guided several post-graduation students for their
dissertation. He currently works at Project Elephant Cell, Ministry of
Environment, Forest & Climate Change, New Delhi as Scientist D’ cum Joint
Director. Dr. R. Nagarajan was Commonwealth Scholar and an Early
Career Awardee of Leverhulme Trust, London, and
currently is Principal in A.V.C. College (Autonomous) & Head, PG and
Research Department of Zoology and Wildlife Biology, conducting research in behavioural ecology of wildlife pertaining to foraging and
nesting and research theme includes factors influencing population of wildlife,
nest-selection and life-history strategies of birds, role of Barn-Owl in rodent
pest control. He has published more than 100 research articles in peer-reviewed
national and international journals and also guided PhD scholars in Wildlife
Biology.
Author contribution: Conceptualization – BMK, KMS and RN; methodology -
KMS, and RN; formal analysis – BMK, KMS and RN; conducting field work – BMK;
Preparing manuscript – All author contributed equally.
Acknowledgements: We are grateful to Tamil Nadu Forest Department for
granting the permission to conduct the study at KMTR. We are grateful to
DST-INSPIRE faculty program for financial support. We give special thanks to
Head of the Department of Ecology and Environmental Science, Pondicherry
University. This research is part of first author’s PhD dissertation in
Wildlife Biology and he extend his sincere thanks to the Management and
Principal of A.V.C. College (Autonomous) for necessary support to this study.
We thank the Chief Conservator of Forest and Field Director of Kalakkad-Mundanthurai Tiger Reserve for permission to carry
out research at KMTR. We are extremely thankful to all administrative staff in
KMTR, in particular, Mr. Velladurai, Mr. Ilango and Mr. Karthikeyan for all the support and
hospitality during our stints in KMTR. We extend a special word of thanks to
Mr. M. Manikandan, Mr. P. Radhakrishnan, Mr. Jeyakumar,
Mr.Madasami for assisting in the field and invaluable
field expertise. Thanks are also due to Dr. Venkateswara Sharma, Department of Biotechnology,
Pondicherry University, for the use of his laboratory facilities. We would like to thank three anonymous
reviewers for their constructive comments and suggestions on an earlier draft
of this article that greatly improved our manuscript.
Abstract: The Endangered Tiger Panthera tigris is
the largest felid, distributed over 1.1 million km2 globally. Conservation of Tigers largely depends on the
preservation of its natural prey base and habitats. Therefore, the availability of prey and its
selection play a major role in the sustainable future of Tigers in the given
landscape. The current study assesses
the prey selection patterns by Tigers in tropical evergreen forest of the Kalakkad-Mundanthurai Tiger Reserve (KMTR), southern
Western Ghats, India. Density of ungulates
was assessed by distance sampling (line transect, N = 21) and diet composition
of Tigers was evaluated by analysing their faecal samples (N = 66).
The study estimated very low ungulate density (26.87 ± 7.41 individuals
km-2) with highest density of Gaur Bos gaurus
(9.04 individuals km-2) followed by Wild Boar Sus scrofa (8.79 ±
2.73 individuals km-2), whereas, primate density was quite high
(45.89 ± 12.48 individuals km-2), with Nilgiri
Langur Semnopithecus johnii
having the highest density (38.05 ± 10.22 individuals km-2). About 74.62% of the biomass of Gaur
constituted in the Tiger’s diet, consumed lesser than its availability, whereas
Sambar constituted 16.73% of the Tiger diet consumed proportionally to its
availability. Chital Axis axis, Muntjac Muntiacus
muntjak, and Indian Chevrotain Moschiola indica were
not represented in the Tiger’s diet. The
current study is the first scientific information on prey selection of the
Tiger in KMTR landscape, which will serve as a baseline for its conservation
planning and management.
Keywords: Faecal
analysis, food habits, line transect, prey abundance, prey selection.
Introduction
The Tiger Panthera tigris, is the largest among five big cats in the genus
(Sunquist 2010), distributed across the heterogeneous
habitats of Asia (Hayward et al. 2012).
Globally, Tiger population has precipitously declined,
and its range has extensively diminished over the past century (Kerley et al.
2015). Poaching for Tiger body
parts, habitat loss, and degradation and
depletion of prey base have been the major causes for its decline (Karanth et al. 2004; Miquelle et
al. 2010). Despite existence of large
tracts of suitable habitats across Asia, Tigers are absent in many of the
areas, probably due to lack of adequate prey base (Rabinowitz 1993; Check
2006), however, previous studies have emphasised that Tigers are flexible and
recover when their habitat and adequate prey species are well protected
(O’Brien et al. 2003).
Tigers are obligate
terrestrial carnivores, generally preying upon ungulates (Seidensticker
1997), including diverse ranges of species that differ in size such as cervids, bovids, and suids (Andheria et al. 2007; Miquelle et
al. 2010; Hayward et al. 2012). Prey
availability, season, topography, and forest types are some of the significant
ecological variables that influence the dietary habits of Tigers (Sunquist & Sunquist
1999). Studies have also suggested that
predators play a major role in regulating the abundance of herbivore population
in an environment of tropical forest (Karanth et al.
2004), which further results in the cascading effect at each trophic level
(Polis & Strong 1996). Therefore,
understanding of the dietary habits of the Tiger in relation to its prey base
availability is essential for efficient management of wildlife and natural
habitats (Biswas & Sankar 2002; Bagchi et al. 2003).
Most of the information on prey selection of Tiger comes from studies
carried out in semi-arid dry thorn and dry deciduous forests of central India (Bagchi et al. 2003; Biswas & Sankar
2002; Sankar et al. 2010) and tropical moist
deciduous forests of southern India (Karanth & Suquist 1995; Ramesh et al. 2012a; Kumaraguru
et al. 2011). In those areas, Chital was
the dominant prey species in the Tiger’s diet (Johnsingh
1992; Karanth & Sunquist
1995; Venkataraman et al. 1995; Andheria et al.
2007), however, no comprehensive study has been conducted to estimate the
abundance of prey and its selection by Tigers in their distribution range in
the southern Western Ghats. There is
scanty information about predator-prey selection at Kalakkad-Mundanthurai
Tiger Reserve (KMTR) and the lack of such information can be a major limitation
in designing and implementing site-specific conservation measures (Karanth et al. 2003).
Understanding the principal constituents of the Tiger diet is essential
for planning effective conservation policies (Kerley et al. 2015). Thus, the current research aims to assess the
prey selection patterns by the Tiger in the tropical evergreen forest of KMTR.
Study
area
The current study was
carried out between July 2015 and May 2018 in four administrative ranges,
namely, Mundanthurai, Papanasam,
Ambasamudram, and Upper Kodhayar
(Intensive study area, henceforth ISA) of 588km2 in KMTR (900km2),
located in the southern Western Ghats (8.357–8.883 0N &
77.169–77.574 0E) in Tamil Nadu, India (Figure 1). The terrain KMTR is mountainous (the
elevation ranges 100–1,866 m), and the vegetation ranges from dry thorn scrub
to montane wet tropical forest and grassland at high altitudes (Ramesh et al.
2012b). KMTR receives rainfall from both
the south-west (June to September) and the north-east (October to January)
monsoons (Sarkar 2012). The annual
rainfall is about 3,000mm, and the temperature fluctuates between 17°C and 37°C
during the year. This reserve is
bordered by agricultural lands with human settlements (about 145 villages) in
the east (Arjunan et al. 2006), and with forest
tracts of the Neyyar, Peppara,
and Shendurni wildlife sanctuaries in the Ashambu Hill range (Naniwadekar
& Vasudevan 2006) in the west. The
rivers Peyar, Karaiyar, Kavuthalaiyar, Servalar, Chithar, and Pambar and their
tributaries drain into a perennial river called
Tamiraparani.
The sympatric carnivore species found here are the Tiger, the Leopard Panthera pardus,
and the Wild Dog Cuon alpinus. Sambar, Gaur, Chital, Wild Boar, Barking
Deer, and Indian Chevrotain are some of the major prey species that occur in
this reserve. In addition, Asian
Elephant Elephas maximus, Indian Hare Lepus nigricollis,
Bonnet Macaque Macaca radiata, Tufted
Grey Langur Semnopithecus priam, Lion-tailed Macaque Macaca
silenus, Nilgiri Tahr Hemitragus hylocrius, Indian Crested Porcupine Hystrix
indica, Indian Giant Squirrel Ratufa
indica, Grey Jungle Fowl Gallus sonneratii, Red Spurfowl Galloperdix
spadicea, and Indian Peafowl Pavo
cristatus are also found in the reserve.
Field
Methods
Density and biomass
estimation of prey species
The densities of wild prey
were estimated by using the line transect sampling technique (Burnham et al.
1980; Buckland et al. 1993, 2001). The
line transect method has been extensively applied to estimate animal densities
in the tropical forests of southern Asia (Karanth
& Sunquist 1992, 1995; Biswas & Sankar 2002; Jathanna et al.
2003; Bagchi et al. 2003; Edgaonkar
2008; Paliwal 2008; Malla
2009). Permanent transect lines (n=21)
were randomly laid across different habitat types of KMTR by the Tamil Nadu
Forest Department. The transect length
vary from 1.5 (n=3) to 2 (n=18) km. The
total length and sampling effort was 40.50 and 243km, respectively. Six replicates of 21 transects were walked at
dawn (06.30–08.30 h) between January and May 2016 and at dusk (16.30–18.30 h)
between January and May 2017 within the ISA area. Data were collected by a researcher and two
trained observers on every transect walk.
For each detection, the animal bearings were recorded using a look
through compass (KB 20, SUNNTO, Vantaa, Finland), while angular sighting
distance were recorded using a laser range finder (Yardage Pro 850, Bushnell,
Overland Park, Kansas USA). Group size
was also recorded during the transect sampling.
Necessary care was taken while walking on transects to maximize
detectability of animals before they disappeared from sight.
Faecal sample collection
As cryptic and nocturnal
behaviour of the carnivores limit the direct observation of their predatory
behaviour in the wild, faecal samples were collected to determine their food
habits. Large carnivores generally prefer
to travel along forest roads and trails, and as they travel they defecate to
mark their presence and passage (Sunquist 1981; Johnsingh 1983; Smith et al. 1989; Karanth
& Sunquist 2000).
Therefore, faecal samples of Tiger were collected by intensively searching
along such trails, river beds, and open glades from July 2015 to May 2017. All trails were revisited after about two
months for consecutive collection.
Faeces of Tigers were collected only when they were associated with
scraps and tracks. We distinguished
faecal samples between Leopard and Tiger by their diameter and supplementary
evidence such as pugmarks and scrapes (Karanth & Sunquist 1995).
Leopard faeces are much larger, twisted, more coiled between
constriction and deposited on the grassy stripes at the centre or the edges of
forest road (Andheria et al. 2007), whereas, Tiger
faeces appear to be less coiled and have larger distance between two successive
constrictions within a single piece of a faeces (Ramesh 2010). Once a faeces was encountered, a large
portion was collected in a paper envelope for diet analysis. One-fourth of the
faeces was left uncollected to avoid disturbances in Tigers’ territorial
marking. The collected faecal samples
were washed in running water through a nylon mesh (<1mm), later sun-dried in
thin paper pages (Andheria et al. 2007). Following that, the dried faecal samples were
stored in airtight bags individually labelled with date and location for
further identification.
Analytical
Methods
Density and biomass estimation
of prey species
The density of major prey
species of Tiger was estimated using the program ‘DISTANCE’ version 7.2 (Thomas
et al. 2010). To maximise the number of
the sighting, the temporal replicates of each of the line transects were pooled
together and were considered as a single spatial sample (n=21). Different detection functions were fitted to
the observed data and the appropriate model was selected based on the lowest
Akaike information criterion (AIC) values (Burnham et al. 1980; Buckland et al.
1996). Parameters such as effective
strip width (ESW), cluster density (Dg), cluster size (Gs), and animal prey individual density (Di)
were also estimated using program DISTANCE 7.2 (Burnham et al. 1980; Buckland
et al. 1993).
The density of ungulate
commonly represented as the biomass of ungulates available in the
ecosystem. The biomass (kg km-2)
of major prey species was calculated by multiplying the individual density (Di)
of prey species by its average estimated unit weight (Tamang 1982; Wegge et al. 2009) from the available information for major
prey species (Karanth & Sunquist
1992, 1995) (see Appendix 1).
Identification of prey
species
Examination of indigestible
parts of animals and plants found in a predator’s faeces is the primary source
of information about its food habits (Andheria et al.
2007). The prey species were identified
by microscopic examination of the medullary pattern (colour, length, and
thickness of the medulla) in 20 hairs, collected randomly from each faecal
sample (Mukherjee et al. 1994), and later corroborated with reference guides of
Bahuguna et al. (2010) and Chakraborty & De
(2010).
Estimation of frequency of
occurrence and relative biomass of prey consumed
A most commonly used measure
of the frequency of occurrence (henceforth FO) for each prey type was to
estimate the prey intake and composition (Andheria et
al. 2007). The FO, however, does not
provide the best approximation of the true dietary patterns of a predator, as
the biomass consumed to faeces excreted is not alike for all prey species due
to their variation in surface area: volume ratio, described by Floyd et al.
(1978) and Ackerman et al. (1984). To
preclude such bias, we have used the biomass calculation model recently
developed for obligate carnivores by Chakrabarti et al. (2016).
Y = ((0.033- (0.025 × exp (-4.284(X/PBM)))) ×PBM
Where, Y is the mass of prey
consumed per collectable faecal sample, X is the prey body mass, and PBM is the
predator body mass. The mean body weight
of each prey consumed by Tiger was based on Karanth
& Sunquist (1995).
The adequacy of the sample
size was calculated using the Brillouin diversity index (Brillouin 1956).
HB = InN!
– ∑ Inni!/N
Where HB is diversity,
N is the number of the prey taxa in all the samples, and ni is the number of individual prey taxa in the i th category.
Analysis of prey selection
To assess the prey selection
patterns of Tigers for different prey species in KMTR, Jacobs’ index (1974) of
preference (D) was used:
D = (ri-pi)
/ (ri + pi - 2ripi)
Where, ri
is the proportion of a prey remains in faecal sample, and pi is the
proportional density of prey species in the population. The resulting values ranges from +1 (strongly
selected) to -1 (strongly avoided). Prey
selection assessment was restricted to those prey species whose density
information was available.
Results
Density and biomass of prey
species
The overall densities of
ungulates and primates were 26.87± 7.41 km-2 and 45.89 ± 12.48 km-2,
respectively, whereas, densities of Indian Giant Squirrel and Grey Jungle Fowl
were 3.20 ± 1.32 km-2 and 25.32 ± 5.09 km2,
respectively. The estimated individual
and cluster density for potential prey species of a large carnivore is given in
Table 1 along with cluster size and their percentage of the coefficient variation,
and effective stripe width (Appendix 2). Half-normal-cosine was the best fit
model that had resulted in the lowest AIC value for all the species. The major prey species of Tigers are
classified into groups such as ungulates (Chital, Sambar, Mouse Deer, Gaur,
Wild Boar) and primates (Tufted Grey Langur, Nilgiri
Langur, Bonnet Macaque), while Grey Jungle Fowl was also consumed by them. In terms of density of clusters in ungulates,
Wild Boar (3.26 ± 1.29 km-2) were most abundant, followed by Sambar
(2.79 ± 0.57km-2), Gaur (1.88± 0.47 km-2), and Chital
(0.94± 0.32 km-2), whereas density of individual Gaur (9.04 ± 2.03
km-2) was the highest among all the ungulates, followed by Wild Boar
(8.79± 2.73 km-2), Sambar (4.80± 1.04 km-2), Chital
(2.50± 0.92 km-2) and Mouse Deer (1.74± 0.69 km-2). The number of detections for elephants was
too low to permit useful analysis. Total
estimated biomass for ungulates and primates in KMTR was 5,115.20 kg km-2
and 404.51 kg km-2, respectively.
KMTR harboured high density
of primates as individual densities for Nilgiri
Langur, Tufted Grey Langur and Bonnet Macaque were 38.05 ± 10.22 individuals km-2,
6.14 ± 1.73 individuals km-2, and 1.70 ± 0.53 individuals km-2,
where the density of cluster was 4.86 ± 1.15, 0.20 ± 0.01, and 0.22 ± 0.11
clusters km-2, respectively.
Substantial observations of Indian Giant Squirrel (3.20 ± 1.32 km-2)
and Grey Jungle Fowl (25.32 ± 5.09 km-2) were obtained on transects
during the study period.
Prey composition and
selection
After excluding faecal
samples (n = 6) which had an unidentifiable object and were loose/viscous in
consistency, we had a total of 66 Tiger faecal samples. The Brillouin diversity index value for the
estimation of adequacy of the sample size reached 15th faecal, indicating
that we had sampled adequately (Figure 2).
Four species of mammals were identified in the Tiger faecal sample
(Table 2). All faecal samples contained
single prey items. Out of the prey
species identified in the Tiger faeces, Gaur constituted 74.2% followed by
Sambar (16.6%), Sloth Bear (6.06%), and Nilgiri Tahr (3.0%). No
remains of Chital, Muntjac, Mouse Deer, Wild Boar, and primates were found in
the Tiger faeces.
The prey selectivity of a
Tiger was tested by comparing with the individual density of the prey
species. Prey selection analysis was
restricted to seven prey species (Gaur, Sambar, Chital, Mouse Deer, Wild Boar, Nilgiri Langur, and Tufted Grey Langur), whose density
information was available. The Jacobs’
index value showed that Tigers displayed strongest selection of Gaur followed
by Sambar (Figure 3) and apparently avoided other prey in KMTR.
Discussion
Density and prey biomass
Comparative account of total
ungulate densities estimated in the present study (Table 3) with that of other
tropical forests in southern Asia revealed that KMTR harboured lower density of
ungulates than most of them but higher than the Tiger reserves such as Bori-Satpura, Pakke, and
Bhadra. The possible reason for the low
density of ungulates might be the majority rocky outcrops and highly
precipitous terrain. Mid-elevation
forest is dry in most of the place coupled with contiguous tracts (c. 440km2)
of tropical rainforest in KMTR which is unfavourable for ungulates (Johnsingh 2001).
Gaur was found to be most abundant species in the ISA and was comparable
with other Tiger reserves of Western Ghats such as Mudumalai
(Ramesh 2010) and Nagarahole (Karanth
& Sunquist 1992).
Nevertheless, most observation of Gaur were in grassland due to
increased visibility compared to heavily vegetated habitat types in KMTR,
therefore, we presume that this might have influenced the overall density of
Gaur. Therefore, we speculated that true
density of Gaur would be closer to the lower confidence limit of 3.08km-2
and it is similar to the previous study in KMTR by Ramesh et al. (2012b). Gaurs were mostly recorded in the morning
within the wet grasslands of higher altitude, whilst they were observed in the
dry thorn and teak forest during the dusk hours. The density of Wild Boar appears to be
closely comparable to Ranthambore (Bagchi et al. 2003), Barida (Stoen & Wegge 1996), and Katka-Kochikahali of Sundarbans (Reza et al. 2002) but
different from Anamalai Tiger Reserve (Kumaraguru et al. 2011).
Estimated density of Sambar in the current study was comparable to
tropical dry moist deciduous (Bori-Satpura, Badhra, Nagarahole) and tropical
dry thorn, dry deciduous, and evergreen forest habitat of Mudumalai
and Bandipur (Table 3). Sambar density, however, was quite low
compared to Anamalai (Kumaraguru
et al. 2011) and Pench (Acharya 2007).
The density of chital
estimated (2.5 individuals km2) was very low compared to other
tropical forests in southern Asia. In
ISA Chital distribution was restricted to 60km2 of the Mundanthurai plateau (Sathyakumar
2000), which was covered with dry thorny and deciduous vegetation interspersed
with the overgrown teak plantation.
Plateau is dominated by unpalatable tall-grass species Cymbopogon flexuosus (Sankaran
2005) and invasive thickets, such as Lantana Lantana
camara and Eupatorium Eupatorium
glandulosum (Uma et al. 1999). Though cattle grazing has been prohibited in
KMTR since 2000 (Venkatesh et al. 2017), there were substantial number of
cattle grazing in the reservoir (Karaiyar and Manimuthar) and Mundanthurai
Plateau. Thus, the cattle grazing and
lack of suitable grassland might be a potential factor explaining the low
density of Chital. Despite being
nocturnal in nature, we sighted Mouse Deer on transect line, however, no
further analysis could be done as it was a solitary sighting.
We compared the density of
Bonnet Macaque with the estimates available from other tropical forests in
India (Table 4). The density of Bonnet
Macaque was available only for Mudumalai (Ramesh
2010), Nagarahole (Karanth
& Sunquist 1992), Bilgiri
Rangaswamy Tiger Reserve (Kumara
et al. 2012), and Srisi-Honnavar (Babureddy
et al. 2015). Bonnet Macaque density in
KMTR was lower than that of the aforesaid parks. The specialist folivore Tufted Grey Langur
was in low densities but their density was found to be comparable with Bilgiri Rangasamy Tiger
Reserve. In terms of density amongst
ungulates and primates, Nilgiri Langur was found in
high density (38.05 individuals km-2) in ISA. The present study has reported that the
densities have increased as compared to a previous study (Ramesh et al. 2012b).
Prey composition and
selection of tiger
In the current study, the
Tiger preyed on three large ungulates, including Gaur, Sambar, and Nilgiri Tahr. We did not find multiple prey species in a
single sample which is contrary to the prediction of Bekoff
et al. (1984). Gaur accounted for 74.6%
of the Tiger diet by biomass. Such
selective predation towards large body mass was also reported in Anamalai Tiger Reserve (Kumaraguru
et al. 2011), Nagarahole (Karanth
& Sunquist 1995), Bandipur
Tiger Reserve (Andheria et al. 2007), and Pakke Tiger Reserve (Selvan et al. 2013a). Carnivores tend to prefer the most abundant
prey (Breuer 2005). Tiger’s selective
predation for Gaur in the present study area indicates selection for a large
ungulate. Thus, in ISA of KMTR, Gaur
occurred in higher densities (9.04km-2) at wet grassland in high
altitudes interspersed with reed brakes (Ochlandra
sp.), majority of collected faecal samples were found from such habitat, which
suggests that the Tiger prefers habitat where Gaur occur more commonly. Such spatial correlation might have increased
their encounter with the predator.
Crepuscular and poor eyesight of Gaur could have enabled the Tiger to
stalk Gaur easily (Karanth 1993). On the other hand, this selective predation
could also be related to optimal foraging theory (Stephens & Krebs 1987),
which suggests that the selected prey could provide higher benefits in terms of
net biomass intake whilst reduce the cost of handling (stalking, subduing, and disemboweling prey) and injury risks (Scheel 1993). Hence, the predator must shift to profitable
species, which may be either medium-size or high density that make them easier
to be captured (Lamichhane & Jha 2015).
In the current study, Sambar
biomass constituted relatively lesser (16.73%) proportion in the Tiger diet
than other tropical forests of India such as Nagarhole
(Karanth & Sunquist
1992), Sariska (Sankar
& Johnsingh 2002), Ranthambhore
(Bagchi et al. 2003), Bandipur
(Andheria et al. 2007), Satpura
(Edgaongar 2008), and Mudumalai
(Ramesh 2010). This may be due to
spatial distance from the Tiger, as Sambar mostly forage around tea plantation
(personal observations), near human habitation, and dry deciduous and thorn
forest of low elevation. Such spatial
segregation between them might have strengthened the predation on Gaur. Chital, being a common prey for the Tiger in
other protected areas (McDougal 1977; Sunquist 1981; Johnsingh 1983; Karanth & Sunquist 1995; Stoen & Wegge 1996; Biswas & Sankar
2002) was absent in the faecal samples of Tigers in KMTR. This is due to scarce and restricted
distribution of Chital in Mundanthurai Plateau with
low density (Selvan et al. 2013b). This
spatial segregation has compelled the Tiger to depend on Gaur. During the current study, we did not see any
sign of Tigers in Mundanthurai Plateau, which also
corroborates a previous study by Uma et al. (1999).
Presence of Sloth Bear
remains in the Tiger’s faeces reflected the occasional predation on this
species. Predation on bear is not a new
phenomenon, as other investigators also reported the same (Biswas & Sankar 2002; Swaminathan et al. 2002; Harsha et al. 2004; Andheria et al. 2007).
Though the bear remains a relatively minor component of the Tiger diet
relative to Gaur and Sambar, this was more than Nilgiri
Tahr in the current study. One possible explanation is the density of
Sloth Bear and Nilgiri Tahr
in the study area. In addition, Nilgiri Tahr occur only in
restricted cliffs in the present study area (Hopeland
et al. 2016). Conversely, bears are
spread across the study area and are mostly nocturnal and crepuscular (Chauhan
et al. 2004; Yoganand et al. 2005). Such spatial segregation between the Tiger
and the Nilgiri Tahr, while
spatial and temporal overlap between the Tiger and the Sloth Bear, could have
increased encounter rate and led to high predation on Sloth Bear compared to Nigliri Tahr in our study
area. Unfortunately, we could not
determine density of Sloth Bear, Nilgiri Tahr, and their activity pattern on our study site;
therefore, future research is needed to confirm the relationships among
density, prey selection, spatial, and temporal overlap.
The present study revealed
that the moderate prey availability is enough to preserve the Tiger in the long
run in this landscape. Management of
relatively few ungulates, primarily Gaur may be critical for Tiger conservation
in this region.
Table 1. Estimated density of
major prey species of large carnivore in Kalakkad-Mundanthurai
Tiger Reserve. Total sampling effort was 243km.
|
Species |
Model (AIC) |
Min AIC |
Cluster size (SE) |
ESW in meter (SE) |
Dikm-2
(SE) |
%CV (D) (km–2)
95% |
95% CI |
Dg km-2
(SE) |
Biomass kg km-2 |
|
Bonnet Macaque |
Half-normal /
Cosine |
15.913 |
7.88 (2.79) |
48.77 (8.46) |
1.70 (0.53) |
21.15 |
0.50 – 5.81 |
0.22 (0.11) |
6.8 |
|
Tufted Grey Langur |
Half-normal /
Cosine |
9.272 |
30.74 (10.22) |
31.52 (6.77) |
6.14 (1.73) |
23.34 |
1.12 – 33.74 |
0.20 (0.01) |
55.26 |
|
Nilgiri Langur |
Half-normal /
Cosine |
212.77 |
7.82 (0.97) |
34.93 (2.86) |
38.05 (10.22) |
26.82 |
22.33 – 64.87 |
4.86 (1.15) |
342.45 |
|
Total primates |
|
|
|
|
45.89 |
|
|
|
404.51 |
|
Chital |
Half-normal /
Cosine |
15.982 |
2.65 (0.70) |
15.61 (4.37) |
2.50 (0.92) |
18.44 |
0.65 – 9.60 |
0.94 (0.32) |
117.5 |
|
Sambar |
Half-normal /
Cosine |
40.157 |
1.72 (0.24) |
15.03 (2.50) |
4.80 (1.04) |
21.70 |
2.06 – 11.17 |
2.80 (0.57) |
643.2 |
|
Mouse Deer |
Half-normal /
Cosine |
7.79 |
* |
(8.42) (2.65) |
1.74 (0.69) |
19.82 |
0.42 – 7.35 |
* |
5.22 |
|
Gaur |
Half-normal /
Cosine |
37.98 |
4.79 (1.31) |
25.55 (4.08) |
9.04 (2.03) |
28.55 |
3.08 – 26.52 |
1.88 (0.47) |
4068 |
|
Wild Boar |
Half-normal /
Cosine |
33.95 |
2.70 (0.43) |
12.87 (2.18) |
8.79 (2.73) |
31.07 |
2.72 – 28.42 |
3.26 (1.29) |
281.28 |
|
Total ungulates |
|
|
|
|
26.87 |
|
|
|
5115.20 |
|
Indian Giant
Squirrel |
Half-normal /
Cosine |
46.82 |
1.57 (0.19) |
19.33 (3.27) |
3.19 (1.32) |
20.8 |
1.42 – 7.20 |
2.03 (0.81) |
|
|
Grey Jungle Fowl |
Half-normal /
Cosine |
201.98 |
1.6 (0.43) |
15.38 (0.79) |
25.32 (5.09) |
20.12 |
16.82 – 38.11 |
15.82 (3.07) |
|
CV—Coefficient of Variation |
Dg—Density of cluster size | D—Density of individuals | ESW—Effective Stripe
Width | Min AIC—Minimum Akaike information criterion | SE—Standard Error |
CI—95% Confident Interval | *—data not analysed.
Table 2. Food
habit of the Tiger in Kalakkad-Mundanthurai Tiger
Reserve based on faecal analysis (n=66).
|
Prey
species |
Relative
frequency of occurrence % (RFO) |
Mean
body weight (kg) |
Biomass
consumed/faeces |
Biomass
consumed (kg) |
Relative
biomass consumed |
|
Gaur |
74.24 |
287
(Karanth & Sunquist
1995) |
4.95 |
242.50 |
74.62 |
|
Sambar |
16.67 |
212
(Karanth & Sunquist
1995) |
4.94 |
54.35 |
16.73 |
|
Nilgiri
Tahr |
3.03 |
100
(Kumaraguru et al. 2011) |
4.73 |
9.47 |
2.91 |
|
Sloth
Bear |
6.03 |
90
(Biswas & Sankar 2002) |
4.66 |
18.65 |
5.74 |
Table 3.
Comparison of ungulate densities and their biomass (Individuals km-2)
from different protected areas in southern
Asia.
|
Study area |
Chital |
Gaur |
Sambar |
Wild Boar |
Muntjac |
Mouse Deer |
Nilgiri Tahr |
Total ungulate
density |
Total ungulate
biomass |
Source |
|
Current Study |
2.5 |
9.04 |
4.8 |
8.79 |
… |
1.74 |
- |
26.87 |
5115.20 |
|
|
Mudumalai Tiger Reserve |
25.4 |
9.4 |
4.8 |
1.3 |
1.2 |
… |
NP |
42.1 |
6133.8 |
Ramesh (2010) |
|
Keoladeo National Park |
52.37 |
NP |
0.32 |
3.21 |
NP |
… |
NP |
69.58 |
5069.39 |
Aakrithi et al. (2017) |
|
Nagarahole National Park |
50.6 |
9.6 |
5.5 |
4.2 |
4.2 |
… |
NP |
74.1 |
7657.8 |
Karanth & Sunquist (1992) |
|
Anamalai Tiger Reserve |
20.54 |
12.34 |
6.54 |
20.61 |
0.28 |
… |
13.67 |
73.98 |
9181.08 |
Kumaraguru et al. (2011) |
|
Bilgiri Rangasamy Tiger Reserve |
13.96 |
5.08 |
6.01 |
5.33 |
3.7 |
… |
NP |
34.08 |
3995.72 |
Kumara et al. (2012) |
|
Kalakkad-Mundanthurai Tiger Reserve |
… |
3.6 |
7.0 |
1.3 |
… |
… |
NP |
11.9 |
2599.6 |
Ramesh et al.
(2012b) |
|
Bandipur Tiger Reserve |
20.1 |
7 |
5.6 |
… |
0.7 |
… |
NP |
33.4 |
4859.8 |
Karanth & Nichols
(1998) |
|
Bhadra Tiger
Reserve |
8.88 |
3.86 |
4.4 |
2.46 |
4.35 |
… |
NP |
23.95 |
2914.03 |
Gopalaswamy et al. (2012) |
|
Pench National Park |
115.6 |
0.4 |
12.2 |
20.3 |
- |
… |
NP |
149.4 |
8059.6 |
Acharya (2007) |
|
Kanha National Park |
469.7 |
… |
1.5 |
2.5 |
0.6 |
… |
NP |
57.3 |
3103.5 |
Karanth & Nichols
(1998) |
|
Bardia National Park |
77.7 |
NP |
… |
8.8 |
1.7 |
NP |
NP |
99.2 |
4786.5 |
Stoen & Wegge (1996) |
|
Bori-Satpura Tiger Reserve |
5.4 |
… |
4 |
1.8 |
0.8 |
NP |
NP |
13.6 |
1152.2 |
Edganokar (2008) |
|
Ranthambore National Park |
31 |
… |
17.1 |
9.7 |
… |
NP |
NP |
74.8 |
6228.4 |
Bagchi et al. (2003) |
|
Gir National Park |
50.8 |
NP |
2 |
- |
… |
NP |
NP |
56.2 |
2819.22 |
Khan et al. (1996) |
|
Sariska Tiger Reserve |
33.88 |
NP |
26.38 |
54.12 |
NP |
NP |
NP |
157.1 |
14548.72 |
Mondal et al.
(2011) |
|
Chitwan National
Park |
61.8 |
… |
20 |
3.6 |
… |
NP |
NP |
85.4 |
5699.8 |
Sunquist (1981) |
|
Kaziranga National Park |
NP |
… |
… |
2.6 |
… |
NP |
NP |
58.1 |
4815.6 |
Karanth & Nichols
(1998) |
|
Rajaji National
Park |
49.9 |
NP |
14.6 |
1.9 |
NP |
NP |
NP |
68.8 |
4794.5 |
Harihar et al.
(2009) |
|
Pakke Tiger Reserve |
NP |
3.5 |
3.8 |
6.7 |
3.9 |
NP |
NP |
17.9 |
2380.5 |
Selvan et al.
(2013a) |
|
Sundarbans |
70.4 |
NP |
NP |
7.9 |
NP |
NP |
NP |
78.3 |
3561.6 |
Reza et al. (2002) |
Notes: NP - The respective
species was not found in the respective area; ... - Data were not reported
Table 4. Comparison of arboreal prey densities (individuals km-2) and
biomass from protected areas in Indian subcontinent.
|
Study area |
Tufted Grey Langur
(previously known as common langur) |
Nilgiri Langur |
Bonnet Macaque |
Total primate
density |
Total primate
biomass |
Reference |
|
KMTR (Present
study) |
6.14 |
38.05 |
1.7 |
45.89 |
404.51 |
|
|
Mudumalai Tiger Reserve |
35.4 |
… |
1.9 |
37.3 |
340.6 |
Ramesh (2010) |
|
Nagarahole National Park |
23.8 |
… |
5.5 |
29.3 |
236.2 |
Karanth & Sunquist (1992) |
|
Bilgiri Rangasamy Tiger reserve |
6.34 |
NP |
6.56 |
12.9 |
83.3 |
Kumara et al. (2012) |
|
Kalakkad–Mundanthurai Tiger Reserve |
… |
9.9 |
… |
9.9 |
89.1 |
Ramesh et al.
(2012) |
|
Sirsi-Honnavar |
25.06 |
NP |
12.4 |
37.46 |
275.14 |
Babureddy et al. (2015) |
|
Badhra Tiger Reserve |
22.6 |
NP |
… |
22.6 |
203.4 |
Jathanna et al. (2003) |
|
Pench Tiger Reserve |
65.8 |
NP |
… |
65.8 |
592.2 |
Acharya (2007) |
|
Bori – Satpura |
28.3 |
NP |
NP |
28.3 |
254.7 |
Edganokar (2008) |
|
Melghat |
42.92 |
NP |
NP |
42.92 |
386.28 |
Narasimmarajan et al. (2014) |
|
Bardia National Park |
2.3 |
NP |
NP |
2.3 |
20.7 |
Stoen & Wegge (1996) |
|
Ranthambore National Park |
21.75 |
NP |
NP |
21.75 |
195.75 |
Bagchi et al. (2004) |
|
Sariska Tiger Reserve |
50.67 |
NP |
NP |
50.67 |
456.03 |
Mondal et al.
(2011) |
|
Chitawan National Park |
3.6 |
NP |
NP |
3.6 |
32.4 |
Sunquist (1981) |
|
Chilla range of Rajaji
National Park |
14.1 |
NP |
NP |
14.1 |
126.9 |
Harihar et al.
(2009) |
Notes: NP - The respective
species was not found in the respective area; ... - Data were not reported
For figures & appendix - - click here
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